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<mods:title id="09D07E03DF426892629E9AC7295AD69B">A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates</mods:title>
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<mods:namePart id="29BEC46CF04C511EF4432543DFDCB91C">Püschel, Hans P.</mods:namePart>
<mods:affiliation id="B2BC737533E634A545F9CC1312844221">Millenium Nucleus Early Evolutionary Transitions of Mammals (EVOTEM), Red Palentológica U-Chile, Santiago, 7800003, Chile &amp; 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="836694D2A838B5C9345C391782BF073E">Shelley, Sarah L.</mods:namePart>
<mods:affiliation id="3A8514E2D5215FC728561537568CF7A6">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="81DD5543250E443A99D74ECE5455370B">Williamson, Thomas E.</mods:namePart>
<mods:affiliation id="BDC47139BD38868368FE0B4C58D901FC">New Mexico Museum of Natural History and Science, Albuquerque, NM 87104, United States</mods:affiliation>
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<mods:namePart id="A9F0BC55F7B5D587E6F8234B4BCA082C">Perini, Fernando A.</mods:namePart>
<mods:affiliation id="65769FCCA507E0FAAF3065B144F0B952">Instituto de Ciência Biológicas, UFMG, BH 31270 - 901, Brazil</mods:affiliation>
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<mods:namePart id="1AF8146AE9045DABE30EA92639FB34CB">Wible, John R.</mods:namePart>
<mods:affiliation id="AF2982CC02795CC7BCEDE6F663378A11">Section of Mammals, Carnegie Museum of Natural History, 5800 Baum Boulevard, Pittsburgh, PA 15206, United States</mods:affiliation>
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<subSubSection id="C30A55A8FFEA0D13C61EFDD78B5FFDBA" box="[284,621,513,539]" pageId="12" pageNumber="13" type="nomenclature">
<paragraph id="8BAF0623FFEA0D13C61EFDD78B5FFDBA" blockId="12.[127,780,513,1641]" box="[284,621,513,539]" pageId="12" pageNumber="13">
<emphasis id="B964DA31FFEA0D13C61EFDD78B5FFDBA" box="[284,621,513,539]" italics="true" pageId="12" pageNumber="13">
<taxonomicName id="4C107DA0FFEA0D13C61EFDD788DBFDBA" authority="Soria, 1980" box="[284,489,513,539]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
(
<figureCitation id="132B1AA6FFEA0D13C6FAFDD48B0FFDBA" box="[504,573,513,539]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="12" pageNumber="13">Figs 4L</figureCitation>
,
<figureCitation id="132B1AA6FFEA0D13C545FDD78B51FDBA" box="[583,611,513,539]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="12" pageNumber="13">5L</figureCitation>
)
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<paragraph id="8BAF0623FFEA0D13C783FDF188E9FB70" blockId="12.[127,780,513,1641]" pageId="12" pageNumber="13">
<taxonomicName id="4C107DA0FFEA0D13C783FDF18851FD9E" box="[129,355,551,575]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
currently has five accepted genera, all with a fossil record exclusively in the Palaeogene (
<figureCitation id="132B1AA6FFEA0D13C54DFD918BBAFDFE" box="[591,648,583,607]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="12" pageNumber="13">Fig. 2</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFEA0D13C609FDB0885BFDDF" box="[267,361,614,638]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="12" pageNumber="13" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table S1</tableCitation>
). The family
<taxonomicName id="4C107DA0FFEA0D13C6FBFDB08BE9FDDF" box="[505,731,614,638]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
was initially proposed by Soria (1980a) to include
<taxonomicName id="4C107DA0FFEA0D13C56AFD5388AAFD1C" authority="Soria, 1980 a" authorityName="Soria" authorityYear="1980" class="Mammalia" family="Sparnotheriodontidae" genus="Sparnotheriodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="epsilonoides">
<emphasis id="B964DA31FFEA0D13C56AFD5389C6FD1C" italics="true" pageId="12" pageNumber="13">Sparnotheriodon epsilonoides</emphasis>
Soria, 1980a
</taxonomicName>
from Cañadón Vaca Member, Sarmiento Formation,
<collectingCountry id="F30746B3FFEA0D13C683FD1288DBFD7D" box="[385,489,708,732]" name="Argentina" pageId="12" pageNumber="13">Argentina</collectingCountry>
, which was classified as
<emphasis id="B964DA31FFEA0D13C783FD328836FD5A" box="[129,260,739,763]" italics="true" pageId="12" pageNumber="13">incertae sedis</emphasis>
within the order
<taxonomicName id="4C107DA0FFEA0D13C6CBFD358B68FD5A" baseAuthorityName="Soria" baseAuthorityYear="2001" box="[457,602,739,763]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="order">Notoungulata</taxonomicName>
. The family was later removed from
<taxonomicName id="4C107DA0FFEA0D13C658FCD588DAFCBA" baseAuthorityName="Soria" baseAuthorityYear="2001" box="[346,488,771,795]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="order">Notoungulata</taxonomicName>
and incorporated into the order
<taxonomicName id="4C107DA0FFEA0D13C7BDFCF4881CFC9B" box="[191,302,802,826]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="order">Litopterna</taxonomicName>
, being expanded by Soria (1980b) adding the genera
<taxonomicName id="4C107DA0FFEA0D13C7D2FC978850FCF8" authorityName="Ameghino" authorityYear="1901" box="[208,354,833,857]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEA0D13C7D2FC978850FCF8" box="[208,354,833,857]" italics="true" pageId="12" pageNumber="13">Victorlemoinea</emphasis>
</taxonomicName>
and
<taxonomicName id="4C107DA0FFEA0D13C6A3FC978A36FCF8" authority="Simpson et al. 1962" authorityName="Simpson" authorityYear="1962" box="[417,772,833,857]" class="Mammalia" family="Proterotheriidae" genus="Phoradiadus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEA0D13C6A3FC978B2FFCF8" box="[417,541,833,857]" italics="true" pageId="12" pageNumber="13">Phoradiadus</emphasis>
Simpson
<emphasis id="B964DA31FFEA0D13C58DFC948BF6FCF8" box="[655,708,833,857]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
1962
</taxonomicName>
.
<taxonomicName id="4C107DA0FFEA0D13C783FCB78821FCD8" authorityName="Ameghino" authorityYear="1901" box="[129,275,865,889]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEA0D13C783FCB78821FCD8" box="[129,275,865,889]" italics="true" pageId="12" pageNumber="13">Victorlemoinea</emphasis>
</taxonomicName>
was previously considered an early macraucheniid (Simpson 1945, 1948), and
<taxonomicName id="4C107DA0FFEA0D13C6B0FC568B1CFC39" box="[434,558,896,920]" class="Mammalia" family="Proterotheriidae" genus="Phoradiadus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEA0D13C6B0FC568B1CFC39" box="[434,558,896,920]" italics="true" pageId="12" pageNumber="13">Phoradiadus</emphasis>
</taxonomicName>
was previously considered with doubts as a proterotheriid (Simpson
<emphasis id="B964DA31FFEA0D13C58EFC768B8CFC16" box="[652,702,927,951]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
1962). Soria’s (1980b) proposal was based on shared similarities in the upper and lower molars, such as the presence of a lophoid metaconule and a closed trigon basin in M2.
<taxonomicName id="4C107DA0FFEA0D13C56AFC2B8BD6FBB4" box="[616,740,1021,1045]" class="Mammalia" family="Proterotheriidae" genus="Phoradiadus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEA0D13C56AFC2B8BD6FBB4" box="[616,740,1021,1045]" italics="true" pageId="12" pageNumber="13">Phoradiadus</emphasis>
</taxonomicName>
includes one species,
<taxonomicName id="4C107DA0FFEA0D13C65AFBCB8B7EFB94" authorityName="Simpson, Minoprio y Patterson" authorityYear="1962" box="[344,588,1053,1077]" class="Mammalia" family="Proterotheriidae" genus="Phoradiadus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="divortiensis">
<emphasis id="B964DA31FFEA0D13C65AFBCB8B7EFB94" box="[344,588,1053,1077]" italics="true" pageId="12" pageNumber="13">Phoradiadus divortiensis</emphasis>
</taxonomicName>
, from Divisadero Largo Formation,
<collectingRegion id="49D4C8C1FFEA0D13C638FBEA88AEFBF5" box="[314,412,1084,1108]" country="Argentina" name="Mendoza" pageId="12" pageNumber="13">Mendoza</collectingRegion>
,
<collectingCountry id="F30746B3FFEA0D13C6A7FBEA8B38FBF5" box="[421,522,1084,1108]" name="Argentina" pageId="12" pageNumber="13">Argentina</collectingCountry>
(age uncertain;
<bibRefCitation id="EF817BD2FFEA0D13C5B2FBEA89D5FBD2" author="Cerdeno" etAl="et al." firstAuthor="Cerdeno" pageId="12" pageNumber="13" pagination="574 - 7" refId="ref40752" refString="Cerdeno E, Lopez GM, Reguero MA. Biostratigraphic considerations of the Divisaderan faunal assemblage. Journal of Vertebrate Paleontology 2008; 28: 574 - 7. https: // doi. org / 10.1671 / 0272 - 4634 (2008) 28 (574: bcotdf) 2.0. co; 2" type="journal article" year="2008">
Cerdeño
<emphasis id="B964DA31FFEA0D13C783FB8A899AFBD2" box="[129,168,1115,1139]" italics="true" pageId="12" pageNumber="13">et al</emphasis>
. 2008
</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFEA0D13C7F3FB8D8858FBD2" author="Lopez" box="[241,362,1115,1139]" firstAuthor="Lopez" pageId="12" pageNumber="13" pagination="410 - 20" refId="ref44025" refString="Lopez GM. Dividaderan: Land Mammal Age or local fauna? In: Madden RH, Carlini AA, Vucetich MG, Kay RF (eds), The paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge: Cambridge University Press, 2010, 410 - 20." type="book chapter" year="2010">López 2010</bibRefCitation>
, Woodburne
<emphasis id="B964DA31FFEA0D13C6F6FB8A8B2EFBD2" box="[500,540,1115,1139]" italics="true" pageId="12" pageNumber="13">et al</emphasis>
. 2014a). An interesting feature of the teeth of the
<taxonomicName id="4C107DA0FFEA0D13C680FBAD8B56FB32" box="[386,612,1147,1171]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
is that they possess vertical Hunter–Schreger bands, a feature unknown in any other litoptern (
<bibRefCitation id="EF817BD2FFEA0D13C624FB6F88F8FB70" author="Bond" box="[294,458,1209,1233]" etAl="et al." firstAuthor="Bond" pageId="12" pageNumber="13" pagination="163 - 76" refId="ref40344" refString="Bond M, Reguero MA, Vizcaino SF et al. A new ' South American ungulate' (Mammalia: Litopterna) from the Eocene of the Antarctic Peninsula. Geological Society, London, Special Publications 2006; 258: 163 - 76. https: // doi. org / 10.1144 / gsl. sp. 2006.258.01.12" type="journal article" year="2006">
Bond
<emphasis id="B964DA31FFEA0D13C661FB6C88A0FB70" box="[355,402,1209,1233]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
2006
</bibRefCitation>
).
</paragraph>
<paragraph id="8BAF0623FFEA0D13C79EFB0F8D68FE22" blockId="12.[127,780,513,1641]" lastBlockId="12.[824,1478,144,1641]" pageId="12" pageNumber="13">
Simpson (1948) tentatively accepted three Patagonian species of the genus
<taxonomicName id="4C107DA0FFEA0D13C64CFB2E88D2FAB1" authorityName="Ameghino" authorityYear="1901" box="[334,480,1272,1296]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEA0D13C64CFB2E88D2FAB1" box="[334,480,1272,1296]" italics="true" pageId="12" pageNumber="13">Victorlemoinea</emphasis>
</taxonomicName>
:
<taxonomicName id="4C107DA0FFEA0D13C6F7FB2E881DFA8E" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="labyrinthica">
<emphasis id="B964DA31FFEA0D13C6F7FB2E8A3BFAB1" box="[501,777,1272,1296]" italics="true" pageId="12" pageNumber="13">Victorlemoinea labyrinthica</emphasis>
<bibRefCitation id="EF817BD2FFEA0D13C783FAC1881DFA8E" author="Ameghino" box="[129,303,1303,1327]" firstAuthor="Ameghino" pageId="12" pageNumber="13" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C107DA0FFEA0D13C63DFAC18A36FA8E" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[319,772,1303,1327]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="emarginata">
<emphasis id="B964DA31FFEA0D13C63DFAC18B79FA8E" box="[319,587,1303,1327]" italics="true" pageId="12" pageNumber="13">Victorlemoinea emarginata</emphasis>
<bibRefCitation id="EF817BD2FFEA0D13C557FAC18A36FA8E" author="Ameghino" box="[597,772,1303,1327]" firstAuthor="Ameghino" pageId="12" pageNumber="13" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C107DA0FFEA0D13C7AFFAE18B62FAEE" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[173,592,1335,1359]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="longidens">
<emphasis id="B964DA31FFEA0D13C7AFFAE188ADFAEE" box="[173,415,1335,1359]" italics="true" pageId="12" pageNumber="13">Victorlemoinea longidens</emphasis>
<bibRefCitation id="EF817BD2FFEA0D13C6A7FAE18B62FAEE" author="Ameghino" box="[421,592,1335,1359]" firstAuthor="Ameghino" pageId="12" pageNumber="13" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
</taxonomicName>
, as they were represented only by isolated teeth, the first two being represented by upper teeth, and the last by lower teeth. This means that
<taxonomicName id="4C107DA0FFEA0D13C783FA4389C5FA0C" authorityName="Ameghino" authorityYear="1901" box="[129,247,1429,1453]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="longidens">
<emphasis id="B964DA31FFEA0D13C783FA4389C5FA0C" box="[129,247,1429,1453]" italics="true" pageId="12" pageNumber="13">V. longidens</emphasis>
</taxonomicName>
could correspond to teeth from the lower dentition of either
<taxonomicName id="4C107DA0FFEA0D13C611FA638891FA6D" authorityName="Ameghino" authorityYear="1901" box="[275,419,1460,1484]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="labyrinthica">
<emphasis id="B964DA31FFEA0D13C611FA638891FA6D" box="[275,419,1460,1484]" italics="true" pageId="12" pageNumber="13">V. labyrinthica</emphasis>
</taxonomicName>
or
<taxonomicName id="4C107DA0FFEA0D13C6C8FA638B66FA6D" authorityName="Ameghino" authorityYear="1901" box="[458,596,1461,1484]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="emarginata">
<emphasis id="B964DA31FFEA0D13C6C8FA638B66FA6D" box="[458,596,1461,1484]" italics="true" pageId="12" pageNumber="13">V. emarginata</emphasis>
</taxonomicName>
, in particular the mandibular fragment with m1–m2 of the
<typeStatus id="54ABB881FFEA0D13C54BFA058B97FA4A" box="[585,677,1491,1515]" pageId="12" pageNumber="13" type="holotype">holotype</typeStatus>
(MACN A-10670), as the allegedly associated premolars of this specimen seem to correspond to an isotemnid notoungulate (
<bibRefCitation id="EF817BD2FFEA0D13C5D3F9C489D9F9E8" author="Bond" etAl="et al." firstAuthor="Bond" pageId="12" pageNumber="13" pagination="163 - 76" refId="ref40344" refString="Bond M, Reguero MA, Vizcaino SF et al. A new ' South American ungulate' (Mammalia: Litopterna) from the Eocene of the Antarctic Peninsula. Geological Society, London, Special Publications 2006; 258: 163 - 76. https: // doi. org / 10.1144 / gsl. sp. 2006.258.01.12" type="journal article" year="2006">
Bond
<emphasis id="B964DA31FFEA0D13C783F9E48983F9E8" box="[129,177,1585,1609]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
2006
</bibRefCitation>
).
<location id="8ECF50F8FFEA0D13C600F9E4882AF9E8" LSID="urn:lsid:plazi:treatment:03B9B735FFEA0D11C61EFDD78B0BFC57:8ECF50F8FFEA0D13C600F9E4882AF9E8" box="[258,280,1586,1609]" country="Brazil" county="Itaborai" municipality="In" name="In" pageId="12" pageNumber="13">In</location>
the genus
<taxonomicName id="4C107DA0FFEA0D13C688F9E78BD5F9E8" authority=", Bond et al. (2006)" authorityName="Bond" authorityYear="2006" box="[394,743,1585,1609]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEA0D13C688F9E78B28F9E8" box="[394,538,1585,1609]" italics="true" pageId="12" pageNumber="13">Victorlemoinea</emphasis>
,
<bibRefCitation id="EF817BD2FFEA0D13C525F9E78BD5F9E8" author="Bond" box="[551,743,1585,1609]" etAl="et al." firstAuthor="Bond" pageId="12" pageNumber="13" pagination="163 - 76" refId="ref40344" refString="Bond M, Reguero MA, Vizcaino SF et al. A new ' South American ungulate' (Mammalia: Litopterna) from the Eocene of the Antarctic Peninsula. Geological Society, London, Special Publications 2006; 258: 163 - 76. https: // doi. org / 10.1144 / gsl. sp. 2006.258.01.12" type="journal article" year="2006">
Bond
<emphasis id="B964DA31FFEA0D13C564F9E48BA4F9E8" box="[614,662,1585,1609]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
(2006)
</bibRefCitation>
</taxonomicName>
included only
<taxonomicName id="4C107DA0FFEA0D13C607F98788A4F9C8" authorityName="Ameghino" authorityYear="1901" box="[261,406,1617,1641]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="labyrinthica">
<emphasis id="B964DA31FFEA0D13C607F98788A4F9C8" box="[261,406,1617,1641]" italics="true" pageId="12" pageNumber="13">V. labyrinthica</emphasis>
</taxonomicName>
in their list of formally recognized sparnotheriodontid species, and more recently some authors have mentioned only
<taxonomicName id="4C107DA0FFEA0D13C311FF668D92FF66" authorityName="Ameghino" authorityYear="1901" box="[1043,1184,175,199]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="labyrinthica">
<emphasis id="B964DA31FFEA0D13C311FF668D92FF66" box="[1043,1184,175,199]" italics="true" pageId="12" pageNumber="13">V. labyrinthica</emphasis>
</taxonomicName>
for the
<location id="8ECF50F8FFEA0D13C3EFFF798C67FF66" LSID="urn:lsid:plazi:treatment:03B9B735FFEA0D11C61EFDD78B0BFC57:8ECF50F8FFEA0D13C3EFFF798C67FF66" box="[1261,1365,175,199]" country="Brazil" county="Itaborai" municipality="In" name="Riochican" pageId="12" pageNumber="13">Riochican</location>
and
<location id="8ECF50F8FFEA0D13C287FF668A98FF46" LSID="urn:lsid:plazi:treatment:03B9B735FFEA0D11C61EFDD78B0BFC57:8ECF50F8FFEA0D13C287FF668A98FF46" country="Brazil" county="Itaborai" municipality="In" name="Vacan Patagonian" pageId="12" pageNumber="13">Vacan Patagonian</location>
faunas, omitting
<taxonomicName id="4C107DA0FFEA0D13C35CFF198DD4FF47" authorityName="Ameghino" authorityYear="1901" box="[1118,1254,207,230]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="emarginata">
<emphasis id="B964DA31FFEA0D13C35CFF198DD4FF47" box="[1118,1254,207,230]" italics="true" pageId="12" pageNumber="13">V. emarginata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C107DA0FFEA0D13C21AFF198CB9FF46" authorityName="Ameghino" authorityYear="1901" box="[1304,1419,207,231]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="longidens">
<emphasis id="B964DA31FFEA0D13C21AFF198CB9FF46" box="[1304,1419,207,231]" italics="true" pageId="12" pageNumber="13">V. longidens</emphasis>
</taxonomicName>
(e.g., Reguero
<emphasis id="B964DA31FFEA0D13C497FF398AF6FEA7" box="[917,964,238,262]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
2014, Gelfo 2016,
<bibRefCitation id="EF817BD2FFEA0D13C381FF388C1AFEA7" author="Gelfo" box="[1155,1320,238,262]" etAl="et al." firstAuthor="Gelfo" pageId="12" pageNumber="13" pagination="274 - 92" refId="ref42638" refString="Gelfo JN, Goin FJ, Bauza N et al. The fossil record of Antarctic land mammals: commented review and hypotheses for future research. Advances in Polar Science 2019; 30: 274 - 92." type="journal article" year="2019">
Gelfo
<emphasis id="B964DA31FFEA0D13C3C0FF398DC2FEA7" box="[1218,1264,238,262]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
2019
</bibRefCitation>
), which can be interpreted as implicitly synonymizing them with
<taxonomicName id="4C107DA0FFEA0D13C232FED88C8EFE84" authorityName="Ameghino" authorityYear="1901" box="[1328,1468,269,293]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="labyrinthica">
<emphasis id="B964DA31FFEA0D13C232FED88C8EFE84" box="[1328,1468,269,293]" italics="true" pageId="12" pageNumber="13">V. labyrinthica</emphasis>
</taxonomicName>
.
<collectingMunicipality id="6BCB9C59FFEA0D13C43BFEFB8A7DFEE5" box="[825,847,301,324]" pageId="12" pageNumber="13">In</collectingMunicipality>
addition, there is one
<taxonomicName id="4C107DA0FFEA0D13C346FEFB8C1AFEE4" box="[1092,1320,301,325]" pageId="12" pageNumber="13">
<emphasis id="B964DA31FFEA0D13C346FEFB8DE4FEE4" box="[1092,1238,301,325]" italics="true" pageId="12" pageNumber="13">Victorlemoinea</emphasis>
species
</taxonomicName>
from
<collectingCounty id="62CE7EAFFFEA0D13C26EFEFB8C8CFEE4" box="[1388,1470,301,325]" pageId="12" pageNumber="13">Itaboraí</collectingCounty>
,
<collectingCountry id="F30746B3FFEA0D13C43BFE9A8A44FEC5" box="[825,886,332,356]" name="Brazil" pageId="12" pageNumber="13">Brazil</collectingCountry>
,
<taxonomicName id="4C107DA0FFEA0D13C47DFE9A8C76FEC5" authority="Paula Couto, 1952" authorityName="Paula Couto" authorityYear="1952" box="[895,1348,332,356]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="prototypica">
<emphasis id="B964DA31FFEA0D13C47DFE9A8DB6FEC5" box="[895,1156,332,356]" italics="true" pageId="12" pageNumber="13">Victorlemoinea prototypica</emphasis>
Paula Couto, 1952
</taxonomicName>
, also known mostly from dental material.
</paragraph>
<paragraph id="8BAF0623FFEA0D13C457FE5D8D43FC57" blockId="12.[824,1478,144,1641]" pageId="12" pageNumber="13">
Cifelli (1983b) indirectly assigned some tarsals (i.e., calcaneum and astragalus) to
<taxonomicName id="4C107DA0FFEA0D13C342FE7D8DF8FE63" authorityName="Paula Couto" authorityYear="1952" box="[1088,1226,427,450]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="prototypica">
<emphasis id="B964DA31FFEA0D13C342FE7D8DF8FE63" box="[1088,1226,427,450]" italics="true" pageId="12" pageNumber="13">V. prototypica</emphasis>
</taxonomicName>
, based on their relative size and abundance. These tarsals were large and scarce, so considering linear regressions with the dentition (m2 area), he argued it could only correspond to either the sparnotheriodontid
<taxonomicName id="4C107DA0FFEA0D13C438FDFE8AE3FD9E" authorityName="Paula Couto" authorityYear="1952" box="[826,977,552,575]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="prototypica">
<emphasis id="B964DA31FFEA0D13C438FDFE8AE3FD9E" box="[826,977,552,575]" italics="true" pageId="12" pageNumber="13">V. prototypica</emphasis>
</taxonomicName>
or the didolodontid
<taxonomicName id="4C107DA0FFEA0D13C3E1FDFE8C8FFD9E" authorityName="Paula Couto" authorityYear="1952" box="[1251,1469,551,575]" class="Mammalia" family="Didolodontidae" genus="Lamegoia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="conodonta">
<emphasis id="B964DA31FFEA0D13C3E1FDFE8C8FFD9E" box="[1251,1469,551,575]" italics="true" pageId="12" pageNumber="13">Lamegoia conodonta</emphasis>
</taxonomicName>
. Considering the similarities of the tarsals assigned to
<taxonomicName id="4C107DA0FFEA0D13C2ADFD918A9AFDDF" authorityName="Paula Couto" authorityYear="1952" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="prototypica">
<emphasis id="B964DA31FFEA0D13C2ADFD918A9AFDDF" italics="true" pageId="12" pageNumber="13">V. prototypica</emphasis>
</taxonomicName>
and others indirectly assigned to didolodontids, such as presenting a medial malleolar facet of the astragalus extending onto the neck and the presence of a dorsal beak on the distal end of the calcaneum,Cifelli (1983a) grouped
<taxonomicName id="4C107DA0FFEA0D13C3E2FD128CF0FD7D" box="[1248,1474,708,732]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
and
<taxonomicName id="4C107DA0FFEA0D13C466FD358D3AFD5A" box="[868,1032,739,763]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Didolodontidae</taxonomicName>
under the superfamily
<taxonomicName id="4C107DA0FFEA0D13C3F7FD358C94FD5A" box="[1269,1446,739,763]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="12" pageNumber="13" phylum="Chordata" rank="superFamily" superFamily="Didolodontoidea">Didolodontoidea</taxonomicName>
in the order
<taxonomicName id="4C107DA0FFEA0D13C4A2FCD58D19FCBA" authorityName="Cope" authorityYear="1881" box="[928,1067,771,795]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="12" pageNumber="13" phylum="Chordata" rank="order">Condylarthra</taxonomicName>
(
<tableCitation id="C6923398FFEA0D13C33EFCD58DBAFCBA" box="[1084,1160,771,795]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="12" pageNumber="13" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table 1</tableCitation>
). This hypothesis found some phylogenetic support (
<bibRefCitation id="EF817BD2FFEA0D13C328FCF48D94FC9B" author="Cifelli" box="[1066,1190,802,826]" firstAuthor="Cifelli" pageId="12" pageNumber="13" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFEA0D13C3B7FCF48C61FC9B" author="Bergqvist" box="[1205,1363,802,826]" firstAuthor="Bergqvist" pageId="12" pageNumber="13" refId="ref39790" refString="Bergqvist LP. Reasociacao de pos-cranio as especies de ungulados da Bacia de S. J. Itaborai (Paleoceno), Estado do Rio de Janeiro, e Filogenia dos ' Condylarthra', e ungulados Sul-Americanos com base no poscranio. D. Phil. Thesis, Universidade Federal do Rio Grande do Sul, Brazil, 1996." type="book" year="1996">Bergqvist 1996</bibRefCitation>
;
<figureCitation id="132B1AA6FFEA0D13C266FCF48C83FC9B" box="[1380,1457,802,826]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="12" pageNumber="13">Fig. 1C</figureCitation>
), and has been followed by some authors (e.g.,
<bibRefCitation id="EF817BD2FFEA0D13C215FC978C83FCF8" author="Bergqvist" box="[1303,1457,833,857]" firstAuthor="Bergqvist" pageId="12" pageNumber="13" pagination="107 - 33" refId="ref39850" refString="Bergqvist LP. Postcranial skeleton of the Upper Paleocene of Itaborai Basin, Brazil. In: Sargis EJ, Dagosto M, (eds), Mammalian Evolutionary Morphology: A Tribute to Frederik S. Szalay. New York, NY: Springer-Verlag, 2008, 107 - 33." type="book chapter" year="2008">Bergqvist 2008</bibRefCitation>
). However, other authors have questioned the indirect anatomical assignment of these tarsals to
<taxonomicName id="4C107DA0FFEA0D13C364FC578DDFFC39" authorityName="Paula Couto" authorityYear="1952" box="[1126,1261,897,920]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="prototypica">
<emphasis id="B964DA31FFEA0D13C364FC578DDFFC39" box="[1126,1261,897,920]" italics="true" pageId="12" pageNumber="13">V. prototypica</emphasis>
</taxonomicName>
(e.g.,
<bibRefCitation id="EF817BD2FFEA0D13C22AFC568A98FC16" author="Hoffstetter and Soria" firstAuthor="Hoffstetter" pageId="12" pageNumber="13" pagination="1619 - 22" refId="ref43229" refString="Hoffstetter R, Soria MF. Neolodus colombianus gen. et sp. nov., un noveau Condylarthre (Mammalia) dans le Miocene de Colombie. Comptes Rendus de la Academie de Sciences, Paris 1986; 303: 1619 - 22." type="journal article" year="1986">Hoffstetter and Soria 1986</bibRefCitation>
, Soria 2001,
<bibRefCitation id="EF817BD2FFEA0D13C337FC768DF0FC16" author="Lorente" box="[1077,1218,927,951]" firstAuthor="Lorente" pageId="12" pageNumber="13" refId="ref44083" refString="Lorente M. Desarrollo de Modelos de Asociacion y Clasificaciones de Restos Postcraneanos Aislados de Ungulados Nativos del Paleoceno- Eoceno de America del Sur. D. Phil. Thesis, Universidad Nacional de la Plata, Argentina, 2015." type="book" year="2015">Lorente 2015</bibRefCitation>
), and even suggested an affinity of these tarsals with
<taxonomicName id="4C107DA0FFEA0D13C35AFC698C43FC76" authority="(Soria 2001)" baseAuthorityName="Soria" baseAuthorityYear="2001" box="[1112,1393,959,983]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="order">Notoungulata (Soria 2001)</taxonomicName>
or with
<taxonomicName id="4C107DA0FFEA0D13C438FC088D5EFC57" authority="(Lorente 2015)" baseAuthorityName="Lorente" baseAuthorityYear="2015" box="[826,1132,990,1014]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="12" pageNumber="13" phylum="Chordata" rank="order">
Astrapotheria (
<bibRefCitation id="EF817BD2FFEA0D13C4D4FC088D52FC57" author="Lorente" box="[982,1120,990,1014]" firstAuthor="Lorente" pageId="12" pageNumber="13" refId="ref44083" refString="Lorente M. Desarrollo de Modelos de Asociacion y Clasificaciones de Restos Postcraneanos Aislados de Ungulados Nativos del Paleoceno- Eoceno de America del Sur. D. Phil. Thesis, Universidad Nacional de la Plata, Argentina, 2015." type="book" year="2015">Lorente 2015</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="8BAF0623FFEA0D13C457FC288CA6FACC" blockId="12.[824,1478,144,1641]" pageId="12" pageNumber="13">
Among the different litoptern families, Soria (2001) considered that sparnotheriodontids were more closely related to the family
<taxonomicName id="4C107DA0FFEA0D13C479FBEA8D10FBF5" authorityName=", Soria" authorityYear="2001" box="[891,1058,1084,1108]" family="Anisolambdidae" pageId="12" pageNumber="13" rank="family">Anisolambdidae</taxonomicName>
due to overall dental similarities between both families, and also considering an isolated M1 (MNRJ
<date id="FFAE20E3FFEA0D13C287FB8D8A7CFB32" bridgedPair="-" pageId="12" pageNumber="13" value="1479-05">1479- V</date>
) from Itaboraí showing an intermediate anatomy between sparnotheriodontids and anisolambdids. Therefore, he created the suborder
<taxonomicName id="4C107DA0FFEA0D13C4C0FB6F8D70FB70" box="[962,1090,1209,1233]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="subOrder" subOrder="Eolitopterna">Eolitopterna</taxonomicName>
to include both families (
<tableCitation id="C6923398FFEA0D13C248FB6F8CA4FB70" box="[1354,1430,1209,1233]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="12" pageNumber="13" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table 1</tableCitation>
). In addition, Soria (2001) tentatively added
<taxonomicName id="4C107DA0FFEA0D13C3CFFB0F8A9CFAB1" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" class="Mammalia" family="Proterotheriidae" genus="Heteroglyphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="dewoletzky">
<emphasis id="B964DA31FFEA0D13C3CFFB0F8CF2FB50" box="[1229,1472,1241,1265]" italics="true" pageId="12" pageNumber="13">Heteroglyphis dewoletzky</emphasis>
Roth, 1899
</taxonomicName>
to
<taxonomicName id="4C107DA0FFEA0D13C4D3FB2E8D81FAB1" box="[977,1203,1272,1296]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
without any detailed anatomical justification; this species is known from a broken upper molar probably from Cerro del Humo,
<collectingCountry id="F30746B3FFEA0D13C3BDFAE18C15FAEE" box="[1215,1319,1335,1359]" name="Argentina" pageId="12" pageNumber="13">Argentina</collectingCountry>
. This taxon was previously referred to
<taxonomicName id="4C107DA0FFEA0D13C31AFA808DF2FACF" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1048,1216,1366,1390]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
by Simpson (1948).
</paragraph>
<paragraph id="8BAF0623FFEA0D11C457FAA088D9FD9E" blockId="12.[824,1478,144,1641]" lastBlockId="14.[128,779,144,1014]" lastPageId="14" lastPageNumber="15" pageId="12" pageNumber="13">
More recently, one genus and two new species from the Eocene La Meseta and Submeseta formations,
<collectingCountry id="F30746B3FFEA0D13C21FFA438CBBFA0C" box="[1309,1417,1429,1453]" name="Antarctica" pageId="12" pageNumber="13">Antarctica</collectingCountry>
, have been added to the family
<taxonomicName id="4C107DA0FFEA0D13C369FA628C7DFA6D" box="[1131,1359,1460,1484]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
,
<taxonomicName id="4C107DA0FFEA0D13C25DFA628DB3FA4A" authority="Bond et al. 2006" authorityName="Bond" authorityYear="2006" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="arquinotiensis">
<emphasis id="B964DA31FFEA0D13C25DFA628A8DFA4A" italics="true" pageId="12" pageNumber="13">Notiolofos arquinotiensis</emphasis>
<bibRefCitation id="EF817BD2FFEA0D13C4C9FA058DB3FA4A" author="Bond" box="[971,1153,1491,1515]" etAl="et al." firstAuthor="Bond" pageId="12" pageNumber="13" pagination="163 - 76" refId="ref40344" refString="Bond M, Reguero MA, Vizcaino SF et al. A new ' South American ungulate' (Mammalia: Litopterna) from the Eocene of the Antarctic Peninsula. Geological Society, London, Special Publications 2006; 258: 163 - 76. https: // doi. org / 10.1144 / gsl. sp. 2006.258.01.12" type="journal article" year="2006">
Bond
<emphasis id="B964DA31FFEA0D13C30CFA028D71FA4A" box="[1038,1091,1491,1515]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
2006
</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C107DA0FFEA0D13C3BFFA058D7EF9AA" authority="Gelfo, Lopez &amp; Santillana, 2017" authorityName="Gelfo, Lopez &amp; Santillana" authorityYear="2017" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="regueroi">
<emphasis id="B964DA31FFEA0D13C3BFFA058C4AFA4A" box="[1213,1400,1491,1515]" italics="true" pageId="12" pageNumber="13">Notiolofos regueroi</emphasis>
<bibRefCitation id="EF817BD2FFEA0D13C286FA058D7EF9AA" author="Gelfo, Lopez &amp; Santillana" firstAuthor="Gelfo" pageId="12" pageNumber="13" pagination="445 - 55" refId="ref42591" refString="Gelfo JN, Lopez GM, Santillana SN. Eocene ungulate mammals from West Antarctica: implications from their fossil record and a new species. Antarctic Science 2017; 29: 445 - 55. https: // doi. org / 10.1017 / s 0954102017000244" type="journal article" year="2017">Gelfo, López &amp; Santillana, 2017</bibRefCitation>
</taxonomicName>
. They are differentiated mainly by their size and represented by isolated teeth (the latter species only by a m3;
<bibRefCitation id="EF817BD2FFEA0D13C4CCF9E78D44F9E8" author="Gelfo" box="[974,1142,1585,1609]" etAl="et al." firstAuthor="Gelfo" pageId="12" pageNumber="13" pagination="274 - 92" refId="ref42638" refString="Gelfo JN, Goin FJ, Bauza N et al. The fossil record of Antarctic land mammals: commented review and hypotheses for future research. Advances in Polar Science 2019; 30: 274 - 92." type="journal article" year="2019">
Gelfo
<emphasis id="B964DA31FFEA0D13C30FF9E48D0FF9E8" box="[1037,1085,1585,1609]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
2019
</bibRefCitation>
).
<taxonomicName id="4C107DA0FFEA0D13C38FF9E78C4EF9E8" box="[1165,1404,1585,1609]" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="arquinotiensis">
<emphasis id="B964DA31FFEA0D13C38FF9E78C4EF9E8" box="[1165,1404,1585,1609]" italics="true" pageId="12" pageNumber="13">Notiolofos arquinotiensis</emphasis>
</taxonomicName>
is particularly interesting as it is the most abundant land mammal in the Palaeogene of the La Meseta and Submeseta formations, presenting a continuous fossil record of at least 10 Mya [44–34 Mya (
<bibRefCitation id="EF817BD2FFE80D11C7BDFF1988B1FF46" author="Douglas" box="[191,387,207,231]" etAl="et al." firstAuthor="Douglas" pageId="14" pageNumber="15" pagination="6582 - 7" refId="ref41674" refString="Douglas PMJ, Affek HP, Ivany LC et al. Pronounced zonal heterogeneity in Eocene southern high-latitude sea surface temperatures. Proceedings of the National Academy of Sciences of the United States of America 2014; 111: 6582 - 7. https: // doi. org / 10.1073 / pnas. 1321441111" type="journal article" year="2014">
Douglas
<emphasis id="B964DA31FFE80D11C618FF198878FF46" box="[282,330,207,231]" italics="true" pageId="14" pageNumber="15">et al.</emphasis>
2014
</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFE80D11C68DFF198B5BFF46" author="Amenabar" box="[399,617,207,231]" etAl="et al." firstAuthor="Amenabar" pageId="14" pageNumber="15" pagination="351 - 66" refId="ref39524" refString="Amenabar CR, Montes M, Nozal F et al. Dinoflagellate cysts of the La Meseta Formation (Middle to Late Eocene), Antarctic Peninsula: implications for biostratigraphy, palaeoceanography and palaeoenvironment. Geological Magazine 2020; 157: 351 - 66. https: // doi. org / 10.1017 / s 0016756819000591" type="journal article" year="2020">
Amenábar
<emphasis id="B964DA31FFE80D11C502FF198B02FF46" box="[512,560,207,231]" italics="true" pageId="14" pageNumber="15">et al.</emphasis>
2020
</bibRefCitation>
), but see Gelfo (2016) for a different age interpretation], which has been interpreted as a case of morphological stasis (Gelfo 2016). The presence of the sparnotheriodontid
<taxonomicName id="4C107DA0FFE80D11C6D6FEFB8B05FEE4" box="[468,567,301,325]" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C6D6FEFB8B05FEE4" box="[468,567,301,325]" italics="true" pageId="14" pageNumber="15">Notiolofos</emphasis>
</taxonomicName>
in
<collectingCountry id="F30746B3FFE80D11C55FFEFB8BF5FEE4" box="[605,711,301,325]" name="Antarctica" pageId="14" pageNumber="15">Antarctica</collectingCountry>
raises important biogeographic questions, as no other litoptern lineage seems to have crossed to
<collectingCountry id="F30746B3FFE80D11C6C7FEBD8B03FE22" box="[453,561,363,387]" name="Antarctica" pageId="14" pageNumber="15">Antarctica</collectingCountry>
, and among SANUs excluding
<taxonomicName id="4C107DA0FFE80D11C7EEFE5D887DFE02" box="[236,335,395,419]" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C7EEFE5D887DFE02" box="[236,335,395,419]" italics="true" pageId="14" pageNumber="15">Notiolofos</emphasis>
</taxonomicName>
, only the astrapothere
<taxonomicName id="4C107DA0FFE80D11C544FE5D8876FE63" authority="(Bond et al. 2011)" baseAuthorityName="Bond" baseAuthorityYear="2011" class="Mammalia" genus="Antarctodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="sobrali">
<emphasis id="B964DA31FFE80D11C544FE5D8A3BFE02" box="[582,777,395,419]" italics="true" pageId="14" pageNumber="15">Antarctodon sobrali</emphasis>
(
<bibRefCitation id="EF817BD2FFE80D11C789FE7C880BFE63" author="Bond" box="[139,313,426,450]" etAl="et al." firstAuthor="Bond" pageId="14" pageNumber="15" pagination="1 - 16" refId="ref40245" refString="Bond M, Kramarz A, Macphee RD, Reguero M. A new astrapothere (Mammalia, Meridiungulata) from La Meseta Formation, Seymour (Marambio) Island, and a reassessment of previous records of Antarctic astrapotheres. American Museum Novitates, 2011 (3718), 1 - 16." type="journal article" year="2011">
Bond
<emphasis id="B964DA31FFE80D11C7CEFE7D89CCFE63" box="[204,254,426,450]" italics="true" pageId="14" pageNumber="15">et al.</emphasis>
2011
</bibRefCitation>
)
</taxonomicName>
has been found thus far on this continent. However, as the preservation of mammals is allochthonous in the marine sediments of La Meseta and Submeseta formations a sampling issue cannot be disregarded for explaining the absence of other SANUs (
<bibRefCitation id="EF817BD2FFE80D11C636FDF188E8FD9E" author="Gelfo" box="[308,474,551,575]" etAl="et al." firstAuthor="Gelfo" pageId="14" pageNumber="15" pagination="274 - 92" refId="ref42638" refString="Gelfo JN, Goin FJ, Bauza N et al. The fossil record of Antarctic land mammals: commented review and hypotheses for future research. Advances in Polar Science 2019; 30: 274 - 92." type="journal article" year="2019">
Gelfo
<emphasis id="B964DA31FFE80D11C671FDFE88A9FD9E" box="[371,411,551,575]" italics="true" pageId="14" pageNumber="15">et al</emphasis>
. 2019
</bibRefCitation>
).
</paragraph>
<caption id="DF6F56ABFFEB0D11C773F92588DFF81B" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14" startId="13.[113,178,1779,1803]" targetBox="[190,1383,148,1748]" targetPageId="13" targetType="figure">
<paragraph id="8BAF0623FFEB0D12C773F9258CA7F812" blockId="13.[113,1456,1779,1971]" pageId="13" pageNumber="14">
<emphasis id="B964DA31FFEB0D12C773F92589FBF8AA" bold="true" box="[113,201,1779,1803]" pageId="13" pageNumber="14">Figure 5.</emphasis>
Right lower molars of
<taxonomicName id="4C107DA0FFEB0D12C69CF9258B31F8AA" box="[414,515,1779,1803]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="order">Litopterna</taxonomicName>
in occlusal view. A, p1–p4 of
<taxonomicName id="4C107DA0FFEB0D12C41BF9258A8DF8AA" authorityName="Roth" authorityYear="1899" box="[793,959,1779,1803]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="lechei">
<emphasis id="B964DA31FFEB0D12C41BF9258A8DF8AA" box="[793,959,1779,1803]" italics="true" pageId="13" pageNumber="14">Polymorphis lechei</emphasis>
</taxonomicName>
[holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of
<taxonomicName id="4C107DA0FFEB0D12C78BF8D98853F886" authorityName="Ameghino" authorityYear="1902" box="[137,353,1807,1831]" class="Mammalia" family="Macraucheniidae" genus="Cramauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="normalis">
<emphasis id="B964DA31FFEB0D12C78BF8D98853F886" box="[137,353,1807,1831]" italics="true" pageId="13" pageNumber="14">Cramauchenia normalis</emphasis>
</taxonomicName>
[MNHN.F.COL181 (mirrored)]. C, m3 of
<taxonomicName id="4C107DA0FFEB0D12C5FCF8D98AE4F886" authorityName="Ameghino" authorityYear="1902" box="[766,982,1807,1831]" class="Mammalia" family="Macraucheniidae" genus="Cramauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="normalis">
<emphasis id="B964DA31FFEB0D12C5FCF8D98AE4F886" box="[766,982,1807,1831]" italics="true" pageId="13" pageNumber="14">Cramauchenia normalis</emphasis>
</taxonomicName>
(AMNH-VP 29753). D, p1–m3 of
<taxonomicName id="4C107DA0FFEB0D12C221F8D989F0F8E2" authorityName="Ameghino" authorityYear="1887" class="Mammalia" family="Macraucheniidae" genus="Theosodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="lydekkeri">
<emphasis id="B964DA31FFEB0D12C221F8D989F0F8E2" italics="true" pageId="13" pageNumber="14">Theosodon lydekkeri</emphasis>
</taxonomicName>
[MACN A 24-90 (mirrored)]. E, m2–m3 of
<taxonomicName id="4C107DA0FFEB0D12C568F8FA8BC2F8E2" box="[618,752,1836,1859]" class="Mammalia" family="Adianthidae" genus="Proectocion" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B964DA31FFEB0D12C568F8FA8BE3F8E2" box="[618,721,1836,1859]" italics="true" pageId="13" pageNumber="14">Proectocion</emphasis>
sp.
</taxonomicName>
(MLP 59-II-28-107). F, p5–m3 of
<taxonomicName id="4C107DA0FFEB0D12C33BF8FD8C27F8E2" authorityName="Ameghino" authorityYear="1897" box="[1081,1301,1835,1859]" class="Mammalia" family="Adianthidae" genus="Tricoelodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="bicuspidatus">
<emphasis id="B964DA31FFEB0D12C33BF8FD8C27F8E2" box="[1081,1301,1835,1859]" italics="true" pageId="13" pageNumber="14">Tricoelodus bicuspidatus</emphasis>
</taxonomicName>
(MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of
<taxonomicName id="4C107DA0FFEB0D12C412F8918AE8F8FE" authorityName="Ameghino" authorityYear="1897" box="[784,986,1863,1887]" class="Mammalia" family="Adianthidae" genus="Proadiantus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="excavatus">
<emphasis id="B964DA31FFEB0D12C412F8918AE8F8FE" box="[784,986,1863,1887]" italics="true" pageId="13" pageNumber="14">Proadiantus excavatus</emphasis>
</taxonomicName>
(MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of
<taxonomicName id="4C107DA0FFEB0D12C78BF8B58877F8DA" authorityName="Ameghino" authorityYear="1897" box="[137,325,1891,1915]" class="Mammalia" family="Proterotheriidae" genus="Lambdaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="suinus">
<emphasis id="B964DA31FFEB0D12C78BF8B58877F8DA" box="[137,325,1891,1915]" italics="true" pageId="13" pageNumber="14">Lambdaconus suinus</emphasis>
</taxonomicName>
(MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of
<taxonomicName id="4C107DA0FFEB0D12C364F8B58C79F8DA" baseAuthorityName="Ameghino" baseAuthorityYear="1887" box="[1126,1355,1891,1915]" class="Mammalia" family="Proterotheriidae" genus="Diadiaphorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="majusculus">
<emphasis id="B964DA31FFEB0D12C364F8B58C79F8DA" box="[1126,1355,1891,1915]" italics="true" pageId="13" pageNumber="14">Diadiaphorus majusculus</emphasis>
</taxonomicName>
(MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of
<taxonomicName id="4C107DA0FFEB0D12C574F8A98A09F836" authorityName="Ameghino" authorityYear="1901" box="[630,827,1919,1943]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="fissidens">
<emphasis id="B964DA31FFEB0D12C574F8A98A09F836" box="[630,827,1919,1943]" italics="true" pageId="13" pageNumber="14">Anisolambda fissidens</emphasis>
</taxonomicName>
(MACN A 10668; holotype). K, m2–m3 of
<taxonomicName id="4C107DA0FFEB0D12C3DDF8A989E0F812" authorityName="Cifelli" authorityYear="1983" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="prodromus">
<emphasis id="B964DA31FFEB0D12C3DDF8A989E0F812" italics="true" pageId="13" pageNumber="14">Paranisolambda prodromus</emphasis>
</taxonomicName>
(cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of
<taxonomicName id="4C107DA0FFEB0D12C4FFF84D8DC3F812" authorityName="Paula Couto" authorityYear="1952" box="[1021,1265,1947,1971]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="prototypica">
<emphasis id="B964DA31FFEB0D12C4FFF84D8DC3F812" box="[1021,1265,1947,1971]" italics="true" pageId="13" pageNumber="14">Victorlemoinea prototypica</emphasis>
</taxonomicName>
(MNRJ 1482V).
</paragraph>
<paragraph id="8BAF0623FFE80D11C79EFD91880BFCF8" blockId="14.[128,779,144,1014]" pageId="14" pageNumber="15">
Apart from early phylogenetic studies (
<bibRefCitation id="EF817BD2FFE80D11C525FD918BACFDFE" author="Cifelli" box="[551,670,583,607]" firstAuthor="Cifelli" pageId="14" pageNumber="15" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFE80D11C5AAFD918987FDDF" author="Bergqvist" firstAuthor="Bergqvist" pageId="14" pageNumber="15" refId="ref39790" refString="Bergqvist LP. Reasociacao de pos-cranio as especies de ungulados da Bacia de S. J. Itaborai (Paleoceno), Estado do Rio de Janeiro, e Filogenia dos ' Condylarthra', e ungulados Sul-Americanos com base no poscranio. D. Phil. Thesis, Universidade Federal do Rio Grande do Sul, Brazil, 1996." type="book" year="1996">Bergqvist 1996</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFE80D11C7C1FDB088C7FDDF" author="Bonaparte and Morales" box="[195,501,614,638]" firstAuthor="Bonaparte" pageId="14" pageNumber="15" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales 1997</bibRefCitation>
;
<figureCitation id="132B1AA6FFE80D11C507FDB08B46FDDF" box="[517,628,614,638]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="14" pageNumber="15">Fig. 1C–D</figureCitation>
), most recent phylogenetic studies have included one sparnotheriodontid species among a wide sample of different SANUs (e.g.,
<bibRefCitation id="EF817BD2FFE80D11C5D2FD7389C3FD7D" author="Gelfo" etAl="et al." firstAuthor="Gelfo" pageId="14" pageNumber="15" pagination="329 - 35" refId="ref42504" refString="Gelfo JN, Lopez GM, Bond M. A new Xenungulata (Mammalia) from the Paleocene of Patagonia, Argentina. Journal of Paleontology 2008; 82: 329 - 35. https: // doi. org / 10.1666 / 06 - 099.1" type="journal article" year="2008">
Gelfo
<emphasis id="B964DA31FFE80D11C783FD138986FD7D" box="[129,180,708,732]" italics="true" pageId="14" pageNumber="15">et al.</emphasis>
2008
</bibRefCitation>
;
<tableCitation id="C6923398FFE80D11C601FD128860FD7D" box="[259,338,708,732]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" 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="14" pageNumber="15" tableUuid="DF6F56ABFFE20D1BC76CFE6E8C76FDA9">Table 2</tableCitation>
), instead of testing their interfamilial relationships. In terms of the phylogenetic relationships within
<taxonomicName id="4C107DA0FFE80D11C783FCD58857FCBA" box="[129,357,771,795]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
, there is currently only one phylogenetic study that used a small taxon and character sample (Reguero
<emphasis id="B964DA31FFE80D11C5FAFCF589ABFCF8" italics="true" pageId="14" pageNumber="15">et al.</emphasis>
2014;
<tableCitation id="C6923398FFE80D11C7DCFC97881AFCF8" box="[222,296,833,857]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" 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="14" pageNumber="15" tableUuid="DF6F56ABFFE20D1BC76CFE6E8C76FDA9">Table 2</tableCitation>
).
</paragraph>
<paragraph id="8BAF0623FFE80D11C79EFCB78B0BFC57" blockId="14.[128,779,144,1014]" pageId="14" pageNumber="15">
The earliest accepted member of
<taxonomicName id="4C107DA0FFE80D11C509FCB78BDFFCD8" box="[523,749,865,889]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
is probably the Early Eocene
<taxonomicName id="4C107DA0FFE80D11C68CFC578B26FC39" authorityName="Paula Couto" authorityYear="1952" box="[398,532,897,920]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="prototypica">
<emphasis id="B964DA31FFE80D11C68CFC578B26FC39" box="[398,532,897,920]" italics="true" pageId="14" pageNumber="15">V. prototypica</emphasis>
</taxonomicName>
with records in Itaboraí,
<collectingCountry id="F30746B3FFE80D11C782FC49898FFC16" box="[128,189,927,951]" name="Brazil" pageId="14" pageNumber="15">Brazil</collectingCountry>
, and the last could be
<taxonomicName id="4C107DA0FFE80D11C6ACFC498B23FC16" box="[430,529,927,951]" class="Mammalia" family="Sparnotheriodontidae" genus="Notiolofos" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C6ACFC498B23FC16" box="[430,529,927,951]" italics="true" pageId="14" pageNumber="15">Notiolofos</emphasis>
</taxonomicName>
from
<collectingCountry id="F30746B3FFE80D11C551FC498B8DFC16" box="[595,703,927,951]" name="Antarctica" pageId="14" pageNumber="15">Antarctica</collectingCountry>
, giving a temporal interval of 56.0–34.0 Mya to this family (
<figureCitation id="132B1AA6FFE80D11C5B4FC698A33FC76" box="[694,769,959,983]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="14" pageNumber="15">Fig. 2B</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFE80D11C67DFC0888E5FC57" box="[383,471,990,1014]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="14" pageNumber="15" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table S1</tableCitation>
, File S2).
