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IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="carolina" checkinTime="1732847695013" checkinUser="plazi" docAuthor="Püschel, Hans P., Shelley, Sarah L., Williamson, Thomas E., Perini, Fernando A., Wible, John R. & Brusatte, Stephen L." docDate="2024" docId="03B9B735FFE00D17C4BFFC858DC3FD9E" docLanguage="en" docName="zlae095.pdf" docOrigin="Zoological Journal of the Linnean Society 202 (1)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlae095" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.9:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="9" docTitle="Macraucheniidae" docType="treatment" docVersion="1" lastPageNumber="15" masterDocId="FF80CF4DFFE60D1FC702FFD68932FFA1" masterDocTitle="A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates" masterLastPageNumber="50" masterPageNumber="1" pageNumber="7" updateTime="1733769861150" updateUser="carolina"> +<mods:mods id="D7D95EC2363A918C3F7321B5DF085776" xmlns:mods="http://www.loc.gov/mods/v3"> +<mods:titleInfo id="EE0403DD4B2B4B0E000E300311D8DCE1"> +<mods:title id="7AC741E94002CD6AC45597B70223791A">A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates</mods:title> +</mods:titleInfo> +<mods:name id="DF1E85D99F1831A8BBAE861F0B5D56D6" type="personal"> +<mods:role id="52F06FD04CA0CEA962D1F054334F0472"> +<mods:roleTerm id="C4D1C500D7D44B360800ECF39CE9FB2A">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="91831FF0DCBCBF484F502315954AA81E">Püschel, Hans P.</mods:namePart> +<mods:affiliation id="7BAB10466E1BDF93FC10AFA8F0106F0C">Millenium Nucleus Early Evolutionary Transitions of Mammals (EVOTEM), Red Palentológica U-Chile, Santiago, 7800003, Chile & School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, EH 9 3 FE, United 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box="[957,1119,851,877]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +( +<figureCitation id="132B1AA6FFE00D19C36CFC828DDCFCCC" box="[1134,1262,851,877]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="6" pageNumber="7">Figs 4A, D–E</figureCitation> +, +<figureCitation id="132B1AA6FFE00D19C3FAFC858C07FCCC" box="[1272,1333,851,877]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="6" pageNumber="7">5A–D</figureCitation> +) +</emphasis> +</paragraph> +</subSubSection> +<subSubSection id="C30A55A8FFE00D17C43BFCAF8DC3FD9E" lastPageId="8" lastPageNumber="9" pageId="6" pageNumber="7" type="multiple"> +<paragraph id="8BAF0623FFE00D19C43BFCAF8ADEF8BF" blockId="6.[824,1476,851,1979]" pageId="6" pageNumber="7"> +With 18 currently accepted genera, +<taxonomicName id="4C107DA0FFE00D19C3C5FCAF8C45FC30" box="[1223,1399,889,913]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +is the second most diverse family of litopterns (Supporting information, +<tableCitation id="C6923398FFE00D19C472FC6E8AF9FC71" box="[880,971,952,976]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="6" pageNumber="7">Table S1</tableCitation> +), being particularly diverse during the Neogene alongside proterotheriids ( +<figureCitation id="132B1AA6FFE00D19C34CFC018D85FC4E" box="[1102,1207,983,1007]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="6" pageNumber="7">Fig. 2B–C</figureCitation> +). It was first proposed by +<bibRefCitation id="EF817BD2FFE00D19C439FC218AEBFBAF" author="Gervais" box="[827,985,1014,1039]" firstAuthor="Gervais" pageId="6" pageNumber="7" pagination="1 - 63" refId="ref42782" refString="Gervais MP. Recherches sur les mammiferes fossiles de l'Amerique Meridonale. In: de Castelnau F (ed.), Animaux Nouveaux ou Rares Recueillis Pendant l'expedition Dans les Parties Centrales de l'Amerique du Sudde Rio de Janeiro a Lima, et de Lima au Para. Paris: P. Bertrand, Libraire-Editeur, 1855, 1 - 63." type="book chapter" year="1855">Gervais (1855)</bibRefCitation> +as a member of the order +<taxonomicName id="4C107DA0FFE00D19C3DEFC218C5CFBAE" box="[1244,1390,1015,1039]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="7" phylum="Arthropoda" rank="order">Perissodactyla</taxonomicName> +(‘Ordre des Jumentés’) to include one species, +<taxonomicName id="4C107DA0FFE00D19C3BFFBC08A88FBEC" authority="Owen, 1838" authorityName="Owen" authorityYear="1838" class="Mammalia" family="Macraucheniidae" genus="Macrauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="patachonica"> +<emphasis id="B964DA31FFE00D19C3BFFBC08CF3FB8F" box="[1213,1473,1046,1070]" italics="true" pageId="6" pageNumber="7">Macrauchenia patachonica</emphasis> +Owen, 1838 +</taxonomicName> +. This taxon was discovered by Charles Darwin in 1834 ( +<bibRefCitation id="EF817BD2FFE00D19C47EFB838D33FBCC" author="Keynes" box="[892,1025,1109,1133]" firstAuthor="Keynes" pageId="6" pageNumber="7" refId="ref43468" refString="Keynes RD. Charles Darwin's Beagle Diary. Cambridge: Cambridge University Press, 2001." type="book" year="2001">Keynes 2001</bibRefCitation> +) and later described by Owen (1838), being the first litoptern known to the scientific community. Later, +<bibRefCitation id="EF817BD2FFE00D19C43BFB458ACDFB0A" author="Ameghino" box="[825,1023,1171,1195]" firstAuthor="Ameghino" pageId="6" pageNumber="7" pagination="71 - 138" refId="ref39328" refString="Ameghino F. Premiere contribution a la connaissance de la faune mammalogique des couches a Colpodon. Boletin de la Academia Nacional de Ciencias en Cordoba 1902 a; 17: 71 - 138." type="journal article" year="1902" yearSuffix="a">Ameghino (1902a)</bibRefCitation> +, based mostly on the position of the external nares in the cranium, subdivided this family into three subfamilies: +<taxonomicName id="4C107DA0FFE00D19C46CFB048D11FB4B" authorityName="Soria" authorityYear="1981" box="[878,1059,1234,1258]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="subFamily" subFamily="Cramaucheniinae">Cramaucheniinae</taxonomicName> +represented by +<taxonomicName id="4C107DA0FFE00D19C3C0FB048A4AFAA8" authority="Ameghino 1902 a" authorityName="Ameghino" authorityYear="1902" class="Mammalia" family="Macraucheniidae" genus="Cramauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C3C0FB048C63FB4B" box="[1218,1361,1234,1258]" italics="true" pageId="6" pageNumber="7">Cramauchenia</emphasis> +<bibRefCitation id="EF817BD2FFE00D19C254FB048A4AFAA8" author="Ameghino" firstAuthor="Ameghino" pageId="6" pageNumber="7" pagination="71 - 138" refId="ref39328" refString="Ameghino F. Premiere contribution a la connaissance de la faune mammalogique des couches a Colpodon. Boletin de la Academia Nacional de Ciencias en Cordoba 1902 a; 17: 71 - 138." type="journal article" year="1902" yearSuffix="a">Ameghino 1902a</bibRefCitation> +</taxonomicName> +from Deseadean to Colhuehuapian South American Land Mammal Ages [hereafter SALMAs; Stages of +<bibRefCitation id="EF817BD2FFE00D19C257FAC78AE3FAE9" author="Cione and Tonni" firstAuthor="Cione" pageId="6" pageNumber="7" pagination="135 - 69" refId="ref41225" refString="Cione A, Tonni E. Chronostratigraphy and ' land-mammal ages' in the Cenozoic of southern South America: principles, practices, and the ' Uquian' problem. Journal of Palentology 1995; 69: 135 - 69." type="journal article" year="1995">Cione and Tonni (1995)</bibRefCitation> +], Theosodontinae represented by +<taxonomicName id="4C107DA0FFE00D19C259FAE68AD4FAC6" authority="Ameghino, 1887" authorityName="Ameghino" authorityYear="1887" class="Mammalia" family="Macraucheniidae" genus="Theosodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C259FAE68CF0FAE9" box="[1371,1474,1328,1352]" italics="true" pageId="6" pageNumber="7">Theosodon</emphasis> +<bibRefCitation id="EF817BD2FFE00D19C43BFA998AD4FAC6" author="Ameghino" box="[825,998,1359,1383]" firstAuthor="Ameghino" pageId="6" pageNumber="7" pagination="1 - 26" refId="ref39043" refString="Ameghino F. Enumeracion sistematica de las especies de mamiferos fosiles coleccionados por Carlos Ameghino en los terrenos Eocenos de la Patagonia Austral y depositados en el Museo de La Plata. Boletin del Museo de La Plata 1887; 1: 1 - 26." type="journal article" year="1887">Ameghino, 1887</bibRefCitation> +</taxonomicName> +(Colhuehuapian to Laventan SALMAs), and +<taxonomicName id="4C107DA0FFE00D19C43BFAB98ADBFA26" authorityName="Gervais" authorityYear="1855" box="[825,1001,1391,1415]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="subFamily" subFamily="Macraucheniinae">Macraucheniinae</taxonomicName> +represented by +<taxonomicName id="4C107DA0FFE00D19C3A6FAB98C1CFA26" authorityName="Owen" authorityYear="1838" box="[1188,1326,1391,1415]" class="Mammalia" family="Macraucheniidae" genus="Macrauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C3A6FAB98C1CFA26" box="[1188,1326,1391,1415]" italics="true" pageId="6" pageNumber="7">Macrauchenia</emphasis> +</taxonomicName> +(Pliocene to Holocene). Simpson (1945) reclassified +<taxonomicName id="4C107DA0FFE00D19C3E3FA588CA3FA07" box="[1249,1425,1422,1446]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +into the subfamilies +<taxonomicName id="4C107DA0FFE00D19C4E9FA7B8DA9FA64" authorityName="Gervais" authorityYear="1855" box="[1003,1179,1453,1477]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="subFamily" subFamily="Macraucheniinae">Macraucheniinae</taxonomicName> +and +<taxonomicName id="4C107DA0FFE00D19C3E2FA7B8C6DFA64" box="[1248,1375,1453,1477]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="subFamily" subFamily="Adianthinae">Adianthinae</taxonomicName> +, without giving an anatomical justification for this change ( +<tableCitation id="C6923398FFE00D19C240FA1B8CBDFA44" box="[1346,1423,1485,1509]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="6" pageNumber="7">Table 1</tableCitation> +). By Simpson’s (1945) definition, +<taxonomicName id="4C107DA0FFE00D19C362FA3A8C22F9A5" authorityName="Gervais" authorityYear="1855" box="[1120,1296,1516,1540]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="subFamily" subFamily="Macraucheniinae">Macraucheniinae</taxonomicName> +was composed of the members of the three subfamilies mentioned by +<bibRefCitation id="EF817BD2FFE00D19C25AF9DD8ABCF9E2" author="Ameghino" firstAuthor="Ameghino" pageId="6" pageNumber="7" pagination="71 - 138" refId="ref39328" refString="Ameghino F. Premiere contribution a la connaissance de la faune mammalogique des couches a Colpodon. Boletin de la Academia Nacional de Ciencias en Cordoba 1902 a; 17: 71 - 138." type="journal article" year="1902" yearSuffix="a">Ameghino (1902a)</bibRefCitation> +and close Neogene relatives discovered since then (e.g., +<taxonomicName id="4C107DA0FFE00D19C475F99C8DDAF9C3" authority="Ameghino, 1904 a" authorityName="Ameghino" authorityYear="1904" box="[887,1256,1610,1634]" class="Mammalia" family="Macraucheniidae" genus="Promacrauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C475F99C8D13F9C3" box="[887,1057,1610,1634]" italics="true" pageId="6" pageNumber="7">Promacrauchenia</emphasis> +<bibRefCitation id="EF817BD2FFE00D19C32FF99C8DDAF9C3" author="Ameghino" box="[1069,1256,1610,1634]" firstAuthor="Ameghino" pageId="6" pageNumber="7" refId="ref39396" refString="Ameghino F. Nuevas especies de mamiferos cretaceos y terciarios de la Republica Argentina. Anales de la Sociedad Cientifica Argentina 1904 a; 58: 561 - 142." type="journal volume" year="1904" yearSuffix="a">Ameghino, 1904a</bibRefCitation> +</taxonomicName> +), but also included +<taxonomicName id="4C107DA0FFE00D19C43BF9BF8DB2F920" authority="Bordas, 1939" authorityName="Bordas" authorityYear="1939" box="[825,1152,1641,1665]" class="Mammalia" family="Macraucheniidae" genus="Paramacrauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C43BF9BF8ADCF920" box="[825,1006,1641,1665]" italics="true" pageId="6" pageNumber="7">Paramacrauchenia</emphasis> +<bibRefCitation id="EF817BD2FFE00D19C4F5F9BF8DB2F920" author="Bordas" box="[1015,1152,1641,1665]" firstAuthor="Bordas" pageId="6" pageNumber="7" pagination="413 - 33" refId="ref40435" refString="Bordas AF. Diagnosis sobre algunos mamiferos de las capas con Colpodon, del valle del rio Chubut (Republica Argentina). Physis 1939; 14: 413 - 33." type="journal article" year="1939">Bordas, 1939</bibRefCitation> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFE00D19C3B5F9BF8A5FF900" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C3B5F9BF8C7BF920" box="[1207,1353,1641,1665]" italics="true" pageId="6" pageNumber="7">Victorlemoinea</emphasis> +<bibRefCitation id="EF817BD2FFE00D19C250F9BF8A5FF900" author="Ameghino" firstAuthor="Ameghino" pageId="6" pageNumber="7" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +, the former now considered a proterotheriid and the latter a sparnotheriodontid (Soria 2001; see +<taxonomicName id="4C107DA0FFE00D19C3E2F97E8CF0F961" box="[1248,1474,1704,1728]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +section for more details). The subfamily +<taxonomicName id="4C107DA0FFE00D19C3DEF9118C6BF97E" box="[1244,1369,1735,1759]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="subFamily" subFamily="Adianthinae">Adianthinae</taxonomicName> +was composed of adianthids such as +<taxonomicName id="4C107DA0FFE00D19C358F9318D8DF95E" authorityName="Ameghino" authorityYear="1891" box="[1114,1215,1767,1791]" class="Mammalia" family="Adianthidae" genus="Adianthus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C358F9318D8DF95E" box="[1114,1215,1767,1791]" italics="true" pageId="6" pageNumber="7">Adianthus</emphasis> +</taxonomicName> +(see +<taxonomicName id="4C107DA0FFE00D19C3F7F9318C40F95E" box="[1269,1394,1767,1791]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +section for more details). +</paragraph> +<paragraph id="8BAF0623FFE00D17C457F8F38896FE63" blockId="6.[824,1476,851,1979]" lastBlockId="8.[126,779,144,1641]" lastPageId="8" lastPageNumber="9" pageId="6" pageNumber="7"> +Similarly to +<bibRefCitation id="EF817BD2FFE00D19C4D2F8F38DA7F89C" author="Ameghino" box="[976,1173,1829,1853]" firstAuthor="Ameghino" pageId="6" pageNumber="7" pagination="71 - 138" refId="ref39328" refString="Ameghino F. Premiere contribution a la connaissance de la faune mammalogique des couches a Colpodon. Boletin de la Academia Nacional de Ciencias en Cordoba 1902 a; 17: 71 - 138." type="journal article" year="1902" yearSuffix="a">Ameghino (1902a)</bibRefCitation> +, Soria (1981) recognized important anatomical differences between macraucheniids before and after the Chasicoan SALMA (Late Miocene) that justified a subfamilial division, separating +<taxonomicName id="4C107DA0FFE00D19C3AAF8558C6AF83A" box="[1192,1368,1923,1947]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +into two subfamilies: +<taxonomicName id="4C107DA0FFE00D19C4C2F8758D47F81A" authorityName="Soria" authorityYear="1981" box="[960,1141,1955,1979]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="subFamily" subFamily="Cramaucheniinae">Cramaucheniinae</taxonomicName> +(pre-Chasicoan SALMA) and +<taxonomicName id="4C107DA0FFEE0D17C783FF468803FF09" authorityName="Gervais" authorityYear="1855" box="[129,305,144,168]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Macraucheniinae">Macraucheniinae</taxonomicName> +(Chasicoan and post-Chasicoan SALMA or post-Santacrucian SALMA). This separation was based on the fact that +<taxonomicName id="4C107DA0FFEE0D17C7DFFF1988BFFF46" authorityName="Gervais" authorityYear="1855" box="[221,397,207,231]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Macraucheniinae">Macraucheniinae</taxonomicName> +tends to have more derived features than +<taxonomicName id="4C107DA0FFEE0D17C7B7FF388859FEA7" authorityName="Soria" authorityYear="1981" box="[181,363,238,262]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Cramaucheniinae">Cramaucheniinae</taxonomicName> +, possessing a retracted nasal aperture to a more centrodorsal position and fused zeugopodial elements in the forelimbs (ulna-radius) and hind limbs (tibia-fibula; Soria 1981). However, of these two subfamilies only +<taxonomicName id="4C107DA0FFEE0D17C55BFE9A8A3BFEC5" authorityName="Gervais" authorityYear="1855" box="[601,777,332,356]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Macraucheniinae">Macraucheniinae</taxonomicName> +is monophyletic according to phylogenetic analyses (Schmidt and Ferrero 2014, +<bibRefCitation id="EF817BD2FFEE0D17C64AFE5D8B2CFE02" author="Forasiepi" box="[328,542,395,419]" etAl="et al." firstAuthor="Forasiepi" pageId="8" pageNumber="9" pagination="1 - 76" refId="ref41925" refString="Forasiepi AM, MacPhee RDE, Hernandez del Pino S et al. Exceptional skull of Huayqueriana (Mammalia, Litopterna, Macraucheniidae) from the Late Miocene of Argentina: anatomy, systematics, and paleobiological implications. Bulletin of the American Museum of Natural History 2016; 404: 1 - 76." type="journal article" year="2016"> +Forasiepi +<emphasis id="B964DA31FFEE0D17C6ADFE5D88D0FE02" box="[431,482,395,419]" italics="true" pageId="8" pageNumber="9">et al.</emphasis> +2016 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFEE0D17C52FFE5D8A36FE02" author="McGrath" box="[557,772,395,419]" etAl="et al." firstAuthor="McGrath" pageId="8" pageNumber="9" refId="ref44292" refString="McGrath AJ, Anaya F, Croft DA. Two new macraucheniids (Mammalia: Litopterna) from the Late Middle Miocene (Laventan South American Land Mammal Age) of Quebrada Honda, Bolivia. Journal of Vertebrate Paleontology 2018; 38: e 1461632. https: // doi. org / 10.10 80 / 02724634.2018.1461632" type="journal volume" year="2018"> +McGrath +<emphasis id="B964DA31FFEE0D17C597FE5D8BF5FE02" box="[661,711,395,419]" italics="true" pageId="8" pageNumber="9">et al.</emphasis> +2018 +</bibRefCitation> +, Püschel +<emphasis id="B964DA31FFEE0D17C7D7FE7D8836FE63" box="[213,260,426,450]" italics="true" pageId="8" pageNumber="9">et al.</emphasis> +2023; +<tableCitation id="C6923398FFEE0D17C64BFE7C88A6FE63" box="[329,404,426,450]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="8" pageNumber="9">Table 2</tableCitation> +). +</paragraph> +<caption id="DF6F56ABFFE00D19C7CDF8A889DAF8C3" box="[207,232,1787,1973]" isContinuationCaption="true" pageId="6" pageNumber="7" startId="6.[207,232,1918,1973]" targetBox="[260,666,147,1973]" targetIsTable="true" targetPageId="6"> +<paragraph id="8BAF0623FFE00D19C7CDF8A889DAF8C3" blockId="6.[207,232,1787,1973]" box="[207,232,1787,1973]" pageId="6" pageNumber="7"> +Table +<emphasis id="B964DA31FFE00D19C7CDF8BA89DAF8CD" bold="true" box="[207,232,1895,1913]" pageId="6" pageNumber="7">2.</emphasis> +Continued +</paragraph> +</caption> +<paragraph id="8BAF0623FFE00D19C606F8B48BA8FC5E" pageId="6" pageNumber="7"> +<table id="F910F483FFE0F2E0C606FF458BA8F814" box="[260,666,147,1973]" gridcols="18" gridrows="5" pageId="6" pageNumber="7" textDirection="bu"> +<tr id="35200461FFE0F2E0C606FF458809F814" box="[260,315,147,1973]" gridrow="0" pageId="6" pageNumber="7" rowspan-12="1" rowspan-13="1" rowspan-16="1" rowspan-7="1" textDirection="bu"> +<th id="76F16D1DFFE0F2E0C606F9F08809F814" box="[260,315,1574,1973]" gridcol="0" gridrow="0" pageId="6" pageNumber="7" textDirection="bu"> +Analysis +<emphasis id="B964DA31FFE00D19C606F900882FF957" bold="true" box="[260,285,1750,1782]" pageId="6" pageNumber="7">No</emphasis> +. +<emphasis id="B964DA31FFE00D19C606F953882FF904" bold="true" box="[260,285,1669,1701]" pageId="6" pageNumber="7">No</emphasis> +. +<emphasis id="B964DA31FFE00D19C606F9E7882FF990" bold="true" box="[260,285,1580,1617]" pageId="6" pageNumber="7">No.</emphasis> +<emphasis id="B964DA31FFE00D19C620F9068809F958" bold="true" box="[290,315,1744,1785]" pageId="6" pageNumber="7">taxa</emphasis> +<emphasis id="B964DA31FFE00D19C620F9AC8809F910" bold="true" box="[290,315,1658,1713]" pageId="6" pageNumber="7">Litop</emphasis> +<emphasis id="B964DA31FFE00D19C620F9A38809F9D8" bold="true" box="[290,315,1653,1657]" pageId="6" pageNumber="7">.</emphasis> +<emphasis id="B964DA31FFE00D19C620F9FC8809F9F7" bold="true" box="[290,315,1578,1622]" pageId="6" pageNumber="7">char</emphasis> +<emphasis id="B964DA31FFE00D19C620F9F08809F98B" bold="true" box="[290,315,1574,1578]" pageId="6" pageNumber="7">.</emphasis> +</th> +<th id="76F16D1DFFE0F2E0C606FA558809F9A9" box="[260,315,1411,1544]" gridcol="1" gridrow="0" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C606FA69882FFA50" bold="true" box="[260,285,1471,1521]" pageId="6" pageNumber="7">Char</emphasis> +<emphasis id="B964DA31FFE00D19C606FA6D882FFA1E" bold="true" box="[260,285,1467,1471]" pageId="6" pageNumber="7">.</emphasis> +<emphasis id="B964DA31FFE00D19C606FA5C882FFA17" bold="true" box="[260,285,1418,1462]" pageId="6" pageNumber="7"> +<typeStatus id="54ABB881FFE00D19C606FA5C882FFA17" box="[260,285,1418,1462]" pageId="6" pageNumber="7">type</typeStatus> +</emphasis> +</th> +<th id="76F16D1DFFE0F2E0C606FBB18809FAC4" box="[260,315,1127,1381]" colspan="5" colspanRight="4" gridcol="2" gridrow="0" pageId="6" pageNumber="7" textDirection="bu">Litoptern families</th> +<th id="76F16D1DFFE0F2E0C606FC808809FBB3" box="[260,315,854,1042]" colspan="4" colspanRight="3" gridcol="8" gridrow="0" pageId="6" pageNumber="7" textDirection="bu">Other families</th> +<th id="76F16D1DFFE0F2E0C606FDB88809FD67" box="[260,315,622,710]" colspan="2" colspanRight="1" gridcol="14" gridrow="0" pageId="6" pageNumber="7" textDirection="bu">Orders</th> +<th id="76F16D1DFFE0F2E0C606FF458809FD80" box="[260,315,147,545]" gridcol="17" gridrow="0" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C606FE55882FFE1F" bold="true" box="[260,285,387,446]" pageId="6" pageNumber="7">Focus</emphasis> +<emphasis id="B964DA31FFE00D19C606FEBC882FFEDF" bold="true" box="[260,285,362,382]" pageId="6" pageNumber="7">of</emphasis> +the analysis +</th> +</tr> +<tr id="35200461FFE0F2E0C64CFF458855F814" box="[334,359,147,1973]" gridrow="1" pageId="6" pageNumber="7" rowspan-0="1" rowspan-17="1" textDirection="bu"> +<td id="76F16D1DFFE0F2E0C64CFA558855F9A9" box="[334,359,1411,1544]" gridcol="1" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFA258855F9A4" bold="true" box="[334,359,1523,1541]" pageId="6" pageNumber="7">D</emphasis> +<emphasis id="B964DA31FFE00D19C64CFA178855FA73" bold="true" box="[334,359,1473,1490]" pageId="6" pageNumber="7">C</emphasis> +<emphasis id="B964DA31FFE00D19C64CFA558855FA02" bold="true" box="[334,359,1411,1443]" pageId="6" pageNumber="7">PC</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFADA8855FAC4" box="[334,359,1292,1381]" colspan="2" colspanRight="1" gridcol="2" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFA9E8855FAC4" bold="true" box="[334,359,1352,1381]" pageId="6" pageNumber="7">Ad</emphasis> +An +</td> +<td id="76F16D1DFFE0F2E0C64CFB1B8855FB4F" box="[334,359,1229,1262]" gridcol="4" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFB1B8855FB4C" bold="true" box="[334,359,1229,1261]" pageId="6" pageNumber="7">Ma</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFB408855FB0E" box="[334,359,1174,1199]" gridcol="5" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFB408855FB0E" bold="true" box="[334,359,1174,1199]" pageId="6" pageNumber="7">Pe</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFBB18855FBD6" box="[334,359,1127,1143]" gridcol="6" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFBBF8855FBD7" bold="true" box="[334,359,1129,1142]" pageId="6" pageNumber="7">S</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFBE68855FBE8" box="[334,359,1072,1097]" gridcol="7" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFBE68855FBE8" bold="true" box="[334,359,1072,1097]" pageId="6" pageNumber="7">Di</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFBD48855FBB3" box="[334,359,1026,1042]" gridcol="8" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFBD48855FBAA" bold="true" box="[334,359,1026,1035]" pageId="6" pageNumber="7">I</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFC1C8855FC7F" box="[334,359,970,990]" gridcol="9" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFC1C8855FC7A" bold="true" box="[334,359,970,987]" pageId="6" pageNumber="7">K</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFC588855FC0B" box="[334,359,910,938]" gridcol="10" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFC588855FC0B" bold="true" box="[334,359,910,938]" pageId="6" pageNumber="7">Mi</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFC808855FCD1" box="[334,359,854,880]" gridcol="11" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFC808855FCD1" bold="true" box="[334,359,854,880]" pageId="6" pageNumber="7">Nt</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFCCA8855FC96" box="[334,359,796,823]" gridcol="12" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFCCA8855FC96" bold="true" box="[334,359,796,823]" pageId="6" pageNumber="7">Po</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFD328855FD5F" box="[334,359,740,766]" gridcol="13" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFD328855FD5F" bold="true" box="[334,359,740,766]" pageId="6" pageNumber="7">As</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFD718855FD67" box="[334,359,679,710]" gridcol="14" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFD718855FD67" bold="true" box="[334,359,679,710]" pageId="6" pageNumber="7">Nu</emphasis> +</td> +<td id="76F16D1DFFE0F2E0C64CFDB88855FD28" box="[334,359,622,649]" gridcol="15" gridrow="1" pageId="6" pageNumber="7" textDirection="bu">Py</td> +<td id="76F16D1DFFE0F2E0C64CFDE98855FDF1" box="[334,359,575,592]" gridcol="16" gridrow="1" pageId="6" pageNumber="7" textDirection="bu"> +<emphasis id="B964DA31FFE00D19C64CFDE98855FDEE" bold="true" box="[334,359,575,591]" pageId="6" pageNumber="7">X</emphasis> +</td> +</tr> +<tr id="35200461FFE0F2E0C679FF458886F814" box="[379,436,147,1973]" gridrow="2" pageId="6" pageNumber="7" rowspan-10="1" rowspan-11="1" rowspan-12="1" rowspan-13="1" rowspan-14="1" rowspan-15="1" rowspan-16="1" rowspan-3="1" rowspan-5="1" rowspan-6="1" rowspan-7="1" rowspan-8="1" rowspan-9="1" textDirection="bu"> +<th id="76F16D1DFFE0F2E0C679F9F08886F814" box="[379,436,1574,1973]" gridcol="0" gridrow="2" pageId="6" pageNumber="7" textDirection="bu"> +Püschel et +<emphasis id="B964DA31FFE00D19C67FF8E988A4F8F3" box="[381,406,1855,1874]" italics="true" pageId="6" pageNumber="7">al</emphasis> +. 21 21 43 (2023) +</th> +<td id="76F16D1DFFE0F2E0C679FA558886F9A9" box="[379,436,1411,1544]" gridcol="1" gridrow="2" pageId="6" pageNumber="7" textDirection="bu">X X X</td> +<td id="76F16D1DFFE0F2E0C679FA998886FAC4" box="[379,436,1359,1381]" gridcol="2" gridrow="2" pageId="6" pageNumber="7" textDirection="bu"> +<superScript id="7C65AB6BFFE00D19C679FA9988B8FAF4" attach="right" box="[379,394,1359,1365]" fontSize="6" pageId="6" pageNumber="7">Xc</superScript> +</td> +<td id="76F16D1DFFE0F2E0C679FB1B8886FB4F" box="[379,436,1229,1262]" gridcol="4" gridrow="2" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C679FF458886FD80" box="[379,436,147,545]" gridcol="17" gridrow="2" pageId="6" pageNumber="7" textDirection="bu"> +Affinities of +<taxonomicName id="4C107DA0FFE00D19C67FFEE588A4FE0F" box="[381,406,307,430]" class="Magnoliopsida" family="Asteraceae" genus="Micrauchenia" kingdom="Plantae" order="Asterales" pageId="6" pageNumber="7" phylum="Tracheophyta" rank="genus"> +<emphasis id="B964DA31FFE00D19C67FFEE588A4FE0F" box="[381,406,307,430]" italics="true" pageId="6" pageNumber="7">Micrauchenia</emphasis> +</taxonomicName> +<emphasis id="B964DA31FFE00D19C67FFF0388A4FE8F" box="[381,406,213,302]" italics="true" pageId="6" pageNumber="7">saladensis</emphasis> +within +<taxonomicName id="4C107DA0FFE00D19C699FEBF8886FDAD" box="[411,436,361,524]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +</td> +</tr> +<tr id="35200461FFE0F2E0C6BEFF4588E7F814" box="[444,469,147,1973]" gridrow="3" pageId="6" pageNumber="7" textDirection="bu"> +<th id="76F16D1DFFE0F2E0C6BEF9F088E7F814" box="[444,469,1574,1973]" gridcol="0" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">This study 38 15 703</th> +<td id="76F16D1DFFE0F2E0C6BEFA5588E7F9A9" box="[444,469,1411,1544]" gridcol="1" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFA9988E7FAC4" box="[444,469,1359,1381]" gridcol="2" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFADA88E7FA88" box="[444,469,1292,1321]" gridcol="3" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFB1B88E7FB4F" box="[444,469,1229,1262]" gridcol="4" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFB4088E7FB0E" box="[444,469,1174,1199]" gridcol="5" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFBB188E7FBD6" box="[444,469,1127,1143]" gridcol="6" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFBE688E7FBE8" box="[444,469,1072,1097]" gridcol="7" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFBD488E7FBB3" box="[444,469,1026,1042]" gridcol="8" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFC1C88E7FC7F" box="[444,469,970,990]" gridcol="9" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFC5888E7FC0B" box="[444,469,910,938]" gridcol="10" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFC8088E7FCD1" box="[444,469,854,880]" gridcol="11" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFCCA88E7FC96" box="[444,469,796,823]" gridcol="12" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFD3288E7FD5F" box="[444,469,740,766]" gridcol="13" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFD7188E7FD67" box="[444,469,679,710]" gridcol="14" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFDB888E7FD28" box="[444,469,622,649]" gridcol="15" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFDE988E7FDF1" box="[444,469,575,592]" gridcol="16" gridrow="3" pageId="6" pageNumber="7" textDirection="bu">X</td> +<td id="76F16D1DFFE0F2E0C6BEFF4588E7FD80" box="[444,469,147,545]" gridcol="17" gridrow="3" pageId="6" pageNumber="7" textDirection="bu"> +Interfamilial affinities of +<taxonomicName id="4C107DA0FFE00D19C6BEFF0388E7FE9B" box="[444,469,213,314]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="order">Litopterna</taxonomicName> +</td> +</tr> +<tr id="35200461FFE0F2E0C6F2FF458BA8F814" box="[496,666,147,1973]" gridrow="4" pageId="6" pageNumber="7" textDirection="bu"> +<th id="76F16D1DFFE0F2E0C6F2FF458BA8F814" box="[496,666,147,1973]" colspan="18" colspanRight="17" gridcol="0" gridrow="4" pageId="6" pageNumber="7" textDirection="bu"> +<subScript id="17940466FFE00D19C6F2F8678B37F810" attach="right" box="[496,517,1926,1973]" fontSize="4" pageId="6" pageNumber="7">aThree</subScript> +families assumed to be monophyletic: +<taxonomicName id="4C107DA0FFE00D19C6F3FA3F8B37F9C5" box="[497,517,1513,1636]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +(probably including proterotheriids and the anisolambdids +<taxonomicName id="4C107DA0FFE00D19C6F3FC618B37FB8A" authorityName=", Cifelli" authorityYear="1983" box="[497,517,951,1067]" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C6F3FC618B37FB8A" box="[497,517,951,1067]" italics="true" pageId="6" pageNumber="7">Paranisolambda</emphasis> +</taxonomicName> +<taxonomicName id="4C107DA0FFE00D19C6F3FCB18B37FC15" authorityName="Cifelli" authorityYear="1983" box="[497,517,871,948]" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="prodromus"> +<emphasis id="B964DA31FFE00D19C6F3FCB18B37FC15" box="[497,517,871,948]" italics="true" pageId="6" pageNumber="7">prodromus</emphasis> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFE00D19C6F3FCD68B37FCFE" authorityName="Ameghino" authorityYear="1901" box="[497,517,768,863]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C6F3FCD68B37FCFE" box="[497,517,768,863]" italics="true" pageId="6" pageNumber="7">Anisolambda</emphasis> +</taxonomicName> +<emphasis id="B964DA31FFE00D19C6F3FD0C8B37FD5D" box="[497,517,730,764]" italics="true" pageId="6" pageNumber="7">amel</emphasis> +, +<taxonomicName id="4C107DA0FFE00D19C6F3FDA28B37FD73" authorityName="Ameghino" authorityYear="1901" box="[497,517,628,722]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C6F3FDA28B37FD73" box="[497,517,628,722]" italics="true" pageId="6" pageNumber="7">Anisolambda</emphasis> +</taxonomicName> +<taxonomicName id="4C107DA0FFE00D19C6F3FDE18B37FDD1" authorityName="Ameghino" authorityYear="1901" box="[497,517,567,624]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="fissidens"> +<emphasis id="B964DA31FFE00D19C6F3FDE18B37FDD1" box="[497,517,567,624]" italics="true" pageId="6" pageNumber="7">fissidens</emphasis> +</taxonomicName> +, and +<emphasis id="B964DA31FFE00D19C6F3FE798B37FE0A" box="[497,517,364,527]" italics="true" pageId="6" pageNumber="7"> +<taxonomicName id="4C107DA0FFE00D19C6F3FE798B37FDAE" authorityName="Ameghino" authorityYear="1897" box="[497,517,431,527]" class="Mammalia" family="Proterotheriidae" genus="Protheosodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<subScript id="17940466FFE00D19C6F3FE798B37FDAE" attach="right" box="[497,517,431,527]" fontSize="8" pageId="6" pageNumber="7">Protheosodon</subScript> +</taxonomicName> +<taxonomicName id="4C107DA0FFE00D19C6F3FEBA8B37FE0A" authorityName="Ameghino" authorityYear="1897" box="[497,517,364,427]" class="Mammalia" family="Proterotheriidae" genus="Protheosodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="coniferus">coniferus</taxonomicName> +</emphasis> +), +<taxonomicName id="4C107DA0FFE00D19C6F3FF0D8B37FEFD" box="[497,517,219,348]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +(probably including +<taxonomicName id="4C107DA0FFE00D19C504F9658B28F8BD" authorityName="Ameghino" authorityYear="1902" box="[518,538,1715,1820]" class="Mammalia" family="Macraucheniidae" genus="Cramauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C504F9658B28F8BD" box="[518,538,1715,1820]" italics="true" pageId="6" pageNumber="7">Cramauchenia</emphasis> +</taxonomicName> +<taxonomicName id="4C107DA0FFE00D19C504F9A48B28F911" authorityName="Ameghino" authorityYear="1902" box="[518,538,1650,1712]" class="Mammalia" family="Macraucheniidae" genus="Cramauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="normalis"> +<emphasis id="B964DA31FFE00D19C504F9A48B28F911" box="[518,538,1650,1712]" italics="true" pageId="6" pageNumber="7">normalis</emphasis> +</taxonomicName> +and later macraucheniids), and +<taxonomicName id="4C107DA0FFE00D19C504FAFF8B28FA24" box="[518,538,1321,1413]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +(probably including +<taxonomicName id="4C107DA0FFE00D19C504FBE38B28FB2D" authorityName="Ameghino" authorityYear="1897" box="[518,538,1077,1164]" class="Mammalia" family="Adianthidae" genus="Proadiantus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C504FBE38B28FB2D" box="[518,538,1077,1164]" italics="true" pageId="6" pageNumber="7">Proadiantus</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFE00D19C504FC168B28FBB0" authorityName="Ameghino" authorityYear="1897" box="[518,538,960,1041]" class="Mammalia" family="Adianthidae" genus="Tricoelodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE00D19C504FC168B28FBB0" box="[518,538,960,1041]" italics="true" pageId="6" pageNumber="7">Tricoelodus</emphasis> +</taxonomicName> +). +<subScript id="17940466FFE00D19C518F8798B02F80E" attach="right" box="[538,560,1929,1973]" fontSize="4" pageId="6" pageNumber="7">bOnly</subScript> +included characters of the lower dentition. +<subScript id="17940466FFE00D19C532F8668B68F82D" attach="both" box="[560,602,1784,1973]" fontSize="4" pageId="6" pageNumber="7">cIncluded as an outgroup. dThis refers</subScript> +to the phylogeny of +<taxonomicName id="4C107DA0FFE00D19C544F9998B68F96B" box="[582,602,1615,1738]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +included in Schmidt (2013). +<subScript id="17940466FFE00D19C558F8678B42F810" attach="both" box="[602,624,1937,1973]" fontSize="4" pageId="6" pageNumber="7">eThis</subScript> +refers to the phylogeny of +<taxonomicName id="4C107DA0FFE00D19C55EF99D8B42F96D" box="[604,624,1611,1740]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="6" pageNumber="7" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +included in Schmidt (2013). +<subScript id="17940466FFE00D19C572F8648BA8F820" attach="both" box="[624,666,1845,1973]" fontSize="4" pageId="6" pageNumber="7">fMolecular study. gStudy with</subScript> +important methodological problems, which raise questions about their conclusions. This was shown in a reply study. +</th> +</tr> +</table> +</paragraph> +<caption id="DF6F56ABFFE10D17C773F8EF8AB2F81B" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8" startId="7.[113,178,1849,1873]" targetBox="[174,1395,148,1822]" targetPageId="7" targetType="figure"> +<paragraph id="8BAF0623FFE10D18C773F8EF8CADF863" blockId="7.[113,1439,1849,1986]" pageId="7" pageNumber="8"> +<emphasis id="B964DA31FFE10D18C773F8EF89FBF8F0" bold="true" box="[113,201,1849,1873]" pageId="7" pageNumber="8">Figure 2.</emphasis> +Diversity of litoptern and associated early SANU families expressed as the number of genera. A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets. In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used. In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated +</paragraph> +<paragraph id="8BAF0623FFEE0D17C79EFE1F8A65FEC5" blockId="8.[126,779,144,1641]" lastBlockId="8.[825,1475,144,575]" pageId="8" pageNumber="9"> +Macraucheniids are distinguished from the two other families, +<taxonomicName id="4C107DA0FFEE0D17C77DFE3F89CEFDA0" box="[127,252,489,513]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFEE0D17C63EFE3F88D5FDA0" box="[316,487,489,513]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +. Among other differences, macraucheniids lack the trilobed m3, which is characteristic of adianthids ( +<bibRefCitation id="EF817BD2FFEE0D17C7FFFDF188DCFD9E" author="Cifelli and Soria" box="[253,494,551,575]" firstAuthor="Cifelli" pageId="8" pageNumber="9" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria 1983a</bibRefCitation> +). Also, macraucheniids are functional tridactyls,unlike proterotheriids that show reduced(or lost) lateral digits II and IV (Soria 2001). However, the origin of +<taxonomicName id="4C107DA0FFEE0D17C782FD538802FD3C" box="[128,304,645,669]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +in the Palaeogene and the affinities of the family within +<taxonomicName id="4C107DA0FFEE0D17C7C4FD738801FD1C" box="[198,307,677,701]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="order">Litopterna</taxonomicName> +are still contentious topics. The oldest uncontroversial members of the family are +<taxonomicName id="4C107DA0FFEE0D17C500FD12898DFD5A" authority="Ameghino 1894 a" authorityName="Ameghino" authorityYear="1894" class="Mammalia" family="Macraucheniidae" genus="Coniopternium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C500FD128BA4FD7A" box="[514,662,708,731]" italics="true" pageId="8" pageNumber="9">Coniopternium</emphasis> +<bibRefCitation id="EF817BD2FFEE0D17C59FFD12898DFD5A" author="Ameghino" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="501 - 660" refId="ref39196" refString="Ameghino F. Sur les oiseaux fossiles de Patagonie et la faune mammalogique des couches a Pyrotherium. Boletin del Instituto Geografico Argentino 1894 a; 15: 501 - 660." type="journal article" year="1894" yearSuffix="a">Ameghino 1894a</bibRefCitation> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFEE0D17C7C9FD358B22FD5A" authority="Ameghino 1902 a" authorityName="Ameghino" authorityYear="1902" box="[203,528,739,763]" class="Mammalia" family="Macraucheniidae" genus="Cramauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C7C9FD358868FD5A" box="[203,346,739,763]" italics="true" pageId="8" pageNumber="9">Cramauchenia</emphasis> +<bibRefCitation id="EF817BD2FFEE0D17C662FD358B22FD5A" author="Ameghino" box="[352,528,739,763]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="71 - 138" refId="ref39328" refString="Ameghino F. Premiere contribution a la connaissance de la faune mammalogique des couches a Colpodon. Boletin de la Academia Nacional de Ciencias en Cordoba 1902 a; 17: 71 - 138." type="journal article" year="1902" yearSuffix="a">Ameghino 1902a</bibRefCitation> +</taxonomicName> +, and +<taxonomicName id="4C107DA0FFEE0D17C54AFD328815FCBA" authority="Cifelli and Soria 1983 b" authorityName="Cifelli and Soria" authorityYear="1983" class="Mammalia" family="Macraucheniidae" genus="Pternoconius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C54AFD328BF7FD5A" box="[584,709,740,763]" italics="true" pageId="8" pageNumber="9">Pternoconius</emphasis> +<bibRefCitation id="EF817BD2FFEE0D17C5C8FD358815FCBA" author="Cifelli and Soria" firstAuthor="Cifelli" pageId="8" pageNumber="9" pagination="141 - 53" refId="ref41096" refString="Cifelli RL, Soria MF. Notes on Deseadan Macraucheniidae. Ameghiniana 1983 b; 20: 141 - 53." type="journal article" year="1983" yearSuffix="b">Cifelli and Soria 1983b</bibRefCitation> +</taxonomicName> +, all with a first appearance datum (FAD) in the Deseadean SALMA (Late Oligocene; +<bibRefCitation id="EF817BD2FFEE0D17C51BFCF48BCBFC9B" author="Dozo and Vera" box="[537,761,802,826]" firstAuthor="Dozo" pageId="8" pageNumber="9" pagination="1818 - 26" refId="ref41729" refString="Dozo MT, Vera B. First skull and associated postcranial bones of Macraucheniidae (Mammalia, Litopterna) from the Deseadan SALMA (Late Oligocene) of Cabeza Blanca (Chubut, Argentina). Journal of Vertebrate Paleontology 2010; 30: 1818 - 26. https: // doi. org / 10.1080 / 02724634.2010.521534" type="journal article" year="2010">Dozo and Vera 2010</bibRefCitation> +). These genera show dental and/or postcranial anatomical features that link them closely to later macraucheniids, like the presence of a mesolophid or cristid that connects the cristid obliqua with the entoconid [=entolophid (Soria and Hoffstetter 1985)] in all or some of the lower molars. Before the Oligocene, +<taxonomicName id="4C107DA0FFEE0D17C592FC6989C5FC57" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C592FC698A38FC76" box="[656,778,959,983]" italics="true" pageId="8" pageNumber="9">Polymorphis</emphasis> +Roth, 1899 +</taxonomicName> +from the Late Eocene of Patagonia has been proposed as the earliest member of +<taxonomicName id="4C107DA0FFEE0D17C6D3FC2B8BB3FBB4" box="[465,641,1021,1045]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +(Soria 1982, Cifelli 1983a), revalidating a previous proposal of +<bibRefCitation id="EF817BD2FFEE0D17C59FFBCB89E5FBF5" author="Ameghino" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="1 - 541" refId="ref39426" refString="Ameghino F. Recherches de morphologic phylogenetique sur les molaires superieures des ongules. Anales del Museo Nacional de Buenos Aires 1904 b; 3: 1 - 541." type="journal article" year="1904" yearSuffix="b">Ameghino (1904b)</bibRefCitation> +. Cifelli (1983a) even considered it as a member of a different subfamily, +<taxonomicName id="4C107DA0FFEE0D17C65BFB8D88CBFBD2" box="[345,505,1115,1139]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Polymorphinae">Polymorphinae</taxonomicName> +, within +<taxonomicName id="4C107DA0FFEE0D17C55BFB8D8A3BFBD2" box="[601,777,1115,1139]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +( +<tableCitation id="C6923398FFEE0D17C789FBAD89EEFB32" box="[139,220,1147,1171]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="8" pageNumber="9">Table 1</tableCitation> +). Previously, +<taxonomicName id="4C107DA0FFEE0D17C671FBAD8B23FB32" box="[371,529,1147,1171]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Polymorphinae">Polymorphinae</taxonomicName> +was also considered a distinct family of the +<taxonomicName id="4C107DA0FFEE0D17C663FB4C8A37FB13" authority="(Odreman Rivas 1969)" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[353,773,1178,1202]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae (Odreman Rivas 1969)</taxonomicName> +. Key features for associating +<taxonomicName id="4C107DA0FFEE0D17C6AFFB6F8B15FB70" authorityName="Roth" authorityYear="1899" box="[429,551,1209,1233]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C6AFFB6F8B15FB70" box="[429,551,1209,1233]" italics="true" pageId="8" pageNumber="9">Polymorphis</emphasis> +</taxonomicName> +with macraucheniids are related to similarities in the upper molars, in particular, the presence of an oblique crest that connects the protocone with the metacone (Soria 1982; +<figureCitation id="132B1AA6FFEE0D17C698FAC188D1FA8E" box="[410,483,1303,1327]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="8" pageNumber="9">Fig. 4A</figureCitation> +; see also +<taxonomicName id="4C107DA0FFEE0D17C54AFAC18BC2FA8E" box="[584,752,1303,1327]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFEE0D17C54AFAC18BF0FA8E" box="[584,706,1303,1327]" italics="true" pageId="8" pageNumber="9">Polymorphis</emphasis> +spp. +</taxonomicName> +in Supporting information, File S2). In contrast, the lower molars of +<taxonomicName id="4C107DA0FFEE0D17C799FA808876FACC" box="[155,324,1366,1390]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFEE0D17C799FA808827FACF" box="[155,277,1366,1390]" italics="true" pageId="8" pageNumber="9">Polymorphis</emphasis> +spp. +</taxonomicName> +show important differences with Deseadean SALMA macraucheniids, such as the complete absence of a mesolophid. The proposal of +<taxonomicName id="4C107DA0FFEE0D17C6B9FA438B07FA0C" authorityName="Roth" authorityYear="1899" box="[443,565,1429,1453]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C6B9FA438B07FA0C" box="[443,565,1429,1453]" italics="true" pageId="8" pageNumber="9">Polymorphis</emphasis> +</taxonomicName> +as a macraucheniid found support in a phylogenetic analysis ( +<bibRefCitation id="EF817BD2FFEE0D17C528FA628B90FA6D" author="Cifelli" box="[554,674,1460,1484]" firstAuthor="Cifelli" pageId="8" pageNumber="9" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation> +; +<figureCitation id="132B1AA6FFEE0D17C5ADFA628BCBFA6D" box="[687,761,1460,1484]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="8" pageNumber="9">Fig. 1C</figureCitation> +), and since then most recent studies have considered this taxon as the earliest member of +<taxonomicName id="4C107DA0FFEE0D17C68BFA258B0BF9AA" box="[393,569,1523,1547]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +without any further testing (e.g., +<bibRefCitation id="EF817BD2FFEE0D17C605F9C488D2F98B" author="Dozo and Vera" box="[263,480,1554,1578]" firstAuthor="Dozo" pageId="8" pageNumber="9" pagination="1818 - 26" refId="ref41729" refString="Dozo MT, Vera B. First skull and associated postcranial bones of Macraucheniidae (Mammalia, Litopterna) from the Deseadan SALMA (Late Oligocene) of Cabeza Blanca (Chubut, Argentina). Journal of Vertebrate Paleontology 2010; 30: 1818 - 26. https: // doi. org / 10.1080 / 02724634.2010.521534" type="journal article" year="2010">Dozo and Vera 2010</bibRefCitation> +, Croft +<emphasis id="B964DA31FFEE0D17C52FF9C58B6DF98B" box="[557,607,1554,1578]" italics="true" pageId="8" pageNumber="9">et al.</emphasis> +2020; +<tableCitation id="C6923398FFEE0D17C5A9F9C48BCBF98B" box="[683,761,1554,1578]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="8" pageNumber="9">Table 2</tableCitation> +). +<bibRefCitation id="EF817BD2FFEE0D17C783F9E78820F9E8" author="Cifelli" box="[129,274,1585,1609]" firstAuthor="Cifelli" pageId="8" pageNumber="9" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli (1993)</bibRefCitation> +also found +<taxonomicName id="4C107DA0FFEE0D17C692F9E78B72F9E8" box="[400,576,1585,1609]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +as the sister group of +<taxonomicName id="4C107DA0FFEE0D17C79CF98788CAF9C8" authority="(Macrauchenioidea)" baseAuthorityName="Macrauchenioidea" box="[158,504,1617,1641]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Adianthidae (Macrauchenioidea)</taxonomicName> +, and both families closely related to +<taxonomicName id="4C107DA0FFEE0D17C4A7FF468D7FFF09" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[933,1101,144,168]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +( +<taxonomicName id="4C107DA0FFEE0D17C35FFF468DCAFF09" box="[1117,1272,144,168]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subOrder" subOrder="Lopholipterna">Lopholipterna</taxonomicName> +; +<figureCitation id="132B1AA6FFEE0D17C207FF468C61FF09" box="[1285,1363,144,168]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="8" pageNumber="9">Fig. 1C</figureCitation> +; +<tableCitation id="C6923398FFEE0D17C261FF468C83FF09" box="[1379,1457,144,168]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="8" pageNumber="9">Table 1</tableCitation> +). Most additional phylogenetic studies with macraucheniids in the taxon sampling have either focused on testing relationships within the family (e.g., Püschel +<emphasis id="B964DA31FFEE0D17C383FF398D80FEA7" box="[1153,1202,238,262]" italics="true" pageId="8" pageNumber="9">et al.</emphasis> +2023) or determining the interordinal relationships within Placentalia (e.g., O’Leary +<emphasis id="B964DA31FFEE0D17C291FED88CF1FE84" box="[1427,1475,269,293]" italics="true" pageId="8" pageNumber="9">et al.</emphasis> +2013, +<bibRefCitation id="EF817BD2FFEE0D17C47AFEFB8D31FEE5" author="Buckley" box="[888,1027,300,325]" firstAuthor="Buckley" pageId="8" pageNumber="9" refId="ref40554" refString="Buckley M. Ancient collagen reveals evolutionary history of the endemic South American ' ungulates'. Proceedings of the Royal Society B: Biological Sciences 2015; 282: 20142671. https: // doi. org / 10.1098 / rspb. 2014.2671" type="journal volume" year="2015">Buckley 2015</bibRefCitation> +) instead of the interfamilial affinities ( +<tableCitation id="C6923398FFEE0D17C28BFEFB8A74FEC5" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="8" pageNumber="9">Table 2</tableCitation> +). +</paragraph> +<paragraph id="8BAF0623FFEE0D17C457FEBD8DC3FD9E" blockId="8.[825,1475,144,575]" pageId="8" pageNumber="9"> +Considering the Late Eocene +<taxonomicName id="4C107DA0FFEE0D17C39EFEBD8C24FE22" authorityName="Roth" authorityYear="1899" box="[1180,1302,363,387]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C39EFEBD8C24FE22" box="[1180,1302,363,387]" italics="true" pageId="8" pageNumber="9">Polymorphis</emphasis> +</taxonomicName> +as the first occurrence of the family and the Pliocene to Late Pleistocene +<taxonomicName id="4C107DA0FFEE0D17C43BFE7C8AF1FE63" authorityName="Owen" authorityYear="1838" box="[825,963,426,450]" class="Mammalia" family="Macraucheniidae" genus="Macrauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C43BFE7C8AF1FE63" box="[825,963,426,450]" italics="true" pageId="8" pageNumber="9">Macrauchenia</emphasis> +</taxonomicName> +( +<bibRefCitation id="EF817BD2FFEE0D17C4D7FE7C8D7AFE63" author="Bond" box="[981,1096,426,450]" firstAuthor="Bond" pageId="8" pageNumber="9" pagination="177 - 205" refId="ref40196" refString="Bond M. Quaternary native ungulates of southern South America. A synthesis. In: Rabassa J, Salemme M, (eds), Quaternary of South America and Antarctic Peninsula. Vol. 12. Rotterdam: CRC Press, 1999, 177 - 205." type="book chapter" year="1999">Bond 1999</bibRefCitation> +, Prado +<emphasis id="B964DA31FFEE0D17C39AFE7D8DFAFE63" box="[1176,1224,426,450]" italics="true" pageId="8" pageNumber="9">et al.</emphasis> +2015) and the probably Late Pleistocene +<taxonomicName id="4C107DA0FFEE0D17C4E5FE1F8C9AFE40" authority="(Cartelle and Lessa 1988)" baseAuthorityName="Cartelle and Lessa" baseAuthorityYear="1988" box="[999,1448,457,481]" class="Mammalia" family="Macraucheniidae" genus="Xenorhinotherium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C4E5FE1F8DABFE40" box="[999,1177,457,481]" italics="true" pageId="8" pageNumber="9">Xenorhinotherium</emphasis> +( +<bibRefCitation id="EF817BD2FFEE0D17C3AAFE1F8CAFFE40" author="Cartelle and Lessa" box="[1192,1437,457,481]" firstAuthor="Cartelle" pageId="8" pageNumber="9" pagination="3 - 26" refId="ref40717" refString="Cartelle C, Lessa G. Descricao de um genero e especie de Macraucheniidae (Mammalia, Litopterna) do Pleistoceno final do Brasil. Paula- Coutiana 1988; 3: 3 - 26." type="journal article" year="1988">Cartelle and Lessa 1988</bibRefCitation> +) +</taxonomicName> +as the last occurrences of +<taxonomicName id="4C107DA0FFEE0D17C329FE3F8DEFFDA0" box="[1067,1245,489,513]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +, the temporal interval for this family would be around 39.00–~0.11 Mya ( +<figureCitation id="132B1AA6FFEE0D17C26FFDDE8C88FD81" box="[1389,1466,520,544]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="8" pageNumber="9">Fig. 2B</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFEE0D17C33AFDF18DA2FD9E" box="[1080,1168,551,575]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="8" pageNumber="9">Table S1</tableCitation> +, File S2). +</paragraph> +<subSubSection id="C30A55A8FFEE0D17C4D7FD898C14FDD8" box="[981,1318,607,633]" pageId="8" pageNumber="9" type="nomenclature"> +<paragraph id="8BAF0623FFEE0D17C4D7FD898C14FDD8" blockId="8.[822,1475,607,1641]" box="[981,1318,607,633]" pageId="8" pageNumber="9"> +<emphasis id="B964DA31FFEE0D17C4D7FD898C14FDD8" box="[981,1318,607,633]" italics="true" pageId="8" pageNumber="9"> +<taxonomicName id="4C107DA0FFEE0D17C4D7FD898D5EFDD8" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[981,1132,607,633]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +( +<figureCitation id="132B1AA6FFEE0D17C379FDB68DEBFDD8" box="[1147,1241,607,633]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="8" pageNumber="9">Figs 4J–K</figureCitation> +, +<figureCitation id="132B1AA6FFEE0D17C3E6FD898C2EFDD8" box="[1252,1308,607,633]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="8" pageNumber="9">5H–I</figureCitation> +) +</emphasis> +</paragraph> +</subSubSection> +<subSubSection id="C30A55A8FFEE0D15C43BFD5388D5FB50" lastPageId="10" lastPageNumber="11" pageId="8" pageNumber="9" type="multiple"> +<paragraph id="8BAF0623FFEE0D17C43BFD538C0EFA0D" blockId="8.[822,1475,607,1641]" pageId="8" pageNumber="9"> +With 28 currently accepted genera, +<taxonomicName id="4C107DA0FFEE0D17C3A6FD538C7EFD3C" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1188,1356,645,669]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +is the most diverse family of litopterns ( +<figureCitation id="132B1AA6FFEE0D17C365FD738D81FD1C" box="[1127,1203,677,701]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="8" pageNumber="9">Fig. 2A</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFEE0D17C43BFD128AA6FD7D" box="[825,916,708,732]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="8" pageNumber="9">Table S1</tableCitation> +), being particularly diverse during the Neogene alongside macraucheniids ( +<figureCitation id="132B1AA6FFEE0D17C327FD358DA6FD5A" box="[1061,1172,739,763]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="8" pageNumber="9">Fig. 2B–C</figureCitation> +). Soria (2001) presented a major revision of this family, so to avoid unnecessary repetition, only the most important taxonomical aspects and the main advances since his revision are discussed here. +<taxonomicName id="4C107DA0FFEE0D17C3F2FC978CAAFCF8" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1264,1432,833,857]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +was named by +<bibRefCitation id="EF817BD2FFEE0D17C4A7FCB78D53FCD8" author="Ameghino" box="[933,1121,865,889]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="1 - 26" refId="ref39043" refString="Ameghino F. Enumeracion sistematica de las especies de mamiferos fosiles coleccionados por Carlos Ameghino en los terrenos Eocenos de la Patagonia Austral y depositados en el Museo de La Plata. Boletin del Museo de La Plata 1887; 1: 1 - 26." type="journal article" year="1887">Ameghino (1887)</bibRefCitation> +with the description of the genera +<taxonomicName id="4C107DA0FFEE0D17C438FC568D4DFC39" authority="Ameghino, 1883" authorityName="Ameghino" authorityYear="1883" box="[826,1151,896,920]" class="Mammalia" family="Proterotheriidae" genus="Proterotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C438FC568AFCFC39" box="[826,974,896,920]" italics="true" pageId="8" pageNumber="9">Proterotherium</emphasis> +<bibRefCitation id="EF817BD2FFEE0D17C4D6FC568D4DFC39" author="Ameghino" box="[980,1151,896,920]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="257 - 306" refId="ref39005" refString="Ameghino F. Sobre una nueva coleccion de mamiferos fosiles recogidos por el profesor Scalabrini en las barrancas del Parana. Boletin de la Academia Nacional de Ciencias en Cordoba 1883; 5: 257 - 306." type="journal article" year="1883">Ameghino, 1883</bibRefCitation> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFEE0D17C38EFC568C8EFC39" authority="Ameghino, 1887" authorityName="Ameghino" authorityYear="1887" box="[1164,1468,896,920]" class="Mammalia" family="Proterotheriidae" genus="Thoatherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C38EFC568C39FC39" box="[1164,1291,896,920]" italics="true" pageId="8" pageNumber="9">Thoatherium</emphasis> +<bibRefCitation id="EF817BD2FFEE0D17C213FC568C8EFC39" author="Ameghino" box="[1297,1468,896,920]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="1 - 26" refId="ref39043" refString="Ameghino F. Enumeracion sistematica de las especies de mamiferos fosiles coleccionados por Carlos Ameghino en los terrenos Eocenos de la Patagonia Austral y depositados en el Museo de La Plata. Boletin del Museo de La Plata 1887; 1: 1 - 26." type="journal article" year="1887">Ameghino, 1887</bibRefCitation> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFEE0D17C43BFC498D59FC16" authority="Ameghino, 1887" authorityName="Ameghino" authorityYear="1887" box="[825,1131,927,951]" class="Mammalia" family="Proterotheriidae" genus="Diadiaphorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C43BFC498AF2FC16" box="[825,960,927,951]" italics="true" pageId="8" pageNumber="9">Diadiaphorus</emphasis> +<bibRefCitation id="EF817BD2FFEE0D17C4C1FC498D59FC16" author="Ameghino" box="[963,1131,927,951]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="1 - 26" refId="ref39043" refString="Ameghino F. Enumeracion sistematica de las especies de mamiferos fosiles coleccionados por Carlos Ameghino en los terrenos Eocenos de la Patagonia Austral y depositados en el Museo de La Plata. Boletin del Museo de La Plata 1887; 1: 1 - 26." type="journal article" year="1887">Ameghino, 1887</bibRefCitation> +</taxonomicName> +, and +<taxonomicName id="4C107DA0FFEE0D17C39FFC498C8EFC16" authority="Ameghino, 1887" authorityName="Ameghino" authorityYear="1887" box="[1181,1468,927,951]" class="Mammalia" family="Proterotheriidae" genus="Licaphrium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C39FFC498C3DFC16" box="[1181,1295,927,951]" italics="true" pageId="8" pageNumber="9">Licaphrium</emphasis> +<bibRefCitation id="EF817BD2FFEE0D17C216FC498C8EFC16" author="Ameghino" box="[1300,1468,927,951]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="1 - 26" refId="ref39043" refString="Ameghino F. Enumeracion sistematica de las especies de mamiferos fosiles coleccionados por Carlos Ameghino en los terrenos Eocenos de la Patagonia Austral y depositados en el Museo de La Plata. Boletin del Museo de La Plata 1887; 1: 1 - 26." type="journal article" year="1887">Ameghino, 1887</bibRefCitation> +</taxonomicName> +. Simpson (1945) subdivided this family into two subfamilies, +<taxonomicName id="4C107DA0FFEE0D17C43BFC088AE5FC57" box="[825,983,990,1014]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Polymorphinae">Polymorphinae</taxonomicName> +and +<taxonomicName id="4C107DA0FFEE0D17C308FC088D86FC57" box="[1034,1204,990,1014]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Proterotheriinae">Proterotheriinae</taxonomicName> +, the former including taxa mostly from the Palaeocene and Eocene (Peligran to Mustersan SALMAs) and the later the taxa from the Late Oligocene onwards (Deseadean SALMA). Although this classification was followed by other authors ( +<bibRefCitation id="EF817BD2FFEE0D17C358FB8A8DD5FBD2" author="Lavocat" box="[1114,1255,1115,1139]" firstAuthor="Lavocat" pageId="8" pageNumber="9" pagination="31 - 59" refId="ref43823" refString="Lavocat R. Litopterna. Traite de Paleontologie 1958; 6: 31 - 59." type="journal article" year="1958">Lavocat 1958</bibRefCitation> +, Paula Couto 1979), Simpson (1948) did not employ this subfamilial division and only used the family +<taxonomicName id="4C107DA0FFEE0D17C304FB4C8D82FB13" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1030,1200,1178,1202]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +, without giving any explanation for this omission. Later, Odreman-Rivas (1969) modified the subfamily +<taxonomicName id="4C107DA0FFEE0D17C4C9FB0F8D5BFB50" box="[971,1129,1241,1265]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Polymorphinae">Polymorphinae</taxonomicName> +from Simpson’s (1945) proposal, keeping +<taxonomicName id="4C107DA0FFEE0D17C493FB2E8DBAFAB1" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" box="[913,1160,1272,1296]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C493FB2E8D39FAB1" box="[913,1035,1272,1296]" italics="true" pageId="8" pageNumber="9">Polymorphis</emphasis> +Roth, 1899 +</taxonomicName> +(including +<taxonomicName id="4C107DA0FFEE0D17C201FB2E8A99FA8E" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" class="Mammalia" family="Proterotheriidae" genus="Polyacrodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="ligatus"> +<emphasis id="B964DA31FFEE0D17C201FB2E8CF0FAB1" box="[1283,1474,1272,1296]" italics="true" pageId="8" pageNumber="9">Polyacrodon ligatus</emphasis> +Roth, 1899 +</taxonomicName> +and +<taxonomicName id="4C107DA0FFEE0D17C4D5FAC18DE1FA8E" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" box="[983,1235,1303,1327]" class="Mammalia" family="Proterotheriidae" genus="Heteroglyphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C4D5FAC18D6EFA8E" box="[983,1116,1303,1327]" italics="true" pageId="8" pageNumber="9">Heteroglyphis</emphasis> +Roth, 1899 +</taxonomicName> +), and excluding the rest of the taxa such as the now considered anisolambdids +<taxonomicName id="4C107DA0FFEE0D17C274FAE18AE5FACF" authority="Simpson, 1935" authorityName="Simpson" authorityYear="1935" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C274FAE18CF0FAEE" box="[1398,1474,1335,1359]" italics="true" pageId="8" pageNumber="9">Wainka</emphasis> +Simpson, 1935 +</taxonomicName> +and +<taxonomicName id="4C107DA0FFEE0D17C310FA808C7FFACF" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[1042,1357,1366,1390]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C310FA808DA1FACF" box="[1042,1171,1366,1390]" italics="true" pageId="8" pageNumber="9">Anisolambda</emphasis> +<bibRefCitation id="EF817BD2FFEE0D17C39CFA808C7FFACF" author="Ameghino" box="[1182,1357,1366,1390]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +(including +<taxonomicName id="4C107DA0FFEE0D17C43BFAA38DAAFA2C" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[825,1176,1397,1421]" class="Mammalia" family="Proterotheriidae" genus="Ricardolydekkeria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C43BFAA38ADBFA2C" box="[825,1001,1397,1421]" italics="true" pageId="8" pageNumber="9">Ricardolydekkeria</emphasis> +<bibRefCitation id="EF817BD2FFEE0D17C4ECFAA38DAAFA2C" author="Ameghino" box="[1006,1176,1397,1421]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFEE0D17C3C8FAA38A5FFA0C" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" class="Mammalia" family="Proterotheriidae" genus="Josepholeidya" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C3C8FAA38C7EFA2C" box="[1226,1356,1397,1421]" italics="true" pageId="8" pageNumber="9">Josepholeidya</emphasis> +<bibRefCitation id="EF817BD2FFEE0D17C250FAA38A5FFA0C" author="Ameghino" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +) and the didolodontid +<taxonomicName id="4C107DA0FFEE0D17C35EFA438C04FA0C" authority="Berg, 1899" authorityName="Berg" authorityYear="1899" box="[1116,1334,1429,1453]" class="Mammalia" family="Proterotheriidae" genus="Xesmodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEE0D17C35EFA438DF3FA0C" box="[1116,1217,1429,1453]" italics="true" pageId="8" pageNumber="9">Xesmodon</emphasis> +<bibRefCitation id="EF817BD2FFEE0D17C3C5FA438C04FA0C" author="Berg" box="[1223,1334,1429,1453]" firstAuthor="Berg" pageId="8" pageNumber="9" pagination="77 - 80" refId="ref39765" refString="Berg C. Substitucion de nombres genericos. III. Comunicaciones del Museo Nacional de Buenos Aires 1899; 1: 77 - 80." type="journal article" year="1899">Berg, 1899</bibRefCitation> +</taxonomicName> +. +</paragraph> +<paragraph id="8BAF0623FFEE0D15C457FA62886AFD3C" blockId="8.[822,1475,607,1641]" lastBlockId="10.[126,779,144,1265]" lastPageId="10" lastPageNumber="11" pageId="8" pageNumber="9"> +Cifelli(1983a)subdividedthefamilyProterotheriidaeintotwo subfamilies, +<taxonomicName id="4C107DA0FFEE0D17C4BAFA058D6DFA4A" box="[952,1119,1491,1515]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Anisolambdinae">Anisolambdinae</taxonomicName> +and +<taxonomicName id="4C107DA0FFEE0D17C39AFA058C70FA4A" box="[1176,1346,1491,1515]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Proterotheriinae">Proterotheriinae</taxonomicName> +, the former subfamily including mostly anisolambdids (see +<taxonomicName id="4C107DA0FFEE0D17C219FA258CF0F9AA" authorityName=", Soria" authorityYear="2001" box="[1307,1474,1523,1547]" family="Anisolambdidae" pageId="8" pageNumber="9" rank="family">Anisolambdidae</taxonomicName> +section for more details), and the latter similar to the original definition of Simpson (1945). In addition, Cifelli (1983a) excluded the subfamily +<taxonomicName id="4C107DA0FFEE0D17C4CFF9878D59F9C8" box="[973,1131,1617,1641]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Polymorphinae">Polymorphinae</taxonomicName> +from +<taxonomicName id="4C107DA0FFEE0D17C3AEF9878C64F9C8" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1196,1366,1617,1641]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +, placing it insteadwithinthefamilyMacraucheniidae (seeMacraucheniidae section for more details). Soria (2001) later removed the subfamily +<taxonomicName id="4C107DA0FFEC0D15C7C4FF19885FFF46" box="[198,365,207,231]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="subFamily" subFamily="Anisolambdinae">Anisolambdinae</taxonomicName> +from +<taxonomicName id="4C107DA0FFEC0D15C6A8FF198B66FF46" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[426,596,207,231]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +, elevating its rank as a distinct family (see +<taxonomicName id="4C107DA0FFEC0D15C680FF388B1BFEA7" authorityName=", Soria" authorityYear="2001" box="[386,553,238,262]" family="Anisolambdidae" pageId="10" pageNumber="11" rank="family">Anisolambdidae</taxonomicName> +section for more details), which leaves the concept of +<taxonomicName id="4C107DA0FFEC0D15C6E0FEDB8BB8FE84" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[482,650,269,293]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +very similar to the subfamily +<taxonomicName id="4C107DA0FFEC0D15C63DFEFB88D5FEE4" box="[319,487,301,325]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="subFamily" subFamily="Proterotheriinae">Proterotheriinae</taxonomicName> +from Simpson (1945), although with numerous new additions to the family since then (e.g., +<bibRefCitation id="EF817BD2FFEC0D15C7B4FEBD885CFE22" author="Corona" box="[182,366,363,387]" etAl="et al." firstAuthor="Corona" pageId="10" pageNumber="11" refId="ref41392" refString="Corona A, Badin AC, Perea D et al. A new genus and species and additional reports of the South American native ungulates Proterotheriidae (Mammalia, Litopterna) in the Late Miocene of Uruguay. Journal of South American Earth Sciences 2020; 102: 102646. https: // doi. org / 10.1016 / j. jsames. 2020.102646" type="journal volume" year="2020"> +Corona +<emphasis id="B964DA31FFEC0D15C60BFEBA8805FE22" box="[265,311,363,387]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2020 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFEC0D15C67AFEBD8B7EFE22" author="McGrath" box="[376,588,363,387]" etAl="et al." firstAuthor="McGrath" pageId="10" pageNumber="11" pagination="159 - 88" refId="ref44350" refString="McGrath AJ, Anaya F, Croft DA. New proterotheriids from the Middle Miocene of Quebrada Honda, Bolivia, and body size and diversity trends in proterotheriid and macraucheniid litopterns (Mammalia). Ameghiniana 2020 a; 57: 159 - 88. https: // doi. org / 10.5710 / amgh. 03.03.2020.3268" type="journal article" year="2020" yearSuffix="a"> +McGrath +<emphasis id="B964DA31FFEC0D15C6D9FEBA8B3BFE22" box="[475,521,363,387]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2020a +</bibRefCitation> +; Supporting information, +<tableCitation id="C6923398FFEC0D15C7D6FE5D881FFE02" box="[212,301,395,419]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="10" pageNumber="11">Table S1</tableCitation> +). Important additions to +<taxonomicName id="4C107DA0FFEC0D15C52EFE5D8BE6FE02" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[556,724,395,419]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +were the bunodont +<taxonomicName id="4C107DA0FFEC0D15C61FFE7C8BEBFE63" authority="McKenna, 1956" authorityName="McKenna" authorityYear="1956" box="[285,729,426,450]" class="Mammalia" family="Didolodontidae" genus="Megadolodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="molariformis"> +<emphasis id="B964DA31FFEC0D15C61FFE7C8B17FE63" box="[285,549,426,450]" italics="true" pageId="10" pageNumber="11">Megadolodus molariformis</emphasis> +<bibRefCitation id="EF817BD2FFEC0D15C52DFE7C8BEBFE63" author="McKenna" box="[559,729,426,450]" firstAuthor="McKenna" pageId="10" pageNumber="11" pagination="736 - 43" refId="ref44469" refString="McKenna MC. Survival of primitive notoungulates and condylarths into the Miocene of Colombia. American Journal of Science 1956; 254: 736 - 43. https: // doi. org / 10.2475 / ajs. 254.12.736" type="journal article" year="1956">McKenna, 1956</bibRefCitation> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFEC0D15C783FE1F8BBBFE40" authority="Hoffstetter & Soria, 1986" authorityName="Hoffstetter & Soria" authorityYear="1986" box="[129,649,457,481]" class="Mammalia" family="Didolodontidae" genus="Neodolodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="colombianus"> +<emphasis id="B964DA31FFEC0D15C783FE1F8847FE40" box="[129,373,457,481]" italics="true" pageId="10" pageNumber="11">Neodolodus colombianus</emphasis> +<bibRefCitation id="EF817BD2FFEC0D15C67FFE1F8BBBFE40" author="Hoffstetter & Soria" box="[381,649,457,481]" firstAuthor="Hoffstetter" pageId="10" pageNumber="11" pagination="1619 - 22" refId="ref43229" refString="Hoffstetter R, Soria MF. Neolodus colombianus gen. et sp. nov., un noveau Condylarthre (Mammalia) dans le Miocene de Colombie. Comptes Rendus de la Academie de Sciences, Paris 1986; 303: 1619 - 22." type="journal article" year="1986">Hoffstetter & Soria, 1986</bibRefCitation> +</taxonomicName> +( +<bibRefCitation id="EF817BD2FFEC0D15C59EFE1F89DEFDA0" author="Cifelli and Diaz" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="222 - 31" refId="ref41021" refString="Cifelli RL, Diaz JG. New remains of Prothoatherium columbianus (Litopterna, Mammalia) from the Miocene of Colombia. Journal of Vertebrate Paleontology 1989; 9: 222 - 31. https: // doi. org / 10.1080 / 02 724634.1989. 10011756" type="journal article" year="1989">Cifelli and Diaz 1989</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFEC0D15C7F8FE3F8B39FDA0" author="Cifelli and Villarroel" box="[250,523,489,513]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="265 - 88" refId="ref41117" refString="Cifelli RL, Villarroel C. Paleobiology and affinities of Megadolodus. In: Kay RF, Madden RH, Cifelli RL, Flynn JJ (eds), Vertebrate Paleontology in the Neotropics: The Miocene Fauna of La Venta, Colombia. Washington and London: Smithsonian Institution Press, 1997, 265 - 88." type="book chapter" year="1997">Cifelli and Villarroel 1997</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFEC0D15C51BFE3F8BCAFDA0" author="McGrath" box="[537,760,489,513]" etAl="et al." firstAuthor="McGrath" pageId="10" pageNumber="11" pagination="717 - 38" refId="ref44408" refString="McGrath AJ, Flynn JJ, Wyss AR. Proterotheriids and macraucheniids (Litopterna: Mammalia) from the Pampa Castillo Fauna, Chile (early Miocene, Santacrucian SALMA) and a new phylogeny of Proterotheriidae. Journal of Systematic Palaeontology 2020 b; 18: 717 - 38. https: // doi. org / 10.1080 / 14772019.2019.1662500" type="journal article" year="2020" yearSuffix="b"> +McGrath +<emphasis id="B964DA31FFEC0D15C582FE3F8B80FDA0" box="[640,690,489,513]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2020b +</bibRefCitation> +), which were previously considered didolodontids ( +<bibRefCitation id="EF817BD2FFEC0D15C5A6FDDE8987FD9E" author="McKenna" firstAuthor="McKenna" pageId="10" pageNumber="11" pagination="736 - 43" refId="ref44469" refString="McKenna MC. Survival of primitive notoungulates and condylarths into the Miocene of Colombia. American Journal of Science 1956; 254: 736 - 43. https: // doi. org / 10.2475 / ajs. 254.12.736" type="journal article" year="1956">McKenna 1956</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFEC0D15C7C0FDF188EBFD9E" author="Hoffstetter and Soria" box="[194,473,551,575]" firstAuthor="Hoffstetter" pageId="10" pageNumber="11" pagination="1619 - 22" refId="ref43229" refString="Hoffstetter R, Soria MF. Neolodus colombianus gen. et sp. nov., un noveau Condylarthre (Mammalia) dans le Miocene de Colombie. Comptes Rendus de la Academie de Sciences, Paris 1986; 303: 1619 - 22." type="journal article" year="1986">Hoffstetter and Soria 1986</bibRefCitation> +). Including both taxa within +<taxonomicName id="4C107DA0FFEC0D15C783FD9189DCFDFE" box="[129,238,583,607]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="order">Litopterna</taxonomicName> +has phylogenetic support, but their affinities within +<taxonomicName id="4C107DA0FFEC0D15C781FDB08819FDDF" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[131,299,614,638]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +and even other litoptern families are unclear (Carrillo +<emphasis id="B964DA31FFEC0D15C7E2FD50883DFD3C" box="[224,271,645,669]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2023). +</paragraph> +<paragraph id="8BAF0623FFEE0D17C783F9098AB2F81B" blockId="8.[129,1459,1759,1979]" pageId="8" pageNumber="9"> +in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, +<tableCitation id="C6923398FFEE0D17C3E2F92D8C01F8B2" box="[1248,1331,1787,1811]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="8" pageNumber="9">Table S1</tableCitation> +, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot. Note that in the Discussion section, we proposed to redefine the families +<taxonomicName id="4C107DA0FFEE0D17C4F4F8E58DA3F8EA" authorityName=", Soria" authorityYear="2001" box="[1014,1169,1843,1867]" family="Anisolambdidae" pageId="8" pageNumber="9" rank="family">Anisolambdidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFEE0D17C3BCF8E58CA2F8EA" box="[1214,1424,1843,1867]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +as the subfamilies +<taxonomicName id="4C107DA0FFEE0D17C616F899889DF8C6" box="[276,431,1871,1895]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Anisolambdinae">Anisolambdinae</taxonomicName> +and +<taxonomicName id="4C107DA0FFEE0D17C6DEF8998B9CF8C6" box="[476,686,1871,1895]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Sparnotheriodontinae">Sparnotheriodontinae</taxonomicName> +of the family +<taxonomicName id="4C107DA0FFEE0D17C42FF8998AF8F8C6" authorityName=", Soria" authorityYear="2001" box="[813,970,1871,1895]" family="Anisolambdidae" pageId="8" pageNumber="9" rank="family">Anisolambdidae</taxonomicName> +. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, +<taxonomicName id="4C107DA0FFEE0D17C6D5F8BD8B59F823" authority="Zone" authorityName="Zone" box="[471,619,1899,1923]" class="Mammalia" family="Carodniidae" genus="Carodnia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Xenungulata" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">Carodnia Zone</taxonomicName> +; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan. +</paragraph> +<caption id="DF6F56ABFFEF0D15C773F93B8D1DF81B" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10" startId="9.[113,178,1773,1797]" targetBox="[220,1352,145,1745]" targetPageId="9" targetType="figure"> +<paragraph id="8BAF0623FFEF0D16C773F93B8C5AF80C" blockId="9.[113,1431,1773,1966]" pageId="9" pageNumber="10"> +<emphasis id="B964DA31FFEF0D16C773F93B89FBF8A4" bold="true" box="[113,201,1773,1797]" pageId="9" pageNumber="10">Figure 3.</emphasis> +Dental nomenclature used in the text and the characters. A, right upper molar in occlusal view. B–D, right lower molars in occlusal view. The main cusps of the molars are highlighted in bold. The molars represent structures sometimes present in SANUs and with relevance in the character scoring of this study. The molars do not represent any particular SANU. Molars in A and B were partially based on drawings from +<bibRefCitation id="EF817BD2FFEF0D16C7A6F8978811F8F8" author="Gelfo" box="[164,291,1857,1881]" firstAuthor="Gelfo" pageId="9" pageNumber="10" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo (2006)</bibRefCitation> +. Abbreviations upper molar: ecg, buccal cingulum or ectocingulum; es, entostyle; hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mt, metastyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; ppc, preparacrista (or paracrista); pplc, preparaconular crista; pprc, preprotocrista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; pshyc, posthypocrista; psmc, postmetacrista (or metacrista); psmlc, postmetaconular +</paragraph> +<paragraph id="8BAF0623FFEC0D15C79EFD7389ACFC57" blockId="10.[126,779,144,1265]" pageId="10" pageNumber="11"> +<taxonomicName id="4C107DA0FFEC0D15C79EFD738876FD1C" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[156,324,677,701]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +has been included by some authors as part of suborder +<taxonomicName id="4C107DA0FFEC0D15C640FD1288E4FD7D" box="[322,470,708,732]" pageId="10" pageNumber="11" rank="subOrder" subOrder="Lopholipterna">Lopholipterna</taxonomicName> +(Cifelli 1983a, Soria 2001; +<tableCitation id="C6923398FFEC0D15C783FD3589E0FD5A" box="[129,210,739,763]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="10" pageNumber="11">Table 1</tableCitation> +), which also includes the families +<taxonomicName id="4C107DA0FFEC0D15C55BFD358BE4FD5A" box="[601,726,739,763]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFEC0D15C783FCD58801FCBA" box="[129,307,771,795]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +, a hypothesis that later found phylogenetic support ( +<bibRefCitation id="EF817BD2FFEC0D15C7E1FCF4886FFC9B" author="Cifelli" box="[227,349,802,826]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation> +; +<figureCitation id="132B1AA6FFEC0D15C66EFCF4888AFC9B" box="[364,440,802,826]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="10" pageNumber="11">Fig. 1C</figureCitation> +). Since then, most phylogenetic studies that included proterotheriids have focused either on resolving the affinities within +<taxonomicName id="4C107DA0FFEC0D15C694FCB78B0CFCD8" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[406,574,865,889]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +(e.g., +<bibRefCitation id="EF817BD2FFEC0D15C57BFCB789F2FC39" author="McGrath" etAl="et al." firstAuthor="McGrath" pageId="10" pageNumber="11" pagination="717 - 38" refId="ref44408" refString="McGrath AJ, Flynn JJ, Wyss AR. Proterotheriids and macraucheniids (Litopterna: Mammalia) from the Pampa Castillo Fauna, Chile (early Miocene, Santacrucian SALMA) and a new phylogeny of Proterotheriidae. Journal of Systematic Palaeontology 2020 b; 18: 717 - 38. https: // doi. org / 10.1080 / 14772019.2019.1662500" type="journal article" year="2020" yearSuffix="b"> +McGrath +<emphasis id="B964DA31FFEC0D15C5DEFCB78A38FCD8" box="[732,778,865,889]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2020b +</bibRefCitation> +) or phylogenies studying interordinal relationships between SANUs (e.g., +<bibRefCitation id="EF817BD2FFEC0D15C64EFC498B2AFC16" author="MacPhee" box="[332,536,927,951]" etAl="et al." firstAuthor="MacPhee" pageId="10" pageNumber="11" pagination="1 - 183" refId="ref44179" refString="MacPhee RDE, Hernandez del Pino S, Kramarz A et al. Cranial morphology and phylogenetic relationships of Trigonostylops wortmani, an Eocene South American Native Ungulate. Bulletin of the American Museum of Natural History 2021; 449: 1 - 183." type="journal article" year="2021"> +MacPhee +<emphasis id="B964DA31FFEC0D15C6B0FC7688D2FC16" box="[434,480,927,951]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2021 +</bibRefCitation> +) instead of phylogenies examining the interfamilial affinities of proterotheriids ( +<tableCitation id="C6923398FFEC0D15C5D3FC6989BCFC57" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="10" pageNumber="11">Table 2</tableCitation> +). +</paragraph> +<paragraph id="8BAF0623FFEC0D15C79EFC2B88D5FB50" blockId="10.[126,779,144,1265]" pageId="10" pageNumber="11"> +Following Soria’s (2001) concept of the family, the earliest proterotheriid is +<taxonomicName id="4C107DA0FFEC0D15C639FBCB8BF9FB94" authority="Ameghino, 1897" authorityName="Ameghino" authorityYear="1897" box="[315,715,1053,1077]" class="Mammalia" family="Proterotheriidae" genus="Lambdaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="suinus"> +<emphasis id="B964DA31FFEC0D15C639FBCB8B3FFB95" box="[315,525,1053,1077]" italics="true" pageId="10" pageNumber="11">Lambdaconus suinus</emphasis> +<bibRefCitation id="EF817BD2FFEC0D15C51BFBCB8BF9FB94" author="Ameghino" box="[537,715,1053,1077]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino, 1897</bibRefCitation> +</taxonomicName> +from different localities from the Sarmiento Formation, +<collectingRegion id="49D4C8C1FFEC0D15C5B6FBEA8A34FBF5" box="[692,774,1084,1108]" country="Argentina" name="Chubut" pageId="10" pageNumber="11">Chubut</collectingRegion> +, +<collectingCountry id="F30746B3FFEC0D15C783FB8D89DBFBD2" box="[129,233,1115,1139]" name="Argentina" pageId="10" pageNumber="11">Argentina</collectingCountry> +, and the last is +<taxonomicName id="4C107DA0FFEC0D15C687FB8D8A3BFBD2" authority="Frenguelli, 1921" authorityName="Frenguelli" authorityYear="1921" box="[389,777,1115,1139]" class="Mammalia" family="Proterotheriidae" genus="Neolicaphrium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="recents"> +<emphasis id="B964DA31FFEC0D15C687FB8D8B6FFBD2" box="[389,605,1115,1139]" italics="true" pageId="10" pageNumber="11">Neolicaphrium recents</emphasis> +<bibRefCitation id="EF817BD2FFEC0D15C560FB8D8A3BFBD2" author="Frenguelli" box="[610,777,1115,1139]" firstAuthor="Frenguelli" pageId="10" pageNumber="11" pagination="7 - 23" refId="ref41980" refString="Frenguelli J. Sobre un proteroterido del Pampeano superior de Cordoba. Actas de la Academia Nacional de Ciencias de la Republica Argentina en Cordoba 1921; 7: 7 - 23." type="journal article" year="1921">Frenguelli, 1921</bibRefCitation> +</taxonomicName> +from the Late Pleistocene from different localities in +<collectingCountry id="F30746B3FFEC0D15C59CFBAD8A34FB32" box="[670,774,1147,1171]" name="Argentina" pageId="10" pageNumber="11">Argentina</collectingCountry> +, +<collectingCountry id="F30746B3FFEC0D15C783FB4C898CFB13" box="[129,190,1178,1202]" name="Brazil" pageId="10" pageNumber="11">Brazil</collectingCountry> +, and +<collectingCountry id="F30746B3FFEC0D15C7FAFB4C8863FB13" box="[248,337,1178,1202]" name="Uruguay" pageId="10" pageNumber="11">Uruguay</collectingCountry> +( +<bibRefCitation id="EF817BD2FFEC0D15C660FB4C8B08FB13" author="Gaudioso" box="[354,570,1178,1202]" etAl="et al." firstAuthor="Gaudioso" pageId="10" pageNumber="11" pagination="23 - 9" refId="ref42056" refString="Gaudioso PJ, Gasparini GM, Herbst R et al. First record of the Neolicaphrium recens Frenguelli, 1921 (Mammalia, Litopterna) in the Pleistocene of Santiago del Estero Province, Argentina. Papeis Avulsos de Zoologia 2017; 57: 23 - 9. https: // doi. org / 10.11606 / 0031 - 1049.2017. 57.03" type="journal article" year="2017"> +Gaudioso +<emphasis id="B964DA31FFEC0D15C6CCFB4D88CDFB13" box="[462,511,1178,1202]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2017 +</bibRefCitation> +), giving a temporal interval of around 29.3–~0.11 Mya to this family ( +<figureCitation id="132B1AA6FFEC0D15C5B1FB6F8A33FB70" box="[691,769,1209,1233]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="10" pageNumber="11">Fig. 2B</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFEC0D15C67DFB0F88EBFB50" box="[383,473,1241,1265]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="10" pageNumber="11">Table S1</tableCitation> +). +</paragraph> +<subSubSection id="C30A55A8FFEC0D15C62AFAC68B50FA8A" box="[296,610,1296,1323]" pageId="10" pageNumber="11" type="nomenclature"> +<paragraph id="8BAF0623FFEC0D15C62AFAC68B50FA8A" blockId="10.[129,779,1296,1735]" box="[296,610,1296,1323]" pageId="10" pageNumber="11"> +<emphasis id="B964DA31FFEC0D15C62AFAC68B50FA8A" box="[296,610,1296,1323]" italics="true" pageId="10" pageNumber="11"> +<taxonomicName id="4C107DA0FFEC0D15C62AFAC688AEFA8B" box="[296,412,1296,1322]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +( +<figureCitation id="132B1AA6FFEC0D15C6A9FAC78B20FA8B" box="[427,530,1296,1322]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="10" pageNumber="11">Figs 4F–H</figureCitation> +, +<figureCitation id="132B1AA6FFEC0D15C519FAC78B6AFA8A" box="[539,600,1297,1323]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="10" pageNumber="11">5E–G</figureCitation> +) +</emphasis> +</paragraph> +</subSubSection> +<subSubSection id="C30A55A8FFEC0D13C783FAE189D8FE40" lastPageId="12" lastPageNumber="13" pageId="10" pageNumber="11" type="multiple"> +<paragraph id="8BAF0623FFEC0D15C783FAE18C3EFEC2" blockId="10.[129,779,1296,1735]" lastBlockId="10.[823,1476,144,1735]" pageId="10" pageNumber="11"> +<taxonomicName id="4C107DA0FFEC0D15C783FAE189CCFAEE" box="[129,254,1335,1359]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +currently includes six genera, being considerably less diverse than proterotheriids and macraucheniids during the Neogene ( +<figureCitation id="132B1AA6FFEC0D15C7EFFAA38817FA2C" box="[237,293,1397,1421]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="10" pageNumber="11">Fig. 2</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFEC0D15C533FAA38BB9FA2C" box="[561,651,1397,1421]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="10" pageNumber="11">Table S1</tableCitation> +). +<bibRefCitation id="EF817BD2FFEC0D15C59DFAA38823FA0C" author="Cifelli and Soria" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria (1983a)</bibRefCitation> +presented a detailed revision of this family, so to avoid unnecessary repetition, only the most relevant taxonomic aspects are discussed here, along with the main advances since their revision. Known as the pygmy litopterns because of their reduced size, the family +<taxonomicName id="4C107DA0FFEC0D15C67DF9C488CEF98B" box="[383,508,1554,1578]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +was named by +<bibRefCitation id="EF817BD2FFEC0D15C59FF9C489E7F9E8" author="Ameghino" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="129 - 67" refId="ref39125" refString="Ameghino F. Caracteres diagnosticos de cincuenta especies nuevas de mamiferos fosiles argentinos. Revista Argentina de Historia Natural 1891 a; 1: 129 - 67." type="journal article" year="1891" yearSuffix="a">Ameghino (1891a)</bibRefCitation> +to place the Early Miocene +<taxonomicName id="4C107DA0FFEC0D15C541F9E78804F9C8" authority="Ameghino 1891 a" authorityName="Ameghino" authorityYear="1891" class="Mammalia" family="Adianthidae" genus="Adianthus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="bucatus"> +<emphasis id="B964DA31FFEC0D15C541F9E78A38F9E8" box="[579,778,1585,1609]" italics="true" pageId="10" pageNumber="11">Adianthus bucatus</emphasis> +<bibRefCitation id="EF817BD2FFEC0D15C783F9878804F9C8" author="Ameghino" box="[129,310,1617,1641]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="129 - 67" refId="ref39125" refString="Ameghino F. Caracteres diagnosticos de cincuenta especies nuevas de mamiferos fosiles argentinos. Revista Argentina de Historia Natural 1891 a; 1: 129 - 67." type="journal article" year="1891" yearSuffix="a">Ameghino 1891a</bibRefCitation> +</taxonomicName> +from the Santa Cruz Formation, +<collectingCountry id="F30746B3FFEC0D15C5A6F9878A3BF9C8" box="[676,777,1617,1641]" name="Argentina" pageId="10" pageNumber="11">Argentina</collectingCountry> +( +<bibRefCitation id="EF817BD2FFEC0D15C789F9A68847F929" author="Cifelli and Soria" box="[139,373,1648,1672]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria 1983a</bibRefCitation> +). Among other distinct anatomical features, adianthids present a trilobed m3 and enamel fossettes in the P4–M3 formed by hypertrophied conular cristae ( +<bibRefCitation id="EF817BD2FFEC0D15C5C8F9798AD0FF09" author="Cifelli and Soria" box="[714,994,144,1735]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria 1983a</bibRefCitation> +). Following new fossil discoveries, +<bibRefCitation id="EF817BD2FFEC0D15C254FF468AB1FF66" author="Ameghino" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="1 - 568" refId="ref39456" refString="Ameghino F. Les formations sedimentaires du cretace superieur et du tertiaire de Patagonie avec un parallele entre leurs faunes mammalogiques et celles de l'ancien continent. Anales del Museo Nacional de Buenos Aires 1906; 15: 1 - 568." type="journal article" year="1906">Ameghino (1906)</bibRefCitation> +added three additional genera to the family +<taxonomicName id="4C107DA0FFEC0D15C23FFF798C8DFF66" box="[1341,1471,175,199]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +: +<taxonomicName id="4C107DA0FFEC0D15C43BFF198D54FF46" authority="Ameghino, 1897" authorityName="Ameghino" authorityYear="1897" box="[825,1126,207,231]" class="Mammalia" family="Adianthidae" genus="Proadiantus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEC0D15C43BFF198A82FF46" box="[825,944,207,231]" italics="true" pageId="10" pageNumber="11">Proadiantus</emphasis> +<bibRefCitation id="EF817BD2FFEC0D15C4BAFF198D54FF46" author="Ameghino" box="[952,1126,207,231]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino, 1897</bibRefCitation> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFEC0D15C39FFF198CF0FF46" authority="Ameghino, 1897" authorityName="Ameghino" authorityYear="1897" box="[1181,1474,207,231]" class="Mammalia" family="Adianthidae" genus="Tricoelodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEC0D15C39FFF198C3EFF46" box="[1181,1292,207,231]" italics="true" pageId="10" pageNumber="11">Tricoelodus</emphasis> +<bibRefCitation id="EF817BD2FFEC0D15C217FF198CF0FF46" author="Ameghino" box="[1301,1474,207,231]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino, 1897</bibRefCitation> +</taxonomicName> +from the Sarmiento Formation (Woodburne +<emphasis id="B964DA31FFEC0D15C20CFF398C0DFEA7" box="[1294,1343,238,262]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2014a), and +<taxonomicName id="4C107DA0FFEC0D15C439FEDB8D47FE84" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[827,1141,269,293]" class="Mammalia" genus="Pseudadiantus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEC0D15C439FEDB8AFAFE84" box="[827,968,269,293]" italics="true" pageId="10" pageNumber="11">Pseudadiantus</emphasis> +<bibRefCitation id="EF817BD2FFEC0D15C4C9FEDB8D47FE84" author="Ameghino" box="[971,1141,269,293]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +. The last was later excluded from the family and formally synonymized with the notopithecine interatheriid +<taxonomicName id="4C107DA0FFEC0D15C4BCFE9A8D08FEC5" authorityName="Ameghino" authorityYear="1901" box="[958,1082,332,356]" class="Mammalia" family="Interatheriidae" genus="Antepithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="10" pageNumber="11" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEC0D15C4BCFE9A8D08FEC5" box="[958,1082,332,356]" italics="true" pageId="10" pageNumber="11">Antepithecus</emphasis> +</taxonomicName> +by Simpson (1967). +</paragraph> +<paragraph id="8BAF0623FFEC0D15C457FEBD8D30FBB4" blockId="10.[823,1476,144,1735]" pageId="10" pageNumber="11"> +<bibRefCitation id="EF817BD2FFEC0D15C457FEBD8ADDFE22" author="Bordas" box="[853,1007,363,387]" firstAuthor="Bordas" pageId="10" pageNumber="11" pagination="413 - 33" refId="ref40435" refString="Bordas AF. Diagnosis sobre algunos mamiferos de las capas con Colpodon, del valle del rio Chubut (Republica Argentina). Physis 1939; 14: 413 - 33." type="journal article" year="1939">Bordas (1939)</bibRefCitation> +added +<taxonomicName id="4C107DA0FFEC0D15C33CFEBD8C6CFE22" authority="Bordas, 1936" authorityName="Bordas" authorityYear="1936" box="[1086,1374,363,387]" class="Mammalia" family="Adianthidae" genus="Proheptaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEC0D15C33CFEBD8DFEFE22" box="[1086,1228,363,387]" italics="true" pageId="10" pageNumber="11">Proheptaconus</emphasis> +<bibRefCitation id="EF817BD2FFEC0D15C3D7FEBD8C6CFE22" author="Bordas" box="[1237,1374,363,387]" firstAuthor="Bordas" pageId="10" pageNumber="11" pagination="110 - 2" refId="ref40403" refString="Bordas AF. Un nuevo mamifero del Colpodon de Gaiman (Proheptaconus trelewense, gen. et sp. nov.). Physis 1936; 12: 110 - 2." type="journal article" year="1936">Bordas, 1936</bibRefCitation> +</taxonomicName> +from the Colhue-Huapi Member of the Sarmiento Formation (Soria 1981) to what he considered the subfamily +<taxonomicName id="4C107DA0FFEC0D15C3F9FE7C8C48FE63" box="[1275,1402,426,450]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="subFamily" subFamily="Adianthinae">Adianthinae</taxonomicName> +, a subfamily of +<taxonomicName id="4C107DA0FFEC0D15C4A0FE1F8D66FE40" box="[930,1108,457,481]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +, which was followed by other authors (Patterson 1940, Simpson and Minoprio 1949). However, Simpson +<emphasis id="B964DA31FFEC0D15C49CFDDF8AE1FD81" box="[926,979,520,544]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +(1962) later restored the family +<taxonomicName id="4C107DA0FFEC0D15C242FDDE8C8DFD81" box="[1344,1471,520,544]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +, but now including a new taxon, +<taxonomicName id="4C107DA0FFEC0D15C398FDF18AEAFDFE" authority="Simpson & Minoprio, 1949" authorityName="Simpson & Minoprio" authorityYear="1949" class="Mammalia" family="Macraucheniidae" genus="Adiantoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="leali"> +<emphasis id="B964DA31FFEC0D15C398FDF18C72FD9E" box="[1178,1344,551,575]" italics="true" pageId="10" pageNumber="11">Adiantoides leali</emphasis> +Simpson & Minoprio, 1949 +</taxonomicName> +from the Divisadero Largo Formation, +<collectingCountry id="F30746B3FFEC0D15C25FFD918CF0FDFE" box="[1373,1474,583,607]" name="Argentina" pageId="10" pageNumber="11">Argentina</collectingCountry> +(age uncertain; +<bibRefCitation id="EF817BD2FFEC0D15C4DDFDB08D9CFDDF" author="Cerdeno" box="[991,1198,614,638]" etAl="et al." firstAuthor="Cerdeno" pageId="10" pageNumber="11" pagination="574 - 7" refId="ref40752" refString="Cerdeno E, Lopez GM, Reguero MA. Biostratigraphic considerations of the Divisaderan faunal assemblage. Journal of Vertebrate Paleontology 2008; 28: 574 - 7. https: // doi. org / 10.1671 / 0272 - 4634 (2008) 28 (574: bcotdf) 2.0. co; 2" type="journal article" year="2008"> +Cerdeño +<emphasis id="B964DA31FFEC0D15C343FDB18D5EFDDF" box="[1089,1132,614,638]" italics="true" pageId="10" pageNumber="11">et al</emphasis> +. 2008 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFEC0D15C3BEFDB08C0BFDDF" author="Lopez" box="[1212,1337,614,638]" firstAuthor="Lopez" pageId="10" pageNumber="11" pagination="410 - 20" refId="ref44025" refString="Lopez GM. Dividaderan: Land Mammal Age or local fauna? In: Madden RH, Carlini AA, Vucetich MG, Kay RF (eds), The paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge: Cambridge University Press, 2010, 410 - 20." type="book chapter" year="2010">López 2010</bibRefCitation> +, Woodburne +<emphasis id="B964DA31FFEC0D15C43BFD508A51FD3C" box="[825,867,645,669]" italics="true" pageId="10" pageNumber="11">et al</emphasis> +. 2014a). +<bibRefCitation id="EF817BD2FFEC0D15C4C7FD538DEEFD3C" author="Bond and Vucetich" box="[965,1244,645,669]" firstAuthor="Bond" pageId="10" pageNumber="11" pagination="107 - 17" refId="ref40299" refString="Bond M, Vucetich MG. Indalecia grandensis gen. et sp. nov. del Eoceno temprano del noroeste argentino, tipo de una nueva subfamilia de los Adianthidae. Revista de la Asociacion Geologica Argentina 1983; 38: 107 - 17." type="journal article" year="1983">Bond and Vucetich (1983)</bibRefCitation> +added to +<taxonomicName id="4C107DA0FFEC0D15C247FD538CF0FD3C" box="[1349,1474,645,669]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +<taxonomicName id="4C107DA0FFEC0D15C439FD738DC5FD1C" authority="Bond & Vucetich, 1983" authorityName="Bond & Vucetich" authorityYear="1983" box="[827,1271,677,701]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="grandensis"> +<emphasis id="B964DA31FFEC0D15C439FD738D32FD1C" box="[827,1024,677,701]" italics="true" pageId="10" pageNumber="11">Indalecia grandensis</emphasis> +<bibRefCitation id="EF817BD2FFEC0D15C306FD738DC5FD1C" author="Bond & Vucetich" box="[1028,1271,677,701]" firstAuthor="Bond" pageId="10" pageNumber="11" pagination="107 - 17" refId="ref40299" refString="Bond M, Vucetich MG. Indalecia grandensis gen. et sp. nov. del Eoceno temprano del noroeste argentino, tipo de una nueva subfamilia de los Adianthidae. Revista de la Asociacion Geologica Argentina 1983; 38: 107 - 17." type="journal article" year="1983">Bond & Vucetich, 1983</bibRefCitation> +</taxonomicName> +from the Lumbrera Formation, +<collectingCountry id="F30746B3FFEC0D15C4BBFD128D2CFD7D" box="[953,1054,708,732]" name="Argentina" pageId="10" pageNumber="11">Argentina</collectingCountry> +(Early Eocene; +<bibRefCitation id="EF817BD2FFEC0D15C3D0FD128C83FD7D" author="Fernicola" box="[1234,1457,708,732]" etAl="et al." firstAuthor="Fernicola" pageId="10" pageNumber="11" pagination="621 - 33" refId="ref41787" refString="Fernicola JC, Zimicz AN, Chornogubsky L et al. The Early Eocene Climatic Optimum at the lower section of the Lumbrera Formation (Ypresian, Salta Province, northwestern Argentina): origin and early diversification of the Cingulata. Journal of Mammalian Evolution 2021; 28: 621 - 33. https: // doi. org / 10.1007 / s 10914 - 021 - 09545 - w" type="journal article" year="2021"> +Fernicola +<emphasis id="B964DA31FFEC0D15C23CFD138C41FD7D" box="[1342,1395,708,732]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2021 +</bibRefCitation> +), based on unique shared anatomical features with +<taxonomicName id="4C107DA0FFEC0D15C24CFD358A53FCBA" authorityName="Simpson & Minoprio" authorityYear="1949" class="Mammalia" family="Macraucheniidae" genus="Adiantoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="leali"> +<emphasis id="B964DA31FFEC0D15C24CFD358A53FCBA" italics="true" pageId="10" pageNumber="11">Adiantoides leali</emphasis> +</taxonomicName> +, such as the absence of a postprotocrista connecting the protocone with the metaconule and a strong parastyle in M1– M2. These features are not present in other adianthids, so they grouped them in the new subfamily +<taxonomicName id="4C107DA0FFEC0D15C3B6FCB78C00FCD8" box="[1204,1330,865,889]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="subFamily" subFamily="Indaleciinae">Indaleciinae</taxonomicName> +, keeping both species tentatively within the family +<taxonomicName id="4C107DA0FFEC0D15C3CAFC568C75FC39" box="[1224,1351,896,920]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +, which was later followed with doubts by +<bibRefCitation id="EF817BD2FFEC0D15C376FC498C45FC16" author="Cifelli and Soria" box="[1140,1399,927,951]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria (1983a)</bibRefCitation> +. In the same work, +<bibRefCitation id="EF817BD2FFEC0D15C4B2FC698D9FFC76" author="Cifelli and Soria" box="[944,1197,959,983]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria (1983a)</bibRefCitation> +added the genus +<taxonomicName id="4C107DA0FFEC0D15C258FC698D3EFC57" authority="Cifelli & Soria, 1983" authorityName="Cifelli & Soria" authorityYear="1983" class="Mammalia" family="Adianthidae" genus="Thadanius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEC0D15C258FC698CF3FC76" box="[1370,1473,959,983]" italics="true" pageId="10" pageNumber="11">Thadanius</emphasis> +Cifelli & Soria, 1983 +</taxonomicName> +from La Salla-Luribay Basin, +<collectingCountry id="F30746B3FFEC0D15C23BFC088CB3FC57" box="[1337,1409,990,1014]" name="Bolivia" pageId="10" pageNumber="11">Bolivia</collectingCountry> +to the family +<taxonomicName id="4C107DA0FFEC0D15C47DFC2B8ACCFBB4" box="[895,1022,1021,1045]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +. +</paragraph> +<paragraph id="8BAF0623FFEC0D13C457FBCB887DFF66" blockId="10.[823,1476,144,1735]" lastBlockId="12.[128,778,144,481]" lastPageId="12" lastPageNumber="13" pageId="10" pageNumber="11"> +Cifelli (1983a) revisited the members of the family, and included +<taxonomicName id="4C107DA0FFEC0D15C48EFBEB8DF0FBF5" authority="Ameghino 1904 b" authorityName="Ameghino" authorityYear="1904" box="[908,1218,1084,1108]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEC0D15C48EFBEB8AC9FBF5" box="[908,1019,1085,1108]" italics="true" pageId="10" pageNumber="11">Proectocion</emphasis> +<bibRefCitation id="EF817BD2FFEC0D15C30AFBEA8DF0FBF5" author="Ameghino" box="[1032,1218,1084,1108]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="1 - 541" refId="ref39426" refString="Ameghino F. Recherches de morphologic phylogenetique sur les molaires superieures des ongules. Anales del Museo Nacional de Buenos Aires 1904 b; 3: 1 - 541." type="journal article" year="1904" yearSuffix="b">Ameghino 1904b</bibRefCitation> +</taxonomicName> +[previously considered a didolodontid by Simpson (1948)] as the earliest member of +<taxonomicName id="4C107DA0FFEC0D15C43BFBAD8A84FB32" box="[825,950,1147,1171]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +based on dental similarities. In addition, he proposed a new superfamily called Macrauchenioidea, to group adianthids and macraucheniids based on dental similarities, such as an absent paraconid and the presence of a paralophid in the lower molars ( +<tableCitation id="C6923398FFEC0D15C31AFB2E8D57FAB1" box="[1048,1125,1272,1296]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="10" pageNumber="11">Table 1</tableCitation> +). Soria (1984a), based on dental similarities with ‘amilnedwarsids’, elevated the rank of the subfamily +<taxonomicName id="4C107DA0FFEC0D15C481FAE18ACDFAEE" box="[899,1023,1335,1359]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="subFamily" subFamily="Indaleciinae">Indaleciinae</taxonomicName> +to the family +<taxonomicName id="4C107DA0FFEC0D15C399FAE18C2BFAEE" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[1179,1305,1335,1359]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Indaleciidae</taxonomicName> +, and later Soria (1989b) classified them as a family of the order +<taxonomicName id="4C107DA0FFEC0D15C248FA808CF0FACC" authorityName="Soria" authorityYear="1989" box="[1354,1474,1366,1389]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="order">Notopterna</taxonomicName> +(see sections of +<taxonomicName id="4C107DA0FFEC0D15C4E3FAA38D95FA2C" box="[993,1191,1397,1421]" class="Mammalia" family="Amilnedwardsiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Amilnedwardsiidae</taxonomicName> +and the order +<taxonomicName id="4C107DA0FFEC0D15C246FAA08C8CFA2C" authorityName="Soria" authorityYear="1989" box="[1348,1470,1398,1421]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="order">Notopterna</taxonomicName> +, and the family +<taxonomicName id="4C107DA0FFEC0D15C4E2FA438D50FA0C" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[992,1122,1429,1453]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Indaleciidae</taxonomicName> +). +<taxonomicName id="4C107DA0FFEC0D15C374FA438DC1FA0C" box="[1142,1267,1429,1453]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +has also been considered by some authors as part of the suborder +<taxonomicName id="4C107DA0FFEC0D15C22CFA628CF0FA6D" box="[1326,1474,1460,1484]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="subOrder" subOrder="Lopholipterna">Lopholipterna</taxonomicName> +(Cifelli 1983a, Soria 2001), which includes the families +<taxonomicName id="4C107DA0FFEC0D15C43BFA258ADBF9AA" box="[825,1001,1523,1547]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFEC0D15C31DFA258DF5F9AA" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1055,1223,1523,1547]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +( +<tableCitation id="C6923398FFEC0D15C3D8FA258C1AF9AA" box="[1242,1320,1523,1547]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="10" pageNumber="11">Table 1</tableCitation> +), a hypothesis that later found phylogenetic support ( +<bibRefCitation id="EF817BD2FFEC0D15C3C4F9C48C0DF98B" author="Cifelli" box="[1222,1343,1554,1578]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation> +; +<figureCitation id="132B1AA6FFEC0D15C24EF9C48CA4F98B" box="[1356,1430,1554,1578]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="10" pageNumber="11">Fig. 1C</figureCitation> +). In the same study, Macrauchenioidea also found support, but only when indaleciids are excluded ( +<bibRefCitation id="EF817BD2FFEC0D15C38AF9878C31F9C8" author="Cifelli" box="[1160,1283,1617,1641]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation> +). Strikingly, there has been not a single study examining the phylogenetic relationships within +<taxonomicName id="4C107DA0FFEC0D15C4BCF9598D09F906" box="[958,1083,1679,1703]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +using modern phylogenetic methods, and most recent phylogenetic studies have included adianthids as outgroups for phylogenies of +<taxonomicName id="4C107DA0FFEA0D13C6C3FF468B43FF09" box="[449,625,144,168]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +(e.g., +<bibRefCitation id="EF817BD2FFEA0D13C5A8FF4689D5FF66" author="Forasiepi" etAl="et al." firstAuthor="Forasiepi" pageId="12" pageNumber="13" pagination="1 - 76" refId="ref41925" refString="Forasiepi AM, MacPhee RDE, Hernandez del Pino S et al. Exceptional skull of Huayqueriana (Mammalia, Litopterna, Macraucheniidae) from the Late Miocene of Argentina: anatomy, systematics, and paleobiological implications. Bulletin of the American Museum of Natural History 2016; 404: 1 - 76." type="journal article" year="2016"> +Forasiepi +<emphasis id="B964DA31FFEA0D13C783FF66899DFF66" box="[129,175,175,199]" italics="true" pageId="12" pageNumber="13">et al.</emphasis> +2016 +</bibRefCitation> +; +<tableCitation id="C6923398FFEA0D13C7F6FF79880DFF66" box="[244,319,175,199]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="12" pageNumber="13">Table 2</tableCitation> +). +</paragraph> +<paragraph id="8BAF0623FFEC0D15C780F8E48D1DF81B" blockId="10.[129,1430,1842,1978]" pageId="10" pageNumber="11">crista; pspc, postparacrista; psplc, postparaconular crista; psprc, postprotocrista. Abbreviations lower molar: co, cristid obliqua; encd, entocristid; end, entoconid; dbgd, distobuccal cingulid; dlgd, distolingual cingulid; hlph, hypolophid; hycd, hypocristid; hyd, hypoconid; hyld, hypoconulid; mbgd, mesiobuccal cingulid; med, metaconid; mlgd, mesiolingual cingulid; mlph, mesolophid; msd, mesoconid; pacd, paracristid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prcd, protocristid; prd, protoconid; prgd, precingulid; pscd, postcristid (or postentocristid); pseld, postentoconulid; psgd, postcingulid; psmcd, postmetacristid.</paragraph> +<caption id="DF6F56ABFFED0D13C773F93C8A36F81A" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12" startId="11.[113,178,1770,1794]" targetBox="[194,1378,147,1740]" targetPageId="11" targetType="figure"> +<paragraph id="8BAF0623FFED0D14C773F93C8CBAF80A" blockId="11.[113,1453,1770,1963]" pageId="11" pageNumber="12"> +<emphasis id="B964DA31FFED0D14C773F93C89FBF8A2" bold="true" box="[113,201,1770,1795]" pageId="11" pageNumber="12">Figure 4.</emphasis> +Right upper molars of +<taxonomicName id="4C107DA0FFED0D14C6A3F93D8B34F8A2" box="[417,518,1771,1795]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="order">Litopterna</taxonomicName> +in occlusal view. A, M3 of +<taxonomicName id="4C107DA0FFED0D14C402F93D8A83F8A2" authorityName="Püschel & Shelley & Williamson & Perini & Wible & Brusatte" authorityYear="2024" baseAuthorityName="Roth" baseAuthorityYear="1899" box="[768,945,1771,1795]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="ligatus"> +<emphasis id="B964DA31FFED0D14C402F93D8A83F8A2" box="[768,945,1771,1795]" italics="true" pageId="11" pageNumber="12">Polymorphis ligatus</emphasis> +</taxonomicName> +(holotype, MLP 12-2169). B, M3 of +<taxonomicName id="4C107DA0FFED0D14C20DF93D8CB2F8A2" authorityName="Roth" authorityYear="1899" box="[1295,1408,1771,1795]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFED0D14C20DF93D8CB2F8A2" box="[1295,1408,1771,1795]" italics="true" pageId="11" pageNumber="12">Polymorphis</emphasis> +</taxonomicName> +? (AMNH-VP 29481). C, M2? of +<taxonomicName id="4C107DA0FFED0D14C6A3F8D18BD5F8BE" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" box="[417,743,1799,1823]" class="Mammalia" family="Proterotheriidae" genus="Polyacrodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="lanciformis"> +<emphasis id="B964DA31FFED0D14C6A3F8D18B4AF8BE" box="[417,632,1799,1823]" italics="true" pageId="11" pageNumber="12">Polyacrodon lanciformis</emphasis> +Roth, 1899 +</taxonomicName> +[MLP 12-2170 (mirrored)]. D, P1–M3 of +<taxonomicName id="4C107DA0FFED0D14C379F8D18C66F8BE" authorityName="Ameghino" authorityYear="1902" box="[1147,1364,1799,1823]" class="Mammalia" family="Macraucheniidae" genus="Cramauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="normalis"> +<emphasis id="B964DA31FFED0D14C379F8D18C66F8BE" box="[1147,1364,1799,1823]" italics="true" pageId="11" pageNumber="12">Cramauchenia normalis</emphasis> +</taxonomicName> +(MLP 85-VII-3-38a). E, P1–M3 of +<taxonomicName id="4C107DA0FFED0D14C67FF8F58B74F89B" authorityName="Scott" authorityYear="1910" box="[381,582,1827,1851]" class="Mammalia" family="Macraucheniidae" genus="Theosodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="garretorum"> +<emphasis id="B964DA31FFED0D14C67FF8F58B74F89B" box="[381,582,1827,1851]" italics="true" pageId="11" pageNumber="12">Theosodon garretorum</emphasis> +</taxonomicName> +(FMNH P 13175). F, P5–M3 of +<taxonomicName id="4C107DA0FFED0D14C483F8F58D01F89B" authorityName="Ameghino" authorityYear="1904" box="[897,1075,1827,1850]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="precisus"> +<emphasis id="B964DA31FFED0D14C483F8F58D01F89B" box="[897,1075,1827,1850]" italics="true" pageId="11" pageNumber="12">Proectocion precisus</emphasis> +</taxonomicName> +(holotype; MACN A 10679). G, P2– M3 of +<taxonomicName id="4C107DA0FFED0D14C7B2F8E988BEF8F6" authorityName="Ameghino" authorityYear="1897" box="[176,396,1855,1879]" class="Mammalia" family="Adianthidae" genus="Tricoelodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="bicuspidatus"> +<emphasis id="B964DA31FFED0D14C7B2F8E988BEF8F6" box="[176,396,1855,1879]" italics="true" pageId="11" pageNumber="12">Tricoelodus bicuspidatus</emphasis> +</taxonomicName> +[cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of +<taxonomicName id="4C107DA0FFED0D14C4A7F8E98D5CF8F7" authorityName="Ameghino" authorityYear="1904" box="[933,1134,1855,1878]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="argentinus"> +<emphasis id="B964DA31FFED0D14C4A7F8E98D5CF8F7" box="[933,1134,1855,1878]" italics="true" pageId="11" pageNumber="12">Proectocion argentinus</emphasis> +</taxonomicName> +(holotype; MACN A 10673). I, M1 of +<taxonomicName id="4C107DA0FFED0D14C7B2F88D8859F8D3" authorityName="Ameghino" authorityYear="1897" box="[176,363,1883,1907]" class="Mammalia" family="Proterotheriidae" genus="Lambdaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="suinus"> +<emphasis id="B964DA31FFED0D14C7B2F88D8859F8D3" box="[176,363,1883,1907]" italics="true" pageId="11" pageNumber="12">Lambdaconus suinus</emphasis> +</taxonomicName> +[MNHN.F.DES162 (mirrored)]. J, P1–M3 of +<taxonomicName id="4C107DA0FFED0D14C421F88D8D3AF8D2" baseAuthorityName="Ameghino" baseAuthorityYear="1887" box="[803,1032,1883,1907]" class="Mammalia" family="Proterotheriidae" genus="Diadiaphorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="majusculus"> +<emphasis id="B964DA31FFED0D14C421F88D8D3AF8D2" box="[803,1032,1883,1907]" italics="true" pageId="11" pageNumber="12">Diadiaphorus majusculus</emphasis> +</taxonomicName> +(MLP 12-304). K, P4–M3 of +<taxonomicName id="4C107DA0FFED0D14C227F88D899BF82F" authorityName="Ameghino" authorityYear="1897" class="Mammalia" family="Proterotheriidae" genus="Lambdaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="suinus"> +<emphasis id="B964DA31FFED0D14C227F88D899BF82F" italics="true" pageId="11" pageNumber="12">Lambdaconus suinus</emphasis> +</taxonomicName> +(MACN A 52-198; P4 was mirrored). L, M1–M3 of +<taxonomicName id="4C107DA0FFED0D14C59FF8A18AA2F82F" authorityName="Paula Couto" authorityYear="1952" box="[669,912,1911,1935]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="prototypica"> +<emphasis id="B964DA31FFED0D14C59FF8A18AA2F82F" box="[669,912,1911,1935]" italics="true" pageId="11" pageNumber="12">Victorlemoinea prototypica</emphasis> +</taxonomicName> +(AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of +<taxonomicName id="4C107DA0FFED0D14C307F8458DCCF80A" authorityName="Cifelli" authorityYear="1983" box="[1029,1278,1939,1963]" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="prodromus"> +<emphasis id="B964DA31FFED0D14C307F8458DCCF80A" box="[1029,1278,1939,1963]" italics="true" pageId="11" pageNumber="12">Paranisolambda prodromus</emphasis> +</taxonomicName> +(cast of DGM +</paragraph> +<paragraph id="8BAF0623FFEA0D13C79EFF1989D8FE40" blockId="12.[128,778,144,481]" pageId="12" pageNumber="13"> +If we follow Cifelli (1983a) and consider +<taxonomicName id="4C107DA0FFEA0D13C539FF198B9EFF47" authorityName="Ameghino" authorityYear="1904" box="[571,684,207,230]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEA0D13C539FF198B9EFF47" box="[571,684,207,230]" italics="true" pageId="12" pageNumber="13">Proectocion</emphasis> +</taxonomicName> +from the Barranca south of Lago Colhue-Huapi of +<collectingCountry id="F30746B3FFEA0D13C540FF388B9BFEA7" box="[578,681,238,262]" name="Argentina" pageId="12" pageNumber="13">Argentina</collectingCountry> +(Middle Eocene; +<bibRefCitation id="EF817BD2FFEA0D13C7D9FED888EEFE84" author="Kramarz and Bond" box="[219,476,269,293]" firstAuthor="Kramarz" pageId="12" pageNumber="13" pagination="203 - 211" refId="ref43514" refString="Kramarz, A., Bond, M. On the status of Isolophodon Roth, 1903 (Mammalia, Astrapotheria) and other little-known Paleogene astrapotheres from central Patagonia. Geobios 2013; 46 (3), 203 - 211." type="journal article" year="2013">Kramarz and Bond 2013</bibRefCitation> +, Woodburne +<emphasis id="B964DA31FFEA0D13C56FFED88BA5FE84" box="[621,663,269,293]" italics="true" pageId="12" pageNumber="13">et al</emphasis> +. 2014a) as the earliest member of +<taxonomicName id="4C107DA0FFEA0D13C671FEFB88C0FEE4" box="[371,498,301,325]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +and considering +<taxonomicName id="4C107DA0FFEA0D13C5A6FEFB89F3FEC5" authorityName="Cifelli" authorityYear="1991" class="Mammalia" family="Adianthidae" genus="Adianthus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="godoyi"> +<emphasis id="B964DA31FFEA0D13C5A6FEFB89F3FEC5" italics="true" pageId="12" pageNumber="13">Adianthus godoyi</emphasis> +</taxonomicName> +from the Río Frías Formation at Alto Río Cisnes, +<collectingCountry id="F30746B3FFEA0D13C5D2FE9A8A3BFEC5" box="[720,777,332,356]" name="Chile" pageId="12" pageNumber="13">Chile</collectingCountry> +as the last member of this family ( +<bibRefCitation id="EF817BD2FFEA0D13C6E5FEBD8B50FE22" author="Cifelli" box="[487,610,363,387]" firstAuthor="Cifelli" pageId="12" pageNumber="13" pagination="119 - 25" refId="ref40949" refString="Cifelli RLA. New adianthid litoptern (Mammalia) from the Miocene of Chile. Revista Chilena de Historia Natural 1991; 64: 119 - 25." type="journal article" year="1991">Cifelli 1991</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFEA0D13C56DFEBD8830FE02" author="de la Cruz and Cortes" firstAuthor="de la Cruz" pageId="12" pageNumber="13" pagination="1 - 70" refId="ref41584" refString="de la Cruz R, Cortes J. Geologia del area oriental de la Hoja Puerto Cisnes, Region Aysen del Gral. Carlos Ibanez del Campo. Serie Geologia Basica 2011; 127: 1 - 70." type="journal article" year="2011">de la Cruz and Cortés 2011</bibRefCitation> +), the family +<taxonomicName id="4C107DA0FFEA0D13C68AFE5D8B35FE02" box="[392,519,395,419]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +encompasses a temporal interval of 42–14.8 Mya ( +<figureCitation id="132B1AA6FFEA0D13C6A1FE7C88C0FE63" box="[419,498,426,450]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="12" pageNumber="13">Fig. 2B</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFEA0D13C783FE1F89EEFE40" box="[129,220,457,481]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="12" pageNumber="13">Table S1</tableCitation> +). +</paragraph> +<subSubSection id="C30A55A8FFEA0D13C61EFDD78B5FFDBA" box="[284,621,513,539]" pageId="12" pageNumber="13" type="nomenclature"> +<paragraph id="8BAF0623FFEA0D13C61EFDD78B5FFDBA" blockId="12.[127,780,513,1641]" box="[284,621,513,539]" pageId="12" pageNumber="13"> +<emphasis id="B964DA31FFEA0D13C61EFDD78B5FFDBA" box="[284,621,513,539]" italics="true" pageId="12" pageNumber="13"> +<taxonomicName id="4C107DA0FFEA0D13C61EFDD788DBFDBA" box="[284,489,513,539]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +( +<figureCitation id="132B1AA6FFEA0D13C6FAFDD48B0FFDBA" box="[504,573,513,539]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="12" pageNumber="13">Figs 4L</figureCitation> +, +<figureCitation id="132B1AA6FFEA0D13C545FDD78B51FDBA" box="[583,611,513,539]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="12" pageNumber="13">5L</figureCitation> +) +</emphasis> +</paragraph> +</subSubSection> +<subSubSection id="C30A55A8FFEA0D11C783FDF18B0BFC57" lastPageId="14" lastPageNumber="15" pageId="12" pageNumber="13" type="multiple"> +<paragraph id="8BAF0623FFEA0D13C783FDF188E9FB70" blockId="12.[127,780,513,1641]" pageId="12" pageNumber="13"> +<taxonomicName id="4C107DA0FFEA0D13C783FDF18851FD9E" box="[129,355,551,575]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +currently has five accepted genera, all with a fossil record exclusively in the Palaeogene ( +<figureCitation id="132B1AA6FFEA0D13C54DFD918BBAFDFE" box="[591,648,583,607]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="12" pageNumber="13">Fig. 2</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFEA0D13C609FDB0885BFDDF" box="[267,361,614,638]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="12" pageNumber="13">Table S1</tableCitation> +). The family +<taxonomicName id="4C107DA0FFEA0D13C6FBFDB08BE9FDDF" box="[505,731,614,638]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +was initially proposed by Soria (1980a) to include +<taxonomicName id="4C107DA0FFEA0D13C56AFD5388AAFD1C" authority="Soria, 1980 a" authorityName="Soria" authorityYear="1980" class="Mammalia" family="Sparnotheriodontidae" genus="Sparnotheriodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="epsilonoides"> +<emphasis id="B964DA31FFEA0D13C56AFD5389C6FD1C" italics="true" pageId="12" pageNumber="13">Sparnotheriodon epsilonoides</emphasis> +Soria, 1980a +</taxonomicName> +from Cañadón Vaca Member, Sarmiento Formation, +<collectingCountry id="F30746B3FFEA0D13C683FD1288DBFD7D" box="[385,489,708,732]" name="Argentina" pageId="12" pageNumber="13">Argentina</collectingCountry> +, which was classified as +<emphasis id="B964DA31FFEA0D13C783FD328836FD5A" box="[129,260,739,763]" italics="true" pageId="12" pageNumber="13">incertae sedis</emphasis> +within the order +<taxonomicName id="4C107DA0FFEA0D13C6CBFD358B68FD5A" baseAuthorityName="Soria" baseAuthorityYear="2001" box="[457,602,739,763]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="order">Notoungulata</taxonomicName> +. The family was later removed from +<taxonomicName id="4C107DA0FFEA0D13C658FCD588DAFCBA" baseAuthorityName="Soria" baseAuthorityYear="2001" box="[346,488,771,795]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="order">Notoungulata</taxonomicName> +and incorporated into the order +<taxonomicName id="4C107DA0FFEA0D13C7BDFCF4881CFC9B" box="[191,302,802,826]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="order">Litopterna</taxonomicName> +, being expanded by Soria (1980b) adding the genera +<taxonomicName id="4C107DA0FFEA0D13C7D2FC978850FCF8" authorityName="Ameghino" authorityYear="1901" box="[208,354,833,857]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEA0D13C7D2FC978850FCF8" box="[208,354,833,857]" italics="true" pageId="12" pageNumber="13">Victorlemoinea</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFEA0D13C6A3FC978A36FCF8" authority="Simpson et al. 1962" authorityName="Simpson" authorityYear="1962" box="[417,772,833,857]" class="Mammalia" family="Proterotheriidae" genus="Phoradiadus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEA0D13C6A3FC978B2FFCF8" box="[417,541,833,857]" italics="true" pageId="12" pageNumber="13">Phoradiadus</emphasis> +Simpson +<emphasis id="B964DA31FFEA0D13C58DFC948BF6FCF8" box="[655,708,833,857]" italics="true" pageId="12" pageNumber="13">et al.</emphasis> +1962 +</taxonomicName> +. +<taxonomicName id="4C107DA0FFEA0D13C783FCB78821FCD8" authorityName="Ameghino" authorityYear="1901" box="[129,275,865,889]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEA0D13C783FCB78821FCD8" box="[129,275,865,889]" italics="true" pageId="12" pageNumber="13">Victorlemoinea</emphasis> +</taxonomicName> +was previously considered an early macraucheniid (Simpson 1945, 1948), and +<taxonomicName id="4C107DA0FFEA0D13C6B0FC568B1CFC39" box="[434,558,896,920]" class="Mammalia" family="Proterotheriidae" genus="Phoradiadus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEA0D13C6B0FC568B1CFC39" box="[434,558,896,920]" italics="true" pageId="12" pageNumber="13">Phoradiadus</emphasis> +</taxonomicName> +was previously considered with doubts as a proterotheriid (Simpson +<emphasis id="B964DA31FFEA0D13C58EFC768B8CFC16" box="[652,702,927,951]" italics="true" pageId="12" pageNumber="13">et al.</emphasis> +1962). Soria’s (1980b) proposal was based on shared similarities in the upper and lower molars, such as the presence of a lophoid metaconule and a closed trigon basin in M2. +<taxonomicName id="4C107DA0FFEA0D13C56AFC2B8BD6FBB4" box="[616,740,1021,1045]" class="Mammalia" family="Proterotheriidae" genus="Phoradiadus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEA0D13C56AFC2B8BD6FBB4" box="[616,740,1021,1045]" italics="true" pageId="12" pageNumber="13">Phoradiadus</emphasis> +</taxonomicName> +includes one species, +<taxonomicName id="4C107DA0FFEA0D13C65AFBCB8B7EFB94" authorityName="Simpson, Minoprio y Patterson" authorityYear="1962" box="[344,588,1053,1077]" class="Mammalia" family="Proterotheriidae" genus="Phoradiadus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="divortiensis"> +<emphasis id="B964DA31FFEA0D13C65AFBCB8B7EFB94" box="[344,588,1053,1077]" italics="true" pageId="12" pageNumber="13">Phoradiadus divortiensis</emphasis> +</taxonomicName> +, from Divisadero Largo Formation, +<collectingRegion id="49D4C8C1FFEA0D13C638FBEA88AEFBF5" box="[314,412,1084,1108]" country="Argentina" name="Mendoza" pageId="12" pageNumber="13">Mendoza</collectingRegion> +, +<collectingCountry id="F30746B3FFEA0D13C6A7FBEA8B38FBF5" box="[421,522,1084,1108]" name="Argentina" pageId="12" pageNumber="13">Argentina</collectingCountry> +(age uncertain; +<bibRefCitation id="EF817BD2FFEA0D13C5B2FBEA89D5FBD2" author="Cerdeno" etAl="et al." firstAuthor="Cerdeno" pageId="12" pageNumber="13" pagination="574 - 7" refId="ref40752" refString="Cerdeno E, Lopez GM, Reguero MA. Biostratigraphic considerations of the Divisaderan faunal assemblage. Journal of Vertebrate Paleontology 2008; 28: 574 - 7. https: // doi. org / 10.1671 / 0272 - 4634 (2008) 28 (574: bcotdf) 2.0. co; 2" type="journal article" year="2008"> +Cerdeño +<emphasis id="B964DA31FFEA0D13C783FB8A899AFBD2" box="[129,168,1115,1139]" italics="true" pageId="12" pageNumber="13">et al</emphasis> +. 2008 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFEA0D13C7F3FB8D8858FBD2" author="Lopez" box="[241,362,1115,1139]" firstAuthor="Lopez" pageId="12" pageNumber="13" pagination="410 - 20" refId="ref44025" refString="Lopez GM. Dividaderan: Land Mammal Age or local fauna? In: Madden RH, Carlini AA, Vucetich MG, Kay RF (eds), The paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge: Cambridge University Press, 2010, 410 - 20." type="book chapter" year="2010">López 2010</bibRefCitation> +, Woodburne +<emphasis id="B964DA31FFEA0D13C6F6FB8A8B2EFBD2" box="[500,540,1115,1139]" italics="true" pageId="12" pageNumber="13">et al</emphasis> +. 2014a). An interesting feature of the teeth of the +<taxonomicName id="4C107DA0FFEA0D13C680FBAD8B56FB32" box="[386,612,1147,1171]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +is that they possess vertical Hunter–Schreger bands, a feature unknown in any other litoptern ( +<bibRefCitation id="EF817BD2FFEA0D13C624FB6F88F8FB70" author="Bond" box="[294,458,1209,1233]" etAl="et al." firstAuthor="Bond" pageId="12" pageNumber="13" pagination="163 - 76" refId="ref40344" refString="Bond M, Reguero MA, Vizcaino SF et al. A new ' South American ungulate' (Mammalia: Litopterna) from the Eocene of the Antarctic Peninsula. Geological Society, London, Special Publications 2006; 258: 163 - 76. https: // doi. org / 10.1144 / gsl. sp. 2006.258.01.12" type="journal article" year="2006"> +Bond +<emphasis id="B964DA31FFEA0D13C661FB6C88A0FB70" box="[355,402,1209,1233]" italics="true" pageId="12" pageNumber="13">et al.</emphasis> +2006 +</bibRefCitation> +). +</paragraph> +<paragraph id="8BAF0623FFEA0D13C79EFB0F8D68FE22" blockId="12.[127,780,513,1641]" lastBlockId="12.[824,1478,144,1641]" pageId="12" pageNumber="13"> +Simpson (1948) tentatively accepted three Patagonian species of the genus +<taxonomicName id="4C107DA0FFEA0D13C64CFB2E88D2FAB1" authorityName="Ameghino" authorityYear="1901" box="[334,480,1272,1296]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEA0D13C64CFB2E88D2FAB1" box="[334,480,1272,1296]" italics="true" pageId="12" pageNumber="13">Victorlemoinea</emphasis> +</taxonomicName> +: +<taxonomicName id="4C107DA0FFEA0D13C6F7FB2E881DFA8E" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="labyrinthica"> +<emphasis id="B964DA31FFEA0D13C6F7FB2E8A3BFAB1" box="[501,777,1272,1296]" italics="true" pageId="12" pageNumber="13">Victorlemoinea labyrinthica</emphasis> +<bibRefCitation id="EF817BD2FFEA0D13C783FAC1881DFA8E" author="Ameghino" box="[129,303,1303,1327]" firstAuthor="Ameghino" pageId="12" pageNumber="13" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFEA0D13C63DFAC18A36FA8E" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[319,772,1303,1327]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="emarginata"> +<emphasis id="B964DA31FFEA0D13C63DFAC18B79FA8E" box="[319,587,1303,1327]" italics="true" pageId="12" pageNumber="13">Victorlemoinea emarginata</emphasis> +<bibRefCitation id="EF817BD2FFEA0D13C557FAC18A36FA8E" author="Ameghino" box="[597,772,1303,1327]" firstAuthor="Ameghino" pageId="12" pageNumber="13" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +, and +<taxonomicName id="4C107DA0FFEA0D13C7AFFAE18B62FAEE" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[173,592,1335,1359]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="longidens"> +<emphasis id="B964DA31FFEA0D13C7AFFAE188ADFAEE" box="[173,415,1335,1359]" italics="true" pageId="12" pageNumber="13">Victorlemoinea longidens</emphasis> +<bibRefCitation id="EF817BD2FFEA0D13C6A7FAE18B62FAEE" author="Ameghino" box="[421,592,1335,1359]" firstAuthor="Ameghino" pageId="12" pageNumber="13" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +, as they were represented only by isolated teeth, the first two being represented by upper teeth, and the last by lower teeth. This means that +<taxonomicName id="4C107DA0FFEA0D13C783FA4389C5FA0C" authorityName="Ameghino" authorityYear="1901" box="[129,247,1429,1453]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="longidens"> +<emphasis id="B964DA31FFEA0D13C783FA4389C5FA0C" box="[129,247,1429,1453]" italics="true" pageId="12" pageNumber="13">V. longidens</emphasis> +</taxonomicName> +could correspond to teeth from the lower dentition of either +<taxonomicName id="4C107DA0FFEA0D13C611FA638891FA6D" authorityName="Ameghino" authorityYear="1901" box="[275,419,1460,1484]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="labyrinthica"> +<emphasis id="B964DA31FFEA0D13C611FA638891FA6D" box="[275,419,1460,1484]" italics="true" pageId="12" pageNumber="13">V. labyrinthica</emphasis> +</taxonomicName> +or +<taxonomicName id="4C107DA0FFEA0D13C6C8FA638B66FA6D" authorityName="Ameghino" authorityYear="1901" box="[458,596,1461,1484]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="emarginata"> +<emphasis id="B964DA31FFEA0D13C6C8FA638B66FA6D" box="[458,596,1461,1484]" italics="true" pageId="12" pageNumber="13">V. emarginata</emphasis> +</taxonomicName> +, in particular the mandibular fragment with m1–m2 of the +<typeStatus id="54ABB881FFEA0D13C54BFA058B97FA4A" box="[585,677,1491,1515]" pageId="12" pageNumber="13" type="holotype">holotype</typeStatus> +(MACN A-10670), as the allegedly associated premolars of this specimen seem to correspond to an isotemnid notoungulate ( +<bibRefCitation id="EF817BD2FFEA0D13C5D3F9C489D9F9E8" author="Bond" etAl="et al." firstAuthor="Bond" pageId="12" pageNumber="13" pagination="163 - 76" refId="ref40344" refString="Bond M, Reguero MA, Vizcaino SF et al. A new ' South American ungulate' (Mammalia: Litopterna) from the Eocene of the Antarctic Peninsula. Geological Society, London, Special Publications 2006; 258: 163 - 76. https: // doi. org / 10.1144 / gsl. sp. 2006.258.01.12" type="journal article" year="2006"> +Bond +<emphasis id="B964DA31FFEA0D13C783F9E48983F9E8" box="[129,177,1585,1609]" italics="true" pageId="12" pageNumber="13">et al.</emphasis> +2006 +</bibRefCitation> +). +<location id="8ECF50F8FFEA0D13C600F9E4882AF9E8" LSID="urn:lsid:plazi:treatment:03B9B735FFE00D17C4BFFC858DC3FD9E:8ECF50F8FFEA0D13C600F9E4882AF9E8" box="[258,280,1586,1609]" country="Brazil" county="Itaborai" municipality="In" name="In" pageId="12" pageNumber="13">In</location> +the genus +<taxonomicName id="4C107DA0FFEA0D13C688F9E78BD5F9E8" authority=", Bond et al. (2006)" authorityName="Bond" authorityYear="2006" box="[394,743,1585,1609]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFEA0D13C688F9E78B28F9E8" box="[394,538,1585,1609]" italics="true" pageId="12" pageNumber="13">Victorlemoinea</emphasis> +, +<bibRefCitation id="EF817BD2FFEA0D13C525F9E78BD5F9E8" author="Bond" box="[551,743,1585,1609]" etAl="et al." firstAuthor="Bond" pageId="12" pageNumber="13" pagination="163 - 76" refId="ref40344" refString="Bond M, Reguero MA, Vizcaino SF et al. A new ' South American ungulate' (Mammalia: Litopterna) from the Eocene of the Antarctic Peninsula. Geological Society, London, Special Publications 2006; 258: 163 - 76. https: // doi. org / 10.1144 / gsl. sp. 2006.258.01.12" type="journal article" year="2006"> +Bond +<emphasis id="B964DA31FFEA0D13C564F9E48BA4F9E8" box="[614,662,1585,1609]" italics="true" pageId="12" pageNumber="13">et al.</emphasis> +(2006) +</bibRefCitation> +</taxonomicName> +included only +<taxonomicName id="4C107DA0FFEA0D13C607F98788A4F9C8" authorityName="Ameghino" authorityYear="1901" box="[261,406,1617,1641]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="labyrinthica"> +<emphasis id="B964DA31FFEA0D13C607F98788A4F9C8" box="[261,406,1617,1641]" italics="true" pageId="12" pageNumber="13">V. labyrinthica</emphasis> +</taxonomicName> +in their list of formally recognized sparnotheriodontid species, and more recently some authors have mentioned only +<taxonomicName id="4C107DA0FFEA0D13C311FF668D92FF66" authorityName="Ameghino" authorityYear="1901" box="[1043,1184,175,199]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="labyrinthica"> +<emphasis id="B964DA31FFEA0D13C311FF668D92FF66" box="[1043,1184,175,199]" italics="true" pageId="12" pageNumber="13">V. labyrinthica</emphasis> +</taxonomicName> +for the +<location id="8ECF50F8FFEA0D13C3EFFF798C67FF66" LSID="urn:lsid:plazi:treatment:03B9B735FFE00D17C4BFFC858DC3FD9E:8ECF50F8FFEA0D13C3EFFF798C67FF66" box="[1261,1365,175,199]" country="Brazil" county="Itaborai" municipality="In" name="Riochican" pageId="12" pageNumber="13">Riochican</location> +and +<location id="8ECF50F8FFEA0D13C287FF668A98FF46" LSID="urn:lsid:plazi:treatment:03B9B735FFE00D17C4BFFC858DC3FD9E:8ECF50F8FFEA0D13C287FF668A98FF46" country="Brazil" county="Itaborai" municipality="In" name="Vacan Patagonian" pageId="12" pageNumber="13">Vacan Patagonian</location> +faunas, omitting +<taxonomicName id="4C107DA0FFEA0D13C35CFF198DD4FF47" authorityName="Ameghino" authorityYear="1901" box="[1118,1254,207,230]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="emarginata"> +<emphasis id="B964DA31FFEA0D13C35CFF198DD4FF47" box="[1118,1254,207,230]" italics="true" pageId="12" pageNumber="13">V. emarginata</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFEA0D13C21AFF198CB9FF46" authorityName="Ameghino" authorityYear="1901" box="[1304,1419,207,231]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="longidens"> +<emphasis id="B964DA31FFEA0D13C21AFF198CB9FF46" box="[1304,1419,207,231]" italics="true" pageId="12" pageNumber="13">V. longidens</emphasis> +</taxonomicName> +(e.g., Reguero +<emphasis id="B964DA31FFEA0D13C497FF398AF6FEA7" box="[917,964,238,262]" italics="true" pageId="12" pageNumber="13">et al.</emphasis> +2014, Gelfo 2016, +<bibRefCitation id="EF817BD2FFEA0D13C381FF388C1AFEA7" author="Gelfo" box="[1155,1320,238,262]" etAl="et al." firstAuthor="Gelfo" pageId="12" pageNumber="13" pagination="274 - 92" refId="ref42638" refString="Gelfo JN, Goin FJ, Bauza N et al. The fossil record of Antarctic land mammals: commented review and hypotheses for future research. Advances in Polar Science 2019; 30: 274 - 92." type="journal article" year="2019"> +Gelfo +<emphasis id="B964DA31FFEA0D13C3C0FF398DC2FEA7" box="[1218,1264,238,262]" italics="true" pageId="12" pageNumber="13">et al.</emphasis> +2019 +</bibRefCitation> +), which can be interpreted as implicitly synonymizing them with +<taxonomicName id="4C107DA0FFEA0D13C232FED88C8EFE84" authorityName="Ameghino" authorityYear="1901" box="[1328,1468,269,293]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="labyrinthica"> +<emphasis id="B964DA31FFEA0D13C232FED88C8EFE84" box="[1328,1468,269,293]" italics="true" pageId="12" pageNumber="13">V. labyrinthica</emphasis> +</taxonomicName> +. +<collectingMunicipality id="6BCB9C59FFEA0D13C43BFEFB8A7DFEE5" box="[825,847,301,324]" pageId="12" pageNumber="13">In</collectingMunicipality> +addition, there is one +<taxonomicName id="4C107DA0FFEA0D13C346FEFB8C1AFEE4" box="[1092,1320,301,325]" pageId="12" pageNumber="13"> +<emphasis id="B964DA31FFEA0D13C346FEFB8DE4FEE4" box="[1092,1238,301,325]" italics="true" pageId="12" pageNumber="13">Victorlemoinea</emphasis> +species +</taxonomicName> +from +<collectingCounty id="62CE7EAFFFEA0D13C26EFEFB8C8CFEE4" box="[1388,1470,301,325]" pageId="12" pageNumber="13">Itaboraí</collectingCounty> +, +<collectingCountry id="F30746B3FFEA0D13C43BFE9A8A44FEC5" box="[825,886,332,356]" name="Brazil" pageId="12" pageNumber="13">Brazil</collectingCountry> +, +<taxonomicName id="4C107DA0FFEA0D13C47DFE9A8C76FEC5" authority="Paula Couto, 1952" authorityName="Paula Couto" authorityYear="1952" box="[895,1348,332,356]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="prototypica"> +<emphasis id="B964DA31FFEA0D13C47DFE9A8DB6FEC5" box="[895,1156,332,356]" italics="true" pageId="12" pageNumber="13">Victorlemoinea prototypica</emphasis> +Paula Couto, 1952 +</taxonomicName> +, also known mostly from dental material. +</paragraph> +<paragraph id="8BAF0623FFEA0D13C457FE5D8D43FC57" blockId="12.[824,1478,144,1641]" pageId="12" pageNumber="13"> +Cifelli (1983b) indirectly assigned some tarsals (i.e., calcaneum and astragalus) to +<taxonomicName id="4C107DA0FFEA0D13C342FE7D8DF8FE63" authorityName="Paula Couto" authorityYear="1952" box="[1088,1226,427,450]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="prototypica"> +<emphasis id="B964DA31FFEA0D13C342FE7D8DF8FE63" box="[1088,1226,427,450]" italics="true" pageId="12" pageNumber="13">V. prototypica</emphasis> +</taxonomicName> +, based on their relative size and abundance. These tarsals were large and scarce, so considering linear regressions with the dentition (m2 area), he argued it could only correspond to either the sparnotheriodontid +<taxonomicName id="4C107DA0FFEA0D13C438FDFE8AE3FD9E" authorityName="Paula Couto" authorityYear="1952" box="[826,977,552,575]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="prototypica"> +<emphasis id="B964DA31FFEA0D13C438FDFE8AE3FD9E" box="[826,977,552,575]" italics="true" pageId="12" pageNumber="13">V. prototypica</emphasis> +</taxonomicName> +or the didolodontid +<taxonomicName id="4C107DA0FFEA0D13C3E1FDFE8C8FFD9E" authorityName="Paula Couto" authorityYear="1952" box="[1251,1469,551,575]" class="Mammalia" family="Didolodontidae" genus="Lamegoia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="conodonta"> +<emphasis id="B964DA31FFEA0D13C3E1FDFE8C8FFD9E" box="[1251,1469,551,575]" italics="true" pageId="12" pageNumber="13">Lamegoia conodonta</emphasis> +</taxonomicName> +. Considering the similarities of the tarsals assigned to +<taxonomicName id="4C107DA0FFEA0D13C2ADFD918A9AFDDF" authorityName="Paula Couto" authorityYear="1952" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="prototypica"> +<emphasis id="B964DA31FFEA0D13C2ADFD918A9AFDDF" italics="true" pageId="12" pageNumber="13">V. prototypica</emphasis> +</taxonomicName> +and others indirectly assigned to didolodontids, such as presenting a medial malleolar facet of the astragalus extending onto the neck and the presence of a dorsal beak on the distal end of the calcaneum,Cifelli (1983a) grouped +<taxonomicName id="4C107DA0FFEA0D13C3E2FD128CF0FD7D" box="[1248,1474,708,732]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFEA0D13C466FD358D3AFD5A" box="[868,1032,739,763]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +under the superfamily +<taxonomicName id="4C107DA0FFEA0D13C3F7FD358C94FD5A" box="[1269,1446,739,763]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="12" pageNumber="13" phylum="Chordata" rank="superFamily" superFamily="Didolodontoidea">Didolodontoidea</taxonomicName> +in the order +<taxonomicName id="4C107DA0FFEA0D13C4A2FCD58D19FCBA" authorityName="Cope" authorityYear="1881" box="[928,1067,771,795]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="12" pageNumber="13" phylum="Chordata" rank="order">Condylarthra</taxonomicName> +( +<tableCitation id="C6923398FFEA0D13C33EFCD58DBAFCBA" box="[1084,1160,771,795]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="12" pageNumber="13">Table 1</tableCitation> +). This hypothesis found some phylogenetic support ( +<bibRefCitation id="EF817BD2FFEA0D13C328FCF48D94FC9B" author="Cifelli" box="[1066,1190,802,826]" firstAuthor="Cifelli" pageId="12" pageNumber="13" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFEA0D13C3B7FCF48C61FC9B" author="Bergqvist" box="[1205,1363,802,826]" firstAuthor="Bergqvist" pageId="12" pageNumber="13" refId="ref39790" refString="Bergqvist LP. Reasociacao de pos-cranio as especies de ungulados da Bacia de S. J. Itaborai (Paleoceno), Estado do Rio de Janeiro, e Filogenia dos ' Condylarthra', e ungulados Sul-Americanos com base no poscranio. D. Phil. Thesis, Universidade Federal do Rio Grande do Sul, Brazil, 1996." type="book" year="1996">Bergqvist 1996</bibRefCitation> +; +<figureCitation id="132B1AA6FFEA0D13C266FCF48C83FC9B" box="[1380,1457,802,826]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="12" pageNumber="13">Fig. 1C</figureCitation> +), and has been followed by some authors (e.g., +<bibRefCitation id="EF817BD2FFEA0D13C215FC978C83FCF8" author="Bergqvist" box="[1303,1457,833,857]" firstAuthor="Bergqvist" pageId="12" pageNumber="13" pagination="107 - 33" refId="ref39850" refString="Bergqvist LP. Postcranial skeleton of the Upper Paleocene of Itaborai Basin, Brazil. In: Sargis EJ, Dagosto M, (eds), Mammalian Evolutionary Morphology: A Tribute to Frederik S. Szalay. New York, NY: Springer-Verlag, 2008, 107 - 33." type="book chapter" year="2008">Bergqvist 2008</bibRefCitation> +). However, other authors have questioned the indirect anatomical assignment of these tarsals to +<taxonomicName id="4C107DA0FFEA0D13C364FC578DDFFC39" authorityName="Paula Couto" authorityYear="1952" box="[1126,1261,897,920]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="prototypica"> +<emphasis id="B964DA31FFEA0D13C364FC578DDFFC39" box="[1126,1261,897,920]" italics="true" pageId="12" pageNumber="13">V. prototypica</emphasis> +</taxonomicName> +(e.g., +<bibRefCitation id="EF817BD2FFEA0D13C22AFC568A98FC16" author="Hoffstetter and Soria" firstAuthor="Hoffstetter" pageId="12" pageNumber="13" pagination="1619 - 22" refId="ref43229" refString="Hoffstetter R, Soria MF. Neolodus colombianus gen. et sp. nov., un noveau Condylarthre (Mammalia) dans le Miocene de Colombie. Comptes Rendus de la Academie de Sciences, Paris 1986; 303: 1619 - 22." type="journal article" year="1986">Hoffstetter and Soria 1986</bibRefCitation> +, Soria 2001, +<bibRefCitation id="EF817BD2FFEA0D13C337FC768DF0FC16" author="Lorente" box="[1077,1218,927,951]" firstAuthor="Lorente" pageId="12" pageNumber="13" refId="ref44083" refString="Lorente M. Desarrollo de Modelos de Asociacion y Clasificaciones de Restos Postcraneanos Aislados de Ungulados Nativos del Paleoceno- Eoceno de America del Sur. D. Phil. Thesis, Universidad Nacional de la Plata, Argentina, 2015." type="book" year="2015">Lorente 2015</bibRefCitation> +), and even suggested an affinity of these tarsals with +<taxonomicName id="4C107DA0FFEA0D13C35AFC698C43FC76" authority="(Soria 2001)" baseAuthorityName="Soria" baseAuthorityYear="2001" box="[1112,1393,959,983]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="order">Notoungulata (Soria 2001)</taxonomicName> +or with +<taxonomicName id="4C107DA0FFEA0D13C438FC088D5EFC57" authority="(Lorente 2015)" baseAuthorityName="Lorente" baseAuthorityYear="2015" box="[826,1132,990,1014]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="12" pageNumber="13" phylum="Chordata" rank="order"> +Astrapotheria ( +<bibRefCitation id="EF817BD2FFEA0D13C4D4FC088D52FC57" author="Lorente" box="[982,1120,990,1014]" firstAuthor="Lorente" pageId="12" pageNumber="13" refId="ref44083" refString="Lorente M. Desarrollo de Modelos de Asociacion y Clasificaciones de Restos Postcraneanos Aislados de Ungulados Nativos del Paleoceno- Eoceno de America del Sur. D. Phil. Thesis, Universidad Nacional de la Plata, Argentina, 2015." type="book" year="2015">Lorente 2015</bibRefCitation> +) +</taxonomicName> +. +</paragraph> +<paragraph id="8BAF0623FFEA0D13C457FC288CA6FACC" blockId="12.[824,1478,144,1641]" pageId="12" pageNumber="13"> +Among the different litoptern families, Soria (2001) considered that sparnotheriodontids were more closely related to the family +<taxonomicName id="4C107DA0FFEA0D13C479FBEA8D10FBF5" authorityName=", Soria" authorityYear="2001" box="[891,1058,1084,1108]" family="Anisolambdidae" pageId="12" pageNumber="13" rank="family">Anisolambdidae</taxonomicName> +due to overall dental similarities between both families, and also considering an isolated M1 (MNRJ +<date id="FFAE20E3FFEA0D13C287FB8D8A7CFB32" bridgedPair="-" pageId="12" pageNumber="13" value="1479-05">1479- V</date> +) from Itaboraí showing an intermediate anatomy between sparnotheriodontids and anisolambdids. Therefore, he created the suborder +<taxonomicName id="4C107DA0FFEA0D13C4C0FB6F8D70FB70" box="[962,1090,1209,1233]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="subOrder" subOrder="Eolitopterna">Eolitopterna</taxonomicName> +to include both families ( +<tableCitation id="C6923398FFEA0D13C248FB6F8CA4FB70" box="[1354,1430,1209,1233]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="12" pageNumber="13">Table 1</tableCitation> +). In addition, Soria (2001) tentatively added +<taxonomicName id="4C107DA0FFEA0D13C3CFFB0F8A9CFAB1" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" class="Mammalia" family="Proterotheriidae" genus="Heteroglyphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="dewoletzky"> +<emphasis id="B964DA31FFEA0D13C3CFFB0F8CF2FB50" box="[1229,1472,1241,1265]" italics="true" pageId="12" pageNumber="13">Heteroglyphis dewoletzky</emphasis> +Roth, 1899 +</taxonomicName> +to +<taxonomicName id="4C107DA0FFEA0D13C4D3FB2E8D81FAB1" box="[977,1203,1272,1296]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +without any detailed anatomical justification; this species is known from a broken upper molar probably from Cerro del Humo, +<collectingCountry id="F30746B3FFEA0D13C3BDFAE18C15FAEE" box="[1215,1319,1335,1359]" name="Argentina" pageId="12" pageNumber="13">Argentina</collectingCountry> +. This taxon was previously referred to +<taxonomicName id="4C107DA0FFEA0D13C31AFA808DF2FACF" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1048,1216,1366,1390]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +by Simpson (1948). +</paragraph> +<paragraph id="8BAF0623FFEA0D13C457FAA08CF0F9C8" blockId="12.[824,1478,144,1641]" pageId="12" pageNumber="13"> +More recently, one genus and two new species from the Eocene La Meseta and Submeseta formations, +<collectingCountry id="F30746B3FFEA0D13C21FFA438CBBFA0C" box="[1309,1417,1429,1453]" name="Antarctica" pageId="12" pageNumber="13">Antarctica</collectingCountry> +, have been added to the family +<taxonomicName id="4C107DA0FFEA0D13C369FA628C7DFA6D" box="[1131,1359,1460,1484]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +, +<taxonomicName id="4C107DA0FFEA0D13C25DFA628DB3FA4A" authority="Bond et al. 2006" authorityName="Bond" authorityYear="2006" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="arquinotiensis"> +<emphasis id="B964DA31FFEA0D13C25DFA628A8DFA4A" italics="true" pageId="12" pageNumber="13">Notiolofos arquinotiensis</emphasis> +<bibRefCitation id="EF817BD2FFEA0D13C4C9FA058DB3FA4A" author="Bond" box="[971,1153,1491,1515]" etAl="et al." firstAuthor="Bond" pageId="12" pageNumber="13" pagination="163 - 76" refId="ref40344" refString="Bond M, Reguero MA, Vizcaino SF et al. A new ' South American ungulate' (Mammalia: Litopterna) from the Eocene of the Antarctic Peninsula. Geological Society, London, Special Publications 2006; 258: 163 - 76. https: // doi. org / 10.1144 / gsl. sp. 2006.258.01.12" type="journal article" year="2006"> +Bond +<emphasis id="B964DA31FFEA0D13C30CFA028D71FA4A" box="[1038,1091,1491,1515]" italics="true" pageId="12" pageNumber="13">et al.</emphasis> +2006 +</bibRefCitation> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFEA0D13C3BFFA058D7EF9AA" authority="Gelfo, Lopez & Santillana, 2017" authorityName="Gelfo, Lopez & Santillana" authorityYear="2017" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="regueroi"> +<emphasis id="B964DA31FFEA0D13C3BFFA058C4AFA4A" box="[1213,1400,1491,1515]" italics="true" pageId="12" pageNumber="13">Notiolofos regueroi</emphasis> +<bibRefCitation id="EF817BD2FFEA0D13C286FA058D7EF9AA" author="Gelfo, Lopez & Santillana" firstAuthor="Gelfo" pageId="12" pageNumber="13" pagination="445 - 55" refId="ref42591" refString="Gelfo JN, Lopez GM, Santillana SN. Eocene ungulate mammals from West Antarctica: implications from their fossil record and a new species. Antarctic Science 2017; 29: 445 - 55. https: // doi. org / 10.1017 / s 0954102017000244" type="journal article" year="2017">Gelfo, López & Santillana, 2017</bibRefCitation> +</taxonomicName> +. They are differentiated mainly by their size and represented by isolated teeth (the latter species only by a m3; +<bibRefCitation id="EF817BD2FFEA0D13C4CCF9E78D44F9E8" author="Gelfo" box="[974,1142,1585,1609]" etAl="et al." firstAuthor="Gelfo" pageId="12" pageNumber="13" pagination="274 - 92" refId="ref42638" refString="Gelfo JN, Goin FJ, Bauza N et al. The fossil record of Antarctic land mammals: commented review and hypotheses for future research. Advances in Polar Science 2019; 30: 274 - 92." type="journal article" year="2019"> +Gelfo +<emphasis id="B964DA31FFEA0D13C30FF9E48D0FF9E8" box="[1037,1085,1585,1609]" italics="true" pageId="12" pageNumber="13">et al.</emphasis> +2019 +</bibRefCitation> +). +<taxonomicName id="4C107DA0FFEA0D13C38FF9E78C4EF9E8" box="[1165,1404,1585,1609]" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="arquinotiensis"> +<emphasis id="B964DA31FFEA0D13C38FF9E78C4EF9E8" box="[1165,1404,1585,1609]" italics="true" pageId="12" pageNumber="13">Notiolofos arquinotiensis</emphasis> +</taxonomicName> +is particularly interesting as it is the most abundant land mammal in +</paragraph> +<paragraph id="8BAF0623FFEA0D13C780F9098CA0F8C6" blockId="12.[129,1473,1759,1979]" pageId="12" pageNumber="13"> +304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 of +<taxonomicName id="4C107DA0FFEA0D13C598F9098A03F956" box="[666,817,1759,1783]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFEA0D13C598F9098A20F956" box="[666,786,1759,1783]" italics="true" pageId="12" pageNumber="13">Anisolambda</emphasis> +sp. +</taxonomicName> +(MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of +<taxonomicName id="4C107DA0FFEA0D13C48CF92D8D13F8B2" authorityName="Simpson" authorityYear="1935" box="[910,1057,1787,1811]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="tshotshe"> +<emphasis id="B964DA31FFEA0D13C48CF92D8D13F8B2" box="[910,1057,1787,1811]" italics="true" pageId="12" pageNumber="13">Wainka tshotshe</emphasis> +</taxonomicName> +( +<typeStatus id="54ABB881FFEA0D13C32FF92D8DBBF8B2" box="[1069,1161,1787,1811]" pageId="12" pageNumber="13" type="holotype">holotype</typeStatus> +; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: +</paragraph> +<paragraph id="8BAF0623FFEA0D13C783F8BD8A36F81A" blockId="12.[129,1473,1759,1979]" pageId="12" pageNumber="13"> +hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal +<quantity id="4CE8ABC6FFEA0D13C5D2F8758A32F81A" box="[720,768,1955,1979]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="12" pageNumber="13" unit="cm" value="1.0">1 cm</quantity> +. +</paragraph> +<caption id="DF6F56ABFFEB0D11C773F92588DFF81B" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14" startId="13.[113,178,1779,1803]" targetBox="[190,1383,148,1748]" targetPageId="13" targetType="figure"> +<paragraph id="8BAF0623FFEB0D12C773F9258CA0F8AA" blockId="13.[113,1456,1779,1971]" box="[113,1426,1779,1803]" pageId="13" pageNumber="14"> +<emphasis id="B964DA31FFEB0D12C773F92589FBF8AA" bold="true" box="[113,201,1779,1803]" pageId="13" pageNumber="14">Figure 5.</emphasis> +Right lower molars of +<taxonomicName id="4C107DA0FFEB0D12C69CF9258B31F8AA" box="[414,515,1779,1803]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="order">Litopterna</taxonomicName> +in occlusal view. A, p1–p4 of +<taxonomicName id="4C107DA0FFEB0D12C41BF9258A8DF8AA" authorityName="Roth" authorityYear="1899" box="[793,959,1779,1803]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="lechei"> +<emphasis id="B964DA31FFEB0D12C41BF9258A8DF8AA" box="[793,959,1779,1803]" italics="true" pageId="13" pageNumber="14">Polymorphis lechei</emphasis> +</taxonomicName> +[holotype; MLP 12-2168 (mirrored)]. B, p1–m3 +</paragraph> +<paragraph id="8BAF0623FFEB0D12C773F8D98CA7F812" blockId="13.[113,1456,1779,1971]" pageId="13" pageNumber="14"> +of +<taxonomicName id="4C107DA0FFEB0D12C78BF8D98853F886" authorityName="Ameghino" authorityYear="1902" box="[137,353,1807,1831]" class="Mammalia" family="Macraucheniidae" genus="Cramauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="normalis"> +<emphasis id="B964DA31FFEB0D12C78BF8D98853F886" box="[137,353,1807,1831]" italics="true" pageId="13" pageNumber="14">Cramauchenia normalis</emphasis> +</taxonomicName> +[MNHN.F.COL181 (mirrored)]. C, m3 of +<taxonomicName id="4C107DA0FFEB0D12C5FCF8D98AE4F886" authorityName="Ameghino" authorityYear="1902" box="[766,982,1807,1831]" class="Mammalia" family="Macraucheniidae" genus="Cramauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="normalis"> +<emphasis id="B964DA31FFEB0D12C5FCF8D98AE4F886" box="[766,982,1807,1831]" italics="true" pageId="13" pageNumber="14">Cramauchenia normalis</emphasis> +</taxonomicName> +(AMNH-VP 29753). D, p1–m3 of +<taxonomicName id="4C107DA0FFEB0D12C221F8D989F0F8E2" authorityName="Ameghino" authorityYear="1887" class="Mammalia" family="Macraucheniidae" genus="Theosodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="lydekkeri"> +<emphasis id="B964DA31FFEB0D12C221F8D989F0F8E2" italics="true" pageId="13" pageNumber="14">Theosodon lydekkeri</emphasis> +</taxonomicName> +[MACN A 24-90 (mirrored)]. E, m2–m3 of +<taxonomicName id="4C107DA0FFEB0D12C568F8FA8BC2F8E2" box="[618,752,1836,1859]" class="Mammalia" family="Adianthidae" genus="Proectocion" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFEB0D12C568F8FA8BE3F8E2" box="[618,721,1836,1859]" italics="true" pageId="13" pageNumber="14">Proectocion</emphasis> +sp. +</taxonomicName> +(MLP 59-II-28-107). F, p5–m3 of +<taxonomicName id="4C107DA0FFEB0D12C33BF8FD8C27F8E2" authorityName="Ameghino" authorityYear="1897" box="[1081,1301,1835,1859]" class="Mammalia" family="Adianthidae" genus="Tricoelodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="bicuspidatus"> +<emphasis id="B964DA31FFEB0D12C33BF8FD8C27F8E2" box="[1081,1301,1835,1859]" italics="true" pageId="13" pageNumber="14">Tricoelodus bicuspidatus</emphasis> +</taxonomicName> +(MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of +<taxonomicName id="4C107DA0FFEB0D12C412F8918AE8F8FE" authorityName="Ameghino" authorityYear="1897" box="[784,986,1863,1887]" class="Mammalia" family="Adianthidae" genus="Proadiantus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="excavatus"> +<emphasis id="B964DA31FFEB0D12C412F8918AE8F8FE" box="[784,986,1863,1887]" italics="true" pageId="13" pageNumber="14">Proadiantus excavatus</emphasis> +</taxonomicName> +(MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of +<taxonomicName id="4C107DA0FFEB0D12C78BF8B58877F8DA" authorityName="Ameghino" authorityYear="1897" box="[137,325,1891,1915]" class="Mammalia" family="Proterotheriidae" genus="Lambdaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="suinus"> +<emphasis id="B964DA31FFEB0D12C78BF8B58877F8DA" box="[137,325,1891,1915]" italics="true" pageId="13" pageNumber="14">Lambdaconus suinus</emphasis> +</taxonomicName> +(MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of +<taxonomicName id="4C107DA0FFEB0D12C364F8B58C79F8DA" baseAuthorityName="Ameghino" baseAuthorityYear="1887" box="[1126,1355,1891,1915]" class="Mammalia" family="Proterotheriidae" genus="Diadiaphorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="majusculus"> +<emphasis id="B964DA31FFEB0D12C364F8B58C79F8DA" box="[1126,1355,1891,1915]" italics="true" pageId="13" pageNumber="14">Diadiaphorus majusculus</emphasis> +</taxonomicName> +(MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of +<taxonomicName id="4C107DA0FFEB0D12C574F8A98A09F836" authorityName="Ameghino" authorityYear="1901" box="[630,827,1919,1943]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="fissidens"> +<emphasis id="B964DA31FFEB0D12C574F8A98A09F836" box="[630,827,1919,1943]" italics="true" pageId="13" pageNumber="14">Anisolambda fissidens</emphasis> +</taxonomicName> +(MACN A 10668; holotype). K, m2–m3 of +<taxonomicName id="4C107DA0FFEB0D12C3DDF8A989E0F812" authorityName="Cifelli" authorityYear="1983" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="prodromus"> +<emphasis id="B964DA31FFEB0D12C3DDF8A989E0F812" italics="true" pageId="13" pageNumber="14">Paranisolambda prodromus</emphasis> +</taxonomicName> +(cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of +<taxonomicName id="4C107DA0FFEB0D12C4FFF84D8DC3F812" authorityName="Paula Couto" authorityYear="1952" box="[1021,1265,1947,1971]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="prototypica"> +<emphasis id="B964DA31FFEB0D12C4FFF84D8DC3F812" box="[1021,1265,1947,1971]" italics="true" pageId="13" pageNumber="14">Victorlemoinea prototypica</emphasis> +</taxonomicName> +(MNRJ 1482V). +</paragraph> +<paragraph id="8BAF0623FFE80D11C783FF4688D9FD9E" blockId="14.[128,779,144,1014]" pageId="14" pageNumber="15"> +the Palaeogene of the La Meseta and Submeseta formations, presenting a continuous fossil record of at least 10 Mya [44–34 Mya ( +<bibRefCitation id="EF817BD2FFE80D11C7BDFF1988B1FF46" author="Douglas" box="[191,387,207,231]" etAl="et al." firstAuthor="Douglas" pageId="14" pageNumber="15" pagination="6582 - 7" refId="ref41674" refString="Douglas PMJ, Affek HP, Ivany LC et al. Pronounced zonal heterogeneity in Eocene southern high-latitude sea surface temperatures. Proceedings of the National Academy of Sciences of the United States of America 2014; 111: 6582 - 7. https: // doi. org / 10.1073 / pnas. 1321441111" type="journal article" year="2014"> +Douglas +<emphasis id="B964DA31FFE80D11C618FF198878FF46" box="[282,330,207,231]" italics="true" pageId="14" pageNumber="15">et al.</emphasis> +2014 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFE80D11C68DFF198B5BFF46" author="Amenabar" box="[399,617,207,231]" etAl="et al." firstAuthor="Amenabar" pageId="14" pageNumber="15" pagination="351 - 66" refId="ref39524" refString="Amenabar CR, Montes M, Nozal F et al. Dinoflagellate cysts of the La Meseta Formation (Middle to Late Eocene), Antarctic Peninsula: implications for biostratigraphy, palaeoceanography and palaeoenvironment. Geological Magazine 2020; 157: 351 - 66. https: // doi. org / 10.1017 / s 0016756819000591" type="journal article" year="2020"> +Amenábar +<emphasis id="B964DA31FFE80D11C502FF198B02FF46" box="[512,560,207,231]" italics="true" pageId="14" pageNumber="15">et al.</emphasis> +2020 +</bibRefCitation> +), but see Gelfo (2016) for a different age interpretation], which has been interpreted as a case of morphological stasis (Gelfo 2016). The presence of the sparnotheriodontid +<taxonomicName id="4C107DA0FFE80D11C6D6FEFB8B05FEE4" box="[468,567,301,325]" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C6D6FEFB8B05FEE4" box="[468,567,301,325]" italics="true" pageId="14" pageNumber="15">Notiolofos</emphasis> +</taxonomicName> +in +<collectingCountry id="F30746B3FFE80D11C55FFEFB8BF5FEE4" box="[605,711,301,325]" name="Antarctica" pageId="14" pageNumber="15">Antarctica</collectingCountry> +raises important biogeographic questions, as no other litoptern lineage seems to have crossed to +<collectingCountry id="F30746B3FFE80D11C6C7FEBD8B03FE22" box="[453,561,363,387]" name="Antarctica" pageId="14" pageNumber="15">Antarctica</collectingCountry> +, and among SANUs excluding +<taxonomicName id="4C107DA0FFE80D11C7EEFE5D887DFE02" box="[236,335,395,419]" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C7EEFE5D887DFE02" box="[236,335,395,419]" italics="true" pageId="14" pageNumber="15">Notiolofos</emphasis> +</taxonomicName> +, only the astrapothere +<taxonomicName id="4C107DA0FFE80D11C544FE5D8876FE63" authority="(Bond et al. 2011)" baseAuthorityName="Bond" baseAuthorityYear="2011" class="Mammalia" genus="Antarctodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="sobrali"> +<emphasis id="B964DA31FFE80D11C544FE5D8A3BFE02" box="[582,777,395,419]" italics="true" pageId="14" pageNumber="15">Antarctodon sobrali</emphasis> +( +<bibRefCitation id="EF817BD2FFE80D11C789FE7C880BFE63" author="Bond" box="[139,313,426,450]" etAl="et al." firstAuthor="Bond" pageId="14" pageNumber="15" pagination="1 - 16" refId="ref40245" refString="Bond M, Kramarz A, Macphee RD, Reguero M. A new astrapothere (Mammalia, Meridiungulata) from La Meseta Formation, Seymour (Marambio) Island, and a reassessment of previous records of Antarctic astrapotheres. American Museum Novitates, 2011 (3718), 1 - 16." type="journal article" year="2011"> +Bond +<emphasis id="B964DA31FFE80D11C7CEFE7D89CCFE63" box="[204,254,426,450]" italics="true" pageId="14" pageNumber="15">et al.</emphasis> +2011 +</bibRefCitation> +) +</taxonomicName> +has been found thus far on this continent. However, as the preservation of mammals is allochthonous in the marine sediments of La Meseta and Submeseta formations a sampling issue cannot be disregarded for explaining the absence of other SANUs ( +<bibRefCitation id="EF817BD2FFE80D11C636FDF188E8FD9E" author="Gelfo" box="[308,474,551,575]" etAl="et al." firstAuthor="Gelfo" pageId="14" pageNumber="15" pagination="274 - 92" refId="ref42638" refString="Gelfo JN, Goin FJ, Bauza N et al. The fossil record of Antarctic land mammals: commented review and hypotheses for future research. Advances in Polar Science 2019; 30: 274 - 92." type="journal article" year="2019"> +Gelfo +<emphasis id="B964DA31FFE80D11C671FDFE88A9FD9E" box="[371,411,551,575]" italics="true" pageId="14" pageNumber="15">et al</emphasis> +. 2019 +</bibRefCitation> +). +</paragraph> +<paragraph id="8BAF0623FFE80D11C79EFD91880BFCF8" blockId="14.[128,779,144,1014]" pageId="14" pageNumber="15"> +Apart from early phylogenetic studies ( +<bibRefCitation id="EF817BD2FFE80D11C525FD918BACFDFE" author="Cifelli" box="[551,670,583,607]" firstAuthor="Cifelli" pageId="14" pageNumber="15" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFE80D11C5AAFD918987FDDF" author="Bergqvist" firstAuthor="Bergqvist" pageId="14" pageNumber="15" refId="ref39790" refString="Bergqvist LP. Reasociacao de pos-cranio as especies de ungulados da Bacia de S. J. Itaborai (Paleoceno), Estado do Rio de Janeiro, e Filogenia dos ' Condylarthra', e ungulados Sul-Americanos com base no poscranio. D. Phil. Thesis, Universidade Federal do Rio Grande do Sul, Brazil, 1996." type="book" year="1996">Bergqvist 1996</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFE80D11C7C1FDB088C7FDDF" author="Bonaparte and Morales" box="[195,501,614,638]" firstAuthor="Bonaparte" pageId="14" pageNumber="15" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales 1997</bibRefCitation> +; +<figureCitation id="132B1AA6FFE80D11C507FDB08B46FDDF" box="[517,628,614,638]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="14" pageNumber="15">Fig. 1C–D</figureCitation> +), most recent phylogenetic studies have included one sparnotheriodontid species among a wide sample of different SANUs (e.g., +<bibRefCitation id="EF817BD2FFE80D11C5D2FD7389C3FD7D" author="Gelfo" etAl="et al." firstAuthor="Gelfo" pageId="14" pageNumber="15" pagination="329 - 35" refId="ref42504" refString="Gelfo JN, Lopez GM, Bond M. A new Xenungulata (Mammalia) from the Paleocene of Patagonia, Argentina. Journal of Paleontology 2008; 82: 329 - 35. https: // doi. org / 10.1666 / 06 - 099.1" type="journal article" year="2008"> +Gelfo +<emphasis id="B964DA31FFE80D11C783FD138986FD7D" box="[129,180,708,732]" italics="true" pageId="14" pageNumber="15">et al.</emphasis> +2008 +</bibRefCitation> +; +<tableCitation id="C6923398FFE80D11C601FD128860FD7D" box="[259,338,708,732]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="14" pageNumber="15">Table 2</tableCitation> +), instead of testing their interfamilial relationships. In terms of the phylogenetic relationships within +<taxonomicName id="4C107DA0FFE80D11C783FCD58857FCBA" box="[129,357,771,795]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +, there is currently only one phylogenetic study that used a small taxon and character sample (Reguero +<emphasis id="B964DA31FFE80D11C5FAFCF589ABFCF8" italics="true" pageId="14" pageNumber="15">et al.</emphasis> +2014; +<tableCitation id="C6923398FFE80D11C7DCFC97881AFCF8" box="[222,296,833,857]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="14" pageNumber="15">Table 2</tableCitation> +). +</paragraph> +<paragraph id="8BAF0623FFE80D11C79EFCB78B0BFC57" blockId="14.[128,779,144,1014]" pageId="14" pageNumber="15"> +The earliest accepted member of +<taxonomicName id="4C107DA0FFE80D11C509FCB78BDFFCD8" box="[523,749,865,889]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +is probably the Early Eocene +<taxonomicName id="4C107DA0FFE80D11C68CFC578B26FC39" authorityName="Paula Couto" authorityYear="1952" box="[398,532,897,920]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="prototypica"> +<emphasis id="B964DA31FFE80D11C68CFC578B26FC39" box="[398,532,897,920]" italics="true" pageId="14" pageNumber="15">V. prototypica</emphasis> +</taxonomicName> +with records in Itaboraí, +<collectingCountry id="F30746B3FFE80D11C782FC49898FFC16" box="[128,189,927,951]" name="Brazil" pageId="14" pageNumber="15">Brazil</collectingCountry> +, and the last could be +<taxonomicName id="4C107DA0FFE80D11C6ACFC498B23FC16" box="[430,529,927,951]" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C6ACFC498B23FC16" box="[430,529,927,951]" italics="true" pageId="14" pageNumber="15">Notiolofos</emphasis> +</taxonomicName> +from +<collectingCountry id="F30746B3FFE80D11C551FC498B8DFC16" box="[595,703,927,951]" name="Antarctica" pageId="14" pageNumber="15">Antarctica</collectingCountry> +, giving a temporal interval of 56.0–34.0 Mya to this family ( +<figureCitation id="132B1AA6FFE80D11C5B4FC698A33FC76" box="[694,769,959,983]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="14" pageNumber="15">Fig. 2B</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFE80D11C67DFC0888E5FC57" box="[383,471,990,1014]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="14" pageNumber="15">Table S1</tableCitation> +, File S2). +</paragraph> +<emphasis id="B964DA31FFE80D11C605FBC38BB1FB91" box="[263,643,1045,1072]" italics="true" pageId="14" pageNumber="15"> +<subSubSection id="C30A55A8FFE80D11C605FBC38892FB8E" box="[263,416,1045,1071]" pageId="14" pageNumber="15" type="nomenclature"> +<paragraph id="8BAF0623FFE80D11C605FBC38892FB8E" blockId="14.[129,779,1045,1766]" box="[263,416,1045,1071]" pageId="14" pageNumber="15"> +<taxonomicName id="4C107DA0FFE80D11C605FBC38892FB8E" authorityName=", Soria" authorityYear="2001" box="[263,416,1045,1071]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName> +</paragraph> +</subSubSection> +<subSubSection id="C30A55A8FFE80D11C6A6FBC08BB1FB91" box="[420,643,1046,1072]" pageId="14" pageNumber="15" type="description"> +<paragraph id="8BAF0623FFE80D11C6A6FBC08BB1FB91" blockId="14.[129,779,1045,1766]" box="[420,643,1046,1072]" pageId="14" pageNumber="15"> +( +<figureCitation id="132B1AA6FFE80D11C6ACFBC08B2CFB91" box="[430,542,1046,1072]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="14" pageNumber="15">Figs 4M–O</figureCitation> +, +<figureCitation id="132B1AA6FFE80D11C525FBC08B4BFB8E" box="[551,633,1046,1072]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="14" pageNumber="15">5J–K, N</figureCitation> +) +</paragraph> +</subSubSection> +</emphasis> +<subSubSection id="C30A55A8FFE80D11C783FBEA8ACFF947" pageId="14" pageNumber="15" type="multiple"> +<paragraph id="8BAF0623FFE80D11C783FBEA8DCFFDFE" blockId="14.[129,779,1045,1766]" lastBlockId="14.[825,1475,144,1766]" pageId="14" pageNumber="15"> +<taxonomicName id="4C107DA0FFE80D11C783FBEA881CFBF5" authorityName=", Soria" authorityYear="2001" box="[129,302,1084,1108]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName> +is currently represented by eight genera with mostly a Palaeogene distribution ( +<figureCitation id="132B1AA6FFE80D11C53CFB8D8B4CFBD2" box="[574,638,1115,1139]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="14" pageNumber="15">Fig. 2</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFE80D11C614FBAD884FFB32" box="[278,381,1147,1171]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="14" pageNumber="15">Table S1</tableCitation> +). The family +<taxonomicName id="4C107DA0FFE80D11C524FBAD8BE1FB32" authorityName=", Soria" authorityYear="2001" box="[550,723,1147,1171]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName> +was proposed by Soria (2001), who elevated the rank of the subfamily +<taxonomicName id="4C107DA0FFE80D11C7F4FB6F8897FB70" box="[246,421,1209,1233]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Anisolambdinae">Anisolambdinae</taxonomicName> +. This subfamily was previously proposed by Cifelli (1983a) to group what he considered early members of the family +<taxonomicName id="4C107DA0FFE80D11C6C4FB2E8B47FAB1" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[454,629,1272,1296]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +with records in the Early Eocene such as +<taxonomicName id="4C107DA0FFE80D11C6C7FAC18B78FA8E" authorityName="Ameghino" authorityYear="1901" box="[453,586,1303,1327]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C6C7FAC18B78FA8E" box="[453,586,1303,1327]" italics="true" pageId="14" pageNumber="15">Anisolambda</emphasis> +</taxonomicName> +, distinctive from proterotheriids with more derived features (subfamily +<taxonomicName id="4C107DA0FFE80D11C783FA808801FACF" box="[129,307,1366,1390]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Proterotheriinae">Proterotheriinae</taxonomicName> +; +<tableCitation id="C6923398FFE80D11C641FA8088A4FACF" box="[323,406,1366,1390]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="14" pageNumber="15">Table 1</tableCitation> +). +<taxonomicName id="4C107DA0FFE80D11C6B6FA808B53FACF" box="[436,609,1366,1390]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Anisolambdinae">Anisolambdinae</taxonomicName> +stands out for the presence of a strong paralophid ending in a large and distinct paraconid. +<taxonomicName id="4C107DA0FFE80D11C62CFA4388E9FA0C" box="[302,475,1429,1453]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Anisolambdinae">Anisolambdinae</taxonomicName> +(currently +<taxonomicName id="4C107DA0FFE80D11C550FA438A30FA0C" authorityName=", Soria" authorityYear="2001" box="[594,770,1429,1453]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName> +) and +<taxonomicName id="4C107DA0FFE80D11C7BAFA628855FA6D" box="[184,359,1460,1484]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Proterotheriinae">Proterotheriinae</taxonomicName> +(currently +<taxonomicName id="4C107DA0FFE80D11C6F2FA628B90FA6D" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[496,674,1460,1484]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +) share a molarized penultimate premolar or P4 (P +<quantity id="4CE8ABC6FFE80D11C537FA058B6FFA4A" box="[565,605,1491,1515]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="14" pageNumber="15" unit="in" value="3.0">3 in</quantity> +Cifelli (1983a); see Material and methods section for tooth position criteria) with a well-developed metacone and mesostyle present, among other features. To a certain extent, Cifelli’s (1983a) proposal was similar to previous arrangements (Simpson 1945, 1948). However, Simpson (1945, 1948) included anisolambdids in the subfamily +<taxonomicName id="4C107DA0FFE80D11C545F9598BDEF906" box="[583,748,1679,1703]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Polymorphinae">Polymorphinae</taxonomicName> +, a subfamily that also included the Late Eocene +<taxonomicName id="4C107DA0FFE80D11C570F9798BC2F966" authorityName="Roth" authorityYear="1899" box="[626,752,1711,1735]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C570F9798BC2F966" box="[626,752,1711,1735]" italics="true" pageId="14" pageNumber="15">Polymorphis</emphasis> +</taxonomicName> +, a taxon later included in +<taxonomicName id="4C107DA0FFE80D11C674F9188B1EF947" box="[374,556,1742,1766]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +(Cifelli 1983a, Soria 2001). In addition, Cifelli (1983a) renamed an anisolambdid from the Early Eocene of Itaboraí, +<collectingCountry id="F30746B3FFE80D11C3B4FF798DC7FF66" box="[1206,1269,175,199]" name="Brazil" pageId="14" pageNumber="15">Brazil</collectingCountry> +, as +<taxonomicName id="4C107DA0FFE80D11C21CFF798D0FFF46" authority="Cifelli 1983 a" authorityName="Cifelli" authorityYear="1983" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="prodromus"> +<emphasis id="B964DA31FFE80D11C21CFF798A97FF46" italics="true" pageId="14" pageNumber="15">Paranisolambda prodromus</emphasis> +Cifelli 1983a +</taxonomicName> +(previously known as +<taxonomicName id="4C107DA0FFE80D11C23FFF198D47FEA7" authority="Paula Couto, 1952" authorityName="Paula Couto" authorityYear="1952" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="prodromus"> +<emphasis id="B964DA31FFE80D11C23FFF198A97FEA7" italics="true" pageId="14" pageNumber="15">Anisolambda prodromus</emphasis> +Paula Couto, 1952 +</taxonomicName> +), based on some dental differences with +<taxonomicName id="4C107DA0FFE80D11C4B6FEDB8D0BFE84" authorityName="Ameghino" authorityYear="1901" box="[948,1081,269,293]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C4B6FEDB8D0BFE84" box="[948,1081,269,293]" italics="true" pageId="14" pageNumber="15">Anisolambda</emphasis> +</taxonomicName> +such as the absence of a postcristid in m3. Based on previous observations (Simpson 1948, Paula Couto 1952) and comparisons with +<taxonomicName id="4C107DA0FFE80D11C3CFFE9A8AA2FE22" authority=", Cifelli (1983 a)" authorityName=", Cifelli" authorityYear="1983" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C3CFFE9A8C43FEC5" box="[1229,1393,332,356]" italics="true" pageId="14" pageNumber="15">Paranisolambda</emphasis> +, Cifelli (1983a) +</taxonomicName> +synonymized the anisolambdids +<taxonomicName id="4C107DA0FFE80D11C20AFEBD8CBDFE22" authorityName="Ameghino" authorityYear="1901" box="[1288,1423,363,387]" class="Mammalia" family="Proterotheriidae" genus="Josepholeidya" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C20AFEBD8CBDFE22" box="[1288,1423,363,387]" italics="true" pageId="14" pageNumber="15">Josepholeidya</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFE80D11C43BFE5D8ADDFE02" authorityName="Ameghino" authorityYear="1901" box="[825,1007,395,419]" class="Mammalia" family="Proterotheriidae" genus="Ricardolydekkeria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C43BFE5D8ADDFE02" box="[825,1007,395,419]" italics="true" pageId="14" pageNumber="15">Ricardolydekkeria</emphasis> +</taxonomicName> +, taxa based on isolated upper teeth, with +<taxonomicName id="4C107DA0FFE80D11C43BFE7C8A8CFE63" authorityName="Ameghino" authorityYear="1901" box="[825,958,426,450]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C43BFE7C8A8CFE63" box="[825,958,426,450]" italics="true" pageId="14" pageNumber="15">Anisolambda</emphasis> +</taxonomicName> +, which was based only on lower dentition. In addition, Cifelli (1983a) included +<taxonomicName id="4C107DA0FFE80D11C3CDFE1F8C4CFE40" authorityName="Simpson" authorityYear="1935" box="[1231,1406,457,481]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="tshotshe"> +<emphasis id="B964DA31FFE80D11C3CDFE1F8C4CFE40" box="[1231,1406,457,481]" italics="true" pageId="14" pageNumber="15">Wainka tshotshe</emphasis> +</taxonomicName> +from Cerro Redondo, +<collectingCountry id="F30746B3FFE80D11C4EFFE3F8D6AFDA0" box="[1005,1112,489,513]" name="Argentina" pageId="14" pageNumber="15">Argentina</collectingCountry> +, in the subfamily +<taxonomicName id="4C107DA0FFE80D11C214FE3F8CF0FDA0" authorityName=", Soria" authorityYear="2001" box="[1302,1474,489,513]" pageId="14" pageNumber="15" rank="subFamily" subFamily="Anisolambdidae">Anisolambdidae</taxonomicName> +based on an m3 with an overall structure very similar to +<taxonomicName id="4C107DA0FFE80D11C43BFDF18A8CFD9E" authorityName="Ameghino" authorityYear="1901" box="[825,958,551,575]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C43BFDF18A8CFD9E" box="[825,958,551,575]" italics="true" pageId="14" pageNumber="15">Anisolambda</emphasis> +</taxonomicName> +. +<taxonomicName id="4C107DA0FFE80D11C4CFFDF18D46FD9E" authorityName="Simpson" authorityYear="1935" box="[973,1140,551,575]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="tshotshe"> +<emphasis id="B964DA31FFE80D11C4CFFDF18D46FD9E" box="[973,1140,551,575]" italics="true" pageId="14" pageNumber="15">Wainka tshotshe</emphasis> +</taxonomicName> +was previously assigned tentatively to +<taxonomicName id="4C107DA0FFE80D11C496FD918DCAFDFE" authority="(Simpson 1948)" baseAuthorityName="Simpson" baseAuthorityYear="1948" box="[916,1272,583,607]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae (Simpson 1948)</taxonomicName> +. +</paragraph> +<paragraph id="8BAF0623FFE80D11C457FDB08AA9FB13" blockId="14.