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<document ID-DOI="10.1038/s41586-020-2234-8" ID-GBIF-Dataset="1a35b0e1-2128-477d-b81f-8c05f3773f36" ID-GBIF-Taxon="163886134" ID-Zenodo-Dep="3784151" LSID="urn:lsid:zoobank.org:act:DA019E3D-6B27-4728-9321-A0502EA0FEC4" checkinTime="1588577580304" checkinUser="plazi" docAuthor="Krause, David W., Hoffmann, Simone, Hu, Yaoming, Wible, John R., Rougier, Guillermo W., Kirk, E. Christopher, Groenke, Joseph R., Rogers, Raymond R., Rossie, James B., Schultz, Julia A., Koenigswald, Wighart von &amp; Rahantarisoa, Lydia J." docDate="2020" docId="C62D8780DE3EFFCB7E532758FDCEFAE0" docLanguage="en" docName="nature.s41586-020-2234-8.pdf" docOrigin="Nature 580 (7805)" docStyle="DocumentStyle{}" docTitle="Adalatheriidae Krause, Hoffmann, Hu, Wible, Rougier, Kirk, Groenke, Rogers, Rossie, Schultz, Koenigswald &amp; Rahantarisoa, 2020, gen. et sp.nov." docType="treatment" docUuid="DA019E3D-6B27-4728-9321-A0502EA0FEC4" docUuidSource="ZooBank" docVersion="5" lastPageId="6" lastPageNumber="7" masterDocId="3A14FFF8DE3FFFCD7E39215AFFCFFFAE" masterDocTitle="Skeleton of a Cretaceous mammal from Madagascar reflects long-term insularity" masterLastPageNumber="21" masterPageNumber="1" pageId="1" pageNumber="2" updateTime="1643459774227" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Skeleton of a Cretaceous mammal from Madagascar reflects long-term insularity</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Krause, David W.</mods:namePart>
<mods:affiliation>Department of Earth Sciences, Denver Museum of Nature and Science, Denver, CO, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Hoffmann, Simone</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Hu, Yaoming</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Wible, John R.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rougier, Guillermo W.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Kirk, E. Christopher</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Groenke, Joseph R.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rogers, Raymond R.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rossie, James B.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Schultz, Julia A.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Koenigswald, Wighart von</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rahantarisoa, Lydia J.</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>Nature</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2020</mods:date>
<mods:detail type="pubDate">
<mods:number>2020-04-29</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>580</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>7805</mods:number>
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<mods:extent unit="page">
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1038/s41586-020-2234-8</mods:identifier>
<mods:identifier type="GBIF-Dataset">1a35b0e1-2128-477d-b81f-8c05f3773f36</mods:identifier>
<mods:identifier type="Zenodo-Dep">3784151</mods:identifier>
<mods:identifier type="ZooBank">A1DB2209-3DB1-46E0-9E0B-2BA801E93BAE</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="163886134" LSID="urn:lsid:zoobank.org:act:DA019E3D-6B27-4728-9321-A0502EA0FEC4" httpUri="http://treatment.plazi.org/id/C62D8780DE3EFFCB7E532758FDCEFAE0" lastPageId="6" lastPageNumber="7" pageId="1" pageNumber="2">
<subSubSection pageId="1" pageNumber="2" type="nomenclature">
<paragraph blockId="1.[106,787,1538,1615]" pageId="1" pageNumber="2">
<taxonomicName box="[106,251,1538,1558]" family="Adalatheriidae" pageId="1" pageNumber="1" rank="family">Adalatheriidae</taxonomicName>
Krause,Hoffmann,Wible,and Rougier,2020,fam.nov.
<taxonomicName authority="Krause,Hoffmann, Wible,and Rougier,2020" authorityName="Krause, Hoffmann, Wible, and Rougier" authorityYear="2020" box="[110,732,1566,1586]" genus="Adalatherium" pageId="1" pageNumber="2" rank="species" species="hui" status="gen. et sp.nov.">
<emphasis box="[110,283,1567,1586]" italics="true" pageId="1" pageNumber="2">Adalatherium hui</emphasis>
Krause,Hoffmann, Wible,and Rougier,2020
</taxonomicName>
,
<taxonomicNameLabel pageId="1" pageNumber="2" rank="species">gen. et sp.nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="2" type="etymology">
<paragraph blockId="1.[106,788,1648,1988]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[106,230,1648,1671]" pageId="1" pageNumber="2">Etymology.</emphasis>
<emphasis box="[233,292,1654,1673]" italics="true" pageId="1" pageNumber="2">Adàla</emphasis>
(Malagasy),crazy;
<emphasis box="[488,567,1654,1673]" italics="true" pageId="1" pageNumber="2">therium</emphasis>
(Latinized form of the Greek
<emphasis box="[170,238,1683,1702]" italics="true" pageId="1" pageNumber="2">θηριoν</emphasis>
), beast;the species name
<taxonomicName authorityName="Krause, Hoffmann, Wible, and Rougier" authorityYear="2020" box="[506,538,1683,1702]" genus="Adalatherium" pageId="1" pageNumber="2" rank="species" species="hui">
<emphasis box="[506,538,1683,1702]" italics="true" pageId="1" pageNumber="2">hui</emphasis>
</taxonomicName>
is in reference to the late Yaoming Hu for his contributions to our knowledge of early mammals.
<materialsCitation ID-GBIF-Occurrence="2609799207" collectionCode="UA" country="Madagascar" location="Berivotra study area" pageId="1" pageNumber="2" specimenCode="UA 9030, MAD99-15" specimenCount="1" typeStatus="holotype">
<emphasis bold="true" box="[106,214,1734,1757]" pageId="1" pageNumber="2">
<typeStatus box="[106,207,1734,1757]" pageId="1" pageNumber="2">Holotype</typeStatus>
.
</emphasis>
<specimenCode box="[217,307,1739,1759]" pageId="1" pageNumber="2">UA 9030</specimenCode>
,skull and postcranial skeleton.
<paragraph blockId="1.[106,788,1648,1988]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[106,387,1763,1786]" pageId="1" pageNumber="2">Type locality and horizon.</emphasis>
<specimenCode box="[391,499,1768,1788]" pageId="1" pageNumber="2">MAD99-15</specimenCode>
,
<location LSID="urn:lsid:plazi:treatment:C62D8780DE3EFFCB7E532758FDCEFAE0:4B5B604DDE3EFFCC7FC027B2FD09F952" box="[505,710,1768,1788]" country="Madagascar" name="Berivotra study area" pageId="1" pageNumber="2">Berivotra study area</location>
(northwestern
<collectingCountry box="[189,316,1796,1816]" name="Madagascar" pageId="1" pageNumber="2">Madagascar</collectingCountry>
).