</paragraph>
<emphasis id="B964DA31FFE80D11C605FBC38BB1FB91" box="[263,643,1045,1072]" italics="true" pageId="14" pageNumber="15">
<subSubSection id="C30A55A8FFE80D11C605FBC38892FB8E" box="[263,416,1045,1071]" pageId="14" pageNumber="15" type="nomenclature">
<paragraph id="8BAF0623FFE80D11C605FBC38892FB8E" blockId="14.[129,779,1045,1766]" box="[263,416,1045,1071]" pageId="14" pageNumber="15">
<taxonomicName id="4C107DA0FFE80D11C605FBC38892FB8E" authorityName=", Soria" authorityYear="2001" box="[263,416,1045,1071]" family="Anisolambdidae" kingdom="Animalia" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="C30A55A8FFE80D11C6A6FBC08BB1FB91" box="[420,643,1046,1072]" pageId="14" pageNumber="15" type="description">
<paragraph id="8BAF0623FFE80D11C6A6FBC08BB1FB91" blockId="14.[129,779,1045,1766]" box="[420,643,1046,1072]" pageId="14" pageNumber="15">
(
<figureCitation id="132B1AA6FFE80D11C6ACFBC08B2CFB91" box="[430,542,1046,1072]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="14" pageNumber="15">Figs 4M–O</figureCitation>
,
<figureCitation id="132B1AA6FFE80D11C525FBC08B4BFB8E" box="[551,633,1046,1072]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="14" pageNumber="15">5J–K, N</figureCitation>
)
</paragraph>
</subSubSection>
</emphasis>
<subSubSection id="C30A55A8FFE80D11C783FBEA8ACFF947" pageId="14" pageNumber="15" type="multiple">
<paragraph id="8BAF0623FFE80D11C783FBEA8DCFFDFE" blockId="14.[129,779,1045,1766]" lastBlockId="14.[825,1475,144,1766]" pageId="14" pageNumber="15">
<taxonomicName id="4C107DA0FFE80D11C783FBEA881CFBF5" authorityName=", Soria" authorityYear="2001" box="[129,302,1084,1108]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName>
is currently represented by eight genera with mostly a Palaeogene distribution (
<figureCitation id="132B1AA6FFE80D11C53CFB8D8B4CFBD2" box="[574,638,1115,1139]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="14" pageNumber="15">Fig. 2</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFE80D11C614FBAD884FFB32" box="[278,381,1147,1171]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="14" pageNumber="15" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table S1</tableCitation>
). The family
<taxonomicName id="4C107DA0FFE80D11C524FBAD8BE1FB32" authorityName=", Soria" authorityYear="2001" box="[550,723,1147,1171]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName>
was proposed by Soria (2001), who elevated the rank of the subfamily
<taxonomicName id="4C107DA0FFE80D11C7F4FB6F8897FB70" box="[246,421,1209,1233]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Anisolambdinae">Anisolambdinae</taxonomicName>
. This subfamily was previously proposed by Cifelli (1983a) to group what he considered early members of the family
<taxonomicName id="4C107DA0FFE80D11C6C4FB2E8B47FAB1" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[454,629,1272,1296]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
with records in the Early Eocene such as
<taxonomicName id="4C107DA0FFE80D11C6C7FAC18B78FA8E" authorityName="Ameghino" authorityYear="1901" box="[453,586,1303,1327]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C6C7FAC18B78FA8E" box="[453,586,1303,1327]" italics="true" pageId="14" pageNumber="15">Anisolambda</emphasis>
</taxonomicName>
, distinctive from proterotheriids with more derived features (subfamily
<taxonomicName id="4C107DA0FFE80D11C783FA808801FACF" box="[129,307,1366,1390]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Proterotheriinae">Proterotheriinae</taxonomicName>
;
<tableCitation id="C6923398FFE80D11C641FA8088A4FACF" box="[323,406,1366,1390]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="14" pageNumber="15" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table 1</tableCitation>
).
<taxonomicName id="4C107DA0FFE80D11C6B6FA808B53FACF" box="[436,609,1366,1390]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Anisolambdinae">Anisolambdinae</taxonomicName>
stands out for the presence of a strong paralophid ending in a large and distinct paraconid.
<taxonomicName id="4C107DA0FFE80D11C62CFA4388E9FA0C" box="[302,475,1429,1453]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Anisolambdinae">Anisolambdinae</taxonomicName>
(currently
<taxonomicName id="4C107DA0FFE80D11C550FA438A30FA0C" authorityName=", Soria" authorityYear="2001" box="[594,770,1429,1453]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName>
) and
<taxonomicName id="4C107DA0FFE80D11C7BAFA628855FA6D" box="[184,359,1460,1484]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Proterotheriinae">Proterotheriinae</taxonomicName>
(currently
<taxonomicName id="4C107DA0FFE80D11C6F2FA628B90FA6D" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[496,674,1460,1484]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
) share a molarized penultimate premolar or P4 (P
<quantity id="4CE8ABC6FFE80D11C537FA058B6FFA4A" box="[565,605,1491,1515]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="14" pageNumber="15" unit="in" value="3.0">3 in</quantity>
Cifelli (1983a); see Material and methods section for tooth position criteria) with a well-developed metacone and mesostyle present, among other features. To a certain extent, Cifelli’s (1983a) proposal was similar to previous arrangements (Simpson 1945, 1948). However, Simpson (1945, 1948) included anisolambdids in the subfamily
<taxonomicName id="4C107DA0FFE80D11C545F9598BDEF906" box="[583,748,1679,1703]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Polymorphinae">Polymorphinae</taxonomicName>
, a subfamily that also included the Late Eocene
<taxonomicName id="4C107DA0FFE80D11C570F9798BC2F966" authorityName="Roth" authorityYear="1899" box="[626,752,1711,1735]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C570F9798BC2F966" box="[626,752,1711,1735]" italics="true" pageId="14" pageNumber="15">Polymorphis</emphasis>
</taxonomicName>
, a taxon later included in
<taxonomicName id="4C107DA0FFE80D11C674F9188B1EF947" box="[374,556,1742,1766]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName>
(Cifelli 1983a, Soria 2001). In addition, Cifelli (1983a) renamed an anisolambdid from the Early Eocene of Itaboraí,
<collectingCountry id="F30746B3FFE80D11C3B4FF798DC7FF66" box="[1206,1269,175,199]" name="Brazil" pageId="14" pageNumber="15">Brazil</collectingCountry>
, as
<taxonomicName id="4C107DA0FFE80D11C21CFF798D0FFF46" authority="Cifelli 1983 a" authorityName="Cifelli" authorityYear="1983" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="prodromus">
<emphasis id="B964DA31FFE80D11C21CFF798A97FF46" italics="true" pageId="14" pageNumber="15">Paranisolambda prodromus</emphasis>
Cifelli 1983a
</taxonomicName>
(previously known as
<taxonomicName id="4C107DA0FFE80D11C23FFF198D47FEA7" authority="Paula Couto, 1952" authorityName="Paula Couto" authorityYear="1952" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="prodromus">
<emphasis id="B964DA31FFE80D11C23FFF198A97FEA7" italics="true" pageId="14" pageNumber="15">Anisolambda prodromus</emphasis>
Paula Couto, 1952
</taxonomicName>
), based on some dental differences with
<taxonomicName id="4C107DA0FFE80D11C4B6FEDB8D0BFE84" authorityName="Ameghino" authorityYear="1901" box="[948,1081,269,293]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C4B6FEDB8D0BFE84" box="[948,1081,269,293]" italics="true" pageId="14" pageNumber="15">Anisolambda</emphasis>
</taxonomicName>
such as the absence of a postcristid in m3. Based on previous observations (Simpson 1948, Paula Couto 1952) and comparisons with
<taxonomicName id="4C107DA0FFE80D11C3CFFE9A8AA2FE22" authority=", Cifelli (1983 a)" authorityName=", Cifelli" authorityYear="1983" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C3CFFE9A8C43FEC5" box="[1229,1393,332,356]" italics="true" pageId="14" pageNumber="15">Paranisolambda</emphasis>
, Cifelli (1983a)
</taxonomicName>
synonymized the anisolambdids
<taxonomicName id="4C107DA0FFE80D11C20AFEBD8CBDFE22" authorityName="Ameghino" authorityYear="1901" box="[1288,1423,363,387]" class="Mammalia" family="Proterotheriidae" genus="Josepholeidya" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C20AFEBD8CBDFE22" box="[1288,1423,363,387]" italics="true" pageId="14" pageNumber="15">Josepholeidya</emphasis>
</taxonomicName>
and
<taxonomicName id="4C107DA0FFE80D11C43BFE5D8ADDFE02" authorityName="Ameghino" authorityYear="1901" box="[825,1007,395,419]" class="Mammalia" family="Proterotheriidae" genus="Ricardolydekkeria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C43BFE5D8ADDFE02" box="[825,1007,395,419]" italics="true" pageId="14" pageNumber="15">Ricardolydekkeria</emphasis>
</taxonomicName>
, taxa based on isolated upper teeth, with
<taxonomicName id="4C107DA0FFE80D11C43BFE7C8A8CFE63" authorityName="Ameghino" authorityYear="1901" box="[825,958,426,450]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C43BFE7C8A8CFE63" box="[825,958,426,450]" italics="true" pageId="14" pageNumber="15">Anisolambda</emphasis>
</taxonomicName>
, which was based only on lower dentition. In addition, Cifelli (1983a) included
<taxonomicName id="4C107DA0FFE80D11C3CDFE1F8C4CFE40" authorityName="Simpson" authorityYear="1935" box="[1231,1406,457,481]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="tshotshe">
<emphasis id="B964DA31FFE80D11C3CDFE1F8C4CFE40" box="[1231,1406,457,481]" italics="true" pageId="14" pageNumber="15">Wainka tshotshe</emphasis>
</taxonomicName>
from Cerro Redondo,
<collectingCountry id="F30746B3FFE80D11C4EFFE3F8D6AFDA0" box="[1005,1112,489,513]" name="Argentina" pageId="14" pageNumber="15">Argentina</collectingCountry>
, in the subfamily
<taxonomicName id="4C107DA0FFE80D11C214FE3F8CF0FDA0" authorityName=", Soria" authorityYear="2001" box="[1302,1474,489,513]" pageId="14" pageNumber="15" rank="subFamily" subFamily="Anisolambdidae">Anisolambdidae</taxonomicName>
based on an m3 with an overall structure very similar to
<taxonomicName id="4C107DA0FFE80D11C43BFDF18A8CFD9E" authorityName="Ameghino" authorityYear="1901" box="[825,958,551,575]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C43BFDF18A8CFD9E" box="[825,958,551,575]" italics="true" pageId="14" pageNumber="15">Anisolambda</emphasis>
</taxonomicName>
.
<taxonomicName id="4C107DA0FFE80D11C4CFFDF18D46FD9E" authorityName="Simpson" authorityYear="1935" box="[973,1140,551,575]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="tshotshe">
<emphasis id="B964DA31FFE80D11C4CFFDF18D46FD9E" box="[973,1140,551,575]" italics="true" pageId="14" pageNumber="15">Wainka tshotshe</emphasis>
</taxonomicName>
was previously assigned tentatively to
<taxonomicName id="4C107DA0FFE80D11C496FD918DCAFDFE" authority="(Simpson 1948)" baseAuthorityName="Simpson" baseAuthorityYear="1948" box="[916,1272,583,607]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae (Simpson 1948)</taxonomicName>
.
</paragraph>
<paragraph id="8BAF0623FFE80D11C457FDB08AA9FB13" blockId="14.[825,1475,144,1766]" pageId="14" pageNumber="15">
Apart from elevating the rank of this subfamily to the family
<taxonomicName id="4C107DA0FFE80D11C439FD538D5DFD3C" authority=", Soria (2001)" authorityName=", Soria" authorityYear="2001" box="[827,1135,645,669]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae, Soria (2001)</taxonomicName>
mostly followed Cifelli’s (1983a) interpretations. However, Soria (2001) removed
<taxonomicName id="4C107DA0FFE80D11C25FFD738CF0FD1C" authorityName="Berg" authorityYear="1899" box="[1373,1474,677,701]" class="Mammalia" family="Proterotheriidae" genus="Xesmodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C25FFD738CF0FD1C" box="[1373,1474,677,701]" italics="true" pageId="14" pageNumber="15">Xesmodon</emphasis>
</taxonomicName>
and
<taxonomicName id="4C107DA0FFE80D11C46BFD128ADCFD7D" authorityName="Roth" authorityYear="1899" box="[873,1006,708,732]" class="Mammalia" family="Proterotheriidae" genus="Heteroglyphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C46BFD128ADCFD7D" box="[873,1006,708,732]" italics="true" pageId="14" pageNumber="15">Heteroglyphis</emphasis>
</taxonomicName>
from the family, tentatively placing them in
<taxonomicName id="4C107DA0FFE80D11C43BFD358AEFFD5A" box="[825,989,739,763]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Didolodontidae</taxonomicName>
and
<taxonomicName id="4C107DA0FFE80D11C30FFD358DC3FD5A" box="[1037,1265,739,763]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
, respectively, as they showed important differences in their dentition compared to other anisolambdids. He also added
<taxonomicName id="4C107DA0FFE80D11C3B1FCF48CF0FC9B" authority="Soria, 2001" authorityName="Soria" authorityYear="2001" box="[1203,1474,802,826]" class="Mammalia" family="Proterotheriidae" genus="Lambdaconops" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C3B1FCF48C75FC9B" box="[1203,1351,802,826]" italics="true" pageId="14" pageNumber="15">Lambdaconops</emphasis>
Soria, 2001
</taxonomicName>
to
<taxonomicName id="4C107DA0FFE80D11C456FC978AC9FCF8" authorityName=", Soria" authorityYear="2001" box="[852,1019,833,857]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName>
due to close dental similarities with the Late Oligocene (Deseadan SALMA) anisolambdid
<taxonomicName id="4C107DA0FFE80D11C23CFCB78D70FC39" authority="Ameghino, 1897" authorityName="Ameghino" authorityYear="1897" class="Mammalia" family="Proterotheriidae" genus="Protheosodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="coniferus">
<emphasis id="B964DA31FFE80D11C23CFCB78ABDFC39" italics="true" pageId="14" pageNumber="15">Protheosodon coniferus</emphasis>
<bibRefCitation id="EF817BD2FFE80D11C494FC568D70FC39" author="Ameghino" box="[918,1090,896,920]" firstAuthor="Ameghino" pageId="14" pageNumber="15" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino, 1897</bibRefCitation>
</taxonomicName>
, the latter being the only member of this family with associated tarsal remains (
<bibRefCitation id="EF817BD2FFE80D11C3F5FC498CB0FC16" author="Loomis" box="[1271,1410,927,951]" firstAuthor="Loomis" pageId="14" pageNumber="15" refId="ref44004" refString="Loomis FB. The Deseado Formation of Patagonia. Amherst, MA: The Trustees of Amherst College, 1914." type="book" year="1914">Loomis 1914</bibRefCitation>
). The validity of the different species of
<taxonomicName id="4C107DA0FFE80D11C3A3FC698C10FC76" authorityName="Ameghino" authorityYear="1901" box="[1185,1314,959,983]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFE80D11C3A3FC698C10FC76" box="[1185,1314,959,983]" italics="true" pageId="14" pageNumber="15">Anisolambda</emphasis>
</taxonomicName>
was not evaluated by Soria (2001). In addition, Soria (2001) erected the suborder
<taxonomicName id="4C107DA0FFE80D11C475FC2B8AC8FBB4" box="[887,1018,1021,1045]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="subOrder" subOrder="Eolitopterna">Eolitopterna</taxonomicName>
, to include the families
<taxonomicName id="4C107DA0FFE80D11C3ECFC2B8CA7FBB4" authorityName=", Soria" authorityYear="2001" box="[1262,1429,1021,1045]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName>
and
<taxonomicName id="4C107DA0FFE80D11C438FBCB8D2EFB94" box="[826,1052,1053,1077]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
due to overall dental similarities between both families (more details in
<taxonomicName id="4C107DA0FFE80D11C387FBEA8C55FBF5" box="[1157,1383,1084,1108]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
section;
<tableCitation id="C6923398FFE80D11C43BFB8D8AB1FBD2" box="[825,899,1115,1139]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="14" pageNumber="15" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table 1</tableCitation>
). Most recent authors have followed Soria’s (2001) taxonomic proposal for
<taxonomicName id="4C107DA0FFE80D11C30AFBAD8D9DFB32" authorityName=", Soria" authorityYear="2001" box="[1032,1199,1147,1171]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName>
(e.g.,
<bibRefCitation id="EF817BD2FFE80D11C3F2FBAD8C57FB32" author="Gelfo" box="[1264,1381,1147,1171]" firstAuthor="Gelfo" pageId="14" pageNumber="15" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo 2006</bibRefCitation>
, Croft
<emphasis id="B964DA31FFE80D11C2B3FBAD8A60FB13" italics="true" pageId="14" pageNumber="15">et al.</emphasis>
2020).
</paragraph>
<paragraph id="8BAF0623FFE80D11C457FB6F8DA3F98B" blockId="14.[825,1475,144,1766]" pageId="14" pageNumber="15">
Early studies examining the interfamilial relationships of
<taxonomicName id="4C107DA0FFE80D11C43BFB0F8A94FB50" box="[825,934,1241,1265]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="order">Litopterna</taxonomicName>
assumed a monophyletic
<taxonomicName id="4C107DA0FFE80D11C3C3FB0F8C5BFB50" baseAuthorityName="Simpson" baseAuthorityYear="1948" box="[1217,1385,1241,1265]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
that included anisolambdids, without testing this hypothesis (
<bibRefCitation id="EF817BD2FFE80D11C281FB2E8A5FFA8E" author="Cifelli" firstAuthor="Cifelli" pageId="14" pageNumber="15" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFE80D11C47AFACE8D93FA8E" author="Bonaparte and Morales" box="[888,1185,1303,1327]" firstAuthor="Bonaparte" pageId="14" pageNumber="15" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales 1997</bibRefCitation>
;
<figureCitation id="132B1AA6FFE80D11C3ACFAC18C2BFA8E" box="[1198,1305,1303,1327]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="14" pageNumber="15">Fig. 1C–D</figureCitation>
). More recently, in phylogenetic studies evaluating phylogenetic affinities within
<taxonomicName id="4C107DA0FFE80D11C438FA808AD0FACF" baseAuthorityName="Simpson" baseAuthorityYear="1948" box="[826,994,1366,1390]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
including some anisolambdids as outgroups, anisolambdids were recovered nested within
<taxonomicName id="4C107DA0FFE80D11C218FAA38CF0FA2C" baseAuthorityName="Simpson" baseAuthorityYear="1948" box="[1306,1474,1397,1421]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
(e.g.,
<bibRefCitation id="EF817BD2FFE80D11C46DFA438D77FA0C" author="McGrath" box="[879,1093,1429,1453]" etAl="et al." firstAuthor="McGrath" pageId="14" pageNumber="15" pagination="717 - 38" refId="ref44408" refString="McGrath AJ, Flynn JJ, Wyss AR. Proterotheriids and macraucheniids (Litopterna: Mammalia) from the Pampa Castillo Fauna, Chile (early Miocene, Santacrucian SALMA) and a new phylogeny of Proterotheriidae. Journal of Systematic Palaeontology 2020 b; 18: 717 - 38. https: // doi. org / 10.1080 / 14772019.2019.1662500" type="journal article" year="2020" yearSuffix="b">
McGrath
<emphasis id="B964DA31FFE80D11C4D0FA438D32FA0C" box="[978,1024,1429,1453]" italics="true" pageId="14" pageNumber="15">et al.</emphasis>
2020b
</bibRefCitation>
;
<tableCitation id="C6923398FFE80D11C352FA438DA8FA0C" box="[1104,1178,1429,1453]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" 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="14" pageNumber="15" tableUuid="DF6F56ABFFE20D1BC76CFE6E8C76FDA9">Table 2</tableCitation>
). However, as not all families of litopterns were included in these analyses, the interfamilial affinities of anisolambdids remain to be tested. In addition, to this date there has been no study examining the phylogenetic relationships within
<taxonomicName id="4C107DA0FFE80D11C4E6F9C48DBFF98B" authorityName=", Soria" authorityYear="2001" box="[996,1165,1554,1578]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName>
.
</paragraph>
<paragraph id="8BAF0623FFE80D11C457F9E78ACFF947" blockId="14.[825,1475,144,1766]" pageId="14" pageNumber="15">
The earliest accepted member of
<taxonomicName id="4C107DA0FFE80D11C3ADF9E78C64F9E8" authorityName=", Soria" authorityYear="2001" box="[1199,1366,1585,1609]" family="Anisolambdidae" pageId="14" pageNumber="15" rank="family">Anisolambdidae</taxonomicName>
is
<taxonomicName id="4C107DA0FFE80D11C274F9E78AB4F9C8" authorityName="Simpson" authorityYear="1935" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="tshotshe">
<emphasis id="B964DA31FFE80D11C274F9E78AB4F9C8" italics="true" pageId="14" pageNumber="15">Wainka tshotshe</emphasis>
</taxonomicName>
from Cerro Redondo,
<collectingRegion id="49D4C8C1FFE80D11C36CF9878DF2F9C8" box="[1134,1216,1617,1641]" country="Argentina" name="Chubut" pageId="14" pageNumber="15">Chubut</collectingRegion>
,
<collectingCountry id="F30746B3FFE80D11C3CAF9878C02F9C8" box="[1224,1328,1617,1641]" name="Argentina" pageId="14" pageNumber="15">Argentina</collectingCountry>
, and the last is
<taxonomicName id="4C107DA0FFE80D11C43BF9A68D2BF929" authorityName="Ameghino" authorityYear="1897" box="[825,1049,1648,1672]" class="Mammalia" family="Proterotheriidae" genus="Protheosodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="coniferus">
<emphasis id="B964DA31FFE80D11C43BF9A68D2BF929" box="[825,1049,1648,1672]" italics="true" pageId="14" pageNumber="15">Protheosodon coniferus</emphasis>
</taxonomicName>
from Cabeza Blanca,
<collectingRegion id="49D4C8C1FFE80D11C3F9F9A68C7FF929" box="[1275,1357,1648,1672]" country="Argentina" name="Chubut" pageId="14" pageNumber="15">Chubut</collectingRegion>
,
<collectingCountry id="F30746B3FFE80D11C255F9A68C8DF929" box="[1367,1471,1648,1672]" name="Argentina" pageId="14" pageNumber="15">Argentina</collectingCountry>
, giving a temporal interval of 62.5–26.0 Mya for this family (
<figureCitation id="132B1AA6FFE80D11C29EF9598A64F966" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="14" pageNumber="15">Fig. 2B</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFE80D11C361F9798D8FF966" box="[1123,1213,1711,1735]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="14" pageNumber="15" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table S1</tableCitation>
; Woodburne
<emphasis id="B964DA31FFE80D11C24BF9798C4AF966" box="[1353,1400,1711,1735]" italics="true" pageId="14" pageNumber="15">et al.</emphasis>
2014a,
<bibRefCitation id="EF817BD2FFE80D11C43BF9188ADFF947" author="Krause" box="[825,1005,1742,1766]" etAl="et al." firstAuthor="Krause" pageId="14" pageNumber="15" pagination="886 - 903" refId="ref43716" refString="Krause JM, Clyde WC, Ibanez-Mejia M et al. New age constraints for Early Paleogene strata of central Patagonia, Argentina: implications for the timing of South American Land Mammal Ages. Geological Society of America Bulletin 2017; 129: 886 - 903. https: // doi. org / 10.1130 / b 31561.1" type="journal article" year="2017">
Krause
<emphasis id="B964DA31FFE80D11C484F9198A86F947" box="[902,948,1742,1766]" italics="true" pageId="14" pageNumber="15">et al.</emphasis>
2017
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).
</paragraph>
</subSubSection>
<paragraph id="8BAF0623FFE80D11C783F8E58CA3F8C6" blockId="14.[129,1463,1843,1979]" pageId="14" pageNumber="15">
M, p1–m3 of
<taxonomicName id="4C107DA0FFE80D11C603F8E58B3AF8EA" authorityName="Soria" authorityYear="1980" box="[257,520,1843,1867]" class="Mammalia" family="Sparnotheriodontidae" genus="Sparnotheriodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="epsilonoides">
<emphasis id="B964DA31FFE80D11C603F8E58B3AF8EA" box="[257,520,1843,1867]" italics="true" pageId="14" pageNumber="15">Sparnotheriodon epsilonoides</emphasis>
</taxonomicName>
(MACN 18225;
<typeStatus id="54ABB881FFE80D11C5AEF8E58A35F8EA" box="[684,775,1843,1867]" pageId="14" pageNumber="15" type="holotype">holotype</typeStatus>
). N, O, m3 of
<taxonomicName id="4C107DA0FFE80D11C48BF8E58D2EF8EA" authorityName="Simpson" authorityYear="1935" box="[905,1052,1843,1867]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="tshotshe">
<emphasis id="B964DA31FFE80D11C48BF8E58D2EF8EA" box="[905,1052,1843,1867]" italics="true" pageId="14" pageNumber="15">Wainka tshotshe</emphasis>
</taxonomicName>
? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File
</paragraph>
<paragraph id="8BAF0623FFE80D11C783F8BD88DFF81B" blockId="14.[129,1463,1843,1979]" pageId="14" pageNumber="15">
S2. For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid. Scale bars equal
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.
</paragraph>
</caption>
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</treatment>
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<taxonomicName id="4C107DA0FFEC0D15C62AFAC688AEFA8B" authority="Ameghino, 1891" box="[296,412,1296,1322]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
(
<figureCitation id="132B1AA6FFEC0D15C6A9FAC78B20FA8B" box="[427,530,1296,1322]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="10" pageNumber="11">Figs 4F–H</figureCitation>
,
<figureCitation id="132B1AA6FFEC0D15C519FAC78B6AFA8A" box="[539,600,1297,1323]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="10" pageNumber="11">5E–G</figureCitation>
)
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C30A55A8FFEC0D13C783FAE189D8FE40" lastPageId="12" lastPageNumber="13" pageId="10" pageNumber="11" type="multiple">
<paragraph id="8BAF0623FFEC0D15C783FAE18C3EFEC2" blockId="10.[129,779,1296,1735]" lastBlockId="10.[823,1476,144,1735]" pageId="10" pageNumber="11">
<taxonomicName id="4C107DA0FFEC0D15C783FAE189CCFAEE" box="[129,254,1335,1359]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
currently includes six genera, being considerably less diverse than proterotheriids and macraucheniids during the Neogene (
<figureCitation id="132B1AA6FFEC0D15C7EFFAA38817FA2C" box="[237,293,1397,1421]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="10" pageNumber="11">Fig. 2</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFEC0D15C533FAA38BB9FA2C" box="[561,651,1397,1421]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="10" pageNumber="11" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table S1</tableCitation>
).
<bibRefCitation id="EF817BD2FFEC0D15C59DFAA38823FA0C" author="Cifelli and Soria" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria (1983a)</bibRefCitation>
presented a detailed revision of this family, so to avoid unnecessary repetition, only the most relevant taxonomic aspects are discussed here, along with the main advances since their revision. Known as the pygmy litopterns because of their reduced size, the family
<taxonomicName id="4C107DA0FFEC0D15C67DF9C488CEF98B" box="[383,508,1554,1578]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
was named by
<bibRefCitation id="EF817BD2FFEC0D15C59FF9C489E7F9E8" author="Ameghino" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="129 - 67" refId="ref39125" refString="Ameghino F. Caracteres diagnosticos de cincuenta especies nuevas de mamiferos fosiles argentinos. Revista Argentina de Historia Natural 1891 a; 1: 129 - 67." type="journal article" year="1891" yearSuffix="a">Ameghino (1891a)</bibRefCitation>
to place the Early Miocene
<taxonomicName id="4C107DA0FFEC0D15C541F9E78804F9C8" authority="Ameghino 1891 a" authorityName="Ameghino" authorityYear="1891" class="Mammalia" family="Adianthidae" genus="Adianthus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="bucatus">
<emphasis id="B964DA31FFEC0D15C541F9E78A38F9E8" box="[579,778,1585,1609]" italics="true" pageId="10" pageNumber="11">Adianthus bucatus</emphasis>
<bibRefCitation id="EF817BD2FFEC0D15C783F9878804F9C8" author="Ameghino" box="[129,310,1617,1641]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="129 - 67" refId="ref39125" refString="Ameghino F. Caracteres diagnosticos de cincuenta especies nuevas de mamiferos fosiles argentinos. Revista Argentina de Historia Natural 1891 a; 1: 129 - 67." type="journal article" year="1891" yearSuffix="a">Ameghino 1891a</bibRefCitation>
</taxonomicName>
from the Santa Cruz Formation,
<collectingCountry id="F30746B3FFEC0D15C5A6F9878A3BF9C8" box="[676,777,1617,1641]" name="Argentina" pageId="10" pageNumber="11">Argentina</collectingCountry>
(
<bibRefCitation id="EF817BD2FFEC0D15C789F9A68847F929" author="Cifelli and Soria" box="[139,373,1648,1672]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria 1983a</bibRefCitation>
). Among other distinct anatomical features, adianthids present a trilobed m3 and enamel fossettes in the P4–M3 formed by hypertrophied conular cristae (
<bibRefCitation id="EF817BD2FFEC0D15C5C8F9798AD0FF09" author="Cifelli and Soria" box="[714,994,144,1735]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria 1983a</bibRefCitation>
). Following new fossil discoveries,
<bibRefCitation id="EF817BD2FFEC0D15C254FF468AB1FF66" author="Ameghino" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="1 - 568" refId="ref39456" refString="Ameghino F. Les formations sedimentaires du cretace superieur et du tertiaire de Patagonie avec un parallele entre leurs faunes mammalogiques et celles de l'ancien continent. Anales del Museo Nacional de Buenos Aires 1906; 15: 1 - 568." type="journal article" year="1906">Ameghino (1906)</bibRefCitation>
added three additional genera to the family
<taxonomicName id="4C107DA0FFEC0D15C23FFF798C8DFF66" box="[1341,1471,175,199]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
:
<taxonomicName id="4C107DA0FFEC0D15C43BFF198D54FF46" authority="Ameghino, 1897" authorityName="Ameghino" authorityYear="1897" box="[825,1126,207,231]" class="Mammalia" family="Adianthidae" genus="Proadiantus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEC0D15C43BFF198A82FF46" box="[825,944,207,231]" italics="true" pageId="10" pageNumber="11">Proadiantus</emphasis>
<bibRefCitation id="EF817BD2FFEC0D15C4BAFF198D54FF46" author="Ameghino" box="[952,1126,207,231]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino, 1897</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C107DA0FFEC0D15C39FFF198CF0FF46" authority="Ameghino, 1897" authorityName="Ameghino" authorityYear="1897" box="[1181,1474,207,231]" class="Mammalia" family="Adianthidae" genus="Tricoelodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEC0D15C39FFF198C3EFF46" box="[1181,1292,207,231]" italics="true" pageId="10" pageNumber="11">Tricoelodus</emphasis>
<bibRefCitation id="EF817BD2FFEC0D15C217FF198CF0FF46" author="Ameghino" box="[1301,1474,207,231]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino, 1897</bibRefCitation>
</taxonomicName>
from the Sarmiento Formation (Woodburne
<emphasis id="B964DA31FFEC0D15C20CFF398C0DFEA7" box="[1294,1343,238,262]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
2014a), and
<taxonomicName id="4C107DA0FFEC0D15C439FEDB8D47FE84" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[827,1141,269,293]" class="Mammalia" genus="Pseudadiantus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEC0D15C439FEDB8AFAFE84" box="[827,968,269,293]" italics="true" pageId="10" pageNumber="11">Pseudadiantus</emphasis>
<bibRefCitation id="EF817BD2FFEC0D15C4C9FEDB8D47FE84" author="Ameghino" box="[971,1141,269,293]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
</taxonomicName>
. The last was later excluded from the family and formally synonymized with the notopithecine interatheriid
<taxonomicName id="4C107DA0FFEC0D15C4BCFE9A8D08FEC5" authorityName="Ameghino" authorityYear="1901" box="[958,1082,332,356]" class="Mammalia" family="Interatheriidae" genus="Antepithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEC0D15C4BCFE9A8D08FEC5" box="[958,1082,332,356]" italics="true" pageId="10" pageNumber="11">Antepithecus</emphasis>
</taxonomicName>
by Simpson (1967).
</paragraph>
<paragraph id="8BAF0623FFEC0D15C457FEBD8D30FBB4" blockId="10.[823,1476,144,1735]" pageId="10" pageNumber="11">
<bibRefCitation id="EF817BD2FFEC0D15C457FEBD8ADDFE22" author="Bordas" box="[853,1007,363,387]" firstAuthor="Bordas" pageId="10" pageNumber="11" pagination="413 - 33" refId="ref40435" refString="Bordas AF. Diagnosis sobre algunos mamiferos de las capas con Colpodon, del valle del rio Chubut (Republica Argentina). Physis 1939; 14: 413 - 33." type="journal article" year="1939">Bordas (1939)</bibRefCitation>
added
<taxonomicName id="4C107DA0FFEC0D15C33CFEBD8C6CFE22" authority="Bordas, 1936" authorityName="Bordas" authorityYear="1936" box="[1086,1374,363,387]" class="Mammalia" family="Adianthidae" genus="Proheptaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEC0D15C33CFEBD8DFEFE22" box="[1086,1228,363,387]" italics="true" pageId="10" pageNumber="11">Proheptaconus</emphasis>
<bibRefCitation id="EF817BD2FFEC0D15C3D7FEBD8C6CFE22" author="Bordas" box="[1237,1374,363,387]" firstAuthor="Bordas" pageId="10" pageNumber="11" pagination="110 - 2" refId="ref40403" refString="Bordas AF. Un nuevo mamifero del Colpodon de Gaiman (Proheptaconus trelewense, gen. et sp. nov.). Physis 1936; 12: 110 - 2." type="journal article" year="1936">Bordas, 1936</bibRefCitation>
</taxonomicName>
from the Colhue-Huapi Member of the Sarmiento Formation (Soria 1981) to what he considered the subfamily
<taxonomicName id="4C107DA0FFEC0D15C3F9FE7C8C48FE63" box="[1275,1402,426,450]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="subFamily" subFamily="Adianthinae">Adianthinae</taxonomicName>
, a subfamily of
<taxonomicName id="4C107DA0FFEC0D15C4A0FE1F8D66FE40" box="[930,1108,457,481]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName>
, which was followed by other authors (Patterson 1940, Simpson and Minoprio 1949). However, Simpson
<emphasis id="B964DA31FFEC0D15C49CFDDF8AE1FD81" box="[926,979,520,544]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
(1962) later restored the family
<taxonomicName id="4C107DA0FFEC0D15C242FDDE8C8DFD81" box="[1344,1471,520,544]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
, but now including a new taxon,
<taxonomicName id="4C107DA0FFEC0D15C398FDF18AEAFDFE" authority="Simpson &amp; Minoprio, 1949" authorityName="Simpson &amp; Minoprio" authorityYear="1949" class="Mammalia" family="Macraucheniidae" genus="Adiantoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="leali">
<emphasis id="B964DA31FFEC0D15C398FDF18C72FD9E" box="[1178,1344,551,575]" italics="true" pageId="10" pageNumber="11">Adiantoides leali</emphasis>
Simpson &amp; Minoprio, 1949
</taxonomicName>
from the Divisadero Largo Formation,
<collectingCountry id="F30746B3FFEC0D15C25FFD918CF0FDFE" box="[1373,1474,583,607]" name="Argentina" pageId="10" pageNumber="11">Argentina</collectingCountry>
(age uncertain;
<bibRefCitation id="EF817BD2FFEC0D15C4DDFDB08D9CFDDF" author="Cerdeno" box="[991,1198,614,638]" etAl="et al." firstAuthor="Cerdeno" pageId="10" pageNumber="11" pagination="574 - 7" refId="ref40752" refString="Cerdeno E, Lopez GM, Reguero MA. Biostratigraphic considerations of the Divisaderan faunal assemblage. Journal of Vertebrate Paleontology 2008; 28: 574 - 7. https: // doi. org / 10.1671 / 0272 - 4634 (2008) 28 (574: bcotdf) 2.0. co; 2" type="journal article" year="2008">
Cerdeño
<emphasis id="B964DA31FFEC0D15C343FDB18D5EFDDF" box="[1089,1132,614,638]" italics="true" pageId="10" pageNumber="11">et al</emphasis>
. 2008
</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFEC0D15C3BEFDB08C0BFDDF" author="Lopez" box="[1212,1337,614,638]" firstAuthor="Lopez" pageId="10" pageNumber="11" pagination="410 - 20" refId="ref44025" refString="Lopez GM. Dividaderan: Land Mammal Age or local fauna? In: Madden RH, Carlini AA, Vucetich MG, Kay RF (eds), The paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge: Cambridge University Press, 2010, 410 - 20." type="book chapter" year="2010">López 2010</bibRefCitation>
, Woodburne
<emphasis id="B964DA31FFEC0D15C43BFD508A51FD3C" box="[825,867,645,669]" italics="true" pageId="10" pageNumber="11">et al</emphasis>
. 2014a).
<bibRefCitation id="EF817BD2FFEC0D15C4C7FD538DEEFD3C" author="Bond and Vucetich" box="[965,1244,645,669]" firstAuthor="Bond" pageId="10" pageNumber="11" pagination="107 - 17" refId="ref40299" refString="Bond M, Vucetich MG. Indalecia grandensis gen. et sp. nov. del Eoceno temprano del noroeste argentino, tipo de una nueva subfamilia de los Adianthidae. Revista de la Asociacion Geologica Argentina 1983; 38: 107 - 17." type="journal article" year="1983">Bond and Vucetich (1983)</bibRefCitation>
added to
<taxonomicName id="4C107DA0FFEC0D15C247FD538CF0FD3C" box="[1349,1474,645,669]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
<taxonomicName id="4C107DA0FFEC0D15C439FD738DC5FD1C" authority="Bond &amp; Vucetich, 1983" authorityName="Bond &amp; Vucetich" authorityYear="1983" box="[827,1271,677,701]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="grandensis">
<emphasis id="B964DA31FFEC0D15C439FD738D32FD1C" box="[827,1024,677,701]" italics="true" pageId="10" pageNumber="11">Indalecia grandensis</emphasis>
<bibRefCitation id="EF817BD2FFEC0D15C306FD738DC5FD1C" author="Bond &amp; Vucetich" box="[1028,1271,677,701]" firstAuthor="Bond" pageId="10" pageNumber="11" pagination="107 - 17" refId="ref40299" refString="Bond M, Vucetich MG. Indalecia grandensis gen. et sp. nov. del Eoceno temprano del noroeste argentino, tipo de una nueva subfamilia de los Adianthidae. Revista de la Asociacion Geologica Argentina 1983; 38: 107 - 17." type="journal article" year="1983">Bond &amp; Vucetich, 1983</bibRefCitation>
</taxonomicName>
from the Lumbrera Formation,
<collectingCountry id="F30746B3FFEC0D15C4BBFD128D2CFD7D" box="[953,1054,708,732]" name="Argentina" pageId="10" pageNumber="11">Argentina</collectingCountry>
(Early Eocene;
<bibRefCitation id="EF817BD2FFEC0D15C3D0FD128C83FD7D" author="Fernicola" box="[1234,1457,708,732]" etAl="et al." firstAuthor="Fernicola" pageId="10" pageNumber="11" pagination="621 - 33" refId="ref41787" refString="Fernicola JC, Zimicz AN, Chornogubsky L et al. The Early Eocene Climatic Optimum at the lower section of the Lumbrera Formation (Ypresian, Salta Province, northwestern Argentina): origin and early diversification of the Cingulata. Journal of Mammalian Evolution 2021; 28: 621 - 33. https: // doi. org / 10.1007 / s 10914 - 021 - 09545 - w" type="journal article" year="2021">
Fernicola
<emphasis id="B964DA31FFEC0D15C23CFD138C41FD7D" box="[1342,1395,708,732]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
2021
</bibRefCitation>
), based on unique shared anatomical features with
<taxonomicName id="4C107DA0FFEC0D15C24CFD358A53FCBA" authorityName="Simpson &amp; Minoprio" authorityYear="1949" class="Mammalia" family="Macraucheniidae" genus="Adiantoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="leali">
<emphasis id="B964DA31FFEC0D15C24CFD358A53FCBA" italics="true" pageId="10" pageNumber="11">Adiantoides leali</emphasis>
</taxonomicName>
, such as the absence of a postprotocrista connecting the protocone with the metaconule and a strong parastyle in M1– M2. These features are not present in other adianthids, so they grouped them in the new subfamily
<taxonomicName id="4C107DA0FFEC0D15C3B6FCB78C00FCD8" box="[1204,1330,865,889]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="subFamily" subFamily="Indaleciinae">Indaleciinae</taxonomicName>
, keeping both species tentatively within the family
<taxonomicName id="4C107DA0FFEC0D15C3CAFC568C75FC39" box="[1224,1351,896,920]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
, which was later followed with doubts by
<bibRefCitation id="EF817BD2FFEC0D15C376FC498C45FC16" author="Cifelli and Soria" box="[1140,1399,927,951]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria (1983a)</bibRefCitation>
. In the same work,
<bibRefCitation id="EF817BD2FFEC0D15C4B2FC698D9FFC76" author="Cifelli and Soria" box="[944,1197,959,983]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="1 - 25" refId="ref41068" refString="Cifelli RL, Soria MF. Systematics of the Adianthidae (Litopterna, Mammalia). American Museum Novitates 1983 a; 2771: 1 - 25." type="journal article" year="1983" yearSuffix="a">Cifelli and Soria (1983a)</bibRefCitation>
added the genus
<taxonomicName id="4C107DA0FFEC0D15C258FC698D3EFC57" authority="Cifelli &amp; Soria, 1983" authorityName="Cifelli &amp; Soria" authorityYear="1983" class="Mammalia" family="Adianthidae" genus="Thadanius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEC0D15C258FC698CF3FC76" box="[1370,1473,959,983]" italics="true" pageId="10" pageNumber="11">Thadanius</emphasis>
Cifelli &amp; Soria, 1983
</taxonomicName>
from La Salla-Luribay Basin,
<collectingCountry id="F30746B3FFEC0D15C23BFC088CB3FC57" box="[1337,1409,990,1014]" name="Bolivia" pageId="10" pageNumber="11">Bolivia</collectingCountry>
to the family
<taxonomicName id="4C107DA0FFEC0D15C47DFC2B8ACCFBB4" box="[895,1022,1021,1045]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
.
</paragraph>
<paragraph id="8BAF0623FFEC0D13C457FBCB887DFF66" blockId="10.[823,1476,144,1735]" lastBlockId="12.[128,778,144,481]" lastPageId="12" lastPageNumber="13" pageId="10" pageNumber="11">
Cifelli (1983a) revisited the members of the family, and included
<taxonomicName id="4C107DA0FFEC0D15C48EFBEB8DF0FBF5" authority="Ameghino 1904 b" authorityName="Ameghino" authorityYear="1904" box="[908,1218,1084,1108]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEC0D15C48EFBEB8AC9FBF5" box="[908,1019,1085,1108]" italics="true" pageId="10" pageNumber="11">Proectocion</emphasis>
<bibRefCitation id="EF817BD2FFEC0D15C30AFBEA8DF0FBF5" author="Ameghino" box="[1032,1218,1084,1108]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="1 - 541" refId="ref39426" refString="Ameghino F. Recherches de morphologic phylogenetique sur les molaires superieures des ongules. Anales del Museo Nacional de Buenos Aires 1904 b; 3: 1 - 541." type="journal article" year="1904" yearSuffix="b">Ameghino 1904b</bibRefCitation>
</taxonomicName>
[previously considered a didolodontid by Simpson (1948)] as the earliest member of
<taxonomicName id="4C107DA0FFEC0D15C43BFBAD8A84FB32" box="[825,950,1147,1171]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
based on dental similarities. In addition, he proposed a new superfamily called Macrauchenioidea, to group adianthids and macraucheniids based on dental similarities, such as an absent paraconid and the presence of a paralophid in the lower molars (
<tableCitation id="C6923398FFEC0D15C31AFB2E8D57FAB1" box="[1048,1125,1272,1296]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="10" pageNumber="11" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table 1</tableCitation>
). Soria (1984a), based on dental similarities with ‘amilnedwarsids’, elevated the rank of the subfamily
<taxonomicName id="4C107DA0FFEC0D15C481FAE18ACDFAEE" box="[899,1023,1335,1359]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="subFamily" subFamily="Indaleciinae">Indaleciinae</taxonomicName>
to the family
<taxonomicName id="4C107DA0FFEC0D15C399FAE18C2BFAEE" baseAuthorityName="Bond &amp; Vucetich" baseAuthorityYear="1983" box="[1179,1305,1335,1359]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Indaleciidae</taxonomicName>
, and later Soria (1989b) classified them as a family of the order
<taxonomicName id="4C107DA0FFEC0D15C248FA808CF0FACC" authorityName="Soria" authorityYear="1989" box="[1354,1474,1366,1389]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="order">Notopterna</taxonomicName>
(see sections of
<taxonomicName id="4C107DA0FFEC0D15C4E3FAA38D95FA2C" box="[993,1191,1397,1421]" class="Mammalia" family="Amilnedwardsiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Amilnedwardsiidae</taxonomicName>
and the order
<taxonomicName id="4C107DA0FFEC0D15C246FAA08C8CFA2C" authorityName="Soria" authorityYear="1989" box="[1348,1470,1398,1421]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="order">Notopterna</taxonomicName>
, and the family
<taxonomicName id="4C107DA0FFEC0D15C4E2FA438D50FA0C" baseAuthorityName="Bond &amp; Vucetich" baseAuthorityYear="1983" box="[992,1122,1429,1453]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Indaleciidae</taxonomicName>
).
<taxonomicName id="4C107DA0FFEC0D15C374FA438DC1FA0C" box="[1142,1267,1429,1453]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
has also been considered by some authors as part of the suborder
<taxonomicName id="4C107DA0FFEC0D15C22CFA628CF0FA6D" box="[1326,1474,1460,1484]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="subOrder" subOrder="Lopholipterna">Lopholipterna</taxonomicName>
(Cifelli 1983a, Soria 2001), which includes the families
<taxonomicName id="4C107DA0FFEC0D15C43BFA258ADBF9AA" box="[825,1001,1523,1547]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName>
and
<taxonomicName id="4C107DA0FFEC0D15C31DFA258DF5F9AA" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1055,1223,1523,1547]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
(
<tableCitation id="C6923398FFEC0D15C3D8FA258C1AF9AA" box="[1242,1320,1523,1547]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="10" pageNumber="11" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table 1</tableCitation>
), a hypothesis that later found phylogenetic support (
<bibRefCitation id="EF817BD2FFEC0D15C3C4F9C48C0DF98B" author="Cifelli" box="[1222,1343,1554,1578]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation>
;
<figureCitation id="132B1AA6FFEC0D15C24EF9C48CA4F98B" box="[1356,1430,1554,1578]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="10" pageNumber="11">Fig. 1C</figureCitation>
). In the same study, Macrauchenioidea also found support, but only when indaleciids are excluded (
<bibRefCitation id="EF817BD2FFEC0D15C38AF9878C31F9C8" author="Cifelli" box="[1160,1283,1617,1641]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation>
). Strikingly, there has been not a single study examining the phylogenetic relationships within
<taxonomicName id="4C107DA0FFEC0D15C4BCF9598D09F906" box="[958,1083,1679,1703]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
using modern phylogenetic methods, and most recent phylogenetic studies have included adianthids as outgroups for phylogenies of
<taxonomicName id="4C107DA0FFEA0D13C6C3FF468B43FF09" box="[449,625,144,168]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName>
(e.g.,
<bibRefCitation id="EF817BD2FFEA0D13C5A8FF4689D5FF66" author="Forasiepi" etAl="et al." firstAuthor="Forasiepi" pageId="12" pageNumber="13" pagination="1 - 76" refId="ref41925" refString="Forasiepi AM, MacPhee RDE, Hernandez del Pino S et al. Exceptional skull of Huayqueriana (Mammalia, Litopterna, Macraucheniidae) from the Late Miocene of Argentina: anatomy, systematics, and paleobiological implications. Bulletin of the American Museum of Natural History 2016; 404: 1 - 76." type="journal article" year="2016">
Forasiepi
<emphasis id="B964DA31FFEA0D13C783FF66899DFF66" box="[129,175,175,199]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
2016
</bibRefCitation>
;
<tableCitation id="C6923398FFEA0D13C7F6FF79880DFF66" box="[244,319,175,199]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" 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="12" pageNumber="13" tableUuid="DF6F56ABFFE20D1BC76CFE6E8C76FDA9">Table 2</tableCitation>
).
</paragraph>
<caption id="DF6F56ABFFED0D13C773F93C8A36F81A" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12" startId="11.[113,178,1770,1794]" targetBox="[194,1378,147,1740]" targetPageId="11" targetType="figure">
<paragraph id="8BAF0623FFED0D13C773F93C8A36F81A" blockId="11.[113,1453,1770,1963]" lastBlockId="12.[129,1473,1759,1979]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
<emphasis id="B964DA31FFED0D14C773F93C89FBF8A2" bold="true" box="[113,201,1770,1795]" pageId="11" pageNumber="12">Figure 4.</emphasis>
Right upper molars of
<taxonomicName id="4C107DA0FFED0D14C6A3F93D8B34F8A2" box="[417,518,1771,1795]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="order">Litopterna</taxonomicName>
in occlusal view. A, M3 of
<taxonomicName id="4C107DA0FFED0D14C402F93D8A83F8A2" authorityName="Püschel &amp; Shelley &amp; Williamson &amp; Perini &amp; Wible &amp; Brusatte" authorityYear="2024" baseAuthorityName="Roth" baseAuthorityYear="1899" box="[768,945,1771,1795]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="ligatus">
<emphasis id="B964DA31FFED0D14C402F93D8A83F8A2" box="[768,945,1771,1795]" italics="true" pageId="11" pageNumber="12">Polymorphis ligatus</emphasis>
</taxonomicName>
(holotype, MLP 12-2169). B, M3 of
<taxonomicName id="4C107DA0FFED0D14C20DF93D8CB2F8A2" authorityName="Roth" authorityYear="1899" box="[1295,1408,1771,1795]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFED0D14C20DF93D8CB2F8A2" box="[1295,1408,1771,1795]" italics="true" pageId="11" pageNumber="12">Polymorphis</emphasis>
</taxonomicName>
? (AMNH-VP 29481). C, M2? of
<taxonomicName id="4C107DA0FFED0D14C6A3F8D18BD5F8BE" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" box="[417,743,1799,1823]" class="Mammalia" family="Proterotheriidae" genus="Polyacrodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="lanciformis">
<emphasis id="B964DA31FFED0D14C6A3F8D18B4AF8BE" box="[417,632,1799,1823]" italics="true" pageId="11" pageNumber="12">Polyacrodon lanciformis</emphasis>
Roth, 1899
</taxonomicName>
[MLP 12-2170 (mirrored)]. D, P1–M3 of
<taxonomicName id="4C107DA0FFED0D14C379F8D18C66F8BE" authorityName="Ameghino" authorityYear="1902" box="[1147,1364,1799,1823]" class="Mammalia" family="Macraucheniidae" genus="Cramauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="normalis">
<emphasis id="B964DA31FFED0D14C379F8D18C66F8BE" box="[1147,1364,1799,1823]" italics="true" pageId="11" pageNumber="12">Cramauchenia normalis</emphasis>
</taxonomicName>
(MLP 85-VII-3-38a). E, P1–M3 of
<taxonomicName id="4C107DA0FFED0D14C67FF8F58B74F89B" authorityName="Scott" authorityYear="1910" box="[381,582,1827,1851]" class="Mammalia" family="Macraucheniidae" genus="Theosodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="garretorum">
<emphasis id="B964DA31FFED0D14C67FF8F58B74F89B" box="[381,582,1827,1851]" italics="true" pageId="11" pageNumber="12">Theosodon garretorum</emphasis>
</taxonomicName>
(FMNH P 13175). F, P5–M3 of
<taxonomicName id="4C107DA0FFED0D14C483F8F58D01F89B" authorityName="Ameghino" authorityYear="1904" box="[897,1075,1827,1850]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="precisus">
<emphasis id="B964DA31FFED0D14C483F8F58D01F89B" box="[897,1075,1827,1850]" italics="true" pageId="11" pageNumber="12">Proectocion precisus</emphasis>
</taxonomicName>
(holotype; MACN A 10679). G, P2– M3 of
<taxonomicName id="4C107DA0FFED0D14C7B2F8E988BEF8F6" authorityName="Ameghino" authorityYear="1897" box="[176,396,1855,1879]" class="Mammalia" family="Adianthidae" genus="Tricoelodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="bicuspidatus">
<emphasis id="B964DA31FFED0D14C7B2F8E988BEF8F6" box="[176,396,1855,1879]" italics="true" pageId="11" pageNumber="12">Tricoelodus bicuspidatus</emphasis>
</taxonomicName>
[cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of
<taxonomicName id="4C107DA0FFED0D14C4A7F8E98D5CF8F7" authorityName="Ameghino" authorityYear="1904" box="[933,1134,1855,1878]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="argentinus">
<emphasis id="B964DA31FFED0D14C4A7F8E98D5CF8F7" box="[933,1134,1855,1878]" italics="true" pageId="11" pageNumber="12">Proectocion argentinus</emphasis>
</taxonomicName>
(holotype; MACN A 10673). I, M1 of
<taxonomicName id="4C107DA0FFED0D14C7B2F88D8859F8D3" authorityName="Ameghino" authorityYear="1897" box="[176,363,1883,1907]" class="Mammalia" family="Proterotheriidae" genus="Lambdaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="suinus">
<emphasis id="B964DA31FFED0D14C7B2F88D8859F8D3" box="[176,363,1883,1907]" italics="true" pageId="11" pageNumber="12">Lambdaconus suinus</emphasis>
</taxonomicName>
[MNHN.F.DES162 (mirrored)]. J, P1–M3 of
<taxonomicName id="4C107DA0FFED0D14C421F88D8D3AF8D2" baseAuthorityName="Ameghino" baseAuthorityYear="1887" box="[803,1032,1883,1907]" class="Mammalia" family="Proterotheriidae" genus="Diadiaphorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="majusculus">
<emphasis id="B964DA31FFED0D14C421F88D8D3AF8D2" box="[803,1032,1883,1907]" italics="true" pageId="11" pageNumber="12">Diadiaphorus majusculus</emphasis>
</taxonomicName>
(MLP 12-304). K, P4–M3 of
<taxonomicName id="4C107DA0FFED0D14C227F88D899BF82F" authorityName="Ameghino" authorityYear="1897" class="Mammalia" family="Proterotheriidae" genus="Lambdaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="suinus">
<emphasis id="B964DA31FFED0D14C227F88D899BF82F" italics="true" pageId="11" pageNumber="12">Lambdaconus suinus</emphasis>
</taxonomicName>
(MACN A 52-198; P4 was mirrored). L, M1–M3 of
<taxonomicName id="4C107DA0FFED0D14C59FF8A18AA2F82F" authorityName="Paula Couto" authorityYear="1952" box="[669,912,1911,1935]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="prototypica">
<emphasis id="B964DA31FFED0D14C59FF8A18AA2F82F" box="[669,912,1911,1935]" italics="true" pageId="11" pageNumber="12">Victorlemoinea prototypica</emphasis>
</taxonomicName>
(AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of
<taxonomicName id="4C107DA0FFED0D14C307F8458DCCF80A" authorityName="Cifelli" authorityYear="1983" box="[1029,1278,1939,1963]" class="Mammalia" family="Proterotheriidae" genus="Paranisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="prodromus">
<emphasis id="B964DA31FFED0D14C307F8458DCCF80A" box="[1029,1278,1939,1963]" italics="true" pageId="11" pageNumber="12">Paranisolambda prodromus</emphasis>
</taxonomicName>
(cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 of
<taxonomicName id="4C107DA0FFEA0D13C598F9098A03F956" box="[666,817,1759,1783]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B964DA31FFEA0D13C598F9098A20F956" box="[666,786,1759,1783]" italics="true" pageId="12" pageNumber="13">Anisolambda</emphasis>
sp.
</taxonomicName>
(MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of
<taxonomicName id="4C107DA0FFEA0D13C48CF92D8D13F8B2" authorityName="Simpson" authorityYear="1935" box="[910,1057,1787,1811]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="tshotshe">
<emphasis id="B964DA31FFEA0D13C48CF92D8D13F8B2" box="[910,1057,1787,1811]" italics="true" pageId="12" pageNumber="13">Wainka tshotshe</emphasis>
</taxonomicName>
(
<typeStatus id="54ABB881FFEA0D13C32FF92D8DBBF8B2" box="[1069,1161,1787,1811]" pageId="12" pageNumber="13" type="holotype">holotype</typeStatus>
; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal
<quantity id="4CE8ABC6FFEA0D13C5D2F8758A32F81A" box="[720,768,1955,1979]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="12" pageNumber="13" unit="cm" value="1.0">1 cm</quantity>
.
</paragraph>
</caption>
<paragraph id="8BAF0623FFEA0D13C79EFF1989D8FE40" blockId="12.[128,778,144,481]" pageId="12" pageNumber="13">
If we follow Cifelli (1983a) and consider
<taxonomicName id="4C107DA0FFEA0D13C539FF198B9EFF47" authorityName="Ameghino" authorityYear="1904" box="[571,684,207,230]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEA0D13C539FF198B9EFF47" box="[571,684,207,230]" italics="true" pageId="12" pageNumber="13">Proectocion</emphasis>
</taxonomicName>
from the Barranca south of Lago Colhue-Huapi of
<collectingCountry id="F30746B3FFEA0D13C540FF388B9BFEA7" box="[578,681,238,262]" name="Argentina" pageId="12" pageNumber="13">Argentina</collectingCountry>
(Middle Eocene;
<bibRefCitation id="EF817BD2FFEA0D13C7D9FED888EEFE84" author="Kramarz and Bond" box="[219,476,269,293]" firstAuthor="Kramarz" pageId="12" pageNumber="13" pagination="203 - 211" refId="ref43514" refString="Kramarz, A., Bond, M. On the status of Isolophodon Roth, 1903 (Mammalia, Astrapotheria) and other little-known Paleogene astrapotheres from central Patagonia. Geobios 2013; 46 (3), 203 - 211." type="journal article" year="2013">Kramarz and Bond 2013</bibRefCitation>
, Woodburne
<emphasis id="B964DA31FFEA0D13C56FFED88BA5FE84" box="[621,663,269,293]" italics="true" pageId="12" pageNumber="13">et al</emphasis>
. 2014a) as the earliest member of
<taxonomicName id="4C107DA0FFEA0D13C671FEFB88C0FEE4" box="[371,498,301,325]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
and considering
<taxonomicName id="4C107DA0FFEA0D13C5A6FEFB89F3FEC5" authorityName="Cifelli" authorityYear="1991" class="Mammalia" family="Adianthidae" genus="Adianthus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="godoyi">
<emphasis id="B964DA31FFEA0D13C5A6FEFB89F3FEC5" italics="true" pageId="12" pageNumber="13">Adianthus godoyi</emphasis>
</taxonomicName>
from the Río Frías Formation at Alto Río Cisnes,
<collectingCountry id="F30746B3FFEA0D13C5D2FE9A8A3BFEC5" box="[720,777,332,356]" name="Chile" pageId="12" pageNumber="13">Chile</collectingCountry>
as the last member of this family (
<bibRefCitation id="EF817BD2FFEA0D13C6E5FEBD8B50FE22" author="Cifelli" box="[487,610,363,387]" firstAuthor="Cifelli" pageId="12" pageNumber="13" pagination="119 - 25" refId="ref40949" refString="Cifelli RLA. New adianthid litoptern (Mammalia) from the Miocene of Chile. Revista Chilena de Historia Natural 1991; 64: 119 - 25." type="journal article" year="1991">Cifelli 1991</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFEA0D13C56DFEBD8830FE02" author="de la Cruz and Cortes" firstAuthor="de la Cruz" pageId="12" pageNumber="13" pagination="1 - 70" refId="ref41584" refString="de la Cruz R, Cortes J. Geologia del area oriental de la Hoja Puerto Cisnes, Region Aysen del Gral. Carlos Ibanez del Campo. Serie Geologia Basica 2011; 127: 1 - 70." type="journal article" year="2011">de la Cruz and Cortés 2011</bibRefCitation>
), the family
<taxonomicName id="4C107DA0FFEA0D13C68AFE5D8B35FE02" box="[392,519,395,419]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
encompasses a temporal interval of 42–14.8 Mya (
<figureCitation id="132B1AA6FFEA0D13C6A1FE7C88C0FE63" box="[419,498,426,450]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="12" pageNumber="13">Fig. 2B</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFEA0D13C783FE1F89EEFE40" box="[129,220,457,481]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="12" pageNumber="13" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table S1</tableCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

322
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(
<figureCitation id="132B1AA6FFEE0D17C379FDB68DEBFDD8" box="[1147,1241,607,633]" captionStart="Figure 4" captionStartId="11.[113,178,1770,1794]" captionTargetBox="[194,1378,147,1740]" captionTargetId="figure-7@11.[191,1381,144,1743]" captionTargetPageId="11" captionText="Figure 4. Right upper molars of Litopterna in occlusal view.A, M3 of Polymorphis ligatus (holotype, MLP 12-2169). B, M3 of Polymorphis? (AMNH-VP 29481). C, M2? of Polyacrodon lanciformis Roth, 1899 [MLP 12-2170 (mirrored)]. D, P1–M3 of Cramauchenia normalis (MLP 85-VII-3-38a). E, P1–M3 of Theosodon garretorum (FMNH P 13175). F, P5–M3 of Proectocion precisus (holotype; MACN A 10679). G, P2– M3 of Tricoelodus bicuspidatus [cast of MLP 61-IV-11-65 (P2–P4 mirrored)]. H, M3 of Proectocion argentinus (holotype; MACN A 10673). I, M1 of Lambdaconus suinus [MNHN.F.DES162 (mirrored)]. J, P1–M3 of Diadiaphorus majusculus (MLP 12-304). K, P4–M3 of Lambdaconus suinus (MACN A 52-198; P4 was mirrored). L, M1–M3 of Victorlemoinea prototypica (AMNH-VP 111963, M1 (cast of MNRJ specimen); cast of MNRJ 1470V, M2 (holotype; mirrored); cast of MNRJ 1472V, M3 (mirrored). M, M1–M3 of Paranisolambda prodromus (cast of DGM 304M, M1; cast of DGM 310M, M2–M3). N, P4–M3 ofAnisolambda sp. (MLP 59-II-28-68, P4-P5 (mirrored); MLP 59-II-24-453, M1; MNHN.F.CAS 486, M2 (mirrored); MNHN.F.CAS488, M3 (mirrored). O, P5? of Wainka tshotshe (holotype; AMNH VP-28505 (mirrored). Relevant anatomical features of the dentition are labelled. Taxa with two prehypocristae are numbered from buccal (phyc1) to lingual (phyc2). When there are two cristae originating from the same cusp, they are numbered. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; psmlc, postmetaconular crista; psplc, postparaconular crista; psprc, postprotocrista. Scale bars equal 1 cm." pageId="8" pageNumber="9">Figs 4J–K</figureCitation>
,
<figureCitation id="132B1AA6FFEE0D17C3E6FD898C2EFDD8" box="[1252,1308,607,633]" captionStart="Figure 5" captionStartId="13.[113,178,1779,1803]" captionTargetBox="[190,1383,148,1748]" captionTargetId="figure-6@13.[187,1386,145,1751]" captionTargetPageId="13" captionText="Figure 5. Right lower molars of Litopterna in occlusal view.A, p1–p4 of Polymorphis lechei [holotype; MLP 12-2168 (mirrored)]. B, p1–m3 of Cramauchenia normalis [MNHN.F.COL181 (mirrored)]. C, m3 of Cramauchenia normalis (AMNH-VP 29753). D, p1–m3 of Theosodon lydekkeri [MACN A 24-90 (mirrored)]. E, m2–m3 of Proectocion sp. (MLP 59-II-28-107). F, p5–m3 of Tricoelodus bicuspidatus (MACN A 52-203, p5–m2 (holotype); cast of FMNH P14696, m3). G, p2–m3 of Proadiantus excavatus (MACN A 52-214). H, p1, dp2–dp5 and m1–m3 of Lambdaconus suinus (MACN A-52-199, p1, dp2–dp5 and m1–m2; MNHN.F.DES159, m3). I, p1–m3 of Diadiaphorus majusculus (MACN A 9180-82, p2–m3; YPM PU 15799, p1). J, m1–m3 of Anisolambda fissidens (MACN A 10668; holotype).K, m2–m3 of Paranisolambda prodromus (cast of MNRJ 1496V, m2 (mirrored); cast of MNRJ 1859V, m3 (mirrored). L, m2 of Victorlemoinea prototypica (MNRJ 1482V). M, p1–m3 of Sparnotheriodon epsilonoides (MACN 18225; holotype). N, O, m3 of Wainka tshotshe? [AMNH VP-29101 (mirrored)]. Relevant anatomical features of the dentition are labelled.More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods. Abbreviations:end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mlph, mesolophid; pad, paraconid; prd, protoconid; prgd, precingulid; psmcd, postmetacristid.Scale bars equal 1 cm." pageId="8" pageNumber="9">5H–I</figureCitation>
)
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C30A55A8FFEE0D15C43BFD5388D5FB50" lastPageId="10" lastPageNumber="11" pageId="8" pageNumber="9" type="multiple">
<paragraph id="8BAF0623FFEE0D17C43BFD538C0EFA0D" blockId="8.[822,1475,607,1641]" pageId="8" pageNumber="9">
With 28 currently accepted genera,
<taxonomicName id="4C107DA0FFEE0D17C3A6FD538C7EFD3C" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1188,1356,645,669]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
is the most diverse family of litopterns (
<figureCitation id="132B1AA6FFEE0D17C365FD738D81FD1C" box="[1127,1203,677,701]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="8" pageNumber="9">Fig. 2A</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFEE0D17C43BFD128AA6FD7D" box="[825,916,708,732]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="8" pageNumber="9" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table S1</tableCitation>
), being particularly diverse during the Neogene alongside macraucheniids (
<figureCitation id="132B1AA6FFEE0D17C327FD358DA6FD5A" box="[1061,1172,739,763]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="8" pageNumber="9">Fig. 2B–C</figureCitation>
). Soria (2001) presented a major revision of this family, so to avoid unnecessary repetition, only the most important taxonomical aspects and the main advances since his revision are discussed here.
<taxonomicName id="4C107DA0FFEE0D17C3F2FC978CAAFCF8" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1264,1432,833,857]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
was named by
<bibRefCitation id="EF817BD2FFEE0D17C4A7FCB78D53FCD8" author="Ameghino" box="[933,1121,865,889]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="1 - 26" refId="ref39043" refString="Ameghino F. Enumeracion sistematica de las especies de mamiferos fosiles coleccionados por Carlos Ameghino en los terrenos Eocenos de la Patagonia Austral y depositados en el Museo de La Plata. Boletin del Museo de La Plata 1887; 1: 1 - 26." type="journal article" year="1887">Ameghino (1887)</bibRefCitation>
with the description of the genera
<taxonomicName id="4C107DA0FFEE0D17C438FC568D4DFC39" authority="Ameghino, 1883" authorityName="Ameghino" authorityYear="1883" box="[826,1151,896,920]" class="Mammalia" family="Proterotheriidae" genus="Proterotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEE0D17C438FC568AFCFC39" box="[826,974,896,920]" italics="true" pageId="8" pageNumber="9">Proterotherium</emphasis>
<bibRefCitation id="EF817BD2FFEE0D17C4D6FC568D4DFC39" author="Ameghino" box="[980,1151,896,920]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="257 - 306" refId="ref39005" refString="Ameghino F. Sobre una nueva coleccion de mamiferos fosiles recogidos por el profesor Scalabrini en las barrancas del Parana. Boletin de la Academia Nacional de Ciencias en Cordoba 1883; 5: 257 - 306." type="journal article" year="1883">Ameghino, 1883</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C107DA0FFEE0D17C38EFC568C8EFC39" authority="Ameghino, 1887" authorityName="Ameghino" authorityYear="1887" box="[1164,1468,896,920]" class="Mammalia" family="Proterotheriidae" genus="Thoatherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEE0D17C38EFC568C39FC39" box="[1164,1291,896,920]" italics="true" pageId="8" pageNumber="9">Thoatherium</emphasis>
<bibRefCitation id="EF817BD2FFEE0D17C213FC568C8EFC39" author="Ameghino" box="[1297,1468,896,920]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="1 - 26" refId="ref39043" refString="Ameghino F. Enumeracion sistematica de las especies de mamiferos fosiles coleccionados por Carlos Ameghino en los terrenos Eocenos de la Patagonia Austral y depositados en el Museo de La Plata. Boletin del Museo de La Plata 1887; 1: 1 - 26." type="journal article" year="1887">Ameghino, 1887</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C107DA0FFEE0D17C43BFC498D59FC16" authority="Ameghino, 1887" authorityName="Ameghino" authorityYear="1887" box="[825,1131,927,951]" class="Mammalia" family="Proterotheriidae" genus="Diadiaphorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEE0D17C43BFC498AF2FC16" box="[825,960,927,951]" italics="true" pageId="8" pageNumber="9">Diadiaphorus</emphasis>
<bibRefCitation id="EF817BD2FFEE0D17C4C1FC498D59FC16" author="Ameghino" box="[963,1131,927,951]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="1 - 26" refId="ref39043" refString="Ameghino F. Enumeracion sistematica de las especies de mamiferos fosiles coleccionados por Carlos Ameghino en los terrenos Eocenos de la Patagonia Austral y depositados en el Museo de La Plata. Boletin del Museo de La Plata 1887; 1: 1 - 26." type="journal article" year="1887">Ameghino, 1887</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C107DA0FFEE0D17C39FFC498C8EFC16" authority="Ameghino, 1887" authorityName="Ameghino" authorityYear="1887" box="[1181,1468,927,951]" class="Mammalia" family="Proterotheriidae" genus="Licaphrium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEE0D17C39FFC498C3DFC16" box="[1181,1295,927,951]" italics="true" pageId="8" pageNumber="9">Licaphrium</emphasis>
<bibRefCitation id="EF817BD2FFEE0D17C216FC498C8EFC16" author="Ameghino" box="[1300,1468,927,951]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="1 - 26" refId="ref39043" refString="Ameghino F. Enumeracion sistematica de las especies de mamiferos fosiles coleccionados por Carlos Ameghino en los terrenos Eocenos de la Patagonia Austral y depositados en el Museo de La Plata. Boletin del Museo de La Plata 1887; 1: 1 - 26." type="journal article" year="1887">Ameghino, 1887</bibRefCitation>
</taxonomicName>
. Simpson (1945) subdivided this family into two subfamilies,
<taxonomicName id="4C107DA0FFEE0D17C43BFC088AE5FC57" box="[825,983,990,1014]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Polymorphinae">Polymorphinae</taxonomicName>
and
<taxonomicName id="4C107DA0FFEE0D17C308FC088D86FC57" box="[1034,1204,990,1014]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Proterotheriinae">Proterotheriinae</taxonomicName>
, the former including taxa mostly from the Palaeocene and Eocene (Peligran to Mustersan SALMAs) and the later the taxa from the Late Oligocene onwards (Deseadean SALMA). Although this classification was followed by other authors (
<bibRefCitation id="EF817BD2FFEE0D17C358FB8A8DD5FBD2" author="Lavocat" box="[1114,1255,1115,1139]" firstAuthor="Lavocat" pageId="8" pageNumber="9" pagination="31 - 59" refId="ref43823" refString="Lavocat R. Litopterna. Traite de Paleontologie 1958; 6: 31 - 59." type="journal article" year="1958">Lavocat 1958</bibRefCitation>
, Paula Couto 1979), Simpson (1948) did not employ this subfamilial division and only used the family
<taxonomicName id="4C107DA0FFEE0D17C304FB4C8D82FB13" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1030,1200,1178,1202]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
, without giving any explanation for this omission. Later, Odreman-Rivas (1969) modified the subfamily
<taxonomicName id="4C107DA0FFEE0D17C4C9FB0F8D5BFB50" box="[971,1129,1241,1265]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Polymorphinae">Polymorphinae</taxonomicName>
from Simpson’s (1945) proposal, keeping
<taxonomicName id="4C107DA0FFEE0D17C493FB2E8DBAFAB1" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" box="[913,1160,1272,1296]" class="Mammalia" family="Proterotheriidae" genus="Polymorphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEE0D17C493FB2E8D39FAB1" box="[913,1035,1272,1296]" italics="true" pageId="8" pageNumber="9">Polymorphis</emphasis>
Roth, 1899
</taxonomicName>
(including
<taxonomicName id="4C107DA0FFEE0D17C201FB2E8A99FA8E" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" class="Mammalia" family="Proterotheriidae" genus="Polyacrodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="ligatus">
<emphasis id="B964DA31FFEE0D17C201FB2E8CF0FAB1" box="[1283,1474,1272,1296]" italics="true" pageId="8" pageNumber="9">Polyacrodon ligatus</emphasis>
Roth, 1899
</taxonomicName>
and
<taxonomicName id="4C107DA0FFEE0D17C4D5FAC18DE1FA8E" authority="Roth, 1899" authorityName="Roth" authorityYear="1899" box="[983,1235,1303,1327]" class="Mammalia" family="Proterotheriidae" genus="Heteroglyphis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEE0D17C4D5FAC18D6EFA8E" box="[983,1116,1303,1327]" italics="true" pageId="8" pageNumber="9">Heteroglyphis</emphasis>
Roth, 1899
</taxonomicName>
), and excluding the rest of the taxa such as the now considered anisolambdids
<taxonomicName id="4C107DA0FFEE0D17C274FAE18AE5FACF" authority="Simpson, 1935" authorityName="Simpson" authorityYear="1935" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEE0D17C274FAE18CF0FAEE" box="[1398,1474,1335,1359]" italics="true" pageId="8" pageNumber="9">Wainka</emphasis>
Simpson, 1935
</taxonomicName>
and
<taxonomicName id="4C107DA0FFEE0D17C310FA808C7FFACF" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[1042,1357,1366,1390]" class="Mammalia" family="Proterotheriidae" genus="Anisolambda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEE0D17C310FA808DA1FACF" box="[1042,1171,1366,1390]" italics="true" pageId="8" pageNumber="9">Anisolambda</emphasis>
<bibRefCitation id="EF817BD2FFEE0D17C39CFA808C7FFACF" author="Ameghino" box="[1182,1357,1366,1390]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
</taxonomicName>
(including
<taxonomicName id="4C107DA0FFEE0D17C43BFAA38DAAFA2C" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[825,1176,1397,1421]" class="Mammalia" family="Proterotheriidae" genus="Ricardolydekkeria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEE0D17C43BFAA38ADBFA2C" box="[825,1001,1397,1421]" italics="true" pageId="8" pageNumber="9">Ricardolydekkeria</emphasis>
<bibRefCitation id="EF817BD2FFEE0D17C4ECFAA38DAAFA2C" author="Ameghino" box="[1006,1176,1397,1421]" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C107DA0FFEE0D17C3C8FAA38A5FFA0C" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" class="Mammalia" family="Proterotheriidae" genus="Josepholeidya" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEE0D17C3C8FAA38C7EFA2C" box="[1226,1356,1397,1421]" italics="true" pageId="8" pageNumber="9">Josepholeidya</emphasis>
<bibRefCitation id="EF817BD2FFEE0D17C250FAA38A5FFA0C" author="Ameghino" firstAuthor="Ameghino" pageId="8" pageNumber="9" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
</taxonomicName>
) and the didolodontid
<taxonomicName id="4C107DA0FFEE0D17C35EFA438C04FA0C" authority="Berg, 1899" authorityName="Berg" authorityYear="1899" box="[1116,1334,1429,1453]" class="Mammalia" family="Proterotheriidae" genus="Xesmodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFEE0D17C35EFA438DF3FA0C" box="[1116,1217,1429,1453]" italics="true" pageId="8" pageNumber="9">Xesmodon</emphasis>
<bibRefCitation id="EF817BD2FFEE0D17C3C5FA438C04FA0C" author="Berg" box="[1223,1334,1429,1453]" firstAuthor="Berg" pageId="8" pageNumber="9" pagination="77 - 80" refId="ref39765" refString="Berg C. Substitucion de nombres genericos. III. Comunicaciones del Museo Nacional de Buenos Aires 1899; 1: 77 - 80." type="journal article" year="1899">Berg, 1899</bibRefCitation>
</taxonomicName>
.
</paragraph>
<paragraph id="8BAF0623FFEE0D15C457FA62886AFD3C" blockId="8.[822,1475,607,1641]" lastBlockId="10.[126,779,144,1265]" lastPageId="10" lastPageNumber="11" pageId="8" pageNumber="9">
Cifelli(1983a)subdividedthefamilyProterotheriidaeintotwo subfamilies,
<taxonomicName id="4C107DA0FFEE0D17C4BAFA058D6DFA4A" box="[952,1119,1491,1515]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Anisolambdinae">Anisolambdinae</taxonomicName>
and
<taxonomicName id="4C107DA0FFEE0D17C39AFA058C70FA4A" box="[1176,1346,1491,1515]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Proterotheriinae">Proterotheriinae</taxonomicName>
, the former subfamily including mostly anisolambdids (see
<taxonomicName id="4C107DA0FFEE0D17C219FA258CF0F9AA" authorityName=", Soria" authorityYear="2001" box="[1307,1474,1523,1547]" family="Anisolambdidae" pageId="8" pageNumber="9" rank="family">Anisolambdidae</taxonomicName>
section for more details), and the latter similar to the original definition of Simpson (1945). In addition, Cifelli (1983a) excluded the subfamily
<taxonomicName id="4C107DA0FFEE0D17C4CFF9878D59F9C8" box="[973,1131,1617,1641]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="subFamily" subFamily="Polymorphinae">Polymorphinae</taxonomicName>
from
<taxonomicName id="4C107DA0FFEE0D17C3AEF9878C64F9C8" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[1196,1366,1617,1641]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="8" pageNumber="9" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
, placing it insteadwithinthefamilyMacraucheniidae (seeMacraucheniidae section for more details). Soria (2001) later removed the subfamily
<taxonomicName id="4C107DA0FFEC0D15C7C4FF19885FFF46" box="[198,365,207,231]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="subFamily" subFamily="Anisolambdinae">Anisolambdinae</taxonomicName>
from
<taxonomicName id="4C107DA0FFEC0D15C6A8FF198B66FF46" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[426,596,207,231]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
, elevating its rank as a distinct family (see
<taxonomicName id="4C107DA0FFEC0D15C680FF388B1BFEA7" authorityName=", Soria" authorityYear="2001" box="[386,553,238,262]" family="Anisolambdidae" pageId="10" pageNumber="11" rank="family">Anisolambdidae</taxonomicName>
section for more details), which leaves the concept of
<taxonomicName id="4C107DA0FFEC0D15C6E0FEDB8BB8FE84" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[482,650,269,293]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
very similar to the subfamily
<taxonomicName id="4C107DA0FFEC0D15C63DFEFB88D5FEE4" box="[319,487,301,325]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="subFamily" subFamily="Proterotheriinae">Proterotheriinae</taxonomicName>
from Simpson (1945), although with numerous new additions to the family since then (e.g.,
<bibRefCitation id="EF817BD2FFEC0D15C7B4FEBD885CFE22" author="Corona" box="[182,366,363,387]" etAl="et al." firstAuthor="Corona" pageId="10" pageNumber="11" refId="ref41392" refString="Corona A, Badin AC, Perea D et al. A new genus and species and additional reports of the South American native ungulates Proterotheriidae (Mammalia, Litopterna) in the Late Miocene of Uruguay. Journal of South American Earth Sciences 2020; 102: 102646. https: // doi. org / 10.1016 / j. jsames. 2020.102646" type="journal volume" year="2020">
Corona
<emphasis id="B964DA31FFEC0D15C60BFEBA8805FE22" box="[265,311,363,387]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
2020
</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFEC0D15C67AFEBD8B7EFE22" author="McGrath" box="[376,588,363,387]" etAl="et al." firstAuthor="McGrath" pageId="10" pageNumber="11" pagination="159 - 88" refId="ref44350" refString="McGrath AJ, Anaya F, Croft DA. New proterotheriids from the Middle Miocene of Quebrada Honda, Bolivia, and body size and diversity trends in proterotheriid and macraucheniid litopterns (Mammalia). Ameghiniana 2020 a; 57: 159 - 88. https: // doi. org / 10.5710 / amgh. 03.03.2020.3268" type="journal article" year="2020" yearSuffix="a">
McGrath
<emphasis id="B964DA31FFEC0D15C6D9FEBA8B3BFE22" box="[475,521,363,387]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
2020a
</bibRefCitation>
; Supporting information,
<tableCitation id="C6923398FFEC0D15C7D6FE5D881FFE02" box="[212,301,395,419]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="10" pageNumber="11" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table S1</tableCitation>
). Important additions to
<taxonomicName id="4C107DA0FFEC0D15C52EFE5D8BE6FE02" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[556,724,395,419]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
were the bunodont
<taxonomicName id="4C107DA0FFEC0D15C61FFE7C8BEBFE63" authority="McKenna, 1956" authorityName="McKenna" authorityYear="1956" box="[285,729,426,450]" class="Mammalia" family="Didolodontidae" genus="Megadolodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="molariformis">
<emphasis id="B964DA31FFEC0D15C61FFE7C8B17FE63" box="[285,549,426,450]" italics="true" pageId="10" pageNumber="11">Megadolodus molariformis</emphasis>
<bibRefCitation id="EF817BD2FFEC0D15C52DFE7C8BEBFE63" author="McKenna" box="[559,729,426,450]" firstAuthor="McKenna" pageId="10" pageNumber="11" pagination="736 - 43" refId="ref44469" refString="McKenna MC. Survival of primitive notoungulates and condylarths into the Miocene of Colombia. American Journal of Science 1956; 254: 736 - 43. https: // doi. org / 10.2475 / ajs. 254.12.736" type="journal article" year="1956">McKenna, 1956</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C107DA0FFEC0D15C783FE1F8BBBFE40" authority="Hoffstetter &amp; Soria, 1986" authorityName="Hoffstetter &amp; Soria" authorityYear="1986" box="[129,649,457,481]" class="Mammalia" family="Didolodontidae" genus="Neodolodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="colombianus">
<emphasis id="B964DA31FFEC0D15C783FE1F8847FE40" box="[129,373,457,481]" italics="true" pageId="10" pageNumber="11">Neodolodus colombianus</emphasis>
<bibRefCitation id="EF817BD2FFEC0D15C67FFE1F8BBBFE40" author="Hoffstetter &amp; Soria" box="[381,649,457,481]" firstAuthor="Hoffstetter" pageId="10" pageNumber="11" pagination="1619 - 22" refId="ref43229" refString="Hoffstetter R, Soria MF. Neolodus colombianus gen. et sp. nov., un noveau Condylarthre (Mammalia) dans le Miocene de Colombie. Comptes Rendus de la Academie de Sciences, Paris 1986; 303: 1619 - 22." type="journal article" year="1986">Hoffstetter &amp; Soria, 1986</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="EF817BD2FFEC0D15C59EFE1F89DEFDA0" author="Cifelli and Diaz" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="222 - 31" refId="ref41021" refString="Cifelli RL, Diaz JG. New remains of Prothoatherium columbianus (Litopterna, Mammalia) from the Miocene of Colombia. Journal of Vertebrate Paleontology 1989; 9: 222 - 31. https: // doi. org / 10.1080 / 02 724634.1989. 10011756" type="journal article" year="1989">Cifelli and Diaz 1989</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFEC0D15C7F8FE3F8B39FDA0" author="Cifelli and Villarroel" box="[250,523,489,513]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="265 - 88" refId="ref41117" refString="Cifelli RL, Villarroel C. Paleobiology and affinities of Megadolodus. In: Kay RF, Madden RH, Cifelli RL, Flynn JJ (eds), Vertebrate Paleontology in the Neotropics: The Miocene Fauna of La Venta, Colombia. Washington and London: Smithsonian Institution Press, 1997, 265 - 88." type="book chapter" year="1997">Cifelli and Villarroel 1997</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFEC0D15C51BFE3F8BCAFDA0" author="McGrath" box="[537,760,489,513]" etAl="et al." firstAuthor="McGrath" pageId="10" pageNumber="11" pagination="717 - 38" refId="ref44408" refString="McGrath AJ, Flynn JJ, Wyss AR. Proterotheriids and macraucheniids (Litopterna: Mammalia) from the Pampa Castillo Fauna, Chile (early Miocene, Santacrucian SALMA) and a new phylogeny of Proterotheriidae. Journal of Systematic Palaeontology 2020 b; 18: 717 - 38. https: // doi. org / 10.1080 / 14772019.2019.1662500" type="journal article" year="2020" yearSuffix="b">
McGrath
<emphasis id="B964DA31FFEC0D15C582FE3F8B80FDA0" box="[640,690,489,513]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
2020b
</bibRefCitation>
), which were previously considered didolodontids (
<bibRefCitation id="EF817BD2FFEC0D15C5A6FDDE8987FD9E" author="McKenna" firstAuthor="McKenna" pageId="10" pageNumber="11" pagination="736 - 43" refId="ref44469" refString="McKenna MC. Survival of primitive notoungulates and condylarths into the Miocene of Colombia. American Journal of Science 1956; 254: 736 - 43. https: // doi. org / 10.2475 / ajs. 254.12.736" type="journal article" year="1956">McKenna 1956</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFEC0D15C7C0FDF188EBFD9E" author="Hoffstetter and Soria" box="[194,473,551,575]" firstAuthor="Hoffstetter" pageId="10" pageNumber="11" pagination="1619 - 22" refId="ref43229" refString="Hoffstetter R, Soria MF. Neolodus colombianus gen. et sp. nov., un noveau Condylarthre (Mammalia) dans le Miocene de Colombie. Comptes Rendus de la Academie de Sciences, Paris 1986; 303: 1619 - 22." type="journal article" year="1986">Hoffstetter and Soria 1986</bibRefCitation>
). Including both taxa within
<taxonomicName id="4C107DA0FFEC0D15C783FD9189DCFDFE" box="[129,238,583,607]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="order">Litopterna</taxonomicName>
has phylogenetic support, but their affinities within
<taxonomicName id="4C107DA0FFEC0D15C781FDB08819FDDF" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[131,299,614,638]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
and even other litoptern families are unclear (Carrillo
<emphasis id="B964DA31FFEC0D15C7E2FD50883DFD3C" box="[224,271,645,669]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
2023).
</paragraph>
<caption id="DF6F56ABFFEF0D15C773F93B8D1DF81B" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10" startId="9.[113,178,1773,1797]" targetBox="[220,1352,145,1745]" targetPageId="9" targetType="figure">
<paragraph id="8BAF0623FFEF0D15C773F93B8D1DF81B" blockId="9.[113,1431,1773,1966]" lastBlockId="10.[129,1430,1842,1978]" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
<emphasis id="B964DA31FFEF0D16C773F93B89FBF8A4" bold="true" box="[113,201,1773,1797]" pageId="9" pageNumber="10">Figure 3.</emphasis>
Dental nomenclature used in the text and the characters. A, right upper molar in occlusal view. B–D, right lower molars in occlusal view. The main cusps of the molars are highlighted in bold. The molars represent structures sometimes present in SANUs and with relevance in the character scoring of this study. The molars do not represent any particular SANU. Molars in A and B were partially based on drawings from
<bibRefCitation id="EF817BD2FFEF0D16C7A6F8978811F8F8" author="Gelfo" box="[164,291,1857,1881]" firstAuthor="Gelfo" pageId="9" pageNumber="10" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo (2006)</bibRefCitation>
. Abbreviations upper molar: ecg, buccal cingulum or ectocingulum; es, entostyle; hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mt, metastyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmc, premetacrista; pmlc, premetaconular crista; ppc, preparacrista (or paracrista); pplc, preparaconular crista; pprc, preprotocrista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; pshyc, posthypocrista; psmc, postmetacrista (or metacrista); psmlc, postmetaconular crista; pspc, postparacrista; psplc, postparaconular crista; psprc, postprotocrista. Abbreviations lower molar: co, cristid obliqua; encd, entocristid; end, entoconid; dbgd, distobuccal cingulid; dlgd, distolingual cingulid; hlph, hypolophid; hycd, hypocristid; hyd, hypoconid; hyld, hypoconulid; mbgd, mesiobuccal cingulid; med, metaconid; mlgd, mesiolingual cingulid; mlph, mesolophid; msd, mesoconid; pacd, paracristid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prcd, protocristid; prd, protoconid; prgd, precingulid; pscd, postcristid (or postentocristid); pseld, postentoconulid; psgd, postcingulid; psmcd, postmetacristid.
</paragraph>
</caption>
<paragraph id="8BAF0623FFEC0D15C79EFD7389ACFC57" blockId="10.[126,779,144,1265]" pageId="10" pageNumber="11">
<taxonomicName id="4C107DA0FFEC0D15C79EFD738876FD1C" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[156,324,677,701]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
has been included by some authors as part of suborder
<taxonomicName id="4C107DA0FFEC0D15C640FD1288E4FD7D" box="[322,470,708,732]" pageId="10" pageNumber="11" rank="subOrder" subOrder="Lopholipterna">Lopholipterna</taxonomicName>
(Cifelli 1983a, Soria 2001;
<tableCitation id="C6923398FFEC0D15C783FD3589E0FD5A" box="[129,210,739,763]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="10" pageNumber="11" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table 1</tableCitation>
), which also includes the families
<taxonomicName id="4C107DA0FFEC0D15C55BFD358BE4FD5A" box="[601,726,739,763]" class="Mammalia" family="Adianthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Adianthidae</taxonomicName>
and
<taxonomicName id="4C107DA0FFEC0D15C783FCD58801FCBA" box="[129,307,771,795]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName>
, a hypothesis that later found phylogenetic support (
<bibRefCitation id="EF817BD2FFEC0D15C7E1FCF4886FFC9B" author="Cifelli" box="[227,349,802,826]" firstAuthor="Cifelli" pageId="10" pageNumber="11" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation>
;
<figureCitation id="132B1AA6FFEC0D15C66EFCF4888AFC9B" box="[364,440,802,826]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="10" pageNumber="11">Fig. 1C</figureCitation>
). Since then, most phylogenetic studies that included proterotheriids have focused either on resolving the affinities within
<taxonomicName id="4C107DA0FFEC0D15C694FCB78B0CFCD8" baseAuthorityName="Odreman Rivas" baseAuthorityYear="1969" box="[406,574,865,889]" class="Mammalia" family="Proterotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Proterotheriidae</taxonomicName>
(e.g.,
<bibRefCitation id="EF817BD2FFEC0D15C57BFCB789F2FC39" author="McGrath" etAl="et al." firstAuthor="McGrath" pageId="10" pageNumber="11" pagination="717 - 38" refId="ref44408" refString="McGrath AJ, Flynn JJ, Wyss AR. Proterotheriids and macraucheniids (Litopterna: Mammalia) from the Pampa Castillo Fauna, Chile (early Miocene, Santacrucian SALMA) and a new phylogeny of Proterotheriidae. Journal of Systematic Palaeontology 2020 b; 18: 717 - 38. https: // doi. org / 10.1080 / 14772019.2019.1662500" type="journal article" year="2020" yearSuffix="b">
McGrath
<emphasis id="B964DA31FFEC0D15C5DEFCB78A38FCD8" box="[732,778,865,889]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
2020b
</bibRefCitation>
) or phylogenies studying interordinal relationships between SANUs (e.g.,
<bibRefCitation id="EF817BD2FFEC0D15C64EFC498B2AFC16" author="MacPhee" box="[332,536,927,951]" etAl="et al." firstAuthor="MacPhee" pageId="10" pageNumber="11" pagination="1 - 183" refId="ref44179" refString="MacPhee RDE, Hernandez del Pino S, Kramarz A et al. Cranial morphology and phylogenetic relationships of Trigonostylops wortmani, an Eocene South American Native Ungulate. Bulletin of the American Museum of Natural History 2021; 449: 1 - 183." type="journal article" year="2021">
MacPhee
<emphasis id="B964DA31FFEC0D15C6B0FC7688D2FC16" box="[434,480,927,951]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
2021
</bibRefCitation>
) instead of phylogenies examining the interfamilial affinities of proterotheriids (
<tableCitation id="C6923398FFEC0D15C5D3FC6989BCFC57" captionStart="Table 2" captionStartId="4.[110,165,440,465]" 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="10" pageNumber="11" tableUuid="DF6F56ABFFE20D1BC76CFE6E8C76FDA9">Table 2</tableCitation>
).
</paragraph>
<paragraph id="8BAF0623FFEC0D15C79EFC2B88D5FB50" blockId="10.[126,779,144,1265]" pageId="10" pageNumber="11">
Following Soria’s (2001) concept of the family, the earliest proterotheriid is
<taxonomicName id="4C107DA0FFEC0D15C639FBCB8BF9FB94" authority="Ameghino, 1897" authorityName="Ameghino" authorityYear="1897" box="[315,715,1053,1077]" class="Mammalia" family="Proterotheriidae" genus="Lambdaconus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="suinus">
<emphasis id="B964DA31FFEC0D15C639FBCB8B3FFB95" box="[315,525,1053,1077]" italics="true" pageId="10" pageNumber="11">Lambdaconus suinus</emphasis>
<bibRefCitation id="EF817BD2FFEC0D15C51BFBCB8BF9FB94" author="Ameghino" box="[537,715,1053,1077]" firstAuthor="Ameghino" pageId="10" pageNumber="11" pagination="406 - 521" refId="ref39261" refString="Ameghino F. Mammiferes cretaces de l'Argentine (Deuxieme contribution a la connaissance de la fauna mammalogique de couches a Pyrotherium). Boletin del Instituto Geografico Argentino 1897; 18: 406 - 521." type="journal article" year="1897">Ameghino, 1897</bibRefCitation>
</taxonomicName>
from different localities from the Sarmiento Formation,
<collectingRegion id="49D4C8C1FFEC0D15C5B6FBEA8A34FBF5" box="[692,774,1084,1108]" country="Argentina" name="Chubut" pageId="10" pageNumber="11">Chubut</collectingRegion>
,
<collectingCountry id="F30746B3FFEC0D15C783FB8D89DBFBD2" box="[129,233,1115,1139]" name="Argentina" pageId="10" pageNumber="11">Argentina</collectingCountry>
, and the last is
<taxonomicName id="4C107DA0FFEC0D15C687FB8D8A3BFBD2" authority="Frenguelli, 1921" authorityName="Frenguelli" authorityYear="1921" box="[389,777,1115,1139]" class="Mammalia" family="Proterotheriidae" genus="Neolicaphrium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="recents">
<emphasis id="B964DA31FFEC0D15C687FB8D8B6FFBD2" box="[389,605,1115,1139]" italics="true" pageId="10" pageNumber="11">Neolicaphrium recents</emphasis>
<bibRefCitation id="EF817BD2FFEC0D15C560FB8D8A3BFBD2" author="Frenguelli" box="[610,777,1115,1139]" firstAuthor="Frenguelli" pageId="10" pageNumber="11" pagination="7 - 23" refId="ref41980" refString="Frenguelli J. Sobre un proteroterido del Pampeano superior de Cordoba. Actas de la Academia Nacional de Ciencias de la Republica Argentina en Cordoba 1921; 7: 7 - 23." type="journal article" year="1921">Frenguelli, 1921</bibRefCitation>
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from the Late Pleistocene from different localities in
<collectingCountry id="F30746B3FFEC0D15C59CFBAD8A34FB32" box="[670,774,1147,1171]" name="Argentina" pageId="10" pageNumber="11">Argentina</collectingCountry>
,
<collectingCountry id="F30746B3FFEC0D15C783FB4C898CFB13" box="[129,190,1178,1202]" name="Brazil" pageId="10" pageNumber="11">Brazil</collectingCountry>
, and
<collectingCountry id="F30746B3FFEC0D15C7FAFB4C8863FB13" box="[248,337,1178,1202]" name="Uruguay" pageId="10" pageNumber="11">Uruguay</collectingCountry>
(
<bibRefCitation id="EF817BD2FFEC0D15C660FB4C8B08FB13" author="Gaudioso" box="[354,570,1178,1202]" etAl="et al." firstAuthor="Gaudioso" pageId="10" pageNumber="11" pagination="23 - 9" refId="ref42056" refString="Gaudioso PJ, Gasparini GM, Herbst R et al. First record of the Neolicaphrium recens Frenguelli, 1921 (Mammalia, Litopterna) in the Pleistocene of Santiago del Estero Province, Argentina. Papeis Avulsos de Zoologia 2017; 57: 23 - 9. https: // doi. org / 10.11606 / 0031 - 1049.2017. 57.03" type="journal article" year="2017">
Gaudioso
<emphasis id="B964DA31FFEC0D15C6CCFB4D88CDFB13" box="[462,511,1178,1202]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
2017
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), giving a temporal interval of around 29.3–~0.11 Mya to this family (
<figureCitation id="132B1AA6FFEC0D15C5B1FB6F8A33FB70" box="[691,769,1209,1233]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="10" pageNumber="11">Fig. 2B</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFEC0D15C67DFB0F88EBFB50" box="[383,473,1241,1265]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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="10" pageNumber="11" tableUuid="DF6F56ABFFE40D1DC76CFE228F67FDAC">Table S1</tableCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

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(
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,
<figureCitation id="132B1AA6FFF30D0AC519FD488B6AFD19" box="[539,600,670,696]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="21" pageNumber="22">9F–H</figureCitation>
)
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<taxonomicName id="4C107DA0FFF30D0AC773FD128829FD7D" box="[113,283,708,732]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Protolipternidae</taxonomicName>
currently includes three genera [or two if we excluded
<taxonomicName id="4C107DA0FFF30D0AC7D5FD3588A5FD5A" authorityName="Ameghino" authorityYear="1901" box="[215,407,739,763]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF30D0AC7D5FD3588A5FD5A" box="[215,407,739,763]" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia</emphasis>
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as
<bibRefCitation id="EF817BD2FFF30D0AC6BFFD358B79FD5A" author="Gelfo" box="[445,587,739,763]" firstAuthor="Gelfo" pageId="21" pageNumber="22" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo (2006)</bibRefCitation>
suggested] with an exclusively Palaeogene distribution (
<figureCitation id="132B1AA6FFF30D0AC506FCD58B09FCBA" box="[516,571,771,795]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="21" pageNumber="22">Fig. 2</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFF30D0AC7C4FCF48810FC9B" box="[198,290,802,826]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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>
). This family was proposed by Cifelli (1983a) to classify specimens from Itaboraí (Itaboraian SALMA) with bunodont didolodontid-like dentition and litoptern-like tarsals. The tarsals show specializations that among SANUs are only present in litopterns, which include a spool-shaped astragalar body and a posterior astragalocalcaneal facet that allows rapid flexion of the crurotarsal joint (Cifelli 1983a). The taxonomic changes generated from this familial proposal included the consideration of
<taxonomicName id="4C107DA0FFF30D0AC7CCFBCB88EEFB95" authorityName="Paula Couto" authorityYear="1952" box="[206,476,1053,1077]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="prototypica">
<emphasis id="B964DA31FFF30D0AC7CCFBCB88EEFB95" box="[206,476,1053,1077]" italics="true" pageId="21" pageNumber="22">Victorlemoinea prototypica</emphasis>
</taxonomicName>
[a former litoptern (Paula Couto 1952)] as a ‘condylarth’ closely related to didolodontids, and
<taxonomicName id="4C107DA0FFF30D0AC7A8FB8D8886FBD2" baseAuthorityName="Cifelli" baseAuthorityYear="1983" box="[170,436,1115,1139]" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="parayirunhor">
<emphasis id="B964DA31FFF30D0AC7A8FB8D8886FBD2" box="[170,436,1115,1139]" italics="true" pageId="21" pageNumber="22">Miguelsoria parayirunhor</emphasis>
</taxonomicName>
[considered a didolodontid
<taxonomicName id="4C107DA0FFF30D0AC773FBAD88ABFB32" baseAuthorityName="Paula Couto" baseAuthorityYear="1952" box="[113,409,1147,1171]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="parayirunhor">
<emphasis id="B964DA31FFF30D0AC773FBAD88ABFB32" box="[113,409,1147,1171]" italics="true" pageId="21" pageNumber="22">Ernestokokenia parayirunhor</emphasis>
</taxonomicName>
by Paula Couto (1952)] with
<taxonomicName id="4C107DA0FFF30D0AC773FB4C885EFB13" authorityName="Paula Couto" authorityYear="1952" box="[113,364,1178,1202]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="scotti">
<emphasis id="B964DA31FFF30D0AC773FB4C885EFB13" box="[113,364,1178,1202]" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia scotti</emphasis>
</taxonomicName>
[considered a didolodontid by Paula Couto (1978)] as litopterns. Cifelli (1983a) grouped the last two, plus a new species
<taxonomicName id="4C107DA0FFF30D0AC687FB0F8983FAB1" authority="Cifelli 1983 a" authorityName="Cifelli" authorityYear="1983" class="Mammalia" family="Protolipternidae" genus="Protolipterna" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="ellipsodontoides">
<emphasis id="B964DA31FFF30D0AC687FB0F8B9CFB50" box="[389,686,1241,1265]" italics="true" pageId="21" pageNumber="22">Protolipterna ellipsodontoides</emphasis>
Cifelli 1983a
</taxonomicName>
, within this family. However, the tarsals referred to
<taxonomicName id="4C107DA0FFF30D0AC773FAC188A6FA8E" authorityName="Cifelli" authorityYear="1983" box="[113,404,1303,1327]" class="Mammalia" family="Protolipternidae" genus="Protolipterna" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="ellipsodontoides">
<emphasis id="B964DA31FFF30D0AC773FAC188A6FA8E" box="[113,404,1303,1327]" italics="true" pageId="21" pageNumber="22">Protolipterna ellipsodontoides</emphasis>
</taxonomicName>
and
<taxonomicName id="4C107DA0FFF30D0AC6C5FAC18BF6FA8E" baseAuthorityName="Cifelli" baseAuthorityYear="1983" box="[455,708,1303,1327]" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="parayirunhor">
<emphasis id="B964DA31FFF30D0AC6C5FAC18BF6FA8E" box="[455,708,1303,1327]" italics="true" pageId="21" pageNumber="22">Miguelsoria parayirunhor</emphasis>
</taxonomicName>
were not found in association with their alleged dentition, being assigned indirectly by their relative size, abundance, and expected morphology (Cifelli 1983b). Some authors have accepted this interpretation (e.g.,
<bibRefCitation id="EF817BD2FFF30D0AC6D2FA438B62FA0D" author="Cifelli" box="[464,592,1428,1453]" firstAuthor="Cifelli" pageId="21" pageNumber="22" pagination="249 - 66" refId="ref40909" refString="Cifelli RL. South American ungulate evolution and extinction. In: Stehli FG, Webb SD (eds), The Great American Biotic Interchange. New York, NY: Springer, 1985, 249 - 66." type="book chapter" year="1985">Cifelli 1985</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFF30D0AC561FA438BA5FA0C" author="Cifelli" box="[611,663,1429,1453]" firstAuthor="Cifelli" pageId="21" pageNumber="22" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">1993</bibRefCitation>
, Muizon and Cifelli 2000,
<bibRefCitation id="EF817BD2FFF30D0AC62AFA6288F4FA6D" author="Bergqvist" box="[296,454,1460,1484]" firstAuthor="Bergqvist" pageId="21" pageNumber="22" pagination="107 - 33" refId="ref39850" refString="Bergqvist LP. Postcranial skeleton of the Upper Paleocene of Itaborai Basin, Brazil. In: Sargis EJ, Dagosto M, (eds), Mammalian Evolutionary Morphology: A Tribute to Frederik S. Szalay. New York, NY: Springer-Verlag, 2008, 107 - 33." type="book chapter" year="2008">Bergqvist 2008</bibRefCitation>
), although others have questioned it (
<bibRefCitation id="EF817BD2FFF30D0AC7DFFA0588C1FA4A" author="Hoffstetter and Soria" box="[221,499,1491,1515]" firstAuthor="Hoffstetter" pageId="21" pageNumber="22" pagination="1619 - 22" refId="ref43229" refString="Hoffstetter R, Soria MF. Neolodus colombianus gen. et sp. nov., un noveau Condylarthre (Mammalia) dans le Miocene de Colombie. Comptes Rendus de la Academie de Sciences, Paris 1986; 303: 1619 - 22." type="journal article" year="1986">Hoffstetter and Soria 1986</bibRefCitation>
, Soria 2001,
<bibRefCitation id="EF817BD2FFF30D0AC57CFA058BC6FA4A" author="Gelfo" box="[638,756,1491,1515]" firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="130 - 42" refId="ref42288" refString="Gelfo JN. The ' condylarth' Didolodontidae from Gran Barranca: history of the bunodont South American mammals until the Eocene- Oligocene transition. In: Madden RH, Carlini AA, Vucetich MG, Kay RF (eds.), The Paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge: Cambridge University Press, 2010, 130 - 42." type="book chapter" year="2010">Gelfo 2010</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFF30D0AC773FA258874F9AA" author="Gelfo and Sige" box="[113,326,1523,1547]" firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="665 - 78" refId="ref42458" refString="Gelfo JN, Sige B. A new didolodontid mammal from the Late Paleocene - earliest Eocene of Laguna Umayo, Peru. Acta Palaeontologica Polonica 2011; 56: 665 - 78. https: // doi. org / 10.4202 / app. 2010.0067" type="journal article" year="2011">Gelfo and Sigé 2011</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFF30D0AC651FA258B7DF9AA" author="Gelfo and Lorente" box="[339,591,1523,1547]" firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="249 - 59" refId="ref42404" refString="Gelfo JN, Lorente M. The alleged astragalar remains of Didolodus Ameghino, 1897 (Mammalia, Panameriungulata) and a critic of isolated bone association models. Bulletin of Geosciences 2012; 87: 249 - 59. https: // doi. org / 10.3140 / bull. geosci. 1301" type="journal article" year="2012">Gelfo and Lorente 2012</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFF30D0AC55EFA258BDBF9AB" author="Lorente" box="[604,745,1522,1546]" firstAuthor="Lorente" pageId="21" pageNumber="22" refId="ref44083" refString="Lorente M. Desarrollo de Modelos de Asociacion y Clasificaciones de Restos Postcraneanos Aislados de Ungulados Nativos del Paleoceno- Eoceno de America del Sur. D. Phil. Thesis, Universidad Nacional de la Plata, Argentina, 2015." type="book" year="2015">Lorente 2015</bibRefCitation>
). For example, Soria (2001) suggested that the astragali and calcanea attributed by Cifelli (1983b) to didolodontids were most likely notoungulate elements. Indeed, a study that indirectly assigned isolated petrosals to the protolipternid
<taxonomicName id="4C107DA0FFF30D0AC584F9A68BC8F929" authorityName="Cifelli" authorityYear="1983" box="[646,762,1648,1672]" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF30D0AC584F9A68BC8F929" box="[646,762,1648,1672]" italics="true" pageId="21" pageNumber="22">Miguelsoria</emphasis>
</taxonomicName>
from Itaboraí failed to recover a monophyletic
<taxonomicName id="4C107DA0FFF30D0AC205FF468C47FF09" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[1287,1397,144,168]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="order">Litopterna</taxonomicName>
when younger and better known litopterns (e.g.,
<taxonomicName id="4C107DA0FFF30D0AC3F5FF798CB1FF66" authorityName="Owen" authorityYear="1838" box="[1271,1411,175,199]" class="Mammalia" family="Macraucheniidae" genus="Macrauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF30D0AC3F5FF798CB1FF66" box="[1271,1411,175,199]" italics="true" pageId="21" pageNumber="22">Macrauchenia</emphasis>
</taxonomicName>
and
<taxonomicName id="4C107DA0FFF30D0AC428FF198A80FF46" authorityName="Ameghino" authorityYear="1887" box="[810,946,207,231]" class="Mammalia" family="Proterotheriidae" genus="Diadiaphorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF30D0AC428FF198A80FF46" box="[810,946,207,231]" italics="true" pageId="21" pageNumber="22">Diadiaphorus</emphasis>
</taxonomicName>
) were included (
<bibRefCitation id="EF817BD2FFF30D0AC36EFF198C2BFF47" author="Billet" box="[1132,1305,206,231]" etAl="et al." firstAuthor="Billet" pageId="21" pageNumber="22" pagination="956 - 87" refId="ref39984" refString="Billet G, de Muizon C, Schellhorn R et al. Petrosal and inner ear anatomy and allometry amongst specimens referred to Litopterna (Placentalia). Zoological Journal of the Linnean Society 2015; 173: 956 - 87. https: // doi. org / 10.1111 / zoj. 12219" type="journal article" year="2015">
Billet
<emphasis id="B964DA31FFF30D0AC3AEFF198DECFF46" box="[1196,1246,207,231]" italics="true" pageId="21" pageNumber="22">et al.</emphasis>
2015
</bibRefCitation>
). Considering that recent association models do not discard the association of litoptern-like tarsals to protolipternids (
<bibRefCitation id="EF817BD2FFF30D0AC3DCFED88C5BFE84" author="Lorente" box="[1246,1385,269,293]" firstAuthor="Lorente" pageId="21" pageNumber="22" refId="ref44083" refString="Lorente M. Desarrollo de Modelos de Asociacion y Clasificaciones de Restos Postcraneanos Aislados de Ungulados Nativos del Paleoceno- Eoceno de America del Sur. D. Phil. Thesis, Universidad Nacional de la Plata, Argentina, 2015." type="book" year="2015">Lorente 2015</bibRefCitation>
) made by Cifelli (1983b), some authors have started to consider the family
<taxonomicName id="4C107DA0FFF30D0AC472FE9A8D28FEC5" box="[880,1050,332,356]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Protolipternidae</taxonomicName>
as a junior synonym of
<taxonomicName id="4C107DA0FFF30D0AC20FFE9A8C80FEC5" box="[1293,1458,332,356]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Didolodontidae</taxonomicName>
(
<bibRefCitation id="EF817BD2FFF30D0AC437FEBD8AD2FE22" author="Gelfo" box="[821,992,363,387]" etAl="et al." firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="285 - 91" refId="ref42550" refString="Gelfo JN, Lopez GM, Lorente M. Los ungulados arcaicos de America del Sur: ' Condylarthra' y Litopterna. Contribuciones Cientificas Del Museo Argentino de Ciencias Naturales ' Bernardino Rivadavia' 2016; 6: 285 - 91." type="journal article" year="2016">
Gelfo
<emphasis id="B964DA31FFF30D0AC474FEBA8A94FE22" box="[886,934,363,387]" italics="true" pageId="21" pageNumber="22">et al.</emphasis>
2016
</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFF30D0AC4EFFEBD8D1FFE22" author="Gelfo" box="[1005,1069,363,387]" firstAuthor="Gelfo" pageId="21" pageNumber="22" pagination="132 - 45" refId="ref42677" refString="Gelfo JN, Alonso RN, Madden RH et al. An Eocene bunodont South American native ungulate (Didolodontidae) from the Lumbrera Formation, Salta Province, Argentina. Ameghiniana 2020 a; 57: 132 - 45. https: // doi. org / 10.5710 / amgh. 29.11.2019.3293" type="journal article" year="2020" yearSuffix="a">
2020
<bibRefCitation id="EF817BD2FFF30D0AC320FEBD8C13FE22" author="Corona A &amp; Badin AC &amp; Perea D" box="[1058,1313,363,387]" firstAuthor="a" pageId="21" pageNumber="22" refId="ref41392" refString="Corona A, Badin AC, Perea D et al. A new genus and species and additional reports of the South American native ungulates Proterotheriidae (Mammalia, Litopterna) in the Late Miocene of Uruguay. Journal of South American Earth Sciences 2020; 102: 102646. https: // doi. org / 10.1016 / j. jsames. 2020.102646" type="journal volume" year="2020">a, Croft and López 2020</bibRefCitation>
</bibRefCitation>
). In addition, based on anatomical observations and a phylogenetic analysis,
<bibRefCitation id="EF817BD2FFF30D0AC42EFE7C8A8BFE63" author="Gelfo" box="[812,953,426,450]" firstAuthor="Gelfo" pageId="21" pageNumber="22" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo (2006)</bibRefCitation>
considered that
<taxonomicName id="4C107DA0FFF30D0AC368FE7C8C56FE63" authorityName="Paula Couto" authorityYear="1952" box="[1130,1380,426,450]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="scotti">
<emphasis id="B964DA31FFF30D0AC368FE7C8C56FE63" box="[1130,1380,426,450]" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia scotti</emphasis>
</taxonomicName>
should be generically renamed as is it not recovered as a sister taxon of the Patagonian
<taxonomicName id="4C107DA0FFF30D0AC4CBFE3F8C94FDA0" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[969,1446,489,513]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="subtrigona">
<emphasis id="B964DA31FFF30D0AC4CBFE3F8DC7FDA0" box="[969,1269,489,513]" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia subtrigona</emphasis>
<bibRefCitation id="EF817BD2FFF30D0AC3F9FE3F8C94FDA0" author="Ameghino" box="[1275,1446,489,513]" firstAuthor="Ameghino" pageId="21" pageNumber="22" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
</taxonomicName>
.