[825,1475,144,1766]" pageId="14" pageNumber="15"> +Apart from elevating the rank of this subfamily to the family +<taxonomicName id="4C107DA0FFE80D11C439FD538D5DFD3C" authority=", Soria (2001)" authorityName=", Soria" authorityYear="2001" box="[827,1135,645,669]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae, Soria (2001)</taxonomicName> +mostly followed Cifelli’s (1983a) interpretations. However, Soria (2001) removed +<taxonomicName id="4C107DA0FFE80D11C25FFD738CF0FD1C" authorityName="Berg" authorityYear="1899" box="[1373,1474,677,701]" class="Mammalia" family="Proterotheriidae" genus="Xesmodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C25FFD738CF0FD1C" box="[1373,1474,677,701]" italics="true" pageId="14" pageNumber="15">Xesmodon</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFE80D11C46BFD128ADCFD7D" authorityName="Roth" authorityYear="1899" box="[873,1006,708,732]" class="Mammalia" family="Proterotheriidae" genus="Heteroglyphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C46BFD128ADCFD7D" box="[873,1006,708,732]" italics="true" pageId="14" pageNumber="15">Heteroglyphis</emphasis> +</taxonomicName> +from the family, tentatively placing them in +<taxonomicName id="4C107DA0FFE80D11C43BFD358AEFFD5A" box="[825,989,739,763]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFE80D11C30FFD358DC3FD5A" box="[1037,1265,739,763]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +, respectively, as they showed important differences in their dentition compared to other anisolambdids. He also added +<taxonomicName id="4C107DA0FFE80D11C3B1FCF48CF0FC9B" authority="Soria, 2001" authorityName="Soria" authorityYear="2001" box="[1203,1474,802,826]" class="Mammalia" family="Proterotheriidae" genus="Lambdaconops" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C3B1FCF48C75FC9B" box="[1203,1351,802,826]" italics="true" pageId="14" pageNumber="15">Lambdaconops</emphasis> +Soria, 2001 +</taxonomicName> +to +<taxonomicName id="4C107DA0FFE80D11C456FC978AC9FCF8" authorityName=", Soria" authorityYear="2001" box="[852,1019,833,857]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName> +due to close dental similarities with the Late Oligocene (Deseadan SALMA) anisolambdid +<taxonomicName id="4C107DA0FFE80D11C23CFCB78D70FC39" authority="Ameghino, 1897" authorityName="Ameghino" authorityYear="1897" class="Mammalia" family="Proterotheriidae" genus="Protheosodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="coniferus"> +<emphasis id="B964DA31FFE80D11C23CFCB78ABDFC39" italics="true" pageId="14" pageNumber="15">Protheosodon coniferus</emphasis> +<bibRefCitation id="EF817BD2FFE80D11C494FC568D70FC39" author="Ameghino" box="[918,1090,896,920]" firstAuthor="Ameghino" pageId="14" pageNumber="15" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino, 1897</bibRefCitation> +</taxonomicName> +, the latter being the only member of this family with associated tarsal remains ( +<bibRefCitation id="EF817BD2FFE80D11C3F5FC498CB0FC16" author="Loomis" box="[1271,1410,927,951]" firstAuthor="Loomis" pageId="14" pageNumber="15" refId="ref44004" refString="Loomis FB. The Deseado Formation of Patagonia. Amherst, MA: The Trustees of Amherst College, 1914." type="book" year="1914">Loomis 1914</bibRefCitation> +). The validity of the different species of +<taxonomicName id="4C107DA0FFE80D11C3A3FC698C10FC76" authorityName="Ameghino" authorityYear="1901" box="[1185,1314,959,983]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE80D11C3A3FC698C10FC76" box="[1185,1314,959,983]" italics="true" pageId="14" pageNumber="15">Anisolambda</emphasis> +</taxonomicName> +was not evaluated by Soria (2001). In addition, Soria (2001) erected the suborder +<taxonomicName id="4C107DA0FFE80D11C475FC2B8AC8FBB4" box="[887,1018,1021,1045]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="subOrder" subOrder="Eolitopterna">Eolitopterna</taxonomicName> +, to include the families +<taxonomicName id="4C107DA0FFE80D11C3ECFC2B8CA7FBB4" authorityName=", Soria" authorityYear="2001" box="[1262,1429,1021,1045]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFE80D11C438FBCB8D2EFB94" box="[826,1052,1053,1077]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +due to overall dental similarities between both families (more details in +<taxonomicName id="4C107DA0FFE80D11C387FBEA8C55FBF5" box="[1157,1383,1084,1108]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +section; +<tableCitation id="C6923398FFE80D11C43BFB8D8AB1FBD2" box="[825,899,1115,1139]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="14" pageNumber="15">Table 1</tableCitation> +). Most recent authors have followed Soria’s (2001) taxonomic proposal for +<taxonomicName id="4C107DA0FFE80D11C30AFBAD8D9DFB32" authorityName=", Soria" authorityYear="2001" box="[1032,1199,1147,1171]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName> +(e.g., +<bibRefCitation id="EF817BD2FFE80D11C3F2FBAD8C57FB32" author="Gelfo" box="[1264,1381,1147,1171]" firstAuthor="Gelfo" pageId="14" pageNumber="15" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo 2006</bibRefCitation> +, Croft +<emphasis id="B964DA31FFE80D11C2B3FBAD8A60FB13" italics="true" pageId="14" pageNumber="15">et al.</emphasis> +2020). +</paragraph> +<paragraph id="8BAF0623FFE80D11C457FB6F8DA3F98B" blockId="14.[825,1475,144,1766]" pageId="14" pageNumber="15"> +Early studies examining the interfamilial relationships of +<taxonomicName id="4C107DA0FFE80D11C43BFB0F8A94FB50" box="[825,934,1241,1265]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="order">Litopterna</taxonomicName> +assumed a monophyletic +<taxonomicName id="4C107DA0FFE80D11C3C3FB0F8C5BFB50" baseAuthorityName="Simpson" baseAuthorityYear="1948" box="[1217,1385,1241,1265]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +that included anisolambdids, without testing this hypothesis ( +<bibRefCitation id="EF817BD2FFE80D11C281FB2E8A5FFA8E" author="Cifelli" firstAuthor="Cifelli" pageId="14" pageNumber="15" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFE80D11C47AFACE8D93FA8E" author="Bonaparte and Morales" box="[888,1185,1303,1327]" firstAuthor="Bonaparte" pageId="14" pageNumber="15" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales 1997</bibRefCitation> +; +<figureCitation id="132B1AA6FFE80D11C3ACFAC18C2BFA8E" box="[1198,1305,1303,1327]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="14" pageNumber="15">Fig. 1C–D</figureCitation> +). More recently, in phylogenetic studies evaluating phylogenetic affinities within +<taxonomicName id="4C107DA0FFE80D11C438FA808AD0FACF" baseAuthorityName="Simpson" baseAuthorityYear="1948" box="[826,994,1366,1390]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +including some anisolambdids as outgroups, anisolambdids were recovered nested within +<taxonomicName id="4C107DA0FFE80D11C218FAA38CF0FA2C" baseAuthorityName="Simpson" baseAuthorityYear="1948" box="[1306,1474,1397,1421]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +(e.g., +<bibRefCitation id="EF817BD2FFE80D11C46DFA438D77FA0C" author="McGrath" box="[879,1093,1429,1453]" etAl="et al." firstAuthor="McGrath" pageId="14" pageNumber="15" pagination="717 - 38" refId="ref44408" refString="McGrath AJ, Flynn JJ, Wyss AR. Proterotheriids and macraucheniids (Litopterna: Mammalia) from the Pampa Castillo Fauna, Chile (early Miocene, Santacrucian SALMA) and a new phylogeny of Proterotheriidae. Journal of Systematic Palaeontology 2020 b; 18: 717 - 38. https: // doi. org / 10.1080 / 14772019.2019.1662500" type="journal article" year="2020" yearSuffix="b"> +McGrath +<emphasis id="B964DA31FFE80D11C4D0FA438D32FA0C" box="[978,1024,1429,1453]" italics="true" pageId="14" pageNumber="15">et al.</emphasis> +2020b +</bibRefCitation> +; +<tableCitation id="C6923398FFE80D11C352FA438DA8FA0C" box="[1104,1178,1429,1453]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="14" pageNumber="15">Table 2</tableCitation> +). However, as not all families of litopterns were included in these analyses, the interfamilial affinities of anisolambdids remain to be tested. In addition, to this date there has been no study examining the phylogenetic relationships within +<taxonomicName id="4C107DA0FFE80D11C4E6F9C48DBFF98B" authorityName=", Soria" authorityYear="2001" box="[996,1165,1554,1578]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName> +. +</paragraph> +<paragraph id="8BAF0623FFE80D11C457F9E78ACFF947" blockId="14.[825,1475,144,1766]" pageId="14" pageNumber="15"> +The earliest accepted member of +<taxonomicName id="4C107DA0FFE80D11C3ADF9E78C64F9E8" authorityName=", Soria" authorityYear="2001" box="[1199,1366,1585,1609]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName> +is +<taxonomicName id="4C107DA0FFE80D11C274F9E78AB4F9C8" authorityName="Simpson" authorityYear="1935" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="tshotshe"> +<emphasis id="B964DA31FFE80D11C274F9E78AB4F9C8" italics="true" pageId="14" pageNumber="15">Wainka tshotshe</emphasis> +</taxonomicName> +from Cerro Redondo, +<collectingRegion id="49D4C8C1FFE80D11C36CF9878DF2F9C8" box="[1134,1216,1617,1641]" country="Argentina" name="Chubut" pageId="14" pageNumber="15">Chubut</collectingRegion> +, +<collectingCountry id="F30746B3FFE80D11C3CAF9878C02F9C8" box="[1224,1328,1617,1641]" name="Argentina" pageId="14" pageNumber="15">Argentina</collectingCountry> +, and the last is +<taxonomicName id="4C107DA0FFE80D11C43BF9A68D2BF929" authorityName="Ameghino" authorityYear="1897" box="[825,1049,1648,1672]" class="Mammalia" family="Proterotheriidae" genus="Protheosodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="coniferus"> +<emphasis id="B964DA31FFE80D11C43BF9A68D2BF929" box="[825,1049,1648,1672]" italics="true" pageId="14" pageNumber="15">Protheosodon coniferus</emphasis> +</taxonomicName> +from Cabeza Blanca, +<collectingRegion id="49D4C8C1FFE80D11C3F9F9A68C7FF929" box="[1275,1357,1648,1672]" country="Argentina" name="Chubut" pageId="14" pageNumber="15">Chubut</collectingRegion> +, +<collectingCountry id="F30746B3FFE80D11C255F9A68C8DF929" box="[1367,1471,1648,1672]" name="Argentina" pageId="14" pageNumber="15">Argentina</collectingCountry> +, giving a temporal interval of 62.5–26.0 Mya for this family ( +<figureCitation id="132B1AA6FFE80D11C29EF9598A64F966" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="14" pageNumber="15">Fig. 2B</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFE80D11C361F9798D8FF966" box="[1123,1213,1711,1735]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="14" pageNumber="15">Table S1</tableCitation> +; Woodburne +<emphasis id="B964DA31FFE80D11C24BF9798C4AF966" box="[1353,1400,1711,1735]" italics="true" pageId="14" pageNumber="15">et al.</emphasis> +2014a, +<bibRefCitation id="EF817BD2FFE80D11C43BF9188ADFF947" author="Krause" box="[825,1005,1742,1766]" etAl="et al." firstAuthor="Krause" pageId="14" pageNumber="15" pagination="886 - 903" refId="ref43716" refString="Krause JM, Clyde WC, Ibanez-Mejia M et al. New age constraints for Early Paleogene strata of central Patagonia, Argentina: implications for the timing of South American Land Mammal Ages. Geological Society of America Bulletin 2017; 129: 886 - 903. https: // doi. org / 10.1130 / b 31561.1" type="journal article" year="2017"> +Krause +<emphasis id="B964DA31FFE80D11C484F9198A86F947" box="[902,948,1742,1766]" italics="true" pageId="14" pageNumber="15">et al.</emphasis> +2017 +</bibRefCitation> +). +</paragraph> +</subSubSection> +<paragraph id="8BAF0623FFE80D11C783F8E58CA3F8C6" blockId="14.[129,1463,1843,1979]" pageId="14" pageNumber="15"> +M, p1–m3 of +<taxonomicName id="4C107DA0FFE80D11C603F8E58B3AF8EA" authorityName="Soria" authorityYear="1980" box="[257,520,1843,1867]" class="Mammalia" family="Sparnotheriodontidae" genus="Sparnotheriodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="epsilonoides"> +<emphasis id="B964DA31FFE80D11C603F8E58B3AF8EA" box="[257,520,1843,1867]" italics="true" pageId="14" pageNumber="15">Sparnotheriodon epsilonoides</emphasis> +</taxonomicName> +(MACN 18225; +<typeStatus id="54ABB881FFE80D11C5AEF8E58A35F8EA" box="[684,775,1843,1867]" pageId="14" pageNumber="15" type="holotype">holotype</typeStatus> +). N, O, m3 of +<taxonomicName id="4C107DA0FFE80D11C48BF8E58D2EF8EA" authorityName="Simpson" authorityYear="1935" box="[905,1052,1843,1867]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="tshotshe"> +<emphasis id="B964DA31FFE80D11C48BF8E58D2EF8EA" box="[905,1052,1843,1867]" italics="true" pageId="14" pageNumber="15">Wainka tshotshe</emphasis> +</taxonomicName> +? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File +</paragraph> +<paragraph id="8BAF0623FFE80D11C783F8BD88DFF81B" blockId="14.[129,1463,1843,1979]" pageId="14" pageNumber="15"> +S2. For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid. Scale bars equal +<quantity id="4CE8ABC6FFE80D11C6BBF87588DAF81B" box="[441,488,1955,1979]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="14" pageNumber="15" unit="cm" value="1.0">1 cm</quantity> +. +</paragraph> +</caption> +</subSubSection> +</caption> +</subSubSection> +</caption> +</subSubSection> +</caption> +</subSubSection> +</treatment> +</document> \ No newline at end of file diff --git a/data/03/B9/B7/03B9B735FFE90D0EC638F9B08B33FD9E.xml b/data/03/B9/B7/03B9B735FFE90D0EC638F9B08B33FD9E.xml new file mode 100644 index 00000000000..abedeca212b --- /dev/null +++ b/data/03/B9/B7/03B9B735FFE90D0EC638F9B08B33FD9E.xml @@ -0,0 +1,2171 @@ +<document id="396B8A729D438E594D7EAEE48C8494E2" ID-DOI="10.1093/zoolinnean/zlae095" ID-ISSN="0024-4082" IM.bibliography_approvedBy="carolina" IM.illustrations_approvedBy="carolina" IM.materialsCitations_approvedBy="carolina" IM.metadata_approvedBy="carolina" IM.tables_requiresApprovalFor="GgImagineBatch,operationResults" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="carolina" checkinTime="1732847695013" checkinUser="plazi" docAuthor="Püschel, Hans P., Shelley, Sarah L., Williamson, Thomas E., Perini, Fernando A., Wible, John R. & Brusatte, Stephen L." docDate="2024" docId="03B9B735FFE90D0EC638F9B08B33FD9E" docLanguage="en" docName="zlae095.pdf" docOrigin="Zoological Journal of the Linnean Society 202 (1)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlae095" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.9:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="9" docTitle="Indaleciidae" docType="treatment" docVersion="1" lastPageNumber="27" masterDocId="FF80CF4DFFE60D1FC702FFD68932FFA1" masterDocTitle="A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates" masterLastPageNumber="50" masterPageNumber="1" pageNumber="16" updateTime="1733769861150" updateUser="carolina"> +<mods:mods id="5B97718148A36E2A4B11289ABF1CF417" xmlns:mods="http://www.loc.gov/mods/v3"> +<mods:titleInfo id="D9A73222374E14CEB15ABDB3A5D2D33E"> +<mods:title id="CFF053801AEBC538B899732294C07E24">A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates</mods:title> +</mods:titleInfo> +<mods:name id="4C83F3CBF1289F1AE2405BA35FB7D9D3" type="personal"> +<mods:role id="4123A8D11931B6DBF46EE177667509D8"> +<mods:roleTerm id="1DBFB3961B9E9C1D9D6EE02EA62626C3">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="5E3EA8A002C86E468ECCE84453C56A7C">Püschel, Hans P.</mods:namePart> +<mods:affiliation id="09E480CE26C95AAD86B4BC9F9C317736">Millenium Nucleus Early Evolutionary Transitions of Mammals (EVOTEM), Red Palentológica U-Chile, Santiago, 7800003, Chile & School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, EH 9 3 FE, United Kingdom</mods:affiliation> +<mods:nameIdentifier id="FD99E0DEBD9D64770DBE581728266613" type="email">hpuschelr@gmail.com</mods:nameIdentifier> +</mods:name> +<mods:name id="DDDF48075515A23CA65B8FA3EB531ACF" type="personal"> +<mods:role id="B844C04199AAB1FCCA7BB3DD974DF207"> +<mods:roleTerm id="AF8D9D2A725400035B336AEAC4143FDC">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="88F5D09E86F0A1BF156A59FE815F3D22">Shelley, Sarah L.</mods:namePart> +<mods:affiliation id="943D5E65BBE8F1AD35DA1C038D00F14C">School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, EH 9 3 FE, United Kingdom</mods:affiliation> +</mods:name> +<mods:name id="336082A90B10CDD7B5EB7124DBF05FC3" type="personal"> +<mods:role id="3A2830FA6703568D723889447CF3BF69"> +<mods:roleTerm id="7905AC1D036B4F5A1AC5C9981CD7CB5E">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="09D218F971F5D04F042B73EC9DB64C5B">Williamson, Thomas E.</mods:namePart> +<mods:affiliation id="6074194580E7CA34D051245743D7D238">New Mexico Museum of Natural History and Science, Albuquerque, NM 87104, United States</mods:affiliation> +</mods:name> +<mods:name id="B473ACDDAD96FC336C6B305AF3F43564" type="personal"> +<mods:role id="1408D2E44A385E06115890B6E01EE070"> +<mods:roleTerm id="543EA6062BBDADCCCA9F256EC97C11ED">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="1D0EF98CD1C6C501879ED1BDAE933F44">Perini, Fernando A.</mods:namePart> +<mods:affiliation id="83ACCF1F7FCF9440E1A9240C56E3814A">Instituto de Ciência Biológicas, UFMG, BH 31270 - 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Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="15" pageNumber="16">Figs 6A</figureCitation> +, +<figureCitation id="132B1AA6FFE90D10C508F9B18B15F920" box="[522,551,1639,1665]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="15" pageNumber="16">7A</figureCitation> +) +</emphasis> +</paragraph> +</subSubSection> +<subSubSection id="C30A55A8FFE90D0EC773F95B8B33FD9E" lastPageId="17" lastPageNumber="18" pageId="15" pageNumber="16" type="multiple"> +<paragraph id="8BAF0623FFE90D10C773F95B8D53FBF5" blockId="15.[113,763,1638,1983]" lastBlockId="15.[808,1461,144,1985]" pageId="15" pageNumber="16"> +<taxonomicName id="4C107DA0FFE90D10C773F95B89C2F904" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[113,240,1677,1701]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Indaleciidae</taxonomicName> +currently includes three accepted genera with an exclusive Palaeogene distribution ( +<figureCitation id="132B1AA6FFE90D10C6DFF97A8B24F965" box="[477,534,1708,1732]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="15" pageNumber="16">Fig. 2</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFE90D10C7A9F91A8839F945" box="[171,267,1740,1764]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="15" pageNumber="16">Table S1</tableCitation> +). +<taxonomicName id="4C107DA0FFE90D10C626F91A8891F945" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[292,419,1740,1764]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Indaleciidae</taxonomicName> +was initially proposed by Soria (1984a) and better justified in a later article (Soria 1989a), in which he elevated the rank of the previously proposed subfamily +<taxonomicName id="4C107DA0FFE90D10C7BFF8FC880CF8E3" box="[189,318,1834,1858]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="subFamily" subFamily="Indaleciinae">Indaleciinae</taxonomicName> +, which included +<taxonomicName id="4C107DA0FFE90D10C6FEF8FC8BFBF8E3" authorityName="Bond & Vucetich" authorityYear="1983" box="[508,713,1834,1858]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="grandensis"> +<emphasis id="B964DA31FFE90D10C6FEF8FC8BFBF8E3" box="[508,713,1834,1858]" italics="true" pageId="15" pageNumber="16">Indalecia grandensis</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFE90D10C773F89F8B67F8C0" authority="(Bond and Vucetich 1983)" baseAuthorityName="Bond and Vucetich" baseAuthorityYear="1983" box="[113,597,1865,1889]" class="Mammalia" family="Macraucheniidae" genus="Adiantoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="leali"> +<emphasis id="B964DA31FFE90D10C773F89F882FF8C0" box="[113,285,1865,1889]" italics="true" pageId="15" pageNumber="16">Adiantoides leali</emphasis> +( +<bibRefCitation id="EF817BD2FFE90D10C636F89F8B7BF8C0" author="Bond and Vucetich" box="[308,585,1865,1889]" firstAuthor="Bond" pageId="15" pageNumber="16" pagination="107 - 17" refId="ref40299" refString="Bond M, Vucetich MG. Indalecia grandensis gen. et sp. nov. del Eoceno temprano del noroeste argentino, tipo de una nueva subfamilia de los Adianthidae. Revista de la Asociacion Geologica Argentina 1983; 38: 107 - 17." type="journal article" year="1983">Bond and Vucetich 1983</bibRefCitation> +) +</taxonomicName> +. Previously, +<taxonomicName id="4C107DA0FFE90D10C5E1F89F89A8F821" baseAuthorityName="Bond and Vucetich" baseAuthorityYear="1983" class="Mammalia" family="Macraucheniidae" genus="Adiantoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="leali"> +<emphasis id="B964DA31FFE90D10C5E1F89F89A8F821" italics="true" pageId="15" pageNumber="16">A. leali</emphasis> +</taxonomicName> +from Divisadero Largo Formation, +<collectingCountry id="F30746B3FFE90D10C52EF8BE8BA6F821" box="[556,660,1896,1920]" name="Argentina" pageId="15" pageNumber="16">Argentina</collectingCountry> +(age uncertain; +<bibRefCitation id="EF817BD2FFE90D10C7C8F85E88AEF801" author="Cerdeno" box="[202,412,1928,1952]" etAl="et al." firstAuthor="Cerdeno" pageId="15" pageNumber="16" pagination="574 - 7" refId="ref40752" refString="Cerdeno E, Lopez GM, Reguero MA. Biostratigraphic considerations of the Divisaderan faunal assemblage. Journal of Vertebrate Paleontology 2008; 28: 574 - 7. https: // doi. org / 10.1671 / 0272 - 4634 (2008) 28 (574: bcotdf) 2.0. co; 2" type="journal article" year="2008"> +Cerdeño +<emphasis id="B964DA31FFE90D10C62FF85E8868F801" box="[301,346,1928,1952]" italics="true" pageId="15" pageNumber="16">et al</emphasis> +. 2008 +</bibRefCitation> +, Woodburne +<emphasis id="B964DA31FFE90D10C532F85E8B6FF801" box="[560,605,1928,1952]" italics="true" pageId="15" pageNumber="16">et al</emphasis> +. 2014a, +<bibRefCitation id="EF817BD2FFE90D10C5BBF85E8994F81E" author="Lopez" firstAuthor="Lopez" pageId="15" pageNumber="16" pagination="410 - 20" refId="ref44025" refString="Lopez GM. Dividaderan: Land Mammal Age or local fauna? In: Madden RH, Carlini AA, Vucetich MG, Kay RF (eds), The paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge: Cambridge University Press, 2010, 410 - 20." type="book chapter" year="2010">López 2010</bibRefCitation> +), was placed tentatively in +<taxonomicName id="4C107DA0FFE90D10C6C9F8718B79F81E" box="[459,587,1959,1983]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +based on dental similarities with the adianthid +<taxonomicName id="4C107DA0FFE90D10C398FF468C33FF09" authorityName="Ameghino" authorityYear="1891" box="[1178,1281,144,168]" class="Mammalia" family="Adianthidae" genus="Adianthus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C398FF468C33FF09" box="[1178,1281,144,168]" italics="true" pageId="15" pageNumber="16">Adianthus</emphasis> +</taxonomicName> +(Simpson and Minoprio 1949, Simpson +<emphasis id="B964DA31FFE90D10C330FF668D68FF66" box="[1074,1114,175,199]" italics="true" pageId="15" pageNumber="16">et al</emphasis> +. 1962). Later, +<bibRefCitation id="EF817BD2FFE90D10C3EFFF798A46FF46" author="Bond and Vucetich" firstAuthor="Bond" pageId="15" pageNumber="16" pagination="107 - 17" refId="ref40299" refString="Bond M, Vucetich MG. Indalecia grandensis gen. et sp. nov. del Eoceno temprano del noroeste argentino, tipo de una nueva subfamilia de los Adianthidae. Revista de la Asociacion Geologica Argentina 1983; 38: 107 - 17." type="journal article" year="1983">Bond and Vucetich (1983)</bibRefCitation> +described +<taxonomicName id="4C107DA0FFE90D10C4FBFF198DF9FF46" authorityName="Bond & Vucetich" authorityYear="1983" box="[1017,1227,207,231]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="grandensis"> +<emphasis id="B964DA31FFE90D10C4FBFF198DF9FF46" box="[1017,1227,207,231]" italics="true" pageId="15" pageNumber="16">Indalecia grandensis</emphasis> +</taxonomicName> +from the Lumbrera Formation, +<collectingCountry id="F30746B3FFE90D10C4A5FF388D3DFEA7" box="[935,1039,238,262]" name="Argentina" pageId="15" pageNumber="16">Argentina</collectingCountry> +[Early Eocene ( +<bibRefCitation id="EF817BD2FFE90D10C3BDFF388CA5FEA7" author="Fernicola" box="[1215,1431,238,262]" etAl="et al." firstAuthor="Fernicola" pageId="15" pageNumber="16" pagination="621 - 33" refId="ref41787" refString="Fernicola JC, Zimicz AN, Chornogubsky L et al. The Early Eocene Climatic Optimum at the lower section of the Lumbrera Formation (Ypresian, Salta Province, northwestern Argentina): origin and early diversification of the Cingulata. Journal of Mammalian Evolution 2021; 28: 621 - 33. https: // doi. org / 10.1007 / s 10914 - 021 - 09545 - w" type="journal article" year="2021"> +Fernicola +<emphasis id="B964DA31FFE90D10C22BFF398C69FEA7" box="[1321,1371,238,262]" italics="true" pageId="15" pageNumber="16">et al.</emphasis> +2021 +</bibRefCitation> +)], that showed dental similarities with +<taxonomicName id="4C107DA0FFE90D10C3ADFEDB8C65FE84" baseAuthorityName="Bond and Vucetich" baseAuthorityYear="1983" box="[1199,1367,269,293]" class="Mammalia" family="Macraucheniidae" genus="Adiantoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="leali"> +<emphasis id="B964DA31FFE90D10C3ADFEDB8C65FE84" box="[1199,1367,269,293]" italics="true" pageId="15" pageNumber="16">Adiantoides leali</emphasis> +</taxonomicName> +, such as a buccolingually transverse crest connecting the hypocone with the metaconule, and the absence of a postprotocrista connecting the protocone with the metaconule on M1–M2, among other features that justified the proposal of a subfamily +<taxonomicName id="4C107DA0FFE90D10C429FE7C8A98FE63" box="[811,938,426,450]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="subFamily" subFamily="Indaleciinae">Indaleciinae</taxonomicName> +within +<taxonomicName id="4C107DA0FFE90D10C4FAFE7C8D48FE63" box="[1016,1146,426,450]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +. +<bibRefCitation id="EF817BD2FFE90D10C383FE7C8CB6FE63" author="Cifelli and Soria" box="[1153,1412,426,450]" firstAuthor="Cifelli" pageId="15" pageNumber="16" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria (1983a)</bibRefCitation> +tentatively followed +<bibRefCitation id="EF817BD2FFE90D10C4E1FE1F8C37FE40" author="Bond and Vucetich" box="[995,1285,457,481]" firstAuthor="Bond" pageId="15" pageNumber="16" pagination="107 - 17" refId="ref40299" refString="Bond M, Vucetich MG. Indalecia grandensis gen. et sp. nov. del Eoceno temprano del noroeste argentino, tipo de una nueva subfamilia de los Adianthidae. Revista de la Asociacion Geologica Argentina 1983; 38: 107 - 17." type="journal article" year="1983">Bond and Vucetich (1983)</bibRefCitation> +considering the shared presence of fossettes formed by hypertrophied conular cristae in the upper molars in members of +<taxonomicName id="4C107DA0FFE90D10C206FDDE8CB1FD81" box="[1284,1411,520,544]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="subFamily" subFamily="Indaleciinae">Indaleciinae</taxonomicName> +and typical adianthids. However, they also noticed some important features that separated them, such as the lack of a mesostyle on the upper molars and the lack of an entolophid (hypolophid in our matrix nomenclature; +<figureCitation id="132B1AA6FFE90D10C343FD538D48FD3C" box="[1089,1146,645,669]" captionStart="Figure 3" captionStartId="9.[113,178,1773,1797]" captionTargetBox="[220,1352,145,1745]" captionTargetId="graphics-399@9.[425,1168,891,1380]" captionTargetPageId="9" captionText="Figure 3. Dental nomenclature used in the text and the characters. A, right upper molar in occlusal view.B–D, right lower molars in occlusal view. The main cusps of the molars are highlighted in bold.The molars represent structures sometimes present in SANUs and with relevance in the character scoring of this study. The molars do not represent any particular SANU.Molars in A and B were partially based on drawings from Gelfo (2006). Abbreviations upper molar: ecg, buccal cingulum or ectocingulum; es, entostyle; hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mt, metastyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; ppc, preparacrista (or paracrista); pplc, preparaconular crista; pprc, preprotocrista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; pshyc, posthypocrista; psmc, postmetacrista (or metacrista); psmlc, postmetaconular crista; pspc, postparacrista; psplc, postparaconular crista; psprc, postprotocrista. Abbreviations lower molar: co, cristid obliqua; encd, entocristid; end, entoconid; dbgd, distobuccal cingulid; dlgd, distolingual cingulid; hlph, hypolophid; hycd, hypocristid; hyd, hypoconid; hyld, hypoconulid; mbgd, mesiobuccal cingulid; med, metaconid; mlgd, mesiolingual cingulid; mlph, mesolophid; msd, mesoconid; pacd, paracristid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prcd, protocristid; prd, protoconid; prgd, precingulid; pscd, postcristid (or postentocristid); pseld, postentoconulid; psgd, postcingulid; psmcd, postmetacristid." pageId="15" pageNumber="16">Fig. 3</figureCitation> +) on the lower molars, among other features. +<bibRefCitation id="EF817BD2FFE90D10C4D2FD738DD4FD1C" author="Cifelli and Soria" box="[976,1254,677,701]" firstAuthor="Cifelli" pageId="15" pageNumber="16" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria (1983a)</bibRefCitation> +also added a new species +<taxonomicName id="4C107DA0FFE90D10C482FD128D7DFD7D" authorityName="Cifelli & Soria" authorityYear="1983" box="[896,1103,708,732]" class="Mammalia" family="Macraucheniidae" genus="Adiantoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="magnus"> +<emphasis id="B964DA31FFE90D10C482FD128D7DFD7D" box="[896,1103,708,732]" italics="true" pageId="15" pageNumber="16">Adiantoides magnus</emphasis> +</taxonomicName> +from Cañadon Vaca, +<collectingCountry id="F30746B3FFE90D10C247FD128C9DFD7D" box="[1349,1455,708,732]" name="Argentina" pageId="15" pageNumber="16">Argentina</collectingCountry> +, to +<taxonomicName id="4C107DA0FFE90D10C44DFD358AE3FD5A" box="[847,977,739,763]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="subFamily" subFamily="Indaleciinae">Indaleciinae</taxonomicName> +. The differences between indaleciines and adianthids and also the dental similarities of the former with ‘amilnedwarsiids’, led Soria (1989a) to propose elevating the subfamily +<taxonomicName id="4C107DA0FFE90D10C499FC978D28FCF8" box="[923,1050,833,857]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="subFamily" subFamily="Indaleciinae">Indaleciinae</taxonomicName> +to the family +<taxonomicName id="4C107DA0FFE90D10C3C2FC978C73FCF8" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[1216,1345,833,857]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Indaleciidae</taxonomicName> +, including indaleciids, ‘amilnedwarsiids’, and notonychopids under the same order +<taxonomicName id="4C107DA0FFE90D10C4DAFC568D60FC36" authorityName="Soria" authorityYear="1989" box="[984,1106,896,919]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Notopterna</taxonomicName> +(see +<taxonomicName id="4C107DA0FFE90D10C390FC568C64FC39" authorityName="Püschel & Shelley & Williamson & Perini & Wible & Brusatte" authorityYear="2024" box="[1170,1366,896,920]" family="Amilnedwardsidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" rank="family">Amilnedwardsidae</taxonomicName> +and the order +<taxonomicName id="4C107DA0FFE90D10C469FC768AD7FC16" authorityName="Soria" authorityYear="1989" box="[875,997,928,951]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Notopterna</taxonomicName> +section). Apart from some common dental features, indaleciids share features of the ear region with notonychopids, such as anteriorly acuminate petrosals; however, they diverge in the presence of enamel fossettes in the P2– M3 and less developed parastyles in indaleciids, among other dental features (Soria 1989b). +</paragraph> +<paragraph id="8BAF0623FFE90D0EC447FB8D8853FEC5" blockId="15.[808,1461,144,1985]" lastBlockId="17.[112,763,144,575]" lastPageId="17" lastPageNumber="18" pageId="15" pageNumber="16"> +<bibRefCitation id="EF817BD2FFE90D10C447FB8D8AE9FBD2" author="Cifelli" box="[837,987,1115,1139]" firstAuthor="Cifelli" pageId="15" pageNumber="16" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli (1993)</bibRefCitation> +conducted a phylogenetic analysis that included indaleciids (i.e., +<taxonomicName id="4C107DA0FFE90D10C344FBAD8DADFB32" authorityName="Bond & Vucetich" authorityYear="1983" box="[1094,1183,1147,1171]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C344FBAD8DADFB32" box="[1094,1183,1147,1171]" italics="true" pageId="15" pageNumber="16">Indalecia</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFE90D10C3EFFBAD8C53FB32" authorityName="Simpson & Minoprio" authorityYear="1949" box="[1261,1377,1147,1171]" class="Mammalia" family="Macraucheniidae" genus="Adiantoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C3EFFBAD8C53FB32" box="[1261,1377,1147,1171]" italics="true" pageId="15" pageNumber="16">Adiantoides</emphasis> +</taxonomicName> +) in a sample of different litoptern families and didolodontids, and found them to form a monophyletic group closely related to sparnotheriodontids (i.e., +<taxonomicName id="4C107DA0FFE90D10C335FB0F8DFBFB50" box="[1079,1225,1241,1265]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C335FB0F8DFBFB50" box="[1079,1225,1241,1265]" italics="true" pageId="15" pageNumber="16">Victorlemoinea</emphasis> +</taxonomicName> +), and these three taxa together were sister to didolodontids ( +<figureCitation id="132B1AA6FFE90D10C3D6FB2E8C17FAB1" box="[1236,1317,1272,1296]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="15" pageNumber="16">Fig. 1C</figureCitation> +). This result was probably influenced by Cifelli’s (1983b) indirect assignment of tarsals for +<taxonomicName id="4C107DA0FFE90D10C4FCFAE18DA2FAEE" box="[1022,1168,1335,1359]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C4FCFAE18DA2FAEE" box="[1022,1168,1335,1359]" italics="true" pageId="15" pageNumber="16">Victorlemoinea</emphasis> +</taxonomicName> +(see +<taxonomicName id="4C107DA0FFE90D10C3D3FAE18C81FAEE" box="[1233,1459,1335,1359]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +section). When +<bibRefCitation id="EF817BD2FFE90D10C4DAFA808C17FACF" author="Bonaparte and Morales" box="[984,1317,1366,1390]" firstAuthor="Bonaparte" pageId="15" pageNumber="16" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation> +repeated the analysis removing didolodontids and adding notonychopids, they similarly found this close relationship between indaleciids and sparnotheriodontids, without a close relationship with notonychopids ( +<figureCitation id="132B1AA6FFE90D10C4D7FA058D2DFA4A" box="[981,1055,1491,1515]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="15" pageNumber="16">Fig. 1D</figureCitation> +). However, in a different analysis based only on lower molars, +<bibRefCitation id="EF817BD2FFE90D10C313FA258DE2F9AA" author="Gelfo" box="[1041,1232,1523,1547]" etAl="et al." firstAuthor="Gelfo" pageId="15" pageNumber="16" pagination="329 - 35" refId="ref42504" refString="Gelfo JN, Lopez GM, Bond M. A new Xenungulata (Mammalia) from the Paleocene of Patagonia, Argentina. Journal of Paleontology 2008; 82: 329 - 35. https: // doi. org / 10.1666 / 06 - 099.1" type="journal article" year="2008"> +Gelfo +<emphasis id="B964DA31FFE90D10C352FA258D4BF9AA" box="[1104,1145,1523,1547]" italics="true" pageId="15" pageNumber="16">et al</emphasis> +. (2008) +</bibRefCitation> +found indaleciids in a polytomy with +<taxonomicName id="4C107DA0FFE90D10C4C9F9C48DA3F98B" authorityName="Soria" authorityYear="1989" box="[971,1169,1554,1578]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="powelli"> +<emphasis id="B964DA31FFE90D10C4C9F9C48DA3F98B" box="[971,1169,1554,1578]" italics="true" pageId="15" pageNumber="16">Notonychops powelli</emphasis> +</taxonomicName> +, and this polytomy as sister of the sparnotheriodontid +<taxonomicName id="4C107DA0FFE90D10C33DF9E78DD3F9E8" authorityName="Soria" authorityYear="1980" box="[1087,1249,1585,1609]" class="Mammalia" family="Sparnotheriodontidae" genus="Sparnotheriodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C33DF9E78DD3F9E8" box="[1087,1249,1585,1609]" italics="true" pageId="15" pageNumber="16">Sparnotheriodon</emphasis> +</taxonomicName> +. Most authors since then have included +<taxonomicName id="4C107DA0FFE90D10C4FBF9878D47F9C8" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[1017,1141,1617,1641]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Indaleciidae</taxonomicName> +as a family of +<taxonomicName id="4C107DA0FFE90D10C20DF9878C4EF9C8" box="[1295,1404,1617,1641]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Litopterna</taxonomicName> +(e.g., Croft +<emphasis id="B964DA31FFE90D10C469F9A78AABF929" box="[875,921,1648,1672]" italics="true" pageId="15" pageNumber="16">et al</emphasis> +. 2020). Nevertheless, a preliminary study that included ear region characters among other craniodental characters found the indaleciid +<taxonomicName id="4C107DA0FFE90D10C306F9798D6FF966" authorityName="Bond & Vucetich" authorityYear="1983" box="[1028,1117,1711,1735]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C306F9798D6FF966" box="[1028,1117,1711,1735]" italics="true" pageId="15" pageNumber="16">Indalecia</emphasis> +</taxonomicName> +as a sister taxon to astrapotheres and notoungulates, instead of having a close relationship with +<taxonomicName id="4C107DA0FFE90D10C428F93B8DDAF8A4" authority="(Garcia-Lopez and Babot 2014)" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[810,1256,1773,1797]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order"> +Litopterna ( +<bibRefCitation id="EF817BD2FFE90D10C4A5F93B8DEEF8A4" author="Garcia-Lopez and Babot" box="[935,1244,1773,1797]" firstAuthor="Garcia-Lopez" pageId="15" pageNumber="16" refId="ref42013" refString="Garcia-Lopez DA, Babot J. The auditory region of the Middle Eocene Litopterna Indalecia grandensis Bond & Vucetich 1983: anatomical and phylogenetic approach. In: IV International Paleontological Congress, Mendoza, Argentina, 2014; p. 183." type="journal volume" year="2014">García-López and Babot 2014</bibRefCitation> +) +</taxonomicName> +. More recently, in a study that included litopterns, kollpanines, and didolodontids, the indaleciid +<taxonomicName id="4C107DA0FFE90D10C4C2F8FA8D06F8E5" authorityName="Simpson & Minoprio" authorityYear="1949" box="[960,1076,1836,1860]" class="Mammalia" family="Macraucheniidae" genus="Adiantoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C4C2F8FA8D06F8E5" box="[960,1076,1836,1860]" italics="true" pageId="15" pageNumber="16">Adiantoides</emphasis> +</taxonomicName> +was found at the stem of litopterns, being more closely related to them than to didolodontids ( +<bibRefCitation id="EF817BD2FFE90D10C437F8BD8D34F822" author="Kramarz" box="[821,1030,1899,1923]" etAl="et al." firstAuthor="Kramarz" pageId="15" pageNumber="16" refId="ref43620" refString="Kramarz AG, Bond M, MacPhee RDE. On the alleged perissodactyl affinities of the ' condylarth' Escribania chubutensis and other endemic South American ungulate-like placentals. Journal of Vertebrate Paleontology 2021; 41: e 1986716." type="journal volume" year="2021"> +Kramarz +<emphasis id="B964DA31FFE90D10C495F8BD8AF1F822" box="[919,963,1899,1923]" italics="true" pageId="15" pageNumber="16">et al</emphasis> +. 2021 +</bibRefCitation> +). In a different study that also included some early notoungulates, the indaleciid +<taxonomicName id="4C107DA0FFE90D10C3D3F85C8C18F803" authorityName="Bond & Vucetich" authorityYear="1983" box="[1233,1322,1930,1954]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C3D3F85C8C18F803" box="[1233,1322,1930,1954]" italics="true" pageId="15" pageNumber="16">Indalecia</emphasis> +</taxonomicName> +was found in a polytomy that includes sparnotheriodontids, protolipternids, didolodontids, and North American phenacodontids (Zimicz +<emphasis id="B964DA31FFF70D0EC5EBFF4789B1FF66" italics="true" pageId="17" pageNumber="18">et al</emphasis> +. 2022). It is important to mention that most of these studies have a relatively limited taxon and character sample, and none of them have included members of all the litoptern families and early members of the various SANU orders ( +<tableCitation id="C6923398FFF70D0EC530FEDB8B4EFE84" box="[562,636,269,293]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="17" pageNumber="18">Table 2</tableCitation> +). Therefore, including +<taxonomicName id="4C107DA0FFF70D0EC7DAFEFB8866FEE4" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[216,340,301,325]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Indaleciidae</taxonomicName> +within the order +<taxonomicName id="4C107DA0FFF70D0EC50FFEFB8B48FEE4" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[525,634,301,325]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="order">Litopterna</taxonomicName> +can only be considered as tentative. +</paragraph> +<caption id="DF6F56ABFFF60D0EC783F8C589ECF81B" lastPageId="17" lastPageNumber="18" pageId="16" pageNumber="17" startId="16.[129,194,1811,1835]" targetBox="[193,1409,146,1780]" targetPageId="16" targetType="figure"> +<paragraph id="8BAF0623FFF60D0FC783F8C58CA3F88A" blockId="16.[129,1465,1811,1976]" box="[129,1425,1811,1836]" pageId="16" pageNumber="17"> +<emphasis id="B964DA31FFF60D0FC783F8C589EAF88A" bold="true" box="[129,216,1811,1835]" pageId="16" pageNumber="17">Figure 6.</emphasis> +Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid +<taxonomicName id="4C107DA0FFF60D0FC455F8C58D3EF88A" authorityName="Bond & Vucetich" authorityYear="1983" box="[855,1036,1811,1835]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="grandensis"> +<emphasis id="B964DA31FFF60D0FC455F8C58D3EF88A" box="[855,1036,1811,1835]" italics="true" pageId="16" pageNumber="17">Indalecia grandensis</emphasis> +</taxonomicName> +in occlusal view [PVL 4186 (mirrored)]. +</paragraph> +<paragraph id="8BAF0623FFF60D0FC783F8E68C85F816" blockId="16.[129,1465,1811,1976]" pageId="16" pageNumber="17"> +B, crownless P1, and P2–M3 of the notonychopid +<taxonomicName id="4C107DA0FFF60D0FC558F8F98A20F8E6" authorityName="Soria" authorityYear="1989" box="[602,786,1839,1863]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="powelli"> +<emphasis id="B964DA31FFF60D0FC558F8F98A20F8E6" box="[602,786,1839,1863]" italics="true" pageId="16" pageNumber="17">Notonychops powelli</emphasis> +</taxonomicName> +in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid +<taxonomicName id="4C107DA0FFF60D0FC599F89A8A06F8C2" authorityName="Bonaparte and Morales" authorityYear="1997" box="[667,820,1867,1891]" class="Mammalia" family="Notonychopidae" genus="Requisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="vidmari"> +<emphasis id="B964DA31FFF60D0FC599F89A8A06F8C2" box="[667,820,1867,1891]" italics="true" pageId="16" pageNumber="17">Requisia vidmari</emphasis> +</taxonomicName> +in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of +<taxonomicName id="4C107DA0FFF60D0FC3C3F8B18C81F8DE" authorityName="Soria & Powell" authorityYear="1981" box="[1217,1459,1895,1919]" class="Mammalia" family="Eoastrapostylopidae" genus="Eoastrapostylops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="riolorense"> +<emphasis id="B964DA31FFF60D0FC3C3F8B18C81F8DE" box="[1217,1459,1895,1919]" italics="true" pageId="16" pageNumber="17">Eoastrapostylops riolorense</emphasis> +</taxonomicName> +in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate +<taxonomicName id="4C107DA0FFF60D0FC428F8528ACCF83A" box="[810,1022,1924,1947]" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="praecursor"> +<emphasis id="B964DA31FFF60D0FC428F8528ACCF83A" box="[810,1022,1924,1947]" italics="true" pageId="16" pageNumber="17">Simpsonotus praecursor</emphasis> +</taxonomicName> +in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate +<taxonomicName id="4C107DA0FFF60D0FC59FF8498A77F816" box="[669,837,1951,1975]" class="Mammalia" family="Henricosborniidae" genus="Henricosbornia" kingdom="Animalia" order="Notoungulata" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFF60D0FC59FF8498A15F816" box="[669,807,1951,1975]" italics="true" pageId="16" pageNumber="17">Henricosbornia</emphasis> +sp. +</taxonomicName> +(MACN A 10717) in occlusal (F) and distoocclusal (G) views. H, +</paragraph> +<paragraph id="8BAF0623FFF70D0EC78FFEBD8B33FD9E" blockId="17.[112,763,144,575]" pageId="17" pageNumber="18"> +The earliest member of +<taxonomicName id="4C107DA0FFF70D0EC69BFEBD8B27FE22" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[409,533,363,387]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Indaleciidae</taxonomicName> +is the Early Eocene +<taxonomicName id="4C107DA0FFF70D0EC773FE5D8808FE02" authorityName="Bond & Vucetich" authorityYear="1983" box="[113,314,395,419]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="grandensis"> +<emphasis id="B964DA31FFF70D0EC773FE5D8808FE02" box="[113,314,395,419]" italics="true" pageId="17" pageNumber="18">Indalecia grandensis</emphasis> +</taxonomicName> +from the Lumbrera Formation, +<collectingCountry id="F30746B3FFF70D0EC597FE5D8BC8FE02" box="[661,762,395,419]" name="Argentina" pageId="17" pageNumber="18">Argentina</collectingCountry> +(55.0–46.2 Mya; +<bibRefCitation id="EF817BD2FFF70D0EC627FE7C88C1FE63" author="Fernicola" box="[293,499,426,450]" etAl="et al." firstAuthor="Fernicola" pageId="17" pageNumber="18" pagination="621 - 33" refId="ref41787" refString="Fernicola JC, Zimicz AN, Chornogubsky L et al. The Early Eocene Climatic Optimum at the lower section of the Lumbrera Formation (Ypresian, Salta Province, northwestern Argentina): origin and early diversification of the Cingulata. Journal of Mammalian Evolution 2021; 28: 621 - 33. https: // doi. org / 10.1007 / s 10914 - 021 - 09545 - w" type="journal article" year="2021"> +Fernicola +<emphasis id="B964DA31FFF70D0EC688FE7D8881FE63" box="[394,435,426,450]" italics="true" pageId="17" pageNumber="18">et al</emphasis> +. 2021 +</bibRefCitation> +) and the youngest would be an undescribed indaleciid from the Abanico Formation, +<collectingCountry id="F30746B3FFF70D0EC5C1FE1F8BC9FE40" box="[707,763,457,481]" name="Chile" pageId="17" pageNumber="18">Chile</collectingCountry> +(Wyss +<emphasis id="B964DA31FFF70D0EC7BAFE3F89D5FDA0" box="[184,231,489,513]" italics="true" pageId="17" pageNumber="18">et al.</emphasis> +1994), which gives the family a temporal interval of 55.0–~31.5 Mya ( +<figureCitation id="132B1AA6FFF70D0EC630FDDE884CFD81" box="[306,382,520,544]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="17" pageNumber="18">Fig. 2B</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFF70D0EC595FDDE8BC7FD81" box="[663,757,520,544]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="17" pageNumber="18">Table S1</tableCitation> +; +<bibRefCitation id="EF817BD2FFF70D0EC773FDF18828FD9E" author="Flynn" box="[113,282,551,575]" etAl="et al." firstAuthor="Flynn" pageId="17" pageNumber="18" pagination="229 - 59" refId="ref41858" refString="Flynn JJ, Wyss AR, Croft DA et al. The Tinguiririca fauna, Chile: biochronology, paleoecology, biogeography, and a new earliest Oligocene South American land mammal ' age'. Palaeogeography, Palaeoclimatology, Palaeoecology 2003; 195: 229 - 59. https: // doi. org / 10.1016 / s 0031 - 0182 (03) 00360 - 2" type="journal article" year="2003"> +Flynn +<emphasis id="B964DA31FFF70D0EC7B1FDFE89D0FD9E" box="[179,226,551,575]" italics="true" pageId="17" pageNumber="18">et al.</emphasis> +2003 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF70D0EC627FDF188C3FD9E" author="Fernicola" box="[293,497,551,575]" etAl="et al." firstAuthor="Fernicola" pageId="17" pageNumber="18" pagination="621 - 33" refId="ref41787" refString="Fernicola JC, Zimicz AN, Chornogubsky L et al. The Early Eocene Climatic Optimum at the lower section of the Lumbrera Formation (Ypresian, Salta Province, northwestern Argentina): origin and early diversification of the Cingulata. Journal of Mammalian Evolution 2021; 28: 621 - 33. https: // doi. org / 10.1007 / s 10914 - 021 - 09545 - w" type="journal article" year="2021"> +Fernicola +<emphasis id="B964DA31FFF70D0EC688FDFE888AFD9E" box="[394,440,551,575]" italics="true" pageId="17" pageNumber="18">et al.</emphasis> +2021 +</bibRefCitation> +). +</paragraph> +<emphasis id="B964DA31FFF70D0EC614FD898B64FDD8" box="[278,598,607,633]" italics="true" pageId="17" pageNumber="18"> +<subSubSection id="C30A55A8FFF70D0EC614FD898882FDD8" box="[278,432,607,633]" pageId="17" pageNumber="18" type="nomenclature"> +<paragraph id="8BAF0623FFF70D0EC614FD898882FDD8" blockId="17.[113,764,607,1547]" box="[278,432,607,633]" pageId="17" pageNumber="18"> +<taxonomicName id="4C107DA0FFF70D0EC614FD898882FDD8" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[278,432,607,633]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName> +</paragraph> +</subSubSection> +<subSubSection id="C30A55A8FFF70D0EC6B7FD898B64FDD8" box="[437,598,607,633]" pageId="17" pageNumber="18" type="description"> +<paragraph id="8BAF0623FFF70D0EC6B7FD898B64FDD8" blockId="17.[113,764,607,1547]" box="[437,598,607,633]" pageId="17" pageNumber="18"> +( +<figureCitation id="132B1AA6FFF70D0EC6BDFDB68B14FDD8" box="[447,550,607,633]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="17" pageNumber="18">Figs 6B–C</figureCitation> +, +<figureCitation id="132B1AA6FFF70D0EC52DFD898B7EFDD8" box="[559,588,607,633]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="17" pageNumber="18">7B</figureCitation> +) +</paragraph> +</subSubSection> +</emphasis> +<subSubSection id="C30A55A8FFF70D0EC773FD538C1AFCBA" pageId="17" pageNumber="18" type="multiple"> +<paragraph id="8BAF0623FFF70D0EC773FD538D4AFF46" blockId="17.[113,764,607,1547]" lastBlockId="17.[809,1461,144,795]" pageId="17" pageNumber="18"> +<taxonomicName id="4C107DA0FFF70D0EC773FD53882FFD3C" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[113,285,645,669]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName> +currently includes two accepted genera with an exclusively Palaeogene distribution ( +<figureCitation id="132B1AA6FFF70D0EC516FD738B62FD1C" box="[532,592,677,701]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="17" pageNumber="18">Fig. 2</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFF70D0EC7E6FD128872FD7D" box="[228,320,708,732]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="17" pageNumber="18">Table S1</tableCitation> +). +<taxonomicName id="4C107DA0FFF70D0EC657FD128B33FD7D" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[341,513,708,732]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName> +was founded to include +<taxonomicName id="4C107DA0FFF70D0EC771FD358888FD5A" authority="Soria, 1989" authorityName="Soria" authorityYear="1989" box="[115,442,739,763]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="powelli"> +<emphasis id="B964DA31FFF70D0EC771FD35880EFD5A" box="[115,316,739,763]" italics="true" pageId="17" pageNumber="18">Notonychops powelli</emphasis> +Soria, 1989 +</taxonomicName> +from the Rio Loro Formation [age uncertain, although probably pre-Itaboraian SALMA age ( +<bibRefCitation id="EF817BD2FFF70D0EC77EFCF48808FC9B" author="Gelfo" box="[124,314,802,826]" etAl="et al." firstAuthor="Gelfo" pageId="17" pageNumber="18" pagination="993 - 1007" refId="ref42730" refString="Gelfo JN, Garcia-Lopez DA, Bergqvist LP. Phylogenetic relationships and palaeobiology of a new xenungulate (Mammalia: Eutheria) from the Palaeogene of Argentina. Journal of Systematic Palaeontology 2020 b; 18: 993 - 1007. https: // doi. org / 10.1080 / 14772019.2020.171 5496" type="journal article" year="2020" yearSuffix="b"> +Gelfo +<emphasis id="B964DA31FFF70D0EC7BDFCF589C0FC9B" box="[191,242,802,826]" italics="true" pageId="17" pageNumber="18">et al.</emphasis> +2020b +</bibRefCitation> +)], an ungulate with great development of the parastyle in M1–M3, which among other features, makes it anatomically distinctive from any other SANU previously described (Soria 1989b). Soria (1989b) considered that +<taxonomicName id="4C107DA0FFF70D0EC5E1FC578987FC16" authorityName="Soria" authorityYear="1989" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="powelli"> +<emphasis id="B964DA31FFF70D0EC5E1FC578987FC16" italics="true" pageId="17" pageNumber="18">N. powelli</emphasis> +</taxonomicName> +was dentally convergent with members of the family +<taxonomicName id="4C107DA0FFF70D0EC773FC6988A1FC76" authority="(Tillodonta)" baseAuthorityName="Tillodonta" box="[113,403,959,983]" class="Mammalia" family="Esthonychidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Esthonychidae (Tillodonta)</taxonomicName> +, particularly with the lower molars being brachydont and selenodont with a subequal trigonid and talonid length. However, +<taxonomicName id="4C107DA0FFF70D0EC670FC2888E1FBB4" authorityName="Soria" authorityYear="1989" box="[370,467,1021,1045]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="powelli"> +<emphasis id="B964DA31FFF70D0EC670FC2888E1FBB4" box="[370,467,1021,1045]" italics="true" pageId="17" pageNumber="18">N. powelli</emphasis> +</taxonomicName> +shares important anatomical features with the dentition of ‘amilnedwarsids’ and indaleciids, and of the ear region with indaleciids, which madeSoria (1989b) place it within the order +<taxonomicName id="4C107DA0FFF70D0EC686FB8A88CEFBD2" authorityName="Soria" authorityYear="1989" box="[388,508,1116,1139]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="order">Notopterna</taxonomicName> +(see +<taxonomicName id="4C107DA0FFF70D0EC539FB8D8BC9FBD2" authorityName="Püschel & Shelley & Williamson & Perini & Wible & Brusatte" authorityYear="2024" box="[571,763,1115,1139]" family="Amilnedwardsidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" rank="family">Amilnedwardsidae</taxonomicName> +and the order +<taxonomicName id="4C107DA0FFF70D0EC606FBAD884EFB33" authorityName="Soria" authorityYear="1989" box="[260,380,1147,1170]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="order">Notopterna</taxonomicName> +section for more details). Key differences between notonychopids with indaleciids are the absence of enamel fossettes in the P2–M3 and a greater development of the parastyle in the former, among other features (Soria 1989b). +<bibRefCitation id="EF817BD2FFF70D0EC773FB2E88B9FAB1" author="McKenna and Bell" box="[113,395,1272,1296]" firstAuthor="McKenna" pageId="17" pageNumber="18" refId="ref44508" refString="McKenna MC, Bell SK. Classification of Mammals Above the Species Level. New York, NY: Columbia University Press, 1997." type="book" year="1997">McKenna and Bell (1997)</bibRefCitation> +later classified +<taxonomicName id="4C107DA0FFF70D0EC532FB2E8BEEFAB1" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[560,732,1272,1296]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName> +as a family within the order +<taxonomicName id="4C107DA0FFF70D0EC68AFAC188C5FA8E" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[392,503,1303,1327]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="order">Litopterna</taxonomicName> +, albeit without any anatomical justification ( +<tableCitation id="C6923398FFF70D0EC653FAE188ACFAEE" box="[337,414,1335,1359]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="17" pageNumber="18">Table 1</tableCitation> +). +<bibRefCitation id="EF817BD2FFF70D0EC6B4FAE18BC8FAEE" author="Bonaparte and Morales" box="[438,762,1335,1359]" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation> +added to +<taxonomicName id="4C107DA0FFF70D0EC7CDFA808849FACF" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[207,379,1366,1390]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName> +the taxon +<taxonomicName id="4C107DA0FFF70D0EC6E4FA81881BFA2C" authority="Bonaparte and Morales 1997" authorityName="Bonaparte and Morales" authorityYear="1997" class="Mammalia" family="Notonychopidae" genus="Requisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="vidmari"> +<emphasis id="B964DA31FFF70D0EC6E4FA818BB8FACF" box="[486,650,1366,1390]" italics="true" pageId="17" pageNumber="18">Requisia vidmari</emphasis> +<bibRefCitation id="EF817BD2FFF70D0EC592FA80881BFA2C" author="Bonaparte and Morales" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales 1997</bibRefCitation> +</taxonomicName> +from Punta Peligro (Peligran SALMA) which they found to be phylogenetically close to +<taxonomicName id="4C107DA0FFF70D0EC527FA438BF6FA0C" authorityName="Simpson" authorityYear="1935" box="[549,708,1429,1453]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="tshotshe"> +<emphasis id="B964DA31FFF70D0EC527FA438BF6FA0C" box="[549,708,1429,1453]" italics="true" pageId="17" pageNumber="18">Wainka tshotshe</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF70D0EC5D6FA4389A3FA6D" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="17" pageNumber="18">Fig. 1D</figureCitation> +). However, since this analysis, +<bibRefCitation id="EF817BD2FFF70D0EC6D2FA628BBBFA6D" author="Goin" box="[464,649,1460,1484]" etAl="et al." firstAuthor="Goin" pageId="17" pageNumber="18" pagination="1029 - 53" refId="ref42920" refString="Goin FJ, Gelfo JN, Ortiz-Jaureguizar E, et al. Mamiferos del Banco Negro Inferior, Formacion Salamanca, Cuenca del Golfo de San Jorge. In: Giacosa RE (ed.), Geologia y Recursos Naturales de la Provincia de Chubut. Puerto Madryn, Relatorio del XXI Congreso Geologico Argentino, 2022, 1029 - 53." type="book chapter" year="2022"> +Goin +<emphasis id="B964DA31FFF70D0EC508FA638B08FA6D" box="[522,570,1460,1484]" italics="true" pageId="17" pageNumber="18">et al.</emphasis> +(2022) +</bibRefCitation> +called into question an isolated m3 (UNP 946) previously assigned to +<taxonomicName id="4C107DA0FFF70D0EC773FA25882BF9AA" authorityName="Bonaparte and Morales" authorityYear="1997" box="[113,281,1523,1547]" class="Mammalia" family="Notonychopidae" genus="Requisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="vidmari"> +<emphasis id="B964DA31FFF70D0EC773FA25882BF9AA" box="[113,281,1523,1547]" italics="true" pageId="17" pageNumber="18">Requisia vidmari</emphasis> +</taxonomicName> +by +<bibRefCitation id="EF817BD2FFF70D0EC643FA258BB4F9AA" author="Bonaparte and Morales" box="[321,646,1523,1547]" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation> +because of its bunodont features and how low and different its cristids are to what would be expected for a notonychopid, suggesting it could be a member of +<taxonomicName id="4C107DA0FFF70D0EC4CCFF198D46FF46" box="[974,1140,207,231]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +. +</paragraph> +<paragraph id="8BAF0623FFF70D0EC447FF388D79FDFE" blockId="17.[809,1461,144,795]" pageId="17" pageNumber="18"> +<bibRefCitation id="EF817BD2FFF70D0EC447FF388DA0FEA7" author="Bonaparte and Morales" box="[837,1170,238,262]" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation> +failed to find a monophyletic +<taxonomicName id="4C107DA0FFF70D0EC462FEDB8D3CFE84" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[864,1038,269,293]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName> +, as +<taxonomicName id="4C107DA0FFF70D0EC339FED88D93FE84" authorityName="Soria" authorityYear="1989" box="[1083,1185,269,293]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="powelli"> +<emphasis id="B964DA31FFF70D0EC339FED88D93FE84" box="[1083,1185,269,293]" italics="true" pageId="17" pageNumber="18">N. powelli</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFF70D0EC3DFFED88C78FE84" authorityName="Bonaparte and Morales" authorityYear="1997" box="[1245,1354,269,293]" class="Mammalia" family="Notonychopidae" genus="Requisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="vidmari"> +<emphasis id="B964DA31FFF70D0EC3DFFED88C78FE84" box="[1245,1354,269,293]" italics="true" pageId="17" pageNumber="18">R. vidmari</emphasis> +</taxonomicName> +were not as closely related in their phylogeny ( +<figureCitation id="132B1AA6FFF70D0EC3B7FEFB8C31FEE4" box="[1205,1283,301,325]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="17" pageNumber="18">Fig. 1D</figureCitation> +). In addition, as notonychopids were found nested within +<taxonomicName id="4C107DA0FFF70D0EC3DCFE9A8C7FFEC5" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[1246,1357,332,356]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="order">Litopterna</taxonomicName> +, most authors now consider them litopterns (e.g., +<bibRefCitation id="EF817BD2FFF70D0EC3C9FEBD8C43FE22" author="Gelfo" box="[1227,1393,363,387]" etAl="et al." firstAuthor="Gelfo" pageId="17" pageNumber="18" pagination="285 - 91" refId="ref42550" refString="Gelfo JN, Lopez GM, Lorente M. Los ungulados arcaicos de America del Sur: ' Condylarthra' y Litopterna. Contribuciones Cientificas Del Museo Argentino de Ciencias Naturales ' Bernardino Rivadavia' 2016; 6: 285 - 91." type="journal article" year="2016"> +Gelfo +<emphasis id="B964DA31FFF70D0EC208FEBA8C0BFE22" box="[1290,1337,363,387]" italics="true" pageId="17" pageNumber="18">et al.</emphasis> +2016 +</bibRefCitation> +, Croft +<emphasis id="B964DA31FFF70D0EC428FE5D8A67FE02" box="[810,853,395,419]" italics="true" pageId="17" pageNumber="18">et al</emphasis> +. 2020, +<bibRefCitation id="EF817BD2FFF70D0EC4A7FE5D8D62FE02" author="Goin" box="[933,1104,395,419]" etAl="et al." firstAuthor="Goin" pageId="17" pageNumber="18" pagination="1029 - 53" refId="ref42920" refString="Goin FJ, Gelfo JN, Ortiz-Jaureguizar E, et al. Mamiferos del Banco Negro Inferior, Formacion Salamanca, Cuenca del Golfo de San Jorge. In: Giacosa RE (ed.), Geologia y Recursos Naturales de la Provincia de Chubut. Puerto Madryn, Relatorio del XXI Congreso Geologico Argentino, 2022, 1029 - 53." type="book chapter" year="2022"> +Goin +<emphasis id="B964DA31FFF70D0EC4E1FE5D8D3CFE02" box="[995,1038,395,419]" italics="true" pageId="17" pageNumber="18">et al</emphasis> +. 2022 +</bibRefCitation> +), which is problematic given that +<bibRefCitation id="EF817BD2FFF70D0EC428FE7C8D5CFE63" author="Bonaparte and Morales" box="[810,1134,426,450]" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation> +used a matrix that includes exclusively litopterns and did not test alternative relationships. Since the early phylogenetic study of +<bibRefCitation id="EF817BD2FFF70D0EC3BFFE3F8A46FD81" author="Bonaparte and Morales" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation> +, there has not been any attempt to test the monophyly of +<taxonomicName id="4C107DA0FFF70D0EC445FDF18AC1FD9E" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[839,1011,551,575]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName> +and to resolve its position among litopterns and other SANUs ( +<tableCitation id="C6923398FFF70D0EC4F2FD918D09FDFE" box="[1008,1083,583,607]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="17" pageNumber="18">Table 2</tableCitation> +). +</paragraph> +<paragraph id="8BAF0623FFF70D0EC447FDB08C1AFCBA" blockId="17.[809,1461,144,795]" pageId="17" pageNumber="18"> +As the only two known members of +<taxonomicName id="4C107DA0FFF70D0EC3C1FDB08C5DFDDF" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[1219,1391,614,638]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName> +are +<taxonomicName id="4C107DA0FFF70D0EC29CFDB18A44FD3C" authorityName="Bonaparte and Morales" authorityYear="1997" class="Mammalia" family="Notonychopidae" genus="Requisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="vidmari"> +<emphasis id="B964DA31FFF70D0EC29CFDB18A44FD3C" italics="true" pageId="17" pageNumber="18">R. vidmari</emphasis> +</taxonomicName> +and the potentially pre-Itaboraian SALMA +<taxonomicName id="4C107DA0FFF70D0EC24BFD508C9FFD3C" authorityName="Soria" authorityYear="1989" box="[1353,1453,645,669]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="powelli"> +<emphasis id="B964DA31FFF70D0EC24BFD508C9FFD3C" box="[1353,1453,645,669]" italics="true" pageId="17" pageNumber="18">N. powelli</emphasis> +</taxonomicName> +, the temporal interval of this family is not well constrained with a current range of ~63.8–51.4 Mya, and is likely much narrower in range ( +<figureCitation id="132B1AA6FFF70D0EC476FD358A8CFD5A" box="[884,958,739,763]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="17" pageNumber="18">Fig. 2B</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFF70D0EC3CDFD358C19FD5A" box="[1231,1323,739,763]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="17" pageNumber="18">Table S1</tableCitation> +; Woodburne +<emphasis id="B964DA31FFF70D0EC428FCD58A6BFCBA" box="[810,857,771,795]" italics="true" pageId="17" pageNumber="18">et al.</emphasis> +2014b, +<bibRefCitation id="EF817BD2FFF70D0EC4AAFCD58D69FCBA" author="Krause" box="[936,1115,771,795]" etAl="et al." firstAuthor="Krause" pageId="17" pageNumber="18" pagination="886 - 903" refId="ref43716" refString="Krause JM, Clyde WC, Ibanez-Mejia M et al. New age constraints for Early Paleogene strata of central Patagonia, Argentina: implications for the timing of South American Land Mammal Ages. Geological Society of America Bulletin 2017; 129: 886 - 903. https: // doi. org / 10.1130 / b 31561.1" type="journal article" year="2017"> +Krause +<emphasis id="B964DA31FFF70D0EC4F7FCD58D11FCBA" box="[1013,1059,771,795]" italics="true" pageId="17" pageNumber="18">et al.</emphasis> +2017 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF70D0EC364FCD58C2AFCBA" author="Gelfo" box="[1126,1304,771,795]" etAl="et al." firstAuthor="Gelfo" pageId="17" pageNumber="18" pagination="993 - 1007" refId="ref42730" refString="Gelfo JN, Garcia-Lopez DA, Bergqvist LP. Phylogenetic relationships and palaeobiology of a new xenungulate (Mammalia: Eutheria) from the Palaeogene of Argentina. Journal of Systematic Palaeontology 2020 b; 18: 993 - 1007. https: // doi. org / 10.1080 / 14772019.2020.171 5496" type="journal article" year="2020" yearSuffix="b"> +Gelfo +<emphasis id="B964DA31FFF70D0EC3A7FCD58DE6FCBA" box="[1189,1236,771,795]" italics="true" pageId="17" pageNumber="18">et al.</emphasis> +2020b +</bibRefCitation> +). +</paragraph> +</subSubSection> +<emphasis id="B964DA31FFF70D0EC4C1FCEC8C28FCF4" box="[963,1306,826,853]" italics="true" pageId="17" pageNumber="18"> +<subSubSection id="C30A55A8FFF70D0EC4C1FCEC8D64FCF5" box="[963,1110,826,852]" pageId="17" pageNumber="18" type="nomenclature"> +<paragraph id="8BAF0623FFF70D0EC4C1FCEC8D64FCF5" blockId="17.[809,1461,826,1547]" box="[963,1110,826,852]" pageId="17" pageNumber="18"> +<taxonomicName id="4C107DA0FFF70D0EC4C1FCEC8D64FCF5" box="[963,1110,826,852]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +</paragraph> +</subSubSection> +<subSubSection id="C30A55A8FFF70D0EC359FCED8C28FCF4" box="[1115,1306,826,853]" pageId="17" pageNumber="18" type="description"> +<paragraph id="8BAF0623FFF70D0EC359FCED8C28FCF4" blockId="17.[809,1461,826,1547]" box="[1115,1306,826,853]" pageId="17" pageNumber="18"> +( +<figureCitation id="132B1AA6FFF70D0EC367FCED8DFFFCF5" box="[1125,1229,826,852]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="17" pageNumber="18">Figs 8B–G</figureCitation> +, +<figureCitation id="132B1AA6FFF70D0EC3D4FCEC8C22FCF5" box="[1238,1296,826,852]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="17" pageNumber="18">9B–E</figureCitation> +) +</paragraph> +</subSubSection> +</emphasis> +<subSubSection id="C30A55A8FFF70D0AC428FCB78839FDDF" lastPageId="21" lastPageNumber="22" pageId="17" pageNumber="18" type="multiple"> +<paragraph id="8BAF0623FFF70D0EC428FCB78C81F9AA" blockId="17.[809,1461,826,1547]" pageId="17" pageNumber="18"> +<taxonomicName id="4C107DA0FFF70D0EC428FCB78AFCFCD8" box="[810,974,865,889]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +currently includes 10 genera [or 12 if we include two unnamed genera (Tejedor +<emphasis id="B964DA31FFF70D0EC371FC578D97FC39" box="[1139,1189,896,920]" italics="true" pageId="17" pageNumber="18">et al.</emphasis> +2009)] with an exclusive Palaeogene distribution ( +<figureCitation id="132B1AA6FFF70D0EC328FC498D43FC16" box="[1066,1137,927,951]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="17" pageNumber="18">Fig. 2B</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFF70D0EC278FC498A74FC76" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="17" pageNumber="18">Table S1</tableCitation> +). There has been recent major detailed revision of this family ( +<bibRefCitation id="EF817BD2FFF70D0EC436FC088A95FC57" author="Gelfo" box="[820,935,990,1014]" firstAuthor="Gelfo" pageId="17" pageNumber="18" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo 2006</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF70D0EC4B0FC088AD4FC57" author="Gelfo" box="[946,998,990,1014]" firstAuthor="Gelfo" pageId="17" pageNumber="18" pagination="130 - 42" refId="ref42288" refString="Gelfo JN. The ' condylarth' Didolodontidae from Gran Barranca: history of the bunodont South American mammals until the Eocene- Oligocene transition. In: Madden RH, Carlini AA, Vucetich MG, Kay RF (eds.), The Paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge: Cambridge University Press, 2010, 130 - 42." type="book chapter" year="2010">2010</bibRefCitation> +), so to avoid unnecessary repetition, only the most relevant taxonomic aspects are discussed here, along with the advances since these revisions. The family +<taxonomicName id="4C107DA0FFF70D0EC212FBCB8C86FB94" box="[1296,1460,1053,1077]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +was proposed by Scott (1913) to classify a group of bunodont ‘condylarths’ (see ‘Condylarthra’ section in Supporting information, File S1) endemic to South America with a Palaeogene fossil record consisting mostly of isolated teeth. Initially, +<bibRefCitation id="EF817BD2FFF70D0EC245FB4C8A46FB70" author="Ameghino" firstAuthor="Ameghino" pageId="17" pageNumber="18" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino (1897)</bibRefCitation> +described the first known Eocene didolodontid species +<taxonomicName id="4C107DA0FFF70D0EC45BFB0F8D03FB50" authorityName="Ameghino" authorityYear="1897" box="[857,1073,1241,1265]" class="Mammalia" family="Didolodontidae" genus="Didolodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="multicuspis"> +<emphasis id="B964DA31FFF70D0EC45BFB0F8D03FB50" box="[857,1073,1241,1265]" italics="true" pageId="17" pageNumber="18">Didolodus multicuspis</emphasis> +</taxonomicName> +as a phenacodontid. Osborn (1910) identified additional South American taxa as ‘Condylarthra’ +<emphasis id="B964DA31FFF70D0EC428FACE8A8AFA8E" box="[810,952,1303,1327]" italics="true" pageId="17" pageNumber="18">incertae sedis</emphasis> +such as +<taxonomicName id="4C107DA0FFF70D0EC33BFACE8DE0FA8E" authorityName="Ameghino" authorityYear="1904" box="[1081,1234,1304,1327]" class="Mammalia" family="Didolodontidae" genus="Notoprotogonia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF70D0EC33BFACE8DE0FA8E" box="[1081,1234,1304,1327]" italics="true" pageId="17" pageNumber="18">Notoprotogonia</emphasis> +</taxonomicName> +( +<bibRefCitation id="EF817BD2FFF70D0EC3F3FAC18C81FA8E" author="Ameghino" box="[1265,1459,1303,1327]" firstAuthor="Ameghino" pageId="17" pageNumber="18" pagination="1 - 541" refId="ref39426" refString="Ameghino F. Recherches de morphologic phylogenetique sur les molaires superieures des ongules. Anales del Museo Nacional de Buenos Aires 1904 b; 3: 1 - 541." type="journal article" year="1904" yearSuffix="b">Ameghino 1904b</bibRefCitation> +[later synonymized with +<taxonomicName id="4C107DA0FFF70D0EC33FFAE18CBCFAEE" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[1085,1422,1335,1359]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="17" pageNumber="18" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF70D0EC33FFAE18DE3FAEE" box="[1085,1233,1335,1359]" italics="true" pageId="17" pageNumber="18">Ernestokokenia</emphasis> +<bibRefCitation id="EF817BD2FFF70D0EC3DFFAE18CBCFAEE" author="Ameghino" box="[1245,1422,1335,1359]" firstAuthor="Ameghino" pageId="17" pageNumber="18" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +by Simpson (1948)], +<taxonomicName id="4C107DA0FFF70D0EC4FEFA808DB7FACF" authorityName="Ameghino" authorityYear="1897" box="[1020,1157,1366,1390]" class="Mammalia" family="Proterotheriidae" genus="Lambdaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF70D0EC4FEFA808DB7FACF" box="[1020,1157,1366,1390]" italics="true" pageId="17" pageNumber="18">Lambdaconus</emphasis> +</taxonomicName> +[later considered an early proterotheriid (Soria 2001)], and +<taxonomicName id="4C107DA0FFF70D0EC38DFAA08DCCFA2C" authorityName="Ameghino" authorityYear="1904" box="[1167,1278,1398,1421]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF70D0EC38DFAA08DCCFA2C" box="[1167,1278,1398,1421]" italics="true" pageId="17" pageNumber="18">Proectocion</emphasis> +</taxonomicName> +[later considered an early adianthid (Cifelli 1983a)]. Scott (1913) later grouped these four taxa as part of the family +<taxonomicName id="4C107DA0FFF70D0EC3BEFA628C0FFA6D" box="[1212,1341,1460,1484]" class="Mammalia" family="Didolodidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Didolodidae</taxonomicName> +within the order +<taxonomicName id="4C107DA0FFF70D0EC46BFA058AE4FA4A" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[873,982,1491,1515]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="order">Litopterna</taxonomicName> +( +<tableCitation id="C6923398FFF70D0EC4EAFA058D07FA4A" box="[1000,1077,1491,1515]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="17" pageNumber="18">Table 1</tableCitation> +). Simpson (1934) subsequently corrected it to +<taxonomicName id="4C107DA0FFF70D0EC4AFFA258D60F9AA" box="[941,1106,1523,1547]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +, and grouped them again within +</paragraph> +<paragraph id="8BAF0623FFF70D0EC773F95D89ECF81B" blockId="17.[113,1454,1675,1979]" pageId="17" pageNumber="18"> +P2–M3 of the astrapothere +<taxonomicName id="4C107DA0FFF70D0EC677F95D8B66F903" authorityName="Ameghino" authorityYear="1897" box="[373,596,1675,1699]" class="Mammalia" family="Trigonostylopidae" genus="Trigonostylops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="wortmani"> +<emphasis id="B964DA31FFF70D0EC677F95D8B66F903" box="[373,596,1675,1699]" italics="true" pageId="17" pageNumber="18">Trigonostylops wortmani</emphasis> +</taxonomicName> +(AMNH VP-28700). I, P5–M3 of the xenungulate +<taxonomicName id="4C107DA0FFF70D0EC338F95D8DFEF903" authorityName="Paula Couto" authorityYear="1952" box="[1082,1228,1675,1699]" class="Mammalia" family="Carodniidae" genus="Carodnia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Xenungulata" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="vieirai"> +<emphasis id="B964DA31FFF70D0EC338F95D8DFEF903" box="[1082,1228,1675,1699]" italics="true" pageId="17" pageNumber="18">Carodnia vieirai</emphasis> +</taxonomicName> +[cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of +<taxonomicName id="4C107DA0FFF70D0EC447F9718D2CF91F" authorityName="Ameghino" authorityYear="1901" box="[837,1054,1703,1727]" class="Mammalia" family="Pyrotheriidae" genus="Propyrotherium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="saxeum"> +<emphasis id="B964DA31FFF70D0EC447F9718D2CF91F" box="[837,1054,1703,1727]" italics="true" pageId="17" pageNumber="18">Propyrotherium saxeum</emphasis> +</taxonomicName> +[AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 ( +<typeStatus id="54ABB881FFF70D0EC77AF90989EAF956" box="[120,216,1759,1783]" pageId="17" pageNumber="18" type="lectotype">lectotype</typeStatus> +), left P5 ( +<typeStatus id="54ABB881FFF70D0EC633F90988A2F956" box="[305,400,1759,1783]" pageId="17" pageNumber="18" type="lectotype">lectotype</typeStatus> +; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods. Abbreviations: efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista. Scale bars equal +<quantity id="4CE8ABC6FFF70D0EC7A8F87589EBF81B" box="[170,217,1955,1979]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="17" pageNumber="18" unit="cm" value="1.0">1 cm</quantity> +. +</paragraph> +<caption id="DF6F56ABFFF40D0CC783F8968A28F81B" lastPageId="19" lastPageNumber="20" pageId="18" pageNumber="19" startId="18.[129,194,1856,1880]" targetBox="[177,1425,147,1825]" targetPageId="18" targetType="figure"> +<paragraph id="8BAF0623FFF40D0DC783F8968CBFF831" blockId="18.[129,1464,1856,1964]" pageId="18" pageNumber="19"> +<emphasis id="B964DA31FFF40D0DC783F89689EAF8F9" bold="true" box="[129,216,1856,1880]" pageId="18" pageNumber="19">Figure 7.</emphasis> +Right lower molars of relevant SANUs in occlusal view. A, p1–m3 of the indaleciid +<taxonomicName id="4C107DA0FFF40D0DC4EAF8968DAFF8F9" authorityName="Bond & Vucetich" authorityYear="1983" box="[1000,1181,1856,1880]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="grandensis"> +<emphasis id="B964DA31FFF40D0DC4EAF8968DAFF8F9" box="[1000,1181,1856,1880]" italics="true" pageId="18" pageNumber="19">Indalecia grandensis</emphasis> +</taxonomicName> +[PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid +<taxonomicName id="4C107DA0FFF40D0DC518F88A8BE0F8D5" authorityName="Soria" authorityYear="1989" box="[538,722,1884,1908]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="powelli"> +<emphasis id="B964DA31FFF40D0DC518F88A8BE0F8D5" box="[538,722,1884,1908]" italics="true" pageId="18" pageNumber="19">Notonychops powelli</emphasis> +</taxonomicName> +[PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate +<taxonomicName id="4C107DA0FFF40D0DC3E6F88A8C8AF8D5" box="[1252,1464,1884,1908]" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="praecursor"> +<emphasis id="B964DA31FFF40D0DC3E6F88A8C8AF8D5" box="[1252,1464,1884,1908]" italics="true" pageId="18" pageNumber="19">Simpsonotus praecursor</emphasis> +</taxonomicName> +[MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere +<taxonomicName id="4C107DA0FFF40D0DC42CF8AE8D3CF831" authorityName="Ameghino" authorityYear="1897" box="[814,1038,1912,1936]" class="Mammalia" family="Trigonostylopidae" genus="Trigonostylops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="wortmani"> +<emphasis id="B964DA31FFF40D0DC42CF8AE8D3CF831" box="[814,1038,1912,1936]" italics="true" pageId="18" pageNumber="19">Trigonostylops wortmani</emphasis> +</taxonomicName> +[MNHN.F.CAS188 (mirrored)]. E, p2– +</paragraph> +<paragraph id="8BAF0623FFF40D0DC783F8428CA0F80D" blockId="18.[129,1464,1856,1964]" box="[129,1426,1940,1964]" pageId="18" pageNumber="19"> +m3 of the xenungulate +<taxonomicName id="4C107DA0FFF40D0DC65AF84388C5F80D" authorityName="Villarroel" authorityYear="1987" box="[344,503,1940,1964]" class="Arthoniomycetes" family="Phaeococcomycetaceae" genus="Etayoa" kingdom="Fungi" order="Lichenostigmatales" pageId="18" pageNumber="19" phylum="Ascomycota" rank="species" species="bacatensis"> +<emphasis id="B964DA31FFF40D0DC65AF84388C5F80D" box="[344,503,1940,1964]" italics="true" pageId="18" pageNumber="19">Etayoa bacatensis</emphasis> +</taxonomicName> +[cast of GM 32 (holotype; mirrored)]. F, p1–m3 of +<taxonomicName id="4C107DA0FFF40D0DC4E1F8428DE7F80D" authorityName="Soria & Powell" authorityYear="1981" box="[995,1237,1940,1964]" class="Mammalia" family="Eoastrapostylopidae" genus="Eoastrapostylops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="riolorense"> +<emphasis id="B964DA31FFF40D0DC4E1F8428DE7F80D" box="[995,1237,1940,1964]" italics="true" pageId="18" pageNumber="19">Eoastrapostylops riolorense</emphasis> +</taxonomicName> +[PVL 4216, p1–m2 +</paragraph> +<paragraph id="8BAF0623FFF50D0CC773FF46886CFDDF" blockId="19.[113,764,144,1703]" pageId="19" pageNumber="20"> +‘Condylarthra’. After adding two new species from the Rio Chico Formation [latest Danian–Late Lutetian in age ( +<bibRefCitation id="EF817BD2FFF50D0CC5B1FF6689DCFF46" author="Krause" etAl="et al." firstAuthor="Krause" pageId="19" pageNumber="20" pagination="886 - 903" refId="ref43716" refString="Krause JM, Clyde WC, Ibanez-Mejia M et al. New age constraints for Early Paleogene strata of central Patagonia, Argentina: implications for the timing of South American Land Mammal Ages. Geological Society of America Bulletin 2017; 129: 886 - 903. https: // doi. org / 10.1130 / b 31561.1" type="journal article" year="2017"> +Krause +<emphasis id="B964DA31FFF50D0CC773FF198999FF46" box="[113,171,207,231]" italics="true" pageId="19" pageNumber="20">et al.</emphasis> +2017 +</bibRefCitation> +)] of the genus +<taxonomicName id="4C107DA0FFF50D0CC6B4FF198B78FF46" authorityName="Ameghino" authorityYear="1901" box="[438,586,207,231]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC6B4FF198B78FF46" box="[438,586,207,231]" italics="true" pageId="19" pageNumber="20">Ernestokokenia</emphasis> +</taxonomicName> +( +<taxonomicName id="4C107DA0FFF50D0CC564FF198858FEA7" authority="Simpson, 1935" authorityName="Simpson" authorityYear="1935" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="yirunhor"> +<emphasis id="B964DA31FFF50D0CC564FF1989F4FEA7" italics="true" pageId="19" pageNumber="20">Ernestokokenia yirunhor</emphasis> +Simpson, 1935 +</taxonomicName> +and +<taxonomicName id="4C107DA0FFF50D0CC6A3FF388997FE84" authority="Simpson 1935" authorityName="Simpson" authorityYear="1935" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="chaishoer"> +<emphasis id="B964DA31FFF50D0CC6A3FF388BAAFEA7" box="[417,664,238,262]" italics="true" pageId="19" pageNumber="20">Ernestokokenia chaishoer</emphasis> +Simpson 1935 +</taxonomicName> +), Simpson (1948) revised the whole group, giving a new definition for +<taxonomicName id="4C107DA0FFF50D0CC644FEFB88D8FEE4" box="[326,490,301,325]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +and emending many of +<bibRefCitation id="EF817BD2FFF50D0CC773FE9A8808FEC5" author="Ameghino's" box="[113,314,332,356]" firstAuthor="Ameghino's" pageId="19" pageNumber="20" pagination="1 - 568" refId="ref39456" refString="Ameghino F. Les formations sedimentaires du cretace superieur et du tertiaire de Patagonie avec un parallele entre leurs faunes mammalogiques et celles de l'ancien continent. Anales del Museo Nacional de Buenos Aires 1906; 15: 1 - 568." type="journal article" year="1906">Ameghino’s (1906)</bibRefCitation> +taxonomic classifications of the Patagonian taxa, including the following genera in the family: +<taxonomicName id="4C107DA0FFF50D0CC593FEBD8BC7FE22" authorityName="Ameghino" authorityYear="1897" box="[657,757,363,387]" class="Mammalia" family="Didolodontidae" genus="Didolodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC593FEBD8BC7FE22" box="[657,757,363,387]" italics="true" pageId="19" pageNumber="20">Didolodus</emphasis> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFF50D0CC773FE5D8837FE02" authorityName="Ameghino" authorityYear="1901" box="[113,261,395,419]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC773FE5D8837FE02" box="[113,261,395,419]" italics="true" pageId="19" pageNumber="20">Ernestokokenia</emphasis> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFF50D0CC616FE5D8B6DFE02" authority="Ameghino 1904 b" authorityName="Ameghino" authorityYear="1904" box="[276,607,395,419]" class="Mammalia" family="Didolodontidae" genus="Argyrolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC616FE5D8890FE02" box="[276,418,395,419]" italics="true" pageId="19" pageNumber="20">Argyrolambda</emphasis> +<bibRefCitation id="EF817BD2FFF50D0CC6A8FE5D8B6DFE02" author="Ameghino" box="[426,607,395,419]" firstAuthor="Ameghino" pageId="19" pageNumber="20" pagination="1 - 541" refId="ref39426" refString="Ameghino F. Recherches de morphologic phylogenetique sur les molaires superieures des ongules. Anales del Museo Nacional de Buenos Aires 1904 b; 3: 1 - 541." type="journal article" year="1904" yearSuffix="b">Ameghino 1904b</bibRefCitation> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFF50D0CC56FFE5D8817FE63" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" class="Mammalia" family="Didolodontidae" genus="Paulogervaisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC56FFE5D8BC8FE02" box="[621,762,395,419]" italics="true" pageId="19" pageNumber="20">Paulogervaisia</emphasis> +<bibRefCitation id="EF817BD2FFF50D0CC773FE7C8817FE63" author="Ameghino" box="[113,293,426,450]" firstAuthor="Ameghino" pageId="19" pageNumber="20" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFF50D0CC638FE7D889BFE63" authorityName="Ameghino" authorityYear="1904" box="[314,425,427,450]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC638FE7D889BFE63" box="[314,425,427,450]" italics="true" pageId="19" pageNumber="20">Proectocion</emphasis> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFF50D0CC6BDFE7D8BC7FE63" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[447,757,426,450]" class="Mammalia" family="Didolodontidae" genus="Enneoconus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC6BDFE7D8B03FE63" box="[447,561,427,450]" italics="true" pageId="19" pageNumber="20">Enneoconus</emphasis> +<bibRefCitation id="EF817BD2FFF50D0CC543FE7C8BC7FE63" author="Ameghino" box="[577,757,426,450]" firstAuthor="Ameghino" pageId="19" pageNumber="20" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +, +<emphasis id="B964DA31FFF50D0CC773FE1F8819FE40" box="[113,299,457,481]" italics="true" pageId="19" pageNumber="20">Asmithwoodwarsia</emphasis> +<bibRefCitation id="EF817BD2FFF50D0CC636FE1F88D3FE40" author="Ameghino" box="[308,481,457,481]" firstAuthor="Ameghino" pageId="19" pageNumber="20" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +, and +<taxonomicName id="4C107DA0FFF50D0CC51CFE1F8828FDA0" authority="Ameghino, 1906" authorityName="Ameghino" authorityYear="1906" class="Mammalia" family="Didolodontidae" genus="Archaeohyracotherium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC51CFE1F8BC8FE40" box="[542,762,457,481]" italics="true" pageId="19" pageNumber="20">Archaeohyracotherium</emphasis> +<bibRefCitation id="EF817BD2FFF50D0CC773FE3F8828FDA0" author="Ameghino" box="[113,282,489,513]" firstAuthor="Ameghino" pageId="19" pageNumber="20" pagination="1 - 568" refId="ref39456" refString="Ameghino F. Les formations sedimentaires du cretace superieur et du tertiaire de Patagonie avec un parallele entre leurs faunes mammalogiques et celles de l'ancien continent. Anales del Museo Nacional de Buenos Aires 1906; 15: 1 - 568." type="journal article" year="1906">Ameghino, 1906</bibRefCitation> +</taxonomicName> +. Simpson soon recognized the close similarities between this group and North American ‘condylarths’, including species of +<taxonomicName id="4C107DA0FFF50D0CC7D8FDF188BFFD9E" box="[218,397,551,575]" class="Mammalia" family="Phenacodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Phenacodontidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFF50D0CC6BDFDF18B79FD9E" box="[447,587,551,575]" class="Mammalia" family="Mioclaenidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Mioclaenidae</taxonomicName> +, stating that they were impossible to separate on a purely morphological basis (Simpson 1948, 1980). +</paragraph> +<paragraph id="8BAF0623FFF50D0CC78FFD538D4DFF66" blockId="19.[113,764,144,1703]" lastBlockId="19.[808,1461,144,1703]" pageId="19" pageNumber="20"> +Paula Couto (1952) added two species of +<taxonomicName id="4C107DA0FFF50D0CC560FD538BC8FD3C" authorityName="Ameghino" authorityYear="1901" box="[610,762,645,669]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC560FD538BC8FD3C" box="[610,762,645,669]" italics="true" pageId="19" pageNumber="20">Ernestokokenia</emphasis> +</taxonomicName> +( +<taxonomicName id="4C107DA0FFF50D0CC77FFD738B87FD1D" authority="Paula Couto, 1952" authorityName="Paula Couto" authorityYear="1952" box="[125,693,676,701]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="protocenica"> +<emphasis id="B964DA31FFF50D0CC77FFD738897FD1D" box="[125,421,677,701]" italics="true" pageId="19" pageNumber="20">Ernestokokenia protocenica</emphasis> +Paula Couto, 1952 +</taxonomicName> +and +<taxonomicName id="4C107DA0FFF50D0CC773FD128B51FD7D" authority="Paula Couto, 1952" authorityName="Paula Couto" authorityYear="1952" box="[113,611,708,732]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="parayiruhnor"> +<emphasis id="B964DA31FFF50D0CC773FD1288A4FD7D" box="[113,406,708,732]" italics="true" pageId="19" pageNumber="20">Ernestokokenia parayiruhnor</emphasis> +Paula Couto, 1952 +</taxonomicName> +) and the new genus +<taxonomicName id="4C107DA0FFF50D0CC7BAFD328828FD5A" authorityName="Paula Couto" authorityYear="1952" box="[184,282,740,763]" class="Mammalia" family="Didolodontidae" genus="Lamegoia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC7BAFD328828FD5A" box="[184,282,740,763]" italics="true" pageId="19" pageNumber="20">Lamegoia</emphasis> +</taxonomicName> +from fissure fills in the Itaboraí Formation, +<collectingCountry id="F30746B3FFF50D0CC773FCD5899DFCBA" box="[113,175,771,795]" name="Brazil" pageId="19" pageNumber="20">Brazil</collectingCountry> +, to the family +<taxonomicName id="4C107DA0FFF50D0CC674FCD58B12FCBA" box="[374,544,771,795]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +. +<bibRefCitation id="EF817BD2FFF50D0CC537FCD58BC8FCBB" author="McKenna" box="[565,762,770,794]" firstAuthor="McKenna" pageId="19" pageNumber="20" pagination="736 - 43" refId="ref44469" refString="McKenna MC. Survival of primitive notoungulates and condylarths into the Miocene of Colombia. American Journal of Science 1956; 254: 736 - 43. https: // doi. org / 10.2475 / ajs. 254.12.736" type="journal article" year="1956">McKenna (1956)</bibRefCitation> +added +<taxonomicName id="4C107DA0FFF50D0CC7BFFCF488FFFC9B" authorityName="McKenna" authorityYear="1956" box="[189,461,802,826]" class="Mammalia" family="Didolodontidae" genus="Megadolodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="molariformis"> +<emphasis id="B964DA31FFF50D0CC7BFFCF488FFFC9B" box="[189,461,802,826]" italics="true" pageId="19" pageNumber="20">Megadolodus molariformis</emphasis> +</taxonomicName> +from La Venta, +<collectingCountry id="F30746B3FFF50D0CC58EFCF48BC5FC9B" box="[652,759,802,826]" name="Colombia" pageId="19" pageNumber="20">Colombia</collectingCountry> +, which +<bibRefCitation id="EF817BD2FFF50D0CC7B9FC9788DCFCF8" author="Cifelli and Villarroel" box="[187,494,833,857]" firstAuthor="Cifelli" pageId="19" pageNumber="20" pagination="265 - 88" refId="ref41117" refString="Cifelli RL, Villarroel C. Paleobiology and affinities of Megadolodus. In: Kay RF, Madden RH, Cifelli RL, Flynn JJ (eds), Vertebrate Paleontology in the Neotropics: The Miocene Fauna of La Venta, Colombia. Washington and London: Smithsonian Institution Press, 1997, 265 - 88." type="book chapter" year="1997">Cifelli and Villarroel (1997)</bibRefCitation> +later considered to be a bunodont proterotheriid, a hypothesis that recently has found some support in phylogenetic analyses (Carrillo +<emphasis id="B964DA31FFF50D0CC578FC578B9FFC39" box="[634,685,896,920]" italics="true" pageId="19" pageNumber="20">et al.</emphasis> +2023). Paula Couto (1978) added +<taxonomicName id="4C107DA0FFF50D0CC6B3FC4989C4FC77" authority="Paula Couto, 1952" authorityName="Paula Couto" authorityYear="1952" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="scotti"> +<emphasis id="B964DA31FFF50D0CC6B3FC498B86FC16" box="[433,692,927,951]" italics="true" pageId="19" pageNumber="20">Asmithwoodwardia scotti</emphasis> +Paula Couto, 1952 +</taxonomicName> +from Itaboraí to the family, a taxon that he previously classified within the North American ‘Condylarthra’ family +<taxonomicName id="4C107DA0FFF50D0CC7BDFC2B885DFBB4" box="[191,367,1021,1045]" class="Mammalia" family="Hyopsodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Hyopsodontidae</taxonomicName> +based on dental similarities (Paula Couto 1952). In a revision of the family +<taxonomicName id="4C107DA0FFF50D0CC54FFBCB8826FBF5" authority=", Cifelli (1983 a)" authorityName=", Cifelli" authorityYear="1983" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae, Cifelli (1983a)</taxonomicName> +, based on an indirect tarsal association, removed +<taxonomicName id="4C107DA0FFF50D0CC7C1FB8D88D8FBD2" baseAuthorityName="Paula Couto" baseAuthorityYear="1952" box="[195,490,1115,1139]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="parayirunhor"> +<emphasis id="B964DA31FFF50D0CC7C1FB8D88D8FBD2" box="[195,490,1115,1139]" italics="true" pageId="19" pageNumber="20">Ernestokokenia parayirunhor</emphasis> +</taxonomicName> +(renamed as +<taxonomicName id="4C107DA0FFF50D0CC587FB8D88AEFB32" authority="Cifelli 1983 a" authorityName="Cifelli" authorityYear="1983" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="parayirunhor"> +<emphasis id="B964DA31FFF50D0CC587FB8D89C7FB32" italics="true" pageId="19" pageNumber="20">Miguelsoria parayirunhor</emphasis> +Cifelli 1983a +</taxonomicName> +) and +<taxonomicName id="4C107DA0FFF50D0CC6F1FBAD8BC9FB33" authorityName="Paula Couto" authorityYear="1952" box="[499,763,1147,1171]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="scotti"> +<emphasis id="B964DA31FFF50D0CC6F1FBAD8BC9FB33" box="[499,763,1147,1171]" italics="true" pageId="19" pageNumber="20">Asmithwoodwardia scotti</emphasis> +</taxonomicName> +from +<taxonomicName id="4C107DA0FFF50D0CC7B0FB4C886BFB13" authorityName=", Cifelli" authorityYear="1983" box="[178,345,1178,1202]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +and placed them in +<taxonomicName id="4C107DA0FFF50D0CC550FB4C8BF3FB13" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[594,705,1178,1202]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="19" pageNumber="20" phylum="Chordata" rank="order">Litopterna</taxonomicName> +(see +<taxonomicName id="4C107DA0FFF50D0CC773FB6F882FFB70" box="[113,285,1209,1233]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Protolipternidae</taxonomicName> +section for more information). This tarsal association was based on the fact that the relative size and abundance of these tarsals in the Itaboraí sites was what you would expect for the size and abundance of the dentition of +<taxonomicName id="4C107DA0FFF50D0CC773FAE18B11FAEE" authority="(Cifelli 1983 b)" baseAuthorityName="Cifelli" baseAuthorityYear="1983" box="[113,547,1335,1359]" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="parayirunhor"> +<emphasis id="B964DA31FFF50D0CC773FAE18847FAEE" box="[113,373,1335,1359]" italics="true" pageId="19" pageNumber="20">Miguelsoria parayirunhor</emphasis> +(Cifelli 1983b) +</taxonomicName> +. Cifelli (1983a) reclassified +<taxonomicName id="4C107DA0FFF50D0CC7D7FA8088D1FACF" authorityName="Paula Couto" authorityYear="1952" box="[213,483,1366,1390]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="protocenica"> +<emphasis id="B964DA31FFF50D0CC7D7FA8088D1FACF" box="[213,483,1366,1390]" italics="true" pageId="19" pageNumber="20">Ernestokokenia protocenica</emphasis> +</taxonomicName> +as +<taxonomicName id="4C107DA0FFF50D0CC500FA808BC8FACF" baseAuthorityName="Paula Couto" baseAuthorityYear="1952" box="[514,762,1366,1390]" class="Mammalia" family="Didolodontidae" genus="Paulacoutoia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="protocenica"> +<emphasis id="B964DA31FFF50D0CC500FA808BC8FACF" box="[514,762,1366,1390]" italics="true" pageId="19" pageNumber="20">Paulacoutoia protocenica</emphasis> +</taxonomicName> +[now known as +<taxonomicName id="4C107DA0FFF50D0CC61CFAA38BE2FA2C" authority="(Mones, 2015)" baseAuthorityName="Mones" baseAuthorityYear="2015" box="[286,720,1397,1421]" class="Mammalia" family="Didolodontidae" genus="Ricardocifellia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="protocenica"> +<emphasis id="B964DA31FFF50D0CC61CFAA38B14FA2C" box="[286,550,1397,1421]" italics="true" pageId="19" pageNumber="20">Ricardocifellia protocenica</emphasis> +( +<bibRefCitation id="EF817BD2FFF50D0CC538FAA08BF7FA2C" author="Mones" box="[570,709,1397,1421]" firstAuthor="Mones" pageId="19" pageNumber="20" refId="ref44574" refString="Mones A. Ricardocifellia, a replacement name for Paulacoutoia Cifelli, 1983, and Depaulacoutoia Cifelli and Ortiz-Jaureguizar, 2014 (Mammalia, ' Condylarthra', Didolodontidae), and the status of Depaulacoutoia Kretzoi and Kretzoi, 2000. Journal of Vertebrate Paleontology 2015; 35: e 973571. https: // doi. org / 10.1080 / 02724634. 2015.973571" type="journal volume" year="2015">Mones, 2015</bibRefCitation> +) +</taxonomicName> +because of preoccupation], because he considered it sufficiently morphologically distinct from the genus +<taxonomicName id="4C107DA0FFF50D0CC55EFA628BC6FA6D" authorityName="Ameghino" authorityYear="1901" box="[604,756,1460,1484]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC55EFA628BC6FA6D" box="[604,756,1460,1484]" italics="true" pageId="19" pageNumber="20">Ernestokokenia</emphasis> +</taxonomicName> +. According to Cifelli (1983a), didolodontids are characterized by having a primitive dentition very similar to mioclaenids, but possessing a hypocone on the M3 (absent in mioclaenids) and a primitive ankle structure similar to many South and North American ‘condylarths’ (i.e., the astragalar facet for the medial malleolus is enlarged anteriorly and the calcaneal cuboid facet is dorsally expanded). Soria (2001), based on dental features, removed +<taxonomicName id="4C107DA0FFF50D0CC496FF468DB6FF09" authority="Berg, 1899" authorityName="Berg" authorityYear="1899" box="[916,1156,144,168]" class="Mammalia" family="Proterotheriidae" genus="Xesmodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC496FF468AC9FF09" box="[916,1019,144,168]" italics="true" pageId="19" pageNumber="20">Xesmodon</emphasis> +<bibRefCitation id="EF817BD2FFF50D0CC30BFF468DB6FF09" author="Berg" box="[1033,1156,144,168]" firstAuthor="Berg" pageId="19" pageNumber="20" pagination="77 - 80" refId="ref39765" refString="Berg C. Substitucion de nombres genericos. III. Comunicaciones del Museo Nacional de Buenos Aires 1899; 1: 77 - 80." type="journal article" year="1899">Berg, 1899</bibRefCitation> +</taxonomicName> +from +<taxonomicName id="4C107DA0FFF50D0CC3D1FF468C4CFF09" authorityName=", Soria" authorityYear="2001" box="[1235,1406,144,168]" family="Anisolambdidae" pageId="19" pageNumber="20" rank="family">Anisolambdidae</taxonomicName> +and placed it within +<taxonomicName id="4C107DA0FFF50D0CC4D3FF798D49FF66" authorityName=", Cifelli" authorityYear="1983" box="[977,1147,175,199]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +. +</paragraph> +<paragraph id="8BAF0623FFF50D0CC447FF198C17FD3C" blockId="19.[808,1461,144,1703]" pageId="19" pageNumber="20"> +In a detailed revision of +<taxonomicName id="4C107DA0FFF50D0CC3B1FF198A47FEA7" authority=", Gelfo (2006)" authorityName=", Gelfo" authorityYear="2006" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family"> +Didolodontidae, +<bibRefCitation id="EF817BD2FFF50D0CC27BFF198A47FEA7" author="Gelfo" firstAuthor="Gelfo" pageId="19" pageNumber="20" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo (2006)</bibRefCitation> +</taxonomicName> +synonymized +<taxonomicName id="4C107DA0FFF50D0CC347FF388DE1FEA7" authorityName="Ameghino" authorityYear="1904" box="[1093,1235,238,262]" class="Mammalia" family="Didolodontidae" genus="Argyrolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC347FF388DE1FEA7" box="[1093,1235,238,262]" italics="true" pageId="19" pageNumber="20">Argyrolambda</emphasis> +</taxonomicName> +with +<taxonomicName id="4C107DA0FFF50D0CC24BFF388C9FFEA7" authorityName="Ameghino" authorityYear="1897" box="[1353,1453,238,262]" class="Mammalia" family="Didolodontidae" genus="Didolodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC24BFF388C9FFEA7" box="[1353,1453,238,262]" italics="true" pageId="19" pageNumber="20">Didolodus</emphasis> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFF50D0CC428FEDB8D34FE84" authorityName="Ameghino" authorityYear="1906" box="[810,1030,269,293]" class="Mammalia" family="Didolodontidae" genus="Archaeohyracotherium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC428FEDB8D34FE84" box="[810,1030,269,293]" italics="true" pageId="19" pageNumber="20">Archaeohyracotherium</emphasis> +</taxonomicName> +with +<taxonomicName id="4C107DA0FFF50D0CC344FEDB8C36FE84" authorityName="Ameghino" authorityYear="1901" box="[1094,1284,269,293]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC344FEDB8C36FE84" box="[1094,1284,269,293]" italics="true" pageId="19" pageNumber="20">Asmithwoodwardia</emphasis> +</taxonomicName> +, and +<taxonomicName id="4C107DA0FFF50D0CC243FED88C81FE84" authorityName="Ameghino" authorityYear="1901" box="[1345,1459,270,293]" class="Mammalia" family="Didolodontidae" genus="Enneoconus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC243FED88C81FE84" box="[1345,1459,270,293]" italics="true" pageId="19" pageNumber="20">Enneoconus</emphasis> +</taxonomicName> +with +<taxonomicName id="4C107DA0FFF50D0CC461FEFB8AC5FEE4" authorityName="Ameghino" authorityYear="1901" box="[867,1015,301,325]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC461FEFB8AC5FEE4" box="[867,1015,301,325]" italics="true" pageId="19" pageNumber="20">Ernestokokenia</emphasis> +</taxonomicName> +, thus reducing the total number of valid didolodontid genera. Later, +<bibRefCitation id="EF817BD2FFF50D0CC344FE9A8DE2FEC5" author="Gelfo" box="[1094,1232,332,356]" firstAuthor="Gelfo" pageId="19" pageNumber="20" pagination="651 - 60" refId="ref42229" refString="Gelfo JN. The ' Condylarth' Raulvaccia peligrensis (Mammalia: Didolodontidae) from the Paleocene of Patagonia, Argentina. Journal of Vertebrate Paleontology 2007; 27: 651 - 60. https: // doi. org / 10.1671 / 0272 - 4634 (2007) 27 (651: tcrpmd) 2.0. co; 2" type="journal article" year="2007">Gelfo (2007)</bibRefCitation> +, based on dental similarities and a phylogenetic analysis, added +<taxonomicName id="4C107DA0FFF50D0CC3DAFEBD8D26FE02" authority="Bonaparte et al., 1993" authorityName="Bonaparte" authorityYear="1993" class="Mammalia" family="Mioclaenidae" genus="Escribania" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="chubutensis"> +<emphasis id="B964DA31FFF50D0CC3DAFEBD8C81FE22" box="[1240,1459,363,387]" italics="true" pageId="19" pageNumber="20">Escribania chubutensis</emphasis> +<bibRefCitation id="EF817BD2FFF50D0CC428FE5D8D26FE02" author="Bonaparte" box="[810,1044,395,419]" etAl="et al." firstAuthor="Bonaparte" pageId="19" pageNumber="20" pagination="3 - 61" refId="ref40125" refString="Bonaparte JF, Van Valen L, Kramarz AG. La fauna local de Punta Peligro, Paleoceno inferior, de la provincia del Chubut, Patagonia, Argentina. Evolutionary Monographs 1993; 14: 3 - 61." type="journal article" year="1993"> +Bonaparte +<emphasis id="B964DA31FFF50D0CC49CFE5D8AE3FE02" box="[926,977,395,419]" italics="true" pageId="19" pageNumber="20">et al.</emphasis> +, 1993 +</bibRefCitation> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFF50D0CC34FFE5D8AB0FE63" authority="Bonaparte et al. 1993" authorityName="Bonaparte" authorityYear="1993" class="Mammalia" family="Mioclaenidae" genus="Raulvaccia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="peligrensis"> +<emphasis id="B964DA31FFF50D0CC34FFE5D8C16FE02" box="[1101,1316,395,419]" italics="true" pageId="19" pageNumber="20">Raulvaccia peligrensis</emphasis> +<bibRefCitation id="EF817BD2FFF50D0CC22CFE5D8AB0FE63" author="Bonaparte" etAl="et al." firstAuthor="Bonaparte" pageId="19" pageNumber="20" pagination="3 - 61" refId="ref40125" refString="Bonaparte JF, Van Valen L, Kramarz AG. La fauna local de Punta Peligro, Paleoceno inferior, de la provincia del Chubut, Patagonia, Argentina. Evolutionary Monographs 1993; 14: 3 - 61." type="journal article" year="1993"> +Bonaparte +<emphasis id="B964DA31FFF50D0CC2A0FE5D8A71FE63" italics="true" pageId="19" pageNumber="20">et al.</emphasis> +1993 +</bibRefCitation> +</taxonomicName> +from Punta Peligro, +<collectingCountry id="F30746B3FFF50D0CC372FE7C8DEAFE63" box="[1136,1240,426,450]" name="Argentina" pageId="19" pageNumber="20">Argentina</collectingCountry> +, to +<taxonomicName id="4C107DA0FFF50D0CC208FE7C8C82FE63" authorityName=", Gelfo" authorityYear="2006" box="[1290,1456,426,450]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +. +<taxonomicName id="4C107DA0FFF50D0CC428FE1F8D3CFE40" box="[810,1038,457,481]" class="Mammalia" family="Mioclaenidae" genus="Escribania" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="chubutensis"> +<emphasis id="B964DA31FFF50D0CC428FE1F8D3CFE40" box="[810,1038,457,481]" italics="true" pageId="19" pageNumber="20">Escribania chubutensis</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFF50D0CC352FE1F8C19FE40" box="[1104,1323,457,481]" class="Mammalia" family="Mioclaenidae" genus="Raulvaccia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="peligrensis"> +<emphasis id="B964DA31FFF50D0CC352FE1F8C19FE40" box="[1104,1323,457,481]" italics="true" pageId="19" pageNumber="20">Raulvaccia peligrensis</emphasis> +</taxonomicName> +were previously classified within the ‘condylarth’ North American family +<taxonomicName id="4C107DA0FFF50D0CC428FDDE8D9CFD81" authority="(Bonaparte et al. 1993)" baseAuthorityName="Bonaparte" baseAuthorityYear="1993" box="[810,1198,520,544]" class="Mammalia" family="Mioclaenidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family"> +Mioclaenidae ( +<bibRefCitation id="EF817BD2FFF50D0CC4C4FDDE8D90FD81" author="Bonaparte" box="[966,1186,520,544]" etAl="et al." firstAuthor="Bonaparte" pageId="19" pageNumber="20" pagination="3 - 61" refId="ref40125" refString="Bonaparte JF, Van Valen L, Kramarz AG. La fauna local de Punta Peligro, Paleoceno inferior, de la provincia del Chubut, Patagonia, Argentina. Evolutionary Monographs 1993; 14: 3 - 61." type="journal article" year="1993"> +Bonaparte +<emphasis id="B964DA31FFF50D0CC335FDDF8D5AFD81" box="[1079,1128,520,544]" italics="true" pageId="19" pageNumber="20">et al.</emphasis> +1993 +</bibRefCitation> +) +</taxonomicName> +. The most recent generic additions to +<taxonomicName id="4C107DA0FFF50D0CC4A9FDF18D7DFD9E" authorityName=", Gelfo" authorityYear="2006" box="[939,1103,551,575]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +have been +<taxonomicName id="4C107DA0FFF50D0CC3BCFDF18A6CFDFE" authority="Gelfo & Sige, 2011" authorityName="Gelfo & Sige" authorityYear="2011" class="Mammalia" family="Didolodontidae" genus="Umayodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC3BCFDF18C14FD9E" box="[1214,1318,551,575]" italics="true" pageId="19" pageNumber="20">Umayodus</emphasis> +<bibRefCitation id="EF817BD2FFF50D0CC229FDF18A6CFDFE" author="Gelfo & Sige" firstAuthor="Gelfo" pageId="19" pageNumber="20" pagination="665 - 78" refId="ref42458" refString="Gelfo JN, Sige B. A new didolodontid mammal from the Late Paleocene - earliest Eocene of Laguna Umayo, Peru. Acta Palaeontologica Polonica 2011; 56: 665 - 78. https: // doi. org / 10.4202 / app. 2010.0067" type="journal article" year="2011">Gelfo & Sigé, 2011</bibRefCitation> +</taxonomicName> +from the Lower Muñani Formation, +<collectingCountry id="F30746B3FFF50D0CC3E5FD918C2BFDFF" box="[1255,1305,583,606]" name="Peru" pageId="19" pageNumber="20">Peru</collectingCountry> +, and +<taxonomicName id="4C107DA0FFF50D0CC256FD918AD2FDDF" authority="Gelfo et al., 2020 a" authorityName="Gelfo" authorityYear="2020" class="Mammalia" family="Didolodontidae" genus="Saltaodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC256FD918C81FDFE" box="[1364,1459,583,607]" italics="true" pageId="19" pageNumber="20">Saltaodus</emphasis> +<bibRefCitation id="EF817BD2FFF50D0CC428FDB08AD2FDDF" author="Gelfo" box="[810,992,614,638]" etAl="et al." firstAuthor="Gelfo" pageId="19" pageNumber="20" pagination="132 - 45" refId="ref42677" refString="Gelfo JN, Alonso RN, Madden RH et al. An Eocene bunodont South American native ungulate (Didolodontidae) from the Lumbrera Formation, Salta Province, Argentina. Ameghiniana 2020 a; 57: 132 - 45. https: // doi. org / 10.5710 / amgh. 29.11.2019.3293" type="journal article" year="2020" yearSuffix="a"> +Gelfo +<emphasis id="B964DA31FFF50D0CC46BFDB18AA5FDDF" box="[873,919,614,638]" italics="true" pageId="19" pageNumber="20">et al.</emphasis> +, 2020a +</bibRefCitation> +</taxonomicName> +from the Lumbrera Formation, both of uncertain age ( +<bibRefCitation id="EF817BD2FFF50D0CC48AFD538D64FD3C" author="Gelfo and Sige" box="[904,1110,645,669]" firstAuthor="Gelfo" pageId="19" pageNumber="20" pagination="665 - 78" refId="ref42458" refString="Gelfo JN, Sige B. A new didolodontid mammal from the Late Paleocene - earliest Eocene of Laguna Umayo, Peru. Acta Palaeontologica Polonica 2011; 56: 665 - 78. https: // doi. org / 10.4202 / app. 2010.0067" type="journal article" year="2011">Gelfo and Sigé 2011</bibRefCitation> +, Zimicz +<emphasis id="B964DA31FFF50D0CC3AFFD508DEEFD3C" box="[1197,1244,645,669]" italics="true" pageId="19" pageNumber="20">et al.</emphasis> +2023). +</paragraph> +<paragraph id="8BAF0623FFF50D0CC447FD738AF4FB13" blockId="19.[808,1461,144,1703]" pageId="19" pageNumber="20"> +In terms of the phylogenetic affinities of +<taxonomicName id="4C107DA0FFF50D0CC208FD738C9DFD1C" authorityName=", Gelfo" authorityYear="2006" box="[1290,1455,677,701]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +, early phylogenetic analyses found a close relationship between didolodontids and sparnotheriodontids ( +<bibRefCitation id="EF817BD2FFF50D0CC3CDFD358C75FD5A" author="Cifelli" box="[1231,1351,739,763]" firstAuthor="Cifelli" pageId="19" pageNumber="20" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF50D0CC250FD358A6CFCBA" author="Bergqvist" firstAuthor="Bergqvist" pageId="19" pageNumber="20" refId="ref39790" refString="Bergqvist LP. Reasociacao de pos-cranio as especies de ungulados da Bacia de S. J. Itaborai (Paleoceno), Estado do Rio de Janeiro, e Filogenia dos ' Condylarthra', e ungulados Sul-Americanos com base no poscranio. D. Phil. Thesis, Universidade Federal do Rio Grande do Sul, Brazil, 1996." type="book" year="1996">Bergqvist 1996</bibRefCitation> +; +<figureCitation id="132B1AA6FFF50D0CC46EFCD58A85FCBA" box="[876,951,771,795]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="19" pageNumber="20">Fig. 1C</figureCitation> +), but these were based on tarsal associations that have been questioned by later authors (e.g.,Soria 2001, +<bibRefCitation id="EF817BD2FFF50D0CC24DFCF48ABCFCF8" author="Gelfo and Sige" firstAuthor="Gelfo" pageId="19" pageNumber="20" pagination="665 - 78" refId="ref42458" refString="Gelfo JN, Sige B. A new didolodontid mammal from the Late Paleocene - earliest Eocene of Laguna Umayo, Peru. Acta Palaeontologica Polonica 2011; 56: 665 - 78. https: // doi. org / 10.4202 / app. 2010.0067" type="journal article" year="2011">Gelfo and Sigé 2011</bibRefCitation> +; see also +<taxonomicName id="4C107DA0FFF50D0CC4F7FC978DE5FCF8" box="[1013,1239,833,857]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFF50D0CC209FC978C81FCF8" box="[1291,1459,833,857]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Protolipternidae</taxonomicName> +sections). Muizon and Cifelli (2000), in a phylogenetic analysis that included North American mioclaenids, South American kollpaniines, didolodontids, and protolipternids, found that protolipternids were nested within +<taxonomicName id="4C107DA0FFF50D0CC3ABFC698C7FFC76" authorityName=", Gelfo" authorityYear="2006" box="[1193,1357,959,983]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +and that both groups together formed the sister group of kollpanines, a result that was also found in a later study ( +<bibRefCitation id="EF817BD2FFF50D0CC3F8FC2B8C5DFBB4" author="Gelfo" box="[1274,1391,1021,1045]" firstAuthor="Gelfo" pageId="19" pageNumber="20" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo 2006</bibRefCitation> +). This means that +<taxonomicName id="4C107DA0FFF50D0CC4A2FBCB8D76FB94" authorityName=", Gelfo" authorityYear="2006" box="[928,1092,1053,1077]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +excluding +<taxonomicName id="4C107DA0FFF50D0CC3B2FBCB8C6AFB94" box="[1200,1368,1053,1077]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Protolipternidae</taxonomicName> +is a paraphyletic group. Other phylogenetic studies that have included didolodontids in very small matrices have focused on their internal relationships, assuming their monophyly (e.g., +<bibRefCitation id="EF817BD2FFF50D0CC27BFBAD8A6CFB13" author="Gelfo" firstAuthor="Gelfo" pageId="19" pageNumber="20" pagination="130 - 42" refId="ref42288" refString="Gelfo JN. The ' condylarth' Didolodontidae from Gran Barranca: history of the bunodont South American mammals until the Eocene- Oligocene transition. In: Madden RH, Carlini AA, Vucetich MG, Kay RF (eds.), The Paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge: Cambridge University Press, 2010, 130 - 42." type="book chapter" year="2010">Gelfo 2010</bibRefCitation> +; +<tableCitation id="C6923398FFF50D0CC469FB4C8A87FB13" box="[875,949,1178,1202]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="19" pageNumber="20">Table 2</tableCitation> +). +</paragraph> +<paragraph id="8BAF0623FFF50D0CC447FB6F8C81F906" blockId="19.[808,1461,144,1703]" pageId="19" pageNumber="20"> +When didolodontids have been included with a limited taxon sample (i.e., usually +<taxonomicName id="4C107DA0FFF50D0CC4F8FB0F8D6FFB50" authorityName="Ameghino" authorityYear="1897" box="[1018,1117,1241,1265]" class="Mammalia" family="Didolodontidae" genus="Didolodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF50D0CC4F8FB0F8D6FFB50" box="[1018,1117,1241,1265]" italics="true" pageId="19" pageNumber="20">Didolodus</emphasis> +</taxonomicName> +) in broader analyses that include other SANUs, they have been found in a branch at the stem of some North American ‘condylarths’ and other SANUs (e.g., +<bibRefCitation id="EF817BD2FFF50D0CC428FAE18AE3FAEF" author="Billet" box="[810,977,1334,1359]" etAl="et al." firstAuthor="Billet" pageId="19" pageNumber="20" pagination="956 - 87" refId="ref39984" refString="Billet G, de Muizon C, Schellhorn R et al. Petrosal and inner ear anatomy and allometry amongst specimens referred to Litopterna (Placentalia). Zoological Journal of the Linnean Society 2015; 173: 956 - 87. https: // doi. org / 10.1111 / zoj. 12219" type="journal article" year="2015"> +Billet +<emphasis id="B964DA31FFF50D0CC465FAE18AA5FAEE" box="[871,919,1335,1359]" italics="true" pageId="19" pageNumber="20">et al.</emphasis> +2015 +</bibRefCitation> +), or in a basal position in Laurasiatheria as the sister group of +<taxonomicName id="4C107DA0FFF50D0CC4CDFA808C57FACF" authority="(O'Leary et al. 2013)" baseAuthorityName="O'Leary" baseAuthorityYear="2013" box="[975,1381,1366,1390]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="family"> +Protolipternidae (O’Leary +<emphasis id="B964DA31FFF50D0CC3EBFA818C2EFACF" box="[1257,1308,1366,1390]" italics="true" pageId="19" pageNumber="20">et al.</emphasis> +2013) +</taxonomicName> +. When a larger sample of didolodontids has been included, they have been found at the stem of litopterns but not forming a monophyletic +<taxonomicName id="4C107DA0FFF50D0CC480FA628C35FA6D" authority="(Kramarz et al. 2021)" baseAuthorityName="Kramarz" baseAuthorityYear="2021" box="[898,1287,1460,1484]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family"> +Didolodontidae ( +<bibRefCitation id="EF817BD2FFF50D0CC334FA628DCEFA6D" author="Kramarz" box="[1078,1276,1460,1484]" etAl="et al." firstAuthor="Kramarz" pageId="19" pageNumber="20" refId="ref43620" refString="Kramarz AG, Bond M, MacPhee RDE. On the alleged perissodactyl affinities of the ' condylarth' Escribania chubutensis and other endemic South American ungulate-like placentals. Journal of Vertebrate Paleontology 2021; 41: e 1986716." type="journal volume" year="2021"> +Kramarz +<emphasis id="B964DA31FFF50D0CC396FA638DF1FA6D" box="[1172,1219,1460,1484]" italics="true" pageId="19" pageNumber="20">et al.</emphasis> +2021 +</bibRefCitation> +) +</taxonomicName> +, or as a paraphyletic group among SANUs and North American ‘condylarths’ (Zimicz +<emphasis id="B964DA31FFF50D0CC481FA258A86F9AA" box="[899,948,1523,1547]" italics="true" pageId="19" pageNumber="20">et al.</emphasis> +2023). In sum, there is still a considerable uncertainty in the phylogenetic position of didolodontids and their relationship among SANUs, and whether they form a monophyletic group. In fact, there has not been any phylogenetic analysis that includes didolodontids alongside representatives of all litoptern families, protolipternids, and the different SANUs’ +</paragraph> +<paragraph id="8BAF0623FFF50D0CC771F92C8A28F81B" blockId="19.[113,1452,1786,1979]" pageId="19" pageNumber="20"> +( +<typeStatus id="54ABB881FFF50D0CC778F92C89E4F8B3" box="[122,214,1786,1810]" pageId="19" pageNumber="20" type="holotype">holotype</typeStatus> +; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 of +<taxonomicName id="4C107DA0FFF50D0CC5C1F92C8AAEF8B3" authorityName="Ameghino" authorityYear="1901" box="[707,924,1786,1810]" class="Mammalia" family="Pyrotheriidae" genus="Propyrotherium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="saxeum"> +<emphasis id="B964DA31FFF50D0CC5C1F92C8AAEF8B3" box="[707,924,1786,1810]" italics="true" pageId="19" pageNumber="20">Propyrotherium saxeum</emphasis> +</taxonomicName> +[LIEB-PV 3200 (mirrored)]. Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H +<quantity id="4CE8ABC6FFF50D0CC5FDF8E58A11F8EB" box="[767,803,1842,1867]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="19" pageNumber="20" unit="in" value="2.0">2 in</quantity> +<taxonomicName id="4C107DA0FFF50D0CC42AF8E58ACEF8EB" box="[808,1020,1843,1866]" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="praecursor"> +<emphasis id="B964DA31FFF50D0CC42AF8E58ACEF8EB" box="[808,1020,1843,1866]" italics="true" pageId="19" pageNumber="20">Simpsonotus praecursor</emphasis> +</taxonomicName> +, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of +<taxonomicName id="4C107DA0FFF50D0CC504F8998BE8F8C7" box="[518,730,1871,1894]" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="praecursor"> +<emphasis id="B964DA31FFF50D0CC504F8998BE8F8C7" box="[518,730,1871,1894]" italics="true" pageId="19" pageNumber="20">Simpsonotus praecursor</emphasis> +</taxonomicName> +is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid. For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal +<quantity id="4CE8ABC6FFF50D0CC5E4F8758A24F81B" box="[742,790,1955,1979]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="19" pageNumber="20" unit="cm" value="1.0">1 cm</quantity> +. +</paragraph> +</caption> +<caption id="DF6F56ABFFF20D0AC781F9238B56F81B" lastPageId="21" lastPageNumber="22" pageId="20" pageNumber="21" startId="20.[131,196,1781,1805]" targetBox="[132,1475,150,1750]" targetPageId="20" targetType="figure"> +<paragraph id="8BAF0623FFF20D0BC781F9238CA0F8E4" blockId="20.[131,1428,1781,1973]" pageId="20" pageNumber="21"> +<emphasis id="B964DA31FFF20D0BC781F92389E8F8AC" bold="true" box="[131,218,1781,1805]" pageId="20" pageNumber="21">Figure 8.</emphasis> +Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of +<taxonomicName id="4C107DA0FFF20D0BC30FF9238D83F8AC" authorityName="Muizon and Marshall" authorityYear="1987" box="[1037,1201,1781,1805]" class="Mammalia" family="Mioclaenidae" genus="Molinodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="suarezi"> +<emphasis id="B964DA31FFF20D0BC30FF9238D83F8AC" box="[1037,1201,1781,1805]" italics="true" pageId="20" pageNumber="21">Molinodus suarezi</emphasis> +</taxonomicName> +in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of +<taxonomicName id="4C107DA0FFF20D0BC4DEF8C78D95F888" box="[988,1191,1809,1833]" class="Mammalia" family="Mioclaenidae" genus="Escribania" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="chubutensis"> +<emphasis id="B964DA31FFF20D0BC4DEF8C78D95F888" box="[988,1191,1809,1833]" italics="true" pageId="20" pageNumber="21">Escribania chubutensis</emphasis> +</taxonomicName> +in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of +<taxonomicName id="4C107DA0FFF20D0BC4A3F8FB8D56F8E4" box="[929,1124,1837,1861]" class="Mammalia" family="Mioclaenidae" genus="Raulvaccia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="peligrensis"> +<emphasis id="B964DA31FFF20D0BC4A3F8FB8D56F8E4" box="[929,1124,1837,1861]" italics="true" pageId="20" pageNumber="21">Raulvaccia peligrensis</emphasis> +</taxonomicName> +[MLP 90-II-12- 70 (mirrored)] +</paragraph> +<paragraph id="8BAF0623FFF20D0BC781F89F8CBBF8DC" blockId="20.[131,1428,1781,1973]" pageId="20" pageNumber="21"> +in occlusal (C) and distoocclusal (D) views. E, P4–M3 of +<taxonomicName id="4C107DA0FFF20D0BC5A0F89F8ABFF8C0" baseAuthorityName="Mones" baseAuthorityYear="2015" box="[674,909,1865,1889]" class="Mammalia" family="Didolodontidae" genus="Ricardocifellia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="protocenica"> +<emphasis id="B964DA31FFF20D0BC5A0F89F8ABFF8C0" box="[674,909,1865,1889]" italics="true" pageId="20" pageNumber="21">Ricardocifellia protocenica</emphasis> +</taxonomicName> +in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of +<taxonomicName id="4C107DA0FFF20D0BC595F8B08D37F8DC" authority="Paula Couto, 1952" authorityName="Paula Couto" authorityYear="1952" box="[663,1029,1893,1917]" class="Mammalia" family="Didolodontidae" genus="Lamegoia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="conodonta"> +<emphasis id="B964DA31FFF20D0BC595F8B08A60F8DC" box="[663,850,1893,1917]" italics="true" pageId="20" pageNumber="21">Lamegoia conodonta</emphasis> +Paula Couto, 1952 +</taxonomicName> +in occlusal view (cast of MNRJ 1465-V). +</paragraph> +<paragraph id="8BAF0623FFF20D0BC781F8578CB5F814" blockId="20.[131,1428,1781,1973]" pageId="20" pageNumber="21"> +G, P1–M3 of +<taxonomicName id="4C107DA0FFF20D0BC601F85788FBF838" authorityName="Ameghino" authorityYear="1897" box="[259,457,1921,1945]" class="Mammalia" family="Didolodontidae" genus="Didolodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="multicuspis"> +<emphasis id="B964DA31FFF20D0BC601F85788FBF838" box="[259,457,1921,1945]" italics="true" pageId="20" pageNumber="21">Didolodus multicuspis</emphasis> +</taxonomicName> +in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of +<taxonomicName id="4C107DA0FFF20D0BC781F84B88BEF814" authorityName="Cifelli" authorityYear="1983" box="[131,396,1949,1973]" class="Mammalia" family="Protolipternidae" genus="Protolipterna" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="ellipsodontoides"> +<emphasis id="B964DA31FFF20D0BC781F84B88BEF814" box="[131,396,1949,1973]" italics="true" pageId="20" pageNumber="21">Protolipterna ellipsodontoides</emphasis> +</taxonomicName> +in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of +<taxonomicName id="4C107DA0FFF20D0BC4C6F84B8D95F814" authorityName="Paula Couto" authorityYear="1952" box="[964,1191,1949,1973]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="scotti"> +<emphasis id="B964DA31FFF20D0BC4C6F84B8D95F814" box="[964,1191,1949,1973]" italics="true" pageId="20" pageNumber="21">Asmithwoodwardia scotti</emphasis> +</taxonomicName> +in occlusal view [DGM +</paragraph> +<paragraph id="8BAF0623FFF30D0AC773FF4688FFFE22" blockId="21.[113,763,144,638]" pageId="21" pageNumber="22"> +orders ( +<tableCitation id="C6923398FFF30D0AC7C6FF46883DFF09" box="[196,271,144,168]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="21" pageNumber="22">Table 2</tableCitation> +). Part of the problem of resolving didolodontid affinities is that their fossil record is limited to mostly dental remains. Nonetheless, recently some authors have started to consider the family +<taxonomicName id="4C107DA0FFF30D0AC648FF3888C0FEA7" box="[330,498,238,262]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Protolipternidae</taxonomicName> +as a junior synonym of +<taxonomicName id="4C107DA0FFF30D0AC773FEDB8824FE84" box="[113,278,269,293]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +, and +<taxonomicName id="4C107DA0FFF30D0AC653FEDB88C7FE84" box="[337,501,269,293]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +as a family of litopterns, but without any phylogenetic analysis supporting this new arrangement ( +<bibRefCitation id="EF817BD2FFF30D0AC7EDFE9A88A4FEC5" author="Gelfo" box="[239,406,332,356]" etAl="et al." firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="285 - 91" refId="ref42550" refString="Gelfo JN, Lopez GM, Lorente M. Los ungulados arcaicos de America del Sur: ' Condylarthra' y Litopterna. Contribuciones Cientificas Del Museo Argentino de Ciencias Naturales ' Bernardino Rivadavia' 2016; 6: 285 - 91." type="journal article" year="2016"> +Gelfo +<emphasis id="B964DA31FFF30D0AC62CFE9B886FFEC5" box="[302,349,332,356]" italics="true" pageId="21" pageNumber="22">et al.</emphasis> +2016 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF30D0AC6A3FE9A88D2FEC5" author="Gelfo" box="[417,480,332,356]" firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="132 - 45" refId="ref42677" refString="Gelfo JN, Alonso RN, Madden RH et al. An Eocene bunodont South American native ungulate (Didolodontidae) from the Lumbrera Formation, Salta Province, Argentina. Ameghiniana 2020 a; 57: 132 - 45. https: // doi. org / 10.5710 / amgh. 29.11.2019.3293" type="journal article" year="2020" yearSuffix="a"> +2020 +<bibRefCitation id="EF817BD2FFF30D0AC6D7FE9A8BFFFEC5" author="Corona A & Badin AC & Perea D" box="[469,717,332,356]" firstAuthor="a" pageId="21" pageNumber="22" refId="ref41392" refString="Corona A, Badin AC, Perea D et al. A new genus and species and additional reports of the South American native ungulates Proterotheriidae (Mammalia, Litopterna) in the Late Miocene of Uruguay. Journal of South American Earth Sciences 2020; 102: 102646. https: // doi. org / 10.1016 / j. jsames. 2020.102646" type="journal volume" year="2020">a, Croft and López 2020</bibRefCitation> +</bibRefCitation> +; see more in +<taxonomicName id="4C107DA0FFF30D0AC7C4FEBD885CFE22" box="[198,366,363,387]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Protolipternidae</taxonomicName> +section). +</paragraph> +<paragraph id="8BAF0623FFF30D0AC78FFE5D8839FDDF" blockId="21.[113,763,144,638]" pageId="21" pageNumber="22"> +The earliest accepted members of +<taxonomicName id="4C107DA0FFF30D0AC527FE5D8BFBFE02" box="[549,713,395,419]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +are +<taxonomicName id="4C107DA0FFF30D0AC773FE7C8867FE63" box="[113,341,426,450]" class="Mammalia" family="Mioclaenidae" genus="Escribania" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="chubutensis"> +<emphasis id="B964DA31FFF30D0AC773FE7C8867FE63" box="[113,341,426,450]" italics="true" pageId="21" pageNumber="22">Escribania chubutensis</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFF30D0AC694FE7C8B42FE63" box="[406,624,426,450]" class="Mammalia" family="Mioclaenidae" genus="Raulvaccia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="peligrensis"> +<emphasis id="B964DA31FFF30D0AC694FE7C8B42FE63" box="[406,624,426,450]" italics="true" pageId="21" pageNumber="22">Raulvaccia peligrensis</emphasis> +</taxonomicName> +from Punta Peligro, +<collectingCountry id="F30746B3FFF30D0AC7D0FE1F8805FE40" box="[210,311,457,481]" name="Argentina" pageId="21" pageNumber="22">Argentina</collectingCountry> +and the youngest are +<taxonomicName id="4C107DA0FFF30D0AC562FE1F89CAFDA0" authority=", Paulogervasia" authorityName="Paulogervasia" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF30D0AC562FE1F8BC6FE40" box="[608,756,457,481]" italics="true" pageId="21" pageNumber="22">Ernestokokenia</emphasis> +, +<emphasis id="B964DA31FFF30D0AC773FE3F89CAFDA0" box="[113,248,489,513]" italics="true" pageId="21" pageNumber="22">Paulogervasia</emphasis> +</taxonomicName> +, and +<taxonomicName id="4C107DA0FFF30D0AC63BFE3F88ACFDA0" authorityName="Berg" authorityYear="1899" box="[313,414,489,513]" class="Mammalia" family="Proterotheriidae" genus="Xesmodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="21" pageNumber="22" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF30D0AC63BFE3F88ACFDA0" box="[313,414,489,513]" italics="true" pageId="21" pageNumber="22">Xesmodon</emphasis> +</taxonomicName> +from different Patagonian localities correlated with the Mustersan SALMA ( +<bibRefCitation id="EF817BD2FFF30D0AC543FDDE8B87FD81" author="Gelfo" box="[577,693,520,544]" firstAuthor="Gelfo" pageId="21" pageNumber="22" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo 2006</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF30D0AC5C3FDDE8BC7FD81" author="Gelfo" box="[705,757,520,544]" firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="651 - 60" refId="ref42229" refString="Gelfo JN. The ' Condylarth' Raulvaccia peligrensis (Mammalia: Didolodontidae) from the Paleocene of Patagonia, Argentina. Journal of Vertebrate Paleontology 2007; 27: 651 - 60. https: // doi. org / 10.1671 / 0272 - 4634 (2007) 27 (651: tcrpmd) 2.0. co; 2" type="journal article" year="2007">2007</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF30D0AC773FDF18997FD9E" author="Gelfo" box="[113,165,551,575]" firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="130 - 42" refId="ref42288" refString="Gelfo JN. The ' condylarth' Didolodontidae from Gran Barranca: history of the bunodont South American mammals until the Eocene- Oligocene transition. In: Madden RH, Carlini AA, Vucetich MG, Kay RF (eds.), The Paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge: Cambridge University Press, 2010, 130 - 42." type="book chapter" year="2010">2010</bibRefCitation> +), which gives a temporal interval of 63.8–36.5 Mya for this family ( +<figureCitation id="132B1AA6FFF30D0AC7C0FD918838FDFE" box="[194,266,583,607]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="21" pageNumber="22">Fig. 2B</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFF30D0AC51AFD918B41FDFE" box="[536,627,583,607]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="21" pageNumber="22">Table S1</tableCitation> +; Woodburne +<emphasis id="B964DA31FFF30D0AC773FDB189A8FDDF" box="[113,154,614,638]" italics="true" pageId="21" pageNumber="22">et al</emphasis> +. 2014a, b). +</paragraph> +<emphasis id="B964DA31FFF30D0AC608FD4B8B53FD19" box="[266,609,669,696]" italics="true" pageId="21" pageNumber="22"> +<subSubSection id="C30A55A8FFF30D0AC608FD4B8893FD16" box="[266,417,669,695]" pageId="21" pageNumber="22" type="nomenclature"> +<paragraph id="8BAF0623FFF30D0AC608FD4B8893FD16" blockId="21.[113,765,669,1672]" box="[266,417,669,695]" pageId="21" pageNumber="22"> +<taxonomicName id="4C107DA0FFF30D0AC608FD4B8893FD16" box="[266,417,669,695]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Protolipternidae</taxonomicName> +</paragraph> +</subSubSection> +<subSubSection id="C30A55A8FFF30D0AC6A4FD488B53FD19" box="[422,609,670,696]" pageId="21" pageNumber="22" type="description"> +<paragraph id="8BAF0623FFF30D0AC6A4FD488B53FD19" blockId="21.[113,765,669,1672]" box="[422,609,670,696]" pageId="21" pageNumber="22"> +( +<figureCitation id="132B1AA6FFF30D0AC6B2FD488B23FD19" box="[432,529,670,696]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="21" pageNumber="22">Figs 8H–J</figureCitation> +, +<figureCitation id="132B1AA6FFF30D0AC519FD488B6AFD19" box="[539,600,670,696]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="21" pageNumber="22">9F–H</figureCitation> +) +</paragraph> +</subSubSection> +</emphasis> +<subSubSection id="C30A55A8FFF30D0AC773FD128AA7FA8E" pageId="21" pageNumber="22" type="multiple"> +<paragraph id="8BAF0623FFF30D0AC773FD128C9FFDA1" blockId="21.[113,765,669,1672]" lastBlockId="21.[808,1461,144,1327]" pageId="21" pageNumber="22"> +<taxonomicName id="4C107DA0FFF30D0AC773FD128829FD7D" box="[113,283,708,732]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Protolipternidae</taxonomicName> +currently includes three genera [or two if we excluded +<taxonomicName id="4C107DA0FFF30D0AC7D5FD3588A5FD5A" authorityName="Ameghino" authorityYear="1901" box="[215,407,739,763]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF30D0AC7D5FD3588A5FD5A" box="[215,407,739,763]" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia</emphasis> +</taxonomicName> +as +<bibRefCitation id="EF817BD2FFF30D0AC6BFFD358B79FD5A" author="Gelfo" box="[445,587,739,763]" firstAuthor="Gelfo" pageId="21" pageNumber="22" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo (2006)</bibRefCitation> +suggested] with an exclusively Palaeogene distribution ( +<figureCitation id="132B1AA6FFF30D0AC506FCD58B09FCBA" box="[516,571,771,795]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="21" pageNumber="22">Fig. 2</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFF30D0AC7C4FCF48810FC9B" box="[198,290,802,826]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="21" pageNumber="22">Table S1</tableCitation> +). This family was proposed by Cifelli (1983a) to classify specimens from Itaboraí (Itaboraian SALMA) with bunodont didolodontid-like dentition and litoptern-like tarsals. The tarsals show specializations that among SANUs are only present in litopterns, which include a spool-shaped astragalar body and a posterior astragalocalcaneal facet that allows rapid flexion of the crurotarsal joint (Cifelli 1983a). The taxonomic changes generated from this familial proposal included the consideration of +<taxonomicName id="4C107DA0FFF30D0AC7CCFBCB88EEFB95" authorityName="Paula Couto" authorityYear="1952" box="[206,476,1053,1077]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="prototypica"> +<emphasis id="B964DA31FFF30D0AC7CCFBCB88EEFB95" box="[206,476,1053,1077]" italics="true" pageId="21" pageNumber="22">Victorlemoinea prototypica</emphasis> +</taxonomicName> +[a former litoptern (Paula Couto 1952)] as a ‘condylarth’ closely related to didolodontids, and +<taxonomicName id="4C107DA0FFF30D0AC7A8FB8D8886FBD2" baseAuthorityName="Cifelli" baseAuthorityYear="1983" box="[170,436,1115,1139]" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="parayirunhor"> +<emphasis id="B964DA31FFF30D0AC7A8FB8D8886FBD2" box="[170,436,1115,1139]" italics="true" pageId="21" pageNumber="22">Miguelsoria parayirunhor</emphasis> +</taxonomicName> +[considered a didolodontid +<taxonomicName id="4C107DA0FFF30D0AC773FBAD88ABFB32" baseAuthorityName="Paula Couto" baseAuthorityYear="1952" box="[113,409,1147,1171]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="parayirunhor"> +<emphasis id="B964DA31FFF30D0AC773FBAD88ABFB32" box="[113,409,1147,1171]" italics="true" pageId="21" pageNumber="22">Ernestokokenia parayirunhor</emphasis> +</taxonomicName> +by Paula Couto (1952)] with +<taxonomicName id="4C107DA0FFF30D0AC773FB4C885EFB13" authorityName="Paula Couto" authorityYear="1952" box="[113,364,1178,1202]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="scotti"> +<emphasis id="B964DA31FFF30D0AC773FB4C885EFB13" box="[113,364,1178,1202]" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia scotti</emphasis> +</taxonomicName> +[considered a didolodontid by Paula Couto (1978)] as litopterns. Cifelli (1983a) grouped the last two, plus a new species +<taxonomicName id="4C107DA0FFF30D0AC687FB0F8983FAB1" authority="Cifelli 1983 a" authorityName="Cifelli" authorityYear="1983" class="Mammalia" family="Protolipternidae" genus="Protolipterna" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="ellipsodontoides"> +<emphasis id="B964DA31FFF30D0AC687FB0F8B9CFB50" box="[389,686,1241,1265]" italics="true" pageId="21" pageNumber="22">Protolipterna ellipsodontoides</emphasis> +Cifelli 1983a +</taxonomicName> +, within this family. However, the tarsals referred to +<taxonomicName id="4C107DA0FFF30D0AC773FAC188A6FA8E" authorityName="Cifelli" authorityYear="1983" box="[113,404,1303,1327]" class="Mammalia" family="Protolipternidae" genus="Protolipterna" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="ellipsodontoides"> +<emphasis id="B964DA31FFF30D0AC773FAC188A6FA8E" box="[113,404,1303,1327]" italics="true" pageId="21" pageNumber="22">Protolipterna ellipsodontoides</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFF30D0AC6C5FAC18BF6FA8E" baseAuthorityName="Cifelli" baseAuthorityYear="1983" box="[455,708,1303,1327]" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="parayirunhor"> +<emphasis id="B964DA31FFF30D0AC6C5FAC18BF6FA8E" box="[455,708,1303,1327]" italics="true" pageId="21" pageNumber="22">Miguelsoria parayirunhor</emphasis> +</taxonomicName> +were not found in association with their alleged dentition, being assigned indirectly by their relative size, abundance, and expected morphology (Cifelli 1983b). Some authors have accepted this interpretation (e.g., +<bibRefCitation id="EF817BD2FFF30D0AC6D2FA438B62FA0D" author="Cifelli" box="[464,592,1428,1453]" firstAuthor="Cifelli" pageId="21" pageNumber="22" pagination="249 - 66" refId="ref40909" refString="Cifelli RL. South American ungulate evolution and extinction. In: Stehli FG, Webb SD (eds), The Great American Biotic Interchange. New York, NY: Springer, 1985, 249 - 66." type="book chapter" year="1985">Cifelli 1985</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF30D0AC561FA438BA5FA0C" author="Cifelli" box="[611,663,1429,1453]" firstAuthor="Cifelli" pageId="21" pageNumber="22" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">1993</bibRefCitation> +, Muizon and Cifelli 2000, +<bibRefCitation id="EF817BD2FFF30D0AC62AFA6288F4FA6D" author="Bergqvist" box="[296,454,1460,1484]" firstAuthor="Bergqvist" pageId="21" pageNumber="22" pagination="107 - 33" refId="ref39850" refString="Bergqvist LP. Postcranial skeleton of the Upper Paleocene of Itaborai Basin, Brazil. In: Sargis EJ, Dagosto M, (eds), Mammalian Evolutionary Morphology: A Tribute to Frederik S. Szalay. New York, NY: Springer-Verlag, 2008, 107 - 33." type="book chapter" year="2008">Bergqvist 2008</bibRefCitation> +), although others have questioned it ( +<bibRefCitation id="EF817BD2FFF30D0AC7DFFA0588C1FA4A" author="Hoffstetter and Soria" box="[221,499,1491,1515]" firstAuthor="Hoffstetter" pageId="21" pageNumber="22" pagination="1619 - 22" refId="ref43229" refString="Hoffstetter R, Soria MF. Neolodus colombianus gen. et sp. nov., un noveau Condylarthre (Mammalia) dans le Miocene de Colombie. Comptes Rendus de la Academie de Sciences, Paris 1986; 303: 1619 - 22." type="journal article" year="1986">Hoffstetter and Soria 1986</bibRefCitation> +, Soria 2001, +<bibRefCitation id="EF817BD2FFF30D0AC57CFA058BC6FA4A" author="Gelfo" box="[638,756,1491,1515]" firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="130 - 42" refId="ref42288" refString="Gelfo JN. The ' condylarth' Didolodontidae from Gran Barranca: history of the bunodont South American mammals until the Eocene- Oligocene transition. In: Madden RH, Carlini AA, Vucetich MG, Kay RF (eds.), The Paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge: Cambridge University Press, 2010, 130 - 42." type="book chapter" year="2010">Gelfo 2010</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF30D0AC773FA258874F9AA" author="Gelfo and Sige" box="[113,326,1523,1547]" firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="665 - 78" refId="ref42458" refString="Gelfo JN, Sige B. A new didolodontid mammal from the Late Paleocene - earliest Eocene of Laguna Umayo, Peru. Acta Palaeontologica Polonica 2011; 56: 665 - 78. https: // doi. org / 10.4202 / app. 2010.0067" type="journal article" year="2011">Gelfo and Sigé 2011</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF30D0AC651FA258B7DF9AA" author="Gelfo and Lorente" box="[339,591,1523,1547]" firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="249 - 59" refId="ref42404" refString="Gelfo JN, Lorente M. The alleged astragalar remains of Didolodus Ameghino, 1897 (Mammalia, Panameriungulata) and a critic of isolated bone association models. Bulletin of Geosciences 2012; 87: 249 - 59. https: // doi. org / 10.3140 / bull. geosci. 1301" type="journal article" year="2012">Gelfo and Lorente 2012</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF30D0AC55EFA258BDBF9AB" author="Lorente" box="[604,745,1522,1546]" firstAuthor="Lorente" pageId="21" pageNumber="22" refId="ref44083" refString="Lorente M. Desarrollo de Modelos de Asociacion y Clasificaciones de Restos Postcraneanos Aislados de Ungulados Nativos del Paleoceno- Eoceno de America del Sur. D. Phil. Thesis, Universidad Nacional de la Plata, Argentina, 2015." type="book" year="2015">Lorente 2015</bibRefCitation> +). For example, Soria (2001) suggested that the astragali and calcanea attributed by Cifelli (1983b) to didolodontids were most likely notoungulate elements. Indeed, a study that indirectly assigned isolated petrosals to the protolipternid +<taxonomicName id="4C107DA0FFF30D0AC584F9A68BC8F929" authorityName="Cifelli" authorityYear="1983" box="[646,762,1648,1672]" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF30D0AC584F9A68BC8F929" box="[646,762,1648,1672]" italics="true" pageId="21" pageNumber="22">Miguelsoria</emphasis> +</taxonomicName> +from Itaboraí failed to recover a monophyletic +<taxonomicName id="4C107DA0FFF30D0AC205FF468C47FF09" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[1287,1397,144,168]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="order">Litopterna</taxonomicName> +when younger and better known litopterns (e.g., +<taxonomicName id="4C107DA0FFF30D0AC3F5FF798CB1FF66" authorityName="Owen" authorityYear="1838" box="[1271,1411,175,199]" class="Mammalia" family="Macraucheniidae" genus="Macrauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF30D0AC3F5FF798CB1FF66" box="[1271,1411,175,199]" italics="true" pageId="21" pageNumber="22">Macrauchenia</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFF30D0AC428FF198A80FF46" authorityName="Ameghino" authorityYear="1887" box="[810,946,207,231]" class="Mammalia" family="Proterotheriidae" genus="Diadiaphorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF30D0AC428FF198A80FF46" box="[810,946,207,231]" italics="true" pageId="21" pageNumber="22">Diadiaphorus</emphasis> +</taxonomicName> +) were included ( +<bibRefCitation id="EF817BD2FFF30D0AC36EFF198C2BFF47" author="Billet" box="[1132,1305,206,231]" etAl="et al." firstAuthor="Billet" pageId="21" pageNumber="22" pagination="956 - 87" refId="ref39984" refString="Billet G, de Muizon C, Schellhorn R et al. Petrosal and inner ear anatomy and allometry amongst specimens referred to Litopterna (Placentalia). Zoological Journal of the Linnean Society 2015; 173: 956 - 87. https: // doi. org / 10.1111 / zoj. 12219" type="journal article" year="2015"> +Billet +<emphasis id="B964DA31FFF30D0AC3AEFF198DECFF46" box="[1196,1246,207,231]" italics="true" pageId="21" pageNumber="22">et al.</emphasis> +2015 +</bibRefCitation> +). Considering that recent association models do not discard the association of litoptern-like tarsals to protolipternids ( +<bibRefCitation id="EF817BD2FFF30D0AC3DCFED88C5BFE84" author="Lorente" box="[1246,1385,269,293]" firstAuthor="Lorente" pageId="21" pageNumber="22" refId="ref44083" refString="Lorente M. Desarrollo de Modelos de Asociacion y Clasificaciones de Restos Postcraneanos Aislados de Ungulados Nativos del Paleoceno- Eoceno de America del Sur. D. Phil. Thesis, Universidad Nacional de la Plata, Argentina, 2015." type="book" year="2015">Lorente 2015</bibRefCitation> +) made by Cifelli (1983b), some authors have started to consider the family +<taxonomicName id="4C107DA0FFF30D0AC472FE9A8D28FEC5" box="[880,1050,332,356]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Protolipternidae</taxonomicName> +as a junior synonym of +<taxonomicName id="4C107DA0FFF30D0AC20FFE9A8C80FEC5" box="[1293,1458,332,356]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +( +<bibRefCitation id="EF817BD2FFF30D0AC437FEBD8AD2FE22" author="Gelfo" box="[821,992,363,387]" etAl="et al." firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="285 - 91" refId="ref42550" refString="Gelfo JN, Lopez GM, Lorente M. Los ungulados arcaicos de America del Sur: ' Condylarthra' y Litopterna. Contribuciones Cientificas Del Museo Argentino de Ciencias Naturales ' Bernardino Rivadavia' 2016; 6: 285 - 91." type="journal article" year="2016"> +Gelfo +<emphasis id="B964DA31FFF30D0AC474FEBA8A94FE22" box="[886,934,363,387]" italics="true" pageId="21" pageNumber="22">et al.</emphasis> +2016 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF30D0AC4EFFEBD8D1FFE22" author="Gelfo" box="[1005,1069,363,387]" firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="132 - 45" refId="ref42677" refString="Gelfo JN, Alonso RN, Madden RH et al. An Eocene bunodont South American native ungulate (Didolodontidae) from the Lumbrera Formation, Salta Province, Argentina. Ameghiniana 2020 a; 57: 132 - 45. https: // doi. org / 10.5710 / amgh. 29.11.2019.3293" type="journal article" year="2020" yearSuffix="a"> +2020 +<bibRefCitation id="EF817BD2FFF30D0AC320FEBD8C13FE22" author="Corona A & Badin AC & Perea D" box="[1058,1313,363,387]" firstAuthor="a" pageId="21" pageNumber="22" refId="ref41392" refString="Corona A, Badin AC, Perea D et al. A new genus and species and additional reports of the South American native ungulates Proterotheriidae (Mammalia, Litopterna) in the Late Miocene of Uruguay. Journal of South American Earth Sciences 2020; 102: 102646. https: // doi. org / 10.1016 / j. jsames. 2020.102646" type="journal volume" year="2020">a, Croft and López 2020</bibRefCitation> +</bibRefCitation> +). In addition, based on anatomical observations and a phylogenetic analysis, +<bibRefCitation id="EF817BD2FFF30D0AC42EFE7C8A8BFE63" author="Gelfo" box="[812,953,426,450]" firstAuthor="Gelfo" pageId="21" pageNumber="22" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo (2006)</bibRefCitation> +considered that +<taxonomicName id="4C107DA0FFF30D0AC368FE7C8C56FE63" authorityName="Paula Couto" authorityYear="1952" box="[1130,1380,426,450]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="scotti"> +<emphasis id="B964DA31FFF30D0AC368FE7C8C56FE63" box="[1130,1380,426,450]" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia scotti</emphasis> +</taxonomicName> +should be generically renamed as is it not recovered as a sister taxon of the Patagonian +<taxonomicName id="4C107DA0FFF30D0AC4CBFE3F8C94FDA0" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[969,1446,489,513]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="subtrigona"> +<emphasis id="B964DA31FFF30D0AC4CBFE3F8DC7FDA0" box="[969,1269,489,513]" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia subtrigona</emphasis> +<bibRefCitation id="EF817BD2FFF30D0AC3F9FE3F8C94FDA0" author="Ameghino" box="[1275,1446,489,513]" firstAuthor="Ameghino" pageId="21" pageNumber="22" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +. +</paragraph> +<paragraph id="8BAF0623FFF30D0AC447FDDE8C6FFB94" blockId="21.[808,1461,144,1327]" pageId="21" pageNumber="22"> +Apartfromearlyphylogeneticstudiesthatfoundprotolipternids closely related to other litopterns based on Cifelli’s (1983b) tarsal associations ( +<bibRefCitation id="EF817BD2FFF30D0AC4B0FD918D15FDFE" author="Cifelli" box="[946,1063,583,607]" firstAuthor="Cifelli" pageId="21" pageNumber="22" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF30D0AC331FD918DF8FDFE" author="Bergqvist" box="[1075,1226,583,607]" firstAuthor="Bergqvist" pageId="21" pageNumber="22" refId="ref39790" refString="Bergqvist LP. Reasociacao de pos-cranio as especies de ungulados da Bacia de S. J. Itaborai (Paleoceno), Estado do Rio de Janeiro, e Filogenia dos ' Condylarthra', e ungulados Sul-Americanos com base no poscranio. D. Phil. Thesis, Universidade Federal do Rio Grande do Sul, Brazil, 1996." type="book" year="1996">Bergqvist 1996</bibRefCitation> +; +<figureCitation id="132B1AA6FFF30D0AC3D5FD918C12FDFE" box="[1239,1312,583,607]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="21" pageNumber="22">Fig. 1C</figureCitation> +), other phylogenetic studies including protolipternids and didolodontids have usually failed to find them as distinctly separate groups [e.g., Muizon and Cifelli (2000); see also +<taxonomicName id="4C107DA0FFF30D0AC3B0FD738C60FD1C" box="[1202,1362,677,701]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +section]. Other phylogenetic studies with a wide taxon sampling have found +<taxonomicName id="4C107DA0FFF30D0AC46DFD358D25FD5A" box="[879,1047,739,763]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Protolipternidae</taxonomicName> +: in a basal position in Laurasiatheria as a sister group of +<taxonomicName id="4C107DA0FFF30D0AC4D8FCD58C52FCBA" authority="(O'Leary et al. 2013)" baseAuthorityName="O'Leary" baseAuthorityYear="2013" box="[986,1376,771,795]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="family"> +Didolodontidae (O’Leary +<emphasis id="B964DA31FFF30D0AC3EBFCD58C26FCBA" box="[1257,1300,771,795]" italics="true" pageId="21" pageNumber="22">et al</emphasis> +. 2013) +</taxonomicName> +; closely related to other more derived litopterns than to other SANUs (Muizon +<emphasis id="B964DA31FFF30D0AC48FFC948A8BFCF8" box="[909,953,833,857]" italics="true" pageId="21" pageNumber="22">et al</emphasis> +. 2015); among didolodontids at the stem of litopterns but not forming a monophyletic group ( +<bibRefCitation id="EF817BD2FFF30D0AC221FCB78A6FFC39" author="Kramarz" etAl="et al." firstAuthor="Kramarz" pageId="21" pageNumber="22" refId="ref43620" refString="Kramarz AG, Bond M, MacPhee RDE. On the alleged perissodactyl affinities of the ' condylarth' Escribania chubutensis and other endemic South American ungulate-like placentals. Journal of Vertebrate Paleontology 2021; 41: e 1986716." type="journal volume" year="2021"> +Kramarz +<emphasis id="B964DA31FFF30D0AC280FCB78C9CFCD8" box="[1410,1454,865,889]" italics="true" pageId="21" pageNumber="22">et al</emphasis> +. 2021 +</bibRefCitation> +); and forming a monophyletic group as part of a polytomy that includes didolodontids, sparnotheriodontids, indaleciids, and North American phenacodontids (Zimicz +<emphasis id="B964DA31FFF30D0AC3D4FC698DCDFC76" box="[1238,1279,959,983]" italics="true" pageId="21" pageNumber="22">et al</emphasis> +. 2022). However, so far there has not been any phylogenetic analysis that includes protolipternids alongside representatives of all litoptern families, didolodontids, and the different SANU orders ( +<tableCitation id="C6923398FFF30D0AC206FBCB8C7FFB94" box="[1284,1357,1053,1077]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="21" pageNumber="22">Table 2</tableCitation> +). +</paragraph> +<paragraph id="8BAF0623FFF30D0AC447FBEA8AA7FA8E" blockId="21.[808,1461,144,1327]" pageId="21" pageNumber="22"> +If we follow Cifelli (1983a) and consider +<taxonomicName id="4C107DA0FFF30D0AC20BFBEA8C81FBF5" box="[1289,1459,1084,1108]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Protolipternidae</taxonomicName> +as a valid family, its earliest members are +<taxonomicName id="4C107DA0FFF30D0AC232FB8D8DF6FB32" authority=", Miguelsoriaparayirunhor" authorityName="Miguelsoriaparayirunhor" class="Mammalia" family="Protolipternidae" genus="Protolipterna" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="ellipsodontoides"> +<emphasis id="B964DA31FFF30D0AC232FB8D8AF6FB32" italics="true" pageId="21" pageNumber="22">Protolipterna ellipsodontoides</emphasis> +, +<emphasis id="B964DA31FFF30D0AC4CEFBAD8DF6FB32" box="[972,1220,1147,1171]" italics="true" pageId="21" pageNumber="22">Miguelsoriaparayirunhor</emphasis> +</taxonomicName> +,and +<taxonomicName id="4C107DA0FFF30D0AC3EDFBAD8A6EFB13" authorityName="Paula Couto" authorityYear="1952" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="scotti"> +<emphasis id="B964DA31FFF30D0AC3EDFBAD8A6EFB13" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia scotti</emphasis> +</taxonomicName> +from Itaboraí, +<collectingCountry id="F30746B3FFF30D0AC4FAFB4C8D04FB13" box="[1016,1078,1178,1202]" name="Brazil" pageId="21" pageNumber="22">Brazil</collectingCountry> +, and the youngest member would be +<taxonomicName id="4C107DA0FFF30D0AC429FB6F8D68FB70" authorityName="Ameghino" authorityYear="1901" box="[811,1114,1209,1233]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="subtrigona"> +<emphasis id="B964DA31FFF30D0AC429FB6F8D68FB70" box="[811,1114,1209,1233]" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia subtrigona</emphasis> +</taxonomicName> +with its last records in Cañadón Vaca, +<collectingCountry id="F30746B3FFF30D0AC46BFB0F8AE2FB50" box="[873,976,1241,1265]" name="Argentina" pageId="21" pageNumber="22">Argentina</collectingCountry> +( +<bibRefCitation id="EF817BD2FFF30D0AC4E6FB0F8D69FB50" author="Gelfo" box="[996,1115,1241,1265]" firstAuthor="Gelfo" pageId="21" pageNumber="22" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo 2006</bibRefCitation> +), which gives the family a temporal interval of 56–42 Mya ( +<figureCitation id="132B1AA6FFF30D0AC35CFB2E8D95FAB1" box="[1118,1191,1272,1296]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="21" pageNumber="22">Fig. 2B</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFF30D0AC429FAC18AB4FA8E" box="[811,902,1303,1327]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="21" pageNumber="22">Table S1</tableCitation> +). +</paragraph> +</subSubSection> +<subSection id="E29F1FC8FFF30D05C4AEFA8B8AB6FC16" lastPageId="26" lastPageNumber="27" pageId="21" pageNumber="22" type="materials_methods"> +<paragraph id="8BAF0623FFF30D0AC4AEFA8B8C03FAD6" blockId="21.[940,1329,1373,1399]" box="[940,1329,1373,1399]" pageId="21" pageNumber="22"> +<heading id="D0E7B14FFFF30D0AC4AEFA8B8C03FAD6" allCaps="true" bold="true" box="[940,1329,1373,1399]" centered="true" fontSize="11" level="1" pageId="21" pageNumber="22" reason="1"> +<emphasis id="B964DA31FFF30D0AC4AEFA8B8C03FAD6" bold="true" box="[940,1329,1373,1399]" pageId="21" pageNumber="22">MATERIALS AND METHODS</emphasis> +</heading> +</paragraph> +<paragraph id="8BAF0623FFF30D0AC318FA5A8DF1FA07" blockId="21.[1050,1219,1420,1446]" box="[1050,1219,1420,1446]" pageId="21" pageNumber="22"> +<emphasis id="B964DA31FFF30D0AC318FA5A8DF1FA07" bold="true" box="[1050,1219,1420,1446]" pageId="21" pageNumber="22">Taxon sampling</emphasis> +</paragraph> +<paragraph id="8BAF0623FFF30D0AC428FA658D89F926" blockId="21.[809,1461,1459,1671]" pageId="21" pageNumber="22"> +As the main objectives of this study are resolving the interfamilial relationship of +<taxonomicName id="4C107DA0FFF30D0AC4C9FA048D0AFA4B" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[971,1080,1490,1514]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="order">Litopterna</taxonomicName> +and testing the monophyly of this order among other SANUs and early placental mammals, our taxon selection includes the earliest members of each litoptern family and also taxa from groups that have been linked to +<taxonomicName id="4C107DA0FFF30D0AC42EF9998AABF9C6" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[812,921,1615,1639]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="order">Litopterna</taxonomicName> +in previous studies. In total, our dataset includes 38 taxa (Supporting information, File S2). +</paragraph> +<paragraph id="8BAF0623FFF30D0AC773F92D8C5CF88E" blockId="21.[113,1431,1787,1979]" pageId="21" pageNumber="22"> +358M ( +<typeStatus id="54ABB881FFF30D0AC7B5F92D8821F8B2" box="[183,275,1787,1811]" pageId="21" pageNumber="22" type="holotype">holotype</typeStatus> +; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of +<taxonomicName id="4C107DA0FFF30D0AC5FEF92D8AD1F8B2" baseAuthorityName="Paula Couto" baseAuthorityYear="1952" box="[764,995,1787,1811]" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="parayiruhnor"> +<emphasis id="B964DA31FFF30D0AC5FEF92D8AD1F8B2" box="[764,995,1787,1811]" italics="true" pageId="21" pageNumber="22">Miguelsoria parayiruhnor</emphasis> +</taxonomicName> +in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of +<taxonomicName id="4C107DA0FFF30D0AC3A3F8C18C77F88F" authorityName="Muizon and Marshall" authorityYear="1987" box="[1185,1349,1815,1839]" class="Mammalia" family="Mioclaenidae" genus="Molinodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="suarezi"> +<emphasis id="B964DA31FFF30D0AC3A3F8C18C77F88F" box="[1185,1349,1815,1839]" italics="true" pageId="21" pageNumber="22">Molinodus suarezi</emphasis> +</taxonomicName> +(A) +</paragraph> +<paragraph id="8BAF0623FFF30D0AC773F8E58B56F81B" blockId="21.[113,1431,1787,1979]" pageId="21" pageNumber="22"> +and the M2 of +<taxonomicName id="4C107DA0FFF30D0AC7F9F8E58884F8EA" authorityName="Paula Couto" authorityYear="1952" box="[251,438,1843,1867]" class="Mammalia" family="Didolodontidae" genus="Lamegoia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="conodonta"> +<emphasis id="B964DA31FFF30D0AC7F9F8E58884F8EA" box="[251,438,1843,1867]" italics="true" pageId="21" pageNumber="22">Lamegoia conodonta</emphasis> +</taxonomicName> +(B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista. Scale bars equal +<quantity id="4CE8ABC6FFF30D0AC532F8758B6DF81B" box="[560,607,1955,1979]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="21" pageNumber="22" unit="cm" value="1.0">1 cm</quantity> +. +</paragraph> +<caption id="DF6F56ABFFF00D08C783F8C888D7F81B" lastPageId="23" lastPageNumber="24" pageId="22" pageNumber="23" startId="22.[129,194,1822,1846]" targetBox="[241,1361,147,1792]" targetPageId="22" targetType="figure"> +<paragraph id="8BAF0623FFF00D09C783F8C88CAEF806" blockId="22.[129,1465,1822,1959]" pageId="22" pageNumber="23"> +<emphasis id="B964DA31FFF00D09C783F8C889EAF896" bold="true" box="[129,216,1822,1847]" pageId="22" pageNumber="23">Figure 9.</emphasis> +Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view. A, p4–m3 of +<taxonomicName id="4C107DA0FFF00D09C39DF8C98C71F897" authorityName="Muizon and Marshall" authorityYear="1987" box="[1183,1347,1823,1847]" class="Mammalia" family="Mioclaenidae" genus="Molinodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="suarezi"> +<emphasis id="B964DA31FFF00D09C39DF8C98C71F897" box="[1183,1347,1823,1847]" italics="true" pageId="22" pageNumber="23">Molinodus suarezi</emphasis> +</taxonomicName> +[cast of MHNC 13867 (mirrored)]. B, m2–m3 of +<taxonomicName id="4C107DA0FFF00D09C527F8ED8BC3F8F2" box="[549,753,1851,1875]" class="Mammalia" family="Mioclaenidae" genus="Escribania" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="chubutensis"> +<emphasis id="B964DA31FFF00D09C527F8ED8BC3F8F2" box="[549,753,1851,1875]" italics="true" pageId="22" pageNumber="23">Escribania chubutensis</emphasis> +</taxonomicName> +[UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of +<taxonomicName id="4C107DA0FFF00D09C783F8818871F8CE" box="[129,323,1879,1903]" class="Mammalia" family="Mioclaenidae" genus="Raulvaccia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="peligrensis"> +<emphasis id="B964DA31FFF00D09C783F8818871F8CE" box="[129,323,1879,1903]" italics="true" pageId="22" pageNumber="23">Raulvaccia peligrensis</emphasis> +</taxonomicName> +[MLP 90−II−12−69 (mirrored)]. D, P5–M3 of +<taxonomicName id="4C107DA0FFF00D09C408F8818AC7F8CF" baseAuthorityName="Mones" baseAuthorityYear="2015" box="[778,1013,1879,1903]" class="Mammalia" family="Didolodontidae" genus="Ricardocifellia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="protocenica"> +<emphasis id="B964DA31FFF00D09C408F8818AC7F8CF" box="[778,1013,1879,1903]" italics="true" pageId="22" pageNumber="23">Ricardocifellia protocenica</emphasis> +</taxonomicName> +[MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of +<taxonomicName id="4C107DA0FFF00D09C6DDF8A58B97F82A" authorityName="Ameghino" authorityYear="1897" box="[479,677,1907,1931]" class="Mammalia" family="Didolodontidae" genus="Didolodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="multicuspis"> +<emphasis id="B964DA31FFF00D09C6DDF8A58B97F82A" box="[479,677,1907,1931]" italics="true" pageId="22" pageNumber="23">Didolodus multicuspis</emphasis> +</taxonomicName> +(MACN A 10689). F, p2–m3 of +<taxonomicName id="4C107DA0FFF00D09C4DFF8A58DD4F82A" box="[989,1254,1907,1931]" class="Mammalia" family="Protolipternidae" genus="Protolipterna" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="ellipsodontoides"> +<emphasis id="B964DA31FFF00D09C4DFF8A58DD4F82A" box="[989,1254,1907,1931]" italics="true" pageId="22" pageNumber="23">Protolipterna ellipsodontoides</emphasis> +</taxonomicName> +[cast of DGM 1308M (mirrored)]. G, p1–m3 of +<taxonomicName id="4C107DA0FFF00D09C67AF8598B69F807" authorityName="Paula Couto" authorityYear="1952" box="[376,603,1935,1959]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="scotti"> +<emphasis id="B964DA31FFF00D09C67AF8598B69F807" box="[376,603,1935,1959]" italics="true" pageId="22" pageNumber="23">Asmithwoodwardia scotti</emphasis> +</taxonomicName> +(cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of +<taxonomicName id="4C107DA0FFF00D09C381F8598C58F806" baseAuthorityName="Paula Couto" baseAuthorityYear="1952" box="[1155,1386,1935,1959]" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="parayiruhnor"> +<emphasis id="B964DA31FFF00D09C381F8598C58F806" box="[1155,1386,1935,1959]" italics="true" pageId="22" pageNumber="23">Miguelsoria parayiruhnor</emphasis> +</taxonomicName> +(cast +</paragraph> +<paragraph id="8BAF0623FFF10D08C78FFF46886CFE63" blockId="23.[112,765,144,1797]" pageId="23" pageNumber="24"> +We chose +<taxonomicName id="4C107DA0FFF10D08C7FFFF468BC8FF09" authority="Sloan & Van Valen, 1965" authorityName="Sloan & Van Valen" authorityYear="1965" box="[253,762,144,168]" class="Mammalia" family="Tupaiidae" genus="Protungulatum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scandentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="donnae"> +<emphasis id="B964DA31FFF10D08C7FFFF4688D0FF09" box="[253,482,144,168]" italics="true" pageId="23" pageNumber="24">Protungulatum donnae</emphasis> +Sloan & Van Valen, 1965 +</taxonomicName> +as the outgroup, as it is one of the earliest archaic ungulates (‘condylarths’) known from the biochronologic zones Pu1 and Pu2 ( +<bibRefCitation id="EF817BD2FFF10D08C7DBFF3888B9FEA7" author="Cifelli" box="[217,395,238,262]" etAl="et al." firstAuthor="Cifelli" pageId="23" pageNumber="24" pagination="21 - 42" refId="ref41175" refString="Cifelli RL, Eberle JJ, Lofgren DL, et al. Mammalian biochronology of the Latest Cretaceous. In: Woodburne MO (ed.), Late Cretaceous and Cenozoic Mammals of North America. New York: Columbia University Press, 2004, 21 - 42." type="book chapter" year="2004"> +Cifelli +<emphasis id="B964DA31FFF10D08C622FF398862FEA7" box="[288,336,238,262]" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +2004 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFF10D08C69AFF388B69FEA7" author="Lofgren" box="[408,603,238,262]" etAl="et al." firstAuthor="Lofgren" pageId="23" pageNumber="24" pagination="43 - 105" refId="ref43948" refString="Lofgren DL, Lillegraven JA, Clemens WA, et al. Paleocene biochronology: the Puercan through Clarkforkian Land Mammal Ages. In: Woodburne MO (ed.), Late Cretaceous and Cenozoic Mammals of North America. New York, NY: Columbia University Press, 2004, 43 - 105." type="book chapter" year="2004"> +Lofgren +<emphasis id="B964DA31FFF10D08C6F2FF398B13FEA7" box="[496,545,238,262]" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +2004 +</bibRefCitation> +) in the earliest Puercan NALMA (North American land mammal ages; Early Palaeocene). The genus +<taxonomicName id="4C107DA0FFF10D08C662FEFB88C6FEE4" authorityName="Sloan & Van Valen" authorityYear="1965" box="[352,500,301,325]" class="Mammalia" family="Tupaiidae" genus="Protungulatum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scandentia" pageId="23" pageNumber="24" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF10D08C662FEFB88C6FEE4" box="[352,500,301,325]" italics="true" pageId="23" pageNumber="24">Protungulatum</emphasis> +</taxonomicName> +has also been found in deposits that are tentatively dated to the Late Cretaceous of North America, making it a potential survivor of the K/Pg extinction and the only record of a ‘condylarth’ in the Late Cretaceous ( +<bibRefCitation id="EF817BD2FFF10D08C77EFE7C887FFE63" author="Archibald" box="[124,333,426,450]" etAl="et al." firstAuthor="Archibald" pageId="23" pageNumber="24" pagination="153 - 61" refId="ref39581" refString="Archibald JD, Zhang Y, Harper T et al. Protungulatum, confirmed Cretaceous occurrence of an otherwise Paleocene eutherian (placental?) mammal. Journal of Mammalian Evolution 2011; 18: 153 - 61. https: // doi. org / 10.1007 / s 10914 - 011 - 9162 - 1" type="journal article" year="2011"> +Archibald +<emphasis id="B964DA31FFF10D08C7E4FE7D8827FE63" box="[230,277,426,450]" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +2011 +</bibRefCitation> +). +</paragraph> +<paragraph id="8BAF0623FFF10D08C78FFE1F8B4FFACF" blockId="23.[112,765,144,1797]" pageId="23" pageNumber="24"> +Considering the proposed phylogenetic affinities of North American mioclaenids with early SANUs (e.g., Muizon and Cifelli 2000), we included four Palaeocene taxa of this group: +<taxonomicName id="4C107DA0FFF10D08C773FDF188F1FD9E" authority="Cope, 1881" authorityName="Cope" authorityYear="1881" box="[113,451,551,575]" class="Mammalia" family="Mioclaenidae" genus="Mioclaenus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="turgidus"> +<emphasis id="B964DA31FFF10D08C773FDF18809FD9E" box="[113,315,551,575]" italics="true" pageId="23" pageNumber="24">Mioclaenus turgidus</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C644FDF188F1FD9E" author="Cope" box="[326,451,551,575]" firstAuthor="Cope" pageId="23" pageNumber="24" pagination="484 - 95" refId="ref41294" refString="Cope ED. On some Mammalia of the Lowest Eocene beds of New Mexico. Proceedings of the American Philosophical Society 1881; 19: 484 - 95." type="journal article" year="1881">Cope, 1881</bibRefCitation> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFF10D08C6D6FDF18997FDFE" authority="Cope, 1882" authorityName="Cope" authorityYear="1882" class="Mammalia" family="Mioclaenidae" genus="Protoselene" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="opisthacus"> +<emphasis id="B964DA31FFF10D08C6D6FDF18B80FD9E" box="[468,690,551,575]" italics="true" pageId="23" pageNumber="24">Protoselene opisthacus</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C5BFFDF18997FDFE" author="Cope" firstAuthor="Cope" pageId="23" pageNumber="24" pagination="979 - 93" refId="ref41323" refString="Cope ED. Some new forms from the Puerco Eocene. The American Naturalist 1882; 16: 979 - 93. https: // doi. org / 10.1086 / 273224" type="journal article" year="1882">Cope, 1882</bibRefCitation> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFF10D08C7ACFD918B08FDFE" authority="Cope, 1888" authorityName="Cope" authorityYear="1888" box="[174,570,583,607]" class="Mammalia" family="Mioclaenidae" genus="Choeroclaenus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="turgidunculus"> +<emphasis id="B964DA31FFF10D08C7ACFD9188F0FDFE" box="[174,450,583,607]" italics="true" pageId="23" pageNumber="24">Choeroclaenus turgidunculus</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C6C4FD918B08FDFE" author="Cope" box="[454,570,583,607]" firstAuthor="Cope" pageId="23" pageNumber="24" pagination="298 - 361" refId="ref41355" refString="Cope ED. Synopsis of the vertebrate fauna of the Puerco series. Transactions of the American Philosophical Society 1888; 16: 298 - 361. https: // doi. org / 10.2307 / 1005393" type="journal article" year="1888">Cope, 1888</bibRefCitation> +</taxonomicName> +, and +<taxonomicName id="4C107DA0FFF10D08C569FD918863FDDC" class="Mammalia" family="Mioclaenidae" genus="Promioclaenus" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFF10D08C569FD918BCAFDFE" box="[619,760,583,607]" italics="true" pageId="23" pageNumber="24">Promioclaenus</emphasis> +Trouessart, 1904 spp. +</taxonomicName> +, which includes +<taxonomicName id="4C107DA0FFF10D08C502FDB0883BFD3C" authority="Matthew, 1897" authorityName="Matthew" authorityYear="1897" class="Mammalia" family="Mioclaenidae" genus="Promioclaenus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="lemuroides"> +<emphasis id="B964DA31FFF10D08C502FDB08BCEFDDF" box="[512,764,614,638]" italics="true" pageId="23" pageNumber="24">Promioclaenus lemuroides</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C773FD53883BFD3C" author="Matthew" box="[113,265,645,669]" firstAuthor="Matthew" pageId="23" pageNumber="24" pagination="259 - 323" refId="ref44224" refString="Matthew WDA. Revision of the Puerco fauna. Bulletin of the American Museum of Natural History 1897; 9: 259 - 323." type="journal article" year="1897">Matthew, 1897</bibRefCitation> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFF10D08C637FD538BBDFD3C" authority="Cope, 1888" authorityName="Cope" authorityYear="1888" box="[309,655,645,669]" class="Mammalia" family="Mioclaenidae" genus="Promioclaenus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="acolytus"> +<emphasis id="B964DA31FFF10D08C637FD538B26FD3C" box="[309,532,645,669]" italics="true" pageId="23" pageNumber="24">Promioclaenus acolytus</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C51BFD538BBDFD3C" author="Cope" box="[537,655,645,669]" firstAuthor="Cope" pageId="23" pageNumber="24" pagination="298 - 361" refId="ref41355" refString="Cope ED. Synopsis of the vertebrate fauna of the Puerco series. Transactions of the American Philosophical Society 1888; 16: 298 - 361. https: // doi. org / 10.2307 / 1005393" type="journal article" year="1888">Cope, 1888</bibRefCitation> +</taxonomicName> +. Including the latter two species within +<taxonomicName id="4C107DA0FFF10D08C69DFD738B69FD1D" box="[415,603,677,701]" class="Mammalia" family="Mioclaenidae" genus="Promioclaenus" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFF10D08C69DFD738B1EFD1C" box="[415,556,677,701]" italics="true" pageId="23" pageNumber="24">Promioclaenus</emphasis> +spp. +</taxonomicName> +maximized the information of this genus, considering that both species have almost the same temporal distribution, and the species level differentiation is based solely on a 20% size difference (Williamson 1996). To represent the earliest SANUs, we included two Tiupampan (probably Early Palaeocene) kollpaniines +<taxonomicName id="4C107DA0FFF10D08C591FC9788E7FCD8" authority="Muizon and Marshall 1987 a" authorityName="Muizon and Marshall" authorityYear="1987" class="Mammalia" family="Mioclaenidae" genus="Molinodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="suarezi"> +<emphasis id="B964DA31FFF10D08C591FC978984FCD9" italics="true" pageId="23" pageNumber="24">Molinodus suarezi</emphasis> +Muizon and Marshall 1987a +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C6E6FCB78B02FCD8" box="[484,560,865,889]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 8A</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C538FCB78B6AFCD8" box="[570,600,865,889]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="23" pageNumber="24">9A</figureCitation> +) and +<taxonomicName id="4C107DA0FFF10D08C593FCB789A9FC36" class="Mammalia" family="Mioclaenidae" genus="Tiuclaenus" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFF10D08C593FCB78BCBFCD8" box="[657,761,865,889]" italics="true" pageId="23" pageNumber="24">Tiuclaenus</emphasis> +spp. +</taxonomicName> +Muizon and Marshall 1987b ( +<taxonomicName id="4C107DA0FFF10D08C6E0FC568870FC16" authority="Muizon and Marshall 1987 b" authorityName="Muizon and Marshall" authorityYear="1987" class="Mammalia" family="Mioclaenidae" genus="Tiuclaenus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="minutus"> +<emphasis id="B964DA31FFF10D08C6E0FC568B90FC39" box="[482,674,896,920]" italics="true" pageId="23" pageNumber="24">Tiuclaenus minutus</emphasis> +Muizon and Marshall 1987b +</taxonomicName> +, +<taxonomicName id="4C107DA0FFF10D08C652FC498BC7FC16" authority="Muizon & Cifelli, 2000" authorityName="Muizon & Cifelli" authorityYear="2000" box="[336,757,927,951]" class="Mammalia" family="Mioclaenidae" genus="Tiuclaenus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="cotasi"> +<emphasis id="B964DA31FFF10D08C652FC4988CAFC16" box="[336,504,927,951]" italics="true" pageId="23" pageNumber="24">Tiuclaenus cotasi</emphasis> +Muizon & Cifelli, 2000 +</taxonomicName> +, and +<taxonomicName id="4C107DA0FFF10D08C7A3FC698B5BFC76" authority="Muizon & Cifelli, 2000" authorityName="Muizon & Cifelli" authorityYear="2000" box="[161,617,959,983]" class="Mammalia" family="Mioclaenidae" genus="Tiuclaenus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="robustus"> +<emphasis id="B964DA31FFF10D08C7A3FC698854FC76" box="[161,358,959,983]" italics="true" pageId="23" pageNumber="24">Tiuclaenus robustus</emphasis> +Muizon & Cifelli, 2000 +</taxonomicName> +), which have been previously included in +<taxonomicName id="4C107DA0FFF10D08C69BFC088983FBB4" authority="(Muizon and Cifelli 2000)" baseAuthorityName="Muizon and Cifelli" baseAuthorityYear="2000" class="Mammalia" family="Mioclaenidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Mioclaenidae (Muizon and Cifelli 2000)</taxonomicName> +. Including three species of +<taxonomicName id="4C107DA0FFF10D08C6D8FC2B8B70FBB4" authorityName="de Muizon & Marshall" authorityYear="1987" box="[474,578,1021,1045]" class="Mammalia" family="Mioclaenidae" genus="Tiuclaenus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF10D08C6D8FC2B8B70FBB4" box="[474,578,1021,1045]" italics="true" pageId="23" pageNumber="24">Tiuclaenus</emphasis> +</taxonomicName> +under +<taxonomicName id="4C107DA0FFF10D08C590FC2B89A9FB95" class="Mammalia" family="Mioclaenidae" genus="Tiuclaenus" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFF10D08C590FC2B8BC8FBB4" box="[658,762,1021,1045]" italics="true" pageId="23" pageNumber="24">Tiuclaenus</emphasis> +spp. +</taxonomicName> +maximized the information of this genus, considering that they have the same temporal distribution and differ mostly in their size and relatively minor features in their dentition (Muizon and Cifelli 2000, Muizon +<emphasis id="B964DA31FFF10D08C67CFBAD889DFB32" box="[382,431,1147,1171]" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +2019). In addition, considering the previously suggested close affinity between the ‘condylarth’ South American family +<taxonomicName id="4C107DA0FFF10D08C676FB6F8B2AFB70" box="[372,536,1209,1233]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Didolodontidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFF10D08C551FB6F8BF2FB70" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[595,704,1209,1233]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="order">Litopterna</taxonomicName> +(e.g., Simpson 1948), we included four taxa of this family: the Early Palaeocene +<taxonomicName id="4C107DA0FFF10D08C7E9FB2E88FAFAB1" box="[235,456,1272,1296]" class="Mammalia" family="Mioclaenidae" genus="Escribania" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="chubutensis"> +<emphasis id="B964DA31FFF10D08C7E9FB2E88FAFAB1" box="[235,456,1272,1296]" italics="true" pageId="23" pageNumber="24">Escribania chubutensis</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C6D8FB2E8B1AFAB1" box="[474,552,1272,1296]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 8B</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C536FB2E8B62FAB1" box="[564,592,1272,1296]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="23" pageNumber="24">9B</figureCitation> +) and +<taxonomicName id="4C107DA0FFF10D08C590FB2E89E1FA8E" class="Mammalia" family="Mioclaenidae" genus="Raulvaccia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="peligrensis"> +<emphasis id="B964DA31FFF10D08C590FB2E89E1FA8E" italics="true" pageId="23" pageNumber="24">Raulvaccia peligrensis</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C7E1FAC18862FA8E" box="[227,336,1303,1327]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 8C–D</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C658FAC1884AFA8E" box="[346,376,1303,1327]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="23" pageNumber="24">9C</figureCitation> +) from Punta Peligro, the Early Eocene +<taxonomicName id="4C107DA0FFF10D08C772FAE18843FAEF" baseAuthorityName="Mones" baseAuthorityYear="2015" box="[112,369,1335,1359]" class="Mammalia" family="Didolodontidae" genus="Ricardocifellia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="protocenica"> +<emphasis id="B964DA31FFF10D08C772FAE18843FAEF" box="[112,369,1335,1359]" italics="true" pageId="23" pageNumber="24">Ricardocifellia protocenica</emphasis> +</taxonomicName> +from Itaboraí ( +<figureCitation id="132B1AA6FFF10D08C515FAE18B57FAEE" box="[535,613,1335,1359]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 8E</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C571FAE18BA0FAEE" box="[627,658,1335,1359]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="23" pageNumber="24">9D</figureCitation> +), and the Middle Eocene +<taxonomicName id="4C107DA0FFF10D08C610FA8088DAFACF" authorityName="Ameghino" authorityYear="1897" box="[274,488,1366,1390]" class="Mammalia" family="Didolodontidae" genus="Didolodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="multicuspis"> +<emphasis id="B964DA31FFF10D08C610FA8088DAFACF" box="[274,488,1366,1390]" italics="true" pageId="23" pageNumber="24">Didolodus multicuspis</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C6FAFA808B74FACF" box="[504,582,1366,1390]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 8G</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C553FA808B5FFACF" box="[593,621,1366,1390]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="23" pageNumber="24">9E</figureCitation> +). +</paragraph> +<paragraph id="8BAF0623FFF10D08C78FFAA08C20FD3C" blockId="23.[112,765,144,1797]" lastBlockId="23.[810,1460,144,1797]" pageId="23" pageNumber="24"> +Because of the molecular evidence supporting close affinities between litopterns and notoungulates ( +<bibRefCitation id="EF817BD2FFF10D08C511FA438B91FA0D" author="Buckley" box="[531,675,1428,1453]" firstAuthor="Buckley" pageId="23" pageNumber="24" refId="ref40554" refString="Buckley M. Ancient collagen reveals evolutionary history of the endemic South American ' ungulates'. Proceedings of the Royal Society B: Biological Sciences 2015; 282: 20142671. https: // doi. org / 10.1098 / rspb. 2014.2671" type="journal volume" year="2015">Buckley 2015</bibRefCitation> +, Welker +<emphasis id="B964DA31FFF10D08C773FA638996FA6D" box="[113,164,1460,1484]" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +2015), and the possibility of a common origin with the other SANU orders ( +<taxonomicName id="4C107DA0FFF10D08C657FA0588DAFA4A" baseAuthorityName="Lorente" baseAuthorityYear="2015" box="[341,488,1491,1515]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="23" pageNumber="24" phylum="Chordata" rank="order">Astrapotheria</taxonomicName> +, +<taxonomicName id="4C107DA0FFF10D08C6F7FA058B4BFA4A" authorityName="Paula Couto" authorityYear="1952" box="[501,633,1491,1515]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Xenungulata" pageId="23" pageNumber="24" phylum="Chordata" rank="order">Xenungulata</taxonomicName> +, +<taxonomicName id="4C107DA0FFF10D08C585FA0588F0F9AB" authority=", Meridiungulata McKenna, 1975" authorityName=", Meridiungulata McKenna" authorityYear="1975" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pyrotheria" pageId="23" pageNumber="24" phylum="Chordata" rank="order"> +Pyrotheria, Meridiungulata +<bibRefCitation id="EF817BD2FFF10D08C61BFA2588F0F9AB" author="McKenna" box="[281,450,1522,1546]" firstAuthor="McKenna" pageId="23" pageNumber="24" pagination="21 - 46" refId="ref44533" refString="McKenna MC. Toward a phylogenetic classification of the Mammalia. In: Luckett WP, Szalay FS (eds), Phylogeny of the Primates. New York, NY: Plenum Press, 1975, 21 - 46." type="book chapter" year="1975">McKenna, 1975</bibRefCitation> +</taxonomicName> +), we included the Middle to Early Palaeocene notoungulate +<taxonomicName id="4C107DA0FFF10D08C6BEF9C489D9F9E8" authority="Pascual et al., 1978" authorityName="Pascual" authorityYear="1978" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="praecursor"> +<emphasis id="B964DA31FFF10D08C6BEF9C48B97F98B" box="[444,677,1554,1578]" italics="true" pageId="23" pageNumber="24">Simpsonotus praecursor</emphasis> +Pascual +<emphasis id="B964DA31FFF10D08C773F9E48995F9E8" box="[113,167,1585,1609]" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +, 1978 +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C603F9E78861F9E8" box="[257,339,1585,1609]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 6E</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C666F9E788B1F9E8" box="[356,387,1585,1609]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="23" pageNumber="24">7C</figureCitation> +), the Middle Eocene astrapothere +<taxonomicName id="4C107DA0FFF10D08C773F9878B10F9C8" authority="Ameghino, 1897" authorityName="Ameghino" authorityYear="1897" box="[113,546,1617,1641]" class="Mammalia" family="Trigonostylopidae" genus="Trigonostylops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="wortmani"> +<emphasis id="B964DA31FFF10D08C773F987885AF9C9" box="[113,360,1617,1641]" italics="true" pageId="23" pageNumber="24">Trigonostylops wortmani</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C671F9878B10F9C8" author="Ameghino" box="[371,546,1617,1641]" firstAuthor="Ameghino" pageId="23" pageNumber="24" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino, 1897</bibRefCitation> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C53AF9878BBFF9C8" box="[568,653,1617,1641]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 6H</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C59CF9878B8FF9C9" box="[670,701,1617,1640]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="23" pageNumber="24">7D</figureCitation> +), the xenungulates +<taxonomicName id="4C107DA0FFF10D08C603F9A78B6DF929" authority="Villarroel, 1987" authorityName="Villarroel" authorityYear="1987" box="[257,607,1648,1672]" class="Arthoniomycetes" family="Phaeococcomycetaceae" genus="Etayoa" kingdom="Fungi" order="Lichenostigmatales" pageId="23" pageNumber="24" phylum="Ascomycota" rank="species" species="bacatensis"> +<emphasis id="B964DA31FFF10D08C603F9A78883F929" box="[257,433,1648,1672]" italics="true" pageId="23" pageNumber="24">Etayoa bacatensis</emphasis> +Villarroel, 1987 +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C576F9A68B8DF926" box="[628,703,1648,1672]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="23" pageNumber="24">Fig. 7E</figureCitation> +) and +<taxonomicName id="4C107DA0FFF10D08C773F95988DDF906" authority="Paula Couto, 1952" authorityName="Paula Couto" authorityYear="1952" box="[113,495,1679,1703]" class="Mammalia" family="Carodniidae" genus="Carodnia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Xenungulata" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="vieirai"> +<emphasis id="B964DA31FFF10D08C773F9598825F906" box="[113,279,1679,1703]" italics="true" pageId="23" pageNumber="24">Carodnia vieirai</emphasis> +Paula Couto, 1952 +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C504F9598B7DF906" box="[518,591,1679,1703]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Fig. 6I</figureCitation> +), and the Late Eocene pyrothere +<taxonomicName id="4C107DA0FFF10D08C643F9798BC8F966" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[321,762,1711,1735]" class="Mammalia" family="Pyrotheriidae" genus="Propyrotherium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="saxeum"> +<emphasis id="B964DA31FFF10D08C643F9798B04F967" box="[321,566,1711,1735]" italics="true" pageId="23" pageNumber="24">Propyrotherium saxeum</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C544F9798BC8F966" author="Ameghino" box="[582,762,1711,1735]" firstAuthor="Ameghino" pageId="23" pageNumber="24" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C77EF91889F7F947" box="[124,197,1742,1766]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 6J</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C7D7F91889C7F947" box="[213,245,1742,1766]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="23" pageNumber="24">7G</figureCitation> +). Other potentially early members of the order +<taxonomicName id="4C107DA0FFF10D08C773F93B89ECF8A4" authorityName=", Meridiungulata McKenna" authorityYear="1975" box="[113,222,1773,1797]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pyrotheria" pageId="23" pageNumber="24" phylum="Chordata" rank="order">Pyrotheria</taxonomicName> +such as +<taxonomicName id="4C107DA0FFF10D08C634F9388B15F8A4" authority="Patterson, 1977" authorityName="Patterson" authorityYear="1977" box="[310,551,1773,1797]" class="Mammalia" family="Pyrotheriidae" genus="Proticia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF10D08C634F93888B0F8A4" box="[310,386,1774,1797]" italics="true" pageId="23" pageNumber="24">Proticia</emphasis> +Patterson, 1977 +</taxonomicName> +and +<taxonomicName id="4C107DA0FFF10D08C559F93B8AECFF09" authority="Hoffstetter, 1970" authorityName="Hoffstetter" authorityYear="1970" box="[603,990,144,1797]" class="Mammalia" family="Pyrotheriidae" genus="Colombitherium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF10D08C559F93B8BC9F8A4" box="[603,763,1773,1797]" italics="true" pageId="23" pageNumber="24">Colombitherium</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C428FF468AECFF09" author="Hoffstetter" box="[810,990,144,168]" firstAuthor="Hoffstetter" pageId="23" pageNumber="24" pagination="149 - 69" refId="ref43200" refString="Hoffstetter R. Colombitherium tolimense, pyrotherien nouveau de la formation Gualanday (Colombie). Annales de Paleontologie, Vertebres 1970; 56: 149 - 69." type="journal article" year="1970">Hoffstetter, 1970</bibRefCitation> +</taxonomicName> +were not included, as their assignment to this order has been challenged (Sánchez-Villagra +<emphasis id="B964DA31FFF10D08C23EFF668C42FF66" box="[1340,1392,175,199]" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +2006, +<bibRefCitation id="EF817BD2FFF10D08C428FF198AFDFF46" author="Billet" box="[810,975,207,231]" etAl="et al." firstAuthor="Billet" pageId="23" pageNumber="24" pagination="319 - 25" refId="ref40038" refString="Billet G, Orliac M, Antoine PO et al. New observations and reinterpretation on the enigmatic taxon Colombitherium (? Pyrotheria, Mammalia) from Colombia. Palaeontology 2010; 53: 319 - 25. https: // doi. org / 10.1111 / j. 1475 - 4983.2010.00936. x" type="journal article" year="2010"> +Billet +<emphasis id="B964DA31FFF10D08C465FF198AA4FF46" box="[871,918,207,231]" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +2010 +</bibRefCitation> +). We also included the Early Eocene indaleciid +<taxonomicName id="4C107DA0FFF10D08C429FF388DDFFEA7" authority="Bond & Vucetich, 1983" authorityName="Bond & Vucetich" authorityYear="1983" box="[811,1261,238,262]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="grandensis"> +<emphasis id="B964DA31FFF10D08C429FF388AC3FEA7" box="[811,1009,238,262]" italics="true" pageId="23" pageNumber="24">Indalecia grandensis</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C4F5FF388DDFFEA7" author="Bond & Vucetich" box="[1015,1261,238,262]" firstAuthor="Bond" pageId="23" pageNumber="24" pagination="107 - 17" refId="ref40299" refString="Bond M, Vucetich MG. Indalecia grandensis gen. et sp. nov. del Eoceno temprano del noroeste argentino, tipo de una nueva subfamilia de los Adianthidae. Revista de la Asociacion Geologica Argentina 1983; 38: 107 - 17." type="journal article" year="1983">Bond & Vucetich, 1983</bibRefCitation> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C3FDFF388C7CFEA7" box="[1279,1358,238,262]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 6A</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C25EFF388C48FEA7" box="[1372,1402,238,262]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="23" pageNumber="24">7A</figureCitation> +) and two notonychopids +<taxonomicName id="4C107DA0FFF10D08C4FEFEDB8C7EFE84" authority="Soria 1989 b" authorityName="Soria" authorityYear="1989" box="[1020,1356,269,293]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="powelli"> +<emphasis id="B964DA31FFF10D08C4FEFEDB8DF4FE84" box="[1020,1222,269,293]" italics="true" pageId="23" pageNumber="24">Notonychops powelli</emphasis> +Soria 1989b +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C25DFEDB8C9FFE84" box="[1375,1453,269,293]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 6B</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C428FEFB8A74FEE5" box="[810,838,301,324]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="23" pageNumber="24">7B</figureCitation> +; probably Early Eocene) and +<taxonomicName id="4C107DA0FFF10D08C383FEFB8AF1FEC5" authority="Bonaparte & Morales, 1997" authorityName="Bonaparte & Morales" authorityYear="1997" class="Mammalia" family="Notonychopidae" genus="Requisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="vidmari"> +<emphasis id="B964DA31FFF10D08C383FEFB8C1AFEE4" box="[1153,1320,301,325]" italics="true" pageId="23" pageNumber="24">Requisia vidmari</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C22CFEFB8AF1FEC5" author="Bonaparte & Morales" firstAuthor="Bonaparte" pageId="23" pageNumber="24" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte & Morales, 1997</bibRefCitation> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C4D4FE9A8D17FEC5" box="[982,1061,332,356]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Fig. 6C</figureCitation> +; Early Palaeocene), members of two families (i.e., +<taxonomicName id="4C107DA0FFF10D08C4C3FEBD8D0FFE22" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[961,1085,363,387]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Indaleciidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFF10D08C382FEBD8C02FE22" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[1152,1328,363,387]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Notonychopidae</taxonomicName> +) sometimes included within +<taxonomicName id="4C107DA0FFF10D08C4D5FE5D8D76FE02" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[983,1092,395,419]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="order">Litopterna</taxonomicName> +(e.g., Croft +<emphasis id="B964DA31FFF10D08C3C7FE5D8DC5FE02" box="[1221,1271,395,419]" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +2020) and sometimes included in a separate order, +<taxonomicName id="4C107DA0FFF10D08C39EFE7C8C9CFE63" authority="(Soria 1989 a)" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[1180,1454,426,450]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="order">Notopterna (Soria 1989a)</taxonomicName> +. Members of +<taxonomicName id="4C107DA0FFF10D08C4AEFE1F8D40FE40" box="[940,1138,457,481]" class="Mammalia" family="Amilnedwardsiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Amilnedwardsiidae</taxonomicName> +were not included because this family is only known from a few isolated upper molars whose positions are tentative (Soria 1989a, Saade +<emphasis id="B964DA31FFF10D08C3DEFDDF8C38FD81" box="[1244,1290,520,544]" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +2023) and some of the specimens from the MACN collections are lost (Saade +<emphasis id="B964DA31FFF10D08C2A0FDFE8A71FDFE" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +2023). In addition, we included the potentially Palaeocene SANU +<taxonomicName id="4C107DA0FFF10D08C476FDB08C63FDDF" authority="Soria & Powell, 1981" authorityName="Soria & Powell" authorityYear="1981" box="[884,1361,614,638]" class="Mammalia" family="Eoastrapostylopidae" genus="Eoastrapostylops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="riolorense"> +<emphasis id="B964DA31FFF10D08C476FDB08D4BFDDF" box="[884,1145,614,638]" italics="true" pageId="23" pageNumber="24">Eoastrapostylops riolorense</emphasis> +Soria & Powell, 1981 +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C262FDB08C9FFDDF" box="[1376,1453,614,638]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 6D</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C428FD508A77FD3C" box="[810,837,646,669]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="23" pageNumber="24">7F</figureCitation> +) of uncertain affinities ( +<bibRefCitation id="EF817BD2FFF10D08C33FFD508C33FD3C" author="Kramarz" box="[1085,1281,645,669]" etAl="et al." firstAuthor="Kramarz" pageId="23" pageNumber="24" pagination="95 - 164" refId="ref43659" refString="Kramarz AG, Bond M, Rougier GW. Re-description of the auditory region of the putative basal astrapothere (Mammalia) Eoastrapostylops riolorense Soria and Powell, 1981. Systematic and phylogenetic considerations. Annals of Carnegie Museum 2017; 84: 95 - 164. https: // doi. org / 10.2992 / 007.084.0204" type="journal article" year="2017"> +Kramarz +<emphasis id="B964DA31FFF10D08C399FD508DFBFD3C" box="[1179,1225,645,669]" italics="true" pageId="23" pageNumber="24">et al.</emphasis> +2017 +</bibRefCitation> +). +</paragraph> +<paragraph id="8BAF0623FFF10D08C447FD738C81F8A4" blockId="23.[810,1460,144,1797]" pageId="23" pageNumber="24"> +We included three members of the family +<taxonomicName id="4C107DA0FFF10D08C206FD738C82FD1C" box="[1284,1456,677,701]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Protolipternidae</taxonomicName> +: the probably Early Eocene +<taxonomicName id="4C107DA0FFF10D08C381FD128C81FD7D" box="[1155,1459,708,732]" class="Mammalia" family="Protolipternidae" genus="Protolipterna" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="ellipsodontoides"> +<emphasis id="B964DA31FFF10D08C381FD128C81FD7D" box="[1155,1459,708,732]" italics="true" pageId="23" pageNumber="24">Protolipterna ellipsodontoides</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C437FD358ABFFD5A" box="[821,909,739,763]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 8H</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C4A3FD358A8EFD5A" box="[929,956,739,763]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="23" pageNumber="24">9F</figureCitation> +), +<taxonomicName id="4C107DA0FFF10D08C4D8FD358DE4FD5A" authorityName="Paula Couto" authorityYear="1952" box="[986,1238,739,763]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="scotti"> +<emphasis id="B964DA31FFF10D08C4D8FD358DE4FD5A" box="[986,1238,739,763]" italics="true" pageId="23" pageNumber="24">Asmithwoodwardia scotti</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C3EDFD358C09FD5A" box="[1263,1339,739,763]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 8I</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C24DFD358C5CFD5A" box="[1359,1390,739,763]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="23" pageNumber="24">9G</figureCitation> +), and +<taxonomicName id="4C107DA0FFF10D08C428FCD58D1AFCBA" baseAuthorityName="Paula Couto" baseAuthorityYear="1952" box="[810,1064,771,795]" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="parayiruhnor"> +<emphasis id="B964DA31FFF10D08C428FCD58D1AFCBA" box="[810,1064,771,795]" italics="true" pageId="23" pageNumber="24">Miguelsoria parayiruhnor</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C33FFCD58DB4FCBA" box="[1085,1158,771,795]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 8J</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C394FCD58D85FCBA" box="[1174,1207,771,795]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="23" pageNumber="24">9H</figureCitation> +), usually placed within +<taxonomicName id="4C107DA0FFF10D08C428FCF48AA5FC9B" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[810,919,802,826]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="order">Litopterna</taxonomicName> +based on tarsal features ( +<bibRefCitation id="EF817BD2FFF10D08C3C0FCF48C71FC9B" author="Cifelli" box="[1218,1347,802,826]" firstAuthor="Cifelli" pageId="23" pageNumber="24" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation> +). In addition, we included members of the five relatively less controversial litoptern families (i.e., +<taxonomicName id="4C107DA0FFF10D08C36DFCB78DDCFCD8" box="[1135,1262,865,889]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +, +<taxonomicName id="4C107DA0FFF10D08C3FCFCB78C82FCD8" box="[1278,1456,865,889]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +, +<taxonomicName id="4C107DA0FFF10D08C428FC568AE6FC39" baseAuthorityName="Simpson" baseAuthorityYear="1948" box="[810,980,896,920]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +, +<taxonomicName id="4C107DA0FFF10D08C4DDFC568DBAFC39" authorityName=", Soria" authorityYear="2001" box="[991,1160,896,920]" family="Anisolambdidae" pageId="23" pageNumber="24" rank="family">Anisolambdidae</taxonomicName> +, and +<taxonomicName id="4C107DA0FFF10D08C3C3FC568C9BFC39" box="[1217,1449,896,920]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +). From +<taxonomicName id="4C107DA0FFF10D08C468FC498ADBFC16" box="[874,1001,927,951]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Adianthidae</taxonomicName> +, we included three taxa: the Middle Eocene +<taxonomicName id="4C107DA0FFF10D08C42EFC698DF8FC76" authority="Ameghino 1904 b" authorityName="Ameghino" authorityYear="1904" box="[812,1226,959,983]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="argentinus"> +<emphasis id="B964DA31FFF10D08C42EFC698D38FC77" box="[812,1034,959,982]" italics="true" pageId="23" pageNumber="24">Proectocion argentinus</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C311FC698DF8FC76" author="Ameghino" box="[1043,1226,959,983]" firstAuthor="Ameghino" pageId="23" pageNumber="24" pagination="1 - 541" refId="ref39426" refString="Ameghino F. Recherches de morphologic phylogenetique sur les molaires superieures des ongules. Anales del Museo Nacional de Buenos Aires 1904 b; 3: 1 - 541." type="journal article" year="1904" yearSuffix="b">Ameghino 1904b</bibRefCitation> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C3D8FC698C79FC77" box="[1242,1355,959,983]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 4F, H</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C25EFC688C45FC77" box="[1372,1399,958,982]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="23" pageNumber="24">5E</figureCitation> +), the Late Oligocene +<taxonomicName id="4C107DA0FFF10D08C4D3FC088C44FC57" authority="Ameghino, 1897" authorityName="Ameghino" authorityYear="1897" box="[977,1398,990,1014]" class="Mammalia" family="Adianthidae" genus="Tricoelodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="bicuspidatus"> +<emphasis id="B964DA31FFF10D08C4D3FC088DF0FC57" box="[977,1218,990,1014]" italics="true" pageId="23" pageNumber="24">Tricoelodus bicuspidatus</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C3C8FC088C44FC57" author="Ameghino" box="[1226,1398,990,1014]" firstAuthor="Ameghino" pageId="23" pageNumber="24" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino, 1897</bibRefCitation> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C28BFC088A78FBB4" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 4G</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C45CFC2B8A4BFBB4" box="[862,889,1021,1045]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="23" pageNumber="24">5F</figureCitation> +), and +<taxonomicName id="4C107DA0FFF10D08C4CCFC2B8C46FBB4" authority="Ameghino, 1897" authorityName="Ameghino" authorityYear="1897" box="[974,1396,1021,1045]" class="Mammalia" family="Adianthidae" genus="Proadiantus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="excavatus"> +<emphasis id="B964DA31FFF10D08C4CCFC2B8D83FBB4" box="[974,1201,1021,1045]" italics="true" pageId="23" pageNumber="24">Proadiantus excavatus</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C3C2FC2B8C46FBB4" author="Ameghino" box="[1216,1396,1021,1045]" firstAuthor="Ameghino" pageId="23" pageNumber="24" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino, 1897</bibRefCitation> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C28FFC2B8A78FB95" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="23" pageNumber="24">Fig. 5G</figureCitation> +). We considered +<taxonomicName id="4C107DA0FFF10D08C30EFBCB8CA6FB94" authority="Ameghino 1904 b" authorityName="Ameghino" authorityYear="1904" box="[1036,1428,1053,1077]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="precisus"> +<emphasis id="B964DA31FFF10D08C30EFBCB8DE3FB95" box="[1036,1233,1053,1076]" italics="true" pageId="23" pageNumber="24">Proectocion precisus</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C3DEFBCB8CA6FB94" author="Ameghino" box="[1244,1428,1053,1077]" firstAuthor="Ameghino" pageId="23" pageNumber="24" pagination="1 - 541" refId="ref39426" refString="Ameghino F. Recherches de morphologic phylogenetique sur les molaires superieures des ongules. Anales del Museo Nacional de Buenos Aires 1904 b; 3: 1 - 541." type="journal article" year="1904" yearSuffix="b">Ameghino 1904b</bibRefCitation> +</taxonomicName> +as a junior synonym of +<taxonomicName id="4C107DA0FFF10D08C315FBEB8DC5FBF5" authorityName="Ameghino" authorityYear="1904" box="[1047,1271,1085,1108]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="argentinus"> +<emphasis id="B964DA31FFF10D08C315FBEB8DC5FBF5" box="[1047,1271,1085,1108]" italics="true" pageId="23" pageNumber="24">Proectocion argentinus</emphasis> +</taxonomicName> +(see justification in Supporting information, File S2). From +<taxonomicName id="4C107DA0FFF10D08C3FCFB8D8C82FBD2" box="[1278,1456,1115,1139]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +, we included four taxa: the Late Eocene +<taxonomicName id="4C107DA0FFF10D08C3CFFBAD8C4AFB33" box="[1229,1400,1147,1171]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFF10D08C3CFFBAD8C75FB32" box="[1229,1351,1147,1171]" italics="true" pageId="23" pageNumber="24">Polymorphis</emphasis> +spp. +</taxonomicName> +[ +<figureCitation id="132B1AA6FFF10D08C28BFBAD8A7AFB13" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 4A</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C455FB4C8A47FB13" box="[855,885,1178,1202]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="23" pageNumber="24">5A</figureCitation> +; +<taxonomicName id="4C107DA0FFF10D08C487FB4C8D89FB13" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" box="[901,1211,1178,1202]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="lechei"> +<emphasis id="B964DA31FFF10D08C487FB4C8D0EFB13" box="[901,1084,1178,1202]" italics="true" pageId="23" pageNumber="24">Polymorphis lechei</emphasis> +Roth, 1899 +</taxonomicName> +and +<taxonomicName id="4C107DA0FFF10D08C3F0FB4C8A82FB70" authority="(Roth 1899)" authorityName="Püschel & Shelley & Williamson & Perini & Wible & Brusatte" authorityYear="2024" baseAuthorityName="Roth" baseAuthorityYear="1899" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="ligatus" status="comb. nov."> +<emphasis id="B964DA31FFF10D08C3F0FB4C8C81FB13" box="[1266,1459,1178,1202]" italics="true" pageId="23" pageNumber="24">Polymorphis ligatus</emphasis> +(Roth 1899) +</taxonomicName> +<taxonomicNameLabel id="A257674AFFF10D08C4BBFB6F8D19FB70" box="[953,1067,1209,1233]" pageId="23" pageNumber="24" rank="species">comb. nov.</taxonomicNameLabel> +(see Supporting information, File S2 for more details of this assignment)], the Late Oligocene to Early Miocene +<taxonomicName id="4C107DA0FFF10D08C4CEFB2E8C47FAB1" authority="Ameghino 1902 a" authorityName="Ameghino" authorityYear="1902" box="[972,1397,1272,1296]" class="Mammalia" family="Macraucheniidae" genus="Cramauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="normalis"> +<emphasis id="B964DA31FFF10D08C4CEFB2E8D8BFAB1" box="[972,1209,1272,1296]" italics="true" pageId="23" pageNumber="24">Cramauchenia normalis</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C3C3FB2E8C47FAB1" author="Ameghino" box="[1217,1397,1272,1296]" firstAuthor="Ameghino" pageId="23" pageNumber="24" pagination="71 - 138" refId="ref39328" refString="Ameghino F. Premiere contribution a la connaissance de la faune mammalogique des couches a Colpodon. Boletin de la Academia Nacional de Ciencias en Cordoba 1902 a; 17: 71 - 138." type="journal article" year="1902" yearSuffix="a">Ameghino 1902a</bibRefCitation> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C28BFB2E8A7BFA8E" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 4D</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C45BFAC18AA4FA8E" box="[857,918,1303,1327]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="23" pageNumber="24">5B–C</figureCitation> +), the Early Miocene +<taxonomicName id="4C107DA0FFF10D08C37CFAC18C2AFA8E" box="[1150,1304,1303,1327]" class="Mammalia" family="Macraucheniidae" genus="Theosodon" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFF10D08C37CFAC18DD7FA8E" box="[1150,1253,1303,1327]" italics="true" pageId="23" pageNumber="24">Theosodon</emphasis> +spp. +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C22EFAC18C4EFA8E" box="[1324,1404,1303,1327]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 4E</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C289FAC18C99FA8E" box="[1419,1451,1303,1327]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="23" pageNumber="24">5D</figureCitation> +; +<taxonomicName id="4C107DA0FFF10D08C428FAE18D88FAEE" authority="Ameghino, 1887" authorityName="Ameghino" authorityYear="1887" box="[810,1210,1335,1359]" class="Mammalia" family="Macraucheniidae" genus="Theosodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="lydekkeri"> +<emphasis id="B964DA31FFF10D08C428FAE18ACAFAEE" box="[810,1016,1335,1359]" italics="true" pageId="23" pageNumber="24">Theosodon lydekkeri</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C305FAE18D88FAEE" author="Ameghino" box="[1031,1210,1335,1359]" firstAuthor="Ameghino" pageId="23" pageNumber="24" pagination="1 - 26" refId="ref39043" refString="Ameghino F. Enumeracion sistematica de las especies de mamiferos fosiles coleccionados por Carlos Ameghino en los terrenos Eocenos de la Patagonia Austral y depositados en el Museo de La Plata. Boletin del Museo de La Plata 1887; 1: 1 - 26." type="journal article" year="1887">Ameghino, 1887</bibRefCitation> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFF10D08C3CDFAE18A90FACF" authority="Scott, 1910" authorityName="Scott" authorityYear="1910" class="Mammalia" family="Macraucheniidae" genus="Theosodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="garretorum"> +<emphasis id="B964DA31FFF10D08C3CDFAE18C81FAEF" box="[1231,1459,1335,1359]" italics="true" pageId="23" pageNumber="24">Theosodon garretorum</emphasis> +Scott, 1910 +</taxonomicName> +, and +<taxonomicName id="4C107DA0FFF10D08C4E2FA808C61FACF" authority="Ameghino 1891 b" authorityName="Ameghino" authorityYear="1891" box="[992,1363,1366,1390]" class="Mammalia" family="Macraucheniidae" genus="Theosodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="gracilis"> +<emphasis id="B964DA31FFF10D08C4E2FA808DA7FACF" box="[992,1173,1366,1390]" italics="true" pageId="23" pageNumber="24">Theosodon gracilis</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C39CFA808C61FACF" author="Ameghino" box="[1182,1363,1366,1390]" firstAuthor="Ameghino" pageId="23" pageNumber="24" pagination="289 - 328" refId="ref39153" refString="Ameghino F. Nuevos restos de mamiferos fosiles descubiertos por Carlos Ameghino en el Eoceno inferior de la Patagonia austral. - Especies nuevas, adiciones y correcciones. Revista Argentina de Historia Natural 1891 b; 1: 289 - 328." type="journal article" year="1891" yearSuffix="b">Ameghino 1891b</bibRefCitation> +</taxonomicName> +, and the Late Pleistocene to Holocene +<taxonomicName id="4C107DA0FFF10D08C377FAA38CB2FA2C" authorityName="Owen" authorityYear="1838" box="[1141,1408,1397,1421]" class="Mammalia" family="Macraucheniidae" genus="Macrauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="patachonica"> +<emphasis id="B964DA31FFF10D08C377FAA38CB2FA2C" box="[1141,1408,1397,1421]" italics="true" pageId="23" pageNumber="24">Macrauchenia patachonica</emphasis> +</taxonomicName> +. We used three species to represent the genus +<taxonomicName id="4C107DA0FFF10D08C3E8FA438C63FA0C" authorityName="Ameghino" authorityYear="1887" box="[1258,1361,1429,1453]" class="Mammalia" family="Macraucheniidae" genus="Theosodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFF10D08C3E8FA438C63FA0C" box="[1258,1361,1429,1453]" italics="true" pageId="23" pageNumber="24">Theosodon</emphasis> +</taxonomicName> +to maximize the information of the genus, considering that these are all species of the same temporal and geographical distribution and that present relatively minor differences in size and anatomical features (Scott 1910). From +<taxonomicName id="4C107DA0FFF10D08C3BAF9C48C50F98B" baseAuthorityName="Simpson" baseAuthorityYear="1948" box="[1208,1378,1554,1578]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName> +, we included two taxa: the Late Oligocene +<taxonomicName id="4C107DA0FFF10D08C3AFF9E78C4BF9E8" authorityName="Ameghino" authorityYear="1897" box="[1197,1401,1585,1609]" class="Mammalia" family="Proterotheriidae" genus="Lambdaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="suinus"> +<emphasis id="B964DA31FFF10D08C3AFF9E78C4BF9E8" box="[1197,1401,1585,1609]" italics="true" pageId="23" pageNumber="24">Lambdaconus suinus</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C28BF9E78A56F9C9" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 4I, K</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C474F9868AA5F9C9" box="[886,919,1616,1640]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="23" pageNumber="24">5H</figureCitation> +) and the Early Miocene +<taxonomicName id="4C107DA0FFF10D08C3B7F9878C81F9C8" baseAuthorityName="Ameghino" baseAuthorityYear="1887" box="[1205,1459,1617,1641]" class="Mammalia" family="Proterotheriidae" genus="Diadiaphorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="majusculus"> +<emphasis id="B964DA31FFF10D08C3B7F9878C81F9C8" box="[1205,1459,1617,1641]" italics="true" pageId="23" pageNumber="24">Diadiaphorus majusculus</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C437F9A68A4BF929" box="[821,889,1648,1672]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 4J</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C486F9A68AABF929" box="[900,921,1648,1672]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="23" pageNumber="24">5I</figureCitation> +). From +<taxonomicName id="4C107DA0FFF10D08C4EFF9A68DA4F929" authorityName=", Soria" authorityYear="2001" box="[1005,1174,1648,1672]" family="Anisolambdidae" pageId="23" pageNumber="24" rank="family">Anisolambdidae</taxonomicName> +, we included three taxa: the Early Palaeocene +<taxonomicName id="4C107DA0FFF10D08C4E3F9598DB0F906" authorityName="Simpson" authorityYear="1935" box="[993,1154,1679,1703]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="tshotshe"> +<emphasis id="B964DA31FFF10D08C4E3F9598DB0F906" box="[993,1154,1679,1703]" italics="true" pageId="23" pageNumber="24">Wainka tshotshe</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C396F9598DE9F906" box="[1172,1243,1679,1703]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="23" pageNumber="24">Fig.5N</figureCitation> +), the probably Early Eocene +<taxonomicName id="4C107DA0FFF10D08C479F9798DBBF966" authorityName="Cifelli" authorityYear="1983" box="[891,1161,1711,1735]" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="prodromus"> +<emphasis id="B964DA31FFF10D08C479F9798DBBF966" box="[891,1161,1711,1735]" italics="true" pageId="23" pageNumber="24">Paranisolambda prodromus</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C39BF9798DDEF966" box="[1177,1260,1711,1735]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 4M</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C3F5F9788C27F967" box="[1271,1301,1710,1734]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="23" pageNumber="24">5K</figureCitation> +), and the Early to Middle Eocene +<taxonomicName id="4C107DA0FFF10D08C4F3F9188D96F944" box="[1009,1188,1742,1766]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFF10D08C4F3F9188D40F947" box="[1009,1138,1742,1766]" italics="true" pageId="23" pageNumber="24">Anisolambda</emphasis> +spp. +</taxonomicName> +( +<figureCitation id="132B1AA6FFF10D08C3B5F9188C3BF947" box="[1207,1289,1742,1766]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="23" pageNumber="24">Figs 4N</figureCitation> +, +<figureCitation id="132B1AA6FFF10D08C21AF9188C1FF947" box="[1304,1325,1742,1766]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="23" pageNumber="24">5J</figureCitation> +; +<emphasis id="B964DA31FFF10D08C23CF9188C81F947" box="[1342,1459,1742,1766]" italics="true" pageId="23" pageNumber="24">Anisolamda</emphasis> +<taxonomicName id="4C107DA0FFF10D08C428F93B8D1AF8A4" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[810,1064,1773,1797]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="fissidens"> +<emphasis id="B964DA31FFF10D08C428F93B8A4AF8A4" box="[810,888,1773,1797]" italics="true" pageId="23" pageNumber="24">fissidens</emphasis> +<bibRefCitation id="EF817BD2FFF10D08C47CF93B8D1AF8A4" author="Ameghino" box="[894,1064,1773,1797]" firstAuthor="Ameghino" pageId="23" pageNumber="24" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +and +<emphasis id="B964DA31FFF10D08C358F93B8C36F8A4" box="[1114,1284,1773,1797]" italics="true" pageId="23" pageNumber="24">Anisolamda amel</emphasis> +Simpson, 1948). +</paragraph> +<paragraph id="8BAF0623FFF10D08C773F8998C4EF822" blockId="23.[113,1432,1871,1979]" pageId="23" pageNumber="24"> +of MNRJ +<date id="FFAE20E3FFF10D08C7CDF899883DF8C6" box="[207,271,1871,1895]" pageId="23" pageNumber="24" value="1468-05">1468V</date> +( +<typeStatus id="54ABB881FFF10D08C619F899884EF8C6" box="[283,380,1871,1895]" pageId="23" pageNumber="24" type="holotype">holotype</typeStatus> +)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: encd, entocristid; end, entoconid; hyd, hypoconid; +</paragraph> +<paragraph id="8BAF0623FFF10D08C773F85188D7F81B" blockId="23.[113,1432,1871,1979]" pageId="23" pageNumber="24"> +hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal +<quantity id="4CE8ABC6FFF10D08C6B3F87588D3F81B" box="[433,481,1955,1979]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="23" pageNumber="24" unit="cm" value="1.0">1 cm</quantity> +. +</paragraph> +</caption> +<paragraph id="8BAF0623FFFE0D07C783FF4689D9FDA0" blockId="24.[128,778,144,513]" pageId="24" pageNumber="25"> +We included two species to represent +<taxonomicName id="4C107DA0FFFE0D07C507FF468BB4FF09" authorityName="Ameghino" authorityYear="1901" box="[517,646,144,168]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C507FF468BB4FF09" box="[517,646,144,168]" italics="true" pageId="24" pageNumber="25">Anisolambda</emphasis> +</taxonomicName> +to maximize the information available for this genus, because these species differ mostly in size (Simpson 1948), and the upper dentition has not been clearly assigned to any of these species (see more in +<taxonomicName id="4C107DA0FFFE0D07C79EFEDB887FFE84" box="[156,333,269,293]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" kingdom="Animalia" order="Litopterna" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="B964DA31FFFE0D07C79EFEDB882FFE84" box="[156,285,269,293]" italics="true" pageId="24" pageNumber="25">Anisolambda</emphasis> +spp. +</taxonomicName> +taxonomic comments in Supporting information, File S2). We only scored the preserved m3 (AMNH-VP 29101) of +<taxonomicName id="4C107DA0FFFE0D07C7EEFE9A88B8FEC5" authorityName="Simpson" authorityYear="1935" box="[236,394,332,356]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="tshotshe"> +<emphasis id="B964DA31FFFE0D07C7EEFE9A88B8FEC5" box="[236,394,332,356]" italics="true" pageId="24" pageNumber="25">Wainka tshotshe</emphasis> +</taxonomicName> +(see justification in Supporting information, File S2). From +<taxonomicName id="4C107DA0FFFE0D07C668FEBD8B7CFE22" box="[362,590,363,387]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="24" pageNumber="25" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +, we included three taxa: the probably Early Eocene +<taxonomicName id="4C107DA0FFFE0D07C6C8FE5D8BE2FE03" authorityName="Paula Couto" authorityYear="1952" box="[458,720,395,419]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="prototypica"> +<emphasis id="B964DA31FFFE0D07C6C8FE5D8BE2FE03" box="[458,720,395,419]" italics="true" pageId="24" pageNumber="25">Victorlemoinea prototypica</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFFE0D07C5E2FE5D89AFFE63" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="24" pageNumber="25">Figs 4L</figureCitation> +, +<figureCitation id="132B1AA6FFFE0D07C7ABFE7C89F6FE63" box="[169,196,426,450]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="24" pageNumber="25">5L</figureCitation> +), the Early to Late Eocene +<taxonomicName id="4C107DA0FFFE0D07C6E0FE7C8BE0FE63" box="[482,722,426,450]" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="arquinotiensis"> +<emphasis id="B964DA31FFFE0D07C6E0FE7C8BE0FE63" box="[482,722,426,450]" italics="true" pageId="24" pageNumber="25">Notiolofos arquinotiensis</emphasis> +</taxonomicName> +from +<collectingCountry id="F30746B3FFFE0D07C780FE1F89DCFE40" box="[130,238,457,481]" name="Antarctica" pageId="24" pageNumber="25">Antarctica</collectingCountry> +, and the Middle Eocene +<taxonomicName id="4C107DA0FFFE0D07C6ECFE1F8A38FE40" authorityName="Soria" authorityYear="1980" box="[494,778,457,481]" class="Mammalia" family="Sparnotheriodontidae" genus="Sparnotheriodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="epsilonoides"> +<emphasis id="B964DA31FFFE0D07C6ECFE1F8A38FE40" box="[494,778,457,481]" italics="true" pageId="24" pageNumber="25">Sparnotheriodon epsilonoides</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFFE0D07C789FE3F89E8FDA1" box="[139,218,488,513]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="24" pageNumber="25">Fig. 5M</figureCitation> +). +</paragraph> +<paragraph id="8BAF0623FFFE0D07C609FDF68B60FDFB" blockId="24.[127,779,544,1986]" pageId="24" pageNumber="25"> +<emphasis id="B964DA31FFFE0D07C609FDF68B60FDFB" italics="true" pageId="24" pageNumber="25">Dental nomenclature and considerations about the hypocone in SANUs</emphasis> +</paragraph> +<paragraph id="8BAF0623FFFE0D07C783FDB0883BFBF5" blockId="24.[127,779,544,1986]" pageId="24" pageNumber="25"> +In terms of tooth position, we used the ancestral dental formula for Placentalia of I1/i1 I2/i2 I3/i3 C/c P1/p1 P2/p2 P4/p4 P5/ p5 M1/m1 M2/m2 M3/m3, and thus we used these positions for character scoring. This is justified because of the probable loss of the P3/p +<quantity id="4CE8ABC6FFFE0D07C7F5FD35882FFD5A" box="[247,285,739,763]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="24" pageNumber="25" unit="in" value="3.0">3 in</quantity> +the common ancestor of Placentalia ( +<bibRefCitation id="EF817BD2FFFE0D07C5A6FD328987FCBB" author="McKenna" firstAuthor="McKenna" pageId="24" pageNumber="25" pagination="21 - 46" refId="ref44533" refString="McKenna MC. Toward a phylogenetic classification of the Mammalia. In: Luckett WP, Szalay FS (eds), Phylogeny of the Primates. New York, NY: Plenum Press, 1975, 21 - 46." type="book chapter" year="1975">McKenna 1975</bibRefCitation> +, Novacek 1986, O’Leary +<emphasis id="B964DA31FFFE0D07C6C3FCD588C0FCBA" box="[449,498,771,795]" italics="true" pageId="24" pageNumber="25">et al.</emphasis> +2013). As SANUs are placentals, most previous workers (e.g., +<bibRefCitation id="EF817BD2FFFE0D07C51EFCF48BCCFC9B" author="Forasiepi" box="[540,766,802,826]" etAl="et al." firstAuthor="Forasiepi" pageId="24" pageNumber="25" pagination="1 - 76" refId="ref41925" refString="Forasiepi AM, MacPhee RDE, Hernandez del Pino S et al. Exceptional skull of Huayqueriana (Mammalia, Litopterna, Macraucheniidae) from the Late Miocene of Argentina: anatomy, systematics, and paleobiological implications. Bulletin of the American Museum of Natural History 2016; 404: 1 - 76." type="journal article" year="2016"> +Forasiepi +<emphasis id="B964DA31FFFE0D07C585FCF58B8CFC9B" box="[647,702,802,826]" italics="true" pageId="24" pageNumber="25">et al.</emphasis> +2016 +</bibRefCitation> +) have considered a formula for SANUs counting the premolars mesiodistally (anteroposteriorly) from one to four (i.e., P1/p1 to P4/p4). Therefore, attention must be given when reading about the last two premolars in the main text. The benefit of using the ancestral placental dental formula is to make easier homologous comparisons between different therians, and is useful for transposing the anatomical information presented in this study into larger-scale projects that include older members of Theria that retain P3/p3. +</paragraph> +<paragraph id="8BAF0623FFFE0D07C79EFB8A8BF7FA2C" blockId="24.[127,779,544,1986]" pageId="24" pageNumber="25"> +In relation to the dental nomenclature, we, in most cases, followed the terminology of previous authors (e.g., +<bibRefCitation id="EF817BD2FFFE0D07C58EFBAD8A36FB32" author="Gelfo" box="[652,772,1147,1171]" firstAuthor="Gelfo" pageId="24" pageNumber="25" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo 2006</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFFE0D07C783FB4C88A0FB13" author="Carrillo and Puschel" box="[129,402,1178,1202]" firstAuthor="Carrillo" pageId="24" pageNumber="25" refId="ref40629" refString="Carrillo JD, Puschel HP. Pleistocene South American native ungulates (Notoungulata and Litopterna) of the historical Roth collections in Switzerland, from the Pampean Region of Argentina. Swiss Journal of Palaeontology 2023; 142: 28. https: // doi. org / 10.1186 / s 13358 - 023 - 00291 - 5" type="journal volume" year="2023">Carrillo and Püschel 2023</bibRefCitation> +) although with some modifications and additions (see +<figureCitation id="132B1AA6FFFE0D07C63CFB6F8844FB70" box="[318,374,1209,1233]" captionStart="Figure 3" captionStartId="9.[113,178,1773,1797]" captionTargetBox="[220,1352,145,1745]" captionTargetId="graphics-399@9.[425,1168,891,1380]" captionTargetPageId="9" captionText="Figure 3. Dental nomenclature used in the text and the characters. A, right upper molar in occlusal view.B–D, right lower molars in occlusal view. The main cusps of the molars are highlighted in bold.The molars represent structures sometimes present in SANUs and with relevance in the character scoring of this study. The molars do not represent any particular SANU.Molars in A and B were partially based on drawings from Gelfo (2006). Abbreviations upper molar: ecg, buccal cingulum or ectocingulum; es, entostyle; hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mt, metastyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; ppc, preparacrista (or paracrista); pplc, preparaconular crista; pprc, preprotocrista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; pshyc, posthypocrista; psmc, postmetacrista (or metacrista); psmlc, postmetaconular crista; pspc, postparacrista; psplc, postparaconular crista; psprc, postprotocrista. Abbreviations lower molar: co, cristid obliqua; encd, entocristid; end, entoconid; dbgd, distobuccal cingulid; dlgd, distolingual cingulid; hlph, hypolophid; hycd, hypocristid; hyd, hypoconid; hyld, hypoconulid; mbgd, mesiobuccal cingulid; med, metaconid; mlgd, mesiolingual cingulid; mlph, mesolophid; msd, mesoconid; pacd, paracristid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prcd, protocristid; prd, protoconid; prgd, precingulid; pscd, postcristid (or postentocristid); pseld, postentoconulid; psgd, postcingulid; psmcd, postmetacristid." pageId="24" pageNumber="25">Fig. 3</figureCitation> +). In those cases, a definition of the term is included as a comment in the character list (Supporting information, File S3). Our interpretation for the upper molars of notoungulates was based on an unworn right M1 or M2 of a +<taxonomicName id="4C107DA0FFFE0D07C791FAE1881AFAEE" authorityName="Ameghino" authorityYear="1901" box="[147,296,1335,1359]" class="Mammalia" family="Henricosborniidae" genus="Henricosbornia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C791FAE1881AFAEE" box="[147,296,1335,1359]" italics="true" pageId="24" pageNumber="25">Henricosbornia</emphasis> +</taxonomicName> +specimen (MACN A 107717; see +<figureCitation id="132B1AA6FFFE0D07C595FAE18BCCFAEE" box="[663,766,1335,1359]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="24" pageNumber="25">Fig. 6F–G</figureCitation> +) which helped in the interpretation of homologous dental structures that presented some wear in +<taxonomicName id="4C107DA0FFFE0D07C6D8FAA08B8DFA2C" box="[474,703,1398,1421]" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="praecursor"> +<emphasis id="B964DA31FFFE0D07C6D8FAA08B8DFA2C" box="[474,703,1398,1421]" italics="true" pageId="24" pageNumber="25">Simpsonotus praecursor</emphasis> +</taxonomicName> +. +</paragraph> +<paragraph id="8BAF0623FFFE0D07C79EFA438ACFFBF5" blockId="24.[127,779,544,1986]" lastBlockId="24.[824,1476,144,1108]" pageId="24" pageNumber="25"> +In the text and also in the character list, the term hypocone does not contain any a priori assumption of the origin of this upper dentition distolingual cusp. This criterion was applied in the character scoring when judging if this cusp was present or absent. There are different pathways to generate this cusp in placentals ( +<bibRefCitation id="EF817BD2FFFE0D07C614F9E48B11F9E8" author="Hunter and Jernvall" box="[278,547,1585,1609]" firstAuthor="Hunter" pageId="24" pageNumber="25" pagination="10718 - 22" refId="ref43329" refString="Hunter JP, Jernvall J. The hypocone as a key innovation in mammalian evolution. Proceedings of the National Academy of Sciences of the United States of America 1995; 92: 10718 - 22. https: // doi. org / 10.1073 / pnas. 92.23.10718" type="journal article" year="1995">Hunter and Jernvall 1995</bibRefCitation> +, Muizon +<emphasis id="B964DA31FFFE0D07C588F9E48B8FF9E8" box="[650,701,1585,1609]" italics="true" pageId="24" pageNumber="25">et al.</emphasis> +2019), and two of them have been previously suggested to explain the distolingual cusp present in some SANUs: (1) a postcingulum-derived hypocone or ‘true’ hypocone, and (2) a protoconeduplication-derived (PPD) hypocone or pseudohypocone (Muizon +<emphasis id="B964DA31FFFE0D07C7EAF919882CF947" box="[232,286,1742,1766]" italics="true" pageId="24" pageNumber="25">et al.</emphasis> +2019). Most SANUs (litopterns, indaleciids, notonychopids, xenungulates, and most ‘didolodontids’) and also some North American mioclaenids present a postcingulum-derived hypocone ( +<figureCitation id="132B1AA6FFFE0D07C64FF8FA88B9F8E5" box="[333,395,1836,1860]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="24" pageNumber="25">Figs 4</figureCitation> +, +<figureCitation id="132B1AA6FFFE0D07C69AF8FA88DEF8E2" box="[408,492,1836,1860]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="24" pageNumber="25">6A–C, I</figureCitation> +, +<figureCitation id="132B1AA6FFFE0D07C6FBF8FA8B58F8E5" box="[505,618,1836,1860]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="24" pageNumber="25">8B–E, G–J</figureCitation> +). Muizon +<emphasis id="B964DA31FFFE0D07C5DBF8FB8A38F8E5" box="[729,778,1836,1860]" italics="true" pageId="24" pageNumber="25">et al.</emphasis> +(2019) suggested that the incipient protocone duplication seen in +<taxonomicName id="4C107DA0FFFE0D07C798F8BD8833F822" authorityName="de Muizon & Marshall" authorityYear="1987" box="[154,257,1899,1923]" class="Mammalia" family="Mioclaenidae" genus="Molinodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C798F8BD8833F822" box="[154,257,1899,1923]" italics="true" pageId="24" pageNumber="25">Molinodus</emphasis> +</taxonomicName> +and other kollpaniines ( +<figureCitation id="132B1AA6FFFE0D07C6FBF8BD8B70F822" box="[505,578,1899,1923]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="24" pageNumber="25">Fig. 8A</figureCitation> +) is the precursor of the hypocone seen in other SANUs such as +<taxonomicName id="4C107DA0FFFE0D07C52CF85C8BABF803" box="[558,665,1930,1954]" class="Mammalia" family="Mioclaenidae" genus="Raulvaccia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C52CF85C8BABF803" box="[558,665,1930,1954]" italics="true" pageId="24" pageNumber="25">Raulvaccia</emphasis> +</taxonomicName> +, +<taxonomicName id="4C107DA0FFFE0D07C5A6F85D8A36F803" authorityName="Paula Couto" authorityYear="1952" box="[676,772,1931,1954]" class="Mammalia" family="Didolodontidae" genus="Lamegoia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C5A6F85D8A36F803" box="[676,772,1931,1954]" italics="true" pageId="24" pageNumber="25">Lamegoia</emphasis> +</taxonomicName> +, and notoungulates. From these taxa, we agree with Muizon +<emphasis id="B964DA31FFFE0D07C5DEF87C8A38F860" box="[732,778,1961,1985]" italics="true" pageId="24" pageNumber="25">et al.</emphasis> +(2019) only in the condition displayed in the M2 of +<taxonomicName id="4C107DA0FFFE0D07C25CFF478C8EFF09" authorityName="Paula Couto" authorityYear="1952" box="[1374,1468,145,168]" class="Mammalia" family="Didolodontidae" genus="Lamegoia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C25CFF478C8EFF09" box="[1374,1468,145,168]" italics="true" pageId="24" pageNumber="25">Lamegoia</emphasis> +</taxonomicName> +, which seems to have a PPD hypocone, considering the appressed bases and position very close to the protocone, and also the disassociation of this distolingual cusp with the low postcingulum present in this taxon ( +<figureCitation id="132B1AA6FFFE0D07C311FEDB8D6AFE84" box="[1043,1112,269,293]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="24" pageNumber="25">Fig. 8F</figureCitation> +). In the case of +<taxonomicName id="4C107DA0FFFE0D07C3F0FEDB8C6FFE84" box="[1266,1373,269,293]" class="Mammalia" family="Mioclaenidae" genus="Raulvaccia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C3F0FEDB8C6FFE84" box="[1266,1373,269,293]" italics="true" pageId="24" pageNumber="25">Raulvaccia</emphasis> +</taxonomicName> +, although its M2 has a distolingual cusp appressed to the protocone, this cusp is clearly derived from the postcingulum ( +<emphasis id="B964DA31FFFE0D07C22FFE9B8C58FEC5" box="[1325,1386,333,356]" italics="true" pageId="24" pageNumber="25">contra</emphasis> +Muizon +<emphasis id="B964DA31FFFE0D07C43BFEBA8A5AFE22" box="[825,872,363,387]" italics="true" pageId="24" pageNumber="25">et al</emphasis> +. 2019) because it is associated and continuous with it ( +<figureCitation id="132B1AA6FFFE0D07C446FE5D8A80FE02" box="[836,946,395,419]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="24" pageNumber="25">Fig. 8C–D</figureCitation> +). The cusp configuration of the M2 of +<taxonomicName id="4C107DA0FFFE0D07C255FE5D8CF0FE02" box="[1367,1474,395,419]" class="Mammalia" family="Mioclaenidae" genus="Raulvaccia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C255FE5D8CF0FE02" box="[1367,1474,395,419]" italics="true" pageId="24" pageNumber="25">Raulvaccia</emphasis> +</taxonomicName> +is very similar to the M2 of +<taxonomicName id="4C107DA0FFFE0D07C36FFE7C8DE6FE63" box="[1133,1236,426,450]" class="Mammalia" family="Mioclaenidae" genus="Escribania" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C36FFE7C8DE6FE63" box="[1133,1236,426,450]" italics="true" pageId="24" pageNumber="25">Escribania</emphasis> +</taxonomicName> +, which also exhibits a postcingulum-derived hypocone ( +<figureCitation id="132B1AA6FFFE0D07C3A0FE1F8DDFFE40" box="[1186,1261,457,481]" captionStart="Figure 8" captionStartId="20.[131,196,1781,1805]" captionTargetBox="[132,1475,150,1750]" captionTargetId="figure-8@20.[129,1479,147,1753]" captionTargetPageId="20" captionText="Figure 8. Right upper molars of South American ‘condylarths’ and ‘didolodontids’. A, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="24" pageNumber="25">Fig. 8B</figureCitation> +). The small lingual cusp mentioned by Muizon +<emphasis id="B964DA31FFFE0D07C366FE3F8DA5FDA0" box="[1124,1175,489,513]" italics="true" pageId="24" pageNumber="25">et al.</emphasis> +(2019) is a hypostyle (additional cusp distal to the hypocone; +<figureCitation id="132B1AA6FFFE0D07C3A5FDDE8DEDFD81" box="[1191,1247,520,544]" captionStart="Figure 3" captionStartId="9.[113,178,1773,1797]" captionTargetBox="[220,1352,145,1745]" captionTargetId="graphics-399@9.[425,1168,891,1380]" captionTargetPageId="9" captionText="Figure 3. Dental nomenclature used in the text and the characters. A, right upper molar in occlusal view.B–D, right lower molars in occlusal view. The main cusps of the molars are highlighted in bold.The molars represent structures sometimes present in SANUs and with relevance in the character scoring of this study. The molars do not represent any particular SANU.Molars in A and B were partially based on drawings from Gelfo (2006). Abbreviations upper molar: ecg, buccal cingulum or ectocingulum; es, entostyle; hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mt, metastyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; ppc, preparacrista (or paracrista); pplc, preparaconular crista; pprc, preprotocrista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; pshyc, posthypocrista; psmc, postmetacrista (or metacrista); psmlc, postmetaconular crista; pspc, postparacrista; psplc, postparaconular crista; psprc, postprotocrista. Abbreviations lower molar: co, cristid obliqua; encd, entocristid; end, entoconid; dbgd, distobuccal cingulid; dlgd, distolingual cingulid; hlph, hypolophid; hycd, hypocristid; hyd, hypoconid; hyld, hypoconulid; mbgd, mesiobuccal cingulid; med, metaconid; mlgd, mesiolingual cingulid; mlph, mesolophid; msd, mesoconid; pacd, paracristid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prcd, protocristid; prd, protoconid; prgd, precingulid; pscd, postcristid (or postentocristid); pseld, postentoconulid; psgd, postcingulid; psmcd, postmetacristid." pageId="24" pageNumber="25">Fig. 3</figureCitation> +) in the postcingulum, which is not uncommon in litopterns such as anisolambdids ( +<figureCitation id="132B1AA6FFFE0D07C446FD918AA1FDFE" box="[836,915,583,607]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="24" pageNumber="25">Fig. 4N</figureCitation> +). Regarding notoungulates, we agree with Muizon +<emphasis id="B964DA31FFFE0D07C2B3FD918A60FDDF" italics="true" pageId="24" pageNumber="25">et al.</emphasis> +(2019) that the postcingulum is low and unconnected to the distolingual cusp, a condition seen clearly in +<taxonomicName id="4C107DA0FFFE0D07C21AFD508CA1FD3C" authorityName="Pascual, Vucetich & Fernandez" authorityYear="1979" box="[1304,1427,646,669]" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C21AFD508CA1FD3C" box="[1304,1427,646,669]" italics="true" pageId="24" pageNumber="25">Simpsonotus</emphasis> +</taxonomicName> +and +<taxonomicName id="4C107DA0FFFE0D07C43BFD738AFCFD1C" authorityName="Ameghino" authorityYear="1901" box="[825,974,677,701]" class="Mammalia" family="Henricosborniidae" genus="Henricosbornia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C43BFD738AFCFD1C" box="[825,974,677,701]" italics="true" pageId="24" pageNumber="25">Henricosbornia</emphasis> +</taxonomicName> +( +<figureCitation id="132B1AA6FFFE0D07C4E3FD738D7FFD1C" box="[993,1101,677,701]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="24" pageNumber="25">Fig. 6E–G</figureCitation> +). However, the protocone is separated from the hypocone by a sulcus, and the hypocone is more closely connected with the metaconular region. This is clear in the +<taxonomicName id="4C107DA0FFFE0D07C45DFCD58AC6FCBA" authorityName="Ameghino" authorityYear="1901" box="[863,1012,771,795]" class="Mammalia" family="Henricosborniidae" genus="Henricosbornia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="24" pageNumber="25" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFE0D07C45DFCD58AC6FCBA" box="[863,1012,771,795]" italics="true" pageId="24" pageNumber="25">Henricosbornia</emphasis> +</taxonomicName> +specimen MACN A 107717, an unworn right M1 or M2, which shows a metaconule, and very close and associated with it by a crista, a hypocone ( +<figureCitation id="132B1AA6FFFE0D07C3BFFC978C17FCF8" box="[1213,1317,833,857]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="24" pageNumber="25">Fig. 6F–G</figureCitation> +). Therefore, an alternative hypothesis is that the hypocone in notoungulates derives from a secondary hypertrophied metaconule that migrated lingually. Without an earlier notoungulate showing a less derived condition, it is difficult to assess these hypotheses. In any case, even a postcingulum-derived hypocone has been shown to have arisen independently in multiple lineages ( +<bibRefCitation id="EF817BD2FFFE0D07C3F4FC288A5FFB95" author="Hunter and Jernvall" firstAuthor="Hunter" pageId="24" pageNumber="25" pagination="10718 - 22" refId="ref43329" refString="Hunter JP, Jernvall J. The hypocone as a key innovation in mammalian evolution. Proceedings of the National Academy of Sciences of the United States of America 1995; 92: 10718 - 22. https: // doi. org / 10.1073 / pnas. 92.23.10718" type="journal article" year="1995">Hunter and Jernvall 1995</bibRefCitation> +), and therefore its presence or absence is not necessarily a keystone character. +</paragraph> +<paragraph id="8BAF0623FFFE0D07C4E8FBA58C23FB2C" blockId="24.[1002,1297,1139,1165]" box="[1002,1297,1139,1165]" pageId="24" pageNumber="25"> +<emphasis id="B964DA31FFFE0D07C4E8FBA58C23FB2C" box="[1002,1297,1139,1165]" italics="true" pageId="24" pageNumber="25">Character sampling and scoring</emphasis> +</paragraph> +<paragraph id="8BAF0623FFFE0D07C43BFB4C8D76F967" blockId="24.[824,1476,1178,1985]" pageId="24" pageNumber="25"> +Based on an inclusive matrix currently being constructed by our team to test the relationships within Placentalia broadly, we built a new morphological matrix with 703 characters, of which 667 concern dental anatomy and 36 mandibular anatomy. Some of the characters were based on observations and characters of previous phylogenetic studies including litopterns ( +<tableCitation id="C6923398FFFE0D07C236FAE18CB1FAEE" box="[1332,1411,1335,1359]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" captionText="Table 2. Previous phylogenetic analyses that included Litopterna and Didolodontidae. Abbreviations: Ad, Adianthidae; An, Anisolambdidae; As, Astrapotheria; C, cranial; D, dental; Di, Didolodontidae; I, Indaleciidae; K, Kollpaniinae; Ma, Macraucheniidae; Mi, North American Mioclaenidae; No. litopterns, number of uncontroversial litopterns; Nt, Notonychopidae; Nu, Notoungulata; PC, postcranial; Pe, Proterotheriidae; Po, Protolipternidae; Py, Pyrotheria; S, Sparnotheriodontidae; X, Xenungulata" pageId="24" pageNumber="25">Table 2</tableCitation> +), and others were proposed +<emphasis id="B964DA31FFFE0D07C32FFA808D4EFACF" box="[1069,1148,1366,1390]" italics="true" pageId="24" pageNumber="25">de novo</emphasis> +from anatomical observations made in this study (Supporting information, File S3). We actively identified and scored autapomorphies, as these have an impact on branch length estimates and evolutionary rates when using the Mk model in Bayesian phylogenetic analyses ( +<bibRefCitation id="EF817BD2FFFE0D07C287FA058A5FF9AA" author="Lewis" firstAuthor="Lewis" pageId="24" pageNumber="25" pagination="913 - 25" refId="ref43877" refString="Lewis P. A likelihood approach to estimating phylogeny from discrete morphological character data. Society of Systematic Biologists 2001; 50: 913 - 25." type="journal article" year="2001">Lewis 2001</bibRefCitation> +, Müller and Reisz 2006, +<bibRefCitation id="EF817BD2FFFE0D07C376FA258C09F9AB" author="Lee and Palci" box="[1140,1339,1522,1547]" firstAuthor="Lee" pageId="24" pageNumber="25" pagination="922 - 9" refId="ref43839" refString="Lee MSY, Palci A. Morphological phylogenetics in the genomic age. Current Biology 2015; 25: R 922 - 9. https: // doi. org / 10.1016 / j. cub. 2015.07.009" type="journal article" year="2015">Lee and Palci 2015</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFFE0D07C24AFA258A9AF98B" author="Matzke and Irmis" firstAuthor="Matzke" pageId="24" pageNumber="25" refId="ref44249" refString="Matzke NJ, Irmis RB. Including autapomorphies is important for paleontological tip-dating with clocklike data, but not with non-clock data. PeerJ 2018; 6: e 4553. https: // doi. org / 10.7717 / peerj. 4553" type="journal volume" year="2018">Matzke and Irmis 2018</bibRefCitation> +). In maximum parsimony, retaining autapomorphies does not affect the topology of the trees obtained (Yeates 1992), so the same matrix was used in parsimony and Bayesian analyses. The matrix is available online on Morphobank (morphobank. org) under project number 4441 and as Nexus files (Supporting information, Files S4, S5). +</paragraph> +<paragraph id="8BAF0623FFFE0D06C457F91888C5FD9E" blockId="24.[824,1476,1178,1985]" lastBlockId="25.[112,764,144,1202]" lastPageId="25" lastPageNumber="26" pageId="24" pageNumber="25"> +<taxonomicName id="4C107DA0FFFE0D07C457F9188AE2F947" box="[853,976,1742,1766]" class="Prions" family="Mammalian" higherTaxonomySource="GBIF" kingdom="Viruses" pageId="24" pageNumber="25" rank="family">Mammalian</taxonomicName> +teeth display serial homology and correlation between their features ( +<bibRefCitation id="EF817BD2FFFE0D07C336F93B8D9DF8A4" author="Butler" box="[1076,1199,1773,1797]" firstAuthor="Butler" pageId="24" pageNumber="25" pagination="843 - 50" refId="ref40597" refString="Butler PM. Dental merism and tooth development. Journal of Dental Research 1967; 46: 843 - 50. https: // doi. org / 10.1177 / 002203456704 60053801" type="journal article" year="1967">Butler 1967</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFFE0D07C3BFF93B8A5FF884" author="Gomez-Robles and Polly" firstAuthor="Gomez-Robles" pageId="24" pageNumber="25" pagination="1024 - 43" refId="ref43022" refString="Gomez-Robles A, Polly PD. Morphological integration in the hominin dentition: evolutionary, developmental, and functional factors. Evolution 2012; 66: 1024 - 43. https: // doi. org / 10.1111 / j. 1558 - 5646.2011.01508. x" type="journal article" year="2012">Gómez-Robles and Polly 2012</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFFE0D07C478F8DB8D74F884" author="Labonne" box="[890,1094,1805,1829]" etAl="et al." firstAuthor="Labonne" pageId="24" pageNumber="25" pagination="224 - 32" refId="ref43775" refString="Labonne G, Navarro N, Laffont R et al. Developmental integration in a functional unit: deciphering processes from adult dental morphology. Evolution & Development 2014; 16: 224 - 32. https: // doi. org / 10.1111 / ede. 12085" type="journal article" year="2014"> +Labonne +<emphasis id="B964DA31FFFE0D07C4DEF8DB8D3EF884" box="[988,1036,1805,1829]" italics="true" pageId="24" pageNumber="25">et al.</emphasis> +2014 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFFE0D07C350F8DB8C09F884" author="Billet and Bardin" box="[1106,1339,1805,1829]" firstAuthor="Billet" pageId="24" pageNumber="25" pagination="267 - 80" refId="ref39937" refString="Billet G, Bardin J. Serial homology and correlated characters in morphological phylogenetics: modeling the evolution of dental crests in placentals. Systematic Biology 2019; 68: 267 - 80. https: // doi. org / 10.1093 / sysbio / syy 071" type="journal article" year="2019">Billet and Bardin 2019</bibRefCitation> +). In the case of SANUs and ‘condylarths’, we observed that the majority of dental structures were consistent between the first and second molar in the upper and lower series (M1/m1 and M2/m2), showing only minor variations. Therefore, to reduce the weight in the analyses of these clearly correlated tooth positions, M1/ m1 and M2/m2 were grouped as M1–M2/m1–m2 characters. For the few anatomical differences between M1/m1 and M2/ m2, these features were scored as polymorphic in the matrix. The remainder of the teeth loci were treated with a separate set of characters because merging them would most likely reduce relevant anatomical information for resolving the litoptern phylogeny. In addition, we observed in SANUs and ‘condylarths’ that the height of some cristae/cristids of certain tooth positions was consistently correlated with the height of contiguous cristae/cristids (e.g., entocristid, postcristid, hypocristid). In those cases, we considered a single character for describing the overall height of the correlated cristae/cristids (e.g., Character 394: m1–m2 talonid cristids, height) to reduce the weight in the analyses of these correlated structures. +</paragraph> +<paragraph id="8BAF0623FFFF0D06C78FFD91881FFB13" blockId="25.[112,764,144,1202]" pageId="25" pageNumber="26"> +When possible, we scored dental characters from unworn specimens that allowed us to examine the anatomy in detail of each taxon. In cases in which all the specimens of a taxon were worn, we scored as question marks (?) the characters that we could not examine. In general, we used question marks (?) for missing information about the state of a character and also for inapplicable characters. For scoring the upper molars of macraucheniids, we followed the cusps and cristae homology proposal of +<bibRefCitation id="EF817BD2FFFF0D06C5A8FCF4880BFCF8" author="Carrillo and Puschel" firstAuthor="Carrillo" pageId="25" pageNumber="26" refId="ref40629" refString="Carrillo JD, Puschel HP. Pleistocene South American native ungulates (Notoungulata and Litopterna) of the historical Roth collections in Switzerland, from the Pampean Region of Argentina. Swiss Journal of Palaeontology 2023; 142: 28. https: // doi. org / 10.1186 / s 13358 - 023 - 00291 - 5" type="journal volume" year="2023">Carrillo and Püschel (2023)</bibRefCitation> +. The scores were based on direct observation of museum specimens, photographs of the specimens, and images from the literature. A full list of museum abbreviations, the scored specimens and their taxonomy, anatomical observations and interpretations, and the justification of tooth position assignments that influenced the character scorings is present in Supporting information, File S2. Considering +<bibRefCitation id="EF817BD2FFFF0D06C549FC2B882BFB94" author="Brocklehurst and Haridy's" firstAuthor="Brocklehurst" pageId="25" pageNumber="26" pagination="707 - 18" refId="ref40512" refString="Brocklehurst N, Haridy Y. Do meristic characters used in phylogenetic analysis evolve in an ordered manner? Systematic Biology 2021; 70: 707 - 18. https: // doi. org / 10.1093 / sysbio / syaa 078" type="journal article" year="2021">Brocklehurst and Haridy’s (2021)</bibRefCitation> +models fitting results for heterodont dentition, we treated all the characters as unordered in the phylogenetic analyses as we did not have any +<emphasis id="B964DA31FFFF0D06C6B9FB8A8B37FBD2" box="[443,517,1116,1139]" italics="true" pageId="25" pageNumber="26">a priori</emphasis> +knowlegde of the ontogenetic trajectory of each continous or meristic character present in our matrix. +</paragraph> +<paragraph id="8BAF0623FFFF0D06C79BFB048BE1FB4A" blockId="25.[112,764,1233,1985]" box="[153,723,1233,1260]" pageId="25" pageNumber="26"> +<emphasis id="B964DA31FFFF0D06C79BFB048BE1FB4A" box="[153,723,1233,1260]" italics="true" pageId="25" pageNumber="26">Testing different hypotheses for the paraconid of notoungulates</emphasis> +</paragraph> +<paragraph id="8BAF0623FFFF0D06C773FB2E8BE3F884" blockId="25.[112,764,1233,1985]" pageId="25" pageNumber="26"> +In early notoungulates of different suborders, there is a small mesiolingual cusp on the lower molars, which is appressed to the metaconid ( +<figureCitation id="132B1AA6FFFF0D06C7F3FAE1880FFAEE" box="[241,317,1335,1359]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="25" pageNumber="26">Fig. 7C</figureCitation> +). Simpson (1967) considered this cusp to be a twinned or duplicated metaconid. However, most later authors have interpreted this cusp to be a paraconid in which the connection with the paracristid is lost (e.g., +<bibRefCitation id="EF817BD2FFFF0D06C540FA438B85FA0C" author="Bond" box="[578,695,1429,1453]" firstAuthor="Bond" pageId="25" pageNumber="26" pagination="76 - 7" refId="ref40165" refString="Bond M. Consideraciones sobre la morfologia de los molariformes inferiores de los Notoungulata. Resumenes de las V Jornadas Argentinas de Paleontologia de Vertebrados 1988; 76 - 7." type="book chapter" year="1988">Bond 1988</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFFF0D06C5C7FA438982FA6D" author="Corona A & Badin AC & Perea D" firstAuthor="Soria" pageId="25" pageNumber="26" refId="ref41392" refString="Corona A, Badin AC, Perea D et al. A new genus and species and additional reports of the South American native ungulates Proterotheriidae (Mammalia, Litopterna) in the Late Miocene of Uruguay. Journal of South American Earth Sciences 2020; 102: 102646. https: // doi. org / 10.1016 / j. jsames. 2020.102646" type="journal volume" year="2020" yearSuffix="a"> +Soria 1989 +<bibRefCitation id="EF817BD2FFFF0D06C7A7FA628858FA6D" author="a, Bauza" box="[165,362,1460,1484]" etAl="et al." firstAuthor="a" pageId="25" pageNumber="26" refId="ref41392" refString="Corona A, Badin AC, Perea D et al. A new genus and species and additional reports of the South American native ungulates Proterotheriidae (Mammalia, Litopterna) in the Late Miocene of Uruguay. Journal of South American Earth Sciences 2020; 102: 102646. https: // doi. org / 10.1016 / j. jsames. 2020.102646" type="journal volume" year="2020"> +a, Bauzá +<emphasis id="B964DA31FFFF0D06C603FA638802FA6D" box="[257,304,1460,1484]" italics="true" pageId="25" pageNumber="26">et al.</emphasis> +2020 +</bibRefCitation> +</bibRefCitation> +). This is consistent with the paraconid position and size in kollpanines, didolodontids, xenungulates, protolipternids, and anisolambdid litopterns ( +<figureCitation id="132B1AA6FFFF0D06C55FFA258BF4F9AB" box="[605,710,1522,1547]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="25" pageNumber="26">Figs 5J–K</figureCitation> +, +<figureCitation id="132B1AA6FFFF0D06C5DAFA258BC6F9AB" box="[728,756,1523,1546]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="25" pageNumber="26">7E</figureCitation> +, +<figureCitation id="132B1AA6FFFF0D06C773F9C4894CF98B" box="[113,126,1554,1578]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="25" pageNumber="26">9</figureCitation> +). In addition, the loss of contact between the paracristid and the paraconid is observed in m1–m2 of the anisolambdid +<taxonomicName id="4C107DA0FFFF0D06C773F9878822F9C8" authorityName=", Cifelli" authorityYear="1983" box="[113,272,1617,1641]" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="25" pageNumber="26" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFFF0D06C773F9878822F9C8" box="[113,272,1617,1641]" italics="true" pageId="25" pageNumber="26">Paranisolambda</emphasis> +</taxonomicName> +(Cifelli 1983a; +<figureCitation id="132B1AA6FFFF0D06C6B4F9878B30F9C9" box="[438,514,1616,1641]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="25" pageNumber="26">Fig. 5K</figureCitation> +), most likely independently emerging in this lineage. In sum, there are two contrasting hypotheses to explain the mesiolingual cusp present on the lower molars of notoungulates: (i) the mesiolingual cusp is a reduced paraconid and (ii) the mesiolingual cusp is a twinned or duplicated metaconid. In the second hypothesis, the paraconid would be extremely reduced at the end of the paracristid ( +<figureCitation id="132B1AA6FFFF0D06C575F8DB8BF3F884" box="[631,705,1805,1829]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="25" pageNumber="26">Fig. 7C</figureCitation> +). +</paragraph> +<paragraph id="8BAF0623FFFF0D06C78FF8FA8D58FEE4" blockId="25.[112,764,1233,1985]" lastBlockId="25.[810,1461,144,325]" pageId="25" pageNumber="26"> +To test the effect of these two different anatomical hypotheses in relation to the paraconid of notoungulates, we scored the notoungulate +<taxonomicName id="4C107DA0FFFF0D06C600F8BD88D9F823" box="[258,491,1899,1922]" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="praecursor"> +<emphasis id="B964DA31FFFF0D06C600F8BD88D9F823" box="[258,491,1899,1922]" italics="true" pageId="25" pageNumber="26">Simpsonotus praecursor</emphasis> +</taxonomicName> +twice, and thus have two different matrix versions (Supporting information, Files S4, S5). In one matrix, +<taxonomicName id="4C107DA0FFFF0D06C60CF87C88C7F860" box="[270,501,1962,1985]" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="praecursor"> +<emphasis id="B964DA31FFFF0D06C60CF87C88C7F860" box="[270,501,1962,1985]" italics="true" pageId="25" pageNumber="26">Simpsonotus praecursor</emphasis> +</taxonomicName> +was scored following the first hypothesis, and in a different matrix following the second hypothesis. Our objective was to test the influence of each hypothesis in the position of +<taxonomicName id="4C107DA0FFFF0D06C340FF198D9DFF46" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[1090,1199,207,231]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="25" pageNumber="26" phylum="Chordata" rank="order">Litopterna</taxonomicName> +among ‘condylarths’ and other SANUs. Therefore, the following phylogenetic analyses were conducted twice, once for each hypothesis, and are denoted in the text as H1 and H2. +</paragraph> +<paragraph id="8BAF0623FFFF0D06C4DDFEB38DCDFEDF" blockId="25.[991,1279,356,382]" box="[991,1279,356,382]" pageId="25" pageNumber="26"> +<emphasis id="B964DA31FFFF0D06C4DDFEB38DCDFEDF" box="[991,1279,356,382]" italics="true" pageId="25" pageNumber="26">Maximum parsimony analyses</emphasis> +</paragraph> +<paragraph id="8BAF0623FFFF0D06C428FE5D8AD6FCD9" blockId="25.[809,1460,395,889]" pageId="25" pageNumber="26"> +We used TNT v.1.6 ( +<bibRefCitation id="EF817BD2FFFF0D06C319FE5D8C60FE02" author="Goloboff and Catalano" box="[1051,1362,395,419]" firstAuthor="Goloboff" pageId="25" pageNumber="26" pagination="221 - 38" refId="ref42983" refString="Goloboff PA, Catalano SA. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics 2016; 32: 221 - 38. https: // doi. org / 10.1111 / cla. 12160" type="journal article" year="2016">Goloboff and Catalano 2016</bibRefCitation> +), setting the outgroup as +<taxonomicName id="4C107DA0FFFF0D06C4DBFE7C8D8FFE63" authorityName="Sloan & Van Valen" authorityYear="1965" box="[985,1213,426,450]" class="Mammalia" family="Tupaiidae" genus="Protungulatum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scandentia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="donnae"> +<emphasis id="B964DA31FFFF0D06C4DBFE7C8D8FFE63" box="[985,1213,426,450]" italics="true" pageId="25" pageNumber="26">Protungulatum donnae</emphasis> +</taxonomicName> +and employing a ‘New Technology’ driven search with sectorial search, ratchet, drift, and tree fusing, using default settings for each search algorithm, and set to find the minimum length tree or best score 100 times. After completion, we conducted an additional ‘traditional search’ using the tree bisection and reconnection ( +<collectingRegion id="49D4C8C1FFFF0D06C271FD918C99FDFE" box="[1395,1451,583,607]" country="Russia" name="Tver'" pageId="25" pageNumber="26">TBR</collectingRegion> +) branch-swapping algorithm on the trees obtained from the New Technology search. In the case of obtaining more than one most most parsimonious tree (MPT), a strict consensus tree was computed. For each hypothesis (H1 and H2), the absolute Bremer supports (BS) and Jackknife resampling support (JS; 1000 replicates, 36%-character removal) for the nodes were calculated to quantify clade support. The consistency index (CI) and the retention index (RI) were estimated using the ‘STATS.RUN’ script provided by TNT. +</paragraph> +<paragraph id="8BAF0623FFFF0D06C4F2FC4E8DDFFC13" blockId="25.[1008,1261,920,946]" box="[1008,1261,920,946]" pageId="25" pageNumber="26"> +<emphasis id="B964DA31FFFF0D06C4F2FC4E8DDFFC13" box="[1008,1261,920,946]" italics="true" pageId="25" pageNumber="26">Undated Bayesian analyses</emphasis> +</paragraph> +<paragraph id="8BAF0623FFFF0D06C428FC698CB2FA0C" blockId="25.[810,1461,959,1453]" pageId="25" pageNumber="26"> +The undated Bayesian analyses were conducted in MrBayes v.3.2.7a (Ronquist +<emphasis id="B964DA31FFFF0D06C4F0FC098D16FC57" box="[1010,1060,990,1014]" italics="true" pageId="25" pageNumber="26">et al.</emphasis> +2012) using +<taxonomicName id="4C107DA0FFFF0D06C3B0FC088CA4FC57" authorityName="Sloan & Van Valen" authorityYear="1965" box="[1202,1430,990,1014]" class="Mammalia" family="Tupaiidae" genus="Protungulatum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scandentia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="donnae"> +<emphasis id="B964DA31FFFF0D06C3B0FC088CA4FC57" box="[1202,1430,990,1014]" italics="true" pageId="25" pageNumber="26">Protungulatum donnae</emphasis> +</taxonomicName> +as the outgroup. We set one partition for the morphological data using the Mkv+Γ model of morphological evolution ( +<bibRefCitation id="EF817BD2FFFF0D06C274FBCB8A6CFBF5" author="Lewis" firstAuthor="Lewis" pageId="25" pageNumber="26" pagination="913 - 25" refId="ref43877" refString="Lewis P. A likelihood approach to estimating phylogeny from discrete morphological character data. Society of Systematic Biologists 2001; 50: 913 - 25." type="journal article" year="2001">Lewis 2001</bibRefCitation> +) with four rate categories following +<bibRefCitation id="EF817BD2FFFF0D06C3D6FBEA8A47FBD2" author="Harrison and Larsson" firstAuthor="Harrison" pageId="25" pageNumber="26" pagination="307 - 24" refId="ref43103" refString="Harrison LB, Larsson HCE. Among-character rate variation distributions in phylogenetic analysis of discrete morphological characters. Systematic Biology 2015; 64: 307 - 24. https: // doi. org / 10.1093 / sysbio / syu 098" type="journal article" year="2015">Harrison and Larsson (2015)</bibRefCitation> +. The analyses were run using two independent runs of four chains for 10 million Markov chain Monte Carlo (MCMC) generations, removing the first 25% of the samples as burn-in. At the end of the analyses, the effective sample size for the parameters was> 200, and the deviation of split frequencies was below 0.01. In addition, the runs were visually inspected using Tracer v.1.7.1 (Rambaut +<emphasis id="B964DA31FFFF0D06C32AFACE8D63FA8E" box="[1064,1105,1303,1327]" italics="true" pageId="25" pageNumber="26">et al</emphasis> +. 2018) to ensure convergence and stationarity were achieved. A 50% majority rule was computed at the end of the analyses and the posterior probabilities for the nodes (prob) were estimated. The same procedure using the same priors and parameters was conducted for H1 and H2. +</paragraph> +<paragraph id="8BAF0623FFFF0D06C4E9FA1A8DC0FA47" blockId="25.[1003,1266,1484,1510]" box="[1003,1266,1484,1510]" pageId="25" pageNumber="26"> +<emphasis id="B964DA31FFFF0D06C4E9FA1A8DC0FA47" box="[1003,1266,1484,1510]" italics="true" pageId="25" pageNumber="26">Tip-dated Bayesian analyses</emphasis> +</paragraph> +<paragraph id="8BAF0623FFFF0D06C428FA258AEBF8E5" blockId="25.[809,1461,1523,1985]" pageId="25" pageNumber="26"> +The tip-dated Bayesian analyses were also conducted in MrBayes v.3.2.7a (Ronquist +<emphasis id="B964DA31FFFF0D06C4EFF9C58D2FF98B" box="[1005,1053,1554,1578]" italics="true" pageId="25" pageNumber="26">et al.</emphasis> +2012) using the same outgroup, model of morphological evolution, and sampling parameters as the undated Bayesian analyses (see above). To calibrate the root of the tree, a truncated normal prior was used with a minimum age of 66.061 Mya which is 0.001 Myr older than the maximum age of the oldest taxon +<taxonomicName id="4C107DA0FFFF0D06C4F1F9798DE7F966" authorityName="Sloan & Van Valen" authorityYear="1965" box="[1011,1237,1711,1735]" class="Mammalia" family="Tupaiidae" genus="Protungulatum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scandentia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="donnae"> +<emphasis id="B964DA31FFFF0D06C4F1F9798DE7F966" box="[1011,1237,1711,1735]" italics="true" pageId="25" pageNumber="26">Protungulatum donnae</emphasis> +</taxonomicName> +(66.06 Mya), a mean age 0.1 Myr older than maximum age of +<taxonomicName id="4C107DA0FFFF0D06C3D1F9188C87F947" authorityName="Sloan & Van Valen" authorityYear="1965" box="[1235,1461,1742,1766]" class="Mammalia" family="Tupaiidae" genus="Protungulatum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scandentia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="donnae"> +<emphasis id="B964DA31FFFF0D06C3D1F9188C87F947" box="[1235,1461,1742,1766]" italics="true" pageId="25" pageNumber="26">Protungulatum donnae</emphasis> +</taxonomicName> +(66.16 Mya), and a standard deviation of 1.0 Myr following an approach used in previous studies (Sallam and Seiffert 2020, +<bibRefCitation id="EF817BD2FFFF0D06C428F8FA8AFBF8E5" author="Beck" box="[810,969,1836,1860]" etAl="et al." firstAuthor="Beck" pageId="25" pageNumber="26" refId="ref39714" refString="Beck RMD, de Vries D, Janiak MC et al. Total evidence phylogeny of platyrrhine primates and a comparison of undated and tip-dating approaches. Journal of Human Evolution 2023; 174: 103293. https: // doi. org / 10.1016 / j. jhevol. 2022.103293" type="journal volume" year="2023"> +Beck +<emphasis id="B964DA31FFFF0D06C460F8FB8AA2F8E5" box="[866,912,1836,1860]" italics="true" pageId="25" pageNumber="26">et al.</emphasis> +2023 +</bibRefCitation> +). +</paragraph> +<paragraph id="8BAF0623FFFF0D05C447F89D89D1FDDF" blockId="25.[809,1461,1523,1985]" lastBlockId="26.[128,779,144,1985]" lastPageId="26" lastPageNumber="27" pageId="25" pageNumber="26"> +When defining the calibration bounds for each taxon, we used a uniform distribution that includes the minimum and maximum ages of the taxa that we scored in the matrix following previous recommendations (Püschel +<emphasis id="B964DA31FFFF0D06C350F87C8DB4F860" box="[1106,1158,1961,1985]" italics="true" pageId="25" pageNumber="26">et al.</emphasis> +2020). This means that the temporal intervals may or may not correspond with the whole temporal interval between the first appearance datum (FAD) and last appearance datum (LAD) of a particular species, as the ages of specimens that we could not examine were excluded. In cases in which there was uncertainty about the stratigraphic location of a specific specimen (e.g., exact location not reported), this uncertainty was accounted for by incorporating the whole known temporal interval for that taxon (between the FAD and LAD). In terms of the exact ages used for each taxon, priority was given to specific radiometric dates of the strata associated with the scored specimens. However, in most cases, the age of a taxon is estimated from faunal correlations from sites with radiometric dates available and associated broadly with one or more SALMAs. See Supporting information, File S2 for information on the specific temporal interval used for each taxon and its justification. +</paragraph> +<paragraph id="8BAF0623FFFC0D05C79EFD538BAFFAB1" blockId="26.[128,779,144,1985]" pageId="26" pageNumber="27"> +The prior distribution and the parameters of the clock rate were derived from the methodology of +<bibRefCitation id="EF817BD2FFFC0D05C525FD738A3BFD1C" author="Gunnell" box="[551,777,677,701]" etAl="et al." firstAuthor="Gunnell" pageId="26" pageNumber="27" pagination="1 - 12" refId="ref43068" refString="Gunnell GF, Boyer DM, Friscia AR et al. Fossil lemurs from Egypt and Kenya suggest an African origin for Madagascar's aye-aye. Nature Communications 2018; 9: 1 - 12." type="journal article" year="2018"> +Gunnell +<emphasis id="B964DA31FFFC0D05C586FD738B85FD1C" box="[644,695,677,701]" italics="true" pageId="26" pageNumber="27">et al.</emphasis> +(2018) +</bibRefCitation> +using the packages ape (Paradis +<emphasis id="B964DA31FFFC0D05C6E0FD138B24FD7D" box="[482,534,708,732]" italics="true" pageId="26" pageNumber="27">et al.</emphasis> +2004) and fitdistrplus ( +<bibRefCitation id="EF817BD2FFFC0D05C789FD358B35FD5A" author="Delignette-Muller and Dutang" box="[139,519,739,764]" firstAuthor="Delignette-Muller" pageId="26" pageNumber="27" pagination="1 - 34" refId="ref41623" refString="Delignette-Muller ML, Dutang C. fitdistrplus: an R package for fitting distributions. Journal of Statistical Software 2004; 64: 1 - 34." type="journal article" year="2004">Delignette-Muller and Dutang 2004</bibRefCitation> +) from R v.2.2 (https:// www.R-project.org/) and the undated Bayesian consensus trees previously obtained. We selected the best model considering the Bayesian information criterion, which was a lognormal distributed clock rate prior with a mean of -3.1566009 and a standard deviation of 0.5451562. We modelled Branch rate variation employing the Independent Gamma Rate (IGR) relaxed clock model with an exponential distribution of rate 10 (default MrBayes setting). The fossilized birth-death (FBD) model ( +<bibRefCitation id="EF817BD2FFFC0D05C789FC2B88B4FBB4" author="Gavryushkina" box="[139,390,1021,1045]" etAl="et al." firstAuthor="Gavryushkina" pageId="26" pageNumber="27" refId="ref42118" refString="Gavryushkina A, Welch D, Stadler T et al. Bayesian inference of sampled ancestor trees for epidemiology and fossil calibration. PLoS Computational Biology 2014; 10: e 1003919. https: // doi. org / 10.1371 / journal. pcbi. 1003919" type="journal volume" year="2014"> +Gavryushkina +<emphasis id="B964DA31FFFC0D05C61DFC28887CFBB4" box="[287,334,1021,1045]" italics="true" pageId="26" pageNumber="27">et al.</emphasis> +2014 +</bibRefCitation> +, +<bibRefCitation id="EF817BD2FFFC0D05C693FC2B8B0EFBB4" author="Heath" box="[401,572,1021,1045]" etAl="et al." firstAuthor="Heath" pageId="26" pageNumber="27" pagination="2957 - 66" refId="ref43144" refString="Heath TA, Huelsenbeck JP, Stadler T. The fossilized birth - death process for coherent calibration of divergence-time estimates. Proceedings of the National Academy of Sciences of the United States of America 2014; 111: E 2957 - 66. https: // doi. org / 10.1073 / pnas. 1319091111" type="journal article" year="2014"> +Heath +<emphasis id="B964DA31FFFC0D05C6D7FC288B36FBB4" box="[469,516,1021,1045]" italics="true" pageId="26" pageNumber="27">et al.</emphasis> +2014 +</bibRefCitation> +) was used as a prior on divergence times with an exponential net diversification prior with rate 1, a beta turnover prior with shape parameters α = 1 and β = 1, a beta fossil sampling proportion prior with shape parameters α = 1 and β = 1, an extant sampling proportion of 1, and allowing fossil taxa to be sampled ancestors (samplestrat = random; Zhang +<emphasis id="B964DA31FFFC0D05C639FB6C8859FB70" box="[315,363,1209,1233]" italics="true" pageId="26" pageNumber="27">et al.</emphasis> +2016). The priors in the FBD model are diffuse priors that reflect the uncertainty in our prior expectation of how these parameters are distributed (Yang 2014). +</paragraph> +<paragraph id="8BAF0623FFFC0D05C79EFACE88A1F98B" blockId="26.[128,779,144,1985]" pageId="26" pageNumber="27"> +We ran the analyses using two independent runs of four chains for 40 million MCMC generations, removing the first 25% of the samples as burn-in. At the end of the analyses, the effective sample size for the parameters was> 200 and the deviation of split frequencies was below 0.01. Additionally, the runs were visually inspected using Tracer v.1.7.1 (Rambaut +<emphasis id="B964DA31FFFC0D05C5D9FA638A38FA6D" box="[731,778,1460,1484]" italics="true" pageId="26" pageNumber="27">et al.</emphasis> +2018) to make sure convergence and stationarity were achieved. The analyses were performed twice using the same priors and parameters for H1 and H2. +</paragraph> +<paragraph id="8BAF0623FFFC0D05C79EF9E78AB6FC16" blockId="26.[128,779,144,1985]" lastBlockId="26.[824,1477,144,951]" pageId="26" pageNumber="27"> +Preliminary results of the tip-dated Bayesian analyses showed that the node for the common ancestor of SANUs was potentially too old considering the fossil evidence, being dated around 68–69 Mya within the Maastrichtian (latest Cretaceous). This is not impossible and it corresponds well with recent molecular studies that have posited that interordinal origination and diversification of Placentalia occurred prior to the K/Pg boundary (long fuse model of placental diversification; Springer +<emphasis id="B964DA31FFFC0D05C5D6F8DB8A38F884" box="[724,778,1805,1829]" italics="true" pageId="26" pageNumber="27">et al.</emphasis> +2017). However, most workers consider that SANUs arrived to South America in the earliest Palaeocene (e.g., Reguero and Goin 2021), considering that well-sampled Maastrichtian fossil localities in South America do not have any SANU or other placental representatives (Rougier +<emphasis id="B964DA31FFFC0D05C6C7F87C88C4F860" box="[453,502,1961,1985]" italics="true" pageId="26" pageNumber="27">et al.</emphasis> +2021). Therefore, it might be expected that the age of their common ancestor is between the K/Pg boundary and the earliest Palaeocene, particularly if they are monophyletic. This is consistent with the fact that from the Early Eocene, different SANU orders diversified, showing an enormous disparity (Croft +<emphasis id="B964DA31FFFC0D05C376FED88D94FE84" box="[1140,1190,269,293]" italics="true" pageId="26" pageNumber="27">et al.</emphasis> +2020). Accordingly, we repeated the analyses constraining the age of the common ancestor of SANUs between 65.075 and 66 Mya, the minimum being the oldest age estimated for the kollpaniines (Muizon and Ladevèze 2020) and the maximum the age of the K/Pg boundary. For these new analyses, we tried four different approaches: (i) using the H1 matrix with no constraints on the topology, (ii) using the H2 matrix with no constraints on the topology, (iii) using the H1 matrix constraining the topology to be the same as in in the H1 undated tree, and (iv) using the H2 matrix constraining the topology to be the same as in the H2 undated tree. The last two approaches (iii and iv) were conducted to limit the influence of the fossil ages in the topology. This is because it is still an open question whether it is desirable to allow the age of fossils to have an influence on the topology of the tree (O’Reilly +<emphasis id="B964DA31FFFC0D05C221FD138C62FD7D" box="[1315,1360,708,732]" italics="true" pageId="26" pageNumber="27">et al.</emphasis> +2015), particularly in cases of rapid diversification in which the ages of many taxa are similar. Therefore, we used soft constraints in all cases except for the SANU common ancestor node in which we used a hard constraint because it is required by MrBayes to time-calibrate a specific node. The remainder of the parameters of the analyses were exactly the same as the other tip-dated Bayesian analyses (see above). +</paragraph> +</subSection> +</caption> +</subSubSection> +</caption> +</subSubSection> +</treatment> +</document> \ No newline at end of file diff --git a/data/03/B9/B7/03B9B735FFE90D10C7E1FF2E88DCF99D.xml b/data/03/B9/B7/03B9B735FFE90D10C7E1FF2E88DCF99D.xml new file mode 100644 index 00000000000..e0cb9a12fa2 --- /dev/null +++ b/data/03/B9/B7/03B9B735FFE90D10C7E1FF2E88DCF99D.xml @@ -0,0 +1,170 @@ +<document id="E2CF0296210FBB8FC54EEC5308EE5D2B" ID-DOI="10.1093/zoolinnean/zlae095" ID-ISSN="0024-4082" IM.bibliography_approvedBy="carolina" IM.illustrations_approvedBy="carolina" IM.materialsCitations_approvedBy="carolina" IM.metadata_approvedBy="carolina" IM.tables_requiresApprovalFor="GgImagineBatch,operationResults" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="carolina" checkinTime="1732847695013" checkinUser="plazi" docAuthor="Püschel, Hans P., Shelley, Sarah L., Williamson, Thomas E., Perini, Fernando A., Wible, John R. & Brusatte, Stephen L." docDate="2024" docId="03B9B735FFE90D10C7E1FF2E88DCF99D" docLanguage="en" docName="zlae095.pdf" docOrigin="Zoological Journal of the Linnean Society 202 (1)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlae095" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.9:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="9" docTitle="Amilnedwardsiidae" docType="treatment" docVersion="1" lastPageNumber="16" masterDocId="FF80CF4DFFE60D1FC702FFD68932FFA1" masterDocTitle="A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates" masterLastPageNumber="50" masterPageNumber="1" pageNumber="16" updateTime="1733769861150" updateUser="carolina"> +<mods:mods id="34E5D8C7CE12B6A52A04958BB276B84A" xmlns:mods="http://www.loc.gov/mods/v3"> +<mods:titleInfo id="C4444849331632AEF4DF7ECFD9F0054A"> +<mods:title id="3DB043A802034F55439D6EDF2B761F78">A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates</mods:title> +</mods:titleInfo> +<mods:name id="AAF6798E83D797FC73778F05E2C60B3B" type="personal"> +<mods:role id="5372D08F03600357DD5F0846A01DC94F"> +<mods:roleTerm id="3417C5DE0645E5FBD770E42C09535C3D">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="780B31BD1A885A9E8D3FC014E702BBF2">Püschel, Hans P.</mods:namePart> +<mods:affiliation id="965BAEE2CBFE86E5F0320052CEE55B7B">Millenium Nucleus Early Evolutionary Transitions of Mammals (EVOTEM), Red Palentológica U-Chile, Santiago, 7800003, Chile & School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, EH 9 3 FE, United Kingdom</mods:affiliation> +<mods:nameIdentifier id="1C5FA41D8A84500A5810D6A8B5B39547" type="email">hpuschelr@gmail.com</mods:nameIdentifier> +</mods:name> +<mods:name id="B7BFB9E8E6CF86B26B619DD0F427CA6F" type="personal"> +<mods:role id="C63F57CB91354DC48E6BC21863F541E4"> +<mods:roleTerm id="8CC69FAFF2C9841566C2AD613AE5DC91">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="3A3D3F7B0E97452AF8D2F2EF334E83BD">Shelley, Sarah L.</mods:namePart> +<mods:affiliation id="06345BAB95F3298983D445DC62A6CF49">School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, EH 9 3 FE, United Kingdom</mods:affiliation> +</mods:name> +<mods:name id="0E80C7BFC016CB3388019B21D7EDC195" type="personal"> +<mods:role id="122FBB9882843759521A368FBD199F20"> +<mods:roleTerm id="CB82373B3CF5625FFE7248173FDCD836">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="75DB5082C8B967B325212D46F754B33B">Williamson, Thomas E.</mods:namePart> +<mods:affiliation id="FFC2B8B720D15B27481860E2D58DE66B">New Mexico Museum of Natural History and Science, Albuquerque, NM 87104, United States</mods:affiliation> +</mods:name> +<mods:name id="846CE226853DB5B7AB2CA0EDF4E99818" type="personal"> +<mods:role id="EB735016112EF3F67550D998BABB4CBF"> +<mods:roleTerm id="F8A7C5BF3925D092D9A3F1DCD9D1228B">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="A9183B2F8B22A19F8DFFACCAB8995A01">Perini, Fernando A.</mods:namePart> +<mods:affiliation id="75DAB92D587FC39A042D70C4A0279408">Instituto de Ciência Biológicas, UFMG, BH 31270 - 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It is based on two genera (i.e., +<emphasis id="B964DA31FFE90D10C687FE888B25FED7" box="[389,535,350,374]" italics="true" pageId="15" pageNumber="16">Amilnedwarsia</emphasis> +<bibRefCitation id="EF817BD2FFE90D10C522FE888BFEFED7" author="Ameghino" box="[544,716,350,374]" firstAuthor="Ameghino" pageId="15" pageNumber="16" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +and +<taxonomicName id="4C107DA0FFE90D10C773FEA88890FE34" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[113,418,381,405]" class="Mammalia" family="Amilnedwardsiidae" genus="Rutimeyeria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C773FEA889DAFE34" box="[113,232,382,405]" italics="true" pageId="15" pageNumber="16">Rutimeyeria</emphasis> +<bibRefCitation id="EF817BD2FFE90D10C7F1FEAB8890FE34" author="Ameghino" box="[243,418,381,405]" firstAuthor="Ameghino" pageId="15" pageNumber="16" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation> +</taxonomicName> +) and three species named after small bunodont isolated upper molars whose positions were assigned tentatively by Soria (1989a) by comparing them with the indaleciid +<taxonomicName id="4C107DA0FFE90D10C606FE0D88FAFE52" authorityName="Bond & Vucetich" authorityYear="1983" box="[260,456,475,499]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="grandensis"> +<emphasis id="B964DA31FFE90D10C606FE0D88FAFE52" box="[260,456,475,499]" italics="true" pageId="15" pageNumber="16">Indalecia grandensis</emphasis> +</taxonomicName> +. +<taxonomicName id="4C107DA0FFE90D10C6D6FE0D8BA8FE52" box="[468,666,475,499]" class="Mammalia" family="Amilnedwardsiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Amilnedwardsiidae</taxonomicName> +was considered to be the most basal family of the order +<taxonomicName id="4C107DA0FFE90D10C580FE2D89D9FD93" authority="Soria, 1989" authorityName="Soria" authorityYear="1989" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Notopterna Soria, 1989</taxonomicName> +, which also includes the families +<taxonomicName id="4C107DA0FFE90D10C54DFDCC8BF9FD93" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[591,715,538,562]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Indaleciidae</taxonomicName> +and +<taxonomicName id="4C107DA0FFE90D10C773FDEF888BFDF0" authority="(Soria 1989 b)" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[113,441,569,593]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Notonychopidae (Soria 1989b)</taxonomicName> +. +<taxonomicName id="4C107DA0FFE90D10C6C4FDEF8B0CFDF1" authorityName="Soria" authorityYear="1989" box="[454,574,569,592]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Notopterna</taxonomicName> +is diagnosed by a well-developed and buccally projected parastyle and the absence of a mesostyle in the upper molars, bicrescentic and selenodont lower molars, and anteriorly acuminate petrosals, among other anatomical features that distinguish them from other SANUs (Soria 1989a). Soria (1989a, b) argued that, considering their dental similarities, ‘amilnedwarsids’ could be the structural ancestor of both indaleciids and notonychopids. However, a phylogenetic analysis failed to recover the members of +<taxonomicName id="4C107DA0FFE90D10C558FCE28BE0FCEA" authorityName="Soria" authorityYear="1989" box="[602,722,820,843]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Notopterna</taxonomicName> +as a monophyletic group ( +<bibRefCitation id="EF817BD2FFE90D10C656FC858B4FFCCA" author="Bonaparte and Morales" box="[340,637,851,875]" firstAuthor="Bonaparte" pageId="15" pageNumber="16" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales 1997</bibRefCitation> +), which has led to some authors to include its members tentatively within +<taxonomicName id="4C107DA0FFE90D10C773FC4489ECFC0B" box="[113,222,914,938]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Litopterna</taxonomicName> +(e.g., +<bibRefCitation id="EF817BD2FFE90D10C622FC4488F8FC0B" author="Gelfo" box="[288,458,914,938]" etAl="et al." firstAuthor="Gelfo" pageId="15" pageNumber="16" pagination="285 - 91" refId="ref42550" refString="Gelfo JN, Lopez GM, Lorente M. Los ungulados arcaicos de America del Sur: ' Condylarthra' y Litopterna. Contribuciones Cientificas Del Museo Argentino de Ciencias Naturales ' Bernardino Rivadavia' 2016; 6: 285 - 91." type="journal article" year="2016"> +Gelfo +<emphasis id="B964DA31FFE90D10C65DFC4488A2FC0B" box="[351,400,914,938]" italics="true" pageId="15" pageNumber="16">et al.</emphasis> +2016 +</bibRefCitation> +, Croft +<emphasis id="B964DA31FFE90D10C514FC448B75FC0B" box="[534,583,914,938]" italics="true" pageId="15" pageNumber="16">et al.</emphasis> +2020, +<bibRefCitation id="EF817BD2FFE90D10C58CFC448997FC68" author="Goin" etAl="et al." firstAuthor="Goin" pageId="15" pageNumber="16" pagination="1029 - 53" refId="ref42920" refString="Goin FJ, Gelfo JN, Ortiz-Jaureguizar E, et al. Mamiferos del Banco Negro Inferior, Formacion Salamanca, Cuenca del Golfo de San Jorge. In: Giacosa RE (ed.), Geologia y Recursos Naturales de la Provincia de Chubut. Puerto Madryn, Relatorio del XXI Congreso Geologico Argentino, 2022, 1029 - 53." type="book chapter" year="2022"> +Goin +<emphasis id="B964DA31FFE90D10C5C8FC448BC9FC0B" box="[714,763,914,938]" italics="true" pageId="15" pageNumber="16">et al.</emphasis> +2022 +</bibRefCitation> +). However, this is a problematic assumption as +<bibRefCitation id="EF817BD2FFE90D10C593FC678873FC49" author="Bonaparte and Morales" firstAuthor="Bonaparte" pageId="15" pageNumber="16" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation> +used a matrix that includes only litopterns, so the broader relationships of notopterns have not yet been tested in an analysis that includes other SANUs such as notoungulates and astrapotheres. +</paragraph> +<paragraph id="8BAF0623FFE90D10C78FFB988B46FA3E" blockId="15.[112,764,287,1596]" pageId="15" pageNumber="16"> +<emphasis id="B964DA31FFE90D10C78FFB988811FBC7" box="[141,291,1102,1126]" italics="true" pageId="15" pageNumber="16">Amilnedwardia</emphasis> +and +<taxonomicName id="4C107DA0FFE90D10C655FB9888FCFBC4" authorityName="Ameghino" authorityYear="1901" box="[343,462,1102,1125]" class="Mammalia" family="Amilnedwardsiidae" genus="Rutimeyeria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C655FB9888FCFBC4" box="[343,462,1102,1125]" italics="true" pageId="15" pageNumber="16">Rutimeyeria</emphasis> +</taxonomicName> +were previously referred tentatively to the family +<taxonomicName id="4C107DA0FFE90D10C64EFBBB88CEFB24" box="[332,508,1133,1157]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName> +and regarded as close to +<taxonomicName id="4C107DA0FFE90D10C771FB5A8837FB05" box="[115,261,1164,1188]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="genus"> +<emphasis id="B964DA31FFE90D10C771FB5A8837FB05" box="[115,261,1164,1188]" italics="true" pageId="15" pageNumber="16">Victorlemoinea</emphasis> +</taxonomicName> +(Simpson 1945, 1948), although the latter now is now considered to be part of the family +<taxonomicName id="4C107DA0FFE90D10C511FB7A8BC5FB65" box="[531,759,1196,1220]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName> +. Cifelli (1983a)considered referred specimens of these taxa to be too fragmentary to determine a clear affinity,but suggested the referred specimens represent deciduous teeth of a henricosborniid notoungulate based on the presence of a metaconule-derived crest connecting with the ectoloph wall. Indeed, probably due to their limited fossil record, thus far no members of this family have ever been included in a phylogenetic analysis. +</paragraph> +<paragraph id="8BAF0623FFE90D10C78FFA7088DCF99D" blockId="15.[112,764,287,1596]" pageId="15" pageNumber="16"> +If we consider +<taxonomicName id="4C107DA0FFE90D10C633FA7088C5FA1F" box="[305,503,1446,1470]" class="Mammalia" family="Amilnedwardsiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Amilnedwardsiidae</taxonomicName> +to be a valid family, as its three species all come from Patagonian fossil localities of a similar age that correlate with the Vacan SALMA, the temporal interval of this family is ~45–42 Mya ( +<figureCitation id="132B1AA6FFE90D10C6F6F9D28B09F9BD" box="[500,571,1540,1564]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="15" pageNumber="16">Fig. 2B</figureCitation> +; Supporting information, +<tableCitation id="C6923398FFE90D10C7C7F9F2882DF99D" box="[197,287,1572,1596]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="15" pageNumber="16">Table S1</tableCitation> +; +<bibRefCitation id="EF817BD2FFE90D10C628F9F288EFF99D" author="Krause" box="[298,477,1572,1596]" etAl="et al." firstAuthor="Krause" pageId="15" pageNumber="16" pagination="886 - 903" refId="ref43716" refString="Krause JM, Clyde WC, Ibanez-Mejia M et al. New age constraints for Early Paleogene strata of central Patagonia, Argentina: implications for the timing of South American Land Mammal Ages. 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Pérez, +<date id="9473104BFFC1A77BFC85FC69FC2BFB9E" box="[892,1017,1023,1047]" pageId="10" pageNumber="11" value="2020-06-02">2 June 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC1A77BFCACFB88FC2EFBFC" blockId="10.[825,1475,692,1987]" pageId="10" pageNumber="11"> +20200602-MRS-MTP6, Pharillons de Maïre, cliff at +<quantity id="27359B6EFFC1A77BFA70FB89FA10FBBF" box="[1417,1474,1055,1079]" metricMagnitude="1" metricUnit="m" metricValue="2.0" pageId="10" pageNumber="11" unit="m" value="20.0">20 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC1A77BFC12FBA8FBE2FBDC" box="[1003,1072,1086,1109]" name="France" pageId="10" pageNumber="11">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC1A77BFBC7FBA8FB68FBDC" box="[1086,1210,1084,1110]" degrees="43" direction="north" minutes="12" orientation="latitude" pageId="10" pageNumber="11" precision="15" seconds="27" value="43.2075"> +43°12 +<emphasis id="D2B9EA99FFC1A77BFB84FBAAFB51FBDC" box="[1149,1155,1084,1109]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +27″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC1A77BFB3DFBA8FAFDFBDC" box="[1220,1327,1084,1110]" degrees="5" direction="east" minutes="20" orientation="longitude" pageId="10" pageNumber="11" precision="15" seconds="17" value="5.3380556"> +5°20 +<emphasis id="D2B9EA99FFC1A77BFB0AFBAAFB2BFBDC" box="[1267,1273,1084,1109]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +17″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC1A77BFC85FBCBFC2BFBFC" box="[892,1017,1117,1141]" pageId="10" pageNumber="11" value="2020-06-02">2 June 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC1A77BFCACFBEAFC2EFB5A" blockId="10.[825,1475,692,1987]" pageId="10" pageNumber="11"> +20200602-MRS-MTP7, Pharillons de Maïre, cliff at +<quantity id="27359B6EFFC1A77BFA70FBEBFA10FB1D" box="[1417,1474,1149,1173]" metricMagnitude="1" metricUnit="m" metricValue="2.0" pageId="10" pageNumber="11" unit="m" value="20.0">20 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC1A77BFC12FB0AFBE2FB3A" box="[1003,1072,1180,1203]" name="France" pageId="10" pageNumber="11">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC1A77BFBC7FB0AFB68FB3A" box="[1086,1210,1178,1204]" degrees="43" direction="north" minutes="12" orientation="latitude" pageId="10" pageNumber="11" precision="15" seconds="27" value="43.2075"> +43°12 +<emphasis id="D2B9EA99FFC1A77BFB84FB0CFB51FB3A" box="[1149,1155,1178,1203]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +27″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC1A77BFB3DFB0AFAFDFB3A" box="[1220,1327,1178,1204]" degrees="5" direction="east" minutes="20" orientation="longitude" pageId="10" pageNumber="11" precision="15" seconds="17" value="5.3380556"> +5°20 +<emphasis id="D2B9EA99FFC1A77BFB0AFB0CFB2BFB3A" box="[1267,1273,1178,1203]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +17″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC1A77BFC85FB2DFC2BFB5A" box="[892,1017,1211,1235]" pageId="10" pageNumber="11" value="2020-06-02">2 June 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC1A77BFCACFB4CFC2EFAB8" blockId="10.[825,1475,692,1987]" pageId="10" pageNumber="11"> +20200602-MRS-MTP8, Pharillons de Maïre, cliff at +<quantity id="27359B6EFFC1A77BFA70FB4DFA10FB7B" box="[1417,1474,1243,1267]" metricMagnitude="1" metricUnit="m" metricValue="2.0" pageId="10" pageNumber="11" unit="m" value="20.0">20 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC1A77BFC12FB6CFBE2FA98" box="[1003,1072,1274,1297]" name="France" pageId="10" pageNumber="11">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC1A77BFBC7FB6CFB68FA98" box="[1086,1210,1272,1298]" degrees="43" direction="north" minutes="12" orientation="latitude" pageId="10" pageNumber="11" precision="15" seconds="27" value="43.2075"> +43°12 +<emphasis id="D2B9EA99FFC1A77BFB84FB6EFB51FA98" box="[1149,1155,1272,1297]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +27″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC1A77BFB3DFB6CFAFDFA98" box="[1220,1327,1272,1298]" degrees="5" direction="east" minutes="20" orientation="longitude" pageId="10" pageNumber="11" precision="15" seconds="17" value="5.3380556"> +5°20 +<emphasis id="D2B9EA99FFC1A77BFB0AFB6EFB2BFA98" box="[1267,1273,1272,1297]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +17″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC1A77BFC85FA8FFC2BFAB8" box="[892,1017,1305,1329]" pageId="10" pageNumber="11" value="2020-06-02">2 June 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC1A77BFCACFAAEFBCDFA06" blockId="10.[825,1475,692,1987]" pageId="10" pageNumber="11"> +20160607-GIJ-JC1, Las Gemelas island, rock substrate at +<quantity id="27359B6EFFC1A77BFA51FAAFFC9CFAE6" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="10" pageNumber="11" unit="m" value="12.0">12 m</quantity> +depth, Gijon, +<collectingCountry id="98DA761BFFC1A77BFC10FACEFBF0FAF9" box="[1001,1058,1368,1392]" name="Spain" pageId="10" pageNumber="11">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFC1A77BFBCBFACEFB7DFAE6" box="[1074,1199,1366,1392]" degrees="43" direction="north" minutes="33" orientation="latitude" pageId="10" pageNumber="11" precision="15" seconds="22" value="43.55611"> +43°33 +<emphasis id="D2B9EA99FFC1A77BFB88FAC0FBA5FAE6" box="[1137,1143,1366,1391]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC1A77BFB43FACEFAE3FAE6" box="[1210,1329,1366,1392]" degrees="5" direction="west" minutes="36" orientation="longitude" pageId="10" pageNumber="11" precision="15" seconds="31" value="-5.608611"> +5°36 +<emphasis id="D2B9EA99FFC1A77BFB13FAC0FB22FAE6" box="[1258,1264,1366,1391]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +31″W +</geoCoordinate> +). Collector: J. Cristobo, +<date id="9473104BFFC1A77BFC67FAEEFBC9FA06" box="[926,1051,1399,1423]" pageId="10" pageNumber="11" value="2016-06-07">7 June 2016</date> +. +</paragraph> +<paragraph id="E072368BFFC1A77BFCACFA00FBECFA64" blockId="10.[825,1475,692,1987]" pageId="10" pageNumber="11"> +20200624-HEN-BMNC2, Belhara, on reef flat at +<quantity id="27359B6EFFC1A77BFAB1FA01FAAEFA27" box="[1352,1404,1431,1455]" metricMagnitude="1" metricUnit="m" metricValue="2.2" pageId="10" pageNumber="11" unit="m" value="22.0">22 m</quantity> +depth, Hendaye, +<collectingCountry id="98DA761BFFC1A77BFC66FA20FC36FA44" box="[927,996,1462,1485]" name="France" pageId="10" pageNumber="11">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC1A77BFC08FA20FBBCFA44" box="[1009,1134,1460,1486]" degrees="43" direction="north" minutes="24" orientation="latitude" pageId="10" pageNumber="11" precision="15" seconds="00" value="43.4"> +43°24 +<emphasis id="D2B9EA99FFC1A77BFBC9FA22FBE4FA44" box="[1072,1078,1460,1485]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +00″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC1A77BFB8FFA20FB3CFA44" box="[1142,1262,1460,1486]" degrees="1" direction="west" minutes="43" orientation="longitude" pageId="10" pageNumber="11" precision="15" seconds="03" value="-1.7175001"> +1°43 +<emphasis id="D2B9EA99FFC1A77BFB5CFA22FB79FA44" box="[1189,1195,1460,1485]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +03″W +</geoCoordinate> +). Collector: M. N. de Casamajor, +<date id="9473104BFFC1A77BFC48FA43FBE8FA64" box="[945,1082,1493,1517]" pageId="10" pageNumber="11" value="2020-06-24">24 June 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC1A77BFCACFA63FB65F9C2" blockId="10.[825,1475,692,1987]" pageId="10" pageNumber="11"> +20200814-HEN-AMNC3, Abbadia, on reef flat at +<quantity id="27359B6EFFC1A77BFA7FFA63FA10F985" box="[1414,1474,1525,1549]" metricMagnitude="1" metricUnit="m" metricValue="1.3" pageId="10" pageNumber="11" unit="m" value="13.0">13 m</quantity> +depth, Hendaye, +<collectingCountry id="98DA761BFFC1A77BFC17F982FBE1F9A2" box="[1006,1075,1556,1579]" name="France" pageId="10" pageNumber="11">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC1A77BFBBCF982FB10F9A2" box="[1093,1218,1554,1580]" degrees="43" direction="north" minutes="23" orientation="latitude" pageId="10" pageNumber="11" precision="15" seconds="56" value="43.39889"> +43°23 +<emphasis id="D2B9EA99FFC1A77BFB7DF984FB58F9A2" box="[1156,1162,1554,1579]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +56″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC1A77BFB36F982FA94F9A2" box="[1231,1350,1554,1580]" degrees="1" direction="west" minutes="45" orientation="longitude" pageId="10" pageNumber="11" precision="15" seconds="28" value="-1.7577778"> +1°45 +<emphasis id="D2B9EA99FFC1A77BFB07F984FAD6F9A2" box="[1278,1284,1554,1579]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +28″W +</geoCoordinate> +). Collector: M. N. de Casamajor, +<date id="9473104BFFC1A77BFBF6F9A5FB61F9C2" box="[1039,1203,1587,1611]" pageId="10" pageNumber="11" value="2020-08-14">14 August 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC1A77BFCACF9C4FB89F920" blockId="10.[825,1475,692,1987]" pageId="10" pageNumber="11"> +20210630-CIB-BPC1, Baloo tunnel at +<quantity id="27359B6EFFC1A77BFADCF9C5FAB9F9E3" box="[1317,1387,1619,1643]" metricMagnitude="1" metricUnit="m" metricValue="1.4" pageId="10" pageNumber="11" unit="m" value="14.0">14 m</quantity> +depth, Ciboure, +<collectingCountry id="98DA761BFFC1A77BFC59F9E4FC37F900" box="[928,997,1650,1673]" name="France" pageId="10" pageNumber="11">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC1A77BFC03F9E4FB58F900" box="[1018,1162,1648,1674]" degrees="43" direction="north" minutes="24" orientation="latitude" pageId="10" pageNumber="11" precision="1" seconds="59.0" value="43.41639"> +43°24 +<emphasis id="D2B9EA99FFC1A77BFBC3F9E6FB92F900" box="[1082,1088,1648,1673]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +59.0″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC1A77BFB63F9E4FAF6F900" box="[1178,1316,1648,1674]" degrees="1" direction="west" minutes="39" orientation="longitude" pageId="10" pageNumber="11" precision="1" seconds="52.2" value="-1.6645"> +1°39 +<emphasis id="D2B9EA99FFC1A77BFB30F9E6FB1DF900" box="[1225,1231,1648,1673]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +52.2″W +</geoCoordinate> +). Collector: P. Chevaldonné, +<date id="9473104BFFC1A77BFC34F907FB84F920" box="[973,1110,1681,1705]" pageId="10" pageNumber="11" value="2021-06-30">30 June 2021</date> +. +</paragraph> +<paragraph id="E072368BFFC1A77BFCACF926FB89F88E" blockId="10.[825,1475,692,1987]" pageId="10" pageNumber="11"> +20210630-CIB-BPC3, Baloo tunnel at +<quantity id="27359B6EFFC1A77BFADCF926FAB9F941" box="[1317,1387,1712,1736]" metricMagnitude="1" metricUnit="m" metricValue="1.4" pageId="10" pageNumber="11" unit="m" value="14.0">14 m</quantity> +depth, Ciboure, +<collectingCountry id="98DA761BFFC1A77BFC59F946FC37F96E" box="[928,997,1744,1767]" name="France" pageId="10" pageNumber="11">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC1A77BFC03F946FB58F96E" box="[1018,1162,1742,1768]" degrees="43" direction="north" minutes="24" orientation="latitude" pageId="10" pageNumber="11" precision="1" seconds="59.0" value="43.41639"> +43°24 +<emphasis id="D2B9EA99FFC1A77BFBC3F958FB92F96E" box="[1082,1088,1742,1767]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +59.0″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC1A77BFB63F946FAF6F96E" box="[1178,1316,1742,1768]" degrees="1" direction="west" minutes="39" orientation="longitude" pageId="10" pageNumber="11" precision="1" seconds="52.2" value="-1.6645"> +1°39 +<emphasis id="D2B9EA99FFC1A77BFB30F958FB1DF96E" box="[1225,1231,1742,1767]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +52.2″W +</geoCoordinate> +). Collector: P. Chevaldonné, +<date id="9473104BFFC1A77BFC34F979FB84F88E" box="[973,1110,1775,1799]" pageId="10" pageNumber="11" value="2021-06-30">30 June 2021</date> +. +</paragraph> +<paragraph id="E072368BFFC1A77BFCACF898FC72F8EC" blockId="10.[825,1475,692,1987]" pageId="10" pageNumber="11"> +20210726-CIB-BTP1, Baloo tunnel at +<quantity id="27359B6EFFC1A77BFB1DF898FAC8F8AF" box="[1252,1306,1806,1830]" metricMagnitude="1" metricUnit="m" metricValue="1.4" pageId="10" pageNumber="11" unit="m" value="14.0">14 m</quantity> +depth, Ciboure, +<collectingCountry id="98DA761BFFC1A77BFCC0F8B8FCACF8CC" box="[825,894,1838,1861]" name="France" pageId="10" pageNumber="11">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC1A77BFC68F8B8FBF3F8CC" box="[913,1057,1836,1862]" degrees="43" direction="north" minutes="24" orientation="latitude" pageId="10" pageNumber="11" precision="1" seconds="59.0" value="43.41639"> +43°24 +<emphasis id="D2B9EA99FFC1A77BFC28F8BAFC05F8CC" box="[977,983,1836,1861]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +59.0″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC1A77BFBD6F8B8FB6BF8CC" box="[1071,1209,1836,1862]" degrees="1" direction="west" minutes="39" orientation="longitude" pageId="10" pageNumber="11" precision="1" seconds="52.2" value="-1.6645"> +1°39 +<emphasis id="D2B9EA99FFC1A77BFBA7F8BAFBB6F8CC" box="[1118,1124,1836,1861]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +52.2″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC1A77BFA51F8B8FC4EF8EC" pageId="10" pageNumber="11" value="2021-07-26">26 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFC1A77BFCACF8FAFC72F84A" blockId="10.[825,1475,692,1987]" pageId="10" pageNumber="11"> +20210726-CIB-BTP2, Baloo tunnel at +<quantity id="27359B6EFFC1A77BFB1DF8FAFAC8F80D" box="[1252,1306,1900,1924]" metricMagnitude="1" metricUnit="m" metricValue="1.4" pageId="10" pageNumber="11" unit="m" value="14.0">14 m</quantity> +depth, Ciboure, +<collectingCountry id="98DA761BFFC1A77BFCC0F81AFCACF82A" box="[825,894,1932,1955]" name="France" pageId="10" pageNumber="11">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC1A77BFC68F81AFBF3F82A" box="[913,1057,1930,1956]" degrees="43" direction="north" minutes="24" orientation="latitude" pageId="10" pageNumber="11" precision="1" seconds="59.0" value="43.41639"> +43°24 +<emphasis id="D2B9EA99FFC1A77BFC28F81CFC05F82A" box="[977,983,1930,1955]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +59.0″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC1A77BFBD6F81AFB6BF82A" box="[1071,1209,1930,1956]" degrees="1" direction="west" minutes="39" orientation="longitude" pageId="10" pageNumber="11" precision="1" seconds="52.2" value="-1.6645"> +1°39 +<emphasis id="D2B9EA99FFC1A77BFBA7F81CFBB6F82A" box="[1118,1124,1930,1955]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis> +52.2″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC1A77BFA51F81AFC4EF84A" pageId="10" pageNumber="11" value="2021-07-26">26 July 2021</date> +. +</paragraph> +<caption id="B4B26603FFC0A77AFF88FB42FDB5FA81" pageId="11" pageNumber="12" startId="11.[113,178,1236,1260]" targetBox="[117,1456,146,1201]" targetPageId="11" targetType="figure"> +<paragraph id="E072368BFFC0A77AFF88FB42FDB5FA81" blockId="11.[113,1456,1236,1288]" pageId="11" pageNumber="12"> +<emphasis id="D2B9EA99FFC0A77AFF88FB42FF1BFB65" bold="true" box="[113,201,1236,1260]" pageId="11" pageNumber="12">Figure 4.</emphasis> +<taxonomicName id="27CD4D08FFC0A77AFF37FB43FED4FB65" box="[206,262,1237,1260]" class="Demospongiae" family="Irciniidae" genus="Ircinia" kingdom="Animalia" order="Dictyoceratida" pageId="11" pageNumber="12" phylum="Porifera" rank="species" species="situ"> +<emphasis id="D2B9EA99FFC0A77AFF37FB43FED4FB65" box="[206,262,1237,1260]" italics="true" pageId="11" pageNumber="12">In situ</emphasis> +</taxonomicName> +pictures of +<taxonomicName id="27CD4D08FFC0A77AFE8AFB43FDD1FB65" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[371,515,1236,1260]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="11" pageNumber="12" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC0A77AFE8AFB43FDD1FB65" box="[371,515,1236,1260]" italics="true" pageId="11" pageNumber="12">Spongia lamella</emphasis> +</taxonomicName> +in the Northeast Atlantic Ocean and the Mediterranean Sea, A, Baloo tunnel, Ciboure, France. B, C, Mouro Island, Santander, Spain. D, Marseille, France. +</paragraph> +</caption> +<paragraph id="E072368BFFC0A77AFF74FAAEFF0AFA06" blockId="11.[113,763,1336,1987]" pageId="11" pageNumber="12"> +20210726-CIB-BTP3, Baloo tunnel at +<quantity id="27359B6EFFC0A77AFDE5FAAEFD80FAD9" box="[540,594,1336,1360]" metricMagnitude="1" metricUnit="m" metricValue="1.4" pageId="11" pageNumber="12" unit="m" value="14.0">14 m</quantity> +depth, Ciboure, +<collectingCountry id="98DA761BFFC0A77AFF88FACEFF64FAE6" box="[113,182,1368,1391]" name="France" pageId="11" pageNumber="12">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC0A77AFF30FACEFE8BFAE6" box="[201,345,1366,1392]" degrees="43" direction="north" minutes="24" orientation="latitude" pageId="11" pageNumber="12" precision="1" seconds="59.0" value="43.41639"> +43°24 +<emphasis id="D2B9EA99FFC0A77AFEF0FAC0FEDDFAE6" box="[265,271,1366,1391]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +59.0″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC0A77AFE9EFACEFE23FAE6" box="[359,497,1366,1392]" degrees="1" direction="west" minutes="39" orientation="longitude" pageId="11" pageNumber="12" precision="1" seconds="52.2" value="-1.6645"> +1°39 +<emphasis id="D2B9EA99FFC0A77AFE6FFAC0FE4EFAE6" box="[406,412,1366,1391]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +52.2″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC0A77AFD19FAC1FF06FA06" pageId="11" pageNumber="12" value="2021-07-26">26 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFC0A77AFF74FA00FF0AFA64" blockId="11.[113,763,1336,1987]" pageId="11" pageNumber="12"> +20210726-CIB-BTP4, Baloo tunnel at +<quantity id="27359B6EFFC0A77AFDE5FA00FD80FA27" box="[540,594,1430,1454]" metricMagnitude="1" metricUnit="m" metricValue="1.4" pageId="11" pageNumber="12" unit="m" value="14.0">14 m</quantity> +depth, Ciboure, +<collectingCountry id="98DA761BFFC0A77AFF88FA20FF64FA44" box="[113,182,1462,1485]" name="France" pageId="11" pageNumber="12">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC0A77AFF30FA20FE8BFA44" box="[201,345,1460,1486]" degrees="43" direction="north" minutes="24" orientation="latitude" pageId="11" pageNumber="12" precision="1" seconds="59.0" value="43.41639"> +43°24 +<emphasis id="D2B9EA99FFC0A77AFEF0FA22FEDDFA44" box="[265,271,1460,1485]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +59.0″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC0A77AFE9EFA20FE23FA44" box="[359,497,1460,1486]" degrees="1" direction="west" minutes="39" orientation="longitude" pageId="11" pageNumber="12" precision="1" seconds="52.2" value="-1.6645"> +1°39 +<emphasis id="D2B9EA99FFC0A77AFE6FFA22FE4EFA44" box="[406,412,1460,1485]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +52.2″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC0A77AFD19FA23FF06FA64" pageId="11" pageNumber="12" value="2021-07-26">26 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFC0A77AFF74FA62FF0AF9C2" blockId="11.[113,763,1336,1987]" pageId="11" pageNumber="12"> +20210726-CIB-BTP7, Baloo tunnel at +<quantity id="27359B6EFFC0A77AFDE5FA62FD80F985" box="[540,594,1524,1548]" metricMagnitude="1" metricUnit="m" metricValue="1.4" pageId="11" pageNumber="12" unit="m" value="14.0">14 m</quantity> +depth, Ciboure, +<collectingCountry id="98DA761BFFC0A77AFF88F982FF64F9A2" box="[113,182,1556,1579]" name="France" pageId="11" pageNumber="12">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC0A77AFF30F982FE8BF9A2" box="[201,345,1554,1580]" degrees="43" direction="north" minutes="24" orientation="latitude" pageId="11" pageNumber="12" precision="1" seconds="59.0" value="43.41639"> +43°24 +<emphasis id="D2B9EA99FFC0A77AFEF0F984FEDDF9A2" box="[265,271,1554,1579]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +59.0″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC0A77AFE9EF982FE23F9A2" box="[359,497,1554,1580]" degrees="1" direction="west" minutes="39" orientation="longitude" pageId="11" pageNumber="12" precision="1" seconds="52.2" value="-1.6645"> +1°39 +<emphasis id="D2B9EA99FFC0A77AFE6FF984FE4EF9A2" box="[406,412,1554,1579]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +52.2″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC0A77AFD19F985FF06F9C2" pageId="11" pageNumber="12" value="2021-07-26">26 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFC0A77AFF74F9C4FF0AF920" blockId="11.[113,763,1336,1987]" pageId="11" pageNumber="12"> +20210726-CIB-BTP10, Baloo tunnel at +<quantity id="27359B6EFFC0A77AFDDBF9C4FD84F9E3" box="[546,598,1618,1642]" metricMagnitude="1" metricUnit="m" metricValue="1.4" pageId="11" pageNumber="12" unit="m" value="14.0">14 m</quantity> +depth, Ciboure, +<collectingCountry id="98DA761BFFC0A77AFF88F9E4FF64F900" box="[113,182,1650,1673]" name="France" pageId="11" pageNumber="12">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC0A77AFF30F9E4FE8BF900" box="[201,345,1648,1674]" degrees="43" direction="north" minutes="24" orientation="latitude" pageId="11" pageNumber="12" precision="1" seconds="59.0" value="43.41639"> +43°24 +<emphasis id="D2B9EA99FFC0A77AFEF0F9E6FEDDF900" box="[265,271,1648,1673]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +59.0″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC0A77AFE9EF9E4FE23F900" box="[359,497,1648,1674]" degrees="1" direction="west" minutes="39" orientation="longitude" pageId="11" pageNumber="12" precision="1" seconds="52.2" value="-1.6645"> +1°39 +<emphasis id="D2B9EA99FFC0A77AFE6FF9E6FE4EF900" box="[406,412,1648,1673]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +52.2″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC0A77AFD19F9E7FF06F920" pageId="11" pageNumber="12" value="2021-07-26">26 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFC0A77AFF74F926FD29F88E" blockId="11.[113,763,1336,1987]" pageId="11" pageNumber="12"> +20210729-ESP-MTP5, western part of Mouro island, semi-dark to dark cave, between 3 and +<quantity id="27359B6EFFC0A77AFE32F946FDD2F96E" box="[459,512,1744,1768]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="11" pageNumber="12" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFC0A77AFD38F959FD28F96E" box="[705,762,1743,1767]" name="Spain" pageId="11" pageNumber="12">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFC0A77AFF80F979FEDAF88F" box="[121,264,1773,1799]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="11" pageNumber="12" precision="1" seconds="22.0" value="43.47278"> +43°28 +<emphasis id="D2B9EA99FFC0A77AFF40F97BFF6DF88F" box="[185,191,1773,1798]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +22.0″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC0A77AFEE8F979FE49F88F" box="[273,411,1773,1799]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="11" pageNumber="12" precision="1" seconds="21.7" value="-3.7560277"> +3°45 +<emphasis id="D2B9EA99FFC0A77AFEB9F97BFE94F88F" box="[320,326,1773,1798]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +21.7″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC0A77AFD8DF979FD24F88E" box="[628,758,1775,1799]" pageId="11" pageNumber="12" value="2021-07-29">29 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFC0A77AFF74F898FE13F80D" blockId="11.[113,763,1336,1987]" pageId="11" pageNumber="12"> +20210730-ESP-M2TP2, eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and +<quantity id="27359B6EFFC0A77AFF65F8DBFF1DF8ED" box="[156,207,1869,1893]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="11" pageNumber="12" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFC0A77AFE70F8DBFE10F8EC" box="[393,450,1869,1893]" name="Spain" pageId="11" pageNumber="12">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFC0A77AFE37F8DBFD8CF8ED" box="[462,606,1867,1893]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="11" pageNumber="12" precision="1" seconds="21.1" value="43.472527"> +43°28 +<emphasis id="D2B9EA99FFC0A77AFDF7F8DDFDC6F8ED" box="[526,532,1867,1892]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +21.1″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC0A77AFD9FF8DBFD3DF8ED" box="[614,751,1867,1893]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="11" pageNumber="12" precision="1" seconds="18.3" value="-3.7550833"> +3°45 +<emphasis id="D2B9EA99FFC0A77AFD6CF8DDFD49F8ED" box="[661,667,1867,1892]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +18.3″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC0A77AFEC2F8FAFE6FF80D" box="[315,445,1900,1924]" pageId="11" pageNumber="12" value="2021-07-30">30 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFC0A77AFF74F81AFBA8FAF9" blockId="11.[113,763,1336,1987]" lastBlockId="11.[810,1460,1335,1517]" pageId="11" pageNumber="12"> +20210730-ESP-M2TP3, eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and +<quantity id="27359B6EFFC0A77AFCACFAAFFC5AFAD9" box="[853,904,1337,1361]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="11" pageNumber="12" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFC0A77AFBBBFAAFFBA9FAD8" box="[1090,1147,1337,1361]" name="Spain" pageId="11" pageNumber="12">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFC0A77AFB7EFAAFFAC4FAD9" box="[1159,1302,1335,1361]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="11" pageNumber="12" precision="1" seconds="21.1" value="43.472527"> +43°28 +<emphasis id="D2B9EA99FFC0A77AFB3EFAA1FB1FFAD9" box="[1223,1229,1335,1360]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +21.1″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC0A77AFAE7FAAEFA7AFAD9" box="[1310,1448,1335,1360]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="11" pageNumber="12" precision="1" seconds="18.3" value="-3.7550833"> +3°45 +<emphasis id="D2B9EA99FFC0A77AFAB4FAA1FA81FAD9" box="[1357,1363,1335,1360]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +18.3″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC0A77AFC0DFACEFBA7FAF9" box="[1012,1141,1368,1392]" pageId="11" pageNumber="12" value="2021-07-30">30 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFC0A77AFCBCFAE1FBA8FA64" blockId="11.[810,1460,1335,1517]" pageId="11" pageNumber="12"> +20210730-ESP-M2TP12, eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and +<quantity id="27359B6EFFC0A77AFCACFA20FC5AFA44" box="[853,904,1462,1486]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="11" pageNumber="12" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFC0A77AFBBBFA20FBA9FA47" box="[1090,1147,1462,1486]" name="Spain" pageId="11" pageNumber="12">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFC0A77AFB7EFA20FAC4FA44" box="[1159,1302,1460,1486]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="11" pageNumber="12" precision="1" seconds="21.1" value="43.472527"> +43°28 +<emphasis id="D2B9EA99FFC0A77AFB3EFA22FB1FFA44" box="[1223,1229,1460,1485]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +21.1″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC0A77AFAE7FA20FA7AFA44" box="[1310,1448,1460,1486]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="11" pageNumber="12" precision="1" seconds="18.3" value="-3.7550833"> +3°45 +<emphasis id="D2B9EA99FFC0A77AFAB4FA22FA81FA44" box="[1357,1363,1460,1485]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +18.3″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC0A77AFC0DFA43FBA7FA64" box="[1012,1141,1493,1517]" pageId="11" pageNumber="12" value="2021-07-30">30 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFC0A77AFC2EF99BFAD4F9AE" blockId="11.[983,1286,1549,1575]" box="[983,1286,1549,1575]" pageId="11" pageNumber="12"> +<emphasis id="D2B9EA99FFC0A77AFC2EF99BFAD4F9AE" box="[983,1286,1549,1575]" italics="true" pageId="11" pageNumber="12">Comparative material examined</emphasis> +</paragraph> +<paragraph id="E072368BFFC0A77AFC44F9DAFAF2F9EC" blockId="11.[810,1460,1611,1737]" box="[957,1312,1611,1638]" pageId="11" pageNumber="12"> +<taxonomicName id="27CD4D08FFC0A77AFC44F9DAFAF2F9EC" authority="(Schmidt, 1868)" baseAuthorityName="Schmidt" baseAuthorityYear="1868" box="[957,1312,1611,1638]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="11" pageNumber="12" phylum="Porifera" rank="species" species="virgultosa"> +<emphasis id="D2B9EA99FFC0A77AFC44F9DAFAF2F9EC" box="[957,1312,1611,1638]" italics="true" pageId="11" pageNumber="12">Spongia virgultosa (Schmidt, 1868)</emphasis> +</taxonomicName> +</paragraph> +<paragraph id="E072368BFFC0A77AFCD3F9E4FBDFF940" blockId="11.[810,1460,1611,1737]" pageId="11" pageNumber="12"> +20070718-MRS-ECITP5, Jarre, cave in the semi-dark part at +<quantity id="27359B6EFFC0A77AFA60F9E4FCEDF920" metricMagnitude="1" metricUnit="m" metricValue="1.5" pageId="11" pageNumber="12" unit="m" value="15.0">15 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC0A77AFC0FF904FBE9F920" box="[1014,1083,1682,1705]" name="France" pageId="11" pageNumber="12">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC0A77AFBB0F904FB17F920" box="[1097,1221,1680,1706]" degrees="43" direction="north" minutes="11" orientation="latitude" pageId="11" pageNumber="12" precision="15" seconds="46" value="43.196114"> +43°11 +<emphasis id="D2B9EA99FFC0A77AFB71F906FB5CF920" box="[1160,1166,1680,1705]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +46″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC0A77AFB36F907FAE8F920" box="[1231,1338,1680,1705]" degrees="5" direction="east" minutes="21" orientation="longitude" pageId="11" pageNumber="12" precision="15" seconds="55" value="5.365278"> +5°21 +<emphasis id="D2B9EA99FFC0A77AFB07F906FAD6F920" box="[1278,1284,1680,1705]" italics="true" pageId="11" pageNumber="12">ʹ</emphasis> +55″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC0A77AFC7FF927FBDAF940" box="[902,1032,1713,1737]" pageId="11" pageNumber="12" value="2007-07-18">18 July 2007</date> +. +</paragraph> +<paragraph id="E072368BFFC0A77AFCB9F97EFB13F8AB" blockId="11.[810,1436,1768,1862]" pageId="11" pageNumber="12"> +<taxonomicName id="27CD4D08FFC0A77AFCB9F97EFB13F8AB" authority="Voultsiadou-Koukoura, van Soest & Koukouras, 1991" authorityName="Voultsiadou-Koukoura, van Soest & Koukouras" authorityYear="1991" class="Demospongiae" family="Spongiidae" genus="Coscinoderma" kingdom="Animalia" order="Dictyoceratida" pageId="11" pageNumber="12" phylum="Porifera" rank="species" species="sporadense"> +<emphasis id="D2B9EA99FFC0A77AFCB9F97EFB13F8AB" italics="true" pageId="11" pageNumber="12">Coscinoderma sporadense Voultsiadou-Koukoura, van Soest & Koukouras, 1991</emphasis> +</taxonomicName> +</paragraph> +<paragraph id="E072368BFFC0A77AFCD3F8B8FB34F8CF" blockId="11.[810,1436,1768,1862]" box="[810,1254,1838,1862]" pageId="11" pageNumber="12"> +Po.25932, +<collectingRegion id="2209F869FFC0A77AFC6FF8B8FC3EF8CF" box="[918,1004,1838,1862]" country="Israel" name="Tel Aviv" pageId="11" pageNumber="12">Tel-Aviv</collectingRegion> +, +<collectingCountry id="98DA761BFFC0A77AFC0DF8B8FBF9F8CF" box="[1012,1067,1838,1862]" name="Israel" pageId="11" pageNumber="12">Israel</collectingCountry> +( +<bibRefCitation id="845C4B7AFFC0A77AFBC2F8B8FB07F8CF" author="Idan" box="[1083,1237,1838,1862]" etAl="et al." firstAuthor="Idan" pageId="11" pageNumber="12" pagination="84 - 106" refId="ref17579" refString="Idan T, Shefer S, Feldstein T et al. Shedding light on an East-Mediterranean mesophotic sponge ground community and the regional sponge fauna. Mediterranean Marine Science 2018; 19: 84 - 106. https: // doi. org / 10.12681 / mms. 13853" type="journal article" year="2018"> +Idan +<emphasis id="D2B9EA99FFC0A77AFB97F8B9FB4FF8CF" box="[1134,1181,1838,1862]" italics="true" pageId="11" pageNumber="12">et al.</emphasis> +2018 +</bibRefCitation> +). +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC0A77DFBC1F8F0FDB8FC5F" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12" type="description"> +<paragraph id="E072368BFFC0A77AFBC1F8F0FB76F8F6" blockId="11.[810,1459,1894,1988]" box="[1080,1188,1894,1919]" pageId="11" pageNumber="12"> +<emphasis id="D2B9EA99FFC0A77AFBC1F8F0FB76F8F6" box="[1080,1188,1894,1919]" italics="true" pageId="11" pageNumber="12">Description</emphasis> +</paragraph> +<paragraph id="E072368BFFC0A77DFCD3F81AFEBDFDB6" blockId="11.[810,1459,1894,1988]" lastBlockId="12.[128,778,144,983]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12"> +This sponge can be massive and irregular, with lobes or protuberances, fan- or vase-shaped when it becomes very big ( +<figureCitation id="78F62A0EFFC0A77AFAC2F83DFA70F84D" box="[1339,1442,1963,1988]" captionStart="Figure 4" captionStartId="11.[113,178,1236,1260]" captionTargetBox="[117,1456,146,1201]" captionTargetId="figure-302@11.[114,1458,144,1208]" captionTargetPageId="11" captionText="Figure 4. In situ pictures of Spongia lamella in the Northeast Atlantic Ocean and the Mediterranean Sea, A, Baloo tunnel, Ciboure, France.B, C, Mouro Island, Santander, Spain.D, Marseille, France." pageId="11" pageNumber="12">Fig. 4A–D</figureCitation> +). The largest studied specimens measured +<quantity id="27359B6EFFC7A77DFDD3FF06FDBEFF2E" box="[554,620,144,168]" metricMagnitude="-1" metricUnit="m" metricValue="5.0" pageId="12" pageNumber="13" unit="cm" value="50.0">50 cm</quantity> +in their largest diameter, but this dimension can be> +<quantity id="27359B6EFFC7A77DFDFEFF39FDFEFF4E" box="[519,556,175,199]" metricMagnitude="0" metricUnit="m" metricValue="1.0" pageId="12" pageNumber="13" unit="m" value="1.0">1 m</quantity> +in the Mediterranean Sea ( +<figureCitation id="78F62A0EFFC7A77DFF4EFF59FED7FF6E" box="[183,261,207,231]" captionStart="Figure 4" captionStartId="11.[113,178,1236,1260]" captionTargetBox="[117,1456,146,1201]" captionTargetId="figure-302@11.[114,1458,144,1208]" captionTargetPageId="11" captionText="Figure 4. In situ pictures of Spongia lamella in the Northeast Atlantic Ocean and the Mediterranean Sea, A, Baloo tunnel, Ciboure, France.B, C, Mouro Island, Santander, Spain.D, Marseille, France." pageId="12" pageNumber="13">Fig. 4D</figureCitation> +). No true vase- or fan-shaped specimen was observed among our Atlantic specimens. This sponge presents a light to dark grey external colour, and the internal tissue is usually light or tawny yellow. The colour does not change upon fixation in 95% ethanol. The sponge consistency is flexible and hardly tearable. The surface is finely covered with small, evenly distributed conules. Oscules can be grouped in large individuals, or more randomly distributed in smaller ones, always on the outer sides in large individuals or on the top of each lobe or protuberance in smaller ones. They measure +<quantity id="27359B6EFFC7A77DFDABFE7FFD79FD89" box="[594,683,489,513]" metricMagnitude="-3" metricUnit="m" metricValue="2.0" metricValueMax="3.0" metricValueMin="1.0" pageId="12" pageNumber="13" unit="mm" value="2.0" valueMax="3.0" valueMin="1.0">1–3 mm</quantity> +in diameter and are slightly raised, with a small whitish membrane delimiting the opening. +</paragraph> +<paragraph id="E072368BFFC7A77DFF65FDD1FDB8FC5F" blockId="12.[128,778,144,983]" pageId="12" pageNumber="13"> +The ectosomal skeleton harbours an epidermal skin, 150– 600 µm thick, easily detachable, with a star-like appearance. It is made of a network of abundant foreign spicules and debris ( +<figureCitation id="78F62A0EFFC7A77DFF72FD32FF25FD35" box="[139,247,676,700]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." pageId="12" pageNumber="13">Fig. 5A–C</figureCitation> +). Small canals can be observed under the surface. In the choanosome, debris and foreign spicules are still abundant at the periphery of canals, and thus throughout the sponge body ( +<figureCitation id="78F62A0EFFC7A77DFF3CFC94FEC3FC93" box="[197,273,770,794]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." pageId="12" pageNumber="13">Fig. 5D</figureCitation> +). The primary fibres, 30–120 µm in diameter, are common, simple, irregular, and cored with numerous inclusions of foreign spicules and debris ( +<figureCitation id="78F62A0EFFC7A77DFE36FCD7FDE9FCD0" box="[463,571,833,857]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." pageId="12" pageNumber="13">Fig. 5A, B</figureCitation> +). Near the surface, they can be ramified. Secondary fibres, 20–40 µm in diameter, without inclusions or pith, form a dense ramified and irregular network ( +<figureCitation id="78F62A0EFFC7A77DFF1CFC09FE96FC3E" box="[229,324,927,951]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." pageId="12" pageNumber="13">Fig. 5B, E</figureCitation> +). Tertiary fibres are present or absent. When they exist, their thickness is ≤10 µm in diameter. +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC7A77DFE72FC60FEA8FBFA" pageId="12" pageNumber="13" type="distribution"> +<paragraph id="E072368BFFC7A77DFE72FC60FE2CFB99" blockId="12.[129,778,1014,1139]" box="[395,510,1014,1040]" pageId="12" pageNumber="13"> +<emphasis id="D2B9EA99FFC7A77DFE72FC60FE2CFB99" box="[395,510,1014,1040]" italics="true" pageId="12" pageNumber="13">Distribution</emphasis> +</paragraph> +<paragraph id="E072368BFFC7A77DFF78FB8BFEA8FBFA" blockId="12.[129,778,1014,1139]" pageId="12" pageNumber="13"> +Mediterranean Sea and South European Atlantic Shelf (e.g. +<bibRefCitation id="845C4B7AFFC7A77DFF78FBAAFE53FBDD" author="Noyer and Becerro" box="[129,385,1084,1108]" firstAuthor="Noyer" pageId="12" pageNumber="13" pagination="85 - 99" refId="ref18110" refString="Noyer C, Becerro MA. Relationship between genetic, chemical, and bacterial diversity in the Atlanto-Mediterranean bath sponge Spongia lamella. In: Maldonado M, Turon X, Becerro M et al. (eds.), Ancient Animals, New Challenges. Developments in Hodrobiologo, Vol. 219. Dordrecht: Springer, 2012, 85 - 99." type="book chapter" year="2012">Noyer and Becerro 2012</bibRefCitation> +). This study expands the distribution of +<taxonomicName id="27CD4D08FFC7A77DFF63FBCAFF2BFBFA" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[154,249,1115,1139]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC7A77DFF63FBCAFF2BFBFA" box="[154,249,1115,1139]" italics="true" pageId="12" pageNumber="13">S. lamella</emphasis> +</taxonomicName> +northwards. +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC7A77DFE59FB05FDF5F901" pageId="12" pageNumber="13" type="biology_ecology"> +<paragraph id="E072368BFFC7A77DFE59FB05FE38FB24" blockId="12.[128,778,1171,1672]" box="[416,490,1171,1197]" pageId="12" pageNumber="13"> +<emphasis id="D2B9EA99FFC7A77DFE59FB05FE38FB24" box="[416,490,1171,1197]" italics="true" pageId="12" pageNumber="13">Ecology</emphasis> +</paragraph> +<paragraph id="E072368BFFC7A77DFF78FB2CFDF5F901" blockId="12.[128,778,1171,1672]" pageId="12" pageNumber="13"> +In the Mediterranean Sea, +<taxonomicName id="27CD4D08FFC7A77DFE76FB2CFE3FFB58" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[399,493,1209,1233]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC7A77DFE76FB2CFE3FFB58" box="[399,493,1209,1233]" italics="true" pageId="12" pageNumber="13">S. lamella</emphasis> +</taxonomicName> +is well known to be distributed mainly along coralligenous cliffs or on deep mesophotic bottoms ( +<bibRefCitation id="845C4B7AFFC7A77DFF13FB6EFEA3FA99" author="Vacelet" box="[234,369,1272,1296]" firstAuthor="Vacelet" pageId="12" pageNumber="13" pagination="39 - 101" refId="ref18958" refString="Vacelet J. Repartition generale des eponges et systematique des eponges cornees de la region de Marseille et de quelques stations Mediterraneennes. Recueil des travaux de la Station Marine d'Endoume 1959; 16: 39 - 101." type="journal article" year="1959">Vacelet 1959</bibRefCitation> +, +<bibRefCitation id="845C4B7AFFC7A77DFE79FB6EFD7DFA99" author="Pronzato and Manconi" box="[384,687,1272,1296]" firstAuthor="Pronzato" pageId="12" pageNumber="13" pagination="146 - 66" refId="ref18376" refString="Pronzato R, Manconi R. Mediterranean commercial sponges: over 5000 years of natural history and cultural heritage. Marine Ecologo 2008; 29: 146 - 66. https: // doi. org / 10.1111 / j. 1439 - 0485.2008.00235. x" type="journal article" year="2008">Pronzato and Manconi 2008</bibRefCitation> +). In the Northeast Atlantic, +<taxonomicName id="27CD4D08FFC7A77DFEA9FA8EFE60FAA6" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[336,434,1303,1327]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC7A77DFEA9FA8EFE60FAA6" box="[336,434,1303,1327]" italics="true" pageId="12" pageNumber="13">S. lamella</emphasis> +</taxonomicName> +has mostly been found in semi-dark and dark cavities, sometimes in crevices found along shallow water cliffs or under rocks. All specimens reported here were found in habitats with very high wave energy, which might explain why no large or fan-shaped individuals were observed. This sponge has a patchy distribution, and it lives in syntopy with +<taxonomicName id="27CD4D08FFC7A77DFF40FA42FEE9FA62" box="[185,315,1492,1515]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC7A77DFF40FA42FEE9FA62" box="[185,315,1492,1515]" italics="true" pageId="12" pageNumber="13">S. maitasuna</emphasis> +</taxonomicName> +. In caves, this species is commonly associated with numerous other sponges, solitary scleractinians, such as +<taxonomicName id="27CD4D08FFC7A77DFF64F985FDBEF9A3" authority="Lacaze-Duthiers, 1897" authorityName="Lacaze-Duthiers" authorityYear="1897" box="[157,620,1554,1578]" class="Anthozoa" family="Dendrophylliidae" genus="Leptopsammia" kingdom="Animalia" order="Scleractinia" pageId="12" pageNumber="13" phylum="Cnidaria" rank="species" species="pruvoti"> +<emphasis id="D2B9EA99FFC7A77DFF64F985FEA9F9A3" box="[157,379,1555,1578]" italics="true" pageId="12" pageNumber="13">Leptopsammia pruvoti</emphasis> +Lacaze-Duthiers, 1897 +</taxonomicName> +<emphasis id="D2B9EA99FFC7A77DFD95F984FDA4F9A0" box="[620,630,1554,1577]" italics="true" pageId="12" pageNumber="13">?</emphasis> +, or zoanthids, such as +<taxonomicName id="27CD4D08FFC7A77DFF25F9A7FD56F9C0" authority="(Schmidt, 1862)" baseAuthorityName="Schmidt" baseAuthorityYear="1862" box="[220,644,1585,1609]" class="Anthozoa" family="Parazoanthidae" genus="Parazoanthus" kingdom="Animalia" order="Zoantharia" pageId="12" pageNumber="13" phylum="Cnidaria" rank="species" species="axinellae"> +<emphasis id="D2B9EA99FFC7A77DFF25F9A7FE17F9C0" box="[220,453,1585,1609]" italics="true" pageId="12" pageNumber="13">Parazoanthus axinellae</emphasis> +( +<bibRefCitation id="845C4B7AFFC7A77DFE21F9A7FDAEF9C0" author="Schmidt" box="[472,636,1585,1609]" firstAuthor="Schmidt" pageId="12" pageNumber="13" refId="ref18646" refString="Schmidt O. Die Spongien des adriatischen Meeres. Leipzig: Wilhelm Engelmann, 1862." type="book" year="1862">Schmidt, 1862</bibRefCitation> +) +</taxonomicName> +. No sign of epibiosis or predation was observed. No reproductive element was recorded in the processed specimens. +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC7A77FFE9EF93EFB45F848" lastPageId="14" lastPageNumber="15" pageId="12" pageNumber="13" type="discussion"> +<paragraph id="E072368BFFC7A77DFE9EF93EFDF1F948" blockId="12.[129,777,1703,1985]" box="[359,547,1703,1729]" pageId="12" pageNumber="13"> +<emphasis id="D2B9EA99FFC7A77DFE9EF93EFDF1F948" box="[359,547,1703,1729]" italics="true" pageId="12" pageNumber="13">Taxonomic remarks</emphasis> +</paragraph> +<paragraph id="E072368BFFC7A77DFF78F958FAEAFD52" blockId="12.[129,777,1703,1985]" lastBlockId="12.[825,1475,144,920]" pageId="12" pageNumber="13"> +The +<specimenCount id="F6CBFD02FFC7A77DFF56F958FEE9F96F" box="[175,315,1742,1766]" count="15" pageId="12" pageNumber="13" type="generic">15 specimens</specimenCount> +examined from the Northeast Atlantic differ slightly in their growth form from +<taxonomicName id="27CD4D08FFC7A77DFE2DF978FDE3F88C" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[468,561,1773,1797]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC7A77DFE2DF978FDE3F88C" box="[468,561,1773,1797]" italics="true" pageId="12" pageNumber="13">S. lamella</emphasis> +</taxonomicName> +in the Mediterranean Sea, but their skeletons are identical. Openings of the oscula are also twice as large in the Atlantic specimens. They all present in their ectosome a rather thick layer of foreign spicules and debris, which is an unusual structure among +<taxonomicName id="27CD4D08FFC7A77DFDF1F8FDFD84F80B" authorityName="Linnaeus" authorityYear="1759" box="[520,598,1899,1922]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC7A77DFDF1F8FDFD84F80B" box="[520,598,1899,1922]" italics="true" pageId="12" pageNumber="13">Spongia</emphasis> +</taxonomicName> +that was not observed in +<taxonomicName id="27CD4D08FFC7A77DFF1FF81CFEB4F82B" box="[230,358,1930,1954]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC7A77DFF1FF81CFEB4F82B" box="[230,358,1930,1954]" italics="true" pageId="12" pageNumber="13">S. maitasuna</emphasis> +</taxonomicName> +. Moreover, the overall network of fibres cannot be confounded with the specific honeycomb-shaped skeleton of the new species. A rather organized ectosomal skeleton made of foreign spicules and debris is known only in +<taxonomicName id="27CD4D08FFC7A77DFCC0FF59FC16FF6E" authorityName="Carter" authorityYear="1883" box="[825,964,207,231]" class="Demospongiae" family="Spongiidae" genus="Coscinoderma" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC7A77DFCC0FF59FC16FF6E" box="[825,964,207,231]" italics="true" pageId="12" pageNumber="13">Coscinoderma</emphasis> +</taxonomicName> +and +<taxonomicName id="27CD4D08FFC7A77DFC06FF59FB81FF6E" authorityName="Lendenfeld" authorityYear="1888" box="[1023,1107,207,231]" class="Demospongiae" family="Spongiidae" genus="Leiosella" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC7A77DFC06FF59FB81FF6E" box="[1023,1107,207,231]" italics="true" pageId="12" pageNumber="13">Leiosella</emphasis> +</taxonomicName> +, but in these cases, this structure is much thicker and the organization of the choanosome skeleton very different (e.g. intertwined secondary fibres in the Mediterranean +<taxonomicName id="27CD4D08FFC7A77DFC2EFEBBFB8CFECD" authorityName="Voultsiadou-Koukoura, van Soest & Koukouras" authorityYear="1991" box="[983,1118,300,324]" class="Demospongiae" family="Spongiidae" genus="Coscinoderma" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="sporadense"> +<emphasis id="D2B9EA99FFC7A77DFC2EFEBBFB8CFECD" box="[983,1118,300,324]" italics="true" pageId="12" pageNumber="13">C. sporadense</emphasis> +</taxonomicName> +). Atlantic and Mediterranean specimens of +<taxonomicName id="27CD4D08FFC7A77DFC66FEDAFC2EFEED" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[927,1020,332,356]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC7A77DFC66FEDAFC2EFEED" box="[927,1020,332,356]" italics="true" pageId="12" pageNumber="13">S. lamella</emphasis> +</taxonomicName> +share the same common simple and irregular primary fibres, with inclusions of foreign spicules and debris, easily observable from the inner part of the sponge to the surface. They remain simple until the formation of conules, whereas in other +<taxonomicName id="27CD4D08FFC7A77DFC68FE5CFBFCFE68" box="[913,1070,457,481]" pageId="12" pageNumber="13"> +<emphasis id="D2B9EA99FFC7A77DFC68FE5CFC0DFE68" box="[913,991,458,481]" italics="true" pageId="12" pageNumber="13">Spongia</emphasis> +species +</taxonomicName> +they become anastomosed at this level. The size of the primary fibres appears a little more variable in the Atlantic specimens (30–120 µm in diameter) than in the Mediterranean specimens (50–100 µm in diameter). Foreign spicules and debris seem to be more abundant in Atlantic specimens. In comparison, the primary fibres of +<taxonomicName id="27CD4D08FFC7A77DFAFBFDF0FAAAFDF7" baseAuthorityName="Schmidt" baseAuthorityYear="1868" box="[1282,1400,614,638]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="virgultosa"> +<emphasis id="D2B9EA99FFC7A77DFAFBFDF0FAAAFDF7" box="[1282,1400,614,638]" italics="true" pageId="12" pageNumber="13">S. virgultosa</emphasis> +</taxonomicName> +, which has an unusual encrusting papillate growth form for +<taxonomicName id="27CD4D08FFC7A77DFA97FD10FA6EFD14" authorityName="Linnaeus" authorityYear="1759" box="[1390,1468,646,669]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC7A77DFA97FD10FA6EFD14" box="[1390,1468,646,669]" italics="true" pageId="12" pageNumber="13">Spongia</emphasis> +</taxonomicName> +, are also full of mineral debris, but they are rare and significantly thinner (40–50 µm in diameter) than in +<taxonomicName id="27CD4D08FFC7A77DFB2AFD52FAE3FD55" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[1235,1329,708,732]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC7A77DFB2AFD52FAE3FD55" box="[1235,1329,708,732]" italics="true" pageId="12" pageNumber="13">S. lamella</emphasis> +</taxonomicName> +. +</paragraph> +<paragraph id="E072368BFFC7A77DFCACFD75FA9BFC1E" blockId="12.[825,1475,144,920]" pageId="12" pageNumber="13">Therefore, the slight morphological differences recorded between Atlantic and Mediterranean specimens (growth form, abundance of debris, and variation in the size of primary fibres) do not seem to us to be sufficient to consider the Atlantic specimens as a different species and are thus interpreted as variations attributable to the different environmental contexts.</paragraph> +<paragraph id="E072368BFFC7A77DFC2FFC21FAF7FC58" blockId="12.[825,1475,951,1985]" box="[982,1317,951,977]" pageId="12" pageNumber="13"> +<emphasis id="D2B9EA99FFC7A77DFC2FFC21FAF7FC58" bold="true" box="[982,1317,951,977]" pageId="12" pageNumber="13">DNA analysis of studied species</emphasis> +</paragraph> +<paragraph id="E072368BFFC7A77DFCC0FC48FB51FB78" blockId="12.[825,1475,951,1985]" pageId="12" pageNumber="13"> +GenBank accession numbers for all sequences generated or downloaded in this study are presented in +<tableCitation id="AD4F0330FFC7A77DFB09FC6BFAEEFB9C" box="[1264,1340,1021,1045]" captionStart="Table 1" captionStartId="3.[113,168,143,167]" captionTargetPageId="3" captionText="Table 1. Species names, voucher codes and GenBank accession numbers of Spongiidae used in this study." pageId="12" pageNumber="13">Table 1</tableCitation> +. For the 28S (C2–D2 fragment) and +<emphasis id="D2B9EA99FFC7A77DFBCBFB8BFBB1FBBC" box="[1074,1123,1053,1077]" italics="true" pageId="12" pageNumber="13">CO1</emphasis> +(dgLCO1490–dgHCO2198 fragment) molecular markers, we obtained, respectively, 18 and 3 sequences of +<taxonomicName id="27CD4D08FFC7A77DFC31FBCAFB9AFBFA" box="[968,1096,1116,1139]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC7A77DFC31FBCAFB9AFBFA" box="[968,1096,1116,1139]" italics="true" pageId="12" pageNumber="13">S. maitasuna</emphasis> +</taxonomicName> +, 21 and 14 sequences of +<taxonomicName id="27CD4D08FFC7A77DFAA3FBCAFA6EFBFA" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[1370,1468,1115,1139]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC7A77DFAA3FBCAFA6EFBFA" box="[1370,1468,1115,1139]" italics="true" pageId="12" pageNumber="13">S. lamella</emphasis> +</taxonomicName> +, 14 and 5 sequences for +<taxonomicName id="27CD4D08FFC7A77DFBC4FBEDFB61FB1A" authorityName="Linnaeus" authorityYear="1759" box="[1085,1203,1147,1171]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC7A77DFBC4FBEDFB61FB1A" box="[1085,1203,1147,1171]" italics="true" pageId="12" pageNumber="13">S. officinalis</emphasis> +</taxonomicName> +, 8 and 2 sequences of +<taxonomicName id="27CD4D08FFC7A77DFA49FBEDFC72FB3B" authorityName="Schmidt" authorityYear="1862" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="mollissima"> +<emphasis id="D2B9EA99FFC7A77DFA49FBEDFC72FB3B" italics="true" pageId="12" pageNumber="13">S. mollissima</emphasis> +</taxonomicName> +, 4 and 3 sequences of +<taxonomicName id="27CD4D08FFC7A77DFB7CFB0DFAD4FB3B" baseAuthorityName="Lamarck" baseAuthorityYear="1814" box="[1157,1286,1179,1202]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="communis"> +<emphasis id="D2B9EA99FFC7A77DFB7CFB0DFAD4FB3B" box="[1157,1286,1179,1202]" italics="true" pageId="12" pageNumber="13">H. communis</emphasis> +</taxonomicName> +, 1 and 1 sequence of +<taxonomicName id="27CD4D08FFC7A77DFCACFB2CFC1CFB58" baseAuthorityName="Schmidt" baseAuthorityYear="1868" box="[853,974,1209,1233]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="virgultosa"> +<emphasis id="D2B9EA99FFC7A77DFCACFB2CFC1CFB58" box="[853,974,1209,1233]" italics="true" pageId="12" pageNumber="13">S. virgultosa</emphasis> +</taxonomicName> +, and only 1 sequence of +<taxonomicName id="27CD4D08FFC7A77DFB2DFB2CFAEEFB58" authorityName="Schmidt" authorityYear="1862" box="[1236,1340,1210,1233]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC7A77DFB2DFB2CFAEEFB58" box="[1236,1340,1210,1233]" italics="true" pageId="12" pageNumber="13">S. zimocca</emphasis> +</taxonomicName> +(28S) and 1 sequence of +<taxonomicName id="27CD4D08FFC7A77DFC4EFB4FFBEEFB78" authorityName="Voultsiadou-Koukoura, van Soest & Koukouras" authorityYear="1991" box="[951,1084,1241,1265]" class="Demospongiae" family="Spongiidae" genus="Coscinoderma" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="sporadense"> +<emphasis id="D2B9EA99FFC7A77DFC4EFB4FFBEEFB78" box="[951,1084,1241,1265]" italics="true" pageId="12" pageNumber="13">C. sporadense</emphasis> +</taxonomicName> +(28S). +</paragraph> +<paragraph id="E072368BFFC7A77FFCACFB6EFE48FE68" blockId="12.[825,1475,951,1985]" lastBlockId="14.[128,778,144,1014]" lastPageId="14" lastPageNumber="15" pageId="12" pageNumber="13"> +Molecular markers allowed us to build highly congruent trees, although the phylogenetic reconstruction with 28S gave a better resolution ( +<figureCitation id="78F62A0EFFC7A77DFC48FAA1FC3BFAC7" box="[945,1001,1334,1359]" captionStart="Figure 6" captionStartId="15.[113,178,1836,1860]" captionTargetBox="[173,1400,146,1806]" captionTargetId="figure-7@15.[171,1402,144,1808]" captionTargetPageId="15" captionText="Figure 6. Phylogenetic reconstruction of Northeast Atlantic and Mediterranean Spongiidae using sequences of the C2–D2 region of 28S, and both the maximum-likelihood (ML) and neigbour-joining (NJ) methods. Bootstrap values are given for both analyses (1000 replicates). The NJ values are in parentheses. Each voucher code is associated with a species name. The ML bootstrap values are given only if>500." pageId="12" pageNumber="13">Fig. 6</figureCitation> +) than +<emphasis id="D2B9EA99FFC7A77DFBD5FAA1FB8FFAC6" box="[1068,1117,1335,1359]" italics="true" pageId="12" pageNumber="13">CO1</emphasis> +(Supporting Information, +<figureCitation id="78F62A0EFFC7A77DFA95FAA1FA63FAC6" box="[1388,1457,1335,1359]" captionStart="Figure 1" captionStartId="2.[129,194,1954,1978]" captionTargetBox="[131,1471,1210,1924]" captionTargetId="figure-610@2.[129,1473,1208,1926]" captionTargetPageId="2" captionText="Figure 1. Study area.Black stars indicate the sampling locations of Spongiidae from this work." pageId="12" pageNumber="13">Fig. S1</figureCitation> +). Both methods (NJ and ML) of phylogenetic reconstruction using 28S produced the same topology: 70 +<taxonomicName id="27CD4D08FFC7A77DFAF2FAE3FAAFFA04" authorityName="Gray" authorityYear="1867" box="[1291,1405,1397,1421]" class="Demospongiae" family="Spongiidae" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="family">Spongiidae</taxonomicName> +specimens divided in two, with only one sequence of +<taxonomicName id="27CD4D08FFC7A77DFAC2FA03FA10FA25" authorityName="Voultsiadou-Koukoura, van Soest & Koukouras" authorityYear="1991" box="[1339,1474,1428,1452]" class="Demospongiae" family="Spongiidae" genus="Coscinoderma" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="sporadense"> +<emphasis id="D2B9EA99FFC7A77DFAC2FA03FA10FA25" box="[1339,1474,1428,1452]" italics="true" pageId="12" pageNumber="13">C. sporadense</emphasis> +</taxonomicName> +on one side and +<specimenCount id="F6CBFD02FFC7A77DFC1EFA22FBA1FA45" box="[999,1139,1460,1484]" count="69" pageId="12" pageNumber="13" type="generic">69 specimens</specimenCount> +of +<taxonomicName id="27CD4D08FFC7A77DFB6FFA22FB36FA42" authorityName="Linnaeus" authorityYear="1759" box="[1174,1252,1460,1483]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC7A77DFB6FFA22FB36FA42" box="[1174,1252,1460,1483]" italics="true" pageId="12" pageNumber="13">Spongia</emphasis> +</taxonomicName> +and +<taxonomicName id="27CD4D08FFC7A77DFAE1FA23FA4DFA45" authorityName="Schulze" authorityYear="1879" box="[1304,1439,1461,1484]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC7A77DFAE1FA23FA4DFA45" box="[1304,1439,1461,1484]" italics="true" pageId="12" pageNumber="13">Hippospongia</emphasis> +</taxonomicName> +on the other side ( +<figureCitation id="78F62A0EFFC7A77DFC2CFA45FBDFFA62" box="[981,1037,1491,1515]" captionStart="Figure 6" captionStartId="15.[113,178,1836,1860]" captionTargetBox="[173,1400,146,1806]" captionTargetId="figure-7@15.[171,1402,144,1808]" captionTargetPageId="15" captionText="Figure 6. Phylogenetic reconstruction of Northeast Atlantic and Mediterranean Spongiidae using sequences of the C2–D2 region of 28S, and both the maximum-likelihood (ML) and neigbour-joining (NJ) methods. Bootstrap values are given for both analyses (1000 replicates). The NJ values are in parentheses. Each voucher code is associated with a species name. The ML bootstrap values are given only if>500." pageId="12" pageNumber="13">Fig. 6</figureCitation> +). This set of specimens can be divided into three different groups. The first group is well supported by both methods of phylogenetic constructions (NJ and ML). It is composed by the proposed new species +<taxonomicName id="27CD4D08FFC7A77DFB46F9A4FA93F9C0" box="[1215,1345,1586,1609]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC7A77DFB46F9A4FA93F9C0" box="[1215,1345,1586,1609]" italics="true" pageId="12" pageNumber="13">S. maitasuna</emphasis> +</taxonomicName> +, individuals from the two sampled localities presenting the same sequence of 28S, and by a well-supported group made of one sequence of +<taxonomicName id="27CD4D08FFC7A77DFCADF919FC6EF92E" authorityName="Schmidt" authorityYear="1862" box="[852,956,1679,1703]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC7A77DFCADF919FC6EF92E" box="[852,956,1679,1703]" italics="true" pageId="12" pageNumber="13">S. zimocca</emphasis> +</taxonomicName> +, two downloaded sequences of +<taxonomicName id="27CD4D08FFC7A77DFAFDF919FA87F92E" baseAuthorityName="Schmidt" baseAuthorityYear="1862" box="[1284,1365,1679,1703]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="nitens"> +<emphasis id="D2B9EA99FFC7A77DFAFDF919FA87F92E" box="[1284,1365,1679,1703]" italics="true" pageId="12" pageNumber="13">S. nitens</emphasis> +</taxonomicName> +, and three sequences obtained in the present study of +<taxonomicName id="27CD4D08FFC7A77DFB01F939FA5BF94F" authorityName="Linnaeus" authorityYear="1759" box="[1272,1417,1710,1734]" class="Demospongiae" family="Spongiidae" genus="Spongia" isUncertain="true" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC7A77DFB01F939FAD8F94F" box="[1272,1290,1711,1734]" italics="true" pageId="12" pageNumber="13">S.</emphasis> +cf. +<emphasis id="D2B9EA99FFC7A77DFAD7F938FA5BF94F" box="[1326,1417,1710,1734]" italics="true" pageId="12" pageNumber="13">officinalis</emphasis> +</taxonomicName> +from +<collectingCountry id="98DA761BFFC7A77DFCC0F958FCABF96F" box="[825,889,1742,1766]" name="Spain" pageId="12" pageNumber="13">Ceuta</collectingCountry> +. +<taxonomicName id="27CD4D08FFC7A77DFC7EF958FB97F96C" box="[903,1093,1742,1765]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC7A77DFC7EF958FB97F96C" box="[903,1093,1742,1765]" italics="true" pageId="12" pageNumber="13">Spongia maitasuna</emphasis> +</taxonomicName> +thus appears closely related to this group, with the p-distance being very low between the proposed new species and +<taxonomicName id="27CD4D08FFC7A77DFC1BF89BFB9BF8AD" authorityName="Schmidt" authorityYear="1862" box="[994,1097,1805,1828]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC7A77DFC1BF89BFB9BF8AD" box="[994,1097,1805,1828]" italics="true" pageId="12" pageNumber="13">S. zimocca</emphasis> +</taxonomicName> +(1.05%, four substitutions), +<taxonomicName id="27CD4D08FFC7A77DFA94F89BFA6FF8AD" baseAuthorityName="Schmidt" baseAuthorityYear="1862" box="[1389,1469,1805,1828]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="nitens"> +<emphasis id="D2B9EA99FFC7A77DFA94F89BFA6FF8AD" box="[1389,1469,1805,1828]" italics="true" pageId="12" pageNumber="13">S. nitens</emphasis> +</taxonomicName> +, and +<taxonomicName id="27CD4D08FFC7A77DFC9DF8BAFC21F8CD" authorityName="Linnaeus" authorityYear="1759" box="[868,1011,1836,1860]" class="Demospongiae" family="Spongiidae" genus="Spongia" isUncertain="true" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC7A77DFC9DF8BAFCA5F8CA" box="[868,887,1836,1859]" italics="true" pageId="12" pageNumber="13">S.</emphasis> +cf. +<emphasis id="D2B9EA99FFC7A77DFC61F8BAFC21F8CD" box="[920,1011,1836,1860]" italics="true" pageId="12" pageNumber="13">officinalis</emphasis> +</taxonomicName> +(0.79%, the substitutions). The second group is weakly supported, whatever the method of construction. It includes +<taxonomicName id="27CD4D08FFC7A77DFC7BF8FDFC28F80B" baseAuthorityName="Schmidt" baseAuthorityYear="1868" box="[898,1018,1898,1922]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="virgultosa"> +<emphasis id="D2B9EA99FFC7A77DFC7BF8FDFC28F80B" box="[898,1018,1898,1922]" italics="true" pageId="12" pageNumber="13">S. virgultosa</emphasis> +</taxonomicName> +and a well-supported group of all +<taxonomicName id="27CD4D08FFC7A77DFA98F8FDFA10F80B" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[1377,1474,1898,1922]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC7A77DFA98F8FDFA10F80B" box="[1377,1474,1898,1922]" italics="true" pageId="12" pageNumber="13">S. lamella</emphasis> +</taxonomicName> +from the Mediterranean and Northeast Atlantic. With only one exception,thesequencedindividualsaregroupedbygeographical origin. The Mediterranean sequence (20200602-MRS-MTP8) mixed with the Atlantic presents a p-distance of 0.26% (one substitution) from the rest of the Mediterranean +<taxonomicName id="27CD4D08FFC5A77FFD7DFF59FD37FF6E" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[644,741,207,231]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC5A77FFD7DFF59FD37FF6E" box="[644,741,207,231]" italics="true" pageId="14" pageNumber="15">S. lamella</emphasis> +</taxonomicName> +. In comparison, +<taxonomicName id="27CD4D08FFC5A77FFEFDFF78FEA8FE8F" baseAuthorityName="Schmidt" baseAuthorityYear="1868" box="[260,378,238,262]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="virgultosa"> +<emphasis id="D2B9EA99FFC5A77FFEFDFF78FEA8FE8F" box="[260,378,238,262]" italics="true" pageId="14" pageNumber="15">S. virgultosa</emphasis> +</taxonomicName> +presents a p-distance of 13.4% (51 substitutions) from all +<taxonomicName id="27CD4D08FFC5A77FFEB5FE98FE7FFEAC" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[332,429,269,293]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC5A77FFEB5FE98FE7FFEAC" box="[332,429,269,293]" italics="true" pageId="14" pageNumber="15">S. lamella</emphasis> +</taxonomicName> +. The third group is composed by a weakly supported set of sponges, including all +<taxonomicName id="27CD4D08FFC5A77FFD7BFEBBFCD6FECC" authorityName="Schmidt" authorityYear="1862" box="[642,772,301,325]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="mollissima"> +<emphasis id="D2B9EA99FFC5A77FFD7BFEBBFCD6FECC" box="[642,772,301,325]" italics="true" pageId="14" pageNumber="15">S. mollissima</emphasis> +</taxonomicName> +, +<taxonomicName id="27CD4D08FFC5A77FFF78FEDAFF25FEED" authorityName="Linnaeus" authorityYear="1759" box="[129,247,332,356]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC5A77FFF78FEDAFF25FEED" box="[129,247,332,356]" italics="true" pageId="14" pageNumber="15">S. officinalis</emphasis> +</taxonomicName> +, and +<taxonomicName id="27CD4D08FFC5A77FFECCFEDBFE6BFEED" baseAuthorityName="Lamarck" baseAuthorityYear="1814" box="[309,441,333,356]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="communis"> +<emphasis id="D2B9EA99FFC5A77FFECCFEDBFE6BFEED" box="[309,441,333,356]" italics="true" pageId="14" pageNumber="15">H. communis</emphasis> +</taxonomicName> +. The genetic distances between these three species are low, with a minimal p-distance of 0.26% between +<taxonomicName id="27CD4D08FFC5A77FFF24FE1DFE8EFE2A" authorityName="Schmidt" authorityYear="1862" box="[221,348,395,419]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="mollissima"> +<emphasis id="D2B9EA99FFC5A77FFF24FE1DFE8EFE2A" box="[221,348,395,419]" italics="true" pageId="14" pageNumber="15">S. mollissima</emphasis> +</taxonomicName> +and +<taxonomicName id="27CD4D08FFC5A77FFE74FE1DFDDCFE2B" baseAuthorityName="Lamarck" baseAuthorityYear="1814" box="[397,526,395,418]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="communis"> +<emphasis id="D2B9EA99FFC5A77FFE74FE1DFDDCFE2B" box="[397,526,395,418]" italics="true" pageId="14" pageNumber="15">H. communis</emphasis> +</taxonomicName> +from +<collectingCountry id="98DA761BFFC5A77FFDB5FE1DFD48FE2A" box="[588,666,395,419]" name="Tunisia" pageId="14" pageNumber="15">Tunisia</collectingCountry> +, and 0.79– 1.57% (three to six substitutions) between +<taxonomicName id="27CD4D08FFC5A77FFDACFE3CFD0AFE4B" authorityName="Schmidt" authorityYear="1862" box="[597,728,426,450]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="mollissima"> +<emphasis id="D2B9EA99FFC5A77FFDACFE3CFD0AFE4B" box="[597,728,426,450]" italics="true" pageId="14" pageNumber="15">S. mollissima</emphasis> +</taxonomicName> +and +<taxonomicName id="27CD4D08FFC5A77FFF78FE5CFF21FE68" authorityName="Linnaeus" authorityYear="1759" box="[129,243,457,481]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC5A77FFF78FE5CFF21FE68" box="[129,243,457,481]" italics="true" pageId="14" pageNumber="15">S. officinalis</emphasis> +</taxonomicName> +or +<taxonomicName id="27CD4D08FFC5A77FFEEAFE5CFE46FE68" baseAuthorityName="Lamarck" baseAuthorityYear="1814" box="[275,404,458,481]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="communis"> +<emphasis id="D2B9EA99FFC5A77FFEEAFE5CFE46FE68" box="[275,404,458,481]" italics="true" pageId="14" pageNumber="15">H. communis</emphasis> +</taxonomicName> +. +</paragraph> +<caption id="B4B26603FFC6A77CFF88F8BAFE9FF811" pageId="13" pageNumber="14" startId="13.[113,178,1836,1860]" targetBox="[158,1413,146,1802]" targetPageId="13" targetType="figure"> +<paragraph id="E072368BFFC6A77CFF88F8BAFE9FF811" blockId="13.[113,1450,1836,1944]" pageId="13" pageNumber="14"> +<emphasis id="D2B9EA99FFC6A77CFF88F8BAFF1BF8CD" bold="true" box="[113,201,1836,1860]" pageId="13" pageNumber="14">Figure 5.</emphasis> +Sections of the spongin skeleton of +<taxonomicName id="27CD4D08FFC6A77CFDE2F8BAFD78F8CD" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[539,682,1836,1860]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="13" pageNumber="14" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC6A77CFDE2F8BAFD78F8CD" box="[539,682,1836,1860]" italics="true" pageId="13" pageNumber="14">Spongia lamella</emphasis> +</taxonomicName> +. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres. B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust. D, foreign spicules and debris around canals. E, details of the secondary fibres. +</paragraph> +</caption> +<paragraph id="E072368BFFC5A77FFF65FE7FFED7FC7C" blockId="14.[128,778,144,1014]" pageId="14" pageNumber="15"> +A phylogenetic tree was made by concatenating all sequences available for both 28S and +<emphasis id="D2B9EA99FFC5A77FFE61FD9EFE1AFDA9" box="[408,456,520,544]" italics="true" pageId="14" pageNumber="15">CO1</emphasis> +( +<figureCitation id="78F62A0EFFC5A77FFE23FD9EFDC6FD96" box="[474,532,520,544]" captionStart="Figure 7" captionStartId="16.