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<mods:title id="7045782E10A3C52C80E48C45B2E1D0BD">A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and archaic South American ungulates</mods:title>
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<mods:namePart id="DA894FDC28434E29414F209A01D8E773">Püschel, Hans P.</mods:namePart>
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<mods:affiliation id="C69A5C012CFDF2F9C50DDD2905C99F8D">Instituto de Ciência Biológicas, UFMG, BH 31270 - 901, Brazil</mods:affiliation>
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<mods:namePart id="5C2E909B8AD0C07D84C57A50B82DDEAD">Wible, John R.</mods:namePart>
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<taxonomicName id="4C107DA0FFF70D0EC4C1FCEC8D64FCF5" authority="Savage, 1951" 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>
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<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, P4M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2M3 of Escribania chubutensis in occlusal view (MLP 90II-1268, roots of M1 and M2 (mirrored); MLP 90II-1263, M3). CD, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2M3 (holotype; mirrored); MACN A 10738, P1]. H, P5M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4P5, and M1M3 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 8BG</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, p4m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90II1269 (mirrored)]. D, P5M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2m3 of Didolodus multicuspis (MACN A 10689). F, p2m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4m3 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">9BE</figureCitation>
)
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<paragraph id="8BAF0623FFF70D0CC428FCB7886CFDDF" blockId="17.[809,1461,826,1547]" lastBlockId="19.[113,764,144,1703]" lastPageId="19" lastPageNumber="20" 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]" 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>
). 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]" 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 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 Condylarthra. After adding two new species from the Rio Chico Formation [latest DanianLate 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">Ameghinos (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>
<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, p1m3 of the indaleciid
<taxonomicName id="4C107DA0FFF40D0DC4EAF8968DAFF8F9" authorityName="Bond &amp; 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 (p1p2 mirrored)]. B, p2m3 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, p1m3 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, p1m3 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="8BAF0623FFF40D0CC783F8428A28F81B" blockId="18.[129,1464,1856,1964]" lastBlockId="19.[113,1452,1786,1979]" lastPageId="19" lastPageNumber="20" 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, p1m3 of
<taxonomicName id="4C107DA0FFF40D0DC4E1F8428DE7F80D" authorityName="Soria &amp; 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, p1m2 (
<typeStatus id="54ABB881FFF50D0CC778F92C89E4F8B3" box="[122,214,1786,1810]" pageId="19" pageNumber="20" type="holotype">holotype</typeStatus>
; p1p5, m2 mirrored); PVL 4216, m3. G, p4m3 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>
<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 &amp; Sige, 2011" authorityName="Gelfo &amp; 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 &amp; 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 &amp; 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 (AB) and its interfamilial relationships (CD). 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]" 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="19" pageNumber="20" tableUuid="DF6F56ABFFE20D1BC76CFE6E8C76FDA9">Table 2</tableCitation>
).
</paragraph>
<paragraph id="8BAF0623FFF50D0AC447FB6F88FFFE22" blockId="19.[808,1461,144,1703]" lastBlockId="21.[113,763,144,638]" lastPageId="21" lastPageNumber="22" 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 (OLeary
<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 orders (
<tableCitation id="C6923398FFF30D0AC7C6FF46883DFF09" box="[196,271,144,168]" 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="21" pageNumber="22" tableUuid="DF6F56ABFFE20D1BC76CFE6E8C76FDA9">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 &amp; Badin AC &amp; 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>
<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="8BAF0623FFF20D0AC781F9238B56F81B" blockId="20.[131,1428,1781,1973]" lastBlockId="21.[113,1431,1787,1979]" lastPageId="21" lastPageNumber="22" 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, P4M3 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, P4M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2M3 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 90II-1268, roots of M1 and M2 (mirrored); MLP 90II-1263, M3). CD, 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)] in occlusal (C) and distoocclusal (D) views. E, P4M3 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, P5M3 (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). G, P1M3 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, P2M3 (holotype; mirrored); MACN A 10738, P1]. H, P5M3 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, P4M3 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 358M (
<typeStatus id="54ABB881FFF30D0AC7B5F92D8821F8B2" box="[183,275,1787,1811]" pageId="21" pageNumber="22" type="holotype">holotype</typeStatus>
; mirrored)]. J, P2, alveoli of P4P5, and M1M3 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) 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>
<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 63.836.5 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]" 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="21" pageNumber="22" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table S1</tableCitation>
; Woodburne
<emphasis id="B964DA31FFF30D0AC773FDB189A8FDDF" box="[113,154,614,638]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. 2014a, b).
</paragraph>
</subSubSection>
</treatment>
</document>