</paragraph>
<paragraph id="8BAF0623FFF30D0AC447FDDE8C6FFB94" blockId="21.[808,1461,144,1327]" pageId="21" pageNumber="22">
Apartfromearlyphylogeneticstudiesthatfoundprotolipternids closely related to other litopterns based on Cifelli’s (1983b) tarsal associations (
<bibRefCitation id="EF817BD2FFF30D0AC4B0FD918D15FDFE" author="Cifelli" box="[946,1063,583,607]" firstAuthor="Cifelli" pageId="21" pageNumber="22" pagination="195 - 216" refId="ref40977" refString="Cifelli RL. The phylogeny of the native South American ungulates. In: Szalay FS, Novacek MJ, McKenna MC (eds), Mammal Phylogeny: Placentals. New York, NY: Springer-Verlag, 1993, 195 - 216." type="book chapter" year="1993">Cifelli 1993</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFF30D0AC331FD918DF8FDFE" author="Bergqvist" box="[1075,1226,583,607]" firstAuthor="Bergqvist" pageId="21" pageNumber="22" refId="ref39790" refString="Bergqvist LP. Reasociacao de pos-cranio as especies de ungulados da Bacia de S. J. Itaborai (Paleoceno), Estado do Rio de Janeiro, e Filogenia dos ' Condylarthra', e ungulados Sul-Americanos com base no poscranio. D. Phil. Thesis, Universidade Federal do Rio Grande do Sul, Brazil, 1996." type="book" year="1996">Bergqvist 1996</bibRefCitation>
;
<figureCitation id="132B1AA6FFF30D0AC3D5FD918C12FDFE" box="[1239,1312,583,607]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="21" pageNumber="22">Fig. 1C</figureCitation>
), other phylogenetic studies including protolipternids and didolodontids have usually failed to find them as distinctly separate groups [e.g., Muizon and Cifelli (2000); see also
<taxonomicName id="4C107DA0FFF30D0AC3B0FD738C60FD1C" box="[1202,1362,677,701]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Didolodontidae</taxonomicName>
section]. Other phylogenetic studies with a wide taxon sampling have found
<taxonomicName id="4C107DA0FFF30D0AC46DFD358D25FD5A" box="[879,1047,739,763]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Protolipternidae</taxonomicName>
: in a basal position in Laurasiatheria as a sister group of
<taxonomicName id="4C107DA0FFF30D0AC4D8FCD58C52FCBA" authority="(O'Leary et al. 2013)" baseAuthorityName="O'Leary" baseAuthorityYear="2013" box="[986,1376,771,795]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="family">
Didolodontidae (O’Leary
<emphasis id="B964DA31FFF30D0AC3EBFCD58C26FCBA" box="[1257,1300,771,795]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. 2013)
</taxonomicName>
; closely related to other more derived litopterns than to other SANUs (Muizon
<emphasis id="B964DA31FFF30D0AC48FFC948A8BFCF8" box="[909,953,833,857]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. 2015); among didolodontids at the stem of litopterns but not forming a monophyletic group (
<bibRefCitation id="EF817BD2FFF30D0AC221FCB78A6FFC39" author="Kramarz" etAl="et al." firstAuthor="Kramarz" pageId="21" pageNumber="22" refId="ref43620" refString="Kramarz AG, Bond M, MacPhee RDE. On the alleged perissodactyl affinities of the ' condylarth' Escribania chubutensis and other endemic South American ungulate-like placentals. Journal of Vertebrate Paleontology 2021; 41: e 1986716." type="journal volume" year="2021">
Kramarz
<emphasis id="B964DA31FFF30D0AC280FCB78C9CFCD8" box="[1410,1454,865,889]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. 2021
</bibRefCitation>
); and forming a monophyletic group as part of a polytomy that includes didolodontids, sparnotheriodontids, indaleciids, and North American phenacodontids (Zimicz
<emphasis id="B964DA31FFF30D0AC3D4FC698DCDFC76" box="[1238,1279,959,983]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. 2022). However, so far there has not been any phylogenetic analysis that includes protolipternids alongside representatives of all litoptern families, didolodontids, and the different SANU orders (
<tableCitation id="C6923398FFF30D0AC206FBCB8C7FFB94" box="[1284,1357,1053,1077]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" 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>
).
</paragraph>
<paragraph id="8BAF0623FFF30D0AC447FBEA8AA7FA8E" blockId="21.[808,1461,144,1327]" pageId="21" pageNumber="22">
If we follow Cifelli (1983a) and consider
<taxonomicName id="4C107DA0FFF30D0AC20BFBEA8C81FBF5" box="[1289,1459,1084,1108]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="family">Protolipternidae</taxonomicName>
as a valid family, its earliest members are
<taxonomicName id="4C107DA0FFF30D0AC232FB8D8DF6FB32" authority=", Miguelsoriaparayirunhor" authorityName="Miguelsoriaparayirunhor" class="Mammalia" family="Protolipternidae" genus="Protolipterna" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="ellipsodontoides">
<emphasis id="B964DA31FFF30D0AC232FB8D8AF6FB32" italics="true" pageId="21" pageNumber="22">Protolipterna ellipsodontoides</emphasis>
,
<emphasis id="B964DA31FFF30D0AC4CEFBAD8DF6FB32" box="[972,1220,1147,1171]" italics="true" pageId="21" pageNumber="22">Miguelsoriaparayirunhor</emphasis>
</taxonomicName>
,and
<taxonomicName id="4C107DA0FFF30D0AC3EDFBAD8A6EFB13" authorityName="Paula Couto" authorityYear="1952" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="scotti">
<emphasis id="B964DA31FFF30D0AC3EDFBAD8A6EFB13" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia scotti</emphasis>
</taxonomicName>
from Itaboraí,
<collectingCountry id="F30746B3FFF30D0AC4FAFB4C8D04FB13" box="[1016,1078,1178,1202]" name="Brazil" pageId="21" pageNumber="22">Brazil</collectingCountry>
, and the youngest member would be
<taxonomicName id="4C107DA0FFF30D0AC429FB6F8D68FB70" authorityName="Ameghino" authorityYear="1901" box="[811,1114,1209,1233]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="subtrigona">
<emphasis id="B964DA31FFF30D0AC429FB6F8D68FB70" box="[811,1114,1209,1233]" italics="true" pageId="21" pageNumber="22">Asmithwoodwardia subtrigona</emphasis>
</taxonomicName>
with its last records in Cañadón Vaca,
<collectingCountry id="F30746B3FFF30D0AC46BFB0F8AE2FB50" box="[873,976,1241,1265]" name="Argentina" pageId="21" pageNumber="22">Argentina</collectingCountry>
(
<bibRefCitation id="EF817BD2FFF30D0AC4E6FB0F8D69FB50" author="Gelfo" box="[996,1115,1241,1265]" firstAuthor="Gelfo" pageId="21" pageNumber="22" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo 2006</bibRefCitation>
), which gives the family a temporal interval of 56–42 Mya (
<figureCitation id="132B1AA6FFF30D0AC35CFB2E8D95FAB1" box="[1118,1191,1272,1296]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="21" pageNumber="22">Fig. 2B</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFF30D0AC429FAC18AB4FA8E" box="[811,902,1303,1327]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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>
).
</paragraph>
</subSubSection>
</treatment>
</document>

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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, P4–M3 of Molinodus suarezi in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of Escribania chubutensis in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of Raulvaccia peligrensis [MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of Ricardocifellia protocenica in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of Lamegoia conodonta Paula Couto, 1952 in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of Didolodus multicuspis in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of Protolipterna ellipsodontoides in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of Asmithwoodwardia scotti in occlusal view [DGM 358M (holotype; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of Miguelsoria parayiruhnor in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of Molinodus suarezi (A) and the M2 of Lamegoia conodonta (B) present a duplication of their protocone (pr1 and pr2); in the former, this is insipient, and in the latter, it forms a pseudohypocone. For information about the tooth position convention, check the Material and methods. Abbreviations: hy, hypocone; hys, hypostyle; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; phy, pseudohypocone; phyc, prehypocrista; pshyc, posthypocrista; pmlc, premetaconular crista; pplc, preparaconular crista; pr, protocone; prl, paraconule; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista.Scale bars equal 1 cm." pageId="17" pageNumber="18">Figs 8B–G</figureCitation>
,
<figureCitation id="132B1AA6FFF70D0EC3D4FCEC8C22FCF5" box="[1238,1296,826,852]" captionStart="Figure 9" captionStartId="22.[129,194,1822,1846]" captionTargetBox="[241,1361,147,1792]" captionTargetId="figure-7@22.[238,1364,144,1795]" captionTargetPageId="22" captionText="Figure 9. Right lower molars of South American ‘condylarths’ and ‘didolodontids’ in occlusal view.A, p4–m3 of Molinodus suarezi [cast of MHNC 13867 (mirrored)]. B, m2–m3 of Escribania chubutensis [UNPSJB PV 916 (holotype; mirrored)]. C, talonid of m2, and m3 of Raulvaccia peligrensis [MLP 90−II−12−69 (mirrored)]. D, P5–M3 of Ricardocifellia protocenica [MNRJ 1431-V, m1–m3 (mirrored); MNRJ 1450-V, p5 (mirrored)]. E, p2–m3 of Didolodus multicuspis (MACN A 10689). F, p2–m3 of Protolipterna ellipsodontoides [cast of DGM 1308M (mirrored)]. G, p1–m3 of Asmithwoodwardia scotti (cast of DGM 358M (holotype; mirrored)]. H, p4–m3 of Miguelsoria parayiruhnor (cast of MNRJ 1468V (holotype)]. More information on the specimens and observations are in Supporting information, File S2. For information about the tooth position convention, check the Material and methods.Abbreviations:encd, entocristid; end, entoconid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; msd, mesoconid; pad, paraconid; peld, preentoconulid; pmcd, premetacristid; prd, protoconid; pseld, postentoconulid. Scale bars equal 1 cm." pageId="17" pageNumber="18">9B–E</figureCitation>
)
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<subSubSection id="C30A55A8FFF70D0AC428FCB78839FDDF" lastPageId="21" lastPageNumber="22" pageId="17" pageNumber="18" type="multiple">
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<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 Danian–Late Lutetian in age (
<bibRefCitation id="EF817BD2FFF50D0CC5B1FF6689DCFF46" author="Krause" etAl="et al." firstAuthor="Krause" pageId="19" pageNumber="20" pagination="886 - 903" refId="ref43716" refString="Krause JM, Clyde WC, Ibanez-Mejia M et al. New age constraints for Early Paleogene strata of central Patagonia, Argentina: implications for the timing of South American Land Mammal Ages. Geological Society of America Bulletin 2017; 129: 886 - 903. https: // doi. org / 10.1130 / b 31561.1" type="journal article" year="2017">
Krause
<emphasis id="B964DA31FFF50D0CC773FF198999FF46" box="[113,171,207,231]" italics="true" pageId="19" pageNumber="20">et al.</emphasis>
2017
</bibRefCitation>
)] of the genus
<taxonomicName id="4C107DA0FFF50D0CC6B4FF198B78FF46" authorityName="Ameghino" authorityYear="1901" box="[438,586,207,231]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF50D0CC6B4FF198B78FF46" box="[438,586,207,231]" italics="true" pageId="19" pageNumber="20">Ernestokokenia</emphasis>
</taxonomicName>
(
<taxonomicName id="4C107DA0FFF50D0CC564FF198858FEA7" authority="Simpson, 1935" authorityName="Simpson" authorityYear="1935" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="yirunhor">
<emphasis id="B964DA31FFF50D0CC564FF1989F4FEA7" italics="true" pageId="19" pageNumber="20">Ernestokokenia yirunhor</emphasis>
Simpson, 1935
</taxonomicName>
and
<taxonomicName id="4C107DA0FFF50D0CC6A3FF388997FE84" authority="Simpson 1935" authorityName="Simpson" authorityYear="1935" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="chaishoer">
<emphasis id="B964DA31FFF50D0CC6A3FF388BAAFEA7" box="[417,664,238,262]" italics="true" pageId="19" pageNumber="20">Ernestokokenia chaishoer</emphasis>
Simpson 1935
</taxonomicName>
), Simpson (1948) revised the whole group, giving a new definition for
<taxonomicName id="4C107DA0FFF50D0CC644FEFB88D8FEE4" box="[326,490,301,325]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName>
and emending many of
<bibRefCitation id="EF817BD2FFF50D0CC773FE9A8808FEC5" author="Ameghino's" box="[113,314,332,356]" firstAuthor="Ameghino's" pageId="19" pageNumber="20" pagination="1 - 568" refId="ref39456" refString="Ameghino F. Les formations sedimentaires du cretace superieur et du tertiaire de Patagonie avec un parallele entre leurs faunes mammalogiques et celles de l'ancien continent. Anales del Museo Nacional de Buenos Aires 1906; 15: 1 - 568." type="journal article" year="1906">Ameghino’s (1906)</bibRefCitation>
taxonomic classifications of the Patagonian taxa, including the following genera in the family:
<taxonomicName id="4C107DA0FFF50D0CC593FEBD8BC7FE22" authorityName="Ameghino" authorityYear="1897" box="[657,757,363,387]" class="Mammalia" family="Didolodontidae" genus="Didolodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF50D0CC593FEBD8BC7FE22" box="[657,757,363,387]" italics="true" pageId="19" pageNumber="20">Didolodus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C107DA0FFF50D0CC773FE5D8837FE02" authorityName="Ameghino" authorityYear="1901" box="[113,261,395,419]" class="Mammalia" family="Didolodontidae" genus="Ernestokokenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF50D0CC773FE5D8837FE02" box="[113,261,395,419]" italics="true" pageId="19" pageNumber="20">Ernestokokenia</emphasis>
</taxonomicName>
,
<taxonomicName id="4C107DA0FFF50D0CC616FE5D8B6DFE02" authority="Ameghino 1904 b" authorityName="Ameghino" authorityYear="1904" box="[276,607,395,419]" class="Mammalia" family="Didolodontidae" genus="Argyrolambda" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF50D0CC616FE5D8890FE02" box="[276,418,395,419]" italics="true" pageId="19" pageNumber="20">Argyrolambda</emphasis>
<bibRefCitation id="EF817BD2FFF50D0CC6A8FE5D8B6DFE02" author="Ameghino" box="[426,607,395,419]" firstAuthor="Ameghino" pageId="19" pageNumber="20" pagination="1 - 541" refId="ref39426" refString="Ameghino F. Recherches de morphologic phylogenetique sur les molaires superieures des ongules. Anales del Museo Nacional de Buenos Aires 1904 b; 3: 1 - 541." type="journal article" year="1904" yearSuffix="b">Ameghino 1904b</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C107DA0FFF50D0CC56FFE5D8817FE63" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" class="Mammalia" family="Didolodontidae" genus="Paulogervaisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF50D0CC56FFE5D8BC8FE02" box="[621,762,395,419]" italics="true" pageId="19" pageNumber="20">Paulogervaisia</emphasis>
<bibRefCitation id="EF817BD2FFF50D0CC773FE7C8817FE63" author="Ameghino" box="[113,293,426,450]" firstAuthor="Ameghino" pageId="19" pageNumber="20" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C107DA0FFF50D0CC638FE7D889BFE63" authorityName="Ameghino" authorityYear="1904" box="[314,425,427,450]" class="Mammalia" family="Adianthidae" genus="Proectocion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF50D0CC638FE7D889BFE63" box="[314,425,427,450]" italics="true" pageId="19" pageNumber="20">Proectocion</emphasis>
</taxonomicName>
,
<taxonomicName id="4C107DA0FFF50D0CC6BDFE7D8BC7FE63" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[447,757,426,450]" class="Mammalia" family="Didolodontidae" genus="Enneoconus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF50D0CC6BDFE7D8B03FE63" box="[447,561,427,450]" italics="true" pageId="19" pageNumber="20">Enneoconus</emphasis>
<bibRefCitation id="EF817BD2FFF50D0CC543FE7C8BC7FE63" author="Ameghino" box="[577,757,426,450]" firstAuthor="Ameghino" pageId="19" pageNumber="20" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
</taxonomicName>
,
<emphasis id="B964DA31FFF50D0CC773FE1F8819FE40" box="[113,299,457,481]" italics="true" pageId="19" pageNumber="20">Asmithwoodwarsia</emphasis>
<bibRefCitation id="EF817BD2FFF50D0CC636FE1F88D3FE40" author="Ameghino" box="[308,481,457,481]" firstAuthor="Ameghino" pageId="19" pageNumber="20" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
, and
<taxonomicName id="4C107DA0FFF50D0CC51CFE1F8828FDA0" authority="Ameghino, 1906" authorityName="Ameghino" authorityYear="1906" class="Mammalia" family="Didolodontidae" genus="Archaeohyracotherium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="B964DA31FFF50D0CC51CFE1F8BC8FE40" box="[542,762,457,481]" italics="true" pageId="19" pageNumber="20">Archaeohyracotherium</emphasis>
<bibRefCitation id="EF817BD2FFF50D0CC773FE3F8828FDA0" author="Ameghino" box="[113,282,489,513]" firstAuthor="Ameghino" pageId="19" pageNumber="20" pagination="1 - 568" refId="ref39456" refString="Ameghino F. Les formations sedimentaires du cretace superieur et du tertiaire de Patagonie avec un parallele entre leurs faunes mammalogiques et celles de l'ancien continent. Anales del Museo Nacional de Buenos Aires 1906; 15: 1 - 568." type="journal article" year="1906">Ameghino, 1906</bibRefCitation>
</taxonomicName>
. Simpson soon recognized the close similarities between this group and North American ‘condylarths’, including species of
<taxonomicName id="4C107DA0FFF50D0CC7D8FDF188BFFD9E" box="[218,397,551,575]" class="Mammalia" family="Phenacodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Phenacodontidae</taxonomicName>
and
<taxonomicName id="4C107DA0FFF50D0CC6BDFDF18B79FD9E" box="[447,587,551,575]" class="Mammalia" family="Mioclaenidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Mioclaenidae</taxonomicName>
, stating that they were impossible to separate on a purely morphological basis (Simpson 1948, 1980).
</paragraph>
<caption id="DF6F56ABFFF40D0CC783F8968A28F81B" lastPageId="19" lastPageNumber="20" pageId="18" pageNumber="19" startId="18.[129,194,1856,1880]" targetBox="[177,1425,147,1825]" targetPageId="18" targetType="figure">
<paragraph id="8BAF0623FFF40D0DC783F8968CBFF831" blockId="18.[129,1464,1856,1964]" pageId="18" pageNumber="19">
<emphasis id="B964DA31FFF40D0DC783F89689EAF8F9" bold="true" box="[129,216,1856,1880]" pageId="18" pageNumber="19">Figure 7.</emphasis>
Right lower molars of relevant SANUs in occlusal view. A, p1–m3 of the indaleciid
<taxonomicName id="4C107DA0FFF40D0DC4EAF8968DAFF8F9" authorityName="Bond &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 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid
<taxonomicName id="4C107DA0FFF40D0DC518F88A8BE0F8D5" authorityName="Soria" authorityYear="1989" box="[538,722,1884,1908]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="powelli">
<emphasis id="B964DA31FFF40D0DC518F88A8BE0F8D5" box="[538,722,1884,1908]" italics="true" pageId="18" pageNumber="19">Notonychops powelli</emphasis>
</taxonomicName>
[PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate
<taxonomicName id="4C107DA0FFF40D0DC3E6F88A8C8AF8D5" box="[1252,1464,1884,1908]" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="praecursor">
<emphasis id="B964DA31FFF40D0DC3E6F88A8C8AF8D5" box="[1252,1464,1884,1908]" italics="true" pageId="18" pageNumber="19">Simpsonotus praecursor</emphasis>
</taxonomicName>
[MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere
<taxonomicName id="4C107DA0FFF40D0DC42CF8AE8D3CF831" authorityName="Ameghino" authorityYear="1897" box="[814,1038,1912,1936]" class="Mammalia" family="Trigonostylopidae" genus="Trigonostylops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astrapotheria" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="wortmani">
<emphasis id="B964DA31FFF40D0DC42CF8AE8D3CF831" box="[814,1038,1912,1936]" italics="true" pageId="18" pageNumber="19">Trigonostylops wortmani</emphasis>
</taxonomicName>
[MNHN.F.CAS188 (mirrored)]. E, p2–
</paragraph>
<paragraph id="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, p1–m3 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, p1–m2 (
<typeStatus id="54ABB881FFF50D0CC778F92C89E4F8B3" box="[122,214,1786,1810]" pageId="19" pageNumber="20" type="holotype">holotype</typeStatus>
; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 of
<taxonomicName id="4C107DA0FFF50D0CC5C1F92C8AAEF8B3" authorityName="Ameghino" authorityYear="1901" box="[707,924,1786,1810]" class="Mammalia" family="Pyrotheriidae" genus="Propyrotherium" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="saxeum">
<emphasis id="B964DA31FFF50D0CC5C1F92C8AAEF8B3" box="[707,924,1786,1810]" italics="true" pageId="19" pageNumber="20">Propyrotherium saxeum</emphasis>
</taxonomicName>
[LIEB-PV 3200 (mirrored)]. Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H
<quantity id="4CE8ABC6FFF50D0CC5FDF8E58A11F8EB" box="[767,803,1842,1867]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="19" pageNumber="20" unit="in" value="2.0">2 in</quantity>
<taxonomicName id="4C107DA0FFF50D0CC42AF8E58ACEF8EB" box="[808,1020,1843,1866]" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="praecursor">
<emphasis id="B964DA31FFF50D0CC42AF8E58ACEF8EB" box="[808,1020,1843,1866]" italics="true" pageId="19" pageNumber="20">Simpsonotus praecursor</emphasis>
</taxonomicName>
, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of
<taxonomicName id="4C107DA0FFF50D0CC504F8998BE8F8C7" box="[518,730,1871,1894]" class="Mammalia" family="Henricosborniidae" genus="Simpsonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="praecursor">
<emphasis id="B964DA31FFF50D0CC504F8998BE8F8C7" box="[518,730,1871,1894]" italics="true" pageId="19" pageNumber="20">Simpsonotus praecursor</emphasis>
</taxonomicName>
is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid. For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal
<quantity id="4CE8ABC6FFF50D0CC5E4F8758A24F81B" box="[742,790,1955,1979]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="19" pageNumber="20" unit="cm" value="1.0">1 cm</quantity>
.
</paragraph>
</caption>
<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 (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="19" pageNumber="20">Fig. 1C</figureCitation>
), but these were based on tarsal associations that have been questioned by later authors (e.g.,Soria 2001,
<bibRefCitation id="EF817BD2FFF50D0CC24DFCF48ABCFCF8" author="Gelfo and Sige" firstAuthor="Gelfo" pageId="19" pageNumber="20" pagination="665 - 78" refId="ref42458" refString="Gelfo JN, Sige B. A new didolodontid mammal from the Late Paleocene - earliest Eocene of Laguna Umayo, Peru. Acta Palaeontologica Polonica 2011; 56: 665 - 78. https: // doi. org / 10.4202 / app. 2010.0067" type="journal article" year="2011">Gelfo and Sigé 2011</bibRefCitation>
; see also
<taxonomicName id="4C107DA0FFF50D0CC4F7FC978DE5FCF8" box="[1013,1239,833,857]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
and
<taxonomicName id="4C107DA0FFF50D0CC209FC978C81FCF8" box="[1291,1459,833,857]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Protolipternidae</taxonomicName>
sections). Muizon and Cifelli (2000), in a phylogenetic analysis that included North American mioclaenids, South American kollpaniines, didolodontids, and protolipternids, found that protolipternids were nested within
<taxonomicName id="4C107DA0FFF50D0CC3ABFC698C7FFC76" authorityName=", Gelfo" authorityYear="2006" box="[1193,1357,959,983]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName>
and that both groups together formed the sister group of kollpanines, a result that was also found in a later study (
<bibRefCitation id="EF817BD2FFF50D0CC3F8FC2B8C5DFBB4" author="Gelfo" box="[1274,1391,1021,1045]" firstAuthor="Gelfo" pageId="19" pageNumber="20" refId="ref42195" refString="Gelfo JN. Los Didolodontidae (Mammalia: Ungulatomorpha) del Terciario Sudamericano. Sistematica, Origen y Evolucion. D. Phil. Thesis, Universidad Nacional de la Plata. 2006." type="book" year="2006">Gelfo 2006</bibRefCitation>
). This means that
<taxonomicName id="4C107DA0FFF50D0CC4A2FBCB8D76FB94" authorityName=", Gelfo" authorityYear="2006" box="[928,1092,1053,1077]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Didolodontidae</taxonomicName>
excluding
<taxonomicName id="4C107DA0FFF50D0CC3B2FBCB8C6AFB94" box="[1200,1368,1053,1077]" class="Mammalia" family="Protolipternidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="19" pageNumber="20" phylum="Chordata" rank="family">Protolipternidae</taxonomicName>
is a paraphyletic group. Other phylogenetic studies that have included didolodontids in very small matrices have focused on their internal relationships, assuming their monophyly (e.g.,
<bibRefCitation id="EF817BD2FFF50D0CC27BFBAD8A6CFB13" author="Gelfo" firstAuthor="Gelfo" pageId="19" pageNumber="20" pagination="130 - 42" refId="ref42288" refString="Gelfo JN. The ' condylarth' Didolodontidae from Gran Barranca: history of the bunodont South American mammals until the Eocene- Oligocene transition. In: Madden RH, Carlini AA, Vucetich MG, Kay RF (eds.), The Paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge: Cambridge University Press, 2010, 130 - 42." type="book chapter" year="2010">Gelfo 2010</bibRefCitation>
;
<tableCitation id="C6923398FFF50D0CC469FB4C8A87FB13" box="[875,949,1178,1202]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" 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 (O’Leary
<emphasis id="B964DA31FFF50D0CC3EBFA818C2EFACF" box="[1257,1308,1366,1390]" italics="true" pageId="19" pageNumber="20">et al.</emphasis>
2013)
</taxonomicName>
. When a larger sample of didolodontids has been included, they have been found at the stem of litopterns but not forming a monophyletic
<taxonomicName id="4C107DA0FFF50D0CC480FA628C35FA6D" authority="(Kramarz et al. 2021)" baseAuthorityName="Kramarz" baseAuthorityYear="2021" box="[898,1287,1460,1484]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="19" pageNumber="20" phylum="Chordata" rank="family">
Didolodontidae (
<bibRefCitation id="EF817BD2FFF50D0CC334FA628DCEFA6D" author="Kramarz" box="[1078,1276,1460,1484]" etAl="et al." firstAuthor="Kramarz" pageId="19" pageNumber="20" refId="ref43620" refString="Kramarz AG, Bond M, MacPhee RDE. On the alleged perissodactyl affinities of the ' condylarth' Escribania chubutensis and other endemic South American ungulate-like placentals. Journal of Vertebrate Paleontology 2021; 41: e 1986716." type="journal volume" year="2021">
Kramarz
<emphasis id="B964DA31FFF50D0CC396FA638DF1FA6D" box="[1172,1219,1460,1484]" italics="true" pageId="19" pageNumber="20">et al.</emphasis>
2021
</bibRefCitation>
)
</taxonomicName>
, or as a paraphyletic group among SANUs and North American ‘condylarths’ (Zimicz
<emphasis id="B964DA31FFF50D0CC481FA258A86F9AA" box="[899,948,1523,1547]" italics="true" pageId="19" pageNumber="20">et al.</emphasis>
2023). In sum, there is still a considerable uncertainty in the phylogenetic position of didolodontids and their relationship among SANUs, and whether they form a monophyletic group. In fact, there has not been any phylogenetic analysis that includes didolodontids alongside representatives of all litoptern families, protolipternids, and the different SANUs’ 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, P4–M3 of
<taxonomicName id="4C107DA0FFF20D0BC30FF9238D83F8AC" authorityName="Muizon and Marshall" authorityYear="1987" box="[1037,1201,1781,1805]" class="Mammalia" family="Mioclaenidae" genus="Molinodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="suarezi">
<emphasis id="B964DA31FFF20D0BC30FF9238D83F8AC" box="[1037,1201,1781,1805]" italics="true" pageId="20" pageNumber="21">Molinodus suarezi</emphasis>
</taxonomicName>
in occlusal view [cast of MHNC 1247, P4–M2; cast of MNHN 13870, M3 (mirrored)]. B, roots of M1, M2–M3 of
<taxonomicName id="4C107DA0FFF20D0BC4DEF8C78D95F888" box="[988,1191,1809,1833]" class="Mammalia" family="Mioclaenidae" genus="Escribania" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="chubutensis">
<emphasis id="B964DA31FFF20D0BC4DEF8C78D95F888" box="[988,1191,1809,1833]" italics="true" pageId="20" pageNumber="21">Escribania chubutensis</emphasis>
</taxonomicName>
in occlusal view (MLP 90–II-12–68, roots of M1 and M2 (mirrored); MLP 90–II-12–63, M3). C–D, M2 of
<taxonomicName id="4C107DA0FFF20D0BC4A3F8FB8D56F8E4" box="[929,1124,1837,1861]" class="Mammalia" family="Mioclaenidae" genus="Raulvaccia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="peligrensis">
<emphasis id="B964DA31FFF20D0BC4A3F8FB8D56F8E4" box="[929,1124,1837,1861]" italics="true" pageId="20" pageNumber="21">Raulvaccia peligrensis</emphasis>
</taxonomicName>
[MLP 90-II-12- 70 (mirrored)] in occlusal (C) and distoocclusal (D) views. E, P4–M3 of
<taxonomicName id="4C107DA0FFF20D0BC5A0F89F8ABFF8C0" baseAuthorityName="Mones" baseAuthorityYear="2015" box="[674,909,1865,1889]" class="Mammalia" family="Didolodontidae" genus="Ricardocifellia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="protocenica">
<emphasis id="B964DA31FFF20D0BC5A0F89F8ABFF8C0" box="[674,909,1865,1889]" italics="true" pageId="20" pageNumber="21">Ricardocifellia protocenica</emphasis>
</taxonomicName>
in occlusal view [cast of DNPM 908M, P5–M3 (mirrored); cast of MNRJ 1461-V (mirrored)]. F, M2 of
<taxonomicName id="4C107DA0FFF20D0BC595F8B08D37F8DC" authority="Paula Couto, 1952" authorityName="Paula Couto" authorityYear="1952" box="[663,1029,1893,1917]" class="Mammalia" family="Didolodontidae" genus="Lamegoia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="conodonta">
<emphasis id="B964DA31FFF20D0BC595F8B08A60F8DC" box="[663,850,1893,1917]" italics="true" pageId="20" pageNumber="21">Lamegoia conodonta</emphasis>
Paula Couto, 1952
</taxonomicName>
in occlusal view (cast of MNRJ 1465-V). G, P1–M3 of
<taxonomicName id="4C107DA0FFF20D0BC601F85788FBF838" authorityName="Ameghino" authorityYear="1897" box="[259,457,1921,1945]" class="Mammalia" family="Didolodontidae" genus="Didolodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="multicuspis">
<emphasis id="B964DA31FFF20D0BC601F85788FBF838" box="[259,457,1921,1945]" italics="true" pageId="20" pageNumber="21">Didolodus multicuspis</emphasis>
</taxonomicName>
in occlusal view [MACN 10690, P2–M3 (holotype; mirrored); MACN A 10738, P1]. H, P5–M3 of
<taxonomicName id="4C107DA0FFF20D0BC781F84B88BEF814" authorityName="Cifelli" authorityYear="1983" box="[131,396,1949,1973]" class="Mammalia" family="Protolipternidae" genus="Protolipterna" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="ellipsodontoides">
<emphasis id="B964DA31FFF20D0BC781F84B88BEF814" box="[131,396,1949,1973]" italics="true" pageId="20" pageNumber="21">Protolipterna ellipsodontoides</emphasis>
</taxonomicName>
in occlusal view [DNPM LE 444I (mirrored)]. I, P4–M3 of
<taxonomicName id="4C107DA0FFF20D0BC4C6F84B8D95F814" authorityName="Paula Couto" authorityYear="1952" box="[964,1191,1949,1973]" class="Mammalia" family="Protolipternidae" genus="Asmithwoodwardia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="scotti">
<emphasis id="B964DA31FFF20D0BC4C6F84B8D95F814" box="[964,1191,1949,1973]" italics="true" pageId="20" pageNumber="21">Asmithwoodwardia scotti</emphasis>
</taxonomicName>
in occlusal view [DGM 358M (
<typeStatus id="54ABB881FFF30D0AC7B5F92D8821F8B2" box="[183,275,1787,1811]" pageId="21" pageNumber="22" type="holotype">holotype</typeStatus>
; mirrored)]. J, P2, alveoli of P4–P5, and M1–M3 of
<taxonomicName id="4C107DA0FFF30D0AC5FEF92D8AD1F8B2" baseAuthorityName="Paula Couto" baseAuthorityYear="1952" box="[764,995,1787,1811]" class="Mammalia" family="Protolipternidae" genus="Miguelsoria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="parayiruhnor">
<emphasis id="B964DA31FFF30D0AC5FEF92D8AD1F8B2" box="[764,995,1787,1811]" italics="true" pageId="21" pageNumber="22">Miguelsoria parayiruhnor</emphasis>
</taxonomicName>
in occlusal view [MNRJ 4094V (mirrored)]. More information on the specimens and observations are in Supporting information, File S2. Note that the M2 of
<taxonomicName id="4C107DA0FFF30D0AC3A3F8C18C77F88F" authorityName="Muizon and Marshall" authorityYear="1987" box="[1185,1349,1815,1839]" class="Mammalia" family="Mioclaenidae" genus="Molinodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="suarezi">
<emphasis id="B964DA31FFF30D0AC3A3F8C18C77F88F" box="[1185,1349,1815,1839]" italics="true" pageId="21" pageNumber="22">Molinodus suarezi</emphasis>
</taxonomicName>
(A) 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 63.8–36.5 Mya for this family (
<figureCitation id="132B1AA6FFF30D0AC7C0FD918838FDFE" box="[194,266,583,607]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="21" pageNumber="22">Fig. 2B</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFF30D0AC51AFD918B41FDFE" box="[536,627,583,607]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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>

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<mods:affiliation id="D33C800582BEC8D88903F7369C5EE084">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:title id="6039ED8BC04B4053EC82C11C75E03C2B">Zoological Journal of the Linnean Society</mods:title>
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<mods:part id="A789A05EC5097D46F8960074C5075AF0">
<mods:date id="087FA6D96E8C7880D08F5C11202C45B5">2024</mods:date>
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<mods:number id="CC7602A0432CB7A140263C7CFC9F978F">2024-09-07</mods:number>
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<mods:number id="5AF9E54E1E5216E8EB58112911057B59">202</mods:number>
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<mods:identifier id="8888FBE246630837DFAA09E6651C5798" type="DOI">10.1093/zoolinnean/zlae095</mods:identifier>
<mods:identifier id="FFBCF1E03A2B78AB3F9F7EEC19DF6FE1" type="ISSN">0024-4082</mods:identifier>
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<emphasis id="B964DA31FFF70D0EC614FD898B64FDD8" box="[278,598,607,633]" italics="true" pageId="17" pageNumber="18">
<subSubSection id="C30A55A8FFF70D0EC614FD898882FDD8" box="[278,432,607,633]" pageId="17" pageNumber="18" type="nomenclature">
<paragraph id="8BAF0623FFF70D0EC614FD898882FDD8" blockId="17.[113,764,607,1547]" box="[278,432,607,633]" pageId="17" pageNumber="18">
<taxonomicName id="4C107DA0FFF70D0EC614FD898882FDD8" authority="(Soria, 1989)" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[278,432,607,633]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="C30A55A8FFF70D0EC6B7FD898B64FDD8" box="[437,598,607,633]" pageId="17" pageNumber="18" type="description">
<paragraph id="8BAF0623FFF70D0EC6B7FD898B64FDD8" blockId="17.[113,764,607,1547]" box="[437,598,607,633]" pageId="17" pageNumber="18">
(
<figureCitation id="132B1AA6FFF70D0EC6BDFDB68B14FDD8" box="[447,550,607,633]" captionStart="Figure 6" captionStartId="16.[129,194,1811,1835]" captionTargetBox="[193,1409,146,1780]" captionTargetId="figure-7@16.[190,1412,143,1783]" captionTargetPageId="16" captionText="Figure 6. Right upper molars of relevant SANUs. A, P1–M3 of the indaleciid Indalecia grandensis in occlusal view [PVL 4186 (mirrored)]. B, crownless P1, and P2–M3 of the notonychopid Notonychops powelli in occlusal view [PVL 4298 (M1–M3 mirrored)]. C, crownless and broken M2, broken M1, and P5–P2 of the notonychopid Requisia vidmari in occlusal view (UNPSJB PV 944, crownless and broken M2, broken parastyle of M1, and P4–P5 (holotype); UNPSJB PV 947, broken M1; UNPSJB PV 945, P2). D, P1–M3 of Eoastrapostylops riolorense in occlusal view [PVL 4216 (holotype)]. E, P1–M3 of the notoungulate Simpsonotus praecursor in occlusal view [MLP 73-VII-3-11 (holotype; mirrored)]. F–G, unworn M1 or M2 of the notoungulate Henricosbornia sp. (MACN A 10717) in occlusal (F) and distoocclusal (G) views.H, P2–M3 of the astrapothere Trigonostylops wortmani (AMNH VP-28700). I, P5–M3 of the xenungulate Carodnia vieirai [cast of DGM 335M, P5–M2 (mirrored); cast of AMNH VP-49828, M3 (mirrored)]. J, P5–M3 of Propyrotherium saxeum [AMNH unnumbered (labelled as ‘O3’)], right M3 (there is a mesiobuccal portion missing); MLP 55-III-10-1a, right M1 or M2 (duplicated for illustrating M1–M2); MACN A 10929 (lectotype), left P5 (lectotype; mirrored). Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisks mark the hypocone and prehypocrista in notoungulates, as they could potentially not be homologous with the hypocone and prehypocrista in other SANUs, considering a different origin of the hypocone in notoungulates (see more in the Material and methods and in Supporting information, File S2). For information about the tooth position convention, check the Material and methods.Abbreviations:efx, ectoflexus; hy, hypocone; me, metacone; ms, mesostyle; mtl, metaconule; pa, paracone; pcg, precingulum; phyc, prehypocrista; pmlc, premetaconular crista (or crochet in notoungulates); pplc, preparaconular crista; pr, protocone; prl, paraconule; prt, protostyle; ps, parastyle; pscg, postcingulum; psmlc, postmetaconular crista; psprc, postprotocrista.Scale bars equal 1 cm." pageId="17" pageNumber="18">Figs 6B–C</figureCitation>
,
<figureCitation id="132B1AA6FFF70D0EC52DFD898B7EFDD8" box="[559,588,607,633]" captionStart="Figure 7" captionStartId="18.[129,194,1856,1880]" captionTargetBox="[177,1425,147,1825]" captionTargetId="figure-8@18.[174,1428,144,1828]" captionTargetPageId="18" captionText="Figure 7. Right lower molars of relevant SANUs in occlusal view.A, p1–m3 of the indaleciid Indalecia grandensis [PVL 4186 (p1–p2 mirrored)]. B, p2–m3 of the notonychopid Notonychops powelli [PVL 4298 (mirrored)]. C, p1–m3 of the notoungulate Simpsonotus praecursor [MLP 73-VII-3-11 (holotype; mirrored)]. D, p1–m3 of the astrapothere Trigonostylops wortmani [MNHN.F.CAS188 (mirrored)]. E, p2– m3 of the xenungulate Etayoa bacatensis [cast of GM 32 (holotype; mirrored)]. F, p1–m3 of Eoastrapostylops riolorense [PVL 4216, p1–m2 (holotype; p1–p5, m2 mirrored); PVL 4216, m3. G, p4–m3 ofPropyrotherium saxeum [LIEB-PV 3200 (mirrored)].Relevant anatomical features of the dentition are labelled. More information on the specimens and observations are in Supporting information, File S2. The asterisk is labelling either a paracristid in H1 or a very reduced paraconid in H2 in Simpsonotus praecursor, the tested hypotheses in our analyses. The question mark (?) next to the paraconid of Simpsonotus praecursor is to reflect that in the H1 matrix, this cusp was scored as a paraconid, whereas in the H2 matrix it was scored as a twinned metaconid.For information about the tooth position convention, check the Material and methods. Abbreviations: end, entoconid; hlph, hypolophid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; pad, paraconid; prd, protoconid; pscd, postcristid (or postentocristid). Scale bars equal 1 cm." pageId="17" pageNumber="18">7B</figureCitation>
)
</paragraph>
</subSubSection>
</emphasis>
<subSubSection id="C30A55A8FFF70D0EC773FD538C1AFCBA" pageId="17" pageNumber="18" type="multiple">
<paragraph id="8BAF0623FFF70D0EC773FD538D4AFF46" blockId="17.[113,764,607,1547]" lastBlockId="17.[809,1461,144,795]" pageId="17" pageNumber="18">
<taxonomicName id="4C107DA0FFF70D0EC773FD53882FFD3C" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[113,285,645,669]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName>
currently includes two accepted genera with an exclusively Palaeogene distribution (
<figureCitation id="132B1AA6FFF70D0EC516FD738B62FD1C" box="[532,592,677,701]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="17" pageNumber="18">Fig. 2</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFF70D0EC7E6FD128872FD7D" box="[228,320,708,732]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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>
).
<taxonomicName id="4C107DA0FFF70D0EC657FD128B33FD7D" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[341,513,708,732]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName>
was founded to include
<taxonomicName id="4C107DA0FFF70D0EC771FD358888FD5A" authority="Soria, 1989" authorityName="Soria" authorityYear="1989" box="[115,442,739,763]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="powelli">
<emphasis id="B964DA31FFF70D0EC771FD35880EFD5A" box="[115,316,739,763]" italics="true" pageId="17" pageNumber="18">Notonychops powelli</emphasis>
Soria, 1989
</taxonomicName>
from the Rio Loro Formation [age uncertain, although probably pre-Itaboraian SALMA age (
<bibRefCitation id="EF817BD2FFF70D0EC77EFCF48808FC9B" author="Gelfo" box="[124,314,802,826]" etAl="et al." firstAuthor="Gelfo" pageId="17" pageNumber="18" pagination="993 - 1007" refId="ref42730" refString="Gelfo JN, Garcia-Lopez DA, Bergqvist LP. Phylogenetic relationships and palaeobiology of a new xenungulate (Mammalia: Eutheria) from the Palaeogene of Argentina. Journal of Systematic Palaeontology 2020 b; 18: 993 - 1007. https: // doi. org / 10.1080 / 14772019.2020.171 5496" type="journal article" year="2020" yearSuffix="b">
Gelfo
<emphasis id="B964DA31FFF70D0EC7BDFCF589C0FC9B" box="[191,242,802,826]" italics="true" pageId="17" pageNumber="18">et al.</emphasis>
2020b
</bibRefCitation>
)], an ungulate with great development of the parastyle in M1–M3, which among other features, makes it anatomically distinctive from any other SANU previously described (Soria 1989b). Soria (1989b) considered that
<taxonomicName id="4C107DA0FFF70D0EC5E1FC578987FC16" authorityName="Soria" authorityYear="1989" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="powelli">
<emphasis id="B964DA31FFF70D0EC5E1FC578987FC16" italics="true" pageId="17" pageNumber="18">N. powelli</emphasis>
</taxonomicName>
was dentally convergent with members of the family
<taxonomicName id="4C107DA0FFF70D0EC773FC6988A1FC76" authority="(Tillodonta)" baseAuthorityName="Tillodonta" box="[113,403,959,983]" class="Mammalia" family="Esthonychidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Esthonychidae (Tillodonta)</taxonomicName>
, particularly with the lower molars being brachydont and selenodont with a subequal trigonid and talonid length. However,
<taxonomicName id="4C107DA0FFF70D0EC670FC2888E1FBB4" authorityName="Soria" authorityYear="1989" box="[370,467,1021,1045]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="powelli">
<emphasis id="B964DA31FFF70D0EC670FC2888E1FBB4" box="[370,467,1021,1045]" italics="true" pageId="17" pageNumber="18">N. powelli</emphasis>
</taxonomicName>
shares important anatomical features with the dentition of ‘amilnedwarsids’ and indaleciids, and of the ear region with indaleciids, which madeSoria (1989b) place it within the order
<taxonomicName id="4C107DA0FFF70D0EC686FB8A88CEFBD2" authorityName="Soria" authorityYear="1989" box="[388,508,1116,1139]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="order">Notopterna</taxonomicName>
(see
<taxonomicName id="4C107DA0FFF70D0EC539FB8D8BC9FBD2" authorityName="Püschel &amp; Shelley &amp; Williamson &amp; Perini &amp; Wible &amp; Brusatte" authorityYear="2024" box="[571,763,1115,1139]" family="Amilnedwardsidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" rank="family">Amilnedwardsidae</taxonomicName>
and the order
<taxonomicName id="4C107DA0FFF70D0EC606FBAD884EFB33" authorityName="Soria" authorityYear="1989" box="[260,380,1147,1170]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="order">Notopterna</taxonomicName>
section for more details). Key differences between notonychopids with indaleciids are the absence of enamel fossettes in the P2–M3 and a greater development of the parastyle in the former, among other features (Soria 1989b).
<bibRefCitation id="EF817BD2FFF70D0EC773FB2E88B9FAB1" author="McKenna and Bell" box="[113,395,1272,1296]" firstAuthor="McKenna" pageId="17" pageNumber="18" refId="ref44508" refString="McKenna MC, Bell SK. Classification of Mammals Above the Species Level. New York, NY: Columbia University Press, 1997." type="book" year="1997">McKenna and Bell (1997)</bibRefCitation>
later classified
<taxonomicName id="4C107DA0FFF70D0EC532FB2E8BEEFAB1" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[560,732,1272,1296]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName>
as a family within the order
<taxonomicName id="4C107DA0FFF70D0EC68AFAC188C5FA8E" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[392,503,1303,1327]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="order">Litopterna</taxonomicName>
, albeit without any anatomical justification (
<tableCitation id="C6923398FFF70D0EC653FAE188ACFAEE" box="[337,414,1335,1359]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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>
).
<bibRefCitation id="EF817BD2FFF70D0EC6B4FAE18BC8FAEE" author="Bonaparte and Morales" box="[438,762,1335,1359]" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation>
added to
<taxonomicName id="4C107DA0FFF70D0EC7CDFA808849FACF" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[207,379,1366,1390]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName>
the taxon
<taxonomicName id="4C107DA0FFF70D0EC6E4FA81881BFA2C" authority="Bonaparte and Morales 1997" authorityName="Bonaparte and Morales" authorityYear="1997" class="Mammalia" family="Notonychopidae" genus="Requisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="vidmari">
<emphasis id="B964DA31FFF70D0EC6E4FA818BB8FACF" box="[486,650,1366,1390]" italics="true" pageId="17" pageNumber="18">Requisia vidmari</emphasis>
<bibRefCitation id="EF817BD2FFF70D0EC592FA80881BFA2C" author="Bonaparte and Morales" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales 1997</bibRefCitation>
</taxonomicName>
from Punta Peligro (Peligran SALMA) which they found to be phylogenetically close to
<taxonomicName id="4C107DA0FFF70D0EC527FA438BF6FA0C" authorityName="Simpson" authorityYear="1935" box="[549,708,1429,1453]" class="Mammalia" family="Proterotheriidae" genus="Wainka" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="tshotshe">
<emphasis id="B964DA31FFF70D0EC527FA438BF6FA0C" box="[549,708,1429,1453]" italics="true" pageId="17" pageNumber="18">Wainka tshotshe</emphasis>
</taxonomicName>
(
<figureCitation id="132B1AA6FFF70D0EC5D6FA4389A3FA6D" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="17" pageNumber="18">Fig. 1D</figureCitation>
). However, since this analysis,
<bibRefCitation id="EF817BD2FFF70D0EC6D2FA628BBBFA6D" author="Goin" box="[464,649,1460,1484]" etAl="et al." firstAuthor="Goin" pageId="17" pageNumber="18" pagination="1029 - 53" refId="ref42920" refString="Goin FJ, Gelfo JN, Ortiz-Jaureguizar E, et al. Mamiferos del Banco Negro Inferior, Formacion Salamanca, Cuenca del Golfo de San Jorge. In: Giacosa RE (ed.), Geologia y Recursos Naturales de la Provincia de Chubut. Puerto Madryn, Relatorio del XXI Congreso Geologico Argentino, 2022, 1029 - 53." type="book chapter" year="2022">
Goin
<emphasis id="B964DA31FFF70D0EC508FA638B08FA6D" box="[522,570,1460,1484]" italics="true" pageId="17" pageNumber="18">et al.</emphasis>
(2022)
</bibRefCitation>
called into question an isolated m3 (UNP 946) previously assigned to
<taxonomicName id="4C107DA0FFF70D0EC773FA25882BF9AA" authorityName="Bonaparte and Morales" authorityYear="1997" box="[113,281,1523,1547]" class="Mammalia" family="Notonychopidae" genus="Requisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="vidmari">
<emphasis id="B964DA31FFF70D0EC773FA25882BF9AA" box="[113,281,1523,1547]" italics="true" pageId="17" pageNumber="18">Requisia vidmari</emphasis>
</taxonomicName>
by
<bibRefCitation id="EF817BD2FFF70D0EC643FA258BB4F9AA" author="Bonaparte and Morales" box="[321,646,1523,1547]" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation>
because of its bunodont features and how low and different its cristids are to what would be expected for a notonychopid, suggesting it could be a member of
<taxonomicName id="4C107DA0FFF70D0EC4CCFF198D46FF46" box="[974,1140,207,231]" class="Mammalia" family="Didolodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Didolodontidae</taxonomicName>
.
</paragraph>
<paragraph id="8BAF0623FFF70D0EC447FF388D79FDFE" blockId="17.[809,1461,144,795]" pageId="17" pageNumber="18">
<bibRefCitation id="EF817BD2FFF70D0EC447FF388DA0FEA7" author="Bonaparte and Morales" box="[837,1170,238,262]" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation>
failed to find a monophyletic
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, as
<taxonomicName id="4C107DA0FFF70D0EC339FED88D93FE84" authorityName="Soria" authorityYear="1989" box="[1083,1185,269,293]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="powelli">
<emphasis id="B964DA31FFF70D0EC339FED88D93FE84" box="[1083,1185,269,293]" italics="true" pageId="17" pageNumber="18">N. powelli</emphasis>
</taxonomicName>
and
<taxonomicName id="4C107DA0FFF70D0EC3DFFED88C78FE84" authorityName="Bonaparte and Morales" authorityYear="1997" box="[1245,1354,269,293]" class="Mammalia" family="Notonychopidae" genus="Requisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="vidmari">
<emphasis id="B964DA31FFF70D0EC3DFFED88C78FE84" box="[1245,1354,269,293]" italics="true" pageId="17" pageNumber="18">R. vidmari</emphasis>
</taxonomicName>
were not as closely related in their phylogeny (
<figureCitation id="132B1AA6FFF70D0EC3B7FEFB8C31FEE4" box="[1205,1283,301,325]" captionStart="Figure 1" captionStartId="3.[113,178,1691,1715]" captionTargetBox="[114,1459,150,1662]" captionTargetPageId="3" captionText="Figure 1. Phylogenies of the order Litopterna illustrating its interordinal relationships (A–B) and its interfamilial relationships (C–D). A, Buckley (2015) phylogeny.B, Westbury et al. (2017) phylogeny. C, Cifelli (1993) phylogeny. D, Bonaparte and Morales (1997) phylogeny. Different colours in the branches indicate different families or orders: blue Adianthidae; brown, Notopterna; green, Proterotheriidae; red, Macraucheniidae; yellow, Sparnotheriodontidae. Litopterna is indicated with a star, but other nodes or tips of relevance are indicated in circles: blue, orders; orange, suborders; pink, superfamilies; black, families; green, non-defined rank.Abbreviations:Ad, Adianthidae; An, Anisolambdidae; Di, Didolodontidae; I, Indaleciidae; L, Litopterna; Lo, Lopholipterna; Ma, Macraucheniidae; Mo, Macrauchenoidea; Nu, Notoungulata; Pa, Panperissodactyla; Pe, Proterotheriidae; Po, Protolipternidae; S, Sparnotheriodontidae." pageId="17" pageNumber="18">Fig. 1D</figureCitation>
). In addition, as notonychopids were found nested within
<taxonomicName id="4C107DA0FFF70D0EC3DCFE9A8C7FFEC5" baseAuthorityName="Garcia-Lopez and Babot" baseAuthorityYear="2014" box="[1246,1357,332,356]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="order">Litopterna</taxonomicName>
, most authors now consider them litopterns (e.g.,
<bibRefCitation id="EF817BD2FFF70D0EC3C9FEBD8C43FE22" author="Gelfo" box="[1227,1393,363,387]" etAl="et al." firstAuthor="Gelfo" pageId="17" pageNumber="18" pagination="285 - 91" refId="ref42550" refString="Gelfo JN, Lopez GM, Lorente M. Los ungulados arcaicos de America del Sur: ' Condylarthra' y Litopterna. Contribuciones Cientificas Del Museo Argentino de Ciencias Naturales ' Bernardino Rivadavia' 2016; 6: 285 - 91." type="journal article" year="2016">
Gelfo
<emphasis id="B964DA31FFF70D0EC208FEBA8C0BFE22" box="[1290,1337,363,387]" italics="true" pageId="17" pageNumber="18">et al.</emphasis>
2016
</bibRefCitation>
, Croft
<emphasis id="B964DA31FFF70D0EC428FE5D8A67FE02" box="[810,853,395,419]" italics="true" pageId="17" pageNumber="18">et al</emphasis>
. 2020,
<bibRefCitation id="EF817BD2FFF70D0EC4A7FE5D8D62FE02" author="Goin" box="[933,1104,395,419]" etAl="et al." firstAuthor="Goin" pageId="17" pageNumber="18" pagination="1029 - 53" refId="ref42920" refString="Goin FJ, Gelfo JN, Ortiz-Jaureguizar E, et al. Mamiferos del Banco Negro Inferior, Formacion Salamanca, Cuenca del Golfo de San Jorge. In: Giacosa RE (ed.), Geologia y Recursos Naturales de la Provincia de Chubut. Puerto Madryn, Relatorio del XXI Congreso Geologico Argentino, 2022, 1029 - 53." type="book chapter" year="2022">
Goin
<emphasis id="B964DA31FFF70D0EC4E1FE5D8D3CFE02" box="[995,1038,395,419]" italics="true" pageId="17" pageNumber="18">et al</emphasis>
. 2022
</bibRefCitation>
), which is problematic given that
<bibRefCitation id="EF817BD2FFF70D0EC428FE7C8D5CFE63" author="Bonaparte and Morales" box="[810,1134,426,450]" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation>
used a matrix that includes exclusively litopterns and did not test alternative relationships. Since the early phylogenetic study of
<bibRefCitation id="EF817BD2FFF70D0EC3BFFE3F8A46FD81" author="Bonaparte and Morales" firstAuthor="Bonaparte" pageId="17" pageNumber="18" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation>
, there has not been any attempt to test the monophyly of
<taxonomicName id="4C107DA0FFF70D0EC445FDF18AC1FD9E" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[839,1011,551,575]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName>
and to resolve its position among litopterns and other SANUs (
<tableCitation id="C6923398FFF70D0EC4F2FD918D09FDFE" box="[1008,1083,583,607]" captionStart="Table 2" captionStartId="4.[110,165,440,465]" 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>
).
</paragraph>
<paragraph id="8BAF0623FFF70D0EC447FDB08C1AFCBA" blockId="17.[809,1461,144,795]" pageId="17" pageNumber="18">
As the only two known members of
<taxonomicName id="4C107DA0FFF70D0EC3C1FDB08C5DFDDF" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[1219,1391,614,638]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="family">Notonychopidae</taxonomicName>
are
<taxonomicName id="4C107DA0FFF70D0EC29CFDB18A44FD3C" authorityName="Bonaparte and Morales" authorityYear="1997" class="Mammalia" family="Notonychopidae" genus="Requisia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="vidmari">
<emphasis id="B964DA31FFF70D0EC29CFDB18A44FD3C" italics="true" pageId="17" pageNumber="18">R. vidmari</emphasis>
</taxonomicName>
and the potentially pre-Itaboraian SALMA
<taxonomicName id="4C107DA0FFF70D0EC24BFD508C9FFD3C" authorityName="Soria" authorityYear="1989" box="[1353,1453,645,669]" class="Mammalia" family="Notonychopidae" genus="Notonychops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="powelli">
<emphasis id="B964DA31FFF70D0EC24BFD508C9FFD3C" box="[1353,1453,645,669]" italics="true" pageId="17" pageNumber="18">N. powelli</emphasis>
</taxonomicName>
, the temporal interval of this family is not well constrained with a current range of ~63.8–51.4 Mya, and is likely much narrower in range (
<figureCitation id="132B1AA6FFF70D0EC476FD358A8CFD5A" box="[884,958,739,763]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="17" pageNumber="18">Fig. 2B</figureCitation>
; Supporting information,
<tableCitation id="C6923398FFF70D0EC3CDFD358C19FD5A" box="[1231,1323,739,763]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" 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>
; Woodburne
<emphasis id="B964DA31FFF70D0EC428FCD58A6BFCBA" box="[810,857,771,795]" italics="true" pageId="17" pageNumber="18">et al.</emphasis>
2014b,
<bibRefCitation id="EF817BD2FFF70D0EC4AAFCD58D69FCBA" author="Krause" box="[936,1115,771,795]" etAl="et al." firstAuthor="Krause" pageId="17" pageNumber="18" pagination="886 - 903" refId="ref43716" refString="Krause JM, Clyde WC, Ibanez-Mejia M et al. New age constraints for Early Paleogene strata of central Patagonia, Argentina: implications for the timing of South American Land Mammal Ages. Geological Society of America Bulletin 2017; 129: 886 - 903. https: // doi. org / 10.1130 / b 31561.1" type="journal article" year="2017">
Krause
<emphasis id="B964DA31FFF70D0EC4F7FCD58D11FCBA" box="[1013,1059,771,795]" italics="true" pageId="17" pageNumber="18">et al.</emphasis>
2017
</bibRefCitation>
,
<bibRefCitation id="EF817BD2FFF70D0EC364FCD58C2AFCBA" author="Gelfo" box="[1126,1304,771,795]" etAl="et al." firstAuthor="Gelfo" pageId="17" pageNumber="18" pagination="993 - 1007" refId="ref42730" refString="Gelfo JN, Garcia-Lopez DA, Bergqvist LP. Phylogenetic relationships and palaeobiology of a new xenungulate (Mammalia: Eutheria) from the Palaeogene of Argentina. Journal of Systematic Palaeontology 2020 b; 18: 993 - 1007. https: // doi. org / 10.1080 / 14772019.2020.171 5496" type="journal article" year="2020" yearSuffix="b">
Gelfo
<emphasis id="B964DA31FFF70D0EC3A7FCD58DE6FCBA" box="[1189,1236,771,795]" italics="true" pageId="17" pageNumber="18">et al.</emphasis>
2020b
</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

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@ -1,177 +1,177 @@
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<mods:affiliation id="0D63135DBF0A29EAD9D1C143C2FEADB7">Natural History Museum, Cromwell Road, London SW 7 5 BD, UK.</mods:affiliation>
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@ -185,7 +185,7 @@ von Dalla Torre, 1880
</heading>
</paragraph>
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We associate a total of 186 published formal names with species of the subgenus
<taxonomicName id="6A894D5F1F5D9526FB45FE5FD90DFDF3" authorityName="von Dalla Torre" authorityYear="1880" box="[1218,1399,510,536]" class="Insecta" genus="Melanobombus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="31" pageNumber="32" phylum="Arthropoda" rank="genus">
@ -262,8 +262,6 @@ and by
</taxonomicName>
-group from our estimate of phylogeny in order to facilitate discussion:
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<taxonomicName id="6A894D5F1F5D9526FF3AFC40DD77FC10" ID-CoL="MFTW" authorityName="Lepeletier de Saint-Fargeau" authorityYear="1835" box="[189,269,993,1019]" class="Insecta" family="Apidae" genus="Bombus" kingdom="Animalia" order="Hymenoptera" pageId="31" pageNumber="32" phylum="Arthropoda" rank="species" species="rufipes">
<emphasis id="9FFDEACE1F5D9526FF3AFC40DD77FC10" box="[189,269,993,1019]" italics="true" pageId="31" pageNumber="32">rufipes</emphasis>
@ -597,7 +595,7 @@ S.-F.
syn. nov.
</paragraph>
</subSubSection>
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<paragraph id="AD3636DC1F629519FF3AFB8FDF23FBA2" blockId="32.[189,857,1069,1098]" box="[189,857,1069,1098]" pageId="32" pageNumber="33">
<emphasis id="9FFDEACE1F629519FF3AFB8FDF23FBA2" bold="true" box="[189,857,1069,1098]" pageId="32" pageNumber="33">
<emphasis id="9FFDEACE1F629519FF3AFB8FDE06FBA1" bold="true" box="[189,636,1069,1098]" italics="true" pageId="32" pageNumber="33">Key to the species of the subgenus</emphasis>
@ -625,7 +623,7 @@ Diagrams showing the major aspects of variation in the colour-patterns of the do
.
</paragraph>
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<emphasis id="9FFDEACE1F629519FF3AF90FDFF1F923" bold="true" box="[189,907,1709,1736]" pageId="32" pageNumber="33">
@ -636,8 +634,6 @@ Key to species for females of the subgenus
</emphasis>
</heading>
</paragraph>
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<paragraph id="AD3636DC1F629519FF3AF97CD90DF8F2" blockId="32.[189,1399,1756,1818]" pageId="32" pageNumber="33">
1. Hair of the thoracic dorsum entirely black ....................................................................................... 2 – Hair of the thoracic dorsum with some pale hair that is
<emphasis id="9FFDEACE1F629519FCF5F95EDFCCF8F2" box="[882,950,1791,1817]" italics="true" pageId="32" pageNumber="33">either</emphasis>
@ -1475,8 +1471,6 @@ eastwards ......................................................................
<bibRefCitation id="C9184B2D1F64951FFB3CFB3CD90DFB5C" author="Morawitz F. F." box="[1211,1399,1181,1207]" pageId="38" pageNumber="39" pagination="195 - 229" refId="ref68420" refString="Morawitz F. F. 1887. Insecta in itinere cl. N. Przewalskii in Asia centrali novissime lecta. I. Apidae. Trudy Russkago entomologicheskago obshchestva 20 (1886): 195 - 229." type="journal article" year="1887">Morawitz, 1887</bibRefCitation>
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<emphasis id="9FFDEACE1F64951FFF3AFB7EDDA9FB10" bold="true" box="[189,467,1247,1276]" italics="true" pageId="38" pageNumber="39">Review of the species</emphasis>
</paragraph>

239
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<collectingRegion id="7722D86D20E1D0FC7DA2726C19A26678" country="Taiwan" name="Kaohsiung">Kaohsiung</collectingRegion>
, southwestern
<collectingCountry id="6C23294E65B149216E6D6C91ABE18925" name="Taiwan">Taiwan</collectingCountry>
,
<location id="AF505EEC3807311D41B57DBCA9749243" LSID="urn:lsid:plazi:treatment:436BC03087585134AC4B88D34B1B346A:AF505EEC3807311D41B57DBCA9749243" country="Taiwan" county="Ho" latitude="22.708334" longLatPrecision="125" longitude="120.18" municipality="Sea" name="northern South" stateProvince="Kaohsiung">northern South</location>
<collectingCountry id="2A3C7864CA1ED85B68DF35DC69C77FE5" name="China">China</collectingCountry>
<collectingMunicipality id="3218962BEB5778E185B743B3F1AA8945">Sea</collectingMunicipality>
<collectingCounty id="81EDAF6896A66E739B1395CB29FAC599">northern South China Sea</collectingCounty>
,
<date id="D7BCBE022F9AAE93EF6551AE996CF4B8" value="2019-09-04">
<collectingDate id="09F02A1940B15813F28C30D7E1A57FBD" value="2019-09-04">4 Sep. 2019</collectingDate>
</date>
, collected by H. - C,
<collectingCounty id="F6977C07BBF9542B1D65C2E7313F640C">Ho</collectingCounty>
, collected by
<collectorName id="AACA4AFE76F88150482F44DC88E18B45">H.-C, Ho</collectorName>
</materialsCitation>
.
<materialsCitation id="26757992E85EEEB65290A5223DED7869" collectingDate="2020-07-25" collectingDateMax="2023-01-09" collectingDateMin="2020-07-25" collectorName="J. - F. Huang" country="Taiwan" county="southern East China Sea" latitude="24.941668" location="Daxi fish landing port" longLatPrecision="124" longitude="121.9" specimenCode="TOU-AE 7802, TOU-AE 8998, TOU-AE 8999, TOU-AE 9294" specimenCount="4" typeStatus="paratype">
<emphasis id="6C218510BF6F41EE3586815535F18B3D" bold="true" italics="true">
<typeStatus id="8A60FAC7DD22597BC66B0D3CDF0C24F9" type="paratype">Paratypes</typeStatus>
</emphasis>
:
<specimenCount id="BF757C4F2CF346B7FF6657FA8D47424C" count="4" type="generic">Four specimens</specimenCount>
, all collected from Daxi fish landing port,
<named-content content-type="dwc:verbatimCoordinates" id="NCID0EVVAG" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[121.900000,24.941667]}">
<geoCoordinate id="8FA4A056CA4C423FCD1B3B66CAFE6338" degrees="24" direction="north" minutes="56.5" orientation="latitude" precision="92" value="24.941668">24 ° 56.5 ' N</geoCoordinate>
, all collected from
<location id="C3EC0884E45A33A434A37C223856EC25" LSID="urn:lsid:plazi:treatment:436BC03087585134AC4B88D34B1B346A:C3EC0884E45A33A434A37C223856EC25" country="Taiwan" county="southern East China Sea" latitude="24.941668" longLatPrecision="124" longitude="121.9" name="Daxi fish landing port">Daxi fish landing port</location>
,
<geoCoordinate id="0D543AF1E10EDCF8B5EE6161671D12E2" degrees="121" direction="east" minutes="54.0" orientation="longitude" precision="92" value="121.9">121 ° 54.0 ' E</geoCoordinate>
<named-content content-type="dwc:verbatimCoordinates" id="NCID0EVVAG" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[121.900000,24.941667]}">
<geoCoordinate id="8FA4A056CA4C423FCD1B3B66CAFE6338" degrees="24" direction="north" minutes="56.5" orientation="latitude" precision="92" value="24.941668">24°56.5'N</geoCoordinate>
,
<geoCoordinate id="0D543AF1E10EDCF8B5EE6161671D12E2" degrees="121" direction="east" minutes="54.0" orientation="longitude" precision="92" value="121.9">121°54.0'E</geoCoordinate>
</named-content>
, northeastern
<collectingCountry id="370AB85561DE9806AB909E42639C8E48" name="Taiwan">Taiwan</collectingCountry>
, southern East
<collectingCountry id="E30E3E1EF609A45083037F88049A9618" name="China">China</collectingCountry>
Sea, collected by J. - F. Huang:
,
<collectingCounty id="251C93A744233229E906AF7E412BA5C6">southern East China Sea</collectingCounty>
, collected by
<collectorName id="19C47D5E8572BD55DDC5BB25A7ABDB27">J.-F. Huang</collectorName>
:
<specimenCode id="8F80A51E8350F658A45CBACA3CE82916">
<abbrev id="ABBRID0E1VAG" xlink_title="National Taiwan Ocean University, Laboratory of Aquatic Ecology, Department of Aquaculture">TOU-AE</abbrev>
<quantity id="394CFEC3B02893FF36F0410263529478" metricMagnitude="3" metricUnit="m" metricValue="7.802518999999999" unit="mm" value="7802519.0">7802, 519 mm</quantity>
7802
</specimenCode>
,
<quantity id="4515CBBCF6C00865A2ABEF037F69BABF" metricMagnitude="-1" metricUnit="m" metricValue="5.19" unit="mm" value="7802519.0">519 mm</quantity>
<abbrev id="ABBRID0E5VAG" xlink_title="Total lengths">TL</abbrev>
,
<date id="7AB7164A5313421D8CD210D2BB5526F1" value="2020-07-25">25 July 2020</date>
<date id="7AB7164A5313421D8CD210D2BB5526F1" value="2020-07-25">
<collectingDate id="FB4BD268CB091BADCC578E40AE394A83" value="2020-07-25">25 July 2020</collectingDate>
</date>
;
<specimenCode id="3548DC39F0DE760310687978F28323A4">
<abbrev id="ABBRID0ECWAG" xlink_title="National Taiwan Ocean University, Laboratory of Aquatic Ecology, Department of Aquaculture">TOU-AE</abbrev>
<quantity id="D2219F66A9CE857B618E67849A8B376F" metricMagnitude="3" metricUnit="m" metricValue="8.998554" unit="mm" value="8998554.0">8998, 554 mm</quantity>
8998
</specimenCode>
,
<quantity id="1EEFC44D71F5BC04F41384332FEEB677" metricMagnitude="-1" metricUnit="m" metricValue="5.54" unit="mm" value="8998554.0">554 mm</quantity>
<abbrev id="ABBRID0EGWAG" xlink_title="Total lengths">TL</abbrev>
,
<specimenCode id="CE213FEF59371F8C8ED06AB7D45C5E57">
<abbrev id="ABBRID0EKWAG" xlink_title="National Taiwan Ocean University, Laboratory of Aquatic Ecology, Department of Aquaculture">TOU-AE</abbrev>
<quantity id="C8714562F6186131486CAAB56ACB8EEC" metricMagnitude="3" metricUnit="m" metricValue="8.999597" unit="mm" value="8999597.0">8999, 597 mm</quantity>
8999
</specimenCode>
,
<quantity id="C82A9F775C10884B8A67C71420DE5F46" metricMagnitude="-1" metricUnit="m" metricValue="5.97" unit="mm" value="8999597.0">597 mm</quantity>
<abbrev id="ABBRID0EOWAG" xlink_title="Total lengths">TL</abbrev>
,
<date id="7AEFFDF87F43C782A5760D99381A8B29" value="2022-11-28">28 Nov. 2022</date>
<date id="7AEFFDF87F43C782A5760D99381A8B29" value="2022-11-28">
<collectingDate id="B65ED8EA999EFE89195AA34B03E02757" value="2022-11-28">28 Nov. 2022</collectingDate>
</date>
;
<specimenCode id="24CAE55379C625EFAB8EE0057D375DD1">
<abbrev id="ABBRID0ESWAG" xlink_title="National Taiwan Ocean University, Laboratory of Aquatic Ecology, Department of Aquaculture">TOU-AE</abbrev>
<quantity id="4BACC7C10CF280FC1D8E7157A37378AE" metricMagnitude="3" metricUnit="m" metricValue="9.294696" unit="mm" value="9294696.0">9294, 696 mm</quantity>
9294
</specimenCode>
,
<quantity id="C1FE1803DEBA6F00D79F6D931A9C3662" metricMagnitude="-1" metricUnit="m" metricValue="6.96" unit="mm" value="9294696.0">696 mm</quantity>
<abbrev id="ABBRID0EWWAG" xlink_title="Total lengths">TL</abbrev>
,
<date id="679DD9F35D6C1DDC41FCD8507AAEFBAF" value="2023-01-09">9 Jan. 2023</date>
<date id="679DD9F35D6C1DDC41FCD8507AAEFBAF" value="2023-01-09">
<collectingDate id="E3669F8090648CCD4870E1A8C840F17F" value="2023-01-09">9 Jan. 2023</collectingDate>
</date>
</materialsCitation>
.
</paragraph>
</subSubSection>
@ -202,11 +231,13 @@ Body elongate, slender (Fig.
<label id="65BA8DB9F2BF0C7DF932A36A0DC65C3D">Figure 3.</label>
</paragraph>
<paragraph id="DC2F9B058C25DDCFDF50FEDC7300EA1E">
<taxonomicName id="7F0DDDA8735B46C852EAD871A18E880E" authorityName="Hibino &amp; Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<taxonomicName id="7F0DDDA8735B46C852EAD871A18E880E" authorityName="Hibino, Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<emphasis id="427A6520C45571FCE64B8BB92F60B72F" italics="true">Ophichthus multidentis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="2E3074AE76BDB68B62B339E4E02BE9D8" rank="species">sp. nov.</taxonomicNameLabel>
, NMMB-P 36205, holotype, 433 mm
,
<specimenCode id="46B6515ED1F0C891D4E6BDECACE67C8A">NMMB-P 36205</specimenCode>
, holotype, 433 mm
<abbrev id="ABBRID0EKZAG" xlink_title="Total lengths">TL</abbrev>
, Ke-tzu-liao, Kaohsiung, Taiwan
<emphasis id="DCCBE7B099DA44146E4FFBF4A68A36B3" bold="true">A</emphasis>
@ -242,9 +273,7 @@ Sensory pores on head (Fig.
) developed but very small and not obvious;
<abbrev id="ABBRID0EH1AG" xlink_title="supraorbital pores">SO</abbrev>
1 + 3, first (ethmoid pore) on underside of snout tip and 3 along dorsal surface of snout, the last above upper margin of eye;
<abbrev id="ABBRID0EL1AG" xlink_title="infraorbital pores">
<collectionCode id="57820C15BCCF3C158302F97DC7619758" country="Portugal" name="Instituto de Oceanografia da Universidade de Lisboa">IO</collectionCode>
</abbrev>
<abbrev id="ABBRID0EL1AG" xlink_title="infraorbital pores">IO</abbrev>
3 + 3, 1 pore behind base of anterior nostril, 2 below eye, and 3 arranged in a vertical row behind eye;
<abbrev id="ABBRID0EP1AG" xlink_title="preoperculomandibular pores">POM</abbrev>
5 or 6 + 2 (
@ -265,11 +294,13 @@ before tail tip; canal on branchial basket weakly arched, 8 on branchial basket
</paragraph>
<paragraph id="F3ACA6F756057D50752D4A33BC32DB67">
Line drawings of
<taxonomicName id="4E3B4460D248AC40D2B2D3635E136EC1" authorityName="Hibino &amp; Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<taxonomicName id="4E3B4460D248AC40D2B2D3635E136EC1" authorityName="Hibino, Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<emphasis id="B4E672A0AE89A959FBD7E584B6A1A6BB" italics="true">O. multidentis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="825DE402A07C3122E92EB8CB5D20CFA2" rank="species">sp. nov.</taxonomicNameLabel>
, NMMB-P 36205, holotype, 433 mm
,
<specimenCode id="AE55409DFC5DB3AA63CA55F26CA00143">NMMB-P 36205</specimenCode>
, holotype, 433 mm
<abbrev id="ABBRID0EO2AG" xlink_title="Total lengths">TL</abbrev>
<emphasis id="5FDA19E6E4DB8CA6247E840E08146759" bold="true">A</emphasis>
sensory pores on head
@ -283,8 +314,11 @@ Teeth numerous, conical, pointed but shape and size variable (Fig.
); multiserial teeth on maxilla and dentary, outermost rows slightly larger and more robust than others, innermost teeth slender and more recurved posteriorly. Those on maxilla in 4 irregular rows in
<typeStatus id="5895E9425C1CA5E395E55D2AE276BE3C" type="holotype">holotype</typeStatus>
, but in larger specimens (
<specimenCode id="CBC865A182A0BA1363FED8D700086024">
<abbrev id="ABBRID0EB3AG" xlink_title="National Taiwan Ocean University, Laboratory of Aquatic Ecology, Department of Aquaculture">TOU-AE</abbrev>
8999 and 9294) 5 rows anteriorly and 4 rows posteriorly; on dentary arranged in 4 rows anteriorly and 3 rows posteriorly; on vomer moderate in size, becoming smaller posteriorly, 4 rows maximum and decreasing to single row posteriorly; 6 or 7 (
8999
</specimenCode>
and 9294) 5 rows anteriorly and 4 rows posteriorly; on dentary arranged in 4 rows anteriorly and 3 rows posteriorly; on vomer moderate in size, becoming smaller posteriorly, 4 rows maximum and decreasing to single row posteriorly; 6 or 7 (
<quantity id="1E1BA75B6391A443009DC3BB318BBFCC" metricMagnitude="-1" metricUnit="m" metricValue="1.524" unit="in" value="6.0">6 in</quantity>
<typeStatus id="D92803CA787CE0EC33F60D24F3F23332" type="holotype">holotype</typeStatus>
) large and robust, close-set teeth on intermaxillary, mostly concealed by lips.
@ -328,7 +362,7 @@ The specific name is derived from the Latin
</paragraph>
<paragraph id="E1F932FE1010E1847FA450F3FB1A3822">
The tooth pattern of
<taxonomicName id="A8C6F80F394EBE61E2E0193AC1A19FED" authorityName="Hibino &amp; Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<taxonomicName id="A8C6F80F394EBE61E2E0193AC1A19FED" authorityName="Hibino, Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<emphasis id="20962794E6F4729A494203E4CF0BD77B" italics="true">Ophichthus multidentis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B26E6483597F66EAD1CE0960D836B5A4" rank="species">sp. nov.</taxonomicNameLabel>
@ -339,7 +373,7 @@ is unique among the congeners. It is the only member of
that possesses up to 5 rows of small teeth on jaws and vomer in the northwestern Pacific region.
</paragraph>
<paragraph id="DF80C143131284994434CBAA7347ED6D">
<taxonomicName id="85183D607C225EB4CD90DC389A7B1B86" authorityName="Hibino &amp; Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<taxonomicName id="85183D607C225EB4CD90DC389A7B1B86" authorityName="Hibino, Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<emphasis id="99ACC49C01C5E5F2F350FA9EB2134AED" italics="true">Ophichthus multidentis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="8A1B26A25304DC44E09F6A7EA36B5CD1" rank="species">sp. nov.</taxonomicNameLabel>
@ -348,7 +382,7 @@ is similar to a number of species that have the dorsal-fin origin situated behin
: table 1;
<bibRefCitation id="F3E906E9DE9FFA8B3E99960181179EAF" author="Ho" etAl="et al." firstAuthor="Ho" journalOrPublisher="The Raffles Bulletin of Zoology" pagination="312-319" refId="B10" refString="Ho H-C, Ng S-L, Lin T-Y (2022) Description of a new Ophichthus eel from Dongsha Atoll, South China Sea, and a range extension of Ophichthus kusanagi Hibino, McCosker &amp; Tashiro, 2019. The Raffles Bulletin of Zoology 70: 312–319." title="Description of a new Ophichthus eel from Dongsha Atoll, South China Sea, and a range extension of Ophichthus kusanagi Hibino, McCosker &amp; Tashiro, 2019." volume="70" year="2022">Ho et al. 2022</bibRefCitation>
).
<taxonomicName id="ED8C28F6423AD28AD4F93E7D510BD214" authorityName="Hibino &amp; Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<taxonomicName id="ED8C28F6423AD28AD4F93E7D510BD214" authorityName="Hibino, Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<emphasis id="BFE68ED952800F8CABC00F8694F61B52" italics="true">Ophichthus multidentis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="39B077198F900C4548A7AE83614CA4E9" rank="species">sp. nov.</taxonomicNameLabel>
@ -373,7 +407,7 @@ vs 50.0–52.9 %
24-64 - 163 vs 27-68 - 159).
</paragraph>
<paragraph id="826B7A2F6CA1B2C5440CFE9429C2C01D">
<taxonomicName id="49B9021758936FC961A9CE0B24BC5DA3" authorityName="Hibino &amp; Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<taxonomicName id="49B9021758936FC961A9CE0B24BC5DA3" authorityName="Hibino, Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<emphasis id="325353643FE237FF72E30A59D3D77F3A" italics="true">Ophichthus multidentis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="54EA3B6300B64C555DA9EC8A85DBB7B9" rank="species">sp. nov.</taxonomicNameLabel>
@ -400,7 +434,7 @@ and
;
<bibRefCitation id="658C2939D5E9BFE6EBF0AE555433DD76" DOI="10.1007/s10228-018-00677-3" author="Hibino Y &amp; McCosker JE &amp; Tashiro F" issue="2" journalOrPublisher="Ichthyological Research" pagination="289-306" refId="B7" refString="Hibino Y, McCosker JE, Tashiro F (2019 b) Four new deepwater Ophichthus (Anguilliformes: Ophichthidae) from Japan with a redescription of Ophichthus pallens (Richardson 1848). Ichthyological Research 66 (2): 289–306. https://doi.org/10.1007/s10228-018-00677-3" title="Four new deepwater Ophichthus (Anguilliformes: Ophichthidae) from Japan with a redescription of Ophichthus pallens (Richardson 1848)." volume="66">Hibino et al. 2019 b</bibRefCitation>
; this study). We do not have the sufficient information of the molecular evidence of the new species with other congeners, while the partial sequence of
<taxonomicName id="7C5D8906BBAA5598A19C18BC11D0AC40" authorityName="Hibino &amp; Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidens" status="sp. nov.">
<taxonomicName id="7C5D8906BBAA5598A19C18BC11D0AC40" authorityName="Hibino, Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidens" status="sp. nov.">
<emphasis id="CA990262ED3A40A429844EF9F774A7EE" italics="true">O. multidens</emphasis>
</taxonomicName>
<taxonomicNameLabel id="7FD1B7B554C48E87FB3CBB2CED138631" rank="species">sp. nov.</taxonomicNameLabel>

133
data/68/64/87/6864879DFFC1A77FFC2FFD13FB45F848.xml
View file

@ -1,69 +1,72 @@
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(
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<figureCitation id="78F62A0EFFC1A77BFB98FD22FB48FD47" box="[1121,1178,692,718]" captionStart="Figure 4" captionStartId="11.[113,178,1236,1260]" captionTargetBox="[117,1456,146,1201]" captionTargetId="figure-302@11.[114,1458,144,1208]" captionTargetPageId="11" captionText="Figure 4. In situ pictures of Spongia lamella in the Northeast Atlantic Ocean and the Mediterranean Sea, A, Baloo tunnel, Ciboure, France.B, C, Mouro Island, Santander, Spain.D, Marseille, France." figureDoi="http://doi.org/10.5281/zenodo.14340973" httpUri="https://zenodo.org/record/14340973/files/figure.png" pageId="10" pageNumber="11">Fig. 4</figureCitation>
)
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@ -364,7 +367,7 @@ depth, Ciboure,
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.
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<emphasis id="D2B9EA99FFC0A77AFF88FB42FF1BFB65" bold="true" box="[113,201,1236,1260]" pageId="11" pageNumber="12">Figure 4.</emphasis>
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@ -599,26 +602,26 @@ Idan
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This sponge can be massive and irregular, with lobes or protuberances, fan- or vase-shaped when it becomes very big (
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). The largest studied specimens measured
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in their largest diameter, but this dimension can be&gt;
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in the Mediterranean Sea (
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). No true vase- or fan-shaped specimen was observed among our Atlantic specimens. This sponge presents a light to dark grey external colour, and the internal tissue is usually light or tawny yellow. The colour does not change upon fixation in 95% ethanol. The sponge consistency is flexible and hardly tearable. The surface is finely covered with small, evenly distributed conules. Oscules can be grouped in large individuals, or more randomly distributed in smaller ones, always on the outer sides in large individuals or on the top of each lobe or protuberance in smaller ones. They measure
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in diameter and are slightly raised, with a small whitish membrane delimiting the opening.
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<paragraph id="E072368BFFC7A77DFF65FDD1FDB8FC5F" blockId="12.[128,778,144,983]" pageId="12" pageNumber="13">
The ectosomal skeleton harbours an epidermal skin, 150– 600 µm thick, easily detachable, with a star-like appearance. It is made of a network of abundant foreign spicules and debris (
<figureCitation id="78F62A0EFFC7A77DFF72FD32FF25FD35" box="[139,247,676,700]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." pageId="12" pageNumber="13">Fig. 5A–C</figureCitation>
<figureCitation id="78F62A0EFFC7A77DFF72FD32FF25FD35" box="[139,247,676,700]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." figureDoi="http://doi.org/10.5281/zenodo.14340977" httpUri="https://zenodo.org/record/14340977/files/figure.png" pageId="12" pageNumber="13">Fig. 5A–C</figureCitation>
). Small canals can be observed under the surface. In the choanosome, debris and foreign spicules are still abundant at the periphery of canals, and thus throughout the sponge body (
<figureCitation id="78F62A0EFFC7A77DFF3CFC94FEC3FC93" box="[197,273,770,794]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." pageId="12" pageNumber="13">Fig. 5D</figureCitation>
<figureCitation id="78F62A0EFFC7A77DFF3CFC94FEC3FC93" box="[197,273,770,794]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." figureDoi="http://doi.org/10.5281/zenodo.14340977" httpUri="https://zenodo.org/record/14340977/files/figure.png" pageId="12" pageNumber="13">Fig. 5D</figureCitation>
). The primary fibres, 30–120 µm in diameter, are common, simple, irregular, and cored with numerous inclusions of foreign spicules and debris (
<figureCitation id="78F62A0EFFC7A77DFE36FCD7FDE9FCD0" box="[463,571,833,857]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." pageId="12" pageNumber="13">Fig. 5A, B</figureCitation>
<figureCitation id="78F62A0EFFC7A77DFE36FCD7FDE9FCD0" box="[463,571,833,857]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." figureDoi="http://doi.org/10.5281/zenodo.14340977" httpUri="https://zenodo.org/record/14340977/files/figure.png" pageId="12" pageNumber="13">Fig. 5A, B</figureCitation>
). Near the surface, they can be ramified. Secondary fibres, 20–40 µm in diameter, without inclusions or pith, form a dense ramified and irregular network (
<figureCitation id="78F62A0EFFC7A77DFF1CFC09FE96FC3E" box="[229,324,927,951]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." pageId="12" pageNumber="13">Fig. 5B, E</figureCitation>
<figureCitation id="78F62A0EFFC7A77DFF1CFC09FE96FC3E" box="[229,324,927,951]" captionStart="Figure 5" captionStartId="13.[113,178,1836,1860]" captionTargetBox="[158,1413,146,1802]" captionTargetId="figure-7@13.[155,1417,144,1808]" captionTargetPageId="13" captionText="Figure 5. Sections of the spongin skeleton of Spongia lamella. A, general view of the surface and the ectosome, showing the crust made of foreign spicules and debris at the surface, and the simple primary fibres.B, primary fibres including foreign spicules and debris, and secondary fibres forming an irregular and dense network. C, tangential section of the surface crust.D, foreign spicules and debris around canals. E, details of the secondary fibres." figureDoi="http://doi.org/10.5281/zenodo.14340977" httpUri="https://zenodo.org/record/14340977/files/figure.png" pageId="12" pageNumber="13">Fig. 5B, E</figureCitation>
). Tertiary fibres are present or absent. When they exist, their thickness is ≤10 µm in diameter.
</paragraph>
</subSubSection>
@ -733,7 +736,7 @@ they become anastomosed at this level. The size of the primary fibres appears a
</paragraph>
<paragraph id="E072368BFFC7A77DFCC0FC48FB51FB78" blockId="12.[825,1475,951,1985]" pageId="12" pageNumber="13">
GenBank accession numbers for all sequences generated or downloaded in this study are presented in
<tableCitation id="AD4F0330FFC7A77DFB09FC6BFAEEFB9C" box="[1264,1340,1021,1045]" captionStart="Table 1" captionStartId="3.[113,168,143,167]" captionTargetPageId="3" captionText="Table 1. Species names, voucher codes and GenBank accession numbers of Spongiidae used in this study." pageId="12" pageNumber="13">Table 1</tableCitation>
<tableCitation id="AD4F0330FFC7A77DFB09FC6BFAEEFB9C" box="[1264,1340,1021,1045]" captionStart="Table 1" captionStartId="3.[113,168,143,167]" captionTargetPageId="3" captionText="Table 1. Species names, voucher codes and GenBank accession numbers of Spongiidae used in this study." httpUri="http://table.plazi.org/id/B4B26603FFC8A772FF88FF19FB82FF2E" pageId="12" pageNumber="13" tableUuid="B4B26603FFC8A772FF88FF19FB82FF2E">Table 1</tableCitation>
. For the 28S (C2–D2 fragment) and
<emphasis id="D2B9EA99FFC7A77DFBCBFB8BFBB1FBBC" box="[1074,1123,1053,1077]" italics="true" pageId="12" pageNumber="13">CO1</emphasis>
(dgLCO1490–dgHCO2198 fragment) molecular markers, we obtained, respectively, 18 and 3 sequences of
@ -772,11 +775,11 @@ GenBank accession numbers for all sequences generated or downloaded in this stud
</paragraph>
<paragraph id="E072368BFFC7A77FFCACFB6EFE48FE68" blockId="12.[825,1475,951,1985]" lastBlockId="14.[128,778,144,1014]" lastPageId="14" lastPageNumber="15" pageId="12" pageNumber="13">
Molecular markers allowed us to build highly congruent trees, although the phylogenetic reconstruction with 28S gave a better resolution (
<figureCitation id="78F62A0EFFC7A77DFC48FAA1FC3BFAC7" box="[945,1001,1334,1359]" captionStart="Figure 6" captionStartId="15.[113,178,1836,1860]" captionTargetBox="[173,1400,146,1806]" captionTargetId="figure-7@15.[171,1402,144,1808]" captionTargetPageId="15" captionText="Figure 6. Phylogenetic reconstruction of Northeast Atlantic and Mediterranean Spongiidae using sequences of the C2–D2 region of 28S, and both the maximum-likelihood (ML) and neigbour-joining (NJ) methods. Bootstrap values are given for both analyses (1000 replicates). The NJ values are in parentheses. Each voucher code is associated with a species name. The ML bootstrap values are given only if&gt;500." pageId="12" pageNumber="13">Fig. 6</figureCitation>
<figureCitation id="78F62A0EFFC7A77DFC48FAA1FC3BFAC7" box="[945,1001,1334,1359]" captionStart="Figure 6" captionStartId="15.[113,178,1836,1860]" captionTargetBox="[173,1400,146,1806]" captionTargetId="figure-7@15.[171,1402,144,1808]" captionTargetPageId="15" captionText="Figure 6. Phylogenetic reconstruction of Northeast Atlantic and Mediterranean Spongiidae using sequences of the C2–D2 region of 28S, and both the maximum-likelihood (ML) and neigbour-joining (NJ) methods. Bootstrap values are given for both analyses (1000 replicates). The NJ values are in parentheses. Each voucher code is associated with a species name. The ML bootstrap values are given only if&gt;500." figureDoi="http://doi.org/10.5281/zenodo.14340979" httpUri="https://zenodo.org/record/14340979/files/figure.png" pageId="12" pageNumber="13">Fig. 6</figureCitation>
) than
<emphasis id="D2B9EA99FFC7A77DFBD5FAA1FB8FFAC6" box="[1068,1117,1335,1359]" italics="true" pageId="12" pageNumber="13">CO1</emphasis>
(Supporting Information,
<figureCitation id="78F62A0EFFC7A77DFA95FAA1FA63FAC6" box="[1388,1457,1335,1359]" captionStart="Figure 1" captionStartId="2.[129,194,1954,1978]" captionTargetBox="[131,1471,1210,1924]" captionTargetId="figure-610@2.[129,1473,1208,1926]" captionTargetPageId="2" captionText="Figure 1. Study area.Black stars indicate the sampling locations of Spongiidae from this work." pageId="12" pageNumber="13">Fig. S1</figureCitation>
<figureCitation id="78F62A0EFFC7A77DFA95FAA1FA63FAC6" box="[1388,1457,1335,1359]" captionStart="Figure 1" captionStartId="2.[129,194,1954,1978]" captionTargetBox="[131,1471,1210,1924]" captionTargetId="figure-610@2.[129,1473,1208,1926]" captionTargetPageId="2" captionText="Figure 1. Study area.Black stars indicate the sampling locations of Spongiidae from this work." figureDoi="http://doi.org/10.5281/zenodo.14340959" httpUri="https://zenodo.org/record/14340959/files/figure.png" pageId="12" pageNumber="13">Fig. S1</figureCitation>
). Both methods (NJ and ML) of phylogenetic reconstruction using 28S produced the same topology: 70
<taxonomicName id="27CD4D08FFC7A77DFAF2FAE3FAAFFA04" authorityName="Gray" authorityYear="1867" box="[1291,1405,1397,1421]" class="Demospongiae" family="Spongiidae" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="family">Spongiidae</taxonomicName>
specimens divided in two, with only one sequence of
@ -794,7 +797,7 @@ and
<emphasis id="D2B9EA99FFC7A77DFAE1FA23FA4DFA45" box="[1304,1439,1461,1484]" italics="true" pageId="12" pageNumber="13">Hippospongia</emphasis>
</taxonomicName>
on the other side (
<figureCitation id="78F62A0EFFC7A77DFC2CFA45FBDFFA62" box="[981,1037,1491,1515]" captionStart="Figure 6" captionStartId="15.[113,178,1836,1860]" captionTargetBox="[173,1400,146,1806]" captionTargetId="figure-7@15.[171,1402,144,1808]" captionTargetPageId="15" captionText="Figure 6. Phylogenetic reconstruction of Northeast Atlantic and Mediterranean Spongiidae using sequences of the C2–D2 region of 28S, and both the maximum-likelihood (ML) and neigbour-joining (NJ) methods. Bootstrap values are given for both analyses (1000 replicates). The NJ values are in parentheses. Each voucher code is associated with a species name. The ML bootstrap values are given only if&gt;500." pageId="12" pageNumber="13">Fig. 6</figureCitation>
<figureCitation id="78F62A0EFFC7A77DFC2CFA45FBDFFA62" box="[981,1037,1491,1515]" captionStart="Figure 6" captionStartId="15.[113,178,1836,1860]" captionTargetBox="[173,1400,146,1806]" captionTargetId="figure-7@15.[171,1402,144,1808]" captionTargetPageId="15" captionText="Figure 6. Phylogenetic reconstruction of Northeast Atlantic and Mediterranean Spongiidae using sequences of the C2–D2 region of 28S, and both the maximum-likelihood (ML) and neigbour-joining (NJ) methods. Bootstrap values are given for both analyses (1000 replicates). The NJ values are in parentheses. Each voucher code is associated with a species name. The ML bootstrap values are given only if&gt;500." figureDoi="http://doi.org/10.5281/zenodo.14340979" httpUri="https://zenodo.org/record/14340979/files/figure.png" pageId="12" pageNumber="13">Fig. 6</figureCitation>
). This set of specimens can be divided into three different groups. The first group is well supported by both methods of phylogenetic constructions (NJ and ML). It is composed by the proposed new species
<taxonomicName id="27CD4D08FFC7A77DFB46F9A4FA93F9C0" box="[1215,1345,1586,1609]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="12" pageNumber="13" phylum="Porifera" rank="species" species="maitasuna">
<emphasis id="D2B9EA99FFC7A77DFB46F9A4FA93F9C0" box="[1215,1345,1586,1609]" italics="true" pageId="12" pageNumber="13">S. maitasuna</emphasis>
@ -889,7 +892,7 @@ or
</taxonomicName>
.
</paragraph>
<caption id="B4B26603FFC6A77CFF88F8BAFE9FF811" pageId="13" pageNumber="14" startId="13.[113,178,1836,1860]" targetBox="[158,1413,146,1802]" targetPageId="13" targetType="figure">
<caption id="B4B26603FFC6A77CFF88F8BAFE9FF811" ID-DOI="http://doi.org/10.5281/zenodo.14340977" ID-Zenodo-Dep="14340977" httpUri="https://zenodo.org/record/14340977/files/figure.png" pageId="13" pageNumber="14" startId="13.[113,178,1836,1860]" targetBox="[158,1413,146,1802]" targetPageId="13" targetType="figure">
<paragraph id="E072368BFFC6A77CFF88F8BAFE9FF811" blockId="13.[113,1450,1836,1944]" pageId="13" pageNumber="14">
<emphasis id="D2B9EA99FFC6A77CFF88F8BAFF1BF8CD" bold="true" box="[113,201,1836,1860]" pageId="13" pageNumber="14">Figure 5.</emphasis>
Sections of the spongin skeleton of
@ -903,7 +906,7 @@ Sections of the spongin skeleton of
A phylogenetic tree was made by concatenating all sequences available for both 28S and
<emphasis id="D2B9EA99FFC5A77FFE61FD9EFE1AFDA9" box="[408,456,520,544]" italics="true" pageId="14" pageNumber="15">CO1</emphasis>
(
<figureCitation id="78F62A0EFFC5A77FFE23FD9EFDC6FD96" box="[474,532,520,544]" captionStart="Figure 7" captionStartId="16.[129,194,1742,1766]" captionTargetBox="[130,1471,146,1712]" captionTargetId="figure-70@16.[129,1473,144,1714]" captionTargetPageId="16" captionText="Figure 7. Phylogenetic reconstruction of Northeast Atlantic and Mediterranean Spongiidae concatenating the C2–D2 region of 28S and the Folmer fragment of CO1 using both Bayesian inference (BI) and maximum likelihood (ML) methods. Posterior probabilities and bootstraps are given for both analyses (1000 replicates). The ML values (as percentages) are in parentheses. *Sequences obtained only in 28S in this study." pageId="14" pageNumber="15">Fig. 7</figureCitation>
<figureCitation id="78F62A0EFFC5A77FFE23FD9EFDC6FD96" box="[474,532,520,544]" captionStart="Figure 7" captionStartId="16.[129,194,1742,1766]" captionTargetBox="[130,1471,146,1712]" captionTargetId="figure-70@16.[129,1473,144,1714]" captionTargetPageId="16" captionText="Figure 7. Phylogenetic reconstruction of Northeast Atlantic and Mediterranean Spongiidae concatenating the C2–D2 region of 28S and the Folmer fragment of CO1 using both Bayesian inference (BI) and maximum likelihood (ML) methods. Posterior probabilities and bootstraps are given for both analyses (1000 replicates). The ML values (as percentages) are in parentheses. *Sequences obtained only in 28S in this study." figureDoi="http://doi.org/10.5281/zenodo.14340983" httpUri="https://zenodo.org/record/14340983/files/figure.png" pageId="14" pageNumber="15">Fig. 7</figureCitation>
), with the exception of two samples of
<taxonomicName id="27CD4D08FFC5A77FFEDBFDBEFE5BFDB6" authorityName="Schmidt" authorityYear="1862" box="[290,393,552,575]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="zimocca">
<emphasis id="D2B9EA99FFC5A77FFEDBFDBEFE5BFDB6" box="[290,393,552,575]" italics="true" pageId="14" pageNumber="15">S. zimocca</emphasis>
@ -980,7 +983,7 @@ could be organized and grouped based on their chemical fingerprints and whether
</paragraph>
<paragraph id="E072368BFFC5A77FFF65FB6EFED7F8CD" blockId="14.[129,778,1045,1985]" pageId="14" pageNumber="15">
The dendrogram (
<figureCitation id="78F62A0EFFC5A77FFEA7FB6EFE48FA99" box="[350,410,1272,1296]" captionStart="Figure 8" captionStartId="17.[113,178,1836,1860]" captionTargetBox="[118,1452,144,1805]" captionTargetId="figure-7@17.[114,1458,144,1808]" captionTargetPageId="17" captionText="Figure 8. Metabolomic analysis of Northeast Atlantic and Mediterranean Spongiidae.Dendrogram from hierarchical clustering analysis using Euclidean distance calculations under the Ward algorithm. *Individuals for which no genetic data could be obtained." pageId="14" pageNumber="15">Fig. 8</figureCitation>
<figureCitation id="78F62A0EFFC5A77FFEA7FB6EFE48FA99" box="[350,410,1272,1296]" captionStart="Figure 8" captionStartId="17.[113,178,1836,1860]" captionTargetBox="[118,1452,144,1805]" captionTargetId="figure-7@17.[114,1458,144,1808]" captionTargetPageId="17" captionText="Figure 8. Metabolomic analysis of Northeast Atlantic and Mediterranean Spongiidae.Dendrogram from hierarchical clustering analysis using Euclidean distance calculations under the Ward algorithm. *Individuals for which no genetic data could be obtained." figureDoi="http://doi.org/10.5281/zenodo.14340987" httpUri="https://zenodo.org/record/14340987/files/figure.png" pageId="14" pageNumber="15">Fig. 8</figureCitation>
) shows distinct groups of extracts all attributed to three distinct groups of species. The first group (group 1) includes all specimens of the proposed new species,
<taxonomicName id="27CD4D08FFC5A77FFF78FAC0FED2FAE7" box="[129,256,1366,1390]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="maitasuna">
<emphasis id="D2B9EA99FFC5A77FFF78FAC0FED2FAE7" box="[129,256,1366,1390]" italics="true" pageId="14" pageNumber="15">S. maitasuna</emphasis>
@ -1023,7 +1026,7 @@ from
</paragraph>
<paragraph id="E072368BFFC5A77FFF65F8DAFB29FE8F" blockId="14.[129,778,1045,1985]" lastBlockId="14.[825,1475,144,1985]" pageId="14" pageNumber="15">
The heatmap (
<figureCitation id="78F62A0EFFC5A77FFEC9F8DDFEBAF8EA" box="[304,360,1867,1891]" captionStart="Figure 9" captionStartId="18.[129,194,1348,1372]" captionTargetBox="[134,1471,147,1317]" captionTargetId="figure-289@18.[129,1473,144,1321]" captionTargetPageId="18" captionText="Figure 9. Metabolomic analysis of Northeast Atlantic and Mediterranean Spongiidae.Heatmap from hierarchical clustering analysis using Euclidean distance calculations under the Ward algorithm, grouping sponge extracts based on their chemical similarities. Chemical features or signals are depicted as horizontal lines, with their colour indicating the normalized level of detection intensity on a scale from −2 to +2, from dark blue to red. The heatmap shows that each species is characterized by a unique set of chemical features (dark red), collectively defining their chemical signature." pageId="14" pageNumber="15">Fig. 9</figureCitation>
<figureCitation id="78F62A0EFFC5A77FFEC9F8DDFEBAF8EA" box="[304,360,1867,1891]" captionStart="Figure 9" captionStartId="18.[129,194,1348,1372]" captionTargetBox="[134,1471,147,1317]" captionTargetId="figure-289@18.[129,1473,144,1321]" captionTargetPageId="18" captionText="Figure 9. Metabolomic analysis of Northeast Atlantic and Mediterranean Spongiidae.Heatmap from hierarchical clustering analysis using Euclidean distance calculations under the Ward algorithm, grouping sponge extracts based on their chemical similarities. Chemical features or signals are depicted as horizontal lines, with their colour indicating the normalized level of detection intensity on a scale from −2 to +2, from dark blue to red. The heatmap shows that each species is characterized by a unique set of chemical features (dark red), collectively defining their chemical signature." figureDoi="http://doi.org/10.5281/zenodo.14340989" httpUri="https://zenodo.org/record/14340989/files/figure.png" pageId="14" pageNumber="15">Fig. 9</figureCitation>
) complements these results by depicting the whole set of chemical features that participated in the sample classification. This representation illustrates clearly that extracts from
<taxonomicName id="27CD4D08FFC5A77FFF44F83CFEEDF848" box="[189,319,1962,1985]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="14" pageNumber="15" phylum="Porifera" rank="species" species="maitasuna">
<emphasis id="D2B9EA99FFC5A77FFF44F83CFEEDF848" box="[189,319,1962,1985]" italics="true" pageId="14" pageNumber="15">S. maitasuna</emphasis>
@ -1034,7 +1037,7 @@ samples. Next, we assessed the distribution of features within the three identif
</paragraph>
<paragraph id="E072368BFFC5A77FFCACFE98FB03FE68" blockId="14.[825,1475,144,1985]" pageId="14" pageNumber="15">
To that end, a second aligned feature list was generated from pooled extracts from each identified taxonomic unit and contained 240 features. Those features represent the most abundant and redundant chemical signals in each taxonomic unit. Venn diagrams were constructed to depict the number of chemical features shared between species as opposed to those that were detected in one or two species (
<figureCitation id="78F62A0EFFC5A77FFB85FE5FFB13FE68" box="[1148,1217,457,481]" captionStart="Figure 10" captionStartId="19.[115,180,1389,1413]" captionTargetBox="[116,1453,146,1357]" captionTargetId="figure-314@19.[114,1458,144,1361]" captionTargetPageId="19" captionText="Figure 10. Venn diagrams representing the distribution of chemical features between species of the three main hierarchical groups of Spongiidae of the Mediterranean Sea and Northeast Atlantic Ocean. Black numbers indicate a unique set of features for each species. White numbers represent shared chemical signals between species. The number of features shared between all species of a given group are in white bold." pageId="14" pageNumber="15">Fig. 10</figureCitation>
<figureCitation id="78F62A0EFFC5A77FFB85FE5FFB13FE68" box="[1148,1217,457,481]" captionStart="Figure 10" captionStartId="19.[115,180,1389,1413]" captionTargetBox="[116,1453,146,1357]" captionTargetId="figure-314@19.[114,1458,144,1361]" captionTargetPageId="19" captionText="Figure 10. Venn diagrams representing the distribution of chemical features between species of the three main hierarchical groups of Spongiidae of the Mediterranean Sea and Northeast Atlantic Ocean. Black numbers indicate a unique set of features for each species. White numbers represent shared chemical signals between species. The number of features shared between all species of a given group are in white bold." figureDoi="http://doi.org/10.5281/zenodo.14340993" httpUri="https://zenodo.org/record/14340993/files/figure.png" pageId="14" pageNumber="15">Fig. 10</figureCitation>
).
</paragraph>
<paragraph id="E072368BFFC5A77FFCACFE7FFB81FCB3" blockId="14.[825,1475,144,1985]" pageId="14" pageNumber="15">

131
data/68/64/87/6864879DFFCDA77BFF4FFB8FFC59FDB6.xml
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@ -1,69 +1,72 @@
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@ -83,7 +86,7 @@ nov.
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(
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<figureCitation id="78F62A0EFFCDA777FE76FBCEFE29FBFB" box="[399,507,1112,1138]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." figureDoi="http://doi.org/10.5281/zenodo.14340963" httpUri="https://zenodo.org/record/14340963/files/figure.png" pageId="6" pageNumber="7">Fig. 2A–D</figureCitation>
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@ -578,7 +581,7 @@ Spongia officinalis
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(
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<figureCitation id="78F62A0EFFCDA777FBA4F97EFB4CF888" box="[1117,1182,1767,1793]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." figureDoi="http://doi.org/10.5281/zenodo.14340963" httpUri="https://zenodo.org/record/14340963/files/figure.png" pageId="6" pageNumber="7">Fig.2E</figureCitation>
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@ -626,7 +629,7 @@ depth, Marseille,
<date id="9473104BFFCDA777FB87F83CFAD5F84B" box="[1150,1287,1962,1986]" pageId="6" pageNumber="7" value="2020-05-19">19 May 2020</date>
.
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<emphasis id="D2B9EA99FFCCA776FF88F957FF1BF950" bold="true" box="[113,201,1729,1753]" pageId="7" pageNumber="8">Figure 2.</emphasis>
A–D,
@ -889,7 +892,7 @@ depth, Marseille,
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<emphasis id="D2B9EA99FFC3A779FE62FCFCFE3DFC0D" box="[411,495,874,900]" italics="true" pageId="8" pageNumber="9">
(
<figureCitation id="78F62A0EFFC3A779FE5CFCFCFE37FC0D" box="[421,485,874,900]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." pageId="8" pageNumber="9">Fig.2F</figureCitation>
<figureCitation id="78F62A0EFFC3A779FE5CFCFCFE37FC0D" box="[421,485,874,900]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." figureDoi="http://doi.org/10.5281/zenodo.14340963" httpUri="https://zenodo.org/record/14340963/files/figure.png" pageId="8" pageNumber="9">Fig.2F</figureCitation>
)
</emphasis>
</paragraph>
@ -1337,37 +1340,37 @@ language. Collected first in a site of the French
</paragraph>
<paragraph id="E072368BFFC3A779FCC0F9E0FB47F84E" blockId="8.[825,1474,1616,1991]" pageId="8" pageNumber="9">
The new sponge is massive and irregular, sometimes slightly branched, with small lobes or protuberances (
<figureCitation id="78F62A0EFFC3A779FAE1F903FA51F924" box="[1304,1411,1685,1709]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." pageId="8" pageNumber="9">Fig. 2A–D</figureCitation>
<figureCitation id="78F62A0EFFC3A779FAE1F903FA51F924" box="[1304,1411,1685,1709]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." figureDoi="http://doi.org/10.5281/zenodo.14340963" httpUri="https://zenodo.org/record/14340963/files/figure.png" pageId="8" pageNumber="9">Fig. 2A–D</figureCitation>
). The sponge can measure ≤
<quantity id="27359B6EFFC3A779FBDBF922FBB6F945" box="[1058,1124,1716,1740]" metricMagnitude="-1" metricUnit="m" metricValue="2.5" pageId="8" pageNumber="9" unit="cm" value="25.0">25 cm</quantity>
in its largest diameter. Its surface is white, beige, light to dark grey, whereas the internal tissue is usually tawny yellow. No change in colour was detected after fixation in 95% ethanol. The sponge surface is sometimes covered by dirty foreign bodies or epibiotic organisms. The surface is very conulose. Oscules can be grouped or not and are often found on the top of each lobe or protuberance. They are bordered by a thin and rather translucid membrane (
<figureCitation id="78F62A0EFFC3A779FB6FF806FAD2F821" box="[1174,1280,1936,1960]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." pageId="8" pageNumber="9">Fig. 2A–D</figureCitation>
<figureCitation id="78F62A0EFFC3A779FB6FF806FAD2F821" box="[1174,1280,1936,1960]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." figureDoi="http://doi.org/10.5281/zenodo.14340963" httpUri="https://zenodo.org/record/14340963/files/figure.png" pageId="8" pageNumber="9">Fig. 2A–D</figureCitation>
). The sponge consistency is rather soft and tearable.
</paragraph>
<paragraph id="E072368BFFC2A778FF74FF06FD8CFEAC" blockId="9.[113,763,144,450]" pageId="9" pageNumber="10">
The ectosomal skeleton presents a very thin epidermal skin, which has a star-like appearance, is easily detachable, and is made of collagen fibrils, from 60 to 350 µm in thickness (
<figureCitation id="78F62A0EFFC2A778FD2DFF59FF5DFE8F" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="9" pageNumber="10">Fig. 3A</figureCitation>
<figureCitation id="78F62A0EFFC2A778FD2DFF59FF5DFE8F" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." figureDoi="http://doi.org/10.5281/zenodo.14340967" httpUri="https://zenodo.org/record/14340967/files/figure.png" pageId="9" pageNumber="10">Fig. 3A</figureCitation>
) Scattered foreign spicules, debris, or remains of associated invertebrates, such as cirripeds, can be observed.
</paragraph>
<paragraph id="E072368BFFC2A77BFF74FEBBFED3FDA9" blockId="9.[113,763,144,450]" lastBlockId="10.[128,778,144,544]" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
The choanosomal skeleton presents primary fibres, which are irregular and sinuous, sometimes difficult to observe and to distinguish because they appear anastomosed with secondary fibres, and in some places, they can also divide and join again (
<figureCitation id="78F62A0EFFC2A778FF85FE3CFF36FE4B" box="[124,228,426,450]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="9" pageNumber="10">Fig. 3B–D</figureCitation>
<figureCitation id="78F62A0EFFC2A778FF85FE3CFF36FE4B" box="[124,228,426,450]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." figureDoi="http://doi.org/10.5281/zenodo.14340967" httpUri="https://zenodo.org/record/14340967/files/figure.png" pageId="9" pageNumber="10">Fig. 3B–D</figureCitation>
). They measure 30–70 µm in diameter and are cored with inclusions of foreign spicules and/or debris of various abundance. They also include a black or translucid pith. These primary fibres with a pith can be observed near the surface into conule formation and/or across the sponge body (
<figureCitation id="78F62A0EFFC2A778FAC1FF78FA71FE8F" box="[1336,1443,238,262]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="9" pageNumber="10">Fig. 3A, C</figureCitation>
<figureCitation id="78F62A0EFFC2A778FAC1FF78FA71FE8F" box="[1336,1443,238,262]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." figureDoi="http://doi.org/10.5281/zenodo.14340967" httpUri="https://zenodo.org/record/14340967/files/figure.png" pageId="9" pageNumber="10">Fig. 3A, C</figureCitation>
). However, when foreign debris and spicules are widely present, the pith in the primary fibres can be difficult to observe. Near the surface, close to the conule formations, the network of primary fibres is denser in comparison to the rest of the sponge body. The space between two primary fibres, thus between conules, seems to be empty, with very few tertiary fibres and foreign spicules and debris. Perforated plates are inconsistently present close to the primary fibres (
<figureCitation id="78F62A0EFFC1A77BFEB1FF39FE46FF4E" box="[328,404,175,199]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="10" pageNumber="11">Fig. 3D</figureCitation>
<figureCitation id="78F62A0EFFC1A77BFEB1FF39FE46FF4E" box="[328,404,175,199]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." figureDoi="http://doi.org/10.5281/zenodo.14340967" httpUri="https://zenodo.org/record/14340967/files/figure.png" pageId="10" pageNumber="11">Fig. 3D</figureCitation>
). The secondary fibres form a dense network, clearly showing a honeycomb shape of variable mesh size (
<figureCitation id="78F62A0EFFC1A77BFF4FFF78FF29FE8F" box="[182,251,238,262]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="10" pageNumber="11">Fig. 3B</figureCitation>
<figureCitation id="78F62A0EFFC1A77BFF4FFF78FF29FE8F" box="[182,251,238,262]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." figureDoi="http://doi.org/10.5281/zenodo.14340967" httpUri="https://zenodo.org/record/14340967/files/figure.png" pageId="10" pageNumber="11">Fig. 3B</figureCitation>
), from 70 µm × 120 µm to 300 µm × 500 µm. These fibres measure 20–50 µm in diameter and they can contain an irregular pith, black or translucid (
<figureCitation id="78F62A0EFFC1A77BFDFBFEBBFD9DFECC" box="[514,591,301,325]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="10" pageNumber="11">Fig. 3E</figureCitation>
<figureCitation id="78F62A0EFFC1A77BFDFBFEBBFD9DFECC" box="[514,591,301,325]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." figureDoi="http://doi.org/10.5281/zenodo.14340967" httpUri="https://zenodo.org/record/14340967/files/figure.png" pageId="10" pageNumber="11">Fig. 3E</figureCitation>
). Tertiary fibres, ~10 µm in diameter, are found throughout the sponge body (
<figureCitation id="78F62A0EFFC1A77BFF72FEFDFF23FE0A" box="[139,241,363,387]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="10" pageNumber="11">Fig. 3F, G</figureCitation>
<figureCitation id="78F62A0EFFC1A77BFF72FEFDFF23FE0A" box="[139,241,363,387]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." figureDoi="http://doi.org/10.5281/zenodo.14340967" httpUri="https://zenodo.org/record/14340967/files/figure.png" pageId="10" pageNumber="11">Fig. 3F, G</figureCitation>
). Sometimes they form a thinner network that also has a clear honeycomb shape with variable mesh size (50–130 µm in diameter). In some places, these fibres appear to be linked to the thicker secondary fibres (
<figureCitation id="78F62A0EFFC1A77BFE21FE5FFDFAFE68" box="[472,552,457,481]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." pageId="10" pageNumber="11">Fig. 3H</figureCitation>
<figureCitation id="78F62A0EFFC1A77BFE21FE5FFDFAFE68" box="[472,552,457,481]" captionStart="Figure 3" captionStartId="9.[113,178,1870,1894]" captionTargetBox="[119,1456,530,1838]" captionTargetId="figure-214@9.[114,1458,526,1842]" captionTargetPageId="9" captionText="Figure 3. Sections of the spongin skeleton of Spongia maitasuna. A, general view of the surface and the skeleton, showing the epiderm made of collagen fibrils and the primary fibres with a pith (arrows).B, secondary fibres organized in a regular honeycomb network.C, in the primary fibres, the pith can be black (black arrow) or translucent (white arrow). D, perforated plates close to the primary fibres.E, secondary fibres with a pith (arrows). F, spicules and debris in the ectoderm.G, tertiary fibres.H, detail of a junction between secondary and tertiary fibres (arrow)." figureDoi="http://doi.org/10.5281/zenodo.14340967" httpUri="https://zenodo.org/record/14340967/files/figure.png" pageId="10" pageNumber="11">Fig. 3H</figureCitation>
), but in some places they look rather disconnected from the rest of the skeletal architecture.
</paragraph>
</subSubSection>
<caption id="B4B26603FFC2A778FF88F8D8FA71F832" pageId="9" pageNumber="10" startId="9.[113,178,1870,1894]" targetBox="[119,1456,530,1838]" targetPageId="9" targetType="figure">
<caption id="B4B26603FFC2A778FF88F8D8FA71F832" ID-DOI="http://doi.org/10.5281/zenodo.14340967" ID-Zenodo-Dep="14340967" httpUri="https://zenodo.org/record/14340967/files/figure.png" pageId="9" pageNumber="10" startId="9.[113,178,1870,1894]" targetBox="[119,1456,530,1838]" targetPageId="9" targetType="figure">
<paragraph id="E072368BFFC2A778FF88F8D8FA71F832" blockId="9.[113,1458,1870,1979]" pageId="9" pageNumber="10">
<emphasis id="D2B9EA99FFC2A778FF88F8D8FF1BF8EE" bold="true" box="[113,201,1870,1895]" pageId="9" pageNumber="10">Figure 3.</emphasis>
Sections of the spongin skeleton of
@ -1479,7 +1482,7 @@ from
, Strait of
<collectingCountry id="98DA761BFFC1A77BFE89F958FE00F96F" box="[368,466,1742,1766]" name="Gibraltar" pageId="10" pageNumber="11">Gibraltar</collectingCountry>
. The main difference in this case lies in the primary fibres, which are slightly larger than in the new species (70–110 µm), and in the overall growth form (
<figureCitation id="78F62A0EFFC1A77BFF75F8BAFF11F8CD" box="[140,195,1836,1860]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." pageId="10" pageNumber="11">Fig.2</figureCitation>
<figureCitation id="78F62A0EFFC1A77BFF75F8BAFF11F8CD" box="[140,195,1836,1860]" captionStart="Figure 2" captionStartId="7.[113,178,1729,1753]" captionTargetBox="[116,1454,146,1697]" captionTargetId="figure-108@7.[114,1458,144,1701]" captionTargetPageId="7" captionText="Figure 2. A–D, in situ pictures of Spongia maitasuna in Baloo tunnel, Ciboure, France (A, B) and Mouro Island, Santander, Spain (C, D). E, Spongia officinalis from Marseille, France.F, Spongia cf.officinalis from Ceuta, Strait of Gibraltar, Spain." figureDoi="http://doi.org/10.5281/zenodo.14340963" httpUri="https://zenodo.org/record/14340963/files/figure.png" pageId="10" pageNumber="11">Fig.2</figureCitation>
).
<taxonomicName id="27CD4D08FFC1A77BFF2EF8BAFE46F8CA" ID-CoL="BW7B2" box="[215,404,1836,1859]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="10" pageNumber="11" phylum="Porifera" rank="species" species="maitasuna">
<emphasis id="D2B9EA99FFC1A77BFF2EF8BAFE46F8CA" box="[215,404,1836,1859]" italics="true" pageId="10" pageNumber="11">Spongia maitasuna</emphasis>

157
data/6A/3D/D1/6A3DD143066DFF905590FDB365926476.xml
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@ -1,87 +1,90 @@
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centra, the core was filled with dense cancellous bone (identified in CT images so of uncertain tissue composition) and the cortex was relatively thin (
<figureCitation id="7AAF7CD0066DFF9055F3FCD2617A6516" box="[241,332,841,865]" captionStart-0="Figure 2" captionStart-1="Figure 3" captionStart-2="Figure 4" captionStartId-0="3.[113,178,975,999]" captionStartId-1="6.[129,194,1669,1693]" captionStartId-2="7.[113,178,1841,1865]" captionTargetBox-0="[307,1266,157,946]" captionTargetBox-1="[305,1297,144,1642]" captionTargetBox-2="[175,1398,149,1813]" captionTargetId-0="figure-579@3.[305,1268,144,948]" captionTargetId-1="figure-103@6.[305,1297,144,1642]" captionTargetId-2="figure-6@7.[175,1398,149,1813]" captionTargetPageId-0="3" captionTargetPageId-1="6" captionTargetPageId-2="7" captionText-0="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." captionText-1="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." captionText-2="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Figs 2–4</figureCitation>
<figureCitation id="7AAF7CD0066DFF9055F3FCD2617A6516" box="[241,332,841,865]" captionStart-0="Figure 2" captionStart-1="Figure 3" captionStart-2="Figure 4" captionStartId-0="3.[113,178,975,999]" captionStartId-1="6.[129,194,1669,1693]" captionStartId-2="7.[113,178,1841,1865]" captionTargetBox-0="[307,1266,157,946]" captionTargetBox-1="[305,1297,144,1642]" captionTargetBox-2="[175,1398,149,1813]" captionTargetId-0="figure-579@3.[305,1268,144,948]" captionTargetId-1="figure-103@6.[305,1297,144,1642]" captionTargetId-2="figure-6@7.[175,1398,149,1813]" captionTargetPageId-0="3" captionTargetPageId-1="6" captionTargetPageId-2="7" captionText-0="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." captionText-1="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." captionText-2="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." figureDoi-0="http://doi.org/10.5281/zenodo.14340880" figureDoi-1="http://doi.org/10.5281/zenodo.14340884" figureDoi-2="http://doi.org/10.5281/zenodo.14340888" httpUri-0="https://zenodo.org/record/14340880/files/figure.png" httpUri-1="https://zenodo.org/record/14340884/files/figure.png" httpUri-2="https://zenodo.org/record/14340888/files/figure.png" pageId="5" pageNumber="6">Figs 2–4</figureCitation>
), whereas in the middle portion, where the internal cavity was revealed, the cortical bone was distinctly thicker. The transverse sections revealed the changes of the cortical bone thickness and the shape of internal cavities, with the mid-centrum being roughly cylindrical. The cortex was thinner near the anteroposteriorly terminal portions of the vertebrae and its thickness gradually increased up to approximately four times the original thickness (about 10% of bone diameter), near the midpoint (
<figureCitation id="7AAF7CD0066DFF90540AFBDF6176622B" box="[264,320,1092,1116]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">Fig. 3</figureCitation>
<figureCitation id="7AAF7CD0066DFF90540AFBDF6176622B" box="[264,320,1092,1116]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." figureDoi="http://doi.org/10.5281/zenodo.14340884" httpUri="https://zenodo.org/record/14340884/files/figure.png" pageId="5" pageNumber="6">Fig. 3</figureCitation>
). The density of the cancellous bone of the anteroposteriorly terminal portions of the centrum decreased medially, with larger cavities being present closer to the midpoint. The transition to the empty internal cavity was relatively sharp. The anterior and posterior ends of the internal cavity were roughly hemispherical and surrounded circumferentially by marginalized trabeculae (
<figureCitation id="7AAF7CD0066DFF905478FA9B6183636F" box="[378,437,1280,1304]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">Fig. 3</figureCitation>
<figureCitation id="7AAF7CD0066DFF905478FA9B6183636F" box="[378,437,1280,1304]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." figureDoi="http://doi.org/10.5281/zenodo.14340884" httpUri="https://zenodo.org/record/14340884/files/figure.png" pageId="5" pageNumber="6">Fig. 3</figureCitation>
). Long intermittent trabeculae composed of parallel-fibred bone moderately remodelled with endosteal secondary lamellar bone (
<figureCitation id="7AAF7CD0066DFF9054DFFAA462106320" box="[477,550,1343,1367]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4D</figureCitation>
<figureCitation id="7AAF7CD0066DFF9054DFFAA462106320" box="[477,550,1343,1367]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." figureDoi="http://doi.org/10.5281/zenodo.14340888" httpUri="https://zenodo.org/record/14340888/files/figure.png" pageId="5" pageNumber="6">Fig. 4D</figureCitation>
) crossed the internal cavity in varied directions, mainly diagonally. In some vertebrae they divided the internal cavity into several smaller terminus-like pockets (
<figureCitation id="7AAF7CD0066DFF9055D1FA06611763C2" box="[211,289,1437,1461]" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 2B</figureCitation>
<figureCitation id="7AAF7CD0066DFF9055D1FA06611763C2" box="[211,289,1437,1461]" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." figureDoi="http://doi.org/10.5281/zenodo.14340880" httpUri="https://zenodo.org/record/14340880/files/figure.png" pageId="5" pageNumber="6">Figs 2B</figureCitation>
,
<figureCitation id="7AAF7CD0066DFF90542DFA06610A63C2" box="[303,316,1437,1461]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">3</figureCitation>
<figureCitation id="7AAF7CD0066DFF90542DFA06610A63C2" box="[303,316,1437,1461]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." figureDoi="http://doi.org/10.5281/zenodo.14340884" httpUri="https://zenodo.org/record/14340884/files/figure.png" pageId="5" pageNumber="6">3</figureCitation>
). The neural canal was separated ventrally from the internal cavity by a horizontal, plate-like lamella consisting of endosteal lamellar bone (
<figureCitation id="7AAF7CD0066DFF9054D0FA40622A6384" box="[466,540,1499,1523]" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 2B</figureCitation>
<figureCitation id="7AAF7CD0066DFF9054D0FA40622A6384" box="[466,540,1499,1523]" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." figureDoi="http://doi.org/10.5281/zenodo.14340880" httpUri="https://zenodo.org/record/14340880/files/figure.png" pageId="5" pageNumber="6">Figs 2B</figureCitation>
,
<figureCitation id="7AAF7CD0066DFF905725FA4062026384" box="[551,564,1499,1523]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">3</figureCitation>
<figureCitation id="7AAF7CD0066DFF905725FA4062026384" box="[551,564,1499,1523]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." figureDoi="http://doi.org/10.5281/zenodo.14340884" httpUri="https://zenodo.org/record/14340884/files/figure.png" pageId="5" pageNumber="6">3</figureCitation>
,
<figureCitation id="7AAF7CD0066DFF90573CFA40624A6384" box="[574,636,1499,1523]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">4A–C</figureCitation>
<figureCitation id="7AAF7CD0066DFF90573CFA40624A6384" box="[574,636,1499,1523]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." figureDoi="http://doi.org/10.5281/zenodo.14340888" httpUri="https://zenodo.org/record/14340888/files/figure.png" pageId="5" pageNumber="6">4A–C</figureCitation>
), which was generally thinner than the internal trabeculae composed of the same tissue. This separation was not continuous, the neural canal floor was perforated by large openings in some specimens (
<figureCitation id="7AAF7CD0066DFF9057D2F9A260B86006" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 2B</figureCitation>
<figureCitation id="7AAF7CD0066DFF9057D2F9A260B86006" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." figureDoi="http://doi.org/10.5281/zenodo.14340880" httpUri="https://zenodo.org/record/14340880/files/figure.png" pageId="5" pageNumber="6">Figs 2B</figureCitation>
and
<figureCitation id="7AAF7CD0066DFF9055C1F9C261326006" box="[195,260,1625,1649]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">3C–G</figureCitation>
<figureCitation id="7AAF7CD0066DFF9055C1F9C261326006" box="[195,260,1625,1649]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." figureDoi="http://doi.org/10.5281/zenodo.14340884" httpUri="https://zenodo.org/record/14340884/files/figure.png" pageId="5" pageNumber="6">3C–G</figureCitation>
) and seemed to disappear completely in others. For example, in MGUWr 3889s at least two large openings were present in the lamella separating the neural canal from the internal cavity. In MGUWr 3902 and GPIH 5194c there appeared to be no bony separation between the neural canal and the internal cavity for most of the length of the centrum. Because no fragments of broken bone were present anywhere inside of the neural canal or the internal cavity of the centrum, and this condition was present in several specimens approximately in the same area, it appears not to be an artifact of preservation. The cortex was predominately composed of parallel-fibred matrix (
<figureCitation id="7AAF7CD0066DFF9057B2F80962DF61DD" box="[688,745,1938,1962]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4</figureCitation>
<figureCitation id="7AAF7CD0066DFF9057B2F80962DF61DD" box="[688,745,1938,1962]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." figureDoi="http://doi.org/10.5281/zenodo.14340888" httpUri="https://zenodo.org/record/14340888/files/figure.png" pageId="5" pageNumber="6">Fig. 4</figureCitation>
), with locally (especially in the dorsolateral part) highly organized arrays of mineralized collagen fibres. No rest lines were present. The vascularization was moderate, and its pattern was radial, which was especially evident in the ventral part of the vertebrae (
<figureCitation id="7AAF7CD0066DFF905637FE9663E56752" box="[821,979,269,293]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4A–C, H–I</figureCitation>
<figureCitation id="7AAF7CD0066DFF905637FE9663E56752" box="[821,979,269,293]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." figureDoi="http://doi.org/10.5281/zenodo.14340888" httpUri="https://zenodo.org/record/14340888/files/figure.png" pageId="5" pageNumber="6">Fig. 4A–C, H–I</figureCitation>
). Inside the neural arch, mostly dorsally to the neural canal (inside of the base of the neural spine) and in some specimens laterally to the neural canal there was a region of occurrence of secondary trabecular bone (
<figureCitation id="7AAF7CD0066DFF9051C3FEF0653767F4" box="[1217,1281,363,387]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4J</figureCitation>
<figureCitation id="7AAF7CD0066DFF9051C3FEF0653767F4" box="[1217,1281,363,387]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." figureDoi="http://doi.org/10.5281/zenodo.14340888" httpUri="https://zenodo.org/record/14340888/files/figure.png" pageId="5" pageNumber="6">Fig. 4J</figureCitation>
). Both the cavity and the neural canal were clear cut and lined with a thin layer of endosteal lamellar bone (
<figureCitation id="7AAF7CD0066DFF905137FE3164A067B5" box="[1077,1174,426,450]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4E–J</figureCitation>
<figureCitation id="7AAF7CD0066DFF905137FE3164A067B5" box="[1077,1174,426,450]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." figureDoi="http://doi.org/10.5281/zenodo.14340888" httpUri="https://zenodo.org/record/14340888/files/figure.png" pageId="5" pageNumber="6">Fig. 4E–J</figureCitation>
). Sharpey’s fibres could be seen extending throughout the cortex, especially in the dorsolateral and ventrolateral regions of the centrum (
<figureCitation id="7AAF7CD0066DFF905017FE7265A56476" box="[1301,1427,489,513]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4C, G–I</figureCitation>
<figureCitation id="7AAF7CD0066DFF905017FE7265A56476" box="[1301,1427,489,513]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." figureDoi="http://doi.org/10.5281/zenodo.14340888" httpUri="https://zenodo.org/record/14340888/files/figure.png" pageId="5" pageNumber="6">Fig. 4C, G–I</figureCitation>
).
</paragraph>
</subSubSection>

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@ -101,25 +104,25 @@ spp.
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Within all of the studied vertebrae, a large cavity could be identified, surrounded by relatively thick cortex composed of alternating layers of well-organized parallel-fibred to lamellar bone (
<figureCitation id="7AAF7CD0066DFF905637FD7563BD6571" box="[821,907,750,774]" captionStart-0="Figure 5" captionStart-1="Figure 6" captionStart-2="Figure 7" captionStart-3="Figure 8" captionStartId-0="8.[130,195,1138,1162]" captionStartId-1="9.[113,178,1191,1215]" captionStartId-2="10.[129,194,695,719]" captionStartId-3="11.[113,178,1841,1865]" captionTargetBox-0="[289,1313,144,1110]" captionTargetBox-1="[146,1426,144,1163]" captionTargetBox-2="[289,1313,144,667]" captionTargetBox-3="[224,1348,150,1812]" captionTargetId-0="figure-299@8.[289,1313,144,1110]" captionTargetId-1="figure-396@9.[146,1426,144,1163]" captionTargetId-2="figure-733@10.[289,1313,144,667]" captionTargetId-3="figure-6@11.[223,1349,149,1813]" captionTargetPageId-0="8" captionTargetPageId-1="9" captionTargetPageId-2="10" captionTargetPageId-3="11" captionText-0="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." captionText-1="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." captionText-2="Figure 7. Internal cavity volume in the cervical vertebrae of ‘P’.antiquus MGUWr 3895s (A) and Tanystropheus sp. from Miedary ZPAL V. 36/181 (B). Anatomical abbreviation:ic—internal cavity. Images obtained using ‘xray’ shader in MeshLab v.2020.12 (Cignoni et al.2008)." captionText-3="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Figs 5–8</figureCitation>
<figureCitation id="7AAF7CD0066DFF905637FD7563BD6571" box="[821,907,750,774]" captionStart-0="Figure 5" captionStart-1="Figure 6" captionStart-2="Figure 7" captionStart-3="Figure 8" captionStartId-0="8.[130,195,1138,1162]" captionStartId-1="9.[113,178,1191,1215]" captionStartId-2="10.[129,194,695,719]" captionStartId-3="11.[113,178,1841,1865]" captionTargetBox-0="[289,1313,144,1110]" captionTargetBox-1="[146,1426,144,1163]" captionTargetBox-2="[289,1313,144,667]" captionTargetBox-3="[224,1348,150,1812]" captionTargetId-0="figure-299@8.[289,1313,144,1110]" captionTargetId-1="figure-396@9.[146,1426,144,1163]" captionTargetId-2="figure-733@10.[289,1313,144,667]" captionTargetId-3="figure-6@11.[223,1349,149,1813]" captionTargetPageId-0="8" captionTargetPageId-1="9" captionTargetPageId-2="10" captionTargetPageId-3="11" captionText-0="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." captionText-1="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." captionText-2="Figure 7. Internal cavity volume in the cervical vertebrae of ‘P’.antiquus MGUWr 3895s (A) and Tanystropheus sp. from Miedary ZPAL V. 36/181 (B). Anatomical abbreviation:ic—internal cavity. Images obtained using ‘xray’ shader in MeshLab v.2020.12 (Cignoni et al.2008)." captionText-3="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi-0="http://doi.org/10.5281/zenodo.14340890" figureDoi-1="http://doi.org/10.5281/zenodo.14340894" figureDoi-2="http://doi.org/10.5281/zenodo.14340898" figureDoi-3="http://doi.org/10.5281/zenodo.14340900" httpUri-0="https://zenodo.org/record/14340890/files/figure.png" httpUri-1="https://zenodo.org/record/14340894/files/figure.png" httpUri-2="https://zenodo.org/record/14340898/files/figure.png" httpUri-3="https://zenodo.org/record/14340900/files/figure.png" pageId="5" pageNumber="6">Figs 5–8</figureCitation>
). In the anterior and posterior terminal portions of the cervicals, discs of secondary cancellous bone of endosteal origin formed the articular regions of the centrum (
<figureCitation id="7AAF7CD0066DFF9051F1FCB6656D6533" box="[1267,1371,812,837]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Fig. 5B–D</figureCitation>
<figureCitation id="7AAF7CD0066DFF9051F1FCB6656D6533" box="[1267,1371,812,837]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." figureDoi="http://doi.org/10.5281/zenodo.14340890" httpUri="https://zenodo.org/record/14340890/files/figure.png" pageId="5" pageNumber="6">Fig. 5B–D</figureCitation>
). Above them, the openings of the neural canal were present. In some of the middle cervicals the neural canal floor extended only slightly from the openings towards the middle of the vertebra, forming a shelf that partially separated the internal cavity from the neural canal (
<figureCitation id="7AAF7CD0066DFF905673FC5263ED6596" box="[881,987,969,993]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 5B–E</figureCitation>
<figureCitation id="7AAF7CD0066DFF905673FC5263ED6596" box="[881,987,969,993]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." figureDoi="http://doi.org/10.5281/zenodo.14340890" httpUri="https://zenodo.org/record/14340890/files/figure.png" pageId="5" pageNumber="6">Figs 5B–E</figureCitation>
,
<figureCitation id="7AAF7CD0066DFF9056EAFC5264146596" box="[1000,1058,969,993]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="5" pageNumber="6">6C, F</figureCitation>
<figureCitation id="7AAF7CD0066DFF9056EAFC5264146596" box="[1000,1058,969,993]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340894" httpUri="https://zenodo.org/record/14340894/files/figure.png" pageId="5" pageNumber="6">6C, F</figureCitation>
), although in some other of these vertebrae the neural canal transitioned smoothly into the internal cavity (compare
<figureCitation id="7AAF7CD0066DFF9056DBFB9364156257" box="[985,1059,1032,1056]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="5" pageNumber="6">Fig. 6B</figureCitation>
<figureCitation id="7AAF7CD0066DFF9056DBFB9364156257" box="[985,1059,1032,1056]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340894" httpUri="https://zenodo.org/record/14340894/files/figure.png" pageId="5" pageNumber="6">Fig. 6B</figureCitation>
to
<figureCitation id="7AAF7CD0066DFF90514BFB9364A26257" box="[1097,1172,1032,1056]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Fig. 5B</figureCitation>
<figureCitation id="7AAF7CD0066DFF90514BFB9364A26257" box="[1097,1172,1032,1056]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." figureDoi="http://doi.org/10.5281/zenodo.14340890" httpUri="https://zenodo.org/record/14340890/files/figure.png" pageId="5" pageNumber="6">Fig. 5B</figureCitation>
;
<figureCitation id="7AAF7CD0066DFF9051A1FB9364DA6268" box="[1187,1260,1032,1056]" captionStart="Figure 7" captionStartId="10.[129,194,695,719]" captionTargetBox="[289,1313,144,667]" captionTargetId="figure-733@10.[289,1313,144,667]" captionTargetPageId="10" captionText="Figure 7. Internal cavity volume in the cervical vertebrae of ‘P’.antiquus MGUWr 3895s (A) and Tanystropheus sp. from Miedary ZPAL V. 36/181 (B). Anatomical abbreviation:ic—internal cavity. Images obtained using ‘xray’ shader in MeshLab v.2020.12 (Cignoni et al.2008)." pageId="5" pageNumber="6">Fig. 7B</figureCitation>
<figureCitation id="7AAF7CD0066DFF9051A1FB9364DA6268" box="[1187,1260,1032,1056]" captionStart="Figure 7" captionStartId="10.[129,194,695,719]" captionTargetBox="[289,1313,144,667]" captionTargetId="figure-733@10.[289,1313,144,667]" captionTargetPageId="10" captionText="Figure 7. Internal cavity volume in the cervical vertebrae of ‘P’.antiquus MGUWr 3895s (A) and Tanystropheus sp. from Miedary ZPAL V. 36/181 (B). Anatomical abbreviation:ic—internal cavity. Images obtained using ‘xray’ shader in MeshLab v.2020.12 (Cignoni et al.2008)." figureDoi="http://doi.org/10.5281/zenodo.14340898" httpUri="https://zenodo.org/record/14340898/files/figure.png" pageId="5" pageNumber="6">Fig. 7B</figureCitation>
). In the transverse cross section, the inner surface of the bone wall of the internal cavity was generally parallel to the circumferential outline of the vertebra (
<figureCitation id="7AAF7CD0066DFF9056B4FBFD642D6209" box="[950,1051,1126,1150]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8A–E</figureCitation>
<figureCitation id="7AAF7CD0066DFF9056B4FBFD642D6209" box="[950,1051,1126,1150]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="5" pageNumber="6">Fig. 8A–E</figureCitation>
). In most of the studied specimens, the dorsal part of the internal cavity seemed to preserve a semicircular shape in the transverse cross section, whereas tissue layout of its ventrolateral portion was usually much less uniform, with an irregular resorption front cutting through multiple annuli (
<figureCitation id="7AAF7CD0066DFF905637FA986381636D" box="[821,951,1283,1307]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8A, C, E</figureCitation>
<figureCitation id="7AAF7CD0066DFF905637FA986381636D" box="[821,951,1283,1307]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="5" pageNumber="6">Fig. 8A, C, E</figureCitation>
). This was especially evident in larger specimens, whereas in the smallest cervical, the regions of resorption were much less developed, and the outline of the internal cavity in the transverse cross section was more regularly oval (
<figureCitation id="7AAF7CD0066DFF905029FAFA6542630E" box="[1323,1396,1377,1401]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8B</figureCitation>
<figureCitation id="7AAF7CD0066DFF905029FAFA6542630E" box="[1323,1396,1377,1401]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="5" pageNumber="6">Fig. 8B</figureCitation>
). The neural spine extended over two deep fossae, located on both of its anteroposteriorly terminal portions. They were triangular in longitudinal cross section and ventrally delimited by thin bone trabeculae, that also constituted the roof of the neural canal. The posterior cavity was larger, forming the ‘postzygapophyseal trough’ (
<bibRefCitation id="86051DA4066DFF905687F98664396042" author="Rieppel O" box="[901,1039,1565,1589]" pageId="5" pageNumber="6" pagination="271 - 87" refId="ref17778" refString="Rieppel O. A new species of Tanystropheus (Reptilia: Protorosauria) from the Middle Triassic of Makhtesh Ramon, Israel. Neues Jahrbuch fur Geologie und Palaontologie - Abhandlungen 2001; 221: 271 - 87. https: // doi. org / 10.1127 / njgpa / 221 / 2001 / 271" type="journal article" year="2001">Rieppel 2001</bibRefCitation>
). Contrary to what was observed for ‘
@ -136,14 +139,14 @@ spp.
middle cervicals did the internal cavity contain bony trabeculae extending internally from the surface of the bone wall. Some semi-symmetrical, slanted trabeculae of endosteal origin were present only incidentally in the anteroposteriorly terminal sections of the vertebrae (see
<bibRefCitation id="86051DA4066DFF905161F94264E36087" author="Broili F" box="[1123,1237,1752,1777]" pageId="5" pageNumber="6" pagination="51 - 62" refId="ref15553" refString="Broili F. Beobachtungen an Tanystropheus conspicuus H. v. Meyer. Neues Jahrbuch fur Mineralogie Geologie und Palaontologie 1915; 2: 51 - 62." type="journal article" year="1915">Broili 1915</bibRefCitation>
: plate 3, fig. 4c). The roof of the neural canal was generally straight and parallel to the longitudinal axis of the vertebra. The ventral delimitation of the neural canal was seemingly absent for most of its length within a vertebra. Intriguingly, in one of the transverse polished sections from the anterior part of U-MO BT 738.00 (
<figureCitation id="7AAF7CD0066DFF90500AF8EE656761FA" box="[1288,1361,1909,1933]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8F</figureCitation>
<figureCitation id="7AAF7CD0066DFF90500AF8EE656761FA" box="[1288,1361,1909,1933]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="5" pageNumber="6">Fig. 8F</figureCitation>
), minute bony projections appeared symmetrically on each lateral side of the internal cavity. The colour of the limestone infill of the vertebra changed drastically at the same horizontal level, at which these projections were situated. Similar characteristics could be noted for the transverse thin section of ZPAL V. 36/166 (
<figureCitation id="7AAF7CD0066EFF9357E1F8E460AD61C0" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="6" pageNumber="7">Fig. 8E</figureCitation>
<figureCitation id="7AAF7CD0066EFF9357E1F8E460AD61C0" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="6" pageNumber="7">Fig. 8E</figureCitation>
), in which corresponding projections composed of periosteal bone were also present in the dorsolateral regions of the internal cavity. Between them, there was a sharp transition in sediment fraction, with the more coarse-grained infill occupying the ventral-more space. Presence of this feature possibly shows the differences in taphonomical microenvironments of the ventral and dorsal portions of the internal cavity, which may have been caused by the original existence of a no longer preserved barrier between them. Near the anteroposterior midpoint of the middle cervicals, two symmetrically placed foramina were located on the ventral side of the centrum. They entered the internal cavity as straight canals (
<figureCitation id="7AAF7CD00660FF9D5439FA5C61AD63A9" box="[315,411,1479,1503]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="8" pageNumber="9">Fig. 8B, E</figureCitation>
<figureCitation id="7AAF7CD00660FF9D5439FA5C61AD63A9" box="[315,411,1479,1503]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="8" pageNumber="9">Fig. 8B, E</figureCitation>
), perforating the dense cortex.
</paragraph>
<caption id="B6EB30DD066EFF935583F91E618B6086" pageId="6" pageNumber="7" startId="6.[129,194,1669,1693]" targetBox="[305,1297,144,1642]" targetPageId="6" targetType="figure">
<caption id="B6EB30DD066EFF935583F91E618B6086" ID-DOI="http://doi.org/10.5281/zenodo.14340884" ID-Zenodo-Dep="14340884" httpUri="https://zenodo.org/record/14340884/files/figure.png" pageId="6" pageNumber="7" startId="6.[129,194,1669,1693]" targetBox="[305,1297,144,1642]" targetPageId="6" targetType="figure">
<paragraph id="E22B6055066EFF935583F91E618B6086" blockId="6.[129,1449,1669,1777]" pageId="6" pageNumber="7">
<emphasis id="D0E0BC47066EFF935583F91E60EE60EA" bold="true" box="[129,216,1669,1693]" pageId="6" pageNumber="7">Figure 3.</emphasis>
CT transverse cross sections (from anterior to posterior) of ‘
@ -155,7 +158,7 @@ CT transverse cross sections (from anterior to posterior) of ‘
cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum.
</paragraph>
</caption>
<caption id="B6EB30DD066FFF9D5573F8AA634161CD" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8" startId="7.[113,178,1841,1865]" subCaptionStartIDs="8.[809,941,1899,1923]" subCaptionStarts="abbr" targetBox="[175,1398,149,1813]" targetPageId="7" targetType="figure">
<caption id="B6EB30DD066FFF9D5573F8AA634161CD" ID-DOI="http://doi.org/10.5281/zenodo.14340888" ID-Zenodo-Dep="14340888" httpUri="https://zenodo.org/record/14340888/files/figure.png" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8" startId="7.[113,178,1841,1865]" subCaptionStartIDs="8.[809,941,1899,1923]" subCaptionStarts="abbr" targetBox="[175,1398,149,1813]" targetPageId="7" targetType="figure">
<paragraph id="E22B6055066FFF9D5573F8AA634161CD" blockId="7.[113,1457,1841,1978]" lastBlockId="8.[129,1454,1843,1979]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
<emphasis id="D0E0BC47066FFF925573F8AA60FF613E" bold="true" box="[113,201,1841,1865]" pageId="7" pageNumber="8">Figure 4.</emphasis>
Thin sections of ‘
@ -171,7 +174,7 @@ and
for D–J.
</paragraph>
</caption>
<caption id="B6EB30DD0660FF9D5580FBE962FF628D" pageId="8" pageNumber="9" startId="8.[130,195,1138,1162]" targetBox="[289,1313,144,1110]" targetPageId="8" targetType="figure">
<caption id="B6EB30DD0660FF9D5580FBE962FF628D" ID-DOI="http://doi.org/10.5281/zenodo.14340890" ID-Zenodo-Dep="14340890" httpUri="https://zenodo.org/record/14340890/files/figure.png" pageId="8" pageNumber="9" startId="8.[130,195,1138,1162]" targetBox="[289,1313,144,1110]" targetPageId="8" targetType="figure">
<paragraph id="E22B60550660FF9D5580FBE962FF628D" blockId="8.[129,1462,1138,1274]" pageId="8" pageNumber="9">
<emphasis id="D0E0BC470660FF9D5580FBE960EC62FD" bold="true" box="[130,218,1138,1162]" pageId="8" pageNumber="9">Figure 5.</emphasis>
Middle cervicals of
@ -200,11 +203,11 @@ It was impossible to demarcate the border between the neural arch and the verteb
spp.
</taxonomicName>
the outermost of these zones formed complete rings, whereas the ones located innermost were discontinued only due to resorption of the region ventral into the internal cavity, as well as the presence of the neural spine (
<figureCitation id="7AAF7CD00660FF9D5176FA3C64D663B7" box="[1140,1248,1447,1472]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="8" pageNumber="9">Fig. 8A, C</figureCitation>
<figureCitation id="7AAF7CD00660FF9D5176FA3C64D663B7" box="[1140,1248,1447,1472]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="8" pageNumber="9">Fig. 8A, C</figureCitation>
,
<figureCitation id="7AAF7CD00660FF9D51ECFA3C650E63B7" box="[1262,1336,1447,1472]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." pageId="8" pageNumber="9">Fig. 9B</figureCitation>
<figureCitation id="7AAF7CD00660FF9D51ECFA3C650E63B7" box="[1262,1336,1447,1472]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." figureDoi="http://doi.org/10.5281/zenodo.14340904" httpUri="https://zenodo.org/record/14340904/files/figure.png" pageId="8" pageNumber="9">Fig. 9B</figureCitation>
). In some of the studied transverse thin sections, including the smallest specimens, the spinous process was developed only as a thin, vertically oriented plate. It was located directly above the internal cavity and emerged from the parallel-fibred tissue of the bone walls in the dorsalmost portion of the vertebra, projecting slightly from the smooth outline of the transverse cross section. In the smallest specimen from Miedary, ZPAL V. 36/181, the neural spine was never overlain by parallel-fibred bone (
<figureCitation id="7AAF7CD00660FF9D504AF93965A460CD" box="[1352,1426,1698,1722]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="8" pageNumber="9">Fig. 8B</figureCitation>
<figureCitation id="7AAF7CD00660FF9D504AF93965A460CD" box="[1352,1426,1698,1722]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="8" pageNumber="9">Fig. 8B</figureCitation>
). In all of the larger specimens from Miedary, the anteroposteriorly middle part of the spinous process was obscured by thick layers of tissue that formed the walls of the internal cavity. Compared to what was noted for ‘
<emphasis id="D0E0BC470661FF9C5464FA05614263C2" box="[358,372,1438,1461]" italics="true" pageId="9" pageNumber="10">P</emphasis>
’.
@ -217,7 +220,7 @@ the outermost of these zones formed complete rings, whereas the ones located inn
spp.
</taxonomicName>
cervicals were thickened (see
<figureCitation id="7AAF7CD00661FF9C5787FA26628B63A2" box="[645,701,1469,1493]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." pageId="9" pageNumber="10">Fig. 9</figureCitation>
<figureCitation id="7AAF7CD00661FF9C5787FA26628B63A2" box="[645,701,1469,1493]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." figureDoi="http://doi.org/10.5281/zenodo.14340904" httpUri="https://zenodo.org/record/14340904/files/figure.png" pageId="9" pageNumber="10">Fig. 9</figureCitation>
). The neural spine was, thus, not relatively reduced in the latter, but rather embedded within the walls of the tubularly structured vertebra. As a result, the neural canal was located close to the middle of the height of the vertebra, as in ‘
<emphasis id="D0E0BC470661FF9C571DF9A0621B6025" box="[543,557,1595,1618]" italics="true" pageId="9" pageNumber="10">P</emphasis>
’.
@ -230,10 +233,10 @@ cervicals were thickened (see
spp.
</taxonomicName>
there was more tissue dorsolaterally (see
<figureCitation id="7AAF7CD00661FF9C5573F9E2609A60E6" box="[113,172,1657,1681]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." pageId="9" pageNumber="10">Fig. 9</figureCitation>
<figureCitation id="7AAF7CD00661FF9C5573F9E2609A60E6" box="[113,172,1657,1681]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." figureDoi="http://doi.org/10.5281/zenodo.14340904" httpUri="https://zenodo.org/record/14340904/files/figure.png" pageId="9" pageNumber="10">Fig. 9</figureCitation>
), due to the neural spine being hypertrophied along the middle section of its anteroposterior length.
</paragraph>
<caption id="B6EB30DD0661FF9C5573FB3C64746358" pageId="9" pageNumber="10" startId="9.[113,178,1191,1215]" targetBox="[146,1426,144,1163]" targetPageId="9" targetType="figure">
<caption id="B6EB30DD0661FF9C5573FB3C64746358" ID-DOI="http://doi.org/10.5281/zenodo.14340894" ID-Zenodo-Dep="14340894" httpUri="https://zenodo.org/record/14340894/files/figure.png" pageId="9" pageNumber="10" startId="9.[113,178,1191,1215]" targetBox="[146,1426,144,1163]" targetPageId="9" targetType="figure">
<paragraph id="E22B60550661FF9C5573FB3C64746358" blockId="9.[113,1454,1191,1327]" pageId="9" pageNumber="10">
<emphasis id="D0E0BC470661FF9C5573FB3C60FF62C8" bold="true" box="[113,201,1191,1215]" pageId="9" pageNumber="10">Figure 6.</emphasis>
Vertebrae of
@ -254,9 +257,9 @@ In
sp.
</taxonomicName>
from Miedary, the external morphology of the cervicals changed near the posterior subregion of the neck. The 10th and 11th vertebrae were still elongate, but, contrary to the middle cervicals, exhibited a well exposed, relatively tall and continuous neural spine (
<figureCitation id="7AAF7CD00661FF9C54B5F8E861C361FC" box="[439,501,1907,1931]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="9" pageNumber="10">Fig. 6</figureCitation>
<figureCitation id="7AAF7CD00661FF9C54B5F8E861C361FC" box="[439,501,1907,1931]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340894" httpUri="https://zenodo.org/record/14340894/files/figure.png" pageId="9" pageNumber="10">Fig. 6</figureCitation>
). The two last cervicals were relatively much shorter (
<figureCitation id="7AAF7CD00661FF9C54AEF808622361DC" box="[428,533,1939,1963]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="9" pageNumber="10">Fig. 6B, C</figureCitation>
<figureCitation id="7AAF7CD00661FF9C54AEF808622361DC" box="[428,533,1939,1963]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340894" httpUri="https://zenodo.org/record/14340894/files/figure.png" pageId="9" pageNumber="10">Fig. 6B, C</figureCitation>
), and more similar in proportions to the dorsals (
<bibRefCitation id="86051DA40661FF9C513DFAC464CE6300" author="Rieppel O &amp; Jiang DY &amp; Fraser NC" box="[1087,1272,1375,1399]" pageId="9" pageNumber="10" pagination="1082 - 9" refId="ref17833" refString="Rieppel O, Jiang DY, Fraser NC et al. Tanystropheus cf. T. longobardicus from the early Late Triassic of Guizhou Province, Southwestern China. Journal of Vertebrate Paleontology 2010; 30: 1082 - 9. https: // doi. org / 10.1080 / 02724634.2010.483548" type="journal article" year="2010">
Rieppel
@ -270,29 +273,29 @@ Rytel
2024
</bibRefCitation>
). This transition was paired with the modifications in the internal structure of these elements. The relative volume occupied by the trabeculae within the centra increased caudally, especially along the neck-torso transition. In the middle cervicals the centrum was nearly devoid of trabecular bone, but this changed within the more posterior vertebrae, with the last two cervicals and the dorsals exhibiting sparse trabeculae occupying the space between the articular discs. The 10th and 11th vertebrae were generally similar in their anatomy to the middle cervicals, with their internal cavities being surrounded by a dense cortex, forming a tube-like structure (
<figureCitation id="7AAF7CD00661FF9C56F8F92C640B60B8" box="[1018,1085,1719,1743]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="9" pageNumber="10">Fig. 6I</figureCitation>
<figureCitation id="7AAF7CD00661FF9C56F8F92C640B60B8" box="[1018,1085,1719,1743]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340894" httpUri="https://zenodo.org/record/14340894/files/figure.png" pageId="9" pageNumber="10">Fig. 6I</figureCitation>
). However, in the succeeding vertebrae the internal cavity was greatly reduced and exhibited much thicker walls, composed of two regions: a relatively thin cortex and porous, cancellous bone encompassed by it. Especially in the middle portion of the centrum of the last two cervicals, there were several large chambers, similar to those present in the anteroposteriorly terminal portions of the more anteriorly located vertebrae (
<figureCitation id="7AAF7CD00661FF9C56F0F808647761DC" box="[1010,1089,1939,1963]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="9" pageNumber="10">Fig. 8D</figureCitation>
<figureCitation id="7AAF7CD00661FF9C56F0F808647761DC" box="[1010,1089,1939,1963]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="9" pageNumber="10">Fig. 8D</figureCitation>
). Some of them connected to the neural canal through its floor (
<figureCitation id="7AAF7CD00662FF9F54C3FC80626A6544" box="[449,604,795,819]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6E, F, H, I</figureCitation>
<figureCitation id="7AAF7CD00662FF9F54C3FC80626A6544" box="[449,604,795,819]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340894" httpUri="https://zenodo.org/record/14340894/files/figure.png" pageId="10" pageNumber="11">Fig. 6E, F, H, I</figureCitation>
). The transverse cross sections of the posteriormost cervicals were less oval than in the preceding vertebrae, with the centrum being much more ventrally expanded, forming a distinct keel (
<figureCitation id="7AAF7CD00662FF9F5762FCE262F165E6" box="[608,711,889,913]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6J, K</figureCitation>
<figureCitation id="7AAF7CD00662FF9F5762FCE262F165E6" box="[608,711,889,913]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340894" httpUri="https://zenodo.org/record/14340894/files/figure.png" pageId="10" pageNumber="11">Fig. 6J, K</figureCitation>
). The diameter of the neural canal was relatively constant along the length of each of the two last vertebrae. In the longitudinal cross section, it had a sagging appearance, with its middle portion being located more ventrally. In the transverse cross section, it was oval, with its dorsolateral regions being roughly semicircular and symmetrical in outline (
<figureCitation id="7AAF7CD00662FF9F54A3FBAE61CB623A" box="[417,509,1077,1101]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6J, K</figureCitation>
<figureCitation id="7AAF7CD00662FF9F54A3FBAE61CB623A" box="[417,509,1077,1101]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340894" httpUri="https://zenodo.org/record/14340894/files/figure.png" pageId="10" pageNumber="11">Fig. 6J, K</figureCitation>
). A similar shape could be noted in the anteroposteriorly terminal sections of the rest of the postaxial cervicals, in which the neural canal floor was still present (e.g.
<figureCitation id="7AAF7CD00662FF9F55E1FB08611862DC" box="[227,302,1171,1195]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="10" pageNumber="11">Fig. 8D</figureCitation>
<figureCitation id="7AAF7CD00662FF9F55E1FB08611862DC" box="[227,302,1171,1195]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="10" pageNumber="11">Fig. 8D</figureCitation>
), and also in the dorsal vertebrae (
<figureCitation id="7AAF7CD00662FF9F5794FB0862EB62DC" box="[662,733,1171,1195]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6L</figureCitation>
<figureCitation id="7AAF7CD00662FF9F5794FB0862EB62DC" box="[662,733,1171,1195]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340894" httpUri="https://zenodo.org/record/14340894/files/figure.png" pageId="10" pageNumber="11">Fig. 6L</figureCitation>
). In each of the two posteriormost neck vertebrae of
<taxonomicName id="25941BD60662FF9F577FFB2860AA629E" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="D0E0BC470662FF9F577FFB28633F62BC" box="[637,777,1203,1227]" italics="true" pageId="10" pageNumber="11">Tanystropheus</emphasis>
sp.
</taxonomicName>
from Miedary, up to two small subcentral foramina could be traced, with some specimens exhibiting only one subcentral foramen or none at all. In the studied posteriormost cervical ZPAL V. 36/110, a straight, non-branching canal connected the one present foramen with the neural canal (
<figureCitation id="7AAF7CD00662FF9F5712FAD462636310" box="[528,597,1359,1383]" captionStart="Figure 10" captionStartId="13.[113,178,1140,1164]" captionTargetBox="[306,1266,144,1112]" captionTargetId="figure-458@13.[306,1266,144,1112]" captionTargetPageId="13" captionText="Figure 10. Longitudinal cross section of ZPAL V.36/110, 13th (last) cervical vertebra of Tanystropheus sp.from Miedary, showing the course of the arterial canal (locations of its end sections are indicated with arrows). The anterior end of the vertebra points towards the left. Anatomical abbreviations: nc—neural canal; ns—neural spine." pageId="10" pageNumber="11">Fig. 10</figureCitation>
<figureCitation id="7AAF7CD00662FF9F5712FAD462636310" box="[528,597,1359,1383]" captionStart="Figure 10" captionStartId="13.[113,178,1140,1164]" captionTargetBox="[306,1266,144,1112]" captionTargetId="figure-458@13.[306,1266,144,1112]" captionTargetPageId="13" captionText="Figure 10. Longitudinal cross section of ZPAL V.36/110, 13th (last) cervical vertebra of Tanystropheus sp.from Miedary, showing the course of the arterial canal (locations of its end sections are indicated with arrows). The anterior end of the vertebra points towards the left. Anatomical abbreviations: nc—neural canal; ns—neural spine." figureDoi="http://doi.org/10.5281/zenodo.14340906" httpUri="https://zenodo.org/record/14340906/files/figure.png" pageId="10" pageNumber="11">Fig. 10</figureCitation>
).
</paragraph>
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<paragraph id="E22B60550662FF9F5583FD2C65AF649B" blockId="10.[129,1433,695,748]" pageId="10" pageNumber="11">
<emphasis id="D0E0BC470662FF9F5583FD2C60EE64B8" bold="true" box="[129,216,695,719]" pageId="10" pageNumber="11">Figure 7.</emphasis>
Internal cavity volume in the cervical vertebrae of ‘
@ -317,7 +320,7 @@ Cignoni
</caption>
<paragraph id="E22B60550662FF9F559EFAF462AB60B7" blockId="10.[128,779,795,1728]" pageId="10" pageNumber="11">
The dorsal vertebrae were generally similar to the posteriormost cervicals in their internal anatomy. They differed in a complete lack of subcentral foramina. Moreover, the neural canal of the dorsals extended anteroposteriorly straight, with its roof and floor remaining on a relatively constant horizontal level (
<figureCitation id="7AAF7CD00662FF9F5589F99060E16054" box="[139,215,1547,1571]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6H</figureCitation>
<figureCitation id="7AAF7CD00662FF9F5589F99060E16054" box="[139,215,1547,1571]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340894" httpUri="https://zenodo.org/record/14340894/files/figure.png" pageId="10" pageNumber="11">Fig. 6H</figureCitation>
). Its transverse cross section was similar in shape to the posteriormost cervicals, but more regular. Congruently to what could be observed in the 12th and 13th cervical vertebrae, the internal portion of the centrum was composed of sparsely distributed trabeculae containing a large number of cavities, some of which opened to the neural canal through its floor.
</paragraph>
<paragraph id="E22B60550662FF9F546DF944622D608E" blockId="10.[367,539,1759,1785]" box="[367,539,1759,1785]" pageId="10" pageNumber="11">
@ -334,11 +337,11 @@ were congruent with the results of the study performed by
and the supplemental data provided therein). New data presented herein expand our knowledge on the anteroposterior and ontogenetic variation of their structure. The walls of the internal cavity were built of primary compact lamellar to parallel-fibred matrix of periosteal origin. Some intervals with decreased vascularization could be traced within these bone walls—they formed ellipsoidal annuli that alternated with more vascularized zones in parallel to the circumference of the vertebra (see
<bibRefCitation id="86051DA40662FF9F51A4FC02658765C6" author="Jaquier VP &amp; Scheyer TM" box="[1190,1457,921,945]" pageId="10" pageNumber="11" refId="ref16697" refString="Jaquier VP, Scheyer TM. Bone histology of the Middle Triassic long-necked reptiles Tanystropheus and Macrocnemus (Archosauromorpha, Protorosauria). Journal of Vertebrate Paleontology 2017; 37: e 1296456. https: // doi. org / 10.1080 / 02724634.2017.1296456" type="journal volume" year="2017">Jaquier and Scheyer 2017</bibRefCitation>
). Closer to the terminal portions of the anteroposterior length of the cervicals, the tissue composition changed, with the internal cavity being dorsolaterally lined with secondary endosteal lamellar bone (
<figureCitation id="7AAF7CD00662FF9F56C0FB8D64386259" box="[962,1038,1046,1070]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="10" pageNumber="11">Fig. 8H</figureCitation>
<figureCitation id="7AAF7CD00662FF9F56C0FB8D64386259" box="[962,1038,1046,1070]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="10" pageNumber="11">Fig. 8H</figureCitation>
) and the ventrolateral portions of the bone centrum not exhibiting the regular annuli, but rather being nearly completely composed of endosteal lamellar bone scattered with secondary osteons and erosion cavities of varying size, forming a net of sparse trabeculae (
<figureCitation id="7AAF7CD00662FF9F5150FB0864F662DC" box="[1106,1216,1171,1195]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="10" pageNumber="11">Figs 5B–D</figureCitation>
<figureCitation id="7AAF7CD00662FF9F5150FB0864F662DC" box="[1106,1216,1171,1195]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." figureDoi="http://doi.org/10.5281/zenodo.14340890" httpUri="https://zenodo.org/record/14340890/files/figure.png" pageId="10" pageNumber="11">Figs 5B–D</figureCitation>
,
<figureCitation id="7AAF7CD00662FF9F51CEFB08653D62DC" box="[1228,1291,1171,1195]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="10" pageNumber="11">8D, G</figureCitation>
<figureCitation id="7AAF7CD00662FF9F51CEFB08653D62DC" box="[1228,1291,1171,1195]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." figureDoi="http://doi.org/10.5281/zenodo.14340900" httpUri="https://zenodo.org/record/14340900/files/figure.png" pageId="10" pageNumber="11">8D, G</figureCitation>
). The core of the neural spine in larger specimens was highly remodelled with longitudinally oriented secondary osteons. Simple primary vascular canals extended radially in the middle part of the vertebrae, but became more longitudinally oriented, larger, and less organized in the anteroposteriorly terminal portions. The number of vascular canals was also smaller in the more compact sections of the cortex within a single cross section.
</paragraph>
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<treatment id="7ADFD4E0F42D56CE8DA4AAFCB53675A2" ID-arpha="7ADFD4E0-F42D-56CE-8DA4-AAFCB53675A2" ID-zoobank="https://zoobank.org/3A13F9F3-BD1E-4AF5-B1A3-4BBD7D604634" LSID="urn:lsid:plazi:treatment:7ADFD4E0F42D56CE8DA4AAFCB53675A2" LSID-ZBK="urn:lsid:zoobank.org:act:3A13F9F3-BD1E-4AF5-B1A3-4BBD7D604634" httpUri="http://treatment.plazi.org/id/7ADFD4E0F42D56CE8DA4AAFCB53675A2" scope_family="Ophichthidae" scope_order="Anguilliformes">
<subSubSection id="49272E65E3CB457CB5E25903AFBD8854" type="nomenclature">
<paragraph id="069F616248670FCB8FB0D6BE799EE53A">
<taxonomicName id="5C4319FC7D7687F5FDF9A856073B3167" authority="Hibino &amp; Ho" authorityName="Hibino &amp; Ho" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="kbalanensis" status="sp. nov.">
<emphasis id="308CB1EF4ACA2C72B8C3F598E8C8D1CD" italics="true">Ophichthus kbalanensis</emphasis>
Hibino &amp; Ho
</taxonomicName>
<taxonomicNameLabel id="4F866C4572CD8FE7CC8A3EFAE734FBB1" rank="species">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="D440BBA08A184519004C54FF64693F94" type="description">
<paragraph id="39D9910C9394AC259001475FE2EB47A2">
<figureCitation id="4A269C3E8337785D9E20BE6272DE628E" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Ophichthus kbalanensis sp. nov., NMMB-P 26381, holotype, 414 mm TL, Da-xi, Taiwan A fresh condition B preserved condition. Arrows indicate positions of dorsal-fin origin (lefts) and anus (rights)." figureDoi="10.3897/zookeys.1220.126337.figure1" httpUri="https://binary.pensoft.net/fig/1194398">Figs 1</figureCitation>
,
<figureCitation id="C79F8C99C9CDF6EBBE7DF28AC18312E4" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Line drawings of O. kbalanensis sp. nov., NMMB-P 26381, holotype, 414 mm TL A sensory pores on head B teeth on upper (left) and lower jaws (right). Arrows indicate interorbital (left) and mid-temporal pores (right); black solid circle indicates holes of lost tooth; and broken lines indicate concealed teeth by fleshy lips." figureDoi="10.3897/zookeys.1220.126337.figure2" httpUri="https://binary.pensoft.net/fig/1194399">2</figureCitation>
;
<tableCitation id="310B34BD71452AC4402AEF51A49BC00D" captionStart="Table 1" captionStartId="T1" captionText="Table 1. Counts and measurements of two new Ophichthus from Taiwan." httpUri="http://table.plazi.org/id/78971F2C976540034B269033BAC72C47" tableUuid="78971F2C976540034B269033BAC72C47">Tables 1</tableCitation>
,
<tableCitation id="83D5D1B3DB4CFEAD2928FE0599A90BCB" captionStart="Table 2" captionStartId="T2" captionText="Table 2. Selected characters of Ophichthus species reported from Taiwan, except patterned species." httpUri="http://table.plazi.org/id/3CC3B79BF31298505EAC7889535A776B" tableUuid="3CC3B79BF31298505EAC7889535A776B">2</tableCitation>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EMKAC" type="material">
<paragraph id="1AB0291C3DAC7FF7802DB9859F57106A">
<heading id="797DEBD05AD761702A1BF4618598DFD1" reason="title">Material examined.</heading>
</paragraph>
<paragraph id="6BB39522EB83A944F55BF3ECD3CE59A6">
<materialsCitation id="66C30F68E50A57185AFA360D2A6AB762" collectingDate="2017-07-01" collectionCode="NMMB-P" country="Taiwan" latitude="24.9" location="Da-xi" longLatPrecision="124" longitude="121.933334" specimenCode="NMMB-P 26381" specimenCount="1" stateProvince="Yilan" typeStatus="holotype">
<emphasis id="CC80B19FFF15273197BEA10AEF82EF5C" bold="true" italics="true">
<typeStatus id="0BBD6BD16BB26D286F0FAACF3203688C" type="holotype">Holotype</typeStatus>
</emphasis>
:
<specimenCode id="0EB683070F7B995D5C6565DCA5433122">NMMB-P 26381</specimenCode>
,
<quantity id="AE5CDE195D266B14A96BD71ADCFB0D8C" metricMagnitude="-1" metricUnit="m" metricValue="4.14" unit="mm" value="2.6381414E7">414 mm</quantity>
<abbrev id="ABBRID0EXKAC" xlink_title="Total lengths">TL</abbrev>
, ca
<named-content content-type="dwc:verbatimCoordinates" id="NCID0E5KAC" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[121.933333,24.900000]}">
<geoCoordinate id="6170F0B607917395D09A6EDA7A82FA58" degrees="24" direction="north" minutes="54.0" orientation="latitude" precision="92" value="24.9">24°54.0'N</geoCoordinate>
,
<geoCoordinate id="BDBAF0CE8F1C9DF923DE8DE3F261DE39" degrees="121" direction="east" minutes="56.0" orientation="longitude" precision="92" value="121.933334">121°56.0'E</geoCoordinate>
</named-content>
,
<location id="61C120241628A6EA66FE334168115CEC" LSID="urn:lsid:plazi:treatment:7ADFD4E0F42D56CE8DA4AAFCB53675A2:61C120241628A6EA66FE334168115CEC" country="Taiwan" latitude="24.9" longLatPrecision="124" longitude="121.933334" municipality="northwestern Pacific Ocean" name="Da-xi" stateProvince="Yilan">Da-xi</location>
,
<collectingRegion id="E5BFEA673CAAEE1D15B699E9E82F182E" country="Taiwan" name="Yilan">Yilan</collectingRegion>
, northeastern
<collectingCountry id="EAB7E062D77314A6B841FF285BA386F1" name="Taiwan">Taiwan</collectingCountry>
, northwestern Pacific Ocean,
<date id="9BC799DAE997744E0A5F43B2C44C0AA0" value="2017-07-01">
<collectingDate id="CA584F402ABEB2C96A8D01591482BF6E" value="2017-07-01">1 Jul. 2017</collectingDate>
</date>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EDLAC" type="diagnosis">
<paragraph id="B5C516552E8FE7DC513A33819105F358">
<heading id="4A16C889A55100B7851B2397E3095926" reason="title">Diagnosis.</heading>
</paragraph>
<paragraph id="D4A00136C9B0EBB35290D426338ED067">
A relatively short
<taxonomicName id="CAC63708B2560CCE2096F5159238138E" authorityName="Ahl" authorityYear="1789" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="genus">
<emphasis id="5992C52BF42E621B168F56DFD0DC1634" italics="true">Ophichthus</emphasis>
</taxonomicName>
with the following combination of characters: head 10.3 %
<abbrev id="ABBRID0EQLAC" xlink_title="Total lengths">TL</abbrev>
; tail 62.7 %
<abbrev id="ABBRID0EULAC" xlink_title="Total lengths">TL</abbrev>
; dorsal-fin origin above about middle of pectoral fin; tip of lower jaw anterior to anterior-nostril base; two simple, distally pointed protrusions along upper lip;
<abbrev id="ABBRID0EYLAC" xlink_title="supraorbital pores">SO</abbrev>
1 + 4;
<abbrev id="ABBRID0E3LAC" xlink_title="preoperculomandibular pores">POM</abbrev>
6 or 7 + 3; teeth on jaws and vomer mostly uniserial but posterior ends of maxilla and anterior end of symphysis biserial; body dark; median fins with narrow dark margins, except the pale fin origins; 14 predorsal and 53 preanal lateral-line pores; VF 12-55 - 153.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EAMAC" type="description">
<paragraph id="17F0BEC3AA3F31045EAA8CE12A07B7DD">
<heading id="F2D2DAFE55CBCEC012A0BD8B2B46D7B3" reason="title">Description.</heading>
</paragraph>
<paragraph id="D4F5B4E2F5524FFABCD3E3896AFEA6E1">
Counts and measurements are mostly shown in Tables
<tableCitation id="1A1A2851EB4BF29F591F2B37B17DB8FC" captionStart="Table 1" captionStartId="T1" captionText="Table 1. Counts and measurements of two new Ophichthus from Taiwan." httpUri="http://table.plazi.org/id/78971F2C976540034B269033BAC72C47" tableUuid="78971F2C976540034B269033BAC72C47">1</tableCitation>
,
<tableCitation id="0EACD71980719B6A27B4729AB25A24DF" captionStart="Table 2" captionStartId="T2" captionText="Table 2. Selected characters of Ophichthus species reported from Taiwan, except patterned species." httpUri="http://table.plazi.org/id/3CC3B79BF31298505EAC7889535A776B" tableUuid="3CC3B79BF31298505EAC7889535A776B">2</tableCitation>
.
</paragraph>
<caption id="78971F2C976540034B269033BAC72C47" ID-Table-UUID="78971F2C976540034B269033BAC72C47" httpUri="http://table.plazi.org/id/78971F2C976540034B269033BAC72C47" startId="T1" targetIsTable="true">
<paragraph id="C20559085C11CE854F89C46B4EC09CEA">
<label id="0E9E35AC3A866895DC5BE5BCF44FB17E">Table 1.</label>
</paragraph>
<paragraph id="C2CE5444515DCFF4264C111C424FF199">
Counts and measurements of two new
<taxonomicName id="5B81E0264470A5D06FF3C9CA2EF61001" authorityName="Ahl" authorityYear="1789" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="genus">
<emphasis id="4B2C07D27FE973BD85A5AA89D0F4ACED" italics="true">Ophichthus</emphasis>
</taxonomicName>
from Taiwan.
</paragraph>
</caption>
<paragraph id="3F64F2EF10110F73B6400DC23DB11700">
<table id="TID0EHXAG" rules="all">
<tbody id="4D07AE1D2A306D69E20470E00DC2576F">
<tr id="CA642269076BC9398C082DA2EAFA12C9">
<th id="2DB69E507B3F32BF53EB15DDE2EA3F7D" colspan="1" isEmpty="true" rowspan="3">-</th>
<th id="FF5CE73CB037B2C2C8E868A53CAFF464" colspan="1" rowspan="1">
<taxonomicName id="861E7FBF6F15E516A65662F7E3661734" authorityName="Hibino &amp; Ho" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="kbalanensis" status="sp. nov.">
<emphasis id="D654C5437BEB63158DFD1DEEBC7826C0" italics="true">O. kbalanensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="D6D342792769FCE1E5E741E59C948A23" rank="species">sp. nov.</taxonomicNameLabel>
</th>
<th id="E9DD5E491920BBD29F21B0410DCE1044" colspan="5" rowspan="1">
<taxonomicName id="1CB91A80201F83AC3E81E5CF4AB13075" authorityName="Hibino, Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<emphasis id="6999EF8D1824A307FF7EDE7B62A538FA" italics="true">O. multidentis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C31193861093D14E2635A61B837EA249" rank="species">sp. nov.</taxonomicNameLabel>
</th>
</tr>
<tr id="688DCF66869DEBBC4507361BFAB745BF">
<th id="C360F3465C762C0C672D42D27F7FC4ED" colspan="1" rowspan="1">Holotype</th>
<th id="AF8A375E90D6D6EF638F238EC66F8C42" colspan="1" rowspan="1">Holotype</th>
<th id="7E91091729252002113B6780C17E87E6" colspan="4" rowspan="1">Paratypes</th>
</tr>
<tr id="2A74D633FAF0FE80A53CBE3013F5176E">
<th id="0B7C7CB1923272314D0EBC4A0DAAA80F" colspan="1" rowspan="1">
<specimenCode id="4EC08C169D6B2C46168A91EAF5FE2009">NMMB-P 26381</specimenCode>
</th>
<th id="34F80070FBDA8084C21D620624A4E780" colspan="1" rowspan="1">
<specimenCode id="4023CA2E4B4BC7A6996BB4670A695C2A">NMMB-P 36205</specimenCode>
</th>
<th id="8D22344FAEDD9DA9415521995A9F17C9" colspan="1" rowspan="1">
<specimenCode id="9F800BB58EDFB07912E6C1FC1E017496">
<abbrev id="ABBRID0EZOAC" xlink_title="National Taiwan Ocean University, Laboratory of Aquatic Ecology, Department of Aquaculture">TOU-AE</abbrev>
7802
</specimenCode>
</th>
<th id="823FCBBA9AD53FBF30900AFEB98D1AB4" colspan="1" rowspan="1">
<specimenCode id="133E372ED4B1D01D7E92E7FDC399D9BD">
<abbrev id="ABBRID0EBPAC" xlink_title="National Taiwan Ocean University, Laboratory of Aquatic Ecology, Department of Aquaculture">TOU-AE</abbrev>
8998
</specimenCode>
</th>
<th id="479DB61336801654095F85580894D35C" colspan="1" rowspan="1">
<specimenCode id="ABD779BE3B038D17F00FADB95CF89924">
<abbrev id="ABBRID0EJPAC" xlink_title="National Taiwan Ocean University, Laboratory of Aquatic Ecology, Department of Aquaculture">TOU-AE</abbrev>
8999
</specimenCode>
</th>
<th id="40A5B71FF8513ABC6AD8D752CE919A2B" colspan="1" rowspan="1">
<specimenCode id="60A92B3DC95C1C5DEE3CAD4144EFAE02">
<abbrev id="ABBRID0ERPAC" xlink_title="National Taiwan Ocean University, Laboratory of Aquatic Ecology, Department of Aquaculture">TOU-AE</abbrev>
9294
</specimenCode>
</th>
</tr>
<tr id="A627F83A37BAAF75854A797B7D99387D">
<td id="2E2F781515A91994D514E4A4AB9CC07D" colspan="1" rowspan="1">Total length (mm)</td>
<td id="E5C0EE96CAC12661FAD5864F80F479B4" colspan="1" rowspan="1">414</td>
<td id="786AC4FB79B8B11F56064F364243B4EF" colspan="1" rowspan="1">433</td>
<td id="BD697A1739C6BD648DC8D0015700C995" colspan="1" rowspan="1">519</td>
<td id="05969891CE11B2ED3F289414E68366B4" colspan="1" rowspan="1">554</td>
<td id="FFCC247C6A3148638199404ABB510C00" colspan="1" rowspan="1">597</td>
<td id="F004B620E8A821D9415E8CBDE875EBC7" colspan="1" rowspan="1">696</td>
</tr>
<tr id="9B27965AB1B0983299253AD309DFE1C9">
<td id="81731D7A9DEF3BA10C5B60FC6DF5A698" colspan="7" rowspan="1">
As % of
<abbrev id="ABBRID0ERAAE" xlink_title="Total lengths">TL</abbrev>
</td>
</tr>
<tr id="4EC5D35EDF386A9B63E01A24A8CB7A26">
<td id="F0795C0581DE0AE7458D3441B765C20A" colspan="1" rowspan="1">
Head length (
<abbrev id="ABBRID0EZAAE" xlink_title="head lengths">HL</abbrev>
)
</td>
<td id="64E43708F2BD361AA0C431532C43A7D5" colspan="1" rowspan="1">10.3</td>
<td id="CF3296E5827A13ECCC393F76E5C7EFDD" colspan="1" rowspan="1">8.1</td>
<td id="EDBFCCE3C8F11173D1C4C1BAAFC97379" colspan="1" rowspan="1">8.0</td>
<td id="C2EEC02ECE50E3945D690DD1B0A36718" colspan="1" rowspan="1">8.2</td>
<td id="15C6035FAFCB9D4EDA8114FA0A34C0F4" colspan="1" rowspan="1">8.8</td>
<td id="ACC0A793153D3869A80CD62E92220900" colspan="1" rowspan="1">8.5</td>
</tr>
<tr id="54F50E81B4BAF3A310C50A97DE6ACC0F">
<td id="8D52EF48BBED07EDBF9E8444B1013F7D" colspan="1" rowspan="1">Preanal length</td>
<td id="8383E16A6485E0AB245A774773BA77BE" colspan="1" rowspan="1">37.3</td>
<td id="0EC5FFCC55DFF9B5787D6D3693BA98A5" colspan="1" rowspan="1">41.2</td>
<td id="044A98EF54FC2F4668BD2C7691BEC210" colspan="1" rowspan="1">41.5</td>
<td id="641F84E20E509BA3527CB6080FF627E8" colspan="1" rowspan="1">43.2</td>
<td id="B6EC0A80D1CB4278E3A1100129D03D5F" colspan="1" rowspan="1">43.2</td>
<td id="AD2116317FA0D22531871AF6941E4642" colspan="1" rowspan="1">42.9</td>
</tr>
<tr id="CF26CEBD7CEF1EAF15C1F082AD331E6C">
<td id="6096FD41FF9C9C63182C58757843E3DC" colspan="1" rowspan="1">Tail length</td>
<td id="41589AE7EF43C69DAF20C1011F5D6BD7" colspan="1" rowspan="1">62.7</td>
<td id="76FDB229AD24D5388B7973723B58B24D" colspan="1" rowspan="1">58.8</td>
<td id="5F3BB24434953E32D3AF1D94E90675B8" colspan="1" rowspan="1">58.5</td>
<td id="BE1E24209241CE63C68E86D9D1B6D0BC" colspan="1" rowspan="1">56.8</td>
<td id="D344DD638E8FF08943E0539FD87F86DB" colspan="1" rowspan="1">56.8</td>
<td id="CE664E8F4105933E029C296CAFEC8B55" colspan="1" rowspan="1">57.1</td>
</tr>
<tr id="281B7447DA0C5AE47A30775C737FDB27">
<td id="1064F935C1B54E9BAEC11AA6040DE720" colspan="1" rowspan="1">Predorsal length</td>
<td id="15C39CE573BD1BCA19E0A43FAB5FFC19" colspan="1" rowspan="1">12.6</td>
<td id="9111666549DADD0525FE39E0D85A43DF" colspan="1" rowspan="1">16.5</td>
<td id="F5AC463BB3026EFB53BE42C6A2B758B6" colspan="1" rowspan="1">15.1</td>
<td id="50BDD08C98689E7E5C064F44E1463712" colspan="1" rowspan="1">16.1</td>
<td id="3D2B4CE01B781BA1569A54B8366993B2" colspan="1" rowspan="1">17.4</td>
<td id="71B1A52B70C6C1252632D4AA1900D56E" colspan="1" rowspan="1">18.0</td>
</tr>
<tr id="3F204D5D2114171FF29DDB51B4BEDC3F">
<td id="0276415E405AEFC3093D07FA26B38915" colspan="1" rowspan="1">Body depth at gill opening</td>
<td id="987584AF8A151A2733144F4649EC3802" colspan="1" rowspan="1">3.1</td>
<td id="7039B469533E053B74A04675788AB398" colspan="1" rowspan="1">2.2</td>
<td id="60C8E2DDD14AEFC277E11ACC491D91B6" colspan="1" rowspan="1">2.7</td>
<td id="B0141907268CB01FC2A8057D3D7423C5" colspan="1" rowspan="1">2.4</td>
<td id="AD6A71AAE2FC6746D101DAAA68804AC3" colspan="1" rowspan="1">2.7</td>
<td id="48B4371C1DA38BF31BE69AC58B48D328" colspan="1" rowspan="1">3.3</td>
</tr>
<tr id="EA65D885BAC8429D5C372BB22559F1CB">
<td id="E8C5ABEF19AC759FF87FFD95D9A975F2" colspan="1" rowspan="1">Body width at gill opening</td>
<td id="3B09B68EF32296DA79F459D99A83A8BC" colspan="1" rowspan="1">2.0</td>
<td id="267CE819E5B622A91AB30226A984945A" colspan="1" rowspan="1">1.6</td>
<td id="A3012365A2377C1A1D21BCCE98663C03" colspan="1" rowspan="1">2.3</td>
<td id="9A3FC75767E140487980DB7CB50C74F2" colspan="1" rowspan="1">1.7</td>
<td id="01718B9D5A915DF37296B9C84143DA80" colspan="1" rowspan="1">1.9</td>
<td id="FCA872D8BCAEC8EC0B7D41695152E86A" colspan="1" rowspan="1">2.7</td>
</tr>
<tr id="EB6F1CA424300D88A76FB731EF2EC854">
<td id="BE0B229E89A75CACB43DC0E916992EF5" colspan="1" rowspan="1">Body depth at mid-anus</td>
<td id="05B60FDD353BEBF04526CC2813BB6128" colspan="1" rowspan="1">2.5</td>
<td id="43FE8F2280F3F4389F287A1B1F9D333B" colspan="1" rowspan="1">2.1</td>
<td id="25A0CA93D1640A6A43E42CF998CC349B" colspan="1" rowspan="1">2.9</td>
<td id="B162E0A69EFAD380E2521B0FF11C8E11" colspan="1" rowspan="1">2.3</td>
<td id="073323AC562D1601F7C950DCDFEBF0F6" colspan="1" rowspan="1">3.0</td>
<td id="454719C90012C74EBE3400B641DF4128" colspan="1" rowspan="1">2.9</td>
</tr>
<tr id="44C8DCFF83DC79E7AF11B4DD685470F1">
<td id="8BA4F5C37580437BAE3B7D0A93F8C90C" colspan="1" rowspan="1">Body width at mid-anus</td>
<td id="3A5C88092DA6C97A2973F23B512B0BBA" colspan="1" rowspan="1">2.4</td>
<td id="4C7108764F452B156071802443F00D89" colspan="1" rowspan="1">2.1</td>
<td id="E437ACF17A1BCE0838BC9AF98EDDB262" colspan="1" rowspan="1">2.8</td>
<td id="6CC9F473507695BC05344C77A7D833C9" colspan="1" rowspan="1">2.4</td>
<td id="3FCAA2CC5E3592DCF75D152EFE6A98D5" colspan="1" rowspan="1">2.6</td>
<td id="7A82A7BCE690F8B120514659CABFCE47" colspan="1" rowspan="1">3.0</td>
</tr>
<tr id="1BA4B076706E25849005DAB133FA9032">
<td id="F51FDEF7EBE9D8735DF1BFE09A4E4EE4" colspan="7" rowspan="1">
As % of
<abbrev id="ABBRID0EOGAE" xlink_title="head lengths">HL</abbrev>
</td>
</tr>
<tr id="70E29765F6FCDCE69FCFE17E1F327EDA">
<td id="B765B74AD62A0348F1D33C381710DE6E" colspan="1" rowspan="1">Snout length</td>
<td id="30D23D3A3717A0DC14C50F526753DC02" colspan="1" rowspan="1">16.4</td>
<td id="34D7D739EFB2D05483C2436581472C40" colspan="1" rowspan="1">20.6</td>
<td id="FF06D0446A194A2ED547FEBC4A81405A" colspan="1" rowspan="1">20.1</td>
<td id="8863184152ED69F541422769A32C94EC" colspan="1" rowspan="1">21.2</td>
<td id="A4FF707FB7A69EF2E788776A244125B5" colspan="1" rowspan="1">18.8</td>
<td id="5B4CD837FA2C2D3B4EFA6D22D800E974" colspan="1" rowspan="1">20.9</td>
</tr>
<tr id="2F6D9A18D1F7373B8CA24F4E0AD076D6">
<td id="E7E0DB58BAE25EC60F04E0C9C6CE567F" colspan="1" rowspan="1">Eye diameter</td>
<td id="DB37BD97C6EAD2C8AEDAC61DE0075D31" colspan="1" rowspan="1">6.8</td>
<td id="96187B0FC2CA919F3EAD5F4437EAF201" colspan="1" rowspan="1">8.6</td>
<td id="665E9F1A5A0C42302F86C84A75F015A6" colspan="1" rowspan="1">9.2</td>
<td id="3BB506337189EF1B8660BDF014BFAD25" colspan="1" rowspan="1">8.3</td>
<td id="C1012701BA0E6A4E6FF46FA510E9B05F" colspan="1" rowspan="1">8.4</td>
<td id="43A2AA35B66A8AD896A8B5FFF19E13C7" colspan="1" rowspan="1">9.1</td>
</tr>
<tr id="50C31006797FA35E1EC4EE7FCB0611A8">
<td id="31F845DAB0C4ACD5F0B41A65A87E5FDF" colspan="1" rowspan="1">Upper-jaw length</td>
<td id="A47EDEA70F2BCD2F4AF10A9D328F7F05" colspan="1" rowspan="1">27.9</td>
<td id="813FCFABA93823C1D9202A21FE28FE64" colspan="1" rowspan="1">28.6</td>
<td id="7E7AC3295C814070CF04A9D29FDAA981" colspan="1" rowspan="1">28.1</td>
<td id="FE9DE1DB49029ECE8A6F474F28982BA7" colspan="1" rowspan="1">29.5</td>
<td id="3C17C9D2E39A02913436DA4A2B2375E2" colspan="1" rowspan="1">29.8</td>
<td id="C2895137894C557C9286011E147C900A" colspan="1" rowspan="1">28.3</td>
</tr>
<tr id="54E63E478227C7D5324D2E68D835CB9C">
<td id="D839AA20C44183BD50A18160B357C94F" colspan="1" rowspan="1">Gill-opening length</td>
<td id="F4F90A312861AAA4271D370BB240DC55" colspan="1" rowspan="1">9.2</td>
<td id="E07AC597177C3C05030B1D775F65624B" colspan="1" rowspan="1">11.1</td>
<td id="74EBEC247D2CADC957446754A44322B0" colspan="1" rowspan="1">8.2</td>
<td id="B0E9633A8C8FCF7DEECF6A5534F3A24E" colspan="1" rowspan="1">9.4</td>
<td id="0DCCC8BBB0EF901343BAD87DD50DD097" colspan="1" rowspan="1">9.7</td>
<td id="5FA9FD3276C813098072942D748F5E5C" colspan="1" rowspan="1">11.3</td>
</tr>
<tr id="5212133A1D3B55B288E0ECF7782BD278">
<td id="5A91109AD8DD98B4523E391BDDA8F450" colspan="1" rowspan="1">Interorbital width</td>
<td id="427B7ED408792576763563D58E4F1824" colspan="1" rowspan="1">9.9</td>
<td id="8BDDD015D3EC9C0A2F8E9D54F03DA33F" colspan="1" rowspan="1">10.0</td>
<td id="B98CB15E5B1EE9880A03D68649B4F098" colspan="1" rowspan="1">10.2</td>
<td id="75402C28C572CC3B52DABDA019D9477D" colspan="1" rowspan="1">12.7</td>
<td id="C04407C9A9472C5F143BEA6861FFF967" colspan="1" rowspan="1">12.9</td>
<td id="6497876E8890F2B615E1800B3015CB8B" colspan="1" rowspan="1">16.5</td>
</tr>
<tr id="1C9094ADFC276CBCC547D22A7FB8D485">
<td id="12FFF322218AB1C5C1AD4191635929C2" colspan="1" rowspan="1">Isthmus width</td>
<td id="4480078C4D7D8E016B658B90158A4E8B" colspan="1" rowspan="1">15.0</td>
<td id="C54C878D1090380451463DB0D77BC4EE" colspan="1" rowspan="1">16.0</td>
<td id="F4C65FBDB1F8CE4FB327CAD829B66652" colspan="1" rowspan="1">22.8</td>
<td id="5DD95D319B5D4562E4710960AB8C2138" colspan="1" rowspan="1">13.8</td>
<td id="E6739EE1D659D95C792C4176A0FCD2ED" colspan="1" rowspan="1">16.9</td>
<td id="3D6477044A2F6FEA22A8EC390E405D29" colspan="1" rowspan="1">28.2</td>
</tr>
<tr id="FC8DB05CBAB0847D3A2245D6288DD719">
<td id="E7EF7EEC8A21AA51CA79D441B1E19B9C" colspan="1" rowspan="1">Pectoral-fin length</td>
<td id="EAFA776E25CC4451439B4CF850C9DC1D" colspan="1" rowspan="1">27.9</td>
<td id="EFE62C6B222393F4FA95B4CE92315409" colspan="1" rowspan="1">26.0</td>
<td id="BC4D1CF32EF1ADC9D2DBB08E2E1120E1" colspan="1" rowspan="1">24.2</td>
<td id="BCC2E25CF11283883945234A5B452687" colspan="1" rowspan="1">29.8</td>
<td id="54CCC3C8A6E9541F8493E5C806E4321C" colspan="1" rowspan="1">29.5</td>
<td id="CE2BD9B3AE2BCAE8BF46F6E648748BE2" colspan="1" rowspan="1">24.6</td>
</tr>
<tr id="426E8CCFF7B7B0B5E2A3703BA389C472">
<td id="1297D6B6061D82DFC207B44FE84AB35E" colspan="1" rowspan="1">Pectoral-fin base</td>
<td id="C0EAC9BA3AFFCDFF9E48A97E0768651B" colspan="1" rowspan="1">12.0</td>
<td id="F23EBE47ACD331E42FD0C77A6FDEEBF1" colspan="1" rowspan="1">11.7</td>
<td id="4288637C700DEBAE9D8178C928929D91" colspan="1" rowspan="1">9.7</td>
<td id="F8E5687056FFCE0947F4FDBDEBDADC65" colspan="1" rowspan="1">9.6</td>
<td id="6F8B710CEE5391C9DAEFE4156BE37AB1" colspan="1" rowspan="1">10.8</td>
<td id="0EFB6F29754955674C7F6638810B624D" colspan="1" rowspan="1">10.1</td>
</tr>
<tr id="9F3739E469B83756C543AC26A843E62B">
<td id="CA76BD56D6EFEF69D16F6C4DD4AAA9FA" colspan="7" rowspan="1">Counts</td>
</tr>
<tr id="3EDAB83C5410A7E3C14C5A6722B1ED77">
<td id="431692F73DAE556F443AD4C6D1FAE0F7" colspan="1" rowspan="1">Predorsal vertebrae</td>
<td id="6A318E15AE1763EE5DD212E9777A87AB" colspan="1" rowspan="1">13</td>
<td id="C28FD101BB10282099C1471FD8C56C18" colspan="1" rowspan="1">24</td>
<td id="0C84EB9EB02EB56A060B1A708085891E" colspan="1" rowspan="1">23</td>
<td id="9B8E55C9446510B7798E9D0C5C5EA546" colspan="1" rowspan="1">23</td>
<td id="F9C829CDA48CC814741C9A0F4B706BA1" colspan="1" rowspan="1">25</td>
<td id="76B60D31CC0A7E66F7C0B612F730F3FA" colspan="1" rowspan="1">26</td>
</tr>
<tr id="7376ED7223C57B30CE5CF0DDF2FCDF23">
<td id="32FC39D1B5EE763591A786DA1CD9C27B" colspan="1" rowspan="1">Preanal vertebrae</td>
<td id="C3781E2368C764B24CCDFA72A0E91895" colspan="1" rowspan="1">55</td>
<td id="3900AE13B7B9A8906A2870D18B94C07A" colspan="1" rowspan="1">62</td>
<td id="C258A59DAC67996B9221117E60E8F639" colspan="1" rowspan="1">64</td>
<td id="C83B63BF2EE3171BB57505C84508EA8D" colspan="1" rowspan="1">66</td>
<td id="56AE6B444F541517B46588E1D4BE330F" colspan="1" rowspan="1">65</td>
<td id="A19C521EFD4A0F035C3F0FBA4DB9BC7A" colspan="1" rowspan="1">65</td>
</tr>
<tr id="8CF083556D008FF5865445EF555C8A9B">
<td id="B7191C3960C9970C51EF50621CDA395A" colspan="1" rowspan="1">Total vertebrae</td>
<td id="CFEF591E0138A5A49536001D8D48D502" colspan="1" rowspan="1">153</td>
<td id="7D8943130422F6A8B959B8043046AE26" colspan="1" rowspan="1">163</td>
<td id="BBC447C2BFDCEC27EC48AE24651DBEA4" colspan="1" rowspan="1">162</td>
<td id="9ABC6CD2A92DFE9FB5C76E4E0BA505A4" colspan="1" rowspan="1">163</td>
<td id="BFCAD9DFCC60F1AD56BD0A04F3817CE1" colspan="1" rowspan="1">164</td>
<td id="0CE42300C3BFE1226CD2DB525FC13CDA" colspan="1" rowspan="1">162</td>
</tr>
</tbody>
</table>
</paragraph>
<caption id="3CC3B79BF31298505EAC7889535A776B" ID-Table-UUID="3CC3B79BF31298505EAC7889535A776B" httpUri="http://table.plazi.org/id/3CC3B79BF31298505EAC7889535A776B" startId="T2" targetIsTable="true">
<paragraph id="702717ADB52270470225A53785C9D5D6">
<label id="C1E3F6B96B90355F8913085388F32AF9">Table 2.</label>
</paragraph>
<paragraph id="D6D514FDAB1662277B39250C2276323B">
Selected characters of
<taxonomicName id="22C45DF76B7944E06757F1BA67AAE73A">
<emphasis id="B2F9C732CA93CA2B2422065F823C757F" italics="true">Ophichthus</emphasis>
species
</taxonomicName>
reported from Taiwan, except patterned species.
</paragraph>
</caption>
<paragraph id="CE0A783A9D515419CD3657A281B1FDA5">
<table id="TID0EWOBG" rules="all">
<tbody id="489AFF8D27236D73AB68EB9163470C51">
<tr id="6573F72FB4B1C5E6D20EF28A2D105743">
<th id="37C047996C9373F6BB84C2CC150EBA46" colspan="1" isEmpty="true" rowspan="1">-</th>
<th id="FC94844CBABD2403049AD8BBDA378533" colspan="1" rowspan="1">
<abbrev id="ABBRID0EDPAE" xlink_title="supraorbital pores">SO</abbrev>
</th>
<th id="8F66B4F233C224E657A29F74427705D9" colspan="1" rowspan="1">
<abbrev id="ABBRID0ELPAE" xlink_title="preoperculomandibular pores">POM</abbrev>
</th>
<th id="2DEF2353DE9F4E9145B641719213BDF4" colspan="1" rowspan="1">Protrusion number</th>
<th id="ED5D1BBDA44C3A2C90A734320E14661B" colspan="1" rowspan="1">PALL</th>
<th id="D03E9093870FD349A36CFB841E49060D" colspan="1" rowspan="1">PDV</th>
<th id="990A4D95535F6DFDA7A5B8A1A4D20184" colspan="1" rowspan="1">PAV</th>
<th id="D69FF78919BF7DF60849D05F924E0888" colspan="1" rowspan="1">TV</th>
<th id="C4FD492A3C8917F5B7AC92903E5F27ED" colspan="1" rowspan="1">TYPE V</th>
<th id="961CD6E59BBA2B373DBFAB1CA7CA1AD4" colspan="1" rowspan="1">Protrusion shape</th>
</tr>
<tr id="61A4F6F0F61C21378715DD90E1B7662D">
<td id="A898BD6C2A19370F8A553C2EF2816CB7" colspan="1" rowspan="1">
<taxonomicName id="9901BD8448BCF664CA962BE35E69CFFC" authorityName="McCosker &amp; Chen" authorityYear="2000" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="aphotistos">
<emphasis id="330AD5314E08D043E97DDDCD7BB100B9" italics="true">O. aphotistos</emphasis>
</taxonomicName>
</td>
<td id="EF8D32F6AFDE94C32EBEFAFFE94FB673" colspan="1" rowspan="1">1 + 4</td>
<td id="76439C4E4A5722420FB1E9CCEC1B7053" colspan="1" rowspan="1">6 + 2</td>
<td id="C13E232F5D6C379DAB07915DC8B59303" colspan="1" rowspan="1">0</td>
<td id="520974F3D94E9688580A48E7788EDD88" colspan="1" rowspan="1">59–61</td>
<td id="517E3E0E669DF378A1767AA5213E70F4" colspan="1" rowspan="1">16–19</td>
<td id="83DCC335DCE64A55E3CAC7D5239A0BE5" colspan="1" rowspan="1">57–61</td>
<td id="0BCB7DDD3A2628C9EFF252DD228573A5" colspan="1" rowspan="1">157–162</td>
<td id="5CB1912E44EE7E6F3AFCD69C30C694A4" colspan="1" rowspan="1">18 / 59 / 161</td>
<td id="1956C57E8567D8CB1C2CAEBFA6DEB3C4" colspan="1" rowspan="1">Absent</td>
</tr>
<tr id="6A9F4272AE7F74517B171F4FBE2E6F8F">
<td id="7BD274393A9EA9B1AACC3ECDC1F541A6" colspan="1" rowspan="1">
<taxonomicName id="9E9EC160241082E6FC07670B25FC6AE2" baseAuthorityName="Anonymous" baseAuthorityYear="1830" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="apicalis">
<emphasis id="D3EB84862911971C30B538AEBC7E8FA2" italics="true">O. apicalis</emphasis>
</taxonomicName>
</td>
<td id="580ECA8F473438946D145DFE515F26EC" colspan="1" rowspan="1">1 + 4</td>
<td id="5FD25959141712A29594DE95C18161D0" colspan="1" rowspan="1">5 or 6 + 3</td>
<td id="B265C2F2E6CDC584ED5E7A1CEFED723C" colspan="1" rowspan="1">2</td>
<td id="C872945E330AA6BF4142489AF7E97B9C" colspan="1" rowspan="1">NO DATA</td>
<td id="8E0FE6516CA7A1924F60DC587F3E919C" colspan="1" rowspan="1">12–14</td>
<td id="19FD9BCA19B6592F25342E1D1BE370C2" colspan="1" rowspan="1">50–53</td>
<td id="60934396ED420FE7BE5E30D70C7C5060" colspan="1" rowspan="1">138–141</td>
<td id="3E7B5A688A40F014EC5EA4F0D118FFE9" colspan="1" rowspan="1">NO TYPE</td>
<td id="27F28CB6B717B155A8BA068C8FC6BE50" colspan="1" rowspan="1">Small, thorn-shape</td>
</tr>
<tr id="76BED01C96C00AE54E0D1D53BFDCE8DB">
<td id="84A5F6BB5DC705AD0AEA008BAC460D38" colspan="1" rowspan="1">
<taxonomicName id="A460444A40D2B6C5A41C6598C331C5A3" authorityName="Jordan &amp; Snyder" authorityYear="1901" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="asakusae">
<emphasis id="C3AF23F0BC4944A8DB16B116E6E1AE52" italics="true">O. asakusae</emphasis>
</taxonomicName>
</td>
<td id="06621ACA81A35CB69E76BAFAF0DFC8BE" colspan="1" rowspan="1">1 + 4</td>
<td id="28DDF9940AE8B4E2F35794989C15882C" colspan="1" rowspan="1">7-10 + 3</td>
<td id="BCDBA324DE4FCBB5E8C01A35CD03504B" colspan="1" rowspan="1">0 or 1</td>
<td id="8C25CC9C85D9872D567E5812C6309B99" colspan="1" rowspan="1">51–58</td>
<td id="0151E8AE8B157C0AF8F308BF0AA47707" colspan="1" rowspan="1">10–12</td>
<td id="42B936243F3E440A2050F6BAE74BE6E2" colspan="1" rowspan="1">49–57</td>
<td id="81BFFB947C6141A8D5E87E4384849462" colspan="1" rowspan="1">123–132</td>
<td id="34CDA974F4CE79EC18BD7EC0D7952EB4" colspan="1" rowspan="1">11 / 54 / 128</td>
<td id="83F6842BE143F47E292A20DB668D6F7D" colspan="1" rowspan="1">Robust hump shape, weak in smaller specimens</td>
</tr>
<tr id="815C2BD4BFF20081C29F23B267A67ACE">
<td id="7A86EE0CC5A43F4DB0B9C2A29FA687D3" colspan="1" rowspan="1">
<taxonomicName id="043D28D2B97E3772F0554AD5E3362740" authorityName="McCosker &amp; Ho" authorityYear="2015" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="bicolor">
<emphasis id="91DCA0D502887057AC131E34F76F9C6B" italics="true">O. bicolor</emphasis>
</taxonomicName>
</td>
<td id="0828E529D7D29C9725ACAAD9A7742195" colspan="1" rowspan="1">1 + 4</td>
<td id="AF4234F538E052433C6B5DBC92C55A17" colspan="1" rowspan="1">6 + 2</td>
<td id="4EEC3C244480F11E7B3CA386BA284C6C" colspan="1" rowspan="1">0</td>
<td id="9065B2467BC68E8F3BF17DA37FB8628B" colspan="1" rowspan="1">63–67</td>
<td id="3118F4B657DB4BC04DFF7DC12BED1FCD" colspan="1" rowspan="1">15–23</td>
<td id="32F54BBA3BF7E3C580E876F3A7E48647" colspan="1" rowspan="1">61–66</td>
<td id="912C4BA05680063F0A326470803C0E9F" colspan="1" rowspan="1">155–163</td>
<td id="7137DA889992D8A254EEE663DA07887B" colspan="1" rowspan="1">19 / 65 / 160</td>
<td id="657FD31B8EE8C5F116EDE377A968D2BC" colspan="1" rowspan="1">Absent</td>
</tr>
<tr id="C9CC30F380772AEBBCD2C8869AEA11D8">
<td id="16F66A3042AAF659DAD97561C0E76C54" colspan="1" rowspan="1">
<taxonomicName id="6B87F87702EBF72D6B1ACF9CDEF3B398" authorityName="Hibino &amp; Ho" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="kbalanensis" status="sp. nov.">
<emphasis id="0DC38ED510791F3528E73889C16EE3EF" italics="true">O. kbalanensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="02E0B771935F6C92EB59E939DB2C893E" rank="species">sp. nov.</taxonomicNameLabel>
</td>
<td id="2C119FA7B97B424C7583813C205F66E0" colspan="1" rowspan="1">1 + 4</td>
<td id="CA274301047F02A63AFBE4F4242E8800" colspan="1" rowspan="1">6 or 7 + 3</td>
<td id="C8E49D5C49CE433BA08E64B98F5D5193" colspan="1" rowspan="1">2</td>
<td id="5FDB56C6807CF5082B0184FD267E47E4" colspan="1" rowspan="1">53</td>
<td id="650FA7F48C38D0EF1E5F75EBE3196413" colspan="1" rowspan="1">12</td>
<td id="4EEFE6797321754D343C65ED418BD72B" colspan="1" rowspan="1">55</td>
<td id="6B00CD0A52F603D5E7ED5D88EE2072DE" colspan="1" rowspan="1">153</td>
<td id="290B3E90C2D129A9BADF6652AD142AE6" colspan="1" rowspan="1">12 / 55 / 153</td>
<td id="D243BEB9FDB4EF7D80B860619CA8074E" colspan="1" rowspan="1">Simple thorn-shape</td>
</tr>
<tr id="BBF73C949D496B7BA284DDC386A1F31D">
<td id="0D909D5BA4396B77AB70BEAA1974E7A8" colspan="1" rowspan="1">
<taxonomicName id="DD7A95E7587243989357CF0208F6DF83" authorityName="Hibino, McCosker &amp; Tashiro" authorityYear="2019" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="kusanagi">
<emphasis id="DFFA92FBE5A975CEAED655C50C544381" italics="true">O. kusanagi</emphasis>
</taxonomicName>
</td>
<td id="6816D00E4BD6756B0DF6C9562DF4CF84" colspan="1" rowspan="1">1 + 4</td>
<td id="200418873E1B61004AC05D963FB2123C" colspan="1" rowspan="1">6 + 2</td>
<td id="2F74B32D96422F06F51A6225E0851081" colspan="1" rowspan="1">0</td>
<td id="9AD6670A7E6EA8D02B0FBAC457BDB9DA" colspan="1" rowspan="1">61–65</td>
<td id="7A5F0C01B7FA7EDF0E1E774CAEEB4CCA" colspan="1" rowspan="1">17–22</td>
<td id="F9244914B35618AB83F50A1E3B0284F9" colspan="1" rowspan="1">59–62</td>
<td id="EB74BD1ECA0F6311B599F252E6C67828" colspan="1" rowspan="1">158–163</td>
<td id="B82649D4E5A059C306FF5E4F9D0432FA" colspan="1" rowspan="1">18 / 61 / 161</td>
<td id="4F368184173B7C82513E030B56A2992E" colspan="1" rowspan="1">Absent</td>
</tr>
<tr id="CFA7302FCD607078BAA6D3DC30BA14FF">
<td id="D81F0B97FDE39E3F64E03D1B1C666CB0" colspan="1" rowspan="1">
<taxonomicName id="9519E9519DE78D4BE1DF81294A3D2A2E" authorityName="McCosker, Ide &amp; Endo" authorityYear="2012" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="machidai">
<emphasis id="984440830EF6B9CA947AE831C6EE84AA" italics="true">O. machidai</emphasis>
</taxonomicName>
</td>
<td id="C043D120609C2F712E7ECA5C790AAC6F" colspan="1" rowspan="1">1 + 4</td>
<td id="587CEC1A4F2C21B15790178A998D9E4F" colspan="1" rowspan="1">5 or 6 + 2 or 3</td>
<td id="DEB65D1AEA52306B87F950182A5C3B58" colspan="1" rowspan="1">2</td>
<td id="A612ECF153E1280FD225C6A56C83C4BC" colspan="1" rowspan="1">51–59</td>
<td id="84FCD44242C0E6E902E622C8781CB336" colspan="1" rowspan="1">11–16</td>
<td id="F79171F0D9A130F3930B0AFBAC84173B" colspan="1" rowspan="1">52–59</td>
<td id="5C2D738CEDC82E819C4F84FCFDDC7CED" colspan="1" rowspan="1">150–161</td>
<td id="9ACF69B69E8BB01674476CEB60AD8E1D" colspan="1" rowspan="1">16 / 58 / 158</td>
<td id="87ABE525A0BC332E911E80834152F450" colspan="1" rowspan="1">Simple thorn-shape</td>
</tr>
<tr id="C92B199A24DEDD21EFB6B1B690D6B75E">
<td id="16A384619CCDD1EB0B8B21A13FD520E3" colspan="1" rowspan="1">
<taxonomicName id="1A5CB05A8C295F9097A33DAAF627E6D6" baseAuthorityName="Bleeker" baseAuthorityYear="1852" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="macrochir">
<emphasis id="49F4F7A593D7072BC3CE7B447385135B" italics="true">O. macrochir</emphasis>
</taxonomicName>
</td>
<td id="4F176D6063256E0BB17833411EC57ACA" colspan="1" rowspan="1">1 + 4</td>
<td id="24419CC534084BF71CBC5647B7AD59B2" colspan="1" rowspan="1">4-6 + 2</td>
<td id="EA09CB3F630A0D8E4B4C207ABC142D8C" colspan="1" rowspan="1">2</td>
<td id="759ED548CA2FD8F23095CC63A489D078" colspan="1" rowspan="1">68–73</td>
<td id="55468B7E4A2ED298DA45BAC222FC5811" colspan="1" rowspan="1">11–12</td>
<td id="C5AC719E89B6E2DA7ED468F08E38E392" colspan="1" rowspan="1">67–71</td>
<td id="C2D11DBA08A970E41DD499EFB3E7DC81" colspan="1" rowspan="1">207–221</td>
<td id="6F202B08BA9E9D2ABD0E1B56FFF52F5B" colspan="1" rowspan="1">11 / 70 / 221</td>
<td id="D0181A23FAA0D60C3856B82399A1F347" colspan="1" rowspan="1">Simple thorn-shape</td>
</tr>
<tr id="605323E6BAB048F055DF301155BB6216">
<td id="940BD005A00470E89825ACB77F10E336" colspan="1" rowspan="1">
<taxonomicName id="EB12577A74251E1F62FAC0C5BD4FF4D8" authorityName="Asano" authorityYear="1987" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="megalops">
<emphasis id="A77705303941D6CD368ED3847837DD86" italics="true">O. megalops</emphasis>
</taxonomicName>
</td>
<td id="21693C83F9CE93ABA4E8386D82D2AFDD" colspan="1" rowspan="1">1 + 4</td>
<td id="C4F9F0F4D2F3BABF7FCDD4ECF4773E05" colspan="1" rowspan="1">6 + 3</td>
<td id="8BEF794301D41AE9548C70A7BB50CA55" colspan="1" rowspan="1">0</td>
<td id="1B759468062145D8118E1131A35D5CF6" colspan="1" rowspan="1">59–64</td>
<td id="CE6E1D5F571790923EAF1FB71CAE8884" colspan="1" rowspan="1">28–35</td>
<td id="69CE86F20152B55533D670F1C4FE039E" colspan="1" rowspan="1">59–63</td>
<td id="B8FE16CE346B0588359EED014D263893" colspan="1" rowspan="1">157–168</td>
<td id="C1195E7BC6C9B8E7112B2D2F39B51526" colspan="1" rowspan="1">29 / 60 / 160</td>
<td id="EBB2AAE5F68253FD987170FF9F22199B" colspan="1" rowspan="1">Absent</td>
</tr>
<tr id="6E45BFB4DF00E8B58358E954FFE6F4F3">
<td id="D65194E0C03FD115594569EDE793903E" colspan="1" rowspan="1">
<taxonomicName id="E7C6A000EDB2DF2C2715AF562F526ED6" authorityName="Hibino, Ho &amp; Huang" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="multidentis" status="sp. nov.">
<emphasis id="36774CDB416B15E159425737A2E21AF3" italics="true">O. multidentis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="473566066E19131CD4CA5B3AB8AD8CDA" rank="species">sp. nov.</taxonomicNameLabel>
</td>
<td id="3CC5D80DF79686AC18CE23206EADD141" colspan="1" rowspan="1">1 + 3</td>
<td id="16CB3A0D1E7A142DA549C0710D64FB34" colspan="1" rowspan="1">5 + 2</td>
<td id="92D60163DD695511E3463787CE517EE1" colspan="1" rowspan="1">0</td>
<td id="AAAB6DD6470A36E5C29765977F7FFBBF" colspan="1" rowspan="1">63</td>
<td id="114102E2CA009B1F08CCB9DC7A0DA2EF" colspan="1" rowspan="1">24</td>
<td id="35054383F9AE4C7DCF5C99A57224C457" colspan="1" rowspan="1">62</td>
<td id="22FB9B647057A7BEED7E962C8EA0F3DB" colspan="1" rowspan="1">163</td>
<td id="8E768F69D499ED040425D4E963180937" colspan="1" rowspan="1">24 / 62 / 163</td>
<td id="D69AFBC1960A03E8A62F8B70F12646CA" colspan="1" rowspan="1">Absent</td>
</tr>
<tr id="8A32BD8640B1070BBD1F82D22B337978">
<td id="3D9BF9DA92B69EBD47457D6CDD997C14" colspan="1" rowspan="1">
<taxonomicName id="E5F51B94D618113A0FD654F5197FEB80" authorityName="McCosker, Ide &amp; Endo" authorityYear="2012" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="obtusus">
<emphasis id="D4AB41D8BA4A48DE366558CEDECD8ADC" italics="true">O. obtusus</emphasis>
</taxonomicName>
</td>
<td id="EBB70D2B9F2AF4562BF1E92F661B75D7" colspan="1" rowspan="1">1 + 4</td>
<td id="02B16AFC35E5AF14314FF3C3277CB6BE" colspan="1" rowspan="1">4 or 5 + 3</td>
<td id="71AE0AF8DDE03CE95CE5B4EDF7C0412D" colspan="1" rowspan="1">2</td>
<td id="E3E8C37BA529FF87C2B0589941BCB871" colspan="1" rowspan="1">57</td>
<td id="5A913475568660CBC999FD6A2C3A10D0" colspan="1" rowspan="1">11–19</td>
<td id="A26511A07A158529991301C9C0A3B027" colspan="1" rowspan="1">52–57</td>
<td id="14BCDDEB6F7D1B131A9B6337678C2B5A" colspan="1" rowspan="1">148–159</td>
<td id="B83AD7CEE1E340585392F305D8385CC8" colspan="1" rowspan="1">12 / 57 / 151</td>
<td id="FACCAF516D4CC4A524515AEE1BAD9119" colspan="1" rowspan="1">Stout, with wrinkles in larger specimens</td>
</tr>
<tr id="D5F37823483EAD9E5C3FABA611B708BA">
<td id="12971A83E0C0F5874055789540DB8259" colspan="1" rowspan="1">
<taxonomicName id="F209D23AC157C00FE1345EEDCC43E7B1" authorityName="Ho, Ng &amp; Lin" authorityYear="2022" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="pratasensis">
<emphasis id="523147E3366AF7C23981C67325DB90DC" italics="true">O. pratasensis</emphasis>
</taxonomicName>
</td>
<td id="E138D5F2F21E63E1CF3CC8DE6FF8162D" colspan="1" rowspan="1">1 + 4</td>
<td id="AE5BB093BD4286897F5AF649A0628AAC" colspan="1" rowspan="1">6 + 2</td>
<td id="9CB4611EDECCA5F984B9D35AACF753C5" colspan="1" rowspan="1">0</td>
<td id="5276C3263A9BF9D4CFE7E65CC7D63F74" colspan="1" rowspan="1">59–60</td>
<td id="F5FBB62E9A29799DF11110657634DDD3" colspan="1" rowspan="1">20</td>
<td id="CCC5E91FA3560FE39C7C66A6CB56C0F3" colspan="1" rowspan="1">59</td>
<td id="AD12CC9F4E17ABDB4EA77DB942B4E75E" colspan="1" rowspan="1">177</td>
<td id="E3816757414B12E33AC5C3AA54ED2F7C" colspan="1" rowspan="1">20 / 59 / 177</td>
<td id="DB5EF55AEC7F3C48D4EFBDA19639C1CB" colspan="1" rowspan="1">Absent</td>
</tr>
<tr id="4340CFF60457ECBB5D72842C01D12FB4">
<td id="267CE0FE41C8AF2B97E3162182E7B549" colspan="1" rowspan="1">
<taxonomicName id="7B5295103EA8BB617F5AA3FB59DDBE2D" authorityName="Lee &amp; Asano" authorityYear="1997" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="rotundus">
<emphasis id="1ACBFA429EE44E9610E530AAF7DC22BF" italics="true">O. rotundus</emphasis>
</taxonomicName>
</td>
<td id="F4402CB616955CDC22B1BD1F41CBBC57" colspan="1" rowspan="1">1 + 3</td>
<td id="736A2832E9F3DC8C4E9A1A303BF219FA" colspan="1" rowspan="1">5 + 2</td>
<td id="76FD9DBF51CDB1DF3EE6CD5B2DF48AE9" colspan="1" rowspan="1">2</td>
<td id="68E820D88B5A8CB99B61D6A1DBFD2F84" colspan="1" rowspan="1">65–66</td>
<td id="453F92EC8EA604D0A471849D6EE32F27" colspan="1" rowspan="1">14?</td>
<td id="EBF6BCC587EA11FD55BEAF3AC93F3927" colspan="1" rowspan="1">64?</td>
<td id="E72550F047B4265633B5E408325A4B87" colspan="1" rowspan="1">178–184</td>
<td id="A1F905649424DF7881A20B9B057815DA" colspan="1" rowspan="1">14 / 64 / 182</td>
<td id="10348AE575CEDFBCB59ED4DCDD59D71A" colspan="1" rowspan="1">Short, simple thorn-shape</td>
</tr>
<tr id="E0108A1B598C46D5615DAE704DB31521">
<td id="902696731D676FA6184A8BB99DA5E013" colspan="1" rowspan="1">
<taxonomicName id="1E78318641096973D967F56B6934596B" baseAuthorityName="Ji &amp; Kim" baseAuthorityYear="2011" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="sangjuensis">
<emphasis id="EE4A198A99D73312EB23EF9F42E68855" italics="true">O. sangjuensis</emphasis>
</taxonomicName>
</td>
<td id="DBCED3F0A36B37DEEC28C2BEEFFFCE60" colspan="1" rowspan="1">1 + 4</td>
<td id="DFE21D86AF1641C553DDDD0CB6B8A8D8" colspan="1" rowspan="1">5 or 6 + 3</td>
<td id="F0B4570E1AB4A53FFC916B9CFFC716D8" colspan="1" rowspan="1">2</td>
<td id="4D00F61B715DB617E5D98CCEA7F64914" colspan="1" rowspan="1">53</td>
<td id="8840BA660383F643397B523206A324A5" colspan="1" rowspan="1">13–14</td>
<td id="0EB66426D1D71878BB411DAEDEC448B8" colspan="1" rowspan="1">48–52</td>
<td id="46D323374B536CE4699A56FF9520772B" colspan="1" rowspan="1">143–153</td>
<td id="1BF8A602288D623C8632B9F6D802CCD3" colspan="1" rowspan="1">13 / 50 / 153</td>
<td id="A703B3BEF85B6755F0E8EEEAF1CAFAC1" colspan="1" rowspan="1">Simple thorn-shape</td>
</tr>
<tr id="7104D5E6CA6A286513781C5AE9FAEB02">
<td id="8B23EDAAD46916FBC06EBA0FD3F7CA39" colspan="1" rowspan="1">
<taxonomicName id="A1DDDEB0FAA03B5A31B7DA87F73BCBC9" authorityName="Hibino &amp; Chiu" authorityYear="2019" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="semilunatus">
<emphasis id="4A1CA008773A2400527840829DD3D4DD" italics="true">O. semilunatus</emphasis>
</taxonomicName>
</td>
<td id="C9953DBCB082EF06389F6E34F0EABA8C" colspan="1" rowspan="1">1 + 3</td>
<td id="FAB6C4B91674AF38DF302754201E57CF" colspan="1" rowspan="1">7 + 2</td>
<td id="BFFA49B0853989A43C176F3C0EA77AF3" colspan="1" rowspan="1">0</td>
<td id="F60D4464C2D203A2D8DC7FD37D01421A" colspan="1" rowspan="1">65</td>
<td id="99757E0091B85F869E969734A317C2BD" colspan="1" rowspan="1">29</td>
<td id="EA350D4CCE21316BA0E5E6A07785BA66" colspan="1" rowspan="1">64</td>
<td id="F32C4BD83A5363BABBB4349579824584" colspan="1" rowspan="1">176</td>
<td id="33E31EDFCFAA00B081B9DC1BDA92C7D9" colspan="1" rowspan="1">29 / 64 / 176</td>
<td id="EB61E567DCF05730318BB48432F7AA75" colspan="1" rowspan="1">Absent</td>
</tr>
<tr id="24E234BBC45A07DC0B60F639E414669C">
<td id="4B7DB20495A8AF0291738C324BBE18E5" colspan="1" rowspan="1">
<taxonomicName id="870C70BD0BD493955478084BDF272B1E" authorityName="McCosker &amp; Ho" authorityYear="2015" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="shaoi">
<emphasis id="AF9AF9CC6B86D38674C546C597A36022" italics="true">O. shaoi</emphasis>
</taxonomicName>
</td>
<td id="683991F2BD844EF93EDA3E8052E52674" colspan="1" rowspan="1">1 + 4</td>
<td id="CA198CC28B74DC4D633E5665382E6C69" colspan="1" rowspan="1">6 or 7 + 3</td>
<td id="F841FE4C46822F74D39D2573CE2DDCA7" colspan="1" rowspan="1">1</td>
<td id="988C1D35267ADB04220BF5F6A4F1F272" colspan="1" rowspan="1">69–72</td>
<td id="7A1657CE01C64DAED687C2B402BDFBA6" colspan="1" rowspan="1">10–13</td>
<td id="7BE86B399075D4A969005C253AFA81F7" colspan="1" rowspan="1">68–72</td>
<td id="2A85AC68E7370E6BCDD4855076A6C6C1" colspan="1" rowspan="1">155–168</td>
<td id="9EF613933ED9930102240449E2987CB6" colspan="1" rowspan="1">12 / 68 / 155</td>
<td id="FC09205FA2E1B6E726E406769CA8D577" colspan="1" rowspan="1">Small, thorn-shape</td>
</tr>
<tr id="876B6A1A70EADE29770AA01CDF67D88F">
<td id="FD79027F3FA68D5380F2A1BBF01EC0DE" colspan="1" rowspan="1">
<taxonomicName id="B43A12608A269F22639867047AFB859E" baseAuthorityName="Temminck &amp; Schlegel" baseAuthorityYear="1846" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="urolophus">
<emphasis id="3DF92C599F4773C859132637CE55E061" italics="true">O. urolophus</emphasis>
</taxonomicName>
</td>
<td id="5BDE1ADBA11FDCEBB73C8881D8142EAC" colspan="1" rowspan="1">1 + 3 or 4</td>
<td id="B17BB4EB9FD71AE407716F87F34A8DC7" colspan="1" rowspan="1">5-8 + 3</td>
<td id="C0D415BA4D8D170385591CEF6702E585" colspan="1" rowspan="1">1</td>
<td id="3714B9F09D844AA931240583562A29C2" colspan="1" rowspan="1">51–58</td>
<td id="9796B0DF50E22137A2B3EE2FF97CC8B3" colspan="1" rowspan="1">13–18</td>
<td id="2FAD0C8E114499D40EC6CF30C31FF8E9" colspan="1" rowspan="1">51–56</td>
<td id="9D35FBD7F693BDDC8BF817ABCA9EEF29" colspan="1" rowspan="1">134–140</td>
<td id="348D6F1A5B1620725C6A5E73A830F7DA" colspan="1" rowspan="1">16 / 54 / 136</td>
<td id="72D527FBADCA3D16FD52503481ED02F4" colspan="1" rowspan="1">Robust hump shape, weak in smaller specimens</td>
</tr>
<tr id="2F19C042BC19026FA65FCC5DF87291A9">
<td id="D8DC62353E1C8FB7DB3D86A3FADE2BAD" colspan="1" rowspan="1">
<taxonomicName id="6BA82B4AF6CED72915F567535215DC3F" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="zophistius">
<emphasis id="98A5B961ECCB6176DA87EF7F2BA764E5" italics="true">O. zophistius</emphasis>
</taxonomicName>
</td>
<td id="BD2E27D1E8115C353E46FC6D006A8E56" colspan="1" rowspan="1">1 + 4</td>
<td id="B1D6DF172D2FAF70749B35267D8C8904" colspan="1" rowspan="1">5 or 6 + 3</td>
<td id="2FA5E786BE67C8A2667F8DDDBE28D978" colspan="1" rowspan="1">2</td>
<td id="BC514E1EA1DEC6DEECCEA3798D0C577A" colspan="1" rowspan="1">59–64</td>
<td id="0806870EFB468D83EF3F81FC8E23D1B3" colspan="1" rowspan="1">11–13</td>
<td id="32A4C6C5B71EF34FF22BF57EE069B559" colspan="1" rowspan="1">61–63</td>
<td id="179499376262E561CFBD50FA3237BF33" colspan="1" rowspan="1">177–184</td>
<td id="4FB82F12B6EF2BD131799C9F09E5F128" colspan="1" rowspan="1">12 / 62 / 181</td>
<td id="1EFFEA460D029391FF6C8EB7A79F87F4" colspan="1" rowspan="1">Simple thorn-shape</td>
</tr>
</tbody>
</table>
</paragraph>
<paragraph id="399E4A0CF42627ED73CBCF1034C257A4">
Body elongate, but relatively short, subcylindrical, its depth at gill opening
<quantity id="8622F122FC59F0CFD15158A13EC98AD9" metricMagnitude="-1" metricUnit="m" metricValue="3.0226" unit="in" value="11.9">11.9 in</quantity>
head and trunk,
<quantity id="38A3731E25AC771849039C4199B1FC12" metricMagnitude="-1" metricUnit="m" metricValue="8.0772" unit="in" value="31.8">31.8 in</quantity>
<abbrev id="ABBRID0ENIAG" xlink_title="Total lengths">TL</abbrev>
(Fig.
<figureCitation id="86F91FDAEA253344974C3A11D148DCC4" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Ophichthus kbalanensis sp. nov., NMMB-P 26381, holotype, 414 mm TL, Da-xi, Taiwan A fresh condition B preserved condition. Arrows indicate positions of dorsal-fin origin (lefts) and anus (rights)." figureDoi="10.3897/zookeys.1220.126337.figure1" httpUri="https://binary.pensoft.net/fig/1194398">1</figureCitation>
); tail more compressed, tapering slowly towards tip of tail, its length
<quantity id="A8D4FD9729B14ABC2C7FF4D62D876AF5" metricMagnitude="-2" metricUnit="m" metricValue="4.064" unit="in" value="1.6">1.6 in</quantity>
<abbrev id="ABBRID0EVIAG" xlink_title="Total lengths">TL</abbrev>
. Skin of body wrinkled; relatively strong wrinkles on snout, with numerous fine longitudinal wrinkles on remaining head and body.
</paragraph>
<caption id="EB87DB12E9482AD7FEA73599010C7ECB" ID-DOI="10.3897/zookeys.1220.126337.figure1" ID-arpha="1F2BBEAC-F128-59B5-8EC8-8B72C6E6DF4E" httpUri="https://binary.pensoft.net/fig/1194398" startId="F1">
<paragraph id="4A4B303D2F51A0362B964C7884DFFC2D">
<label id="EA8B01ACB178C06EDDCF40BFFFD9D82D">Figure 1.</label>
</paragraph>
<paragraph id="88A5EA95CD58079CF8A65F9A8927FB13">
<taxonomicName id="630351CCB500D988C17B3899FBEC91DD" authorityName="Hibino &amp; Ho" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="kbalanensis" status="sp. nov.">
<emphasis id="D582BAD039F3F4F86377C4FE8C802A8E" italics="true">Ophichthus kbalanensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="F5BB5A61918BADE8AC84584979E1CAF9" rank="species">sp. nov.</taxonomicNameLabel>
,
<specimenCode id="1B6C50263EDBC978B8823E90A69ABE4C">NMMB-P 26381</specimenCode>
, holotype, 414 mm
<abbrev id="ABBRID0EMJAG" xlink_title="Total lengths">TL</abbrev>
, Da-xi, Taiwan
<emphasis id="866704A0D31F748B9624C12CCE555D2C" bold="true">A</emphasis>
fresh condition
<emphasis id="989F7F50D368B93B06BA532A4BA8D0C3" bold="true">B</emphasis>
preserved condition. Arrows indicate positions of dorsal-fin origin (lefts) and anus (rights).
</paragraph>
</caption>
<paragraph id="2AD01B736024297F09D7AA9F9E421049">
Head moderate,
<quantity id="22D340C2CD599225D7AE132F41221616" metricMagnitude="-2" metricUnit="m" metricValue="9.144" unit="in" value="3.6">3.6 in</quantity>
head and trunk and
<quantity id="7ADAB3402D82B43D9236BC7B05B99553" metricMagnitude="-1" metricUnit="m" metricValue="2.4638" unit="in" value="9.7">9.7 in</quantity>
<abbrev id="ABBRID0E2JAG" xlink_title="Total lengths">TL</abbrev>
; dorsal contour of head weakly curved above eye, occipital weakly convex; branchial basket slightly swollen, its maximum depth
<quantity id="2D876A48B4E212513492214BBAF83E92" metricMagnitude="-2" metricUnit="m" metricValue="7.366" unit="in" value="2.9">2.9 in</quantity>
head. Snout tip relatively blunt and robust, moderate in length,
<quantity id="AC9A1690E72D08BBA63F58A76C5E51F0" metricMagnitude="-1" metricUnit="m" metricValue="1.5493999999999999" unit="in" value="6.1">6.1 in</quantity>
<abbrev id="ABBRID0E6JAG" xlink_title="head lengths">HL</abbrev>
and
<quantity id="D423479DD99773207876FB102A70B881" metricMagnitude="-2" metricUnit="m" metricValue="1.016" unit="in" value="0.4">0.4 in</quantity>
eye. Anterior nostril a simple tube opening anteroventrally; posterior nostril a hole at inner margin of upper lip, completely covered by a wide dermal flap. Eye relatively small,
<quantity id="BCDD92E3F1F5EA23CD87E3CF33693A07" metricMagnitude="-2" metricUnit="m" metricValue="6.096" unit="in" value="2.4">2.4 in</quantity>
snout length. Mouth subterminal, tip of lower jaw anterior to anterior base of anterior nostril tube. Rictus well behind posterior margin of eye. Lips smooth without small papillae; two low, small, simple, thorn-shaped protrusions, their tips pointed. Interorbital region smooth, transverse contour rounded, convex. Gill openings located ventrolaterally, upper ends slightly above middle of pectoral fin.
</paragraph>
<paragraph id="11A253839E6EB7CF30643D7CF716D472">
Sensory pores on head (Fig.
<figureCitation id="A018EA72471C219E974ED031F176C237" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Line drawings of O. kbalanensis sp. nov., NMMB-P 26381, holotype, 414 mm TL A sensory pores on head B teeth on upper (left) and lower jaws (right). Arrows indicate interorbital (left) and mid-temporal pores (right); black solid circle indicates holes of lost tooth; and broken lines indicate concealed teeth by fleshy lips." figureDoi="10.3897/zookeys.1220.126337.figure2" httpUri="https://binary.pensoft.net/fig/1194399">2 A</figureCitation>
) developed but very small, not obvious;
<abbrev id="ABBRID0EJKAG" xlink_title="supraorbital pores">SO</abbrev>
1 + 4, first one (ethmoid) on ventral surface of snout;
<abbrev id="ABBRID0ENKAG" xlink_title="infraorbital pores">IO</abbrev>
3 + 3, the first behind anterior nostril base, 2 below eye, and 3 arranged in a vertical row behind eye;
<abbrev id="ABBRID0ERKAG" xlink_title="preoperculomandibular pores">POM</abbrev>
6 (left) or 7 (right) + 3, the 6 (or seventh on right side) below rictus;
<abbrev id="ABBRID0EVKAG" xlink_title="supratemporal pores">ST</abbrev>
3, single pore on mid-temporal; and single interorbital pore. Lateral-line nearly complete, end anterior to about 1 / 2
<abbrev id="ABBRID0EZKAG" xlink_title="head lengths">HL</abbrev>
before tail tip; canal on branchial basket slightly arched, 10 on branchial basket before gill opening, 14 anterior to origin of dorsal fin, 53 anterior to anus, and total 138.
</paragraph>
<caption id="811B848D31E3ABDB27D8B99BC162CBD5" ID-DOI="10.3897/zookeys.1220.126337.figure2" ID-arpha="AD874F42-CC1D-56DF-896E-2774630D0DC6" httpUri="https://binary.pensoft.net/fig/1194399" startId="F2">
<paragraph id="44324B05CAD687754792D39EFB0C1A7C">
<label id="EC215D8F8C9818FCB778AC7E9CFDD239">Figure 2.</label>
</paragraph>
<paragraph id="32C502DE450273B8F86328CEB5FBD006">
Line drawings of
<taxonomicName id="174FEF90C49EB483DB11289D074A10A9" authorityName="Hibino &amp; Ho" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="kbalanensis" status="sp. nov.">
<emphasis id="56986E2CF87300FB6EA210972C0EFC25" italics="true">O. kbalanensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B851D531FCFF068B41E64D145E16382B" rank="species">sp. nov.</taxonomicNameLabel>
,
<specimenCode id="89F33A174150D5D53FECB8C2FFAA4B81">NMMB-P 26381</specimenCode>
, holotype, 414 mm
<abbrev id="ABBRID0EQLAG" xlink_title="Total lengths">TL</abbrev>
<emphasis id="93F0863F4952AA71884289D07E602F37" bold="true">A</emphasis>
sensory pores on head
<emphasis id="78AD35B4C4F85CF2285CF8FCF17B7062" bold="true">B</emphasis>
teeth on upper (left) and lower jaws (right). Arrows indicate interorbital (left) and mid-temporal pores (right); black solid circle indicates holes of lost tooth; and broken lines indicate concealed teeth by fleshy lips.
</paragraph>
</caption>
<paragraph id="1A81E992D20CFC29C45A489BCA580ED8">
Teeth moderate, conical, pointed (Fig.
<figureCitation id="C101D4D1E25201E5687D2AD1C19DF65D" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Line drawings of O. kbalanensis sp. nov., NMMB-P 26381, holotype, 414 mm TL A sensory pores on head B teeth on upper (left) and lower jaws (right). Arrows indicate interorbital (left) and mid-temporal pores (right); black solid circle indicates holes of lost tooth; and broken lines indicate concealed teeth by fleshy lips." figureDoi="10.3897/zookeys.1220.126337.figure2" httpUri="https://binary.pensoft.net/fig/1194399">2 B</figureCitation>
); teeth on maxilla mostly uniserial, ending with an additional row consisting of 5 teeth on inner side; dentary mostly uniserial but an additional row of small teeth in outer region of symphysis anteriorly; vomerine biserial anteriorly and medially, uniserial at posterior end; 6 large teeth on intermaxillary, arranged as chevron-shape.
</paragraph>
<paragraph id="D95B8DFAE789FEE85813A559D81D310F">Dorsal and anal fins low, anal fin slightly higher than dorsal fin; both ending slightly anterior to tail tip. Dorsal-fin origin over about posterior third of pectoral fin. Pectoral fin tip weakly pointed, not lanceolate (somewhat damaged by trawl operation). Caudal fin absent, rear end of tail tip pointed.</paragraph>
</subSubSection>
<subSubSection id="SECID0EEMAG" type="Coloration">
<paragraph id="A6E01CC1442D50DF35EC53A5D5754DE9">
<heading id="ACD408B1CB0DAEA0458C4A5F068A5A5E" reason="title">Coloration.</heading>
</paragraph>
<paragraph id="C8B0D323F82AE7CC28A40C1A7C882871">
Freshly caught specimen has a somewhat purplish body, darker dorsally and paler ventrally; pectoral fin dark brown and anal fin with dark brown to black margin; tail tip relatively pale (Fig.
<figureCitation id="EC6ECD0423FC553489DA6732F1487AB6" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Ophichthus kbalanensis sp. nov., NMMB-P 26381, holotype, 414 mm TL, Da-xi, Taiwan A fresh condition B preserved condition. Arrows indicate positions of dorsal-fin origin (lefts) and anus (rights)." figureDoi="10.3897/zookeys.1220.126337.figure1" httpUri="https://binary.pensoft.net/fig/1194398">1 A</figureCitation>
). Preserved condition in 50 % isopropanol ethanol (Fig.
<figureCitation id="5CD84BEC6C13A0B88AE0327058D8591B" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Ophichthus kbalanensis sp. nov., NMMB-P 26381, holotype, 414 mm TL, Da-xi, Taiwan A fresh condition B preserved condition. Arrows indicate positions of dorsal-fin origin (lefts) and anus (rights)." figureDoi="10.3897/zookeys.1220.126337.figure1" httpUri="https://binary.pensoft.net/fig/1194398">1 B</figureCitation>
): head and body dark brown, abdomen slightly paler, densely covered with melanophores; branchial basket darker and blueish. Anterior nostril tube similar to body color; sensory pores not prominently margined. Mouth cavity dusky brown. Membrane of gill opening paler than body. Anterior portion of dorsal fin creamy white, gradually becoming dark brown similar in color to dorsal surface; narrow black margin along entire dorsal fin. Anterior portion of anal fin creamy white, gradually becoming dark brown on anterior one-third of its length, with black margin becoming broader posteriorly. Pectoral fin yellowish brown, gradually darkening posteriorly.
</paragraph>
</subSubSection>
<subSubSection id="SECID0ESMAG" type="etymology">
<paragraph id="F7868E406D3830AA19E17069ADDAE8AA">
<heading id="DBD63F9075BD73E277BFCA08D947EBA2" reason="title">Etymology.</heading>
</paragraph>
<paragraph id="402D12578C86D94002E94E3EB84E38EB">
The specific name is derived from the type locality “Kbalan”, an old name of Yilan region (Kat-má-lán in Taiwanese or Cabaran in Spanish) dated back to 1300–800 years ago. Kbalan means “ people who live in the plain ” in the Taiwanese aboriginal race Kebalan. The earliest record of Kbalan occurred in the occupation of the Spanish (~ 1632) which was replaced by the Dutch East
<collectingCountry id="E3735DA8AEEEC76A824A426BD86CCEE4" name="India">India</collectingCountry>
Company in 1642.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EXMAG" type="comparisons">
<paragraph id="958042167132E5C49F371FB8AD576EE3">
<heading id="1A5F89709DB03152FB40663CE12F9420" reason="title">Comparison.</heading>
</paragraph>
<paragraph id="5980F601CDC759FF39C2A47B9E683106">
The first distinctive character found in
<taxonomicName id="3BB32935E51CE2DB2BF2EE2FBF80C4A7" authorityName="Hibino &amp; Ho" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="kbalanensis" status="sp. nov.">
<emphasis id="8B7553819C37FA6D6630791E9EADB38D" italics="true">Ophichthus kbalanensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="59742E4DB924B390936DA6DDC0B68E7C" rank="species">sp. nov.</taxonomicNameLabel>
is the unique tooth arrangement. In most species of
<taxonomicName id="A72D4F61472A9CCF562F7C826588AC99" authorityName="Ahl" authorityYear="1789" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="genus">
<emphasis id="09DE089855076FA0549189D106C90276" italics="true">Ophichthus</emphasis>
</taxonomicName>
we examined, the tooth rows on jaws maintained the same number or reduced to fewer row (s) posteriorly. However, in the new species, there is a short additional row of teeth on posterior portion of upper jaw.
</paragraph>
<paragraph id="D4F9BC7858A4D92D4DFB0283CF5D03EF">
Secondly, the tip of lower jaw extends beyond anterior margin of base of anterior nostril tube is also quite distinct among
<taxonomicName id="A2C8DC63AE08C9D6797550AC5E1E6E7D">
<emphasis id="91412B404FFE2635AE373C37F2441216" italics="true">Ophichthus</emphasis>
species
</taxonomicName>
(Hibino pers. obs.).
<taxonomicName id="924D7C7F37F7DFAFD651CAE40C9334D3" authority="McCosker, 2010" authorityName="McCosker" authorityYear="2010" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="ishiyamorum">
<emphasis id="6FF12D6086EBB625EDAED3F7815677D3" italics="true">Ophichthus ishiyamorum</emphasis>
<bibRefCitation id="E8B0FA58623A36E0E7CA1B76B8308893" DOI="10.11646/zootaxa.2505.1.1" author="McCosker JE" issue="1" journalOrPublisher="Zootaxa" pagination="1-39" refId="B12" refString="McCosker JE (2010) Deepwater Indo-Pacific species of the snake-eel genus Ophichthus (Anguilliformes: Ophichthidae), with the description of nine new species. Zootaxa 2505 (1): 1 - 39. https: // doi. org / 10.11646 / zootaxa. 2505.1. 1" title="Deepwater Indo-Pacific species of the snake-eel genus Ophichthus (Anguilliformes: Ophichthidae), with the description of nine new species." volume="2505" year="2010">McCosker, 2010</bibRefCitation>
</taxonomicName>
shares this character with
<taxonomicName id="685940B4C4ABD5DC0136377C84E32B85" authorityName="Hibino &amp; Ho" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="kbalanensis">
<emphasis id="DD74B9FE3761D745228D1BD4AC2544CC" italics="true">O. kbalanensis</emphasis>
</taxonomicName>
, as well as the dorsal-fin origin above middle of the pectoral fin, and similar body coloration. However,
<taxonomicName id="146B4C497DF71D0824375601C4DACF5D" authorityName="Hibino &amp; Ho" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="kbalanensis" status="sp. nov.">
<emphasis id="B01032F1A54A0F4C124976F3C10E097C" italics="true">O. kbalanensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="314A307F68978B8E191D9429F94CBB54" rank="species">sp. nov.</taxonomicNameLabel>
differs from
<taxonomicName id="A097C7A7792FDC2279BBB856A5731473" authorityName="McCosker" authorityYear="2010" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="ishiyamorum">
<emphasis id="BA3088FD772A547EE577EB00C79C7944" italics="true">O. ishiyamorum</emphasis>
</taxonomicName>
in having a smaller head (10.3 %
<abbrev id="ABBRID0EEPAG" xlink_title="Total lengths">TL</abbrev>
vs 14–15 %
<abbrev id="ABBRID0EIPAG" xlink_title="Total lengths">TL</abbrev>
), more vertebrae (153 vs 130–132), maxillary teeth mostly uniserial but ending in biserial (vs mostly uniserial and biserial anteriorly), and median fins with dark margined (vs pale) (
<bibRefCitation id="375C1AD8C356D291B88812210D710A9E" DOI="10.11646/zootaxa.2505.1.1" author="McCosker" firstAuthor="McCosker" issue="1" journalOrPublisher="Zootaxa" pagination="1-39" refId="B12" refString="McCosker JE (2010) Deepwater Indo-Pacific species of the snake-eel genus Ophichthus (Anguilliformes: Ophichthidae), with the description of nine new species. Zootaxa 2505 (1): 1–39. https://doi.org/10.11646/zootaxa.2505.1.1" title="Deepwater Indo-Pacific species of the snake-eel genus Ophichthus (Anguilliformes: Ophichthidae), with the description of nine new species." volume="2505" year="2010">McCosker 2010</bibRefCitation>
).
</paragraph>
<paragraph id="3648DEFD75C1B0D23ED2A4A475167169">
The tip of lower jaw is also before the anterior nostril tube in
<taxonomicName id="76777CD4802984CE0D00E2B5F97D3254" authority="McCosker, 2010" authorityName="McCosker" authorityYear="2010" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="alleni">
<emphasis id="C65A8F7F0A6DA6F4F4DE70524E263534" italics="true">Ophichthus alleni</emphasis>
<bibRefCitation id="25D35F53ED366763278E9442059B5683" DOI="10.11646/zootaxa.2505.1.1" author="McCosker JE" issue="1" journalOrPublisher="Zootaxa" pagination="1-39" refId="B12" refString="McCosker JE (2010) Deepwater Indo-Pacific species of the snake-eel genus Ophichthus (Anguilliformes: Ophichthidae), with the description of nine new species. Zootaxa 2505 (1): 1 - 39. https: // doi. org / 10.11646 / zootaxa. 2505.1. 1" title="Deepwater Indo-Pacific species of the snake-eel genus Ophichthus (Anguilliformes: Ophichthidae), with the description of nine new species." volume="2505" year="2010">McCosker, 2010</bibRefCitation>
</taxonomicName>
, several specimens of
<taxonomicName id="E158809824E5C242252B1E9561215C67" authorityName="Jordan &amp; Snyder" authorityYear="1901" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="asakusae">
<emphasis id="72A1F018ABB69DA5968B82733333356B" italics="true">Ophichthus asakusae</emphasis>
</taxonomicName>
<collectingCountry id="5D09AD47A2BB9EEA86307A177A825AC1" name="Jordan">Jordan</collectingCountry>
&amp; Snyder, 1901 and
<taxonomicName id="BF2DB5D06290B64A869AF7780F544731" authority="(Temminck &amp; Schlegel, 1846)" baseAuthorityName="Temminck &amp; Schlegel" baseAuthorityYear="1846" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="urolophus">
<emphasis id="B23F17BBAD7BEECED5B8FA7652954FEE" italics="true">Ophichthus urolophus</emphasis>
(Temminck &amp; Schlegel, 1846)
</taxonomicName>
; however, they have only one or no protrusions on upper lip, much fewer total vertebrae (
<quantity id="BD2A3377E7E5EB4026D20FFDE9DF7BBE" metricMagnitude="0" metricUnit="m" metricValue="3.3528" metricValueMax="3.3781999999999996" metricValueMin="3.3274" unit="in" value="132.0" valueMax="133.0" valueMin="131.0">131–133 in</quantity>
<taxonomicName id="EC3062058EB7C4322E40CFCBA0236572" authorityName="McCosker" authorityYear="2010" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="alleni">
<emphasis id="03FE430024C8FFE37272703C61929D33" italics="true">O. alleni</emphasis>
</taxonomicName>
,
<quantity id="63641E44859A80DEE4BB6AB222D075FA" metricMagnitude="0" metricUnit="m" metricValue="3.2766000000000006" metricValueMax="3.3528000000000007" metricValueMin="3.2004000000000006" unit="in" value="129.0" valueMax="132.0" valueMin="126.0">126–132 in</quantity>
<taxonomicName id="55C33D8B8EF56C55A730565CD7C78C4C" authorityName="Jordan &amp; Snyder" authorityYear="1901" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="asakusae">
<emphasis id="95BB0B9E376A7F878523B69248500D8D" italics="true">O. asakusae</emphasis>
</taxonomicName>
and
<quantity id="A36966C868F965C438F8C35B61F95649" metricMagnitude="0" metricUnit="m" metricValue="3.4671" metricValueMax="3.5305999999999997" metricValueMin="3.4036" unit="in" value="136.5" valueMax="139.0" valueMin="134.0">134–139 in</quantity>
<taxonomicName id="DF69F3950A172150506D34E290C53875" baseAuthorityName="Temminck &amp; Schlegel" baseAuthorityYear="1846" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="urolophus">
<emphasis id="BB4099D245820E8B9412066C91612595" italics="true">O. urolophus</emphasis>
</taxonomicName>
), and a bicolored body with a mostly pale ventral surface (
<bibRefCitation id="3F79DFF81FD22C7CC34EC6952B991949" DOI="10.11646/zootaxa.2505.1.1" author="McCosker" firstAuthor="McCosker" issue="1" journalOrPublisher="Zootaxa" pagination="1-39" refId="B12" refString="McCosker JE (2010) Deepwater Indo-Pacific species of the snake-eel genus Ophichthus (Anguilliformes: Ophichthidae), with the description of nine new species. Zootaxa 2505 (1): 1–39. https://doi.org/10.11646/zootaxa.2505.1.1" title="Deepwater Indo-Pacific species of the snake-eel genus Ophichthus (Anguilliformes: Ophichthidae), with the description of nine new species." volume="2505" year="2010">McCosker 2010</bibRefCitation>
;
<bibRefCitation id="309837E5F8EABDCCA88B6CED9D07F221" DOI="10.1007/s10228-018-00677-3" author="Hibino Y &amp; McCosker JE &amp; Tashiro F" issue="2" journalOrPublisher="Ichthyological Research" pagination="289-306" refId="B7" refString="Hibino Y, McCosker JE, Tashiro F (2019 b) Four new deepwater Ophichthus (Anguilliformes: Ophichthidae) from Japan with a redescription of Ophichthus pallens (Richardson 1848). Ichthyological Research 66 (2): 289–306. https://doi.org/10.1007/s10228-018-00677-3" title="Four new deepwater Ophichthus (Anguilliformes: Ophichthidae) from Japan with a redescription of Ophichthus pallens (Richardson 1848)." volume="66">Hibino et al. 2019 b</bibRefCitation>
; this study).
</paragraph>
<paragraph id="1A3C2D90C211DCFF8BCB3B8538E0C746">
In
<collectingCountry id="6F69A0585EB6A3736CF8B775C6948D57" name="Taiwan">Taiwan</collectingCountry>
,
<taxonomicName id="B3D3FD7BD58AE891D6DCA5422BF3A3D5" authorityName="Hibino &amp; Ho" authorityYear="2024" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="kbalanensis" status="sp. nov.">
<emphasis id="B478A6A2DDF983F1F715C57868576C60" italics="true">O. kbalanensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C48C868537A17B80A6963F790ED31DCB" rank="species">sp. nov.</taxonomicNameLabel>
is also similar to
<taxonomicName id="285D898F112F0D8E941B81B1A6591733" authorityName="McCosker, Ide &amp; Endo" authorityYear="2012" family="Ophichthidae" genus="Ophichthus" kingdom="Animalia" order="Anguilliformes" phylum="Chordata" rank="species" species="obtusus">
<emphasis id="3CE9DB907923C0F2467DBF58BB1F372E" italics="true">O. obtusus</emphasis>
</taxonomicName>
in the uniformly black body and vertebral count, but it can be distinguished by the different tooth arrangement on the jaws, more mandibular pores (6 or 7 vs 4 or 5), position of the end of the rictus (behind posterior margin of eye vs not behind), and two small, simple, thorn-like labial protrusions on the upper lip (vs at least anterior one fat, cauliflower-shaped protrusion with weak wrinkles) (
<bibRefCitation id="DA2C040254AC7BAD9C55FE3A57B97F20" author="McCosker" etAl="et al." firstAuthor="McCosker" refId="B14" refString="McCosker JE, Ide S, Endo H (2012) Three new species of ophichthid eels (Anguilliformes: Ophichthidae) from Japan. Bulletin of the National Museum of Nature and Science, Series A (Supplement 6): 1–16." year="2012">McCosker et al. 2012</bibRefCitation>
;
<bibRefCitation id="E1B862A70FB1935F8F406761CD950CCC" DOI="10.6119/JMST-013-1220-10" author="Chiu" etAl="et al." firstAuthor="Chiu" refId="B2" refString="Chiu Y-C, Lin J, Chen H-M (2013) One new record genus and three new record species of snake eels (Ophichthidae: Anguilliformes) from Taiwan. Journal of Marine Science and Technology (Supplement 21): 201–206. https://doi.org/10.6119/JMST-013-1220-10" year="2013">Chiu et al. 2013</bibRefCitation>
; this study).
</paragraph>
</subSubSection>
</treatment>
</document>