<location LSID="urn:lsid:plazi:treatment:C62D8780DE3EFFCB7E532758FDCEFAE0:4B5B604DDE3EFFCC7F71265EFE49F8E8" country="Madagascar" name="Upper Cretaceous series (Maastrichtian stage, 72.1 - 66 million years (Myr) ago), Anembalemba Member, Maevarano Formation, MahajangaBasin 18" pageId="1" pageNumber="2">
Upper Cretaceous series (Maastrichtian stage, 72.166million years (Myr) ago),Anembalemba Member,Maevarano Formation,MahajangaBasin
<bibRefCitation author="Rogers, R. R. &amp; Hartman, J. H. &amp; Krause, D. W." box="[377,390,1849,1862]" journalOrPublisher="J. Geol." pageId="1" pageNumber="2" pagination="275 - 301" part="108" refId="ref5006" refString="18. Rogers, R. R., Hartman, J. H. &amp; Krause, D. W. Stratigraphic analysis of Upper Cretaceous rocks in the Mahajanga Basin, northwestern Madagascar: implications for ancient and modern faunas. J. Geol. 108, 275 - 301 (2000)." title="Stratigraphic analysis of Upper Cretaceous rocks in the Mahajanga Basin, northwestern Madagascar: implications for ancient and modern faunas" type="journal article" year="2000">
<superScript attach="left" box="[377,390,1849,1862]" fontSize="5" pageId="1" pageNumber="2">18</superScript>
</bibRefCitation>
</location>
.Additionalinformationonthegeological context is provided in the Supplementary Information.
</paragraph>
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="2" type="diagnosis">
<paragraph blockId="1.[106,788,1648,1988]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[106,220,1906,1929]" pageId="1" pageNumber="2">Diagnosis.</emphasis>
<taxonomicName authorityName="Krause, Hoffmann, Wible, and Rougier" authorityYear="2020" box="[224,280,1912,1931]" genus="Adalatherium" pageId="1" pageNumber="2" rank="species" species="hui">
<emphasis box="[224,280,1912,1931]" italics="true" pageId="1" pageNumber="2">
A.
<taxonomicName box="[247,280,1912,1931]" pageId="1" pageNumber="2" rank="species" species="hui">hui</taxonomicName>
</emphasis>
</taxonomicName>
differs from all other known Mesozoic mammaliaforms in possessing quadrangular upper postcanine tooth crowns with four major cusps and three connecting perimetric ridges mesially,
</paragraph>
<paragraph blockId="1.[816,1497,1538,1702]" pageId="1" pageNumber="2">lingually and distally that border—on threesides—a central valley that opens buccally; and lower postcanine tooth crowns with four major cusps arranged in a diamond pattern and connected by four perimetric crests,and a prominent mesiobuccal basin on the two distalmost lower</paragraph>
<paragraph blockId="1.[816,1497,1538,1702]" pageId="1" pageNumber="2">postcanines. The full diagnosis is provided in the Supplementary Information.</paragraph>
</subSubSection>
<subSubSection lastPageId="3" lastPageNumber="4" pageId="1" pageNumber="2" type="description">
<paragraph blockId="1.[816,1498,1758,1988]" box="[816,926,1758,1784]" pageId="1" pageNumber="2">
<heading bold="true" box="[816,926,1758,1784]" fontSize="10" level="4" pageId="1" pageNumber="2" reason="0">
<emphasis bold="true" box="[816,926,1758,1784]" pageId="1" pageNumber="2">Cranium</emphasis>
</heading>
</paragraph>
<paragraph blockId="1.[816,1498,1758,1988]" pageId="1" pageNumber="2">
The cranium of
<taxonomicName box="[972,1110,1797,1816]" genus="Adalatherium" pageId="1" pageNumber="2" rank="genus">
<emphasis box="[972,1110,1797,1816]" italics="true" pageId="1" pageNumber="2">Adalatherium</emphasis>
</taxonomicName>
reveals a mosaic of plesiomorphic and derived features (
<figureCitation box="[988,1042,1825,1845]" captionStart="Fig" captionStartId="2.[106,137,1614,1632]" captionTargetBox="[90,1497,119,1594]" captionTargetId="figure@2.[589,1024,1257,1589]" captionTargetPageId="2" captionText="Fig.2 |Cranium,lower jaw anddentition of A.hui holotype(UA 9030). ad, Reconstructed craniumin dorsal (a), ventral (b), rightlateral (c) and anterior(d) views.eg, Reconstructed right lowerjaw in lateral (e),dorsal (= occlusal) (f) andmedial (g) views.hk, Micro-computed tomography(ΜCT) digital renderings of rightupper dentition,showing thepostcanine teeth(h), distalincisor (i) and mesialincisor (j) inbuccal views,and thepostcanine teeth in occlusal view (k). ln, ΜCT digitalrenderings of right lowerdentition, showing thepostcanine teeth(l)and incisor (m) inbuccal views,and the postcanine teethin occlusal view (n).Scale bars,2 cm (ag; scale barabove e and f applies toag), 5mm (hn). PC,upper postcanine tooth;pc,lower postcanine tooth." figureDoi="http://doi.org/10.5281/zenodo.3784157" httpUri="https://zenodo.org/record/3784157/files/figure.png" pageId="1" pageNumber="2">Fig. 2</figureCitation>
ad, Extended Data
<figureCitation box="[1237,1291,1825,1845]" captionStart="Fig" captionStartId="4.[106,137,1572,1590]" captionTargetBox="[138,1476,134,1542]" captionTargetId="figure@4.[695,981,710,990]" captionTargetPageId="4" captionText="Fig.3 |Skeleton ofA.huiholotype (UA 9030). ag, Digitally reconstructed skeletonin left lateral view,highlighting the left scapulocoracoidin lateral view (a), thoracic vertebra 6 andlumbar vertebra 7 in anteriorand dorsal views (missingparts mirrored andrendered as semi-transparent) (b), theleft femurin distaland anterior views(c),the left humerus in anteriorand distalviews (d), the left astragalusand navicular inanterior view,and left calcaneusin medial and dorsal views (e),left hind foot indorsal view (f),and theleft tibia inlateral and anterior views(g). L,lumbar vertebra;T,thoracic vertebra.Scale bars,5 cm (main skeleton),1cm (ad, f, g), 5mm (e)." figureDoi="http://doi.org/10.5281/zenodo.3784159" httpUri="https://zenodo.org/record/3784159/files/figure.png" pageId="1" pageNumber="2">Fig.3</figureCitation>
,Supplementary Videos 13). The presence of a very large internasal vacuity,five infraorbi-
</paragraph>
<paragraph blockId="1.[816,1498,1758,1988]" lastBlockId="3.[106,787,134,526]" lastPageId="3" lastPageNumber="4" pageId="1" pageNumber="2">
tal foramina, a large foramen in the lacrimal that is not related to the nasolacrimal duct (probably for the ethmoidal branch of ophthalmic nerve (V
<subScript attach="left" box="[903,909,1949,1962]" fontSize="5" pageId="1" pageNumber="2">1</subScript>
) and associated vessels), numerous nasal foramina and a paranasal sinus that arises from the anterior vestibule of the nasal cavity are particularly unusual for mammaliaforms.The snout region shares several features with that of
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[451,531,1884,1902]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis box="[451,531,1884,1902]" italics="true" pageId="2" pageNumber="3">Vintana</emphasis>
