361 lines
58 KiB
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361 lines
58 KiB
XML
<document id="FED7BB14A2875C55A3069720B94C9CFE" ID-DOI="10.1093/zoolinnean/zlae095" ID-ISSN="0024-4082" ID-Zenodo-Dep="14341218" IM.bibliography_approvedBy="carolina" IM.illustrations_approvedBy="carolina" IM.materialsCitations_approvedBy="carolina" IM.metadata_approvedBy="carolina" IM.tables_requiresApprovalFor="GgImagineBatch,operationResults" IM.taxonomicNames_approvedBy="carolina" 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="4" lastPageNumber="18" masterDocId="FF80CF4DFFE60D1FC702FFD68932FFA1" masterDocTitle="A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates" masterLastPageNumber="50" masterPageNumber="1" pageNumber="16" updateTime="1733771398044" updateUser="ExternalLinkService">
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<mods:title id="830334033709965D898FBF426F98D271">A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates</mods:title>
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<mods:namePart id="CA2056BCD407DC3FCE303B6B830A43B5">Püschel, Hans P.</mods:namePart>
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<mods:affiliation id="48B78FB40568145A0F3AFCE5BE30580A">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="34B3139553E1A41AF73F7D4E523C3132" type="email">hpuschelr@gmail.com</mods:nameIdentifier>
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<mods:namePart id="26CFA6877FBA77BCEE16345D4AD6ED57">Shelley, Sarah L.</mods:namePart>
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<mods:affiliation id="901AE4D6241ACC0E3D18BA4FFA2396A4">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="21E76E49DC1A9D2020F5612BE9F5F76E">Williamson, Thomas E.</mods:namePart>
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<mods:affiliation id="230DF77038E03A9E57E3D4BE5471A03D">New Mexico Museum of Natural History and Science, Albuquerque, NM 87104, United States</mods:affiliation>
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<mods:namePart id="483CD11EB72E87CCF116E0A923F4F761">Perini, Fernando A.</mods:namePart>
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<mods:affiliation id="E2545BDE885F66581E2D12CF31B0FCD7">Instituto de Ciência Biológicas, UFMG, BH 31270 - 901, Brazil</mods:affiliation>
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<mods:namePart id="5C8FC444D0363EF5E3D3228F6BEAC4D6">Wible, John R.</mods:namePart>
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<mods:affiliation id="9E42A3E968B0858E380D69E38A348A15">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="8E636D7D56669508A25DDCBC81F75B09">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="18" 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" authority="(Bond & Vucetich, 1983)" 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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,
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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>
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currently includes three accepted genera with an exclusive Palaeogene distribution (
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<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>
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; Supporting information,
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<tableCitation id="C6923398FFE90D10C7A9F91A8839F945" box="[171,267,1740,1764]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionTargetPageId="2" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" httpUri="http://table.plazi.org/id/DF6F56ABFFE40D1DC76CFE228F67FDAC" pageId="15" pageNumber="16" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table S1</tableCitation>
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).
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<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>
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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
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<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>
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, which included
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<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">
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<emphasis id="B964DA31FFE90D10C6FEF8FC8BFBF8E3" box="[508,713,1834,1858]" italics="true" pageId="15" pageNumber="16">Indalecia grandensis</emphasis>
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</taxonomicName>
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and
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<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">
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<emphasis id="B964DA31FFE90D10C773F89F882FF8C0" box="[113,285,1865,1889]" italics="true" pageId="15" pageNumber="16">Adiantoides leali</emphasis>
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(
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<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>
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)
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</taxonomicName>
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. Previously,
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<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">
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<emphasis id="B964DA31FFE90D10C5E1F89F89A8F821" italics="true" pageId="15" pageNumber="16">A. leali</emphasis>
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</taxonomicName>
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from Divisadero Largo Formation,
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<collectingCountry id="F30746B3FFE90D10C52EF8BE8BA6F821" box="[556,660,1896,1920]" name="Argentina" pageId="15" pageNumber="16">Argentina</collectingCountry>
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(age uncertain;
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<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">
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Cerdeño
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<emphasis id="B964DA31FFE90D10C62FF85E8868F801" box="[301,346,1928,1952]" italics="true" pageId="15" pageNumber="16">et al</emphasis>
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. 2008
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</bibRefCitation>
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, 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]" captionTargetPageId="4" 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" httpUri="http://table.plazi.org/id/DF6F56ABFFE20D1BC76CFE6E8C76FDA9" pageId="17" pageNumber="18" tableUuid="DF6F56ABFFE20D1BC76CFE6E8C76FDA9">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="8BAF0623FFF60D0EC783F8E689ECF81B" blockId="16.[129,1465,1811,1976]" lastBlockId="17.[113,1454,1675,1979]" lastPageId="17" lastPageNumber="18" 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, 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>
|
||
<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]" captionTargetPageId="2" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" httpUri="http://table.plazi.org/id/DF6F56ABFFE40D1DC76CFE228F67FDAC" pageId="17" pageNumber="18" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">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>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |