2171 lines
468 KiB
XML
2171 lines
468 KiB
XML
<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">
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<mods:title id="CFF053801AEBC538B899732294C07E24">A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates</mods:title>
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<mods:namePart id="5E3EA8A002C86E468ECCE84453C56A7C">Püschel, Hans P.</mods:namePart>
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<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>
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<mods:nameIdentifier id="FD99E0DEBD9D64770DBE581728266613" type="email">hpuschelr@gmail.com</mods:nameIdentifier>
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<mods:namePart id="88F5D09E86F0A1BF156A59FE815F3D22">Shelley, Sarah L.</mods:namePart>
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<mods:affiliation id="943D5E65BBE8F1AD35DA1C038D00F14C">School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, EH 9 3 FE, United Kingdom</mods:affiliation>
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<mods:namePart id="09D218F971F5D04F042B73EC9DB64C5B">Williamson, Thomas E.</mods:namePart>
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<mods:affiliation id="6074194580E7CA34D051245743D7D238">New Mexico Museum of Natural History and Science, Albuquerque, NM 87104, United States</mods:affiliation>
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<mods:namePart id="1D0EF98CD1C6C501879ED1BDAE933F44">Perini, Fernando A.</mods:namePart>
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<mods:affiliation id="83ACCF1F7FCF9440E1A9240C56E3814A">Instituto de Ciência Biológicas, UFMG, BH 31270 - 901, Brazil</mods:affiliation>
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<mods:namePart id="30F9A1D868E65755170F51ABE2527F9E">Wible, John R.</mods:namePart>
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<mods:affiliation id="1147DF07496B5630841BFFDA5DD3F933">Section of Mammals, Carnegie Museum of Natural History, 5800 Baum Boulevard, Pittsburgh, PA 15206, United States</mods:affiliation>
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<mods:namePart id="B0555C400C4EAB63400AE51B67E450A9">Brusatte, Stephen L.</mods:namePart>
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<mods:affiliation id="06F33F779C8EFA5D4D494389F448DD01">School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, EH 9 3 FE, United Kingdom</mods:affiliation>
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<mods:date id="1526278939CC04166CEB407226CE1844">2024</mods:date>
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<treatment id="03B9B735FFE90D0EC638F9B08B33FD9E" LSID="urn:lsid:plazi:treatment:03B9B735FFE90D0EC638F9B08B33FD9E" httpUri="http://treatment.plazi.org/id/03B9B735FFE90D0EC638F9B08B33FD9E" lastPageId="17" lastPageNumber="27" pageId="15" pageNumber="16">
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<paragraph id="8BAF0623FFE90D10C638F9B08B00F920" blockId="15.[113,763,1638,1983]" box="[314,562,1638,1665]" pageId="15" pageNumber="16">
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<emphasis id="B964DA31FFE90D10C638F9B08B00F920" box="[314,562,1638,1665]" italics="true" pageId="15" pageNumber="16">
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<taxonomicName id="4C107DA0FFE90D10C638F9B08898F921" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[314,426,1638,1664]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Indaleciidae</taxonomicName>
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(
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<figureCitation id="132B1AA6FFE90D10C6BBF9B188CDF921" box="[441,511,1638,1664]" 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="15" pageNumber="16">Figs 6A</figureCitation>
|
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,
|
||
<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>
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)
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</emphasis>
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</paragraph>
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</subSubSection>
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<subSubSection id="C30A55A8FFE90D0EC773F95B8B33FD9E" lastPageId="17" lastPageNumber="18" pageId="15" pageNumber="16" type="multiple">
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<paragraph id="8BAF0623FFE90D10C773F95B8D53FBF5" blockId="15.[113,763,1638,1983]" lastBlockId="15.[808,1461,144,1985]" pageId="15" pageNumber="16">
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<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
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<emphasis id="B964DA31FFFC0D05C221FD138C62FD7D" box="[1315,1360,708,732]" italics="true" pageId="26" pageNumber="27">et al.</emphasis>
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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).
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</paragraph>
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||
</subSection>
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||
</caption>
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||
</subSubSection>
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||
</caption>
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||
</subSubSection>
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</treatment>
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</document> |