</taxonomicName>
<superScript attach="left" box="[531,550,1878,1891]" fontSize="5" pageId="2" pageNumber="3">
<bibRefCitation author="Krause, D. W." box="[531,538,1878,1891]" journalOrPublisher="Nature" pageId="2" pageNumber="3" pagination="512 - 517" part="515" refId="ref4435" refString="3. Krause, D. W. et al. First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism. Nature 515, 512 - 517 (2014)." title="First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism" type="journal article" year="2014">3</bibRefCitation>
,
<bibRefCitation author="Krause, D. W." box="[542,550,1878,1891]" journalOrPublisher="J. Vertebr. Paleontol." pageId="2" pageNumber="3" pagination="6" part="34" refId="ref4467" refString="4. Krause, D. W. (ed.) Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar. J. Vertebr. Paleontol. 34, suppl. to issue 6 (2014)." title="Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar" type="journal article" year="2014">4</bibRefCitation>
</superScript>
, including the presence of a massive lacrimal bone that excludes the frontal from contacting the maxilla and a large septomaxilla with prominent posterodorsal and intranarial processes.By contrast,
<taxonomicName box="[482,618,1969,1988]" genus="Adalatherium" pageId="2" pageNumber="3" rank="genus">
<emphasis box="[482,618,1969,1988]" italics="true" pageId="2" pageNumber="3">Adalatherium</emphasis>
</taxonomicName>
does not possess several autapomorphic features that are seen in
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[1295,1375,1855,1873]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis box="[1295,1375,1855,1873]" italics="true" pageId="2" pageNumber="3">Vintana</emphasis>
</taxonomicName>
, including a massive jugal flange and contact between the premaxillae and palatines. Although much of the posterior part of the cranium was severely damaged post mortem, the left inner ear of UA 9030 is partially pre- served and exhibits several features that were previously unknown among mammaliaforms (Extended
<taxonomicName authorityName="Walker" authorityYear="1862" box="[474,523,134,154]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="genus">Data</taxonomicName>
<figureCitation box="[528,588,134,154]" captionStart="Fig" captionStartId="5.[106,137,1456,1474]" captionTargetBox="[119,1483,135,1429]" captionTargetId="graphics@5.[170,1432,811,1406]" captionTargetPageId="5" captionText="Fig.4 |Keystages in platetectonic historyof Madagascar.a, Position of Madagascar before rifting betweenWestGondwana (South America and Africa)and EastGondwana (Madagascar,Seychelles,Indiansubcontinent, Sri Lanka,Antarctica and Australia) at183 Myr ago (Early Jurassic epoch). b,Separation of Indo-Madagascarfrom Antarctica andAustralia at 124Myr ago (mid-Early Cretaceous epoch).c, Separation of Indiansubcontinent from Madagascar at 88 Myr ago (mid-Late Cretaceous epoch).d, Approximate time of deposition of MaevaranoFormation at 66 Myrago (latest Cretaceous period).Solid blacklines indicate current coastlines of Madagascar andeast Africa;brown representsPrecambrian terranes;andyellow indicates sedimentary basins along west coast of Madagascar.The discovery siteof UA 9030 isindicated by red star ind.Scale bars,500 km.Maps adapted from Earthworks (www.reeves.nl/gond.com)." figureDoi="http://doi.org/10.5281/zenodo.3784161" httpUri="https://zenodo.org/record/3784161/files/figure.png" pageId="3" pageNumber="4">Fig. 4</figureCitation>
). Most notably, the primary bony lamina is structurally different from that of therians in being single-layered instead of double-layered,and the branches of the cochlear nerve appear to have passed along the surface of—rather than within—the primary lamina.This unique morphology suggests that the primary bony lamina of
<taxonomicName box="[381,519,278,297]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[381,519,278,297]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
evolved convergently with those of therian mammals.The cochleae of
<taxonomicName box="[527,663,307,326]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[527,663,307,326]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[707,787,307,325]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[707,787,307,325]" italics="true" pageId="3" pageNumber="4">Vintana</emphasis>
</taxonomicName>
are unique among mammaliaforms in possessing a secondary bony canal that parallels the cochlear ganglion canal,and probably enclosed a vascular network. Overall,the cochlear canal is curved through at least 210° and possesses, in addition to the primary bony lamina, the base of a secondary bony lamina,a cribriform plate,a well-developed cochlear ganglion canal and a separate canal for the lagenar nerve;this last feature is not present in
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[383,463,508,526]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[383,463,508,526]" italics="true" pageId="3" pageNumber="4">Vintana</emphasis>
</taxonomicName>
<bibRefCitation author="Krause, D. W." box="[463,471,502,515]" journalOrPublisher="J. Vertebr. Paleontol." pageId="3" pageNumber="4" pagination="6" part="34" refId="ref4467" refString="4. Krause, D. W. (ed.) Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar. J. Vertebr. Paleontol. 34, suppl. to issue 6 (2014)." title="Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar" type="journal article" year="2014">
<superScript attach="left" box="[463,471,502,515]" fontSize="5" pageId="3" pageNumber="4">4</superScript>
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.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3784155" ID-Zenodo-Dep="3784155" httpUri="https://zenodo.org/record/3784155/files/figure.png" pageId="2" pageNumber="3" startId="2.[106,137,1614,1632]" targetBox="[90,1497,119,1594]" targetPageId="2">
<paragraph blockId="2.[106,778,1614,1765]" lastBlockId="2.[816,1466,1613,1739]" pageId="2" pageNumber="3">
<caption ID-DOI="http://doi.org/10.5281/zenodo.3784157" ID-Zenodo-Dep="3784157" box="[106,730,1614,1632]" httpUri="https://zenodo.org/record/3784157/files/figure.png" pageId="2" pageNumber="3" startId="2.[106,137,1614,1632]" targetBox="[90,1497,119,1594]" targetPageId="2">
Fig.2 |Cranium,lower jaw and dentition of
<taxonomicName authorityName="Krause, Hoffmann, Wible, and Rougier" authorityYear="2020" box="[492,542,1614,1632]" genus="Adalatherium" pageId="2" pageNumber="3" rank="species" species="hui">
<emphasis bold="true" box="[492,542,1614,1632]" italics="true" pageId="2" pageNumber="3">A.hui</emphasis>
</taxonomicName>
holotype(UA 9030).