[129,194,1742,1766]" captionTargetBox="[130,1471,146,1712]" captionTargetId="figure-70@16.[129,1473,144,1714]" captionTargetPageId="16" captionText="Figure 7. Phylogenetic reconstruction of Northeast Atlantic and Mediterranean Spongiidae concatenating the C2–D2 region of 28S and the Folmer fragment of CO1 using both Bayesian inference (BI) and maximum likelihood (ML) methods. Posterior probabilities and bootstraps are given for both analyses (1000 replicates). The ML values (as percentages) are in parentheses. *Sequences obtained only in 28S in this study." pageId="14" pageNumber="15">Fig. 7</figureCitation> +), with the exception of two samples of +<taxonomicName id="27CD4D08FFC5A77FFEDBFDBEFE5BFDB6" authorityName="Schmidt" authorityYear="1862" box="[290,393,552,575]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC5A77FFEDBFDBEFE5BFDB6" box="[290,393,552,575]" italics="true" pageId="14" pageNumber="15">S. zimocca</emphasis> +</taxonomicName> +and +<taxonomicName id="27CD4D08FFC5A77FFE45FDBEFDECFDB6" baseAuthorityName="Lamarck" baseAuthorityYear="1814" box="[444,574,552,575]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="communis"> +<emphasis id="D2B9EA99FFC5A77FFE45FDBEFDECFDB6" box="[444,574,552,575]" italics="true" pageId="14" pageNumber="15">H. communis</emphasis> +</taxonomicName> +for which only 28S could be amplified. The topology of the concatenated tree is congruent with the 28S tree. Both phylogenetic reconstruction methods used here (BI and ML) presented the same pattern. +<specimenCount id="F6CBFD02FFC5A77FFF78FD32FEE4FD35" box="[129,310,676,700]" count="30" pageId="14" pageNumber="15" type="generic">Thirty specimens</specimenCount> +are grouped into two different groups representing the +<taxonomicName id="27CD4D08FFC5A77FFEE9FD52FE50FD55" authorityName="Gray" authorityYear="1867" box="[272,386,708,732]" class="Demospongiae" family="Spongiidae" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="family">Spongiidae</taxonomicName> +family. Group 1 is composed by +<specimenCount id="F6CBFD02FFC5A77FFD1AFD52FF30FD72" count="1" pageId="14" pageNumber="15" type="generic">one specimen</specimenCount> +of the genus +<taxonomicName id="27CD4D08FFC5A77FFE88FD75FE2EFD72" authorityName="Carter" authorityYear="1883" box="[369,508,739,763]" class="Demospongiae" family="Spongiidae" genus="Coscinoderma" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC5A77FFE88FD75FE2EFD72" box="[369,508,739,763]" italics="true" pageId="14" pageNumber="15">Coscinoderma</emphasis> +</taxonomicName> +, whereas group 2 is composed by +<specimenCount id="F6CBFD02FFC5A77FFF1FFC95FEA1FC93" box="[230,371,770,795]" count="29" pageId="14" pageNumber="15" type="generic">29 specimens</specimenCount> +of +<taxonomicName id="27CD4D08FFC5A77FFE6FFC95FE36FC93" authorityName="Linnaeus" authorityYear="1759" box="[406,484,771,794]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC5A77FFE6FFC95FE36FC93" box="[406,484,771,794]" italics="true" pageId="14" pageNumber="15">Spongia</emphasis> +</taxonomicName> +and +<taxonomicName id="27CD4D08FFC5A77FFDE3FC95FD73FC93" authorityName="Schulze" authorityYear="1879" box="[538,673,771,794]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC5A77FFDE3FC95FD73FC93" box="[538,673,771,794]" italics="true" pageId="14" pageNumber="15">Hippospongia</emphasis> +</taxonomicName> +. The proposed new species, +<taxonomicName id="27CD4D08FFC5A77FFEB2FCB4FE18FCB3" box="[331,458,802,826]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC5A77FFEB2FCB4FE18FCB3" box="[331,458,802,826]" italics="true" pageId="14" pageNumber="15">S. maitasuna</emphasis> +</taxonomicName> +, forms a well-supported group in both phylogenetic analyses and still separate but relatively close to +<taxonomicName id="27CD4D08FFC5A77FFF2FFCF7FEEFFCF1" authorityName="Schmidt" authorityYear="1862" box="[214,317,865,888]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC5A77FFF2FFCF7FEEFFCF1" box="[214,317,865,888]" italics="true" pageId="14" pageNumber="15">S. zimocca</emphasis> +</taxonomicName> +, +<taxonomicName id="27CD4D08FFC5A77FFEB0FCF7FE48FCF1" baseAuthorityName="Schmidt" baseAuthorityYear="1862" box="[329,410,865,888]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="nitens"> +<emphasis id="D2B9EA99FFC5A77FFEB0FCF7FE48FCF1" box="[329,410,865,888]" italics="true" pageId="14" pageNumber="15">S. nitens</emphasis> +</taxonomicName> +, and +<taxonomicName id="27CD4D08FFC5A77FFE2BFCF7FDB1FCF1" authorityName="Linnaeus" authorityYear="1759" box="[466,611,864,888]" class="Demospongiae" family="Spongiidae" genus="Spongia" isUncertain="true" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC5A77FFE2BFCF7FE37FCF1" box="[466,485,865,888]" italics="true" pageId="14" pageNumber="15">S.</emphasis> +cf. +<emphasis id="D2B9EA99FFC5A77FFDF1FCF6FDB1FCF1" box="[520,611,864,888]" italics="true" pageId="14" pageNumber="15">officinalis</emphasis> +</taxonomicName> +from +<collectingCountry id="98DA761BFFC5A77FFD5BFCF7FD30FCF0" box="[674,738,865,889]" name="Spain" pageId="14" pageNumber="15">Ceuta</collectingCountry> +. All +<taxonomicName id="27CD4D08FFC5A77FFF79FC16FF32FC11" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[128,224,896,920]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC5A77FFF79FC16FF32FC11" box="[128,224,896,920]" italics="true" pageId="14" pageNumber="15">S. lamella</emphasis> +</taxonomicName> +form a well-supported group, which still appears separate by geographical origin. In this representation, +<taxonomicName id="27CD4D08FFC5A77FFD6CFC09FCDBFC3E" authorityName="Linnaeus" authorityYear="1759" box="[661,777,927,951]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC5A77FFD6CFC09FCDBFC3E" box="[661,777,927,951]" italics="true" pageId="14" pageNumber="15">S. officinalis</emphasis> +</taxonomicName> +and +<taxonomicName id="27CD4D08FFC5A77FFF56FC29FEE1FC5F" baseAuthorityName="Lamarck" baseAuthorityYear="1814" box="[175,307,959,982]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="communis"> +<emphasis id="D2B9EA99FFC5A77FFF56FC29FEE1FC5F" box="[175,307,959,982]" italics="true" pageId="14" pageNumber="15">H. communis</emphasis> +</taxonomicName> +appear mixed in a subgroup also containing +<taxonomicName id="27CD4D08FFC5A77FFF79FC48FF2DFC7F" authorityName="Schmidt" authorityYear="1862" box="[128,255,990,1014]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="mollissima"> +<emphasis id="D2B9EA99FFC5A77FFF79FC48FF2DFC7F" box="[128,255,990,1014]" italics="true" pageId="14" pageNumber="15">S. mollissima</emphasis> +</taxonomicName> +. +</paragraph> +<paragraph id="E072368BFFC5A77FFF27FB83FD7EFBA6" blockId="14.[129,778,1045,1985]" box="[222,684,1045,1072]" pageId="14" pageNumber="15"> +<emphasis id="D2B9EA99FFC5A77FFF27FB83FD7EFBA6" bold="true" box="[222,684,1045,1072]" pageId="14" pageNumber="15">Chemical fingerprints of the studied species</emphasis> +</paragraph> +<paragraph id="E072368BFFC5A77FFF78FBAAFCD8FB78" blockId="14.[129,778,1045,1985]" pageId="14" pageNumber="15"> +We evaluated whether +<taxonomicName id="27CD4D08FFC5A77FFE7BFBAAFE26FBDD" authorityName="Gray" authorityYear="1867" box="[386,500,1084,1108]" class="Demospongiae" family="Spongiidae" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="family">Spongiidae</taxonomicName> +could be organized and grouped based on their chemical fingerprints and whether this chemical-based classification was congruent with the molecular phylogeny. Using the first aligned feature list, containing 915 chemical signals, we performed hierarchical clustering analysis to group sponge extracts according to their chemical similarities. +</paragraph> +<paragraph id="E072368BFFC5A77FFF65FB6EFED7F8CD" blockId="14.[129,778,1045,1985]" pageId="14" pageNumber="15"> +The dendrogram ( +<figureCitation id="78F62A0EFFC5A77FFEA7FB6EFE48FA99" box="[350,410,1272,1296]" captionStart="Figure 8" captionStartId="17.[113,178,1836,1860]" captionTargetBox="[118,1452,144,1805]" captionTargetId="figure-7@17.[114,1458,144,1808]" captionTargetPageId="17" captionText="Figure 8. Metabolomic analysis of Northeast Atlantic and Mediterranean Spongiidae.Dendrogram from hierarchical clustering analysis using Euclidean distance calculations under the Ward algorithm. *Individuals for which no genetic data could be obtained." pageId="14" pageNumber="15">Fig. 8</figureCitation> +) shows distinct groups of extracts all attributed to three distinct groups of species. The first group (group 1) includes all specimens of the proposed new species, +<taxonomicName id="27CD4D08FFC5A77FFF78FAC0FED2FAE7" box="[129,256,1366,1390]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC5A77FFF78FAC0FED2FAE7" box="[129,256,1366,1390]" italics="true" pageId="14" pageNumber="15">S. maitasuna</emphasis> +</taxonomicName> +, in a subgroup, and +<taxonomicName id="27CD4D08FFC5A77FFE2FFAC0FDFBFAE7" baseAuthorityName="Schmidt" baseAuthorityYear="1862" box="[470,553,1366,1390]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="nitens"> +<emphasis id="D2B9EA99FFC5A77FFE2FFAC0FDFBFAE7" box="[470,553,1366,1390]" italics="true" pageId="14" pageNumber="15">S. nitens</emphasis> +</taxonomicName> +and +<taxonomicName id="27CD4D08FFC5A77FFDA4FAC0FD14FAE7" authorityName="Schmidt" authorityYear="1862" box="[605,710,1366,1390]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC5A77FFDA4FAC0FD14FAE7" box="[605,710,1366,1390]" italics="true" pageId="14" pageNumber="15">S. zimocca</emphasis> +</taxonomicName> +in another subgroup. No sample of +<taxonomicName id="27CD4D08FFC5A77FFE4EFAE0FD95FA04" authorityName="Linnaeus" authorityYear="1759" box="[439,583,1397,1421]" class="Demospongiae" family="Spongiidae" genus="Spongia" isUncertain="true" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC5A77FFE4EFAE0FE1BFA04" box="[439,457,1398,1421]" italics="true" pageId="14" pageNumber="15">S.</emphasis> +cf. +<emphasis id="D2B9EA99FFC5A77FFE15FAE3FD95FA04" box="[492,583,1397,1421]" italics="true" pageId="14" pageNumber="15">officinalis</emphasis> +</taxonomicName> +from +<collectingCountry id="98DA761BFFC5A77FFD7DFAE3FD10FA04" box="[644,706,1397,1421]" name="Spain" pageId="14" pageNumber="15">Ceuta</collectingCountry> +(Strait of +<collectingCountry id="98DA761BFFC5A77FFF64FA03FF2CFA24" box="[157,254,1429,1453]" name="Gibraltar" pageId="14" pageNumber="15">Gibraltar</collectingCountry> +) was available for this analysis. The second group (group 2) includes all specimens of +<taxonomicName id="27CD4D08FFC5A77FFDFEFA22FDB8FA45" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[519,618,1460,1484]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC5A77FFDFEFA22FDB8FA45" box="[519,618,1460,1484]" italics="true" pageId="14" pageNumber="15">S. lamella</emphasis> +</taxonomicName> +, still separated into two subgroups according to their geographical origin. The Mediterranean exception (20200602-MRS-MTP8) that was grouping in the 28S phylogenetic with Atlantic specimens is grouped here with all other Mediterranean samples of the same species. The third main group (group 3) includes +<taxonomicName id="27CD4D08FFC5A77FFD76F9C7FCD6F9E1" authorityName="Linnaeus" authorityYear="1759" box="[655,772,1616,1640]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC5A77FFD76F9C7FCD6F9E1" box="[655,772,1616,1640]" italics="true" pageId="14" pageNumber="15">S. officinalis</emphasis> +</taxonomicName> +, +<taxonomicName id="27CD4D08FFC5A77FFF78F9E6FED8F901" authorityName="Schmidt" authorityYear="1862" box="[129,266,1648,1672]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="mollissima"> +<emphasis id="D2B9EA99FFC5A77FFF78F9E6FED8F901" box="[129,266,1648,1672]" italics="true" pageId="14" pageNumber="15">S. mollissima</emphasis> +</taxonomicName> +, and +<taxonomicName id="27CD4D08FFC5A77FFEAEF9E7FE31F901" baseAuthorityName="Lamarck" baseAuthorityYear="1814" box="[343,483,1649,1672]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="communis"> +<emphasis id="D2B9EA99FFC5A77FFEAEF9E7FE31F901" box="[343,483,1649,1672]" italics="true" pageId="14" pageNumber="15">H. communis</emphasis> +</taxonomicName> +, but in the case of this metabolomic analysis, all species formed well-separated subgroups. Thus, although its typology cannot be compared with those of the phylogenetic trees, the groupings presented in the dendrogram resulting from the metabolomic analysis are well congruent with those permitted by the morphological and genetic analyses. +</paragraph> +<paragraph id="E072368BFFC5A77FFF65F8DAFB29FE8F" blockId="14.[129,778,1045,1985]" lastBlockId="14.[825,1475,144,1985]" pageId="14" pageNumber="15"> +The heatmap ( +<figureCitation id="78F62A0EFFC5A77FFEC9F8DDFEBAF8EA" box="[304,360,1867,1891]" captionStart="Figure 9" captionStartId="18.[129,194,1348,1372]" captionTargetBox="[134,1471,147,1317]" captionTargetId="figure-289@18.[129,1473,144,1321]" captionTargetPageId="18" captionText="Figure 9. Metabolomic analysis of Northeast Atlantic and Mediterranean Spongiidae.Heatmap from hierarchical clustering analysis using Euclidean distance calculations under the Ward algorithm, grouping sponge extracts based on their chemical similarities. Chemical features or signals are depicted as horizontal lines, with their colour indicating the normalized level of detection intensity on a scale from −2 to +2, from dark blue to red. The heatmap shows that each species is characterized by a unique set of chemical features (dark red), collectively defining their chemical signature." pageId="14" pageNumber="15">Fig. 9</figureCitation> +) complements these results by depicting the whole set of chemical features that participated in the sample classification. This representation illustrates clearly that extracts from +<taxonomicName id="27CD4D08FFC5A77FFF44F83CFEEDF848" box="[189,319,1962,1985]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC5A77FFF44F83CFEEDF848" box="[189,319,1962,1985]" italics="true" pageId="14" pageNumber="15">S. maitasuna</emphasis> +</taxonomicName> +contain a distinct set of chemical features compared with other +<taxonomicName id="27CD4D08FFC5A77FFBDCFF06FB45FF21" authorityName="Gray" authorityYear="1867" box="[1061,1175,144,168]" class="Demospongiae" family="Spongiidae" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="family">Spongiidae</taxonomicName> +samples. Next, we assessed the distribution of features within the three identified groups of species to evaluate the chemical diversity and number of unique chemical signals characterizing each species. +</paragraph> +<paragraph id="E072368BFFC5A77FFCACFE98FB03FE68" blockId="14.[825,1475,144,1985]" pageId="14" pageNumber="15"> +To that end, a second aligned feature list was generated from pooled extracts from each identified taxonomic unit and contained 240 features. Those features represent the most abundant and redundant chemical signals in each taxonomic unit. Venn diagrams were constructed to depict the number of chemical features shared between species as opposed to those that were detected in one or two species ( +<figureCitation id="78F62A0EFFC5A77FFB85FE5FFB13FE68" box="[1148,1217,457,481]" captionStart="Figure 10" captionStartId="19.[115,180,1389,1413]" captionTargetBox="[116,1453,146,1357]" captionTargetId="figure-314@19.[114,1458,144,1361]" captionTargetPageId="19" captionText="Figure 10. Venn diagrams representing the distribution of chemical features between species of the three main hierarchical groups of Spongiidae of the Mediterranean Sea and Northeast Atlantic Ocean. Black numbers indicate a unique set of features for each species. White numbers represent shared chemical signals between species. The number of features shared between all species of a given group are in white bold." pageId="14" pageNumber="15">Fig. 10</figureCitation> +). +</paragraph> +<paragraph id="E072368BFFC5A77FFCACFE7FFB81FCB3" blockId="14.[825,1475,144,1985]" pageId="14" pageNumber="15"> +The lowest number of features was detected for group 1, for which 45 chemical signals were shared between all three species ( +<taxonomicName id="27CD4D08FFC5A77FFCBDFDBEFC10FDB6" box="[836,962,552,575]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC5A77FFCBDFDBEFC10FDB6" box="[836,962,552,575]" italics="true" pageId="14" pageNumber="15">S. maitasuna</emphasis> +</taxonomicName> +, +<taxonomicName id="27CD4D08FFC5A77FFC34FDBEFBCFFDB6" baseAuthorityName="Schmidt" baseAuthorityYear="1862" box="[973,1053,552,575]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="nitens"> +<emphasis id="D2B9EA99FFC5A77FFC34FDBEFBCFFDB6" box="[973,1053,552,575]" italics="true" pageId="14" pageNumber="15">S. nitens</emphasis> +</taxonomicName> +, and +<taxonomicName id="27CD4D08FFC5A77FFBAAFDBEFB6BFDB6" authorityName="Schmidt" authorityYear="1862" box="[1107,1209,552,575]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC5A77FFBAAFDBEFB6BFDB6" box="[1107,1209,552,575]" italics="true" pageId="14" pageNumber="15">S. zimocca</emphasis> +</taxonomicName> +). A total of 16 unique signals were detected for +<taxonomicName id="27CD4D08FFC5A77FFBDEFDD1FB75FDD7" box="[1063,1191,583,606]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC5A77FFBDEFDD1FB75FDD7" box="[1063,1191,583,606]" italics="true" pageId="14" pageNumber="15">S. maitasuna</emphasis> +</taxonomicName> +, illustrating a much higher chemical diversity and uniqueness in extracts than in +<taxonomicName id="27CD4D08FFC5A77FFA96FDF0FA10FDF7" baseAuthorityName="Schmidt" baseAuthorityYear="1862" box="[1391,1474,614,638]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="nitens"> +<emphasis id="D2B9EA99FFC5A77FFA96FDF0FA10FDF7" box="[1391,1474,614,638]" italics="true" pageId="14" pageNumber="15">S. nitens</emphasis> +</taxonomicName> +(2) and +<taxonomicName id="27CD4D08FFC5A77FFC69FD10FC25FD14" authorityName="Schmidt" authorityYear="1862" box="[912,1015,646,669]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC5A77FFC69FD10FC25FD14" box="[912,1015,646,669]" italics="true" pageId="14" pageNumber="15">S. zimocca</emphasis> +</taxonomicName> +(1). At this stage, none of these features has been identified formally, because the purpose of this analysis was not to assign the associated molecular structure to each feature. However, it is certain that none of the well-known and referenced compounds from groups 2 and 3 (as shown later) were detected among the signals. +</paragraph> +<paragraph id="E072368BFFC5A77FFCACFCD7FB5EFBDD" blockId="14.[825,1475,144,1985]" pageId="14" pageNumber="15"> +The highest number of features was associated with group 2, represented by +<taxonomicName id="27CD4D08FFC5A77FFC28FCF7FBFFFCF1" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[977,1069,864,888]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC5A77FFC28FCF7FBFFFCF1" box="[977,1069,864,888]" italics="true" pageId="14" pageNumber="15">S. lamella</emphasis> +</taxonomicName> +from different locations (Mediterranean and Atlantic). There were between 80% and 85% similarities of detected signals ( +<quantity id="27359B6EFFC5A77FFC09FC09FBEAFC3E" box="[1008,1080,927,951]" metricMagnitude="0" metricUnit="m" metricValue="2.8194" pageId="14" pageNumber="15" unit="in" value="111.0">111 in</quantity> +total) in all extracts. Among the detected chemical features, nitenin is one of the most abundant and well-known furanoterpenoids from this species ( +<bibRefCitation id="845C4B7AFFC5A77FFA91FC48FC58FB9C" author="Noyer" etAl="et al." firstAuthor="Noyer" pageId="14" pageNumber="15" refId="ref18232" refString="Noyer C, Thomas OP, Becerro MA. Patterns of chemical diversity in the Mediterranean sponge Spongia lamella. PLoS One 2011; 6: e 20844. https: // doi. org / 10.1371 / journal. pone. 0020844" type="journal volume" year="2011"> +Noyer +<emphasis id="D2B9EA99FFC5A77FFA48FC48FC80FB9C" italics="true" pageId="14" pageNumber="15">et al.</emphasis> +2011 +</bibRefCitation> +). Extracts from the Mediterranean specimens present a greater number of unique detected signals (13) than those from the Atlantic specimens (5 and 4). +</paragraph> +<paragraph id="E072368BFFC5A77FFCACFBCDFB7EF92E" blockId="14.[825,1475,144,1985]" pageId="14" pageNumber="15"> +Group 3, represented by +<taxonomicName id="27CD4D08FFC5A77FFB8BFBCDFB3CFBFA" authorityName="Linnaeus" authorityYear="1759" box="[1138,1262,1115,1139]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC5A77FFB8BFBCDFB3CFBFA" box="[1138,1262,1115,1139]" italics="true" pageId="14" pageNumber="15">S. officinalis</emphasis> +</taxonomicName> +, +<taxonomicName id="27CD4D08FFC5A77FFAF9FBCDFA5BFBFA" authorityName="Schmidt" authorityYear="1862" box="[1280,1417,1115,1139]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="mollissima"> +<emphasis id="D2B9EA99FFC5A77FFAF9FBCDFA5BFBFA" box="[1280,1417,1115,1139]" italics="true" pageId="14" pageNumber="15">S. mollissima</emphasis> +</taxonomicName> +, and +<taxonomicName id="27CD4D08FFC5A77FFCC0FBEDFC10FB1B" baseAuthorityName="Lamarck" baseAuthorityYear="1814" box="[825,962,1147,1170]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="communis"> +<emphasis id="D2B9EA99FFC5A77FFCC0FBEDFC10FB1B" box="[825,962,1147,1170]" italics="true" pageId="14" pageNumber="15">H. communis</emphasis> +</taxonomicName> +, also contains a relatively high number of detected features compared with the proposed new species. Extracts from these three species have 98 chemical signals in common. Some other well-known and characteristic furanoterpenoid compounds could be identified among these signals, such as furospongin-1, furospongin-4, and demethylfurospongin-4 ( +<bibRefCitation id="845C4B7AFFC5A77FFBB1FAA0FAC6FAC7" author="Bauvais" box="[1096,1300,1334,1358]" etAl="et al." firstAuthor="Bauvais" pageId="14" pageNumber="15" refId="ref16168" refString="Bauvais C, Bonneau N, Blond A et al. Furanoterpene diversity and variability in the marine sponge Spongia officinalis, from untargeted LC - MS / MS metabolomic profiling to furanolactam derivatives. Metabolites 2017; 7: 27. https: // doi. org / 10.3390 / metabo 7020027" type="journal volume" year="2017"> +Bauvais +<emphasis id="D2B9EA99FFC5A77FFB5AFAA1FB05FAC7" box="[1187,1239,1334,1358]" italics="true" pageId="14" pageNumber="15">et al.</emphasis> +2017 +</bibRefCitation> +). Extracts from +<taxonomicName id="27CD4D08FFC5A77FFCC0FAC0FC6FFAE7" authorityName="Schmidt" authorityYear="1862" box="[825,957,1366,1390]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="mollissima"> +<emphasis id="D2B9EA99FFC5A77FFCC0FAC0FC6FFAE7" box="[825,957,1366,1390]" italics="true" pageId="14" pageNumber="15">S. mollissima</emphasis> +</taxonomicName> +have the highest number of unique detected signals (23), compared with +<taxonomicName id="27CD4D08FFC5A77FFBA0FAE3FB01FA04" authorityName="Linnaeus" authorityYear="1759" box="[1113,1235,1397,1421]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC5A77FFBA0FAE3FB01FA04" box="[1113,1235,1397,1421]" italics="true" pageId="14" pageNumber="15">S. officinalis</emphasis> +</taxonomicName> +(3) and +<taxonomicName id="27CD4D08FFC5A77FFAC3FAE0FA10FA04" baseAuthorityName="Lamarck" baseAuthorityYear="1814" box="[1338,1474,1398,1421]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="communis"> +<emphasis id="D2B9EA99FFC5A77FFAC3FAE0FA10FA04" box="[1338,1474,1398,1421]" italics="true" pageId="14" pageNumber="15">H. communis</emphasis> +</taxonomicName> +(9). Among all +<taxonomicName id="27CD4D08FFC5A77FFC22FA02FB81FA25" authorityName="Gray" authorityYear="1867" box="[987,1107,1428,1452]" class="Demospongiae" family="Spongiidae" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="family">Spongiidae</taxonomicName> +, +<taxonomicName id="27CD4D08FFC5A77FFBA5FA03FB6DFA25" baseAuthorityName="Schulze" baseAuthorityYear="1879" box="[1116,1215,1428,1452]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC5A77FFBA5FA03FB6DFA25" box="[1116,1215,1428,1452]" italics="true" pageId="14" pageNumber="15">S. lamella</emphasis> +</taxonomicName> +is the species presenting the highest chemical diversity, with a total of 170 chemical signals (including its unique chemical signals and others shared with other +<taxonomicName id="27CD4D08FFC5A77FFC54FA64FBFBF983" authorityName="Gray" authorityYear="1867" box="[941,1065,1522,1546]" class="Demospongiae" family="Spongiidae" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="family">Spongiidae</taxonomicName> +). In comparison, extracts from the proposed new species contain 151 chemical features, and>10% of them are unique and thus have unknown molecular structures. After undergoing purification and determination of their chemical/molecular structure, they have the potential to become chemotaxonomic markers. +</paragraph> +<paragraph id="E072368BFFC5A77FFCACF938FB45F848" blockId="14.[825,1475,144,1985]" pageId="14" pageNumber="15">Therefore, the hierarchical classification of species, based on their chemical fingerprints, is congruent with the phylogenetic trees previously presented. Each species can be differentiated by its chemical fingerprints associated with a unique set of chemical features that might serve as chemotaxonomic markers. These putative new phenotypic traits would need to be annotated or identified after isolation, purification, and structural analysis by means of additional spectroscopic (e.g. nuclear magnetic resonance) and spectrometric methods.</paragraph> +</subSubSection> +</treatment> +</document> \ No newline at end of file diff --git a/data/68/64/87/6864879DFFCDA777FEDAFCDFFD66FC73.xml b/data/68/64/87/6864879DFFCDA777FEDAFCDFFD66FC73.xml new file mode 100644 index 00000000000..db0fe7244fb --- /dev/null +++ b/data/68/64/87/6864879DFFCDA777FEDAFCDFFD66FC73.xml @@ -0,0 +1,97 @@ +<document id="9C639FD6B65CD396D67B186907722711" ID-DOI="10.1093/zoolinnean/zlad166" ID-ISSN="0024-4082" ID-ZooBank="ED45D8A-954B-4723-9D0F-714357A7DAB5" IM.bibliography_approvedBy="guilherme" IM.illustrations_approvedBy="guilherme" IM.materialsCitations_approvedBy="guilherme" IM.metadata_approvedBy="guilherme" IM.tables_requiresApprovalFor="existingObjects" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="guilherme" checkinTime="1732842034475" checkinUser="plazi" docAuthor="Grenier, Marie, Simmler, Charlotte, Chevaldonné, Pierre, Callizot, Noëlle & Pérez, Thierry" docDate="2024" docId="6864879DFFCDA777FEDAFCDFFD66FC73" docLanguage="en" docName="zlad166.pdf" docOrigin="Zoological Journal of the Linnean Society 202" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.9:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="9" docTitle="Spongia Linnaeus 1759" docType="treatment" docVersion="3" lastPageNumber="7" masterDocId="945DFFE5FFCBA771FFF9FF96FFD2FF89" masterDocTitle="Searching for Mediterranean bath sponges (Demospongiae: Dictyoceratida: Spongiidae) in the Northeast Atlantic reveals a new species: an integrative taxonomic approach" masterLastPageNumber="23" masterPageNumber="1" pageNumber="7" updateTime="1733770290331" updateUser="guilherme"> +<mods:mods id="5C53154DE4763220D8DB067608F5A1A7" xmlns:mods="http://www.loc.gov/mods/v3"> +<mods:titleInfo id="DF8DE0CE0373DE9A4F3929AF00D90BC6"> +<mods:title id="EDE9AC15468E72DB4D2825E877B46A29">Searching for Mediterranean bath sponges (Demospongiae: Dictyoceratida: Spongiidae) in the Northeast Atlantic reveals a new species: an integrative taxonomic approach</mods:title> +</mods:titleInfo> +<mods:name id="2037FB6A47DD8DD401A2F1AD1A639763" type="personal"> +<mods:role id="51D20E93D4D75B81E5F23017307CD048"> +<mods:roleTerm id="A5E429B1F706755DDD07B263067DF1EF">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="ED35FD0C2E88D2CDCCD4D8487E997B80">Grenier, Marie</mods:namePart> +<mods:affiliation id="27F0F5D385A7385B70C25A79A4FDEAA7">Institut Méditerranéen de Biodiversité et d’Ecologie marine et continentale (IMBE), UMR CNRS 7263, IRD 237 - 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Perez" country="Spain" elevation="12" latitude="43.4725" location="Mouro island" longLatPrecision="19" longitude="-3.755" municipality="Santander" pageId="6" pageNumber="7" specimenCode="MNHN-IP-2018-421" specimenCount="1" typeStatus="paratype"> +<emphasis id="D2B9EA99FFCDA777FF78F981FF20F9A7" box="[129,242,1558,1582]" italics="true" pageId="6" pageNumber="7"> +<typeStatus id="3F768829FFCDA777FF78F981FF0AF9A7" box="[129,216,1559,1582]" pageId="6" pageNumber="7">Paratype</typeStatus> +2: +</emphasis> +<specimenCode id="B06B9EF0FFCDA777FF0EF980FE04F9A7" box="[247,470,1558,1582]" pageId="6" pageNumber="7">MNHN-IP-2018-421</specimenCode> +, Eastern part of +<location id="E5126050FFCDA777FD83F980FCD4F9A7" LSID="urn:lsid:plazi:treatment:6864879DFFCDA77BFF4FFB8FFC59FDB6:E5126050FFCDA777FD83F980FCD4F9A7" box="[634,774,1558,1582]" country="Spain" latitude="43.4725" longLatPrecision="19" longitude="-3.755" municipality="Santander" name="Mouro island" pageId="6" pageNumber="7">Mouro island</location> +, shallow water cliffs and chaos of rocks and crevices, between 3 and +<quantity id="27359B6EFFCDA777FF49F9C3FF3BF9E5" box="[176,233,1621,1645]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0"> +<elevation id="6BE0D1B8FFCDA777FF49F9C3FF3BF9E5" box="[176,233,1621,1645]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</elevation> +</quantity> +depth, +<collectingMunicipality id="0016ACF1FFCDA777FEC4F9C3FE74F9E4" box="[317,422,1621,1645]" pageId="6" pageNumber="7">Santander</collectingMunicipality> +, +<collectingCountry id="98DA761BFFCDA777FE4AF9C3FE3EF9E4" box="[435,492,1621,1645]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FE04F9C3FDA8F9E5" box="[509,634,1619,1645]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="21" value="43.4725"> +43°28 +<emphasis id="D2B9EA99FFCDA777FDC4F9C5FD91F9E5" box="[573,579,1619,1644]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +21″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FD7EF9C2FD2CF9E5" box="[647,766,1619,1644]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="18" value="-3.755"> +3°45 +<emphasis id="D2B9EA99FFCDA777FD4FF9C5FD6EF9E5" box="[694,700,1619,1644]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +18″W +</geoCoordinate> +). Collector: +<collectorName id="4D38535DFFCDA777FF14F9E2FE92F905" box="[237,320,1652,1676]" pageId="6" pageNumber="7">T. Pérez</collectorName> +, +<date id="9473104BFFCDA777FEB1F9E2FE1AF905" box="[328,456,1652,1676]" pageId="6" pageNumber="7" value="2021-07-30"> +<collectingDate id="8437E9A3FFCDA777FEB1F9E2FE1AF905" box="[328,456,1652,1676]" pageId="6" pageNumber="7" value="2021-07-30">30 July 2021</collectingDate> +</date> +. Voucher code: 20210730-ESP-M2TP1. 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Collector: P. Chevaldonné, +<date id="9473104BFFCDA777FF78F8D9FEDBF8EE" box="[129,265,1871,1895]" pageId="6" pageNumber="7" value="2021-06-30">30 June 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FF65F8F9FF69F84C" blockId="6.[129,778,1770,1989]" pageId="6" pageNumber="7"> +20210726-CIB-BTP11, Baloo tunnel at +<quantity id="27359B6EFFCDA777FDC8F8F9FDB7F80F" box="[561,613,1903,1927]" metricMagnitude="1" metricUnit="m" metricValue="1.4" pageId="6" pageNumber="7" unit="m" value="14.0">14 m</quantity> +depth, Ciboure, +<collectingCountry id="98DA761BFFCDA777FF78F818FF14F82C" box="[129,198,1934,1957]" name="France" pageId="6" pageNumber="7">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FF2FF818FE81F82F" box="[214,339,1932,1958]" degrees="43" direction="north" minutes="24" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="59" value="43.41639"> +43°24 +<emphasis id="D2B9EA99FFCDA777FEEFF81AFECEF82C" box="[278,284,1932,1957]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +59″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FE99F818FE05F82F" box="[352,471,1932,1958]" degrees="1" direction="west" minutes="39" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="52" value="-1.6644444"> +1°39 +<emphasis id="D2B9EA99FFCDA777FE76F81AFE47F82C" box="[399,405,1932,1957]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +52″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FD46F818FF65F84C" pageId="6" pageNumber="7" value="2021-07-26">26 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFF06FA11FF4E" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210726-CIB-BTP12, Baloo tunnel at +<quantity id="27359B6EFFCDA777FB1EFF06FACCFF2E" box="[1255,1310,144,168]" metricMagnitude="1" metricUnit="m" metricValue="1.4" pageId="6" pageNumber="7" unit="m" value="14.0">14 m</quantity> +depth, Ciboure, +<collectingCountry id="98DA761BFFCDA777FCC0FF26FCAEFF4E" box="[825,892,176,199]" name="France" pageId="6" pageNumber="7">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FC7EFF39FC2CFF4E" box="[903,1022,174,199]" degrees="43" direction="north" minutes="24" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="59" value="43.41639"> +43°24 +<emphasis id="D2B9EA99FFCDA777FC3DFF38FC18FF4E" box="[964,970,174,199]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +59″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FBFFFF39FBAAFF4E" box="[1030,1144,174,199]" degrees="1" direction="west" minutes="39" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="52" value="-1.6644444"> +1°39 +<emphasis id="D2B9EA99FFCDA777FBCAFF38FBEBFF4E" box="[1075,1081,174,199]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +52″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FABDFF39FA6DFF4E" box="[1348,1471,175,199]" pageId="6" pageNumber="7" value="2021-07-26">26 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFF59FA4FFEAC" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210729-ESP-MTP2, Western part of Mouro island, semi-dark to dark cave, between 3 and +<quantity id="27359B6EFFCDA777FB6BFF78FB1AFE8C" box="[1170,1224,238,262]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FA70FF78FA10FE8F" box="[1417,1474,238,262]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FCB8FE9BFC6CFEAC" box="[833,958,268,293]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="43.47278"> +43°28 +<emphasis id="D2B9EA99FFCDA777FC79FE9AFC54FEAC" box="[896,902,268,293]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FC3EFE9BFBECFEAC" box="[967,1086,268,293]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="-3.7561111"> +3°45 +<emphasis id="D2B9EA99FFCDA777FC0FFE9AFC2EFEAC" box="[1014,1020,268,293]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FAEEFE9BFA4BFEAC" box="[1303,1433,269,293]" pageId="6" pageNumber="7" value="2021-07-29">29 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFEBBFA4FFE0A" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210729-ESP-MTP3, Western part of Mouro island, semi-dark to dark cave, between 3 and +<quantity id="27359B6EFFCDA777FB6BFEDAFB1AFEEA" box="[1170,1224,332,356]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FA70FEDAFA10FEED" box="[1417,1474,332,356]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FCB8FEFDFC6CFE0A" box="[833,958,362,387]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="43.47278"> +43°28 +<emphasis id="D2B9EA99FFCDA777FC79FEFCFC54FE0A" box="[896,902,362,387]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FC3EFEFDFBECFE0A" box="[967,1086,362,387]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="-3.7561111"> +3°45 +<emphasis id="D2B9EA99FFCDA777FC0FFEFCFC2EFE0A" box="[1014,1020,362,387]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FAEEFEFDFA4BFE0A" box="[1303,1433,363,387]" pageId="6" pageNumber="7" value="2021-07-29">29 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFE1DFA4FFE68" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210729-ESP-MTP4, Western part of Mouro island, semi-dark to dark cave, between 3 and +<quantity id="27359B6EFFCDA777FB6BFE3CFB1AFE48" box="[1170,1224,426,450]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FA70FE3CFA10FE4B" box="[1417,1474,426,450]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FCB8FE5FFC6CFE68" box="[833,958,456,481]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="43.47278"> +43°28 +<emphasis id="D2B9EA99FFCDA777FC79FE5EFC54FE68" box="[896,902,456,481]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FC3EFE5FFBECFE68" box="[967,1086,456,481]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="-3.7561111"> +3°45 +<emphasis id="D2B9EA99FFCDA777FC0FFE5EFC2EFE68" box="[1014,1020,456,481]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FAEEFE5FFA4BFE68" box="[1303,1433,457,481]" pageId="6" pageNumber="7" value="2021-07-29">29 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFE7EFA4FFDB6" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210729-ESP-MTP6, Western part of Mouro island, semi-dark to dark cave, between 3 and +<quantity id="27359B6EFFCDA777FB6BFD9EFB1AFD96" box="[1170,1224,520,544]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FA70FD9EFA10FDA9" box="[1417,1474,520,544]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FCB8FDB1FC6CFDB6" box="[833,958,550,575]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="43.47278"> +43°28 +<emphasis id="D2B9EA99FFCDA777FC79FDB0FC54FDB6" box="[896,902,550,575]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FC3EFDB1FBECFDB6" box="[967,1086,550,575]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="-3.7561111"> +3°45 +<emphasis id="D2B9EA99FFCDA777FC0FFDB0FC2EFDB6" box="[1014,1020,550,575]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FAEEFDB1FA4BFDB6" box="[1303,1433,551,575]" pageId="6" pageNumber="7" value="2021-07-29">29 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFDD1FA4FFD14" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210729-ESP-MTP7, Western part of Mouro island, semi-dark to dark cave, between 3 and +<quantity id="27359B6EFFCDA777FB6BFDF0FB1AFDF4" box="[1170,1224,614,638]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FA70FDF0FA10FDF7" box="[1417,1474,614,638]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FCB8FD13FC6CFD14" box="[833,958,644,669]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="43.47278"> +43°28 +<emphasis id="D2B9EA99FFCDA777FC79FD12FC54FD14" box="[896,902,644,669]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FC3EFD13FBECFD14" box="[967,1086,644,669]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="-3.7561111"> +3°45 +<emphasis id="D2B9EA99FFCDA777FC0FFD12FC2EFD14" box="[1014,1020,644,669]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FAEEFD13FA4BFD14" box="[1303,1433,645,669]" pageId="6" pageNumber="7" value="2021-07-29">29 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFD33FA4FFD72" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210729-ESP-MTP8, Western part of Mouro island, semi-dark to dark cave, between 3 and +<quantity id="27359B6EFFCDA777FB6BFD52FB1AFD52" box="[1170,1224,708,732]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FA70FD52FA10FD55" box="[1417,1474,708,732]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FCB8FD75FC6CFD72" box="[833,958,738,763]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="43.47278"> +43°28 +<emphasis id="D2B9EA99FFCDA777FC79FD74FC54FD72" box="[896,902,738,763]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FC3EFD75FBECFD72" box="[967,1086,738,763]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="-3.7561111"> +3°45 +<emphasis id="D2B9EA99FFCDA777FC0FFD74FC2EFD72" box="[1014,1020,738,763]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FAEEFD75FA4BFD72" box="[1303,1433,739,763]" pageId="6" pageNumber="7" value="2021-07-29">29 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFC95FB5BFCF0" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210730-ESP-M2TP7, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and +<quantity id="27359B6EFFCDA777FC90FCD7FC70FCD0" box="[873,930,833,857]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FB95FCD7FB77FCD0" box="[1132,1189,833,857]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FB4FFCD7FAE1FCD0" box="[1206,1331,832,857]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="21" value="43.4725"> +43°28 +<emphasis id="D2B9EA99FFCDA777FB0FFCD6FB2EFCD0" box="[1270,1276,832,857]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +21″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FAB9FCD7FA64FCD0" box="[1344,1462,832,857]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="18" value="-3.755"> +3°45 +<emphasis id="D2B9EA99FFCDA777FA96FCD6FAA7FCD0" box="[1391,1397,832,857]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +18″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FBFAFCF7FB57FCF0" box="[1027,1157,865,889]" pageId="6" pageNumber="7" value="2021-07-30">30 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFC16FB5BFC7F" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210730-ESP-M2TP8, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and +<quantity id="27359B6EFFCDA777FC90FC29FC70FC5F" box="[873,930,959,983]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FB95FC29FB77FC5E" box="[1132,1189,959,983]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FB4FFC29FAE1FC5F" box="[1206,1331,957,983]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="21" value="43.4725"> +43°28 +<emphasis id="D2B9EA99FFCDA777FB0FFC2BFB2EFC5F" box="[1270,1276,957,982]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +21″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FAB9FC28FA64FC5F" box="[1344,1462,957,982]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="18" value="-3.755"> +3°45 +<emphasis id="D2B9EA99FFCDA777FA96FC2BFAA7FC5F" box="[1391,1397,957,982]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +18″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FBFAFC48FB57FC7F" box="[1027,1157,990,1014]" pageId="6" pageNumber="7" value="2021-07-30">30 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFC6BFB5BFBFA" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210730-ESP-M2TP9, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and +<quantity id="27359B6EFFCDA777FC90FBAAFC70FBDA" box="[873,930,1084,1108]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FB95FBAAFB77FBDD" box="[1132,1189,1084,1108]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FB4FFBAAFAE1FBDA" box="[1206,1331,1082,1108]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="21" value="43.4725"> +43°28 +<emphasis id="D2B9EA99FFCDA777FB0FFBACFB2EFBDA" box="[1270,1276,1082,1107]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +21″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FAB9FBAAFA64FBDA" box="[1344,1462,1082,1108]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="18" value="-3.755"> +3°45 +<emphasis id="D2B9EA99FFCDA777FA96FBACFAA7FBDA" box="[1391,1397,1082,1107]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +18″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FBFAFBCDFB57FBFA" box="[1027,1157,1115,1139]" pageId="6" pageNumber="7" value="2021-07-30">30 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFBEDFB5BFB78" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210730-ESP-M2TP10, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and +<quantity id="27359B6EFFCDA777FC90FB2FFC70FB58" box="[873,930,1209,1233]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FB95FB2FFB77FB58" box="[1132,1189,1209,1233]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FB4FFB2FFAE1FB58" box="[1206,1331,1208,1233]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="21" value="43.4725"> +43°28 +<emphasis id="D2B9EA99FFCDA777FB0FFB2EFB2EFB58" box="[1270,1276,1208,1233]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +21″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FAB9FB2FFA64FB58" box="[1344,1462,1208,1233]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="18" value="-3.755"> +3°45 +<emphasis id="D2B9EA99FFCDA777FA96FB2EFAA7FB58" box="[1391,1397,1208,1233]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +18″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FBFAFB4FFB57FB78" box="[1027,1157,1240,1265]" pageId="6" pageNumber="7" value="2021-07-30">30 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFB6EFB5BFAE7" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210730-ESP-M2TP11, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and +<quantity id="27359B6EFFCDA777FC90FAA1FC70FAC7" box="[873,930,1335,1359]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FB95FAA0FB77FAC7" box="[1132,1189,1334,1358]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FB4FFAA1FAE1FAC7" box="[1206,1331,1333,1359]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="21" value="43.4725"> +43°28 +<emphasis id="D2B9EA99FFCDA777FB0FFAA3FB2EFAC7" box="[1270,1276,1333,1358]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +21″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FAB9FAA0FA64FAC7" box="[1344,1462,1333,1358]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="18" value="-3.755"> +3°45 +<emphasis id="D2B9EA99FFCDA777FA96FAA3FAA7FAC7" box="[1391,1397,1333,1358]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +18″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FBFAFAC0FB57FAE7" box="[1027,1157,1366,1390]" pageId="6" pageNumber="7" value="2021-07-30">30 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFAE3FB5BFA62" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210730-ESP-M2TP13, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and +<quantity id="27359B6EFFCDA777FC90FA22FC70FA42" box="[873,930,1460,1484]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FB95FA22FB77FA45" box="[1132,1189,1460,1484]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FB4FFA22FAE1FA42" box="[1206,1331,1458,1484]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="21" value="43.4725"> +43°28 +<emphasis id="D2B9EA99FFCDA777FB0FFA24FB2EFA42" box="[1270,1276,1458,1483]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +21″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FAB9FA22FA64FA42" box="[1344,1462,1458,1484]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="18" value="-3.755"> +3°45 +<emphasis id="D2B9EA99FFCDA777FA96FA24FAA7FA42" box="[1391,1397,1458,1483]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +18″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FBFAFA45FB57FA62" box="[1027,1157,1491,1515]" pageId="6" pageNumber="7" value="2021-07-30">30 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACFA65FB5BF9E0" blockId="6.[825,1475,144,1641]" pageId="6" pageNumber="7"> +20210730-ESP-M2TP14, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and +<quantity id="27359B6EFFCDA777FC90F9A7FC70F9C0" box="[873,930,1585,1609]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="7" unit="m" value="12.0">12 m</quantity> +depth, Santander, +<collectingCountry id="98DA761BFFCDA777FB95F9A7FB77F9C0" box="[1132,1189,1585,1609]" name="Spain" pageId="6" pageNumber="7">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FB4FF9A7FAE1F9C0" box="[1206,1331,1584,1609]" degrees="43" direction="north" minutes="28" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="21" value="43.4725"> +43°28 +<emphasis id="D2B9EA99FFCDA777FB0FF9A6FB2EF9C0" box="[1270,1276,1584,1609]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +21″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FAB9F9A7FA64F9C0" box="[1344,1462,1584,1609]" degrees="3" direction="west" minutes="45" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="18" value="-3.755"> +3°45 +<emphasis id="D2B9EA99FFCDA777FA96F9A6FAA7F9C0" box="[1391,1397,1584,1609]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +18″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FBFAF9C7FB57F9E0" box="[1027,1157,1616,1641]" pageId="6" pageNumber="7" value="2021-07-30">30 July 2021</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FC1FF91FFAC7F92B" blockId="6.[998,1301,1672,1699]" box="[998,1301,1672,1699]" pageId="6" pageNumber="7"> +<emphasis id="D2B9EA99FFCDA777FC1FF91FFAC7F92B" box="[998,1301,1672,1699]" italics="true" pageId="6" pageNumber="7">Comparative material examined</emphasis> +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFCDA777FC2DF92EFAFAF95B" box="[980,1320,1719,1746]" pageId="6" pageNumber="7" type="multiple"> +<paragraph id="E072368BFFCDA777FC2DF92EFAFAF95B" blockId="6.[980,1320,1719,1746]" box="[980,1320,1719,1746]" pageId="6" pageNumber="7"> +<taxonomicName id="27CD4D08FFCDA777FC2DF92EFAFAF95B" ID-CoL="6ZBNB" authority="Linnaeus, 1759" authorityName="Linnaeus" authorityYear="1759" box="[980,1320,1719,1746]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="6" pageNumber="7" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFCDA777FC2DF92EFAFAF95B" box="[980,1320,1719,1746]" italics="true" pageId="6" pageNumber="7"> +Spongia officinalis +<bibRefCitation id="845C4B7AFFCDA777FB6DF92EFAFAF95B" author="Linnaeus" box="[1172,1320,1720,1746]" firstAuthor="Linnaeus" pageId="6" pageNumber="7" pagination="825 - 1384" refId="ref17828" refString="Linnaeus C. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus II, Editio decima, reformata, Holmiae. (L. Salvii) 1759; 825 - 1384. https: // www. biodiversitylibrary. org / page / 587267" type="book chapter" year="1759">Linnaeus, 1759</bibRefCitation> +</emphasis> +</taxonomicName> +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFCDA777FBAAF971FB7AF888" box="[1107,1192,1767,1793]" pageId="6" pageNumber="7" type="description"> +<paragraph id="E072368BFFCDA777FBAAF971FB7AF888" blockId="6.[825,1474,1767,1986]" box="[1107,1192,1767,1793]" pageId="6" pageNumber="7"> +<emphasis id="D2B9EA99FFCDA777FBAAF971FB7AF888" box="[1107,1192,1767,1793]" italics="true" pageId="6" pageNumber="7"> +( +<figureCitation id="78F62A0EFFCDA777FBA4F97EFB4CF888" box="[1117,1182,1767,1793]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." pageId="6" pageNumber="7">Fig.2E</figureCitation> +) +</emphasis> +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFCDA779FCC0F89BFE11FB2A" lastPageId="8" lastPageNumber="9" pageId="6" pageNumber="7" type="materials_examined"> +<paragraph id="E072368BFFCDA777FCC0F89BFAD8F8ED" blockId="6.[825,1474,1767,1986]" pageId="6" pageNumber="7"> +20200519-MRS-ITP3, Impérial de Terre, semi-dark cave, between 10 and +<quantity id="27359B6EFFCDA777FC26F8BBFBC9F8CD" box="[991,1051,1837,1861]" metricMagnitude="1" metricUnit="m" metricValue="2.0" pageId="6" pageNumber="7" unit="m" value="20.0">20 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFCDA777FB1EF8BBFAFEF8CD" box="[1255,1324,1837,1860]" name="France" pageId="6" pageNumber="7">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FABBF8BBFA6CF8CD" box="[1346,1470,1835,1861]" degrees="43" direction="north" minutes="10" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="43.17278"> +43°10 +<emphasis id="D2B9EA99FFCDA777FA78F8BDFA55F8CD" box="[1409,1415,1835,1860]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FCC0F8DAFC77F8ED" box="[825,933,1866,1892]" degrees="5" direction="east" minutes="23" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="35" value="5.3930554"> +5°23 +<emphasis id="D2B9EA99FFCDA777FC91F8DCFCBCF8EA" box="[872,878,1866,1891]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +35″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FB87F8DAFAD5F8ED" box="[1150,1287,1868,1892]" pageId="6" pageNumber="7" value="2020-05-19">19 May 2020</date> +. +</paragraph> +<paragraph id="E072368BFFCDA777FCACF8FDFAD8F84B" blockId="6.[825,1474,1767,1986]" pageId="6" pageNumber="7"> +20200519-MRS-ITP4, Impérial de Terre, semi-dark cave, between 10 and +<quantity id="27359B6EFFCDA777FC17F81DFBF5F82B" box="[1006,1063,1931,1955]" metricMagnitude="1" metricUnit="m" metricValue="2.0" pageId="6" pageNumber="7" unit="m" value="20.0">20 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFCDA777FB13F81DFAFDF82B" box="[1258,1327,1931,1954]" name="France" pageId="6" pageNumber="7">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFCDA777FABBF81DFA6CF82B" box="[1346,1470,1929,1955]" degrees="43" direction="north" minutes="10" orientation="latitude" pageId="6" pageNumber="7" precision="15" seconds="22" value="43.17278"> +43°10 +<emphasis id="D2B9EA99FFCDA777FA78F81FFA55F82B" box="[1409,1415,1929,1954]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCDA777FCC0F83CFC77F84B" box="[825,933,1960,1986]" degrees="5" direction="east" minutes="23" orientation="longitude" pageId="6" pageNumber="7" precision="15" seconds="35" value="5.3930554"> +5°23 +<emphasis id="D2B9EA99FFCDA777FC91F83EFCBCF848" box="[872,878,1960,1985]" italics="true" pageId="6" pageNumber="7">ʹ</emphasis> +35″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCDA777FB87F83CFAD5F84B" box="[1150,1287,1962,1986]" pageId="6" pageNumber="7" value="2020-05-19">19 May 2020</date> +. +</paragraph> +<caption id="B4B26603FFCCA776FF88F957FBFCF97C" pageId="7" pageNumber="8" startId="7.[113,178,1729,1753]" targetBox="[116,1454,146,1697]" targetPageId="7" targetType="figure"> +<paragraph id="E072368BFFCCA776FF88F957FBFCF97C" blockId="7.[113,1430,1729,1781]" pageId="7" pageNumber="8"> +<emphasis id="D2B9EA99FFCCA776FF88F957FF1BF950" bold="true" box="[113,201,1729,1753]" pageId="7" pageNumber="8">Figure 2.</emphasis> +A–D, +<taxonomicName id="27CD4D08FFCCA776FEFDF954FEE9F950" box="[260,315,1730,1753]" class="Demospongiae" family="Irciniidae" genus="Ircinia" kingdom="Animalia" order="Dictyoceratida" pageId="7" pageNumber="12" phylum="Porifera" rank="species" species="situ"> +<emphasis id="D2B9EA99FFCCA776FEFDF954FEE9F950" box="[260,315,1730,1753]" italics="true" pageId="7" pageNumber="8">in situ</emphasis> +</taxonomicName> +pictures of +<taxonomicName id="27CD4D08FFCCA776FE51F957FDB7F950" box="[424,613,1729,1753]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="7" pageNumber="8" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFCCA776FE51F957FDB7F950" bold="true" box="[424,613,1729,1753]" italics="true" pageId="7" pageNumber="8">Spongia maitasuna</emphasis> +</taxonomicName> +in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, +<taxonomicName id="27CD4D08FFCCA776FF88F94BFEC1F97C" authorityName="Linnaeus" authorityYear="1759" box="[113,275,1757,1781]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="7" pageNumber="8" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFCCA776FF88F94BFEC1F97C" box="[113,275,1757,1781]" italics="true" pageId="7" pageNumber="8">Spongia officinalis</emphasis> +</taxonomicName> +from Marseille, France. F, +<taxonomicName id="27CD4D08FFCCA776FDF5F94BFD1AF97C" authorityName="Linnaeus" authorityYear="1759" box="[524,712,1757,1781]" class="Demospongiae" family="Spongiidae" genus="Spongia" isUncertain="true" kingdom="Animalia" order="Dictyoceratida" pageId="7" pageNumber="8" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFCCA776FDF5F94BFD87F97D" box="[524,597,1757,1780]" italics="true" pageId="7" pageNumber="8">Spongia</emphasis> +cf. +<emphasis id="D2B9EA99FFCCA776FD8DF94BFD1AF97C" box="[628,712,1757,1781]" italics="true" pageId="7" pageNumber="8">officinalis</emphasis> +</taxonomicName> +from Ceuta, Strait of Gibraltar, Spain. +</paragraph> +</caption> +<paragraph id="E072368BFFCCA776FF74F8B3FD90F8F5" blockId="7.[113,763,1829,2010]" pageId="7" pageNumber="8"> +20200519-MRS-ITP5, Impérial de Terre, semi-dark cave, between 10 and +<quantity id="27359B6EFFCCA776FEDFF8D2FE8DF8D5" box="[294,351,1860,1884]" metricMagnitude="1" metricUnit="m" metricValue="2.0" pageId="7" pageNumber="8" unit="m" value="20.0">20 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFCCA776FDDAF8D3FDBAF8D5" box="[547,616,1861,1884]" name="France" pageId="7" pageNumber="8">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFCCA776FD83F8D2FD24F8D5" box="[634,758,1859,1884]" degrees="43" direction="north" minutes="10" orientation="latitude" pageId="7" pageNumber="8" precision="15" seconds="22" value="43.17278"> +43°10 +<emphasis id="D2B9EA99FFCCA776FD40F8D5FD6DF8D5" box="[697,703,1859,1884]" italics="true" pageId="7" pageNumber="8">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCCA776FF88F8F5FF0FF8F2" box="[113,221,1890,1915]" degrees="5" direction="east" minutes="23" orientation="longitude" pageId="7" pageNumber="8" precision="15" seconds="35" value="5.3930554"> +5°23 +<emphasis id="D2B9EA99FFCCA776FF59F8F4FF74F8F2" box="[160,166,1890,1915]" italics="true" pageId="7" pageNumber="8">ʹ</emphasis> +35″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCCA776FE4FF8F2FDEDF8F5" box="[438,575,1892,1916]" pageId="7" pageNumber="8" value="2020-05-19">19 May 2020</date> +. +</paragraph> +<paragraph id="E072368BFFCCA776FF74F815FD02F853" blockId="7.[113,763,1829,2010]" pageId="7" pageNumber="8"> +20200522-MRS-FTP2, Fauconnière, semi-dark tunnel, between 5 and +<quantity id="27359B6EFFCCA776FEE7F834FE8BF833" box="[286,345,1954,1978]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="7" pageNumber="8" unit="m" value="10.0">10 m</quantity> +depth, Saint-Cyr Les Lecques, +<collectingCountry id="98DA761BFFCCA776FD4CF835FD28F833" box="[693,762,1955,1978]" name="France" pageId="7" pageNumber="8">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFCCA776FF80F854FF24F850" box="[121,246,1984,2010]" degrees="43" direction="north" minutes="09" orientation="latitude" pageId="7" pageNumber="8" precision="15" seconds="16" value="43.154446"> +43°09 +<emphasis id="D2B9EA99FFCCA776FF40F856FF6DF850" box="[185,191,1984,2009]" italics="true" pageId="7" pageNumber="8">ʹ</emphasis> +16″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCCA776FF06F857FEB9F850" box="[255,363,1984,2009]" degrees="5" direction="east" minutes="40" orientation="longitude" pageId="7" pageNumber="8" precision="15" seconds="59" value="5.6830554"> +5°40 +<emphasis id="D2B9EA99FFCCA776FED7F856FEE6F850" box="[302,308,1984,2009]" italics="true" pageId="7" pageNumber="8">ʹ</emphasis> +59″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCCA776FDBDF854FD1EF853" box="[580,716,1986,2010]" pageId="7" pageNumber="8" value="2020-05-22">22 May 2020</date> +. +</paragraph> +<paragraph id="E072368BFFCCA776FCBCF8B3FA5BF8F5" blockId="7.[810,1460,1829,2010]" pageId="7" pageNumber="8"> +20200522-MRS-FTP3, Fauconnière, semi-dark tunnel, between 5 and +<quantity id="27359B6EFFCCA776FC2EF8D2FBC3F8D5" box="[983,1041,1860,1884]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="7" pageNumber="8" unit="m" value="10.0">10 m</quantity> +depth, Saint-Cyr Les Lecques, +<collectingCountry id="98DA761BFFCCA776FA97F8D3FA61F8D5" box="[1390,1459,1861,1884]" name="France" pageId="7" pageNumber="8">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFCCA776FCCBF8F2FC7DF8F2" box="[818,943,1890,1916]" degrees="43" direction="north" minutes="09" orientation="latitude" pageId="7" pageNumber="8" precision="15" seconds="16" value="43.154446"> +43°09 +<emphasis id="D2B9EA99FFCCA776FC88F8F4FCA5F8F2" box="[881,887,1890,1915]" italics="true" pageId="7" pageNumber="8">ʹ</emphasis> +16″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCCA776FC4EF8F5FBF1F8F2" box="[951,1059,1890,1915]" degrees="5" direction="east" minutes="40" orientation="longitude" pageId="7" pageNumber="8" precision="15" seconds="59" value="5.6830554"> +5°40 +<emphasis id="D2B9EA99FFCCA776FC1EF8F4FC3FF8F2" box="[999,1005,1890,1915]" italics="true" pageId="7" pageNumber="8">ʹ</emphasis> +59″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCCA776FB05F8F2FA57F8F5" box="[1276,1413,1892,1916]" pageId="7" pageNumber="8" value="2020-05-22">22 May 2020</date> +. +</paragraph> +<paragraph id="E072368BFFCCA776FCBCF815FC3FF853" blockId="7.[810,1460,1829,2010]" pageId="7" pageNumber="8"> +20200602-MRS-MTP9, Pharillons de Maïre, cliff at +<quantity id="27359B6EFFCCA776FA83F815FA61F813" box="[1402,1459,1923,1947]" metricMagnitude="1" metricUnit="m" metricValue="2.0" pageId="7" pageNumber="8" unit="m" value="20.0">20 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFCCA776FC22F835FBF2F833" box="[987,1056,1955,1978]" name="France" pageId="7" pageNumber="8">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFCCA776FBD7F834FB79F833" box="[1070,1195,1953,1978]" degrees="43" direction="north" minutes="12" orientation="latitude" pageId="7" pageNumber="8" precision="15" seconds="27" value="43.2075"> +43°12 +<emphasis id="D2B9EA99FFCCA776FB97F837FBA6F833" box="[1134,1140,1953,1978]" italics="true" pageId="7" pageNumber="8">ʹ</emphasis> +27″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFCCA776FB4CF834FAF2F833" box="[1205,1312,1953,1978]" degrees="5" direction="east" minutes="20" orientation="longitude" pageId="7" pageNumber="8" precision="15" seconds="17" value="5.3380556"> +5°20 +<emphasis id="D2B9EA99FFCCA776FB1DF837FB38F833" box="[1252,1258,1953,1978]" italics="true" pageId="7" pageNumber="8">ʹ</emphasis> +17″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFCCA776FC94F854FC38F853" box="[877,1002,1986,2010]" pageId="7" pageNumber="8" value="2020-06-02">2 June 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65FF06FE05FF6E" blockId="8.[129,778,144,795]" pageId="8" pageNumber="9"> +20200617-MRS-NTP1, Niolon, semi-dark cave, between 8 and +<quantity id="27359B6EFFC3A779FF3AFF39FF28FF4E" box="[195,250,175,199]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="8" pageNumber="9" unit="m" value="10.0">10 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC3A779FE4FFF26FE29FF4E" box="[438,507,176,199]" name="France" pageId="8" pageNumber="9">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FDF2FF39FD55FF4E" box="[523,647,174,199]" degrees="43" direction="north" minutes="20" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="15" value="43.337498"> +43°20 +<emphasis id="D2B9EA99FFC3A779FDB3FF38FD82FF4E" box="[586,592,174,199]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +15″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FD6AFF39FD2DFF4E" box="[659,767,174,199]" degrees="5" direction="east" minutes="15" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="22" value="5.256111"> +5°15 +<emphasis id="D2B9EA99FFC3A779FD3BFF38FD1AFF4E" box="[706,712,174,199]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +22″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FEB3FF59FE06FF6E" box="[330,468,207,231]" pageId="8" pageNumber="9" value="2020-06-17">17 June 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65FF78FE56FECC" blockId="8.[129,778,144,795]" pageId="8" pageNumber="9"> +20220909-MRS2-RTP4, Riou island, overhang at +<quantity id="27359B6EFFC3A779FD32FF78FCD8FE8C" box="[715,778,238,262]" metricMagnitude="1" metricUnit="m" metricValue="2.8" pageId="8" pageNumber="9" unit="m" value="28.0">28 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC3A779FECBFE98FEA5FEAC" box="[306,375,270,293]" name="France" pageId="8" pageNumber="9">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE7CFE9BFDD3FEAC" box="[389,513,268,293]" degrees="43" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="23" value="43.173058"> +43°10 +<emphasis id="D2B9EA99FFC3A779FE3DFE9AFE18FEAC" box="[452,458,268,293]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +23″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDF2FE9BFDA5FEAC" box="[523,631,268,293]" degrees="5" direction="east" minutes="23" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="15" value="5.3875"> +5°23 +<emphasis id="D2B9EA99FFC3A779FDC3FE9AFD92FEAC" box="[570,576,268,293]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +15″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF3AFEBBFE52FECC" box="[195,384,301,325]" pageId="8" pageNumber="9" value="2022-09-09">9 September 2022</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65FEDAFE56FE2A" blockId="8.[129,778,144,795]" pageId="8" pageNumber="9"> +20220909-MRS2-RTP5, Riou island, overhang at +<quantity id="27359B6EFFC3A779FD32FEDAFCD8FEEA" box="[715,778,332,356]" metricMagnitude="1" metricUnit="m" metricValue="2.8" pageId="8" pageNumber="9" unit="m" value="28.0">28 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC3A779FECBFEFAFEA5FE0A" box="[306,375,364,387]" name="France" pageId="8" pageNumber="9">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE7CFEFDFDD3FE0A" box="[389,513,362,387]" degrees="43" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="23" value="43.173058"> +43°10 +<emphasis id="D2B9EA99FFC3A779FE3DFEFCFE18FE0A" box="[452,458,362,387]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +23″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDF2FEFDFDA5FE0A" box="[523,631,362,387]" degrees="5" direction="east" minutes="23" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="15" value="5.3875"> +5°23 +<emphasis id="D2B9EA99FFC3A779FDC3FEFCFD92FE0A" box="[570,576,362,387]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +15″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF3AFE1DFE52FE2A" box="[195,384,395,419]" pageId="8" pageNumber="9" value="2022-09-09">9 September 2022</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65FE3CFE56FD88" blockId="8.[129,778,144,795]" pageId="8" pageNumber="9"> +20220909-MRS2-RTP6, Riou island, overhang at +<quantity id="27359B6EFFC3A779FD32FE3CFCD8FE48" box="[715,778,426,450]" metricMagnitude="1" metricUnit="m" metricValue="2.8" pageId="8" pageNumber="9" unit="m" value="28.0">28 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC3A779FECBFE5CFEA5FE68" box="[306,375,458,481]" name="France" pageId="8" pageNumber="9">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE7CFE5FFDD3FE68" box="[389,513,456,481]" degrees="43" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="23" value="43.173058"> +43°10 +<emphasis id="D2B9EA99FFC3A779FE3DFE5EFE18FE68" box="[452,458,456,481]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +23″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDF2FE5FFDA5FE68" box="[523,631,456,481]" degrees="5" direction="east" minutes="23" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="15" value="5.3875"> +5°23 +<emphasis id="D2B9EA99FFC3A779FDC3FE5EFD92FE68" box="[570,576,456,481]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +15″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF3AFE7FFE52FD88" box="[195,384,489,513]" pageId="8" pageNumber="9" value="2022-09-09">9 September 2022</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65FD9EFDCAFDD6" blockId="8.[129,778,144,795]" pageId="8" pageNumber="9"> +20220923-MRS2-ITP25, Impérial de Terre, semi-dark cave at +<quantity id="27359B6EFFC3A779FF58FDB1FF0FFDB6" box="[161,221,551,575]" metricMagnitude="1" metricUnit="m" metricValue="2.0" pageId="8" pageNumber="9" unit="m" value="20.0">20 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC3A779FE52FDBEFE22FDB6" box="[427,496,552,575]" name="France" pageId="8" pageNumber="9">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FDFCFDB1FD50FDB6" box="[517,642,550,575]" degrees="43" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="22" value="43.17278"> +43°10 +<emphasis id="D2B9EA99FFC3A779FDBCFDB0FD99FDB6" box="[581,587,550,575]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FD6AFDB1FD2CFDB6" box="[659,766,550,575]" degrees="5" direction="east" minutes="23" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="35" value="5.3930554"> +5°23 +<emphasis id="D2B9EA99FFC3A779FD3BFDB0FD1AFDB6" box="[706,712,550,575]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +35″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FEB3FDD1FDC6FDD6" box="[330,532,582,607]" pageId="8" pageNumber="9" value="2022-09-23">23 September 2022</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65FDF0FDCAFD34" blockId="8.[129,778,144,795]" pageId="8" pageNumber="9"> +20220923-MRS2-ITP26, Impérial de Terre, semi-dark cave at +<quantity id="27359B6EFFC3A779FF58FD13FF0FFD14" box="[161,221,645,669]" metricMagnitude="1" metricUnit="m" metricValue="2.0" pageId="8" pageNumber="9" unit="m" value="20.0">20 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC3A779FE52FD10FE22FD14" box="[427,496,646,669]" name="France" pageId="8" pageNumber="9">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FDFCFD13FD50FD14" box="[517,642,644,669]" degrees="43" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="22" value="43.17278"> +43°10 +<emphasis id="D2B9EA99FFC3A779FDBCFD12FD99FD14" box="[581,587,644,669]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +22″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FD6AFD13FD2CFD14" box="[659,766,644,669]" degrees="5" direction="east" minutes="23" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="35" value="5.3930554"> +5°23 +<emphasis id="D2B9EA99FFC3A779FD3BFD12FD1AFD14" box="[706,712,644,669]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +35″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FEB3FD33FDC6FD34" box="[330,532,676,701]" pageId="8" pageNumber="9" value="2022-09-23">23 September 2022</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65FD52FE43FC92" blockId="8.[129,778,144,795]" pageId="8" pageNumber="9"> +20220923-MRS2-RTP27, Riou island, overhang at +<quantity id="27359B6EFFC3A779FD34FD52FCD8FD52" box="[717,778,708,732]" metricMagnitude="1" metricUnit="m" metricValue="2.5" pageId="8" pageNumber="9" unit="m" value="25.0">25 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC3A779FECBFD72FEA5FD72" box="[306,375,740,763]" name="France" pageId="8" pageNumber="9">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE7CFD75FDD3FD72" box="[389,513,738,763]" degrees="43" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="23" value="43.173058"> +43°10 +<emphasis id="D2B9EA99FFC3A779FE3DFD74FE18FD72" box="[452,458,738,763]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +23″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDF2FD75FDA5FD72" box="[523,631,738,763]" degrees="5" direction="east" minutes="23" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="15" value="5.3875"> +5°23 +<emphasis id="D2B9EA99FFC3A779FDC3FD74FD92FD72" box="[570,576,738,763]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +15″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF3AFC95FE5FFC92" box="[195,397,770,795]" pageId="8" pageNumber="9" value="2022-09-23">23 September 2022</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FEA5FCADFDFCFCDD" blockId="8.[348,558,826,853]" box="[348,558,826,853]" pageId="8" pageNumber="9"> +<taxonomicName id="27CD4D08FFC3A779FEA5FCADFDFCFCDD" authorityName="Linnaeus" authorityYear="1759" box="[348,558,826,853]" class="Demospongiae" family="Spongiidae" genus="Spongia" isUncertain="true" kingdom="Animalia" order="Dictyoceratida" pageId="8" pageNumber="9" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC3A779FEA5FCADFDFCFCDD" box="[348,558,826,853]" italics="true" pageId="8" pageNumber="9">Spongia cf.officinalis</emphasis> +</taxonomicName> +</paragraph> +<paragraph id="E072368BFFC3A779FE62FCFCFE3DFC0D" blockId="8.[129,778,874,1187]" box="[411,495,874,900]" pageId="8" pageNumber="9"> +<emphasis id="D2B9EA99FFC3A779FE62FCFCFE3DFC0D" box="[411,495,874,900]" italics="true" pageId="8" pageNumber="9"> +( +<figureCitation id="78F62A0EFFC3A779FE5CFCFCFE37FC0D" box="[421,485,874,900]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." pageId="8" pageNumber="9">Fig.2F</figureCitation> +) +</emphasis> +</paragraph> +<paragraph id="E072368BFFC3A779FF78FC06FEB8FC6E" blockId="8.[129,778,874,1187]" pageId="8" pageNumber="9"> +20090617-CEU-ECITP1, ECIMAR campaign, between 3 and +<quantity id="27359B6EFFC3A779FF78FC39FF64FC4E" box="[129,182,943,967]" metricMagnitude="1" metricUnit="m" metricValue="3.0" pageId="8" pageNumber="9" unit="m" value="30.0">30 m</quantity> +depth, +<collectingCountry id="98DA761BFFC3A779FEFAFC39FE91FC4E" box="[259,323,943,967]" name="Spain" pageId="8" pageNumber="9">Ceuta</collectingCountry> +, +<collectingCountry id="98DA761BFFC3A779FEB5FC39FE57FC4E" box="[332,389,943,967]" name="Spain" pageId="8" pageNumber="9">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE6AFC39FDDDFC4E" box="[403,527,942,967]" degrees="35" direction="north" minutes="54" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="25" value="35.906944"> +35°54 +<emphasis id="D2B9EA99FFC3A779FE2BFC38FE0AFC4E" box="[466,472,942,967]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +25″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDE0FC39FD42FC4E" box="[537,656,942,967]" degrees="5" direction="west" minutes="18" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="00" value="-5.3"> +5°18 +<emphasis id="D2B9EA99FFC3A779FDB1FC38FD9CFC4E" box="[584,590,942,967]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +00″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF24FC59FEB4FC6E" box="[221,358,975,999]" pageId="8" pageNumber="9" value="2009-06-17">17 June 2009</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65FC78FE11FBCC" blockId="8.[129,778,874,1187]" pageId="8" pageNumber="9"> +20070701-CEU-ECITP2, ECIMAR campaign, between 3 and +<quantity id="27359B6EFFC3A779FF32FB9BFED4FBAC" box="[203,262,1037,1061]" metricMagnitude="1" metricUnit="m" metricValue="3.0" pageId="8" pageNumber="9" unit="m" value="30.0">30 m</quantity> +depth, +<collectingCountry id="98DA761BFFC3A779FEA6FB9BFE4DFBAC" box="[351,415,1037,1061]" name="Spain" pageId="8" pageNumber="9">Ceuta</collectingCountry> +, +<collectingCountry id="98DA761BFFC3A779FE57FB9BFE35FBAC" box="[430,487,1037,1061]" name="Spain" pageId="8" pageNumber="9">Spain</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE02FB9BFDA5FBAC" box="[507,631,1036,1061]" degrees="35" direction="north" minutes="52" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="38" value="35.87722"> +35°52 +<emphasis id="D2B9EA99FFC3A779FDC2FB9AFD93FBAC" box="[571,577,1036,1061]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +38″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FD7EFB9BFD2CFBAC" box="[647,766,1036,1061]" degrees="5" direction="west" minutes="18" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="20" value="-5.305556"> +5°18 +<emphasis id="D2B9EA99FFC3A779FD4FFB9AFD6EFBAC" box="[694,700,1036,1061]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +20″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FEB3FBBBFE6DFBCC" box="[330,447,1069,1093]" pageId="8" pageNumber="9" value="2007-07-01">1 July 2007</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65FBDAFE11FB2A" blockId="8.[129,778,874,1187]" pageId="8" pageNumber="9"> +20070705-CEU-ECITP3, ECIMAR campaign, between 3 and +<quantity id="27359B6EFFC3A779FF30FBFDFED0FB0A" box="[201,258,1131,1155]" metricMagnitude="1" metricUnit="m" metricValue="3.0" pageId="8" pageNumber="9" unit="m" value="30.0">30 m</quantity> +depth, +<collectingCountry id="98DA761BFFC3A779FEA1FBFDFE4AFB0A" box="[344,408,1131,1155]" name="Spain" pageId="8" pageNumber="9">Ceuta</collectingCountry> +, +<collectingCountry id="98DA761BFFC3A779FE5FFBFDFE0DFB0A" box="[422,479,1131,1155]" name="Spain" pageId="8" pageNumber="9">Spain</collectingCountry> +(( +<geoCoordinate id="85F9504CFFC3A779FE05FBFDFDABFB0A" box="[508,633,1130,1155]" degrees="35" direction="north" minutes="52" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="49" value="35.880276"> +35°52 +<emphasis id="D2B9EA99FFC3A779FDC5FBFCFD90FB0A" box="[572,578,1130,1155]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +49″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FD7EFBFDFD2CFB0A" box="[647,766,1130,1155]" degrees="5" direction="west" minutes="18" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="45" value="-5.3125"> +5°18 +<emphasis id="D2B9EA99FFC3A779FD4FFBFCFD6EFB0A" box="[694,700,1130,1155]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +45″W +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FEB3FB1CFE6DFB2A" box="[330,447,1162,1187]" pageId="8" pageNumber="9" value="2007-07-05">5 July 2007</date> +. +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC3A779FEE0FB55FEA5FAB6" pageId="8" pageNumber="9" type="multiple"> +<paragraph id="E072368BFFC3A779FEE0FB55FEA5FAB6" blockId="8.[128,778,1218,2001]" pageId="8" pageNumber="9"> +<taxonomicName id="27CD4D08FFC3A779FEE0FB55FDA3FB55" ID-CoL="85YK8" authority="Schmidt, 1862" authorityName="Schmidt" authorityYear="1862" box="[281,625,1218,1245]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="8" pageNumber="9" phylum="Porifera" rank="species" species="mollissima"> +<emphasis id="D2B9EA99FFC3A779FEE0FB55FDA3FB55" box="[281,625,1218,1245]" italics="true" pageId="8" pageNumber="9"> +Spongia mollissima +<bibRefCitation id="845C4B7AFFC3A779FE1FFB54FDA3FB55" author="Schmidt" box="[486,625,1218,1244]" firstAuthor="Schmidt" pageId="8" pageNumber="9" refId="ref18646" refString="Schmidt O. Die Spongien des adriatischen Meeres. Leipzig: Wilhelm Engelmann, 1862." type="book" year="1862">Schmidt, 1862</bibRefCitation> +</emphasis> +</taxonomicName> +20200809-IT-NTP1, +<emphasis id="D2B9EA99FFC3A779FE9AFB7FFE6BFA88" box="[355,441,1257,1281]" italics="true" pageId="8" pageNumber="9">Neuralia</emphasis> +wreck, between 30 and +<quantity id="27359B6EFFC3A779FD36FB7EFCD8FA89" box="[719,778,1256,1280]" metricMagnitude="1" metricUnit="m" metricValue="3.5" pageId="8" pageNumber="9" unit="m" value="35.0">35 m</quantity> +depth, Porto Cesareo, +<collectingCountry id="98DA761BFFC3A779FEA7FA9EFE59FAA9" box="[350,395,1288,1312]" name="Italy" pageId="8" pageNumber="9">Italy</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE6FFA9EFDC1FAA9" box="[406,531,1286,1312]" degrees="40" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="27" value="40.174168"> +40°10 +<emphasis id="D2B9EA99FFC3A779FE2FFA90FE0EFA96" box="[470,476,1286,1311]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +27″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDE3FA9EFD40FAA9" box="[538,658,1286,1312]" degrees="17" direction="east" minutes="54" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="19" value="17.905277"> +17°54 +<emphasis id="D2B9EA99FFC3A779FDAFFA90FD8EFA96" box="[598,604,1286,1311]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +19″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF24FAB1FEA6FAB6" box="[221,372,1319,1344]" pageId="8" pageNumber="9" value="2020-08-09">9 August 2020</date> +. +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC3A779FF65FAD1FB78FF6E" pageId="8" pageNumber="9" type="materials_examined"> +<paragraph id="E072368BFFC3A779FF65FAD1FEA5FA14" blockId="8.[128,778,1218,2001]" pageId="8" pageNumber="9"> +20200809-IT-NTP2, +<emphasis id="D2B9EA99FFC3A779FE83FAD1FE02FAD6" box="[378,464,1351,1375]" italics="true" pageId="8" pageNumber="9">Neuralia</emphasis> +wreck, between 30 and +<quantity id="27359B6EFFC3A779FD2AFAD0FCD8FAD7" box="[723,778,1350,1374]" metricMagnitude="1" metricUnit="m" metricValue="3.5" pageId="8" pageNumber="9" unit="m" value="35.0">35 m</quantity> +depth, Porto Cesareo, +<collectingCountry id="98DA761BFFC3A779FEA7FAF0FE59FAF7" box="[350,395,1382,1406]" name="Italy" pageId="8" pageNumber="9">Italy</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE6FFAF0FDC1FAF7" box="[406,531,1380,1406]" degrees="40" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="27" value="40.174168"> +40°10 +<emphasis id="D2B9EA99FFC3A779FE2FFAF2FE0EFAF4" box="[470,476,1380,1405]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +27″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDE3FAF0FD40FAF7" box="[538,658,1380,1406]" degrees="17" direction="east" minutes="54" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="19" value="17.905277"> +17°54 +<emphasis id="D2B9EA99FFC3A779FDAFFAF2FD8EFAF4" box="[598,604,1380,1405]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +19″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF24FA13FEA6FA14" box="[221,372,1413,1437]" pageId="8" pageNumber="9" value="2020-08-09">9 August 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65FA33FEA5FA72" blockId="8.[128,778,1218,2001]" pageId="8" pageNumber="9"> +20200809-IT-NTP3, +<emphasis id="D2B9EA99FFC3A779FE83FA33FE02FA34" box="[378,464,1445,1469]" italics="true" pageId="8" pageNumber="9">Neuralia</emphasis> +wreck, between 30 and +<quantity id="27359B6EFFC3A779FD2AFA32FCD8FA35" box="[723,778,1444,1468]" metricMagnitude="1" metricUnit="m" metricValue="3.5" pageId="8" pageNumber="9" unit="m" value="35.0">35 m</quantity> +depth, Porto Cesareo, +<collectingCountry id="98DA761BFFC3A779FEA7FA52FE59FA55" box="[350,395,1476,1500]" name="Italy" pageId="8" pageNumber="9">Italy</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE6FFA52FDC1FA55" box="[406,531,1474,1500]" degrees="40" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="27" value="40.174168"> +40°10 +<emphasis id="D2B9EA99FFC3A779FE2FFA54FE0EFA52" box="[470,476,1474,1499]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +27″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDE3FA52FD40FA55" box="[538,658,1474,1500]" degrees="17" direction="east" minutes="54" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="19" value="17.905277"> +17°54 +<emphasis id="D2B9EA99FFC3A779FDAEFA54FD8FFA52" box="[599,605,1474,1499]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +19″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF24FA75FEA6FA72" box="[221,372,1507,1531]" pageId="8" pageNumber="9" value="2020-08-09">9 August 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65F995FEAAF9D0" blockId="8.[128,778,1218,2001]" pageId="8" pageNumber="9"> +20200809-IT-NTP4, +<emphasis id="D2B9EA99FFC3A779FE83F995FE02F992" box="[378,464,1539,1563]" italics="true" pageId="8" pageNumber="9">Neuralia</emphasis> +wreck, between 30 and +<quantity id="27359B6EFFC3A779FD2AF994FCD8F993" box="[723,778,1538,1562]" metricMagnitude="1" metricUnit="m" metricValue="3.5" pageId="8" pageNumber="9" unit="m" value="35.0">35 m</quantity> +depth, Porto Cesareo, +<collectingCountry id="98DA761BFFC3A779FEA7F9B4FE59F9B3" box="[350,395,1570,1594]" name="Italy" pageId="8" pageNumber="9">Italy</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE6FF9B4FDC1F9B0" box="[406,531,1568,1594]" degrees="40" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="27" value="40.174168"> +40°10 +<emphasis id="D2B9EA99FFC3A779FE2FF9B6FE0EF9B0" box="[470,476,1568,1593]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +27″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDE3F9B4FD40F9B0" box="[538,658,1568,1594]" degrees="17" direction="east" minutes="54" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="19" value="17.905277"> +17°54 +<emphasis id="D2B9EA99FFC3A779FDAEF9B6FD8FF9B0" box="[599,605,1568,1593]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +19″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF24F9D7FEA6F9D0" box="[221,372,1601,1625]" pageId="8" pageNumber="9" value="2020-08-09">9 August 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65F9F6FEAAF93E" blockId="8.[128,778,1218,2001]" pageId="8" pageNumber="9"> +20200809-IT-NTP5, +<emphasis id="D2B9EA99FFC3A779FE83F9F6FE02F9F1" box="[378,464,1632,1656]" italics="true" pageId="8" pageNumber="9">Neuralia</emphasis> +wreck, between 30 and +<quantity id="27359B6EFFC3A779FD2AF9F6FCD8F9F1" box="[723,778,1632,1656]" metricMagnitude="1" metricUnit="m" metricValue="3.5" pageId="8" pageNumber="9" unit="m" value="35.0">35 m</quantity> +depth, Porto Cesareo, +<collectingCountry id="98DA761BFFC3A779FEA7F916FE59F911" box="[350,395,1664,1688]" name="Italy" pageId="8" pageNumber="9">Italy</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE6FF916FDC1F91E" box="[406,531,1662,1688]" degrees="40" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="27" value="40.174168"> +40°10 +<emphasis id="D2B9EA99FFC3A779FE2FF9E8FE0EF91E" box="[470,476,1662,1687]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +27″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDE3F916FD40F91E" box="[538,658,1662,1688]" degrees="17" direction="east" minutes="54" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="19" value="17.905277"> +17°54 +<emphasis id="D2B9EA99FFC3A779FDAEF9E8FD8FF91E" box="[599,605,1662,1687]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +19″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF24F909FEA6F93E" box="[221,372,1695,1719]" pageId="8" pageNumber="9" value="2020-08-09">9 August 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65F928FEAAF89C" blockId="8.[128,778,1218,2001]" pageId="8" pageNumber="9"> +20200809-IT-NTP6, +<emphasis id="D2B9EA99FFC3A779FE83F928FE02F95F" box="[378,464,1726,1750]" italics="true" pageId="8" pageNumber="9">Neuralia</emphasis> +wreck, between 30 and +<quantity id="27359B6EFFC3A779FD2AF928FCD8F95F" box="[723,778,1726,1750]" metricMagnitude="1" metricUnit="m" metricValue="3.5" pageId="8" pageNumber="9" unit="m" value="35.0">35 m</quantity> +depth, Porto Cesareo, +<collectingCountry id="98DA761BFFC3A779FEA7F948FE59F97F" box="[350,395,1758,1782]" name="Italy" pageId="8" pageNumber="9">Italy</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE6FF948FDC1F97C" box="[406,531,1756,1782]" degrees="40" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="27" value="40.174168"> +40°10 +<emphasis id="D2B9EA99FFC3A779FE2FF94AFE0EF97C" box="[470,476,1756,1781]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +27″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDE2F948FD40F97C" box="[539,658,1756,1782]" degrees="17" direction="east" minutes="54" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="19" value="17.905277"> +17°54 +<emphasis id="D2B9EA99FFC3A779FDAEF94AFD8FF97C" box="[599,605,1756,1781]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +19″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF24F96BFEA6F89C" box="[221,372,1789,1813]" pageId="8" pageNumber="9" value="2020-08-09">9 August 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65F88BFEA5F8FA" blockId="8.[128,778,1218,2001]" pageId="8" pageNumber="9"> +20200809-IT-NTP8, +<emphasis id="D2B9EA99FFC3A779FE83F88BFE02F8BC" box="[378,464,1821,1845]" italics="true" pageId="8" pageNumber="9">Neuralia</emphasis> +wreck, between 30 and +<quantity id="27359B6EFFC3A779FD2AF88AFCD8F8BD" box="[723,778,1820,1844]" metricMagnitude="1" metricUnit="m" metricValue="3.5" pageId="8" pageNumber="9" unit="m" value="35.0">35 m</quantity> +depth, Porto Cesareo, +<collectingCountry id="98DA761BFFC3A779FEA7F8AAFE59F8DD" box="[350,395,1852,1876]" name="Italy" pageId="8" pageNumber="9">Italy</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE6FF8AAFDC1F8DD" box="[406,531,1850,1876]" degrees="40" direction="north" minutes="10" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="27" value="40.174168"> +40°10 +<emphasis id="D2B9EA99FFC3A779FE2FF8ACFE0EF8DA" box="[470,476,1850,1875]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +27″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FDE3F8AAFD40F8DD" box="[538,658,1850,1876]" degrees="17" direction="east" minutes="54" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="19" value="17.905277"> +17°54 +<emphasis id="D2B9EA99FFC3A779FDAFF8ACFD8EF8DA" box="[598,604,1850,1875]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +19″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FF24F8CDFEA6F8FA" box="[221,372,1883,1908]" pageId="8" pageNumber="9" value="2020-08-09">9 August 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FF65F8EDFE20F858" blockId="8.[128,778,1218,2001]" pageId="8" pageNumber="9"> +20200810-IT-CTP1, El Camino, photophilic rocky bottoms at +<quantity id="27359B6EFFC3A779FF62F80CFF03F838" box="[155,209,1946,1970]" metricMagnitude="1" metricUnit="m" metricValue="1.7" pageId="8" pageNumber="9" unit="m" value="17.0">17 m</quantity> +depth, Porto Cesareo, +<collectingCountry id="98DA761BFFC3A779FE3BF80CFE3DF83B" box="[450,495,1946,1970]" name="Italy" pageId="8" pageNumber="9">Italy</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FE07F80CFDA9F83B" box="[510,635,1944,1970]" degrees="40" direction="north" minutes="16" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="12" value="40.27"> +40°16 +<emphasis id="D2B9EA99FFC3A779FDC4F80EFD91F838" box="[573,579,1944,1969]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +12″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FD7FF80CFD2DF83B" box="[646,767,1944,1970]" degrees="17" direction="east" minutes="51" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="15" value="17.854168"> +17°51 +<emphasis id="D2B9EA99FFC3A779FD3BF80EFD1AF838" box="[706,712,1944,1969]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +15″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FEB3F82FFE3CF858" box="[330,494,1977,2001]" pageId="8" pageNumber="9" value="2020-08-10">10 August 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FCACFF06FB78FF6E" blockId="8.[825,1475,144,231]" pageId="8" pageNumber="9"> +20200810-IT-StTP2, La Strea, photophilic rocky bottoms at +<quantity id="27359B6EFFC3A779FCADFF39FC58FF4E" box="[852,906,175,199]" metricMagnitude="1" metricUnit="m" metricValue="1.7" pageId="8" pageNumber="9" unit="m" value="17.0">17 m</quantity> +depth, Porto Cesareo, +<collectingCountry id="98DA761BFFC3A779FB83FF39FB75FF4E" box="[1146,1191,175,199]" name="Italy" pageId="8" pageNumber="9">Italy</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FB4EFF39FAE1FF4E" box="[1207,1331,174,199]" degrees="40" direction="north" minutes="14" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="54" value="40.248333"> +40°14 +<emphasis id="D2B9EA99FFC3A779FB0FFF38FB2EFF4E" box="[1270,1276,174,199]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +54″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FAC6FF39FA65FF4E" box="[1343,1463,174,199]" degrees="17" direction="east" minutes="53" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="38" value="17.893888"> +17°53 +<emphasis id="D2B9EA99FFC3A779FA82FF38FA53FF4E" box="[1403,1409,174,199]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +38″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FBFAFF59FB75FF6E" box="[1027,1191,207,231]" pageId="8" pageNumber="9" value="2020-08-10">10 August 2020</date> +. +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC3A779FC19FE81FBDAFDE6" pageId="8" pageNumber="9" type="multiple"> +<paragraph id="E072368BFFC3A779FC19FE81FAC9FEB8" blockId="8.[825,1307,279,373]" box="[992,1307,279,305]" pageId="8" pageNumber="9"> +<taxonomicName id="27CD4D08FFC3A779FC19FE81FAC9FEB8" ID-CoL="6ZBMW" authority="(Schmidt, 1862)" baseAuthorityName="Schmidt" baseAuthorityYear="1862" box="[992,1307,279,305]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="8" pageNumber="9" phylum="Porifera" rank="species" species="nitens"> +<emphasis id="D2B9EA99FFC3A779FC19FE81FAC9FEB8" box="[992,1307,279,305]" italics="true" pageId="8" pageNumber="9"> +Spongia nitens ( +<bibRefCitation id="845C4B7AFFC3A779FB7AFE81FAC1FEB8" author="Schmidt" box="[1155,1299,279,305]" firstAuthor="Schmidt" pageId="8" pageNumber="9" refId="ref18646" refString="Schmidt O. Die Spongien des adriatischen Meeres. Leipzig: Wilhelm Engelmann, 1862." type="book" year="1862">Schmidt, 1862</bibRefCitation> +) +</emphasis> +</taxonomicName> +</paragraph> +<paragraph id="E072368BFFC3A779FCC0FEABFB21FEDC" blockId="8.[825,1307,279,373]" box="[825,1267,317,341]" pageId="8" pageNumber="9"> +Po.25493, +<collectingRegion id="2209F869FFC3A779FC5CFEABFC2AFEDC" box="[933,1016,317,341]" country="Israel" name="Tel Aviv" pageId="8" pageNumber="9">Tel Aviv</collectingRegion> +, +<collectingCountry id="98DA761BFFC3A779FBF8FEABFBEAFEDC" box="[1025,1080,317,341]" name="Israel" pageId="8" pageNumber="9">Israel</collectingCountry> +( +<bibRefCitation id="845C4B7AFFC3A779FBB1FEABFB30FEDC" author="Idan" box="[1096,1250,317,341]" etAl="et al." firstAuthor="Idan" pageId="8" pageNumber="9" pagination="84 - 106" refId="ref17579" refString="Idan T, Shefer S, Feldstein T et al. Shedding light on an East-Mediterranean mesophotic sponge ground community and the regional sponge fauna. Mediterranean Marine Science 2018; 19: 84 - 106. https: // doi. org / 10.12681 / mms. 13853" type="journal article" year="2018"> +Idan +<emphasis id="D2B9EA99FFC3A779FB82FEA8FB78FEDC" box="[1147,1194,317,341]" italics="true" pageId="8" pageNumber="9">et al.</emphasis> +2018 +</bibRefCitation> +). +</paragraph> +<paragraph id="E072368BFFC3A779FCACFECAFADBFEFC" blockId="8.[825,1307,279,373]" box="[853,1289,348,373]" pageId="8" pageNumber="9"> +Po.25665 +<collectingRegion id="2209F869FFC3A779FC43FECBFBDCFEFC" box="[954,1038,349,373]" country="Israel" name="Tel Aviv" pageId="8" pageNumber="9">Tel Aviv</collectingRegion> +, +<collectingCountry id="98DA761BFFC3A779FBEEFECBFB9CFEFC" box="[1047,1102,349,373]" name="Israel" pageId="8" pageNumber="9">Israel</collectingCountry> +( +<bibRefCitation id="845C4B7AFFC3A779FBA7FECBFB2AFEFC" author="Idan" box="[1118,1272,349,373]" etAl="et al." firstAuthor="Idan" pageId="8" pageNumber="9" pagination="84 - 106" refId="ref17579" refString="Idan T, Shefer S, Feldstein T et al. Shedding light on an East-Mediterranean mesophotic sponge ground community and the regional sponge fauna. Mediterranean Marine Science 2018; 19: 84 - 106. https: // doi. org / 10.12681 / mms. 13853" type="journal article" year="2018"> +Idan +<emphasis id="D2B9EA99FFC3A779FB68FECBFB12FEFC" box="[1169,1216,349,373]" italics="true" pageId="8" pageNumber="9">et al.</emphasis> +2018 +</bibRefCitation> +). +</paragraph> +<paragraph id="E072368BFFC3A779FC27FE03FACFFE27" blockId="8.[825,1309,404,467]" box="[990,1309,404,431]" pageId="8" pageNumber="9"> +<taxonomicName id="27CD4D08FFC3A779FC27FE03FACFFE27" ID-CoL="6ZBPS" authority="Schmidt, 1862" authorityName="Schmidt" authorityYear="1862" box="[990,1309,404,431]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="8" pageNumber="9" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC3A779FC27FE03FACFFE27" box="[990,1309,404,431]" italics="true" pageId="8" pageNumber="9"> +Spongia zimocca +<bibRefCitation id="845C4B7AFFC3A779FB68FE02FACFFE27" author="Schmidt" box="[1169,1309,404,430]" firstAuthor="Schmidt" pageId="8" pageNumber="9" refId="ref18646" refString="Schmidt O. Die Spongien des adriatischen Meeres. Leipzig: Wilhelm Engelmann, 1862." type="book" year="1862">Schmidt, 1862</bibRefCitation> +</emphasis> +</taxonomicName> +</paragraph> +<paragraph id="E072368BFFC3A779FCC0FE2CFB3AFE5A" blockId="8.[825,1309,404,467]" box="[825,1256,442,467]" pageId="8" pageNumber="9"> +Po.25742 +<collectingRegion id="2209F869FFC3A779FC66FE2DFC21FE5A" box="[927,1011,443,467]" country="Israel" name="Tel Aviv" pageId="8" pageNumber="9">Tel Aviv</collectingRegion> +, +<collectingCountry id="98DA761BFFC3A779FC02FE2DFBE0FE5A" box="[1019,1074,443,467]" name="Israel" pageId="8" pageNumber="9">Israel</collectingCountry> +( +<bibRefCitation id="845C4B7AFFC3A779FBBBFE2DFB0FFE5A" author="Idan" box="[1090,1245,443,467]" etAl="et al." firstAuthor="Idan" pageId="8" pageNumber="9" pagination="84 - 106" refId="ref17579" refString="Idan T, Shefer S, Feldstein T et al. Shedding light on an East-Mediterranean mesophotic sponge ground community and the regional sponge fauna. Mediterranean Marine Science 2018; 19: 84 - 106. https: // doi. org / 10.12681 / mms. 13853" type="journal article" year="2018"> +Idan +<emphasis id="D2B9EA99FFC3A779FB8FFE2DFB76FE5A" box="[1142,1188,443,467]" italics="true" pageId="8" pageNumber="9">et al.</emphasis> +2018 +</bibRefCitation> +) +</paragraph> +<paragraph id="E072368BFFC3A779FC5FFE65FA87FD85" blockId="8.[825,1475,498,874]" box="[934,1365,498,524]" pageId="8" pageNumber="9"> +<taxonomicName id="27CD4D08FFC3A779FC5FFE65FA87FD85" authority="(Lamarck, 1814)" baseAuthorityName="Lamarck" baseAuthorityYear="1814" box="[934,1365,498,524]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="8" pageNumber="9" phylum="Porifera" rank="species" species="communis"> +<emphasis id="D2B9EA99FFC3A779FC5FFE65FA87FD85" box="[934,1365,498,524]" italics="true" pageId="8" pageNumber="9">Hippospongia communis (Lamarck, 1814)</emphasis> +</taxonomicName> +</paragraph> +<paragraph id="E072368BFFC3A779FCC0FD8EFBDAFDE6" blockId="8.[825,1475,498,874]" pageId="8" pageNumber="9"> +20200519-MRS-JTP1, Jarre, cave in the semi-dark part at +<quantity id="27359B6EFFC3A779FA74FD8EFA10FDB9" box="[1421,1474,536,560]" metricMagnitude="1" metricUnit="m" metricValue="1.5" pageId="8" pageNumber="9" unit="m" value="15.0">15 m</quantity> +depth, Marseille, +<collectingCountry id="98DA761BFFC3A779FC12FDAEFBE2FDC6" box="[1003,1072,568,591]" name="France" pageId="8" pageNumber="9">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FBC7FDAEFB68FDD9" box="[1086,1210,566,592]" degrees="43" direction="north" minutes="11" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="46" value="43.196114"> +43°11 +<emphasis id="D2B9EA99FFC3A779FB84FDA0FB51FDC6" box="[1149,1155,566,591]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +46″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FB3DFDAEFAFDFDD9" box="[1220,1327,566,592]" degrees="5" direction="east" minutes="21" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="55" value="5.365278"> +5°21 +<emphasis id="D2B9EA99FFC3A779FB0AFDA0FB2BFDC6" box="[1267,1273,566,591]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +55″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FC85FDC1FBD7FDE6" box="[892,1029,599,624]" pageId="8" pageNumber="9" value="2020-05-19">19 May 2020</date> +. +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC3A779FCACFDE0FADEFCE3" pageId="8" pageNumber="9" type="materials_examined"> +<paragraph id="E072368BFFC3A779FCACFDE0FA4AFD44" blockId="8.[825,1475,498,874]" pageId="8" pageNumber="9"> +20200522-MRS-FTP4, Fauconnière, semi-dark tunnel, between 5 and +<quantity id="27359B6EFFC3A779FC1FFD00FBF3FD24" box="[998,1057,662,686]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="8" pageNumber="9" unit="m" value="10.0">10 m</quantity> +depth, Saint-Cyr Les Lecques, +<collectingCountry id="98DA761BFFC3A779FA84FD00FA10FD24" box="[1405,1474,662,685]" name="France" pageId="8" pageNumber="9">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FCB8FD23FC6CFD44" box="[833,958,692,717]" degrees="43" direction="north" minutes="09" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="16" value="43.154446"> +43°09 +<emphasis id="D2B9EA99FFC3A779FC79FD22FC54FD44" box="[896,902,692,717]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +16″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FC3EFD23FBE0FD44" box="[967,1074,692,717]" degrees="5" direction="east" minutes="40" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="59" value="5.6830554"> +5°40 +<emphasis id="D2B9EA99FFC3A779FC0FFD22FC2EFD44" box="[1014,1020,692,717]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +59″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FAF2FD23FA46FD44" box="[1291,1428,693,718]" pageId="8" pageNumber="9" value="2020-05-22">22 May 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FCACFD42FA4AFCA2" blockId="8.[825,1475,498,874]" pageId="8" pageNumber="9"> +20200522-MRS-FTP6, Fauconnière, semi-dark tunnel, between 5 and +<quantity id="27359B6EFFC3A779FC1FFD62FBF3FC82" box="[998,1057,756,780]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="8" pageNumber="9" unit="m" value="10.0">10 m</quantity> +depth, Saint-Cyr Les Lecques, +<collectingCountry id="98DA761BFFC3A779FA84FD62FA10FC82" box="[1405,1474,756,779]" name="France" pageId="8" pageNumber="9">France</collectingCountry> +( +<geoCoordinate id="85F9504CFFC3A779FCB8FC85FC6CFCA2" box="[833,958,786,811]" degrees="43" direction="north" minutes="09" orientation="latitude" pageId="8" pageNumber="9" precision="15" seconds="16" value="43.154446"> +43°09 +<emphasis id="D2B9EA99FFC3A779FC79FC84FC54FCA2" box="[896,902,786,811]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +16″N +</geoCoordinate> +, +<geoCoordinate id="85F9504CFFC3A779FC3EFC85FBE0FCA2" box="[967,1074,786,811]" degrees="5" direction="east" minutes="40" orientation="longitude" pageId="8" pageNumber="9" precision="15" seconds="59" value="5.6830554"> +5°40 +<emphasis id="D2B9EA99FFC3A779FC0FFC84FC2EFCA2" box="[1014,1020,786,811]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis> +59″E +</geoCoordinate> +). Collector: T. Pérez, +<date id="9473104BFFC3A779FAF2FC85FA46FCA2" box="[1291,1428,787,812]" pageId="8" pageNumber="9" value="2020-05-22">22 May 2020</date> +. +</paragraph> +<paragraph id="E072368BFFC3A779FCACFCA4FADEFCE3" blockId="8.[825,1475,498,874]" pageId="8" pageNumber="9"> +20201225-TUN-Ep8, harvested by a fisherman in a seagrass meadow, +<collectingCountry id="98DA761BFFC3A779FC60FCC4FC30FCE3" box="[921,994,850,874]" name="Tunisia" pageId="8" pageNumber="9">Djerba</collectingCountry> +, +<collectingCountry id="98DA761BFFC3A779FC12FCC4FBEBFCE3" box="[1003,1081,850,874]" name="Tunisia" pageId="8" pageNumber="9">Tunisia</collectingCountry> +, +<date id="9473104BFFC3A779FBBBFCC4FADAFCE3" box="[1090,1288,850,874]" pageId="8" pageNumber="9" value="2020-12-25">25 December 2020</date> +. +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC3A779FBB2FC04FBA6FB04" pageId="8" pageNumber="9" type="etymology"> +<paragraph id="E072368BFFC3A779FBB2FC04FB63FC25" blockId="8.[825,1475,914,1165]" box="[1099,1201,914,940]" pageId="8" pageNumber="9"> +<emphasis id="D2B9EA99FFC3A779FBB2FC04FB63FC25" box="[1099,1201,914,940]" italics="true" pageId="8" pageNumber="9">Etymology</emphasis> +</paragraph> +<paragraph id="E072368BFFC3A779FCC0FC2FFBA6FB04" blockId="8.[825,1475,914,1165]" pageId="8" pageNumber="9"> +The species name ‘maitasuna’ comes from ‘maitasun’, which means ‘love’ in Euskara, the +<collectingRegion id="2209F869FFC3A779FBA8FC4EFB2AFC79" box="[1105,1272,984,1008]" country="Spain" name="Pais Vasco" pageId="8" pageNumber="9">Basque Country</collectingRegion> +language. Collected first in a site of the French +<collectingRegion id="2209F869FFC3A779FBB7FC6EFB1FFB86" box="[1102,1229,1016,1040]" country="Spain" name="Pais Vasco" pageId="8" pageNumber="9">Pays Basque</collectingRegion> +, the new sponge is thus dedicated to all beloved members of our families, with special thoughts to those recently departed, especially to Marguerite Grenier, century-old grandmother, who passed away while this manuscript was in preparation. +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC3A779FBB6FB25FA7AF9A3" pageId="8" pageNumber="9" type="diagnosis"> +<paragraph id="E072368BFFC3A779FBB6FB25FB7EFB45" blockId="8.[825,1475,1203,1578]" box="[1103,1196,1203,1228]" pageId="8" pageNumber="9"> +<emphasis id="D2B9EA99FFC3A779FBB6FB25FB7EFB45" box="[1103,1196,1203,1228]" italics="true" pageId="8" pageNumber="9">Diagnosis</emphasis> +</paragraph> +<paragraph id="E072368BFFC3A779FCC0FB4FFA7AF9A3" blockId="8.[825,1475,1203,1578]" pageId="8" pageNumber="9">Massive sponge with unarmoured and non-lacunose surface. Surface covered with evenly disposed conules, sometimes with traces of epibiosis or foreign bodies. Soft and compressible consistency. Skeleton made of primary, secondary, and tertiary fibres. Primary fibres are anastomosed and denser at the surface, piercing the surface, forming conules, and presenting an open space variable in size between each anastomosed primary fibre. Primary fibres harbouring inclusions such as foreign spicules and/or a pith. Secondary fibres containing a pith inclusion and forming a clearly and regular honeycomb network. Tertiary fibres present both at the surface and within the sponge body.</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC3A77BFBB1F9C6FED3FDA9" lastPageId="10" lastPageNumber="11" pageId="8" pageNumber="9" type="description"> +<paragraph id="E072368BFFC3A779FBB1F9C6FB66F9E0" blockId="8.[825,1474,1616,1991]" box="[1096,1204,1616,1641]" pageId="8" pageNumber="9"> +<emphasis id="D2B9EA99FFC3A779FBB1F9C6FB66F9E0" box="[1096,1204,1616,1641]" italics="true" pageId="8" pageNumber="9">Description</emphasis> +</paragraph> +<paragraph id="E072368BFFC3A779FCC0F9E0FB47F84E" blockId="8.[825,1474,1616,1991]" pageId="8" pageNumber="9"> +The new sponge is massive and irregular, sometimes slightly branched, with small lobes or protuberances ( +<figureCitation id="78F62A0EFFC3A779FAE1F903FA51F924" box="[1304,1411,1685,1709]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." pageId="8" pageNumber="9">Fig. 2A–D</figureCitation> +). The sponge can measure ≤ +<quantity id="27359B6EFFC3A779FBDBF922FBB6F945" box="[1058,1124,1716,1740]" metricMagnitude="-1" metricUnit="m" metricValue="2.5" pageId="8" pageNumber="9" unit="cm" value="25.0">25 cm</quantity> +in its largest diameter. Its surface is white, beige, light to dark grey, whereas the internal tissue is usually tawny yellow. No change in colour was detected after fixation in 95% ethanol. The sponge surface is sometimes covered by dirty foreign bodies or epibiotic organisms. The surface is very conulose. Oscules can be grouped or not and are often found on the top of each lobe or protuberance. They are bordered by a thin and rather translucid membrane ( +<figureCitation id="78F62A0EFFC3A779FB6FF806FAD2F821" box="[1174,1280,1936,1960]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." pageId="8" pageNumber="9">Fig. 2A–D</figureCitation> +). The sponge consistency is rather soft and tearable. +</paragraph> +<paragraph id="E072368BFFC2A778FF74FF06FD8CFEAC" blockId="9.[113,763,144,450]" pageId="9" pageNumber="10"> +The ectosomal skeleton presents a very thin epidermal skin, which has a star-like appearance, is easily detachable, and is made of collagen fibrils, from 60 to 350 µm in thickness ( +<figureCitation id="78F62A0EFFC2A778FD2DFF59FF5DFE8F" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="9" pageNumber="10">Fig. 3A</figureCitation> +) Scattered foreign spicules, debris, or remains of associated invertebrates, such as cirripeds, can be observed. +</paragraph> +<paragraph id="E072368BFFC2A77BFF74FEBBFED3FDA9" blockId="9.[113,763,144,450]" lastBlockId="10.[128,778,144,544]" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10"> +The choanosomal skeleton presents primary fibres, which are irregular and sinuous, sometimes difficult to observe and to distinguish because they appear anastomosed with secondary fibres, and in some places, they can also divide and join again ( +<figureCitation id="78F62A0EFFC2A778FF85FE3CFF36FE4B" box="[124,228,426,450]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="9" pageNumber="10">Fig. 3B–D</figureCitation> +). They measure 30–70 µm in diameter and are cored with inclusions of foreign spicules and/or debris of various abundance. They also include a black or translucid pith. These primary fibres with a pith can be observed near the surface into conule formation and/or across the sponge body ( +<figureCitation id="78F62A0EFFC2A778FAC1FF78FA71FE8F" box="[1336,1443,238,262]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="9" pageNumber="10">Fig. 3A, C</figureCitation> +). However, when foreign debris and spicules are widely present, the pith in the primary fibres can be difficult to observe. Near the surface, close to the conule formations, the network of primary fibres is denser in comparison to the rest of the sponge body. The space between two primary fibres, thus between conules, seems to be empty, with very few tertiary fibres and foreign spicules and debris. Perforated plates are inconsistently present close to the primary fibres ( +<figureCitation id="78F62A0EFFC1A77BFEB1FF39FE46FF4E" box="[328,404,175,199]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="10" pageNumber="11">Fig. 3D</figureCitation> +). The secondary fibres form a dense network, clearly showing a honeycomb shape of variable mesh size ( +<figureCitation id="78F62A0EFFC1A77BFF4FFF78FF29FE8F" box="[182,251,238,262]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="10" pageNumber="11">Fig. 3B</figureCitation> +), from 70 µm × 120 µm to 300 µm × 500 µm. These fibres measure 20–50 µm in diameter and they can contain an irregular pith, black or translucid ( +<figureCitation id="78F62A0EFFC1A77BFDFBFEBBFD9DFECC" box="[514,591,301,325]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="10" pageNumber="11">Fig. 3E</figureCitation> +). Tertiary fibres, ~10 µm in diameter, are found throughout the sponge body ( +<figureCitation id="78F62A0EFFC1A77BFF72FEFDFF23FE0A" box="[139,241,363,387]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="10" pageNumber="11">Fig. 3F, G</figureCitation> +). Sometimes they form a thinner network that also has a clear honeycomb shape with variable mesh size (50–130 µm in diameter). In some places, these fibres appear to be linked to the thicker secondary fibres ( +<figureCitation id="78F62A0EFFC1A77BFE21FE5FFDFAFE68" box="[472,552,457,481]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="10" pageNumber="11">Fig. 3H</figureCitation> +), but in some places they look rather disconnected from the rest of the skeletal architecture. +</paragraph> +</subSubSection> +<caption id="B4B26603FFC2A778FF88F8D8FA71F832" pageId="9" pageNumber="10" startId="9.[113,178,1870,1894]" targetBox="[119,1456,530,1838]" targetPageId="9" targetType="figure"> +<paragraph id="E072368BFFC2A778FF88F8D8FA71F832" blockId="9.[113,1458,1870,1979]" pageId="9" pageNumber="10"> +<emphasis id="D2B9EA99FFC2A778FF88F8D8FF1BF8EE" bold="true" box="[113,201,1870,1895]" pageId="9" pageNumber="10">Figure 3.</emphasis> +Sections of the spongin skeleton of +<taxonomicName id="27CD4D08FFC2A778FDE2F8D9FD05F8EE" box="[539,727,1871,1895]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="9" pageNumber="10" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC2A778FDE2F8D9FD05F8EE" bold="true" box="[539,727,1871,1895]" italics="true" pageId="9" pageNumber="10">Spongia maitasuna</emphasis> +</taxonomicName> +. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows). B, secondary fibres organized in a regular honeycomb network. C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres. E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm. G, tertiary fibres. H, detail of a junction between secondary and tertiary fibres (arrow). +</paragraph> +</caption> +<subSubSection id="A8D76500FFC1A77BFE72FDA9FCD6FD14" pageId="10" pageNumber="11" type="distribution"> +<paragraph id="E072368BFFC1A77BFE72FDA9FE2CFDD0" blockId="10.[129,778,575,670]" box="[395,510,575,601]" pageId="10" pageNumber="11"> +<emphasis id="D2B9EA99FFC1A77BFE72FDA9FE2CFDD0" box="[395,510,575,601]" italics="true" pageId="10" pageNumber="11">Distribution</emphasis> +</paragraph> +<paragraph id="E072368BFFC1A77BFF78FDF0FCD6FD14" blockId="10.[129,778,575,670]" pageId="10" pageNumber="11"> +South European Atlantic Shelf ( +<bibRefCitation id="845C4B7AFFC1A77BFE21FDF0FD79FDF7" author="Spalding" box="[472,683,614,638]" etAl="et al." firstAuthor="Spalding" pageId="10" pageNumber="11" pagination="573 - 83" refId="ref18695" refString="Spalding MD, Fox HE, Allen GR et al. Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. BioScience 2007; 57: 573 - 83. https: // doi. org / 10.1641 / b 570707" type="journal article" year="2007"> +Spalding +<emphasis id="D2B9EA99FFC1A77BFDC3FDF1FDBCFDF7" box="[570,622,614,638]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2007 +</bibRefCitation> +), Bay of Biscay, French +<collectingRegion id="2209F869FFC1A77BFEE1FD10FE44FD14" box="[280,406,646,670]" country="Spain" name="Pais Vasco" pageId="10" pageNumber="11">Pays Basque</collectingRegion> +, and Spanish +<collectingRegion id="2209F869FFC1A77BFDD9FD13FD57FD14" box="[544,645,645,669]" country="Spain" name="Cantabria" pageId="10" pageNumber="11">Cantabria</collectingRegion> +(this work). +</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC1A77BFE59FD2BFE69FBBC" pageId="10" pageNumber="11" type="biology_ecology"> +<paragraph id="E072368BFFC1A77BFE59FD2BFE38FD5E" blockId="10.[129,778,701,1077]" box="[416,490,701,727]" pageId="10" pageNumber="11"> +<emphasis id="D2B9EA99FFC1A77BFE59FD2BFE38FD5E" box="[416,490,701,727]" italics="true" pageId="10" pageNumber="11">Ecology</emphasis> +</paragraph> +<paragraph id="E072368BFFC1A77BFF78FD72FDCBFCF0" blockId="10.[129,778,701,1077]" pageId="10" pageNumber="11"> +<taxonomicName id="27CD4D08FFC1A77BFF78FD72FEEEFD72" box="[129,316,740,763]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC1A77BFF78FD72FEEEFD72" box="[129,316,740,763]" italics="true" pageId="10" pageNumber="11">Spongia maitasuna</emphasis> +</taxonomicName> +is found on rocky substrate in shallow water cliffs, boulders, or semi-dark and dark cavities, habitats usually with high wave energy. This sponge presents a patchy distribution and lives in syntopy with +<taxonomicName id="27CD4D08FFC1A77BFE3EFCD4FDFBFCD0" authorityName="Schulze" authorityYear="1879" box="[455,553,833,857]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="lamella"> +<emphasis id="D2B9EA99FFC1A77BFE3EFCD4FDFBFCD0" box="[455,553,833,857]" italics="true" pageId="10" pageNumber="11">S. lamella</emphasis> +</taxonomicName> +. In some cases, both +<taxonomicName id="27CD4D08FFC1A77BFF78FCF7FF21FCF0" authorityName="Gray" authorityYear="1867" box="[129,243,865,889]" class="Demospongiae" family="Spongiidae" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="family">Spongiidae</taxonomicName> +present contact interactions. +</paragraph> +<paragraph id="E072368BFFC1A77BFF65FC16FE69FBBC" blockId="10.[129,778,701,1077]" pageId="10" pageNumber="11">Some specimens may have epibiotic organisms, mostly encrusting sponges and zoanthids. No sign of predation was observed. In the specimens collected between late June and late July, various stages of embryogenesis were observed, including early fertilized eggs, various morula stages, and pre-larvae. No spermatic cysts were observed.</paragraph> +</subSubSection> +<subSubSection id="A8D76500FFC1A77BFE9EFBC3FC59FDB6" pageId="10" pageNumber="11" type="discussion"> +<paragraph id="E072368BFFC1A77BFE9EFBC3FDF1FBE7" blockId="10.[129,778,1108,1985]" box="[359,547,1108,1134]" pageId="10" pageNumber="11"> +<emphasis id="D2B9EA99FFC1A77BFE9EFBC3FDF1FBE7" box="[359,547,1108,1134]" italics="true" pageId="10" pageNumber="11">Taxonomic remarks</emphasis> +</paragraph> +<paragraph id="E072368BFFC1A77BFF78FBEDFF08FA45" blockId="10.[129,778,1108,1985]" pageId="10" pageNumber="11"> +The new species is assigned to +<taxonomicName id="27CD4D08FFC1A77BFE3CFBEDFDC1FB1B" authorityName="Linnaeus" authorityYear="1759" box="[453,531,1147,1170]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC1A77BFE3CFBEDFDC1FB1B" box="[453,531,1147,1170]" italics="true" pageId="10" pageNumber="11">Spongia</emphasis> +</taxonomicName> +because its body is not lacunose, as in +<taxonomicName id="27CD4D08FFC1A77BFEEEFB0DFE4CFB3B" authorityName="Schulze" authorityYear="1879" box="[279,414,1179,1202]" class="Demospongiae" family="Spongiidae" genus="Hippospongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC1A77BFEEEFB0DFE4CFB3B" box="[279,414,1179,1202]" italics="true" pageId="10" pageNumber="11">Hippospongia</emphasis> +</taxonomicName> +or +<emphasis id="D2B9EA99FFC1A77BFE47FB0CFDCAFB3B" box="[446,536,1178,1202]" italics="true" pageId="10" pageNumber="11">Hyatella</emphasis> +, and it does not present an armoured ectosomal skeleton, as in +<taxonomicName id="27CD4D08FFC1A77BFDFCFB2FFD42FB58" authorityName="Carter" authorityYear="1883" box="[517,656,1209,1233]" class="Demospongiae" family="Spongiidae" genus="Coscinoderma" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC1A77BFDFCFB2FFD42FB58" box="[517,656,1209,1233]" italics="true" pageId="10" pageNumber="11">Coscinoderma</emphasis> +</taxonomicName> +or +<taxonomicName id="27CD4D08FFC1A77BFD56FB2FFCD1FB58" authorityName="Lendenfeld" authorityYear="1888" box="[687,771,1209,1233]" class="Demospongiae" family="Spongiidae" genus="Leiosella" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC1A77BFD56FB2FFCD1FB58" box="[687,771,1209,1233]" italics="true" pageId="10" pageNumber="11">Leiosella</emphasis> +</taxonomicName> +. Unlike +<taxonomicName id="27CD4D08FFC1A77BFF32FB4FFE9EFB78" authorityName="Thompson, Murphy, Bergquist & Evans" authorityYear="1987" box="[203,332,1241,1265]" class="Demospongiae" family="Spongiidae" genus="Rhopaloeides" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC1A77BFF32FB4FFE9EFB78" box="[203,332,1241,1265]" italics="true" pageId="10" pageNumber="11">Rhopaloeides</emphasis> +</taxonomicName> +, which presents long and simple fascicles of primary fibres, the primary fibres of the new species are simple and sometimes anastomosed. In +<taxonomicName id="27CD4D08FFC1A77BFE33FA8EFDCAFAA6" authorityName="Linnaeus" authorityYear="1759" box="[458,536,1304,1327]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC1A77BFE33FA8EFDCAFAA6" box="[458,536,1304,1327]" italics="true" pageId="10" pageNumber="11">Spongia</emphasis> +</taxonomicName> +, the new species should belong to the subgenus +<taxonomicName id="27CD4D08FFC1A77BFE7BFAA1FE02FAC7" box="[386,464,1335,1358]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="subGenus" subGenus="Spongia"> +<emphasis id="D2B9EA99FFC1A77BFE7BFAA1FE02FAC7" box="[386,464,1335,1358]" italics="true" pageId="10" pageNumber="11">Spongia</emphasis> +</taxonomicName> +, because it presents a simple primary skeleton with regular and polygonal meshes. However, considering the present status of the genus, we prefer not to use this infrageneric, lower rank classification (see the Discussion section). +</paragraph> +<paragraph id="E072368BFFC1A77BFF65FA42FC59FDB6" blockId="10.[129,778,1108,1985]" lastBlockId="10.[825,1475,144,575]" pageId="10" pageNumber="11"> +<taxonomicName id="27CD4D08FFC1A77BFF65FA42FE88FA62" ID-CoL="BW7B2" box="[156,346,1492,1515]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC1A77BFF65FA42FE88FA62" box="[156,346,1492,1515]" italics="true" pageId="10" pageNumber="11">Spongia maitasuna</emphasis> +</taxonomicName> +shares with the +<typeStatus id="3F768829FFC1A77BFE03FA42FDFBFA65" box="[506,553,1492,1516]" pageId="10" pageNumber="11">type</typeStatus> +species of +<taxonomicName id="27CD4D08FFC1A77BFD6AFA42FD36FA62" ID-CoL="84TSR" authorityName="Linnaeus" authorityYear="1759" box="[659,740,1492,1515]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="genus"> +<emphasis id="D2B9EA99FFC1A77BFD6AFA42FD36FA62" box="[659,740,1492,1515]" italics="true" pageId="10" pageNumber="11">Spongia</emphasis> +</taxonomicName> +the very thin collagenic epidermal skin and the fact that primary fibres are never forming fascicles, in other words they are never fused. In +<taxonomicName id="27CD4D08FFC1A77BFF18F9A4FE85F9C0" ID-CoL="6ZBNB" authorityName="Linnaeus" authorityYear="1759" box="[225,343,1585,1609]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC1A77BFF18F9A4FE85F9C0" box="[225,343,1585,1609]" italics="true" pageId="10" pageNumber="11">S. officinalis</emphasis> +</taxonomicName> +, the primary fibres are thicker than in the new species, ranging from 50 to 100 µm. The network of secondary fibres is much denser in +<taxonomicName id="27CD4D08FFC1A77BFE30F9E6FDECF901" ID-CoL="6ZBNB" authorityName="Linnaeus" authorityYear="1759" box="[457,574,1648,1672]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC1A77BFE30F9E6FDECF901" box="[457,574,1648,1672]" italics="true" pageId="10" pageNumber="11">S. officinalis</emphasis> +</taxonomicName> +, and the meshes do not have the typical honeycomb shape of the new species. This structure of the secondary skeleton is found in +<taxonomicName id="27CD4D08FFC1A77BFD8AF939FCDBF94F" ID-CoL="6ZBNB" authorityName="Linnaeus" authorityYear="1759" box="[627,777,1710,1734]" class="Demospongiae" family="Spongiidae" genus="Spongia" isUncertain="true" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="officinalis"> +<emphasis id="D2B9EA99FFC1A77BFD8AF939FD54F94F" box="[627,646,1711,1734]" italics="true" pageId="10" pageNumber="11">S.</emphasis> +cf. +<emphasis id="D2B9EA99FFC1A77BFD52F938FCDBF94F" box="[683,777,1710,1734]" italics="true" pageId="10" pageNumber="11">officinalis</emphasis> +</taxonomicName> +from +<collectingCountry id="98DA761BFFC1A77BFF47F958FED2F96F" box="[190,256,1742,1766]" name="Spain" pageId="10" pageNumber="11">Ceuta</collectingCountry> +, Strait of +<collectingCountry id="98DA761BFFC1A77BFE89F958FE00F96F" box="[368,466,1742,1766]" name="Gibraltar" pageId="10" pageNumber="11">Gibraltar</collectingCountry> +. The main difference in this case lies in the primary fibres, which are slightly larger than in the new species (70–110 µm), and in the overall growth form ( +<figureCitation id="78F62A0EFFC1A77BFF75F8BAFF11F8CD" box="[140,195,1836,1860]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." pageId="10" pageNumber="11">Fig.2</figureCitation> +). +<taxonomicName id="27CD4D08FFC1A77BFF2EF8BAFE46F8CA" ID-CoL="BW7B2" box="[215,404,1836,1859]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="maitasuna"> +<emphasis id="D2B9EA99FFC1A77BFF2EF8BAFE46F8CA" box="[215,404,1836,1859]" italics="true" pageId="10" pageNumber="11">Spongia maitasuna</emphasis> +</taxonomicName> +has a morphology similar to +<taxonomicName id="27CD4D08FFC1A77BFD4AF8BAFCD6F8CA" ID-CoL="6ZBMW" baseAuthorityName="Schmidt" baseAuthorityYear="1862" box="[691,772,1836,1859]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="nitens"> +<emphasis id="D2B9EA99FFC1A77BFD4AF8BAFCD6F8CA" box="[691,772,1836,1859]" italics="true" pageId="10" pageNumber="11">S. nitens</emphasis> +</taxonomicName> +, and both species display a pith within their primary fibres. But +<taxonomicName id="27CD4D08FFC1A77BFF78F8FDFF04F80B" ID-CoL="6ZBMW" baseAuthorityName="Schmidt" baseAuthorityYear="1862" box="[129,214,1899,1922]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="nitens"> +<emphasis id="D2B9EA99FFC1A77BFF78F8FDFF04F80B" box="[129,214,1899,1922]" italics="true" pageId="10" pageNumber="11">S. nitens</emphasis> +</taxonomicName> +has a denser secondary skeleton, and the secondary fibres are devoid of pith. Moreover, tertiary fibres were present in the two studied specimens of +<taxonomicName id="27CD4D08FFC1A77BFDEAF83FFDBBF848" ID-CoL="6ZBMW" baseAuthorityName="Schmidt" baseAuthorityYear="1862" box="[531,617,1961,1985]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="nitens"> +<emphasis id="D2B9EA99FFC1A77BFDEAF83FFDBBF848" box="[531,617,1961,1985]" italics="true" pageId="10" pageNumber="11">S. nitens</emphasis> +</taxonomicName> +, but according to the literature they can be absent ( +<bibRefCitation id="845C4B7AFFC1A77BFB41FF06FAEDFF21" author="Vacelet" box="[1208,1343,144,168]" firstAuthor="Vacelet" pageId="10" pageNumber="11" pagination="39 - 101" refId="ref18958" refString="Vacelet J. Repartition generale des eponges et systematique des eponges cornees de la region de Marseille et de quelques stations Mediterraneennes. Recueil des travaux de la Station Marine d'Endoume 1959; 16: 39 - 101." type="journal article" year="1959">Vacelet 1959</bibRefCitation> +, +<bibRefCitation id="845C4B7AFFC1A77BFAB5FF06FC5EFF4E" author="Manconi" etAl="et al." firstAuthor="Manconi" pageId="10" pageNumber="11" pagination="1 - 68" refId="ref17923" refString="Manconi R, Cadeddu B, Ledda F et al. An overview of the Mediterranean cave-dwelling horny sponges (Porifera, Demospongiae). ZooKeos 2013; 281: 1 - 68. https: // doi. org / 10.3897 / zookeys. 281.4171" type="journal article" year="2013"> +Manconi +<emphasis id="D2B9EA99FFC1A77BFA48FF06FC80FF4E" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2013 +</bibRefCitation> +). They were recorded in all specimens of the new species. Finally, the new species shares with the studied specimen of +<taxonomicName id="27CD4D08FFC1A77BFCAEFF78FC16FE8C" ID-CoL="6ZBPS" authorityName="Schmidt" authorityYear="1862" box="[855,964,238,261]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC1A77BFCAEFF78FC16FE8C" box="[855,964,238,261]" italics="true" pageId="10" pageNumber="11">S. zimocca</emphasis> +</taxonomicName> +the anastomosed primary fibres and the thickness of the secondary fibres, although they can be reported thicker (50–80 µm) in the literature ( +<bibRefCitation id="845C4B7AFFC1A77BFB34FEBAFA84FECD" author="Vacelet" box="[1229,1366,300,324]" firstAuthor="Vacelet" pageId="10" pageNumber="11" pagination="39 - 101" refId="ref18958" refString="Vacelet J. Repartition generale des eponges et systematique des eponges cornees de la region de Marseille et de quelques stations Mediterraneennes. Recueil des travaux de la Station Marine d'Endoume 1959; 16: 39 - 101." type="journal article" year="1959">Vacelet 1959</bibRefCitation> +, +<bibRefCitation id="845C4B7AFFC1A77BFA9AFEBAFC79FEED" author="Manconi" etAl="et al." firstAuthor="Manconi" pageId="10" pageNumber="11" pagination="1 - 68" refId="ref17923" refString="Manconi R, Cadeddu B, Ledda F et al. An overview of the Mediterranean cave-dwelling horny sponges (Porifera, Demospongiae). ZooKeos 2013; 281: 1 - 68. https: // doi. org / 10.3897 / zookeys. 281.4171" type="journal article" year="2013"> +Manconi +<emphasis id="D2B9EA99FFC1A77BFCC0FEDAFCBFFEED" box="[825,877,332,356]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2013 +</bibRefCitation> +). In the studied specimen of +<taxonomicName id="27CD4D08FFC1A77BFB0DFEDAFAB3FEEA" ID-CoL="6ZBPS" authorityName="Schmidt" authorityYear="1862" box="[1268,1377,332,355]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC1A77BFB0DFEDAFAB3FEEA" box="[1268,1377,332,355]" italics="true" pageId="10" pageNumber="11">S. zimocca</emphasis> +</taxonomicName> +, the primary fibres are much thicker, 50–100 µm (100–200 µm in the literature) than in the new species. These fibres also present a pith, although this trait has not been reported previously ( +<bibRefCitation id="845C4B7AFFC1A77BFCBCFE5FFC1CFE68" author="Vacelet" box="[837,974,457,481]" firstAuthor="Vacelet" pageId="10" pageNumber="11" pagination="39 - 101" refId="ref18958" refString="Vacelet J. Repartition generale des eponges et systematique des eponges cornees de la region de Marseille et de quelques stations Mediterraneennes. Recueil des travaux de la Station Marine d'Endoume 1959; 16: 39 - 101." type="journal article" year="1959">Vacelet 1959</bibRefCitation> +, +<bibRefCitation id="845C4B7AFFC1A77BFC25FE5FFAC9FE68" author="Castritsi-Catharios" box="[988,1307,457,481]" etAl="et al." firstAuthor="Castritsi-Catharios" pageId="10" pageNumber="11" pagination="41 - 62" refId="ref16583" refString="Castritsi-Catharios J, van Soest RWM, Kefalas E et al. Revised description of a poorly known Mediterranean Dictyoceratid bath sponge, Spongia (Spongia) zimocca (Schmidt, 1862) (Porifera: Demospongiae: Dictyoceratida). Zootaxa 2011; 2812: 41 - 62. https: // doi. org / 10.11646 / zootaxa. 2812.1.4" type="journal article" year="2011"> +Castritsi-Catharios +<emphasis id="D2B9EA99FFC1A77BFB54FE5CFB0DFE68" box="[1197,1247,457,481]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2011 +</bibRefCitation> +, +<bibRefCitation id="845C4B7AFFC1A77BFAD3FE5FFCBCFD88" author="Manconi" etAl="et al." firstAuthor="Manconi" pageId="10" pageNumber="11" pagination="1 - 68" refId="ref17923" refString="Manconi R, Cadeddu B, Ledda F et al. An overview of the Mediterranean cave-dwelling horny sponges (Porifera, Demospongiae). ZooKeos 2013; 281: 1 - 68. https: // doi. org / 10.3897 / zookeys. 281.4171" type="journal article" year="2013"> +Manconi +<emphasis id="D2B9EA99FFC1A77BFA69FE5CFA11FE68" box="[1424,1475,457,481]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2013 +</bibRefCitation> +), and rare foreign spicules. Finally, the spongin network is overall irregular and denser in +<taxonomicName id="27CD4D08FFC1A77BFB58FD9EFADCFDA9" ID-CoL="6ZBPS" authorityName="Schmidt" authorityYear="1862" box="[1185,1294,520,544]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="zimocca"> +<emphasis id="D2B9EA99FFC1A77BFB58FD9EFADCFDA9" box="[1185,1294,520,544]" italics="true" pageId="10" pageNumber="11">S. zimocca</emphasis> +</taxonomicName> +than in the new species. +</paragraph> +</subSubSection> +</treatment> +</document> \ No newline at end of file diff --git a/data/6A/3D/D1/6A3DD143066DFF905590FDB365926476.xml b/data/6A/3D/D1/6A3DD143066DFF905590FDB365926476.xml new file mode 100644 index 00000000000..ed9f8144ff5 --- /dev/null +++ b/data/6A/3D/D1/6A3DD143066DFF905590FDB365926476.xml @@ -0,0 +1,144 @@ +<document id="B07CEC2D95D8154653F600C643FF4D0E" ID-DOI="10.1093/zoolinnean/zlae126" ID-ISSN="0024-4082" IM.bibliography_approvedBy="jonas" IM.illustrations_approvedBy="jonas" IM.materialsCitations_approvedBy="jonas" IM.metadata_approvedBy="jonas" IM.taxonomicNames_approvedBy="jonas" IM.treatments_approvedBy="jonas" checkinTime="1732850508520" checkinUser="plazi" docAuthor="Rytel, Adam, Surmik, Dawid, Szczygielski, Tomasz, Spiekman, Stephan N. F., Kamp, Thomas van de, Zuber, Marcus & Scheyer, Torsten M." docDate="2024" docId="6A3DD143066DFF905590FDB365926476" docLanguage="en" docName="zlae126.pdf" docOrigin="Zoological Journal of the Linnean Society 202 (3)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlae126" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.9:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="9" docTitle="Protanystropheus antiquus" docType="treatment" docVersion="1" lastPageNumber="6" masterDocId="9604A93B0668FF955502FF9B60366677" masterDocTitle="Unique internal anatomy of vertebrae as a key factor for neck elongation in Triassic archosauromorphs" masterLastPageNumber="21" masterPageNumber="1" pageNumber="6" updateTime="1733770130157" updateUser="jonas"> +<mods:mods id="CCB63C0710B53916FA4C0982AC9767FD" xmlns:mods="http://www.loc.gov/mods/v3"> +<mods:titleInfo id="BD58ECD91DFBA28B3A81636F82A07781"> +<mods:title id="439AE117AFD1B18D2D7A92AB096D8E36">Unique internal anatomy of vertebrae as a key factor for neck elongation in Triassic archosauromorphs</mods:title> +</mods:titleInfo> +<mods:name id="D0632969D92EAEE88C87FD9290735025" type="personal"> +<mods:role id="C88830E51CF45A5ACB675FABD5C46489"> +<mods:roleTerm id="11B3C428567B0EC445F439F9CC4601E4">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="01EB37F50081F6F11021ED7FA0470404">Rytel, Adam</mods:namePart> +<mods:affiliation id="5ED5263984B4E9A47E0742EDCE3E9BBC">Institute of Paleobiology, Polish Academy of Sciences, Twarda 51 / 55, 00 - 818 Warsaw, Poland & Department of Paleontology, University of Zurich, Karl Schmid-Strasse 4, 8006 Zurich, Switzerland</mods:affiliation> +<mods:nameIdentifier id="D52372958B53E7E1A8FEC0C200DF1065" type="email">adam.rytel@twarda.pan.pl</mods:nameIdentifier> +</mods:name> +<mods:name id="33DF89D6F7ACE4A40E2B575B34D73B87" type="personal"> +<mods:role id="C26C19F15B587172E1EC98BD51C25F21"> +<mods:roleTerm id="3B79D2732F8A277CDB01E8100145B783">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="5241178210D129460778CB1A0E57B1AF">Surmik, Dawid</mods:namePart> +<mods:affiliation id="2AD1F3C58F24E501B49CCE28001EFFA3">Institute of Earth Sciences, Faculty of Natural Sciences, University of Silesia in Katowice, Bedzinska 60, 41 - 200 Sosnowiec, Poland</mods:affiliation> +<mods:nameIdentifier id="D92EFFEDE116540892843942F4F1DC1D" type="email">adam.rytel@twarda.pan.pl</mods:nameIdentifier> +</mods:name> +<mods:name id="CE8F68ACEE7B503602F52B95FB60AFA1" type="personal"> +<mods:role id="1717B63FB3FC3C0EF2B6905C485AF0E3"> +<mods:roleTerm id="6DAF7B02F64226D2B390C6D4B2C78AD4">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="76F4988C98C6CA120C94C4EACE354AF9">Szczygielski, Tomasz</mods:namePart> +<mods:affiliation id="E75C5D53B1783A561D59F2BCD775DACA">Institute of Paleobiology, Polish Academy of Sciences, Twarda 51 / 55, 00 - 818 Warsaw, Poland</mods:affiliation> +</mods:name> +<mods:name id="F40056F39FAEE8DEC05DFB1467328F37" type="personal"> +<mods:role id="30B5576C0223A0A0CFB0537A87F5E9A6"> +<mods:roleTerm id="E95BD60C64FE7D060E8F4C8D5E18FDF3">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="178E86DCE60D750881AA11AB098E1148">Spiekman, Stephan N. F.</mods:namePart> +<mods:affiliation id="5124BDB2DAC4F9A7AF221D28CA9FADE7">Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany</mods:affiliation> +</mods:name> +<mods:name id="6F1FF451B8A352F8968C4C9CBBE4182B" type="personal"> +<mods:role id="380295331E12DB22307CEF03A526895A"> +<mods:roleTerm id="0BACFEDCDEDFF315EC70DCB183F9D517">Author</mods:roleTerm> +</mods:role> +<mods:namePart id="6A268DE664DA90E1F312E7636831268A">Kamp, Thomas van de</mods:namePart> +<mods:affiliation id="32195E158A4F93DEEA28B6E92B7A528B">Institute for Photon Science and Synchrotron Radiation (IPS), Karlsruhe Institute of Technology (KIT), Hermann-von-Helmholtz-Platz 1, 76344 Eggenstein-Leopoldshafen, Germany & Laboratory for Applications of Synchrotron Radiation (LAS), Karlsruhe Institute of Technology (KIT), Kaiserstrasse 12, 76131 Karlsruhe, Germany</mods:affiliation> +</mods:name> +<mods:name id="9EBB5D2651549F1E96C075295CC8D460" type="personal"> +<mods:role 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+<paragraph id="E22B6055066DFF905590FDB361D76416" blockId="5.[146,733,552,609]" pageId="5" pageNumber="6"> +<emphasis id="D0E0BC47066DFF905590FDB361D76416" bold="true" pageId="5" pageNumber="6"> +Bone histology and internal structure of vertebrae of +<taxonomicName id="25941BD6066DFF9057BFFDB361D76416" authority="(Huene, 1905)" baseAuthorityName="Huene" baseAuthorityYear="1905" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="antiquus"> +‘ +<emphasis id="D0E0BC47066DFF9057C1FDB262E46434" bold="true" box="[707,722,553,579]" italics="true" pageId="5" pageNumber="6">P</emphasis> +’. +<emphasis id="D0E0BC47066DFF905488FDDC61D76416" bold="true" box="[394,481,583,609]" italics="true" pageId="5" pageNumber="6">antiquus</emphasis> +</taxonomicName> +</emphasis> +</paragraph> +</subSubSection> +<subSubSection id="AA8E33DE066DFF905573FDF565926476" pageId="5" pageNumber="6" type="description"> +<paragraph id="E22B6055066DFF905573FDF565926476" blockId="5.[112,764,622,1962]" lastBlockId="5.[810,1460,144,513]" pageId="5" pageNumber="6"> +The CT examination of vertebrae was performed to nondestructively investigate the internal structure of each specimen and to assess the thickness of compact and cancellous bone in the different regions of the vertebrae. Within the anteroposteriorly terminal portions of all scanned ‘ +<emphasis id="D0E0BC47066DFF9054D4FD7761D26574" box="[470,484,748,771]" italics="true" pageId="5" pageNumber="6">P</emphasis> +’. +<taxonomicName id="25941BD6066DFF9054F4FD77627C6574" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[502,586,748,771]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="antiquus"> +<emphasis id="D0E0BC47066DFF9054F4FD77627C6574" box="[502,586,748,771]" italics="true" pageId="5" pageNumber="6">antiquus</emphasis> +</taxonomicName> +centra, the core was filled with dense cancellous bone (identified in CT images so of uncertain tissue composition) and the cortex was relatively thin ( +<figureCitation id="7AAF7CD0066DFF9055F3FCD2617A6516" box="[241,332,841,865]" captionStart-0="Figure 2" captionStart-1="Figure 3" captionStart-2="Figure 4" captionStartId-0="3.[113,178,975,999]" captionStartId-1="6.[129,194,1669,1693]" captionStartId-2="7.[113,178,1841,1865]" captionTargetBox-0="[307,1266,157,946]" captionTargetBox-1="[305,1297,144,1642]" captionTargetBox-2="[175,1398,149,1813]" captionTargetId-0="figure-579@3.[305,1268,144,948]" captionTargetId-1="figure-103@6.[305,1297,144,1642]" captionTargetId-2="figure-6@7.[175,1398,149,1813]" captionTargetPageId-0="3" captionTargetPageId-1="6" captionTargetPageId-2="7" captionText-0="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." captionText-1="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." captionText-2="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Figs 2–4</figureCitation> +), whereas in the middle portion, where the internal cavity was revealed, the cortical bone was distinctly thicker. The transverse sections revealed the changes of the cortical bone thickness and the shape of internal cavities, with the mid-centrum being roughly cylindrical. The cortex was thinner near the anteroposteriorly terminal portions of the vertebrae and its thickness gradually increased up to approximately four times the original thickness (about 10% of bone diameter), near the midpoint ( +<figureCitation id="7AAF7CD0066DFF90540AFBDF6176622B" box="[264,320,1092,1116]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">Fig. 3</figureCitation> +). The density of the cancellous bone of the anteroposteriorly terminal portions of the centrum decreased medially, with larger cavities being present closer to the midpoint. The transition to the empty internal cavity was relatively sharp. The anterior and posterior ends of the internal cavity were roughly hemispherical and surrounded circumferentially by marginalized trabeculae ( +<figureCitation id="7AAF7CD0066DFF905478FA9B6183636F" box="[378,437,1280,1304]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">Fig. 3</figureCitation> +). Long intermittent trabeculae composed of parallel-fibred bone moderately remodelled with endosteal secondary lamellar bone ( +<figureCitation id="7AAF7CD0066DFF9054DFFAA462106320" box="[477,550,1343,1367]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4D</figureCitation> +) crossed the internal cavity in varied directions, mainly diagonally. In some vertebrae they divided the internal cavity into several smaller terminus-like pockets ( +<figureCitation id="7AAF7CD0066DFF9055D1FA06611763C2" box="[211,289,1437,1461]" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 2B</figureCitation> +, +<figureCitation id="7AAF7CD0066DFF90542DFA06610A63C2" box="[303,316,1437,1461]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">3</figureCitation> +). The neural canal was separated ventrally from the internal cavity by a horizontal, plate-like lamella consisting of endosteal lamellar bone ( +<figureCitation id="7AAF7CD0066DFF9054D0FA40622A6384" box="[466,540,1499,1523]" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 2B</figureCitation> +, +<figureCitation id="7AAF7CD0066DFF905725FA4062026384" box="[551,564,1499,1523]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">3</figureCitation> +, +<figureCitation id="7AAF7CD0066DFF90573CFA40624A6384" box="[574,636,1499,1523]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">4A–C</figureCitation> +), which was generally thinner than the internal trabeculae composed of the same tissue. This separation was not continuous, the neural canal floor was perforated by large openings in some specimens ( +<figureCitation id="7AAF7CD0066DFF9057D2F9A260B86006" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 2B</figureCitation> +and +<figureCitation id="7AAF7CD0066DFF9055C1F9C261326006" box="[195,260,1625,1649]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">3C–G</figureCitation> +) and seemed to disappear completely in others. For example, in MGUWr 3889s at least two large openings were present in the lamella separating the neural canal from the internal cavity. In MGUWr 3902 and GPIH 5194c there appeared to be no bony separation between the neural canal and the internal cavity for most of the length of the centrum. Because no fragments of broken bone were present anywhere inside of the neural canal or the internal cavity of the centrum, and this condition was present in several specimens approximately in the same area, it appears not to be an artifact of preservation. The cortex was predominately composed of parallel-fibred matrix ( +<figureCitation id="7AAF7CD0066DFF9057B2F80962DF61DD" box="[688,745,1938,1962]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4</figureCitation> +), with locally (especially in the dorsolateral part) highly organized arrays of mineralized collagen fibres. No rest lines were present. The vascularization was moderate, and its pattern was radial, which was especially evident in the ventral part of the vertebrae ( +<figureCitation id="7AAF7CD0066DFF905637FE9663E56752" box="[821,979,269,293]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4A–C, H–I</figureCitation> +). Inside the neural arch, mostly dorsally to the neural canal (inside of the base of the neural spine) and in some specimens laterally to the neural canal there was a region of occurrence of secondary trabecular bone ( +<figureCitation id="7AAF7CD0066DFF9051C3FEF0653767F4" box="[1217,1281,363,387]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4J</figureCitation> +). Both the cavity and the neural canal were clear cut and lined with a thin layer of endosteal lamellar bone ( +<figureCitation id="7AAF7CD0066DFF905137FE3164A067B5" box="[1077,1174,426,450]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4E–J</figureCitation> +). Sharpey’s fibres could be seen extending throughout the cortex, especially in the dorsolateral and ventrolateral regions of the centrum ( +<figureCitation id="7AAF7CD0066DFF905017FE7265A56476" box="[1301,1427,489,513]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4C, G–I</figureCitation> +). +</paragraph> +</subSubSection> +</treatment> +</document> \ No newline at end of file diff --git a/data/6A/3D/D1/6A3DD143066DFF9F5659FDBB64A963F0.xml b/data/6A/3D/D1/6A3DD143066DFF9F5659FDBB64A963F0.xml new file mode 100644 index 00000000000..02c3a52ceb3 --- /dev/null +++ b/data/6A/3D/D1/6A3DD143066DFF9F5659FDBB64A963F0.xml @@ -0,0 +1,346 @@ +<document id="F432FB799579E2FB39BD36A8C6FE82B9" ID-DOI="10.1093/zoolinnean/zlae126" ID-ISSN="0024-4082" IM.bibliography_approvedBy="jonas" IM.illustrations_approvedBy="jonas" IM.materialsCitations_approvedBy="jonas" IM.metadata_approvedBy="jonas" IM.taxonomicNames_approvedBy="jonas" IM.treatments_approvedBy="jonas" checkinTime="1732850508520" checkinUser="plazi" docAuthor="Rytel, Adam, Surmik, Dawid, Szczygielski, Tomasz, Spiekman, Stephan N. 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type="nomenclature"> +<paragraph id="E22B6055066DFF905659FDBB64F9642D" blockId="5.[859,1410,544,602]" pageId="5" pageNumber="6"> +<emphasis id="D0E0BC47066DFF905659FDBB64F9642D" bold="true" pageId="5" pageNumber="6"> +Bone histology and internal structure of vertebrae of +<taxonomicName id="25941BD6066DFF90510CFDA464F9642D" authority="Meyer, 1852" authorityName="Meyer" authorityYear="1852" box="[1038,1231,575,602]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Protorosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus"> +<emphasis id="D0E0BC47066DFF90510CFDA464A9642D" bold="true" box="[1038,1183,575,602]" italics="true" pageId="5" pageNumber="6">Tanystropheus</emphasis> +spp. +</taxonomicName> +</emphasis> +</paragraph> +</subSubSection> +<subSubSection id="AA8E33DE066DFF9F56DEFDF164A963F0" lastPageId="10" lastPageNumber="11" pageId="5" pageNumber="6" type="description"> +<paragraph id="E22B6055066DFF9056DEFDF1653464F4" blockId="5.[809,1463,617,1965]" box="[988,1282,617,644]" pageId="5" pageNumber="6"> +<emphasis id="D0E0BC47066DFF9056DEFDF1653464F4" box="[988,1282,617,644]" italics="true" pageId="5" pageNumber="6">Anatomy of the middle cervicals</emphasis> +</paragraph> +<paragraph id="E22B6055066DFF9D5628FD0B62E063A9" blockId="5.[809,1463,617,1965]" lastBlockId="8.[128,779,1322,1753]" lastPageId="8" lastPageNumber="9" pageId="5" pageNumber="6"> +Within all of the studied vertebrae, a large cavity could be identified, surrounded by relatively thick cortex composed of alternating layers of well-organized parallel-fibred to lamellar bone ( +<figureCitation id="7AAF7CD0066DFF905637FD7563BD6571" box="[821,907,750,774]" captionStart-0="Figure 5" captionStart-1="Figure 6" captionStart-2="Figure 7" captionStart-3="Figure 8" captionStartId-0="8.[130,195,1138,1162]" captionStartId-1="9.[113,178,1191,1215]" captionStartId-2="10.[129,194,695,719]" captionStartId-3="11.[113,178,1841,1865]" captionTargetBox-0="[289,1313,144,1110]" captionTargetBox-1="[146,1426,144,1163]" captionTargetBox-2="[289,1313,144,667]" captionTargetBox-3="[224,1348,150,1812]" captionTargetId-0="figure-299@8.[289,1313,144,1110]" captionTargetId-1="figure-396@9.[146,1426,144,1163]" captionTargetId-2="figure-733@10.[289,1313,144,667]" captionTargetId-3="figure-6@11.[223,1349,149,1813]" captionTargetPageId-0="8" captionTargetPageId-1="9" captionTargetPageId-2="10" captionTargetPageId-3="11" captionText-0="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." captionText-1="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." captionText-2="Figure 7. Internal cavity volume in the cervical vertebrae of ‘P’.antiquus MGUWr 3895s (A) and Tanystropheus sp. from Miedary ZPAL V. 36/181 (B). Anatomical abbreviation:ic—internal cavity. Images obtained using ‘xray’ shader in MeshLab v.2020.12 (Cignoni et al.2008)." captionText-3="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Figs 5–8</figureCitation> +). In the anterior and posterior terminal portions of the cervicals, discs of secondary cancellous bone of endosteal origin formed the articular regions of the centrum ( +<figureCitation id="7AAF7CD0066DFF9051F1FCB6656D6533" box="[1267,1371,812,837]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Fig. 5B–D</figureCitation> +). Above them, the openings of the neural canal were present. In some of the middle cervicals the neural canal floor extended only slightly from the openings towards the middle of the vertebra, forming a shelf that partially separated the internal cavity from the neural canal ( +<figureCitation id="7AAF7CD0066DFF905673FC5263ED6596" box="[881,987,969,993]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 5B–E</figureCitation> +, +<figureCitation id="7AAF7CD0066DFF9056EAFC5264146596" box="[1000,1058,969,993]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="5" pageNumber="6">6C, F</figureCitation> +), although in some other of these vertebrae the neural canal transitioned smoothly into the internal cavity (compare +<figureCitation id="7AAF7CD0066DFF9056DBFB9364156257" box="[985,1059,1032,1056]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="5" pageNumber="6">Fig. 6B</figureCitation> +to +<figureCitation id="7AAF7CD0066DFF90514BFB9364A26257" box="[1097,1172,1032,1056]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Fig. 5B</figureCitation> +; +<figureCitation id="7AAF7CD0066DFF9051A1FB9364DA6268" box="[1187,1260,1032,1056]" captionStart="Figure 7" captionStartId="10.[129,194,695,719]" captionTargetBox="[289,1313,144,667]" captionTargetId="figure-733@10.[289,1313,144,667]" captionTargetPageId="10" captionText="Figure 7. Internal cavity volume in the cervical vertebrae of ‘P’.antiquus MGUWr 3895s (A) and Tanystropheus sp. from Miedary ZPAL V. 36/181 (B). Anatomical abbreviation:ic—internal cavity. Images obtained using ‘xray’ shader in MeshLab v.2020.12 (Cignoni et al.2008)." pageId="5" pageNumber="6">Fig. 7B</figureCitation> +). In the transverse cross section, the inner surface of the bone wall of the internal cavity was generally parallel to the circumferential outline of the vertebra ( +<figureCitation id="7AAF7CD0066DFF9056B4FBFD642D6209" box="[950,1051,1126,1150]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8A–E</figureCitation> +). In most of the studied specimens, the dorsal part of the internal cavity seemed to preserve a semicircular shape in the transverse cross section, whereas tissue layout of its ventrolateral portion was usually much less uniform, with an irregular resorption front cutting through multiple annuli ( +<figureCitation id="7AAF7CD0066DFF905637FA986381636D" box="[821,951,1283,1307]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8A, C, E</figureCitation> +). This was especially evident in larger specimens, whereas in the smallest cervical, the regions of resorption were much less developed, and the outline of the internal cavity in the transverse cross section was more regularly oval ( +<figureCitation id="7AAF7CD0066DFF905029FAFA6542630E" box="[1323,1396,1377,1401]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8B</figureCitation> +). The neural spine extended over two deep fossae, located on both of its anteroposteriorly terminal portions. They were triangular in longitudinal cross section and ventrally delimited by thin bone trabeculae, that also constituted the roof of the neural canal. The posterior cavity was larger, forming the ‘postzygapophyseal trough’ ( +<bibRefCitation id="86051DA4066DFF905687F98664396042" author="Rieppel O" box="[901,1039,1565,1589]" pageId="5" pageNumber="6" pagination="271 - 87" refId="ref17778" refString="Rieppel O. A new species of Tanystropheus (Reptilia: Protorosauria) from the Middle Triassic of Makhtesh Ramon, Israel. Neues Jahrbuch fur Geologie und Palaontologie - Abhandlungen 2001; 221: 271 - 87. https: // doi. org / 10.1127 / njgpa / 221 / 2001 / 271" type="journal article" year="2001">Rieppel 2001</bibRefCitation> +). Contrary to what was observed for ‘ +<emphasis id="D0E0BC47066DFF90509FF986659D6043" box="[1437,1451,1565,1588]" italics="true" pageId="5" pageNumber="6">P</emphasis> +’. +<taxonomicName id="25941BD6066DFF905628F9A663486023" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[810,894,1597,1620]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="antiquus"> +<emphasis id="D0E0BC47066DFF905628F9A663486023" box="[810,894,1597,1620]" italics="true" pageId="5" pageNumber="6">antiquus</emphasis> +</taxonomicName> +, in none of the studied sections of the +<taxonomicName id="25941BD6066DFF905025F9A763626004" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="D0E0BC47066DFF905025F9A765856023" box="[1319,1459,1596,1620]" italics="true" pageId="5" pageNumber="6">Tanystropheus</emphasis> +spp. +</taxonomicName> +middle cervicals did the internal cavity contain bony trabeculae extending internally from the surface of the bone wall. Some semi-symmetrical, slanted trabeculae of endosteal origin were present only incidentally in the anteroposteriorly terminal sections of the vertebrae (see +<bibRefCitation id="86051DA4066DFF905161F94264E36087" author="Broili F" box="[1123,1237,1752,1777]" pageId="5" pageNumber="6" pagination="51 - 62" refId="ref15553" refString="Broili F. Beobachtungen an Tanystropheus conspicuus H. v. Meyer. Neues Jahrbuch fur Mineralogie Geologie und Palaontologie 1915; 2: 51 - 62." type="journal article" year="1915">Broili 1915</bibRefCitation> +: plate 3, fig. 4c). The roof of the neural canal was generally straight and parallel to the longitudinal axis of the vertebra. The ventral delimitation of the neural canal was seemingly absent for most of its length within a vertebra. Intriguingly, in one of the transverse polished sections from the anterior part of U-MO BT 738.00 ( +<figureCitation id="7AAF7CD0066DFF90500AF8EE656761FA" box="[1288,1361,1909,1933]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8F</figureCitation> +), minute bony projections appeared symmetrically on each lateral side of the internal cavity. The colour of the limestone infill of the vertebra changed drastically at the same horizontal level, at which these projections were situated. Similar characteristics could be noted for the transverse thin section of ZPAL V. 36/166 ( +<figureCitation id="7AAF7CD0066EFF9357E1F8E460AD61C0" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="6" pageNumber="7">Fig. 8E</figureCitation> +), in which corresponding projections composed of periosteal bone were also present in the dorsolateral regions of the internal cavity. Between them, there was a sharp transition in sediment fraction, with the more coarse-grained infill occupying the ventral-more space. Presence of this feature possibly shows the differences in taphonomical microenvironments of the ventral and dorsal portions of the internal cavity, which may have been caused by the original existence of a no longer preserved barrier between them. Near the anteroposterior midpoint of the middle cervicals, two symmetrically placed foramina were located on the ventral side of the centrum. They entered the internal cavity as straight canals ( +<figureCitation id="7AAF7CD00660FF9D5439FA5C61AD63A9" box="[315,411,1479,1503]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="8" pageNumber="9">Fig. 8B, E</figureCitation> +), perforating the dense cortex. +</paragraph> +<caption id="B6EB30DD066EFF935583F91E618B6086" pageId="6" pageNumber="7" startId="6.[129,194,1669,1693]" targetBox="[305,1297,144,1642]" targetPageId="6" targetType="figure"> +<paragraph id="E22B6055066EFF935583F91E618B6086" blockId="6.[129,1449,1669,1777]" pageId="6" pageNumber="7"> +<emphasis id="D0E0BC47066EFF935583F91E60EE60EA" bold="true" box="[129,216,1669,1693]" pageId="6" pageNumber="7">Figure 3.</emphasis> +CT transverse cross sections (from anterior to posterior) of ‘ +<emphasis id="D0E0BC47066EFF935616F91D631760EA" box="[788,801,1670,1693]" italics="true" pageId="6" pageNumber="7">P</emphasis> +’. +<taxonomicName id="25941BD6066EFF93562FF91D634D60EA" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[813,891,1670,1693]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="antiquus"> +<emphasis id="D0E0BC47066EFF93562FF91D634D60EA" box="[813,891,1670,1693]" italics="true" pageId="6" pageNumber="7">antiquus</emphasis> +</taxonomicName> +cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum. +</paragraph> +</caption> +<caption id="B6EB30DD066FFF9D5573F8AA634161CD" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8" startId="7.[113,178,1841,1865]" subCaptionStartIDs="8.[809,941,1899,1923]" subCaptionStarts="abbr" targetBox="[175,1398,149,1813]" targetPageId="7" targetType="figure"> +<paragraph id="E22B6055066FFF9D5573F8AA634161CD" blockId="7.[113,1457,1841,1978]" lastBlockId="8.[129,1454,1843,1979]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8"> +<emphasis id="D0E0BC47066FFF925573F8AA60FF613E" bold="true" box="[113,201,1841,1865]" pageId="7" pageNumber="8">Figure 4.</emphasis> +Thin sections of ‘ +<emphasis id="D0E0BC47066FFF92546CF8A9614D613E" box="[366,379,1842,1865]" italics="true" pageId="7" pageNumber="8">P</emphasis> +’. +<taxonomicName id="25941BD6066FFF925485F8A961E3613E" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[391,469,1842,1865]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="antiquus"> +<emphasis id="D0E0BC47066FFF925485F8A961E3613E" box="[391,469,1842,1865]" italics="true" pageId="7" pageNumber="8">antiquus</emphasis> +</taxonomicName> +cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light. Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone. Scale bars for panels A–C equal +<quantity id="256CCDB00660FF9D577BF838628661CD" box="[633,688,1955,1979]" metricMagnitude="-3" metricUnit="m" metricValue="5.0" pageId="8" pageNumber="9" unit="mm" value="5.0">5 mm</quantity> +and +<quantity id="256CCDB00660FF9D57DFF838631361CD" box="[733,805,1955,1979]" metricMagnitude="-4" metricUnit="m" metricValue="5.0" pageId="8" pageNumber="9" unit="mm" value="0.5">0.5 mm</quantity> +for D–J. +</paragraph> +</caption> +<caption id="B6EB30DD0660FF9D5580FBE962FF628D" pageId="8" pageNumber="9" startId="8.[130,195,1138,1162]" targetBox="[289,1313,144,1110]" targetPageId="8" targetType="figure"> +<paragraph id="E22B60550660FF9D5580FBE962FF628D" blockId="8.[129,1462,1138,1274]" pageId="8" pageNumber="9"> +<emphasis id="D0E0BC470660FF9D5580FBE960EC62FD" bold="true" box="[130,218,1138,1162]" pageId="8" pageNumber="9">Figure 5.</emphasis> +Middle cervicals of +<taxonomicName id="25941BD60660FF9D5494FBE9620162FD" box="[406,567,1138,1162]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="D0E0BC470660FF9D5494FBE9622E62FD" box="[406,536,1138,1162]" italics="true" pageId="8" pageNumber="9">Tanystropheus</emphasis> +sp. +</taxonomicName> +from Miedary: A, B, D—ZPAL V. 36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view. B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis. +</paragraph> +</caption> +<paragraph id="E22B60550660FF9C559EFA7D620660C7" blockId="8.[128,779,1322,1753]" lastBlockId="9.[113,763,1375,1712]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9"> +While in ‘ +<emphasis id="D0E0BC470660FF9D5409FA7C612F6389" box="[267,281,1511,1534]" italics="true" pageId="8" pageNumber="9">P</emphasis> +’. +<taxonomicName id="25941BD60660FF9D542EFA7C61B66389" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[300,384,1511,1534]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="antiquus"> +<emphasis id="D0E0BC470660FF9D542EFA7C61B66389" box="[300,384,1511,1534]" italics="true" pageId="8" pageNumber="9">antiquus</emphasis> +</taxonomicName> +the neural spine was easily identifiable, it was much less recognizable in the middle cervicals of +<taxonomicName id="25941BD60660FF9D5583F9BE6109604B" box="[129,319,1573,1597]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="D0E0BC470660FF9D5583F9BE613B604A" box="[129,269,1573,1597]" italics="true" pageId="8" pageNumber="9">Tanystropheus</emphasis> +spp. +</taxonomicName> +It was impossible to demarcate the border between the neural arch and the vertebral centrum—they were completely fused, with no preserved remnants of the suture between them, even in the smallest specimens, and little to no trabeculae within the neural arch. Continuous layers of dense parallel-fibred tissue sheathed the internal cavity. In the middle section of the larger middle cervicals of +<taxonomicName id="25941BD60660FF9D51DFFAB165AC6335" box="[1245,1434,1322,1346]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="D0E0BC470660FF9D51DFFAB1655F6335" box="[1245,1385,1322,1346]" italics="true" pageId="8" pageNumber="9">Tanystropheus</emphasis> +spp. +</taxonomicName> +the outermost of these zones formed complete rings, whereas the ones located innermost were discontinued only due to resorption of the region ventral into the internal cavity, as well as the presence of the neural spine ( +<figureCitation id="7AAF7CD00660FF9D5176FA3C64D663B7" box="[1140,1248,1447,1472]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="8" pageNumber="9">Fig. 8A, C</figureCitation> +, +<figureCitation id="7AAF7CD00660FF9D51ECFA3C650E63B7" box="[1262,1336,1447,1472]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." pageId="8" pageNumber="9">Fig. 9B</figureCitation> +). In some of the studied transverse thin sections, including the smallest specimens, the spinous process was developed only as a thin, vertically oriented plate. It was located directly above the internal cavity and emerged from the parallel-fibred tissue of the bone walls in the dorsalmost portion of the vertebra, projecting slightly from the smooth outline of the transverse cross section. In the smallest specimen from Miedary, ZPAL V. 36/181, the neural spine was never overlain by parallel-fibred bone ( +<figureCitation id="7AAF7CD00660FF9D504AF93965A460CD" box="[1352,1426,1698,1722]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="8" pageNumber="9">Fig. 8B</figureCitation> +). In all of the larger specimens from Miedary, the anteroposteriorly middle part of the spinous process was obscured by thick layers of tissue that formed the walls of the internal cavity. Compared to what was noted for ‘ +<emphasis id="D0E0BC470661FF9C5464FA05614263C2" box="[358,372,1438,1461]" italics="true" pageId="9" pageNumber="10">P</emphasis> +’. +<taxonomicName id="25941BD60661FF9C5485FA0561ED63C2" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[391,475,1438,1461]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="antiquus"> +<emphasis id="D0E0BC470661FF9C5485FA0561ED63C2" box="[391,475,1438,1461]" italics="true" pageId="9" pageNumber="10">antiquus</emphasis> +</taxonomicName> +, the dorsolateral regions of the +<taxonomicName id="25941BD60661FF9C559AFA26616463A3" box="[152,338,1469,1493]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="D0E0BC470661FF9C559AFA26611263A2" box="[152,292,1469,1493]" italics="true" pageId="9" pageNumber="10">Tanystropheus</emphasis> +spp. +</taxonomicName> +cervicals were thickened (see +<figureCitation id="7AAF7CD00661FF9C5787FA26628B63A2" box="[645,701,1469,1493]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." pageId="9" pageNumber="10">Fig. 9</figureCitation> +). The neural spine was, thus, not relatively reduced in the latter, but rather embedded within the walls of the tubularly structured vertebra. As a result, the neural canal was located close to the middle of the height of the vertebra, as in ‘ +<emphasis id="D0E0BC470661FF9C571DF9A0621B6025" box="[543,557,1595,1618]" italics="true" pageId="9" pageNumber="10">P</emphasis> +’. +<taxonomicName id="25941BD60661FF9C5738F9A062B86025" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[570,654,1595,1618]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="antiquus"> +<emphasis id="D0E0BC470661FF9C5738F9A062B86025" box="[570,654,1595,1618]" italics="true" pageId="9" pageNumber="10">antiquus</emphasis> +</taxonomicName> +. However, in +<taxonomicName id="25941BD60661FF9C558FF9C2617D6006" box="[141,331,1625,1649]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="D0E0BC470661FF9C558FF9C2612F6006" box="[141,281,1625,1649]" italics="true" pageId="9" pageNumber="10">Tanystropheus</emphasis> +spp. +</taxonomicName> +there was more tissue dorsolaterally (see +<figureCitation id="7AAF7CD00661FF9C5573F9E2609A60E6" box="[113,172,1657,1681]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." pageId="9" pageNumber="10">Fig. 9</figureCitation> +), due to the neural spine being hypertrophied along the middle section of its anteroposterior length. +</paragraph> +<caption id="B6EB30DD0661FF9C5573FB3C64746358" pageId="9" pageNumber="10" startId="9.[113,178,1191,1215]" targetBox="[146,1426,144,1163]" targetPageId="9" targetType="figure"> +<paragraph id="E22B60550661FF9C5573FB3C64746358" blockId="9.[113,1454,1191,1327]" pageId="9" pageNumber="10"> +<emphasis id="D0E0BC470661FF9C5573FB3C60FF62C8" bold="true" box="[113,201,1191,1215]" pageId="9" pageNumber="10">Figure 6.</emphasis> +Vertebrae of +<taxonomicName id="25941BD60661FF9C5444FB3C61D062C9" box="[326,486,1191,1215]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="D0E0BC470661FF9C5444FB3C61FE62C8" box="[326,456,1191,1215]" italics="true" pageId="9" pageNumber="10">Tanystropheus</emphasis> +sp. +</taxonomicName> +from Miedary: A, E, I—ZPAL V. 36/106, 11th cervical; B, F, J—ZPAL V. 36/108, 12th cervical; C, G, K—ZPAL V. 36/110, 13th (last) cervical; D—ZPAL V. 36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections. White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels. +</paragraph> +</caption> +<paragraph id="E22B60550661FF9C5418F94B6267609D" blockId="9.[113,765,1744,1963]" box="[282,593,1744,1770]" pageId="9" pageNumber="10"> +<emphasis id="D0E0BC470661FF9C5418F94B6267609D" box="[282,593,1744,1770]" italics="true" pageId="9" pageNumber="10">Anatomy of the posterior cervicals</emphasis> +</paragraph> +<paragraph id="E22B60550661FF9F5573F96C62506310" blockId="9.[113,765,1744,1963]" lastBlockId="10.[128,779,795,1728]" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10"> +In +<taxonomicName id="25941BD60661FF9C558CF96D610B6179" box="[142,317,1782,1806]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="D0E0BC470661FF9C558CF96D612C6179" box="[142,282,1782,1806]" italics="true" pageId="9" pageNumber="10">Tanystropheus</emphasis> +sp. +</taxonomicName> +from Miedary, the external morphology of the cervicals changed near the posterior subregion of the neck. The 10th and 11th vertebrae were still elongate, but, contrary to the middle cervicals, exhibited a well exposed, relatively tall and continuous neural spine ( +<figureCitation id="7AAF7CD00661FF9C54B5F8E861C361FC" box="[439,501,1907,1931]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="9" pageNumber="10">Fig. 6</figureCitation> +). The two last cervicals were relatively much shorter ( +<figureCitation id="7AAF7CD00661FF9C54AEF808622361DC" box="[428,533,1939,1963]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="9" pageNumber="10">Fig. 6B, C</figureCitation> +), and more similar in proportions to the dorsals ( +<bibRefCitation id="86051DA40661FF9C513DFAC464CE6300" author="Rieppel O & Jiang DY & Fraser NC" box="[1087,1272,1375,1399]" pageId="9" pageNumber="10" pagination="1082 - 9" refId="ref17833" refString="Rieppel O, Jiang DY, Fraser NC et al. Tanystropheus cf. T. longobardicus from the early Late Triassic of Guizhou Province, Southwestern China. Journal of Vertebrate Paleontology 2010; 30: 1082 - 9. https: // doi. org / 10.1080 / 02724634.2010.483548" type="journal article" year="2010"> +Rieppel +<emphasis id="D0E0BC470661FF9C5191FAC464F76300" box="[1171,1217,1375,1399]" italics="true" pageId="9" pageNumber="10">et al.</emphasis> +2010 +</bibRefCitation> +, +<bibRefCitation id="86051DA40661FF9C5000FAC465946300" author="Rytel A & Bohmer C & Spiekman SNF" box="[1282,1442,1375,1399]" pageId="9" pageNumber="10" refId="ref17884" refString="Rytel A, Bohmer C, Spiekman SNF et al. Extreme neck elongation evolved despite strong developmental constraints in bizarre Triassic reptiles - implications for neck modularity in archosaurs. Royal Society Open Science 2024; 11: 240233. https: // doi. org / 10.1098 / rsos. 240233" type="journal volume" year="2024"> +Rytel +<emphasis id="D0E0BC470661FF9C503FFAC4655D6300" box="[1341,1387,1375,1399]" italics="true" pageId="9" pageNumber="10">et al.</emphasis> +2024 +</bibRefCitation> +). This transition was paired with the modifications in the internal structure of these elements. The relative volume occupied by the trabeculae within the centra increased caudally, especially along the neck-torso transition. In the middle cervicals the centrum was nearly devoid of trabecular bone, but this changed within the more posterior vertebrae, with the last two cervicals and the dorsals exhibiting sparse trabeculae occupying the space between the articular discs. The 10th and 11th vertebrae were generally similar in their anatomy to the middle cervicals, with their internal cavities being surrounded by a dense cortex, forming a tube-like structure ( +<figureCitation id="7AAF7CD00661FF9C56F8F92C640B60B8" box="[1018,1085,1719,1743]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="9" pageNumber="10">Fig. 6I</figureCitation> +). However, in the succeeding vertebrae the internal cavity was greatly reduced and exhibited much thicker walls, composed of two regions: a relatively thin cortex and porous, cancellous bone encompassed by it. Especially in the middle portion of the centrum of the last two cervicals, there were several large chambers, similar to those present in the anteroposteriorly terminal portions of the more anteriorly located vertebrae ( +<figureCitation id="7AAF7CD00661FF9C56F0F808647761DC" box="[1010,1089,1939,1963]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="9" pageNumber="10">Fig. 8D</figureCitation> +). Some of them connected to the neural canal through its floor ( +<figureCitation id="7AAF7CD00662FF9F54C3FC80626A6544" box="[449,604,795,819]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6E, F, H, I</figureCitation> +). The transverse cross sections of the posteriormost cervicals were less oval than in the preceding vertebrae, with the centrum being much more ventrally expanded, forming a distinct keel ( +<figureCitation id="7AAF7CD00662FF9F5762FCE262F165E6" box="[608,711,889,913]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6J, K</figureCitation> +). The diameter of the neural canal was relatively constant along the length of each of the two last vertebrae. In the longitudinal cross section, it had a sagging appearance, with its middle portion being located more ventrally. In the transverse cross section, it was oval, with its dorsolateral regions being roughly semicircular and symmetrical in outline ( +<figureCitation id="7AAF7CD00662FF9F54A3FBAE61CB623A" box="[417,509,1077,1101]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6J, K</figureCitation> +). A similar shape could be noted in the anteroposteriorly terminal sections of the rest of the postaxial cervicals, in which the neural canal floor was still present (e.g. +<figureCitation id="7AAF7CD00662FF9F55E1FB08611862DC" box="[227,302,1171,1195]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="10" pageNumber="11">Fig. 8D</figureCitation> +), and also in the dorsal vertebrae ( +<figureCitation id="7AAF7CD00662FF9F5794FB0862EB62DC" box="[662,733,1171,1195]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6L</figureCitation> +). In each of the two posteriormost neck vertebrae of +<taxonomicName id="25941BD60662FF9F577FFB2860AA629E" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="D0E0BC470662FF9F577FFB28633F62BC" box="[637,777,1203,1227]" italics="true" pageId="10" pageNumber="11">Tanystropheus</emphasis> +sp. +</taxonomicName> +from Miedary, up to two small subcentral foramina could be traced, with some specimens exhibiting only one subcentral foramen or none at all. In the studied posteriormost cervical ZPAL V. 36/110, a straight, non-branching canal connected the one present foramen with the neural canal ( +<figureCitation id="7AAF7CD00662FF9F5712FAD462636310" box="[528,597,1359,1383]" captionStart="Figure 10" captionStartId="13.[113,178,1140,1164]" captionTargetBox="[306,1266,144,1112]" captionTargetId="figure-458@13.[306,1266,144,1112]" captionTargetPageId="13" captionText="Figure 10. Longitudinal cross section of ZPAL V.36/110, 13th (last) cervical vertebra of Tanystropheus sp.from Miedary, showing the course of the arterial canal (locations of its end sections are indicated with arrows). The anterior end of the vertebra points towards the left. Anatomical abbreviations: nc—neural canal; ns—neural spine." pageId="10" pageNumber="11">Fig. 10</figureCitation> +). +</paragraph> +<caption id="B6EB30DD0662FF9F5583FD2C65AF649B" pageId="10" pageNumber="11" startId="10.[129,194,695,719]" targetBox="[289,1313,144,667]" targetPageId="10" targetType="figure"> +<paragraph id="E22B60550662FF9F5583FD2C65AF649B" blockId="10.[129,1433,695,748]" pageId="10" pageNumber="11"> +<emphasis id="D0E0BC470662FF9F5583FD2C60EE64B8" bold="true" box="[129,216,695,719]" pageId="10" pageNumber="11">Figure 7.</emphasis> +Internal cavity volume in the cervical vertebrae of ‘ +<emphasis id="D0E0BC470662FF9F57BAFD2362F364B8" box="[696,709,696,719]" italics="true" pageId="10" pageNumber="11">P</emphasis> +’. +<taxonomicName id="25941BD60662FF9F57D3FD23632964B8" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[721,799,696,719]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="antiquus"> +<emphasis id="D0E0BC470662FF9F57D3FD23632964B8" box="[721,799,696,719]" italics="true" pageId="10" pageNumber="11">antiquus</emphasis> +</taxonomicName> +MGUWr 3895s (A) and +<taxonomicName id="25941BD60662FF9F510CFD2C649864B8" box="[1038,1198,695,719]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="D0E0BC470662FF9F510CFD2C64A664B8" box="[1038,1168,695,719]" italics="true" pageId="10" pageNumber="11">Tanystropheus</emphasis> +sp. +</taxonomicName> +from Miedary ZPAL V. 36/181 (B). Anatomical abbreviation: ic—internal cavity. Images obtained using ‘xray’ shader in MeshLab v.2020.12 ( +<bibRefCitation id="86051DA40662FF9F51DAFD4865BF649C" author="Cignoni P & Callieri M & Corsini M" box="[1240,1417,723,747]" pageId="10" pageNumber="11" pagination="29 - 36" refId="ref15718" refString="Cignoni P, Callieri M, Corsini M et al. MeshLab: an open-source mesh processing tool. In: 6 th Eurographics Italian Chapter Conference 2008 - Proceedings, Salerno, Italy, 2008. p. 29 - 36. Eindhoven, The Netherlands: Eurographics Association, 2008." type="book chapter" year="2008"> +Cignoni +<emphasis id="D0E0BC470662FF9F5028FD4F6560649C" box="[1322,1366,723,747]" italics="true" pageId="10" pageNumber="11">et al.</emphasis> +2008 +</bibRefCitation> +). +</paragraph> +</caption> +<paragraph id="E22B60550662FF9F559EFAF462AB60B7" blockId="10.[128,779,795,1728]" pageId="10" pageNumber="11"> +The dorsal vertebrae were generally similar to the posteriormost cervicals in their internal anatomy. They differed in a complete lack of subcentral foramina. Moreover, the neural canal of the dorsals extended anteroposteriorly straight, with its roof and floor remaining on a relatively constant horizontal level ( +<figureCitation id="7AAF7CD00662FF9F5589F99060E16054" box="[139,215,1547,1571]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6H</figureCitation> +). Its transverse cross section was similar in shape to the posteriormost cervicals, but more regular. Congruently to what could be observed in the 12th and 13th cervical vertebrae, the internal portion of the centrum was composed of sparsely distributed trabeculae containing a large number of cavities, some of which opened to the neural canal through its floor. +</paragraph> +<paragraph id="E22B60550662FF9F546DF944622D608E" blockId="10.[367,539,1759,1785]" box="[367,539,1759,1785]" pageId="10" pageNumber="11"> +<emphasis id="D0E0BC470662FF9F546DF944622D608E" box="[367,539,1759,1785]" italics="true" pageId="10" pageNumber="11">Vertebral histology</emphasis> +</paragraph> +<paragraph id="E22B60550662FF9F5583F89D64A963F0" blockId="10.[129,780,1798,1979]" lastBlockId="10.[824,1476,795,1415]" pageId="10" pageNumber="11"> +The histological characteristics of the studied vertebrae of +<taxonomicName id="25941BD60662FF9F5583F8BE610A614A" box="[129,316,1829,1853]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="undetermined"> +<emphasis id="D0E0BC470662FF9F5583F8BE613B614A" box="[129,269,1829,1853]" italics="true" pageId="10" pageNumber="11">Tanystropheus</emphasis> +spp. +</taxonomicName> +were congruent with the results of the study performed by +<bibRefCitation id="86051DA40662FF9F5417F8DE6210612A" author="Jaquier VP & Scheyer TM" box="[277,550,1861,1885]" pageId="10" pageNumber="11" refId="ref16697" refString="Jaquier VP, Scheyer TM. Bone histology of the Middle Triassic long-necked reptiles Tanystropheus and Macrocnemus (Archosauromorpha, Protorosauria). Journal of Vertebrate Paleontology 2017; 37: e 1296456. https: // doi. org / 10.1080 / 02724634.2017.1296456" type="journal volume" year="2017">Jaquier and Scheyer (2017</bibRefCitation> +and the supplemental data provided therein). New data presented herein expand our knowledge on the anteroposterior and ontogenetic variation of their structure. The walls of the internal cavity were built of primary compact lamellar to parallel-fibred matrix of periosteal origin. Some intervals with decreased vascularization could be traced within these bone walls—they formed ellipsoidal annuli that alternated with more vascularized zones in parallel to the circumference of the vertebra (see +<bibRefCitation id="86051DA40662FF9F51A4FC02658765C6" author="Jaquier VP & Scheyer TM" box="[1190,1457,921,945]" pageId="10" pageNumber="11" refId="ref16697" refString="Jaquier VP, Scheyer TM. Bone histology of the Middle Triassic long-necked reptiles Tanystropheus and Macrocnemus (Archosauromorpha, Protorosauria). Journal of Vertebrate Paleontology 2017; 37: e 1296456. https: // doi. org / 10.1080 / 02724634.2017.1296456" type="journal volume" year="2017">Jaquier and Scheyer 2017</bibRefCitation> +). Closer to the terminal portions of the anteroposterior length of the cervicals, the tissue composition changed, with the internal cavity being dorsolaterally lined with secondary endosteal lamellar bone ( +<figureCitation id="7AAF7CD00662FF9F56C0FB8D64386259" box="[962,1038,1046,1070]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="10" pageNumber="11">Fig. 8H</figureCitation> +) and the ventrolateral portions of the bone centrum not exhibiting the regular annuli, but rather being nearly completely composed of endosteal lamellar bone scattered with secondary osteons and erosion cavities of varying size, forming a net of sparse trabeculae ( +<figureCitation id="7AAF7CD00662FF9F5150FB0864F662DC" box="[1106,1216,1171,1195]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="10" pageNumber="11">Figs 5B–D</figureCitation> +, +<figureCitation id="7AAF7CD00662FF9F51CEFB08653D62DC" box="[1228,1291,1171,1195]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="10" pageNumber="11">8D, G</figureCitation> +). The core of the neural spine in larger specimens was highly remodelled with longitudinally oriented secondary osteons. Simple primary vascular canals extended radially in the middle part of the vertebrae, but became more longitudinally oriented, larger, and less organized in the anteroposteriorly terminal portions. The number of vascular canals was also smaller in the more compact sections of the cortex within a single cross section. +</paragraph> +</subSubSection> +</treatment> +</document> \ No newline at end of file