</caption>
<emphasis bold="true" box="[106,139,1641,1659]" pageId="2" pageNumber="3">ad</emphasis>
, Reconstructed cranium in dorsal (
<emphasis bold="true" box="[440,450,1641,1659]" pageId="2" pageNumber="3">a</emphasis>
), ventral (
<emphasis bold="true" box="[537,549,1641,1659]" pageId="2" pageNumber="3">b</emphasis>
), right lateral (
<emphasis bold="true" box="[674,683,1641,1659]" pageId="2" pageNumber="3">c</emphasis>
) and anterior(
<emphasis bold="true" box="[187,199,1667,1685]" pageId="2" pageNumber="3">d</emphasis>
) views.
<emphasis bold="true" box="[266,298,1667,1685]" pageId="2" pageNumber="3">eg</emphasis>
, Reconstructed right lower jaw in lateral (
<emphasis bold="true" box="[655,665,1667,1685]" pageId="2" pageNumber="3">e</emphasis>
),dorsal (= occlusal) (
<emphasis bold="true" box="[218,225,1694,1712]" pageId="2" pageNumber="3">f</emphasis>
) and medial (
<emphasis bold="true" box="[341,352,1694,1712]" pageId="2" pageNumber="3">g</emphasis>
) views.
<emphasis bold="true" box="[420,453,1694,1712]" pageId="2" pageNumber="3">hk</emphasis>
, Micro-computed tomography(ΜCT) digital renderings of right upper dentition,showing the postcanine teeth(
<emphasis bold="true" box="[743,755,1721,1739]" pageId="2" pageNumber="3">h</emphasis>
), distal incisor (
<emphasis bold="true" box="[228,234,1747,1765]" pageId="2" pageNumber="3">i</emphasis>
) and mesial incisor (
<emphasis bold="true" box="[410,416,1747,1765]" pageId="2" pageNumber="3">j</emphasis>
) in buccal views,and the postcanine teeth in occlusal view (
<emphasis bold="true" box="[961,972,1614,1632]" pageId="2" pageNumber="3">k</emphasis>
).
<emphasis bold="true" box="[987,1016,1614,1632]" pageId="2" pageNumber="3">ln</emphasis>
, ΜCT digital renderings of right lower dentition, showing the postcanine teeth(
<emphasis bold="true" box="[1082,1088,1641,1659]" pageId="2" pageNumber="3">l</emphasis>
)and incisor (
<emphasis bold="true" box="[1203,1220,1641,1659]" pageId="2" pageNumber="3">m</emphasis>
) in buccal views,and the postcanine teeth in occlusal view (
<emphasis bold="true" box="[1110,1122,1667,1685]" pageId="2" pageNumber="3">n</emphasis>
).Scale bars,2 cm (
<emphasis bold="true" box="[1280,1313,1667,1685]" pageId="2" pageNumber="3">ag</emphasis>
; scale bar above
<emphasis bold="true" box="[1456,1466,1667,1685]" pageId="2" pageNumber="3">e</emphasis>
and
<emphasis bold="true" box="[852,859,1694,1712]" pageId="2" pageNumber="3">f</emphasis>
applies to
<emphasis bold="true" box="[949,982,1694,1712]" pageId="2" pageNumber="3">ag</emphasis>
), 5mm (
<emphasis bold="true" box="[1053,1088,1694,1712]" pageId="2" pageNumber="3">hn</emphasis>
). PC,upper postcanine tooth;pc,lower postcanine tooth.
</paragraph>
</caption>
<paragraph blockId="3.[106,788,583,1272]" box="[106,229,583,608]" pageId="3" pageNumber="4">
<heading bold="true" box="[106,229,583,608]" fontSize="10" level="4" pageId="3" pageNumber="4" reason="0">
<emphasis bold="true" box="[106,229,583,608]" pageId="3" pageNumber="4">Lower jaw</emphasis>
</heading>
</paragraph>
<paragraph blockId="3.[106,788,583,1272]" pageId="3" pageNumber="4">
The lower jaw of
<taxonomicName box="[278,421,622,641]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[278,421,622,641]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
is more complete than in any other known gondwanatherian,and is the first to preserve the ascending ramus of the dentary (
<figureCitation box="[321,375,678,698]" captionStart="Fig" captionStartId="2.[106,137,1614,1632]" captionTargetBox="[90,1497,119,1594]" captionTargetId="figure@2.[589,1024,1257,1589]" captionTargetPageId="2" captionText="Fig.2 |Cranium,lower jaw anddentition of A.hui holotype(UA 9030). ad, Reconstructed craniumin dorsal (a), ventral (b), rightlateral (c) and anterior(d) views.eg, Reconstructed right lowerjaw in lateral (e),dorsal (= occlusal) (f) andmedial (g) views.hk, Micro-computed tomography(ΜCT) digital renderings of rightupper dentition,showing thepostcanine teeth(h), distalincisor (i) and mesialincisor (j) inbuccal views,and thepostcanine teeth in occlusal view (k). ln, ΜCT digitalrenderings of right lowerdentition, showing thepostcanine teeth(l)and incisor (m) inbuccal views,and the postcanine teethin occlusal view (n).Scale bars,2 cm (ag; scale barabove e and f applies toag), 5mm (hn). PC,upper postcanine tooth;pc,lower postcanine tooth." figureDoi="http://doi.org/10.5281/zenodo.3784157" httpUri="https://zenodo.org/record/3784157/files/figure.png" pageId="3" pageNumber="4">Fig.2</figureCitation>
eg, Extended
<taxonomicName authorityName="Walker" authorityYear="1862" box="[520,567,678,698]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="genus">Data</taxonomicName>
<figureCitation box="[570,624,678,698]" captionStart="Extended" captionStartId="12.[106,195,1019,1037]" captionTargetBox="[121,1481,133,992]" captionTargetId="figure@12.[121,1481,131,995]" captionText="ExtendedData Fig.5 | Lowerjaw of A.hui holotype(UA 9030).Photographs of left dentary inleft column;photographs of rightdentary in rightcolumn. a,b, Lateral views.c, d, Dorsal (occlusal) views.e, f, Medial views.i, lower incisor;pc,lower postcaninetooth." figureDoi="http://doi.org/10.5281/zenodo.3784171" httpUri="https://zenodo.org/record/3784171/files/figure.png" pageId="3" pageNumber="4">Fig.5</figureCitation>
,Supplementary Videos 47).Among gondwanatherians,the horizontal ramus of
<taxonomicName genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
is essentially identical to that of
<taxonomicName authorityName="Scillato-Yane &amp; Pascual" authorityYear="1984" box="[539,661,737,756]" class="Mammalia" family="Sudamericidae" genus="Sudamerica" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[539,661,737,756]" italics="true" pageId="3" pageNumber="4">Sudamerica</emphasis>
</taxonomicName>
<bibRefCitation author="Pascual, R. &amp; Goin, F. J. &amp; Krause, D. W. &amp; Ortiz-Jaureguizar, E. &amp; Carlini, A. A." box="[661,674,731,744]" journalOrPublisher="J. Vertebr. Paleontol." pageId="3" pageNumber="4" pagination="373 - 382" part="19" refId="ref5062" refString="19. Pascual, R., Goin, F. J., Krause, D. W., Ortiz-Jaureguizar, E. &amp; Carlini, A. A. The first gnathic remains of Sudamerica: implications for gondwanathere relationships. J. Vertebr. Paleontol. 19, 373 - 382 (1999)." title="The first gnathic remains of Sudamerica: implications for gondwanathere relationships" type="journal article" year="1999">
<superScript attach="left" box="[661,674,731,744]" fontSize="5" pageId="3" pageNumber="4">19</superScript>
</bibRefCitation>
but differs from that of
<emphasis box="[226,371,765,784]" italics="true" pageId="3" pageNumber="4">Galulatherium</emphasis>
<bibRefCitation author="O'Connor, P. M." box="[371,377,760,773]" journalOrPublisher="Acta Palaeontol. Pol." pageId="3" pageNumber="4" pagination="65 - 84" part="64" refId="ref4510" refString="5. O'Connor, P. M. et al. A new mammal from the Turonian-Campanian (Upper Cretaceous) Galula Formation, southwestern Tanzania. Acta Palaeontol. Pol. 64, 65 - 84 (2019)." title="A new mammal from the Turonian-Campanian (Upper Cretaceous) Galula Formation, southwestern Tanzania" type="journal article" year="2019">
<superScript attach="left" box="[371,377,760,773]" fontSize="5" pageId="3" pageNumber="4">5</superScript>
</bibRefCitation>
, primarily in having a stepped differential in height between the diastema and the postcanine alveolar portion. The dentary of
<taxonomicName box="[254,392,823,842]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[254,392,823,842]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
is short and deep,and bears a large diastema between the incisor and postcanine teeth,a prominent pterygoid fossa and shelf, and a masseteric fossa that extends anteriorly onto the horizontal ramus.There is no evidence of a postdentary trough, Meckelian sulcus,coronoid bone or angular process.In these features, the dentary of
<taxonomicName box="[252,391,966,985]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[252,391,966,985]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
is similar to those found in members of
<taxonomicName box="[106,285,994,1014]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="order">Multituberculata</taxonomicName>
, which is a largely Laurasian group. The dentaries of euharamiyidans differ from that of
<taxonomicName box="[482,620,1023,1042]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[482,620,1023,1042]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
in possessing an angular process,a coronoid bone and—according to ref.
<bibRefCitation author="Luo, Z. - X." box="[648,664,1046,1059]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="326 - 329" part="548" refId="ref5121" refString="20. Luo, Z. - X. et al. New evidence for mammaliaform ear evolution and feeding adaptation in a Jurassic ecosystem. Nature 548, 326 - 329 (2017)." title="New evidence for mammaliaform ear evolution and feeding adaptation in a Jurassic ecosystem" type="journal article" year="2017">
<superScript attach="right" box="[648,664,1046,1059]" fontSize="5" pageId="3" pageNumber="4">20</superScript>
</bibRefCitation>
—a reduced postdentary trough (although this is disputed in ref.
<bibRefCitation author="Mao, F. - Y. &amp; Meng, J." box="[614,626,1075,1088]" journalOrPublisher="Zool. J. Linn. Soc." pageId="3" pageNumber="4" pagination="529 - 552" part="186" refId="ref5157" refString="21. Mao, F. - Y. &amp; Meng, J. A new haramiyidan mammal from the Jurassic Yanliao Biota and comparisons with other haramiyidans. Zool. J. Linn. Soc. 186, 529 - 552 (2019)." title="A new haramiyidan mammal from the Jurassic Yanliao Biota and comparisons with other haramiyidans" type="journal article" year="2019">
<superScript attach="right" box="[614,626,1075,1088]" fontSize="5" pageId="3" pageNumber="4">21</superScript>
</bibRefCitation>
). The dentary of
<taxonomicName authorityName="Jenkins, Gatesy, Shubin &amp; Amaral" authorityYear="1997" box="[106,235,1110,1128]" class="Mammalia" family="Haramiyidae" genus="Haramiyavia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[106,235,1110,1128]" italics="true" pageId="3" pageNumber="4">Haramiyavia</emphasis>
</taxonomicName>
is much more plesiomorphic than those of both
<taxonomicName genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
and euharamiyidans in retaining a long and shallow horizontal ramus,a fully developed postdentary trough and Meckelian sulcus, and in lacking a pterygoid fossa and shelf
<bibRefCitation author="Luo, Z. X. &amp; Gatesy, S. M. &amp; Jenkins, F. A. Jr &amp; Amaral, W. W. &amp; Shubin, N. H." box="[523,538,1190,1203]" journalOrPublisher="Proc. Natl Acad. Sci. USA" pageId="3" pageNumber="4" pagination="7101 - E 7109" part="112" refId="ref5203" refString="22. Luo, Z. X., Gatesy, S. M., Jenkins, F. A. Jr, Amaral, W. W. &amp; Shubin, N. H. Mandibular and dental characteristics of Late Triassic mammaliaform Haramiyavia and their ramifications for basal mammal evolution. Proc. Natl Acad. Sci. USA 112, E 7101 - E 7109 (2015)." title="Mandibular and dental characteristics of Late Triassic mammaliaform Haramiyavia and their ramifications for basal mammal evolution" type="journal article" year="2015">
<superScript attach="left" box="[523,538,1190,1203]" fontSize="5" pageId="3" pageNumber="4">22</superScript>
</bibRefCitation>
. The masseteric fossa of
<taxonomicName box="[106,244,1224,1243]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[106,244,1224,1243]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
is positioned relatively high dorsally on the dentary,an apparent autapomorphy.
</paragraph>
<paragraph blockId="3.[106,788,1329,1988]" box="[106,228,1329,1354]" pageId="3" pageNumber="4">
<heading bold="true" box="[106,228,1329,1354]" fontSize="10" level="4" pageId="3" pageNumber="4" reason="0">
<emphasis bold="true" box="[106,228,1329,1354]" pageId="3" pageNumber="4">Dentition</emphasis>
</heading>
</paragraph>
<paragraph blockId="3.[106,788,1329,1988]" pageId="3" pageNumber="4">
The dentition of
<taxonomicName box="[274,414,1367,1386]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[274,414,1367,1386]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
is unlike that of any known Mesozoic mammaliaform (
<figureCitation box="[278,335,1395,1415]" captionStart="Fig" captionStartId="2.[106,137,1614,1632]" captionTargetBox="[90,1497,119,1594]" captionTargetId="figure@2.[589,1024,1257,1589]" captionTargetPageId="2" captionText="Fig.2 |Cranium,lower jaw anddentition of A.hui holotype(UA 9030). ad, Reconstructed craniumin dorsal (a), ventral (b), rightlateral (c) and anterior(d) views.eg, Reconstructed right lowerjaw in lateral (e),dorsal (= occlusal) (f) andmedial (g) views.hk, Micro-computed tomography(ΜCT) digital renderings of rightupper dentition,showing thepostcanine teeth(h), distalincisor (i) and mesialincisor (j) inbuccal views,and thepostcanine teeth in occlusal view (k). ln, ΜCT digitalrenderings of right lowerdentition, showing thepostcanine teeth(l)and incisor (m) inbuccal views,and the postcanine teethin occlusal view (n).Scale bars,2 cm (ag; scale barabove e and f applies toag), 5mm (hn). PC,upper postcanine tooth;pc,lower postcanine tooth." figureDoi="http://doi.org/10.5281/zenodo.3784157" httpUri="https://zenodo.org/record/3784157/files/figure.png" pageId="3" pageNumber="4">Fig. 2</figureCitation>
hn, Supplementary Video 8 for upper postcanines). There are two very large,open-rooted upper incisors, each of which bears a buccally restricted band of enamel. The size, shape and positional relationships of the upper incisors are very similar to those discerned from the alveoli of
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[455,535,1511,1529]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[455,535,1511,1529]" italics="true" pageId="3" pageNumber="4">Vintana</emphasis>
</taxonomicName>
<superScript attach="left" box="[535,553,1505,1518]" fontSize="5" pageId="3" pageNumber="4">
<bibRefCitation author="Krause, D. W." box="[535,542,1505,1518]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="512 - 517" part="515" refId="ref4435" refString="3. Krause, D. W. et al. First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism. Nature 515, 512 - 517 (2014)." title="First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism" type="journal article" year="2014">3</bibRefCitation>
,
<bibRefCitation author="Krause, D. W." box="[545,553,1505,1518]" journalOrPublisher="J. Vertebr. Paleontol." pageId="3" pageNumber="4" pagination="6" part="34" refId="ref4467" refString="4. Krause, D. W. (ed.) Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar. J. Vertebr. Paleontol. 34, suppl. to issue 6 (2014)." title="Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar" type="journal article" year="2014">4</bibRefCitation>
</superScript>
. The presence of upper canines in
<taxonomicName box="[211,350,1539,1558]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[211,350,1539,1558]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
is indicated by tiny elliptical alveoli that are separated mesially from the incisors and distally from the postcanines by sizeable diastemata. The first upper postcanine is a small, simple, two-rooted premolariform tooth. The four more-distal postcanines are each supported by five or more roots and are unique among Mesozoic mammaliaforms in bearing four major cusps connected by ridges mesially, lingually and distally that border—on three sides—a central valley that opens buccally.
</paragraph>
<paragraph blockId="3.[106,788,1329,1988]" lastBlockId="3.[816,1498,134,383]" pageId="3" pageNumber="4">
The single lower incisor is large,curved and open-rooted,and bears enamel that is largely restricted to the buccal surface.In these features, the lower incisor resembles those known for other gondwanatherians (except for the enamel-less condition in
<emphasis box="[505,651,1855,1874]" italics="true" pageId="3" pageNumber="4">Galulatherium</emphasis>
<bibRefCitation author="O'Connor, P. M." box="[651,658,1849,1862]" journalOrPublisher="Acta Palaeontol. Pol." pageId="3" pageNumber="4" pagination="65 - 84" part="64" refId="ref4510" refString="5. O'Connor, P. M. et al. A new mammal from the Turonian-Campanian (Upper Cretaceous) Galula Formation, southwestern Tanzania. Acta Palaeontol. Pol. 64, 65 - 84 (2019)." title="A new mammal from the Turonian-Campanian (Upper Cretaceous) Galula Formation, southwestern Tanzania" type="journal article" year="2019">
<superScript attach="left" box="[651,658,1849,1862]" fontSize="5" pageId="3" pageNumber="4">5</superScript>
</bibRefCitation>
). Each of the four lower postcanines has four cusps connected by prominent crests, forming a diamond pattern.The most mesial cusp dominates the crown on all four teeth. The second lower postcanine bears a mesiobuccal bulge,which is developed into a prominent basin on the penultimate and ultimate lower postcanines.The first lower postcanine has two roots, whereas each of the more distal postcanines has at least four. The enamel microstructure of
<emphasis box="[1139,1290,192,211]" italics="true" pageId="3" pageNumber="4">Aadalatherium</emphasis>
consists of relatively plesiomorphic normal radial enamel (Extended Data
<figureCitation box="[1373,1433,220,240]" captionStart="Extended" captionStartId="14.[106,195,133,151]" captionTargetId="figure@13.[169,1433,131,1744]" captionTargetPageId="13" captionText="ExtendedData Fig.6 |Enamelmicrostructure of A. huiholotype(UA 9030). ad, Scanning electron micrographsof single postcanine tooth enamel fragment sectioned invarious planes.a, Transverse section of entire enamel band from the enameldentinejunction (EDJ) tothe outer enamel surface (OES) (about 0.4-mm thick) showingsingle layer of radial enamel and absence of distinctlayer of prismless external enamel.Prism sizeincreases from,on average,2.3 to 2.8 Μm from theenameldentine junction tothe outer enamel surface.Prisms close tothe enameldentinejunction are intersected by interprismatic matrixat slightly higherangles than towardsthe outer enamel surface.b, Transverse section showingthe clear distinction betweenenamel" figureDoi="http://doi.org/10.5281/zenodo.3784173" httpUri="https://zenodo.org/record/3784173/files/figure.png" pageId="3" pageNumber="4">Fig. 6</figureCitation>
), typi- cal of several gondwanatherians from the Late Cretaceous epoch and Palaeogene subperiod of Argentina
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,
<bibRefCitation author="Krause, D. W." box="[1177,1185,272,285]" journalOrPublisher="J. Vertebr. Paleontol." pageId="3" pageNumber="4" pagination="6" part="34" refId="ref4467" refString="4. Krause, D. W. (ed.) Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar. J. Vertebr. Paleontol. 34, suppl. to issue 6 (2014)." title="Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar" type="journal article" year="2014">4</bibRefCitation>
</superScript>
. It is unlike the modified radial enamel,with pronounced inter-row sheets of interprismatic matrix, documented for other gondwanatherians(
<taxonomicName box="[1230,1315,335,354]" class="Mammalia" family="Sudamericidae" genus="Lavanify" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[1230,1315,335,354]" italics="true" pageId="3" pageNumber="4">Lavanify</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[1358,1437,336,354]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[1358,1437,336,354]" italics="true" pageId="3" pageNumber="4">Vintana</emphasis>
</taxonomicName>
) from the Late Cretaceous epoch of India and Madagascar
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,
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</superScript>
.
</paragraph>
<paragraph blockId="3.[816,1499,440,1014]" box="[816,1071,440,465]" pageId="3" pageNumber="4">
<heading bold="true" box="[816,1071,440,465]" fontSize="10" level="4" pageId="3" pageNumber="4" reason="0">
<emphasis bold="true" box="[816,1071,440,465]" pageId="3" pageNumber="4">Postcranial skeleton</emphasis>
</heading>
</paragraph>
<paragraph blockId="3.[816,1499,440,1014]" pageId="3" pageNumber="4">
UA 9030 includes the only postcranial material assigned to a gondwanatherian,and
<taxonomicName box="[956,1088,507,526]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[956,1088,507,526]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
isonlythefourthMesozoicmammaliaform from Gondwana represented by articulated postcranial remains
<bibRefCitation author="Kielan-Jaworowska, Z. &amp; Cifelli, R. L. &amp; Luo, Z. - X." box="[1443,1448,530,543]" journalOrPublisher="Press" pageId="3" pageNumber="4" refId="ref4358" refString="1. Kielan-Jaworowska, Z., Cifelli, R. L. &amp; Luo, Z. - X. Mammals from the Age of Dinosaurs: Origins, Evolution, and Structure (Columbia Univ. Press, 2004)." title="Mammals from the Age of Dinosaurs: Origins, Evolution, and Structure (Columbia Univ" type="book" year="2004">
<superScript attach="left" box="[1443,1448,530,543]" fontSize="5" pageId="3" pageNumber="4">1</superScript>
</bibRefCitation>
. The postcranial skeleton exhibits a number of unusual features,including an anteroposteriorly bowed and mediolaterally compressed tibia,a trochleatedsurfaceonthedistalastragalus,alargenumberoftrunkvertebrae (at least16thoracic and12lumbar vertebrae),and a short tail(24vertebrae, almost all wider than they are long) (
<figureCitation box="[1156,1215,678,698]" captionStart="Fig" captionStartId="1.[106,137,1437,1455]" captionTargetBox="[164,1462,133,1406]" captionTargetId="figure@1.[611,1085,489,848]" captionTargetPageId="1" captionText="Fig.1 |Skull andpostcranial skeletonof A. huiholotype(UA 9030).a, Top view,as preserved.Scale bar,5cm.b, Reconstructionin left lateral view.Left and rightsides indicated as (l) and (r),respectively." figureDoi="http://doi.org/10.5281/zenodo.3784153" httpUri="https://zenodo.org/record/3784153/files/figure.png" pageId="3" pageNumber="4">Figs.1</figureCitation>
,
<figureCitation box="[1224,1235,678,698]" captionStart="Fig" captionStartId="4.[106,137,1572,1590]" captionTargetBox="[138,1476,134,1542]" captionTargetId="figure@4.[695,981,710,990]" captionTargetPageId="4" captionText="Fig.3 |Skeleton ofA.huiholotype (UA 9030). ag, Digitally reconstructed skeletonin left lateral view,highlighting the left scapulocoracoidin lateral view (a), thoracic vertebra 6 andlumbar vertebra 7 in anteriorand dorsal views (missingparts mirrored andrendered as semi-transparent) (b), theleft femurin distaland anterior views(c),the left humerus in anteriorand distalviews (d), the left astragalusand navicular inanterior view,and left calcaneusin medial and dorsal views (e),left hind foot indorsal view (f),and theleft tibia inlateral and anterior views(g). L,lumbar vertebra;T,thoracic vertebra.Scale bars,5 cm (main skeleton),1cm (ad, f, g), 5mm (e)." figureDoi="http://doi.org/10.5281/zenodo.3784159" httpUri="https://zenodo.org/record/3784159/files/figure.png" pageId="3" pageNumber="4">3</figureCitation>
, Extended Data
<figureCitation box="[1387,1445,678,698]" captionStart="Fig" captionStartId="1.[106,137,1437,1455]" captionTargetBox="[164,1462,133,1406]" captionTargetId="figure@1.[611,1085,489,848]" captionTargetPageId="1" captionText="Fig.1 |Skull andpostcranial skeletonof A. huiholotype(UA 9030).a, Top view,as preserved.Scale bar,5cm.b, Reconstructionin left lateral view.Left and rightsides indicated as (l) and (r),respectively." figureDoi="http://doi.org/10.5281/zenodo.3784153" httpUri="https://zenodo.org/record/3784153/files/figure.png" pageId="3" pageNumber="4">Figs.1</figureCitation>
,
<figureCitation box="[1452,1465,678,698]" captionStart="Extended" captionStartId="15.[106,195,1769,1787]" captionTargetBox="[165,1436,133,1740]" captionTargetId="figure@15.[164,1438,131,1744]" captionText="ExtendedData Fig.7 |Selected individual vertebrae ofA.hui holotype (UA 9030). Thoracic (T6and T16),lumbar (L1 andL11) andanterior caudal (Ca8) vertebraeare depictedinanterior,dorsal,left lateral and ventral views.Theleft transverse process of L11 is preserved but wasseparated from the vertebral column duringpreparation and hasnot been CTscanned.Dotted outlines represent theshape of preserved left transverse process,and themirrored reconstructed right transverse process." figureDoi="http://doi.org/10.5281/zenodo.3784175" httpUri="https://zenodo.org/record/3784175/files/figure.png" pageId="3" pageNumber="4">7</figureCitation>
,8). The long spinous and transverse processes of the thoracic and lumbar vertebraesuggestthepresenceofenhancedepaxial(back) musculature. In the pectoral girdle,a procoracoid bone is absent but a separate coracoid is well-developed (
<figureCitation box="[992,1055,793,813]" captionStart="Fig" captionStartId="4.[106,137,1572,1590]" captionTargetBox="[138,1476,134,1542]" captionTargetId="figure@4.[695,981,710,990]" captionTargetPageId="4" captionText="Fig.3 |Skeleton ofA.huiholotype (UA 9030). ag, Digitally reconstructed skeletonin left lateral view,highlighting the left scapulocoracoidin lateral view (a), thoracic vertebra 6 andlumbar vertebra 7 in anteriorand dorsal views (missingparts mirrored andrendered as semi-transparent) (b), theleft femurin distaland anterior views(c),the left humerus in anteriorand distalviews (d), the left astragalusand navicular inanterior view,and left calcaneusin medial and dorsal views (e),left hind foot indorsal view (f),and theleft tibia inlateral and anterior views(g). L,lumbar vertebra;T,thoracic vertebra.Scale bars,5 cm (main skeleton),1cm (ad, f, g), 5mm (e)." figureDoi="http://doi.org/10.5281/zenodo.3784159" httpUri="https://zenodo.org/record/3784159/files/figure.png" pageId="3" pageNumber="4">Fig.3a</figureCitation>
, Extended Data
<figureCitation box="[1208,1294,793,813]" captionStart="Extended" captionStartId="17.[106,195,1160,1178]" captionTargetBox="[135,1471,133,1135]" captionTargetId="figure@17.[131,1472,131,1136]" captionText="ExtendedData Fig.9 |Pectoraland pelvic girdle elements of A.hui holotype(UA9030).ad, ΜCT images.a, b, Left scapulacoracoid,left andrightclavicle and manubrium intop(a)and bottom (b) views(as preserved).c, d, Leftos coxa and epipubic bone inlateral (c)and medial (d) views." figureDoi="http://doi.org/10.5281/zenodo.3784179" httpUri="https://zenodo.org/record/3784179/files/figure.png" pageId="3" pageNumber="4">Fig.9a,b</figureCitation>
). The forelimbs had a moderately parasagittal posture,as indicated by the ventrally directed glenoid fossa and the well-developed humeral trochlea(
<figureCitation box="[1313,1396,850,870]" captionStart="Fig" captionStartId="4.[106,137,1572,1590]" captionTargetBox="[138,1476,134,1542]" captionTargetId="figure@4.[695,981,710,990]" captionTargetPageId="4" captionText="Fig.3 |Skeleton ofA.huiholotype (UA 9030). ag, Digitally reconstructed skeletonin left lateral view,highlighting the left scapulocoracoidin lateral view (a), thoracic vertebra 6 andlumbar vertebra 7 in anteriorand dorsal views (missingparts mirrored andrendered as semi-transparent) (b), theleft femurin distaland anterior views(c),the left humerus in anteriorand distalviews (d), the left astragalusand navicular inanterior view,and left calcaneusin medial and dorsal views (e),left hind foot indorsal view (f),and theleft tibia inlateral and anterior views(g). L,lumbar vertebra;T,thoracic vertebra.Scale bars,5 cm (main skeleton),1cm (ad, f, g), 5mm (e)." figureDoi="http://doi.org/10.5281/zenodo.3784159" httpUri="https://zenodo.org/record/3784159/files/figure.png" pageId="3" pageNumber="4">Fig.3a,d</figureCitation>
,Extended DataFig.8a,b).Bycontrast,theasymmetricalmedialandlateralcondyles of the femur are suggestive of a more sprawled hindlimb posture.Other notable features of the pelvis and hindlimbs include the presence of a largeobturatorforamen (similarinsizetothoseoftherians),anepipubic bone and a large,separate parafibula (
<figureCitation box="[1176,1227,994,1014]" captionStart="Fig" captionStartId="4.[106,137,1572,1590]" captionTargetBox="[138,1476,134,1542]" captionTargetId="figure@4.[695,981,710,990]" captionTargetPageId="4" captionText="Fig.3 |Skeleton ofA.huiholotype (UA 9030). ag, Digitally reconstructed skeletonin left lateral view,highlighting the left scapulocoracoidin lateral view (a), thoracic vertebra 6 andlumbar vertebra 7 in anteriorand dorsal views (missingparts mirrored andrendered as semi-transparent) (b), theleft femurin distaland anterior views(c),the left humerus in anteriorand distalviews (d), the left astragalusand navicular inanterior view,and left calcaneusin medial and dorsal views (e),left hind foot indorsal view (f),and theleft tibia inlateral and anterior views(g). L,lumbar vertebra;T,thoracic vertebra.Scale bars,5 cm (main skeleton),1cm (ad, f, g), 5mm (e)." figureDoi="http://doi.org/10.5281/zenodo.3784159" httpUri="https://zenodo.org/record/3784159/files/figure.png" pageId="3" pageNumber="4">Fig.3</figureCitation>
,Extended Data
<figureCitation box="[1381,1468,994,1014]" captionStart="Extended" captionStartId="17.[106,195,1160,1178]" captionTargetBox="[135,1471,133,1135]" captionTargetId="figure@17.[131,1472,131,1136]" captionText="ExtendedData Fig.9 |Pectoraland pelvic girdle elements of A.hui holotype(UA9030).ad, ΜCT images.a, b, Left scapulacoracoid,left andrightclavicle and manubrium intop(a)and bottom (b) views(as preserved).c, d, Leftos coxa and epipubic bone inlateral (c)and medial (d) views." figureDoi="http://doi.org/10.5281/zenodo.3784179" httpUri="https://zenodo.org/record/3784179/files/figure.png" pageId="3" pageNumber="4">Fig.9c, d</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="6" lastPageNumber="7" pageId="3" pageNumber="4" type="discussion">
<paragraph blockId="3.[816,1498,1071,1644]" box="[816,1153,1071,1096]" pageId="3" pageNumber="4">
<heading bold="true" box="[816,1153,1071,1096]" fontSize="10" level="4" pageId="3" pageNumber="4" reason="0">
<emphasis bold="true" box="[816,1153,1071,1096]" pageId="3" pageNumber="4">Phylogenetic relationships</emphasis>
</heading>
</paragraph>
<paragraph blockId="3.[816,1498,1071,1644]" pageId="3" pageNumber="4">
Our phylogenetic analysis, performed using 84 cynodont taxa and 530 morphological characters, places
<taxonomicName box="[1197,1335,1138,1157]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[1197,1335,1138,1157]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
within Gondwanatheria, which in turn is placed within Allotheria as the sister taxon to
<taxonomicName box="[842,1013,1194,1214]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="order">Multituberculata</taxonomicName>
(Extended Data
<figureCitation box="[1175,1242,1194,1214]" captionStart="Extended" captionStartId="18.[106,195,1761,1779]" captionTargetBox="[193,1407,135,1734]" captionTargetId="figure@18.[192,1411,131,1737]" captionText="ExtendedData Fig.10 | Phylogenetic relationshipsof A. huiand selected mammaliaforms.Strict consensus treeof 16 equallyparsimonious trees(tree length = 2,315,consistencyindex =0.3015 and retentionindex = 0.7001) derived from analysis of 84 cynodont taxaand 530characters,withmultistate characters unorderedand unweighted.Bremer values arelisted next tothe nodes.Adalatheriumis highlighted inred.Allotheria—consisting of Cifelliodon, Euharamiyida,Gondwanatheria (including Adalatherium)and Multituberculata—is highlighted in blue.Taxon andcharacter lists,the data matrix,limitations andassumptions,phylogeneticmethods and amore detailed explanation of theresults are providedin theSupplementary Information." figureDoi="http://doi.org/10.5281/zenodo.3784181" httpUri="https://zenodo.org/record/3784181/files/figure.png" pageId="3" pageNumber="4">Fig. 10</figureCitation>
,Supplementary Information).
<taxonomicName box="[874,1021,1223,1243]" family="Adalatheriidae" pageId="3" pageNumber="1" rank="family">Adalatheriidae</taxonomicName>
(as solely represented by
<taxonomicName box="[1268,1404,1224,1243]" genus="Adalatherium" pageId="3" pageNumber="4" rank="genus">
<emphasis box="[1268,1404,1224,1243]" italics="true" pageId="3" pageNumber="4">Adalatherium</emphasis>
</taxonomicName>
) is recovered as more derived than
<taxonomicName authorityName="Bonaparte" authorityYear="1986" box="[1075,1251,1252,1272]" class="Mammalia" family="Ferugliotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Ferugliotheriidae</taxonomicName>
and stemward relative to
<taxonomicName authorityName="Scillato-Yane &amp; Pascual" authorityYear="1984" box="[816,971,1280,1300]" class="Mammalia" family="Sudamericidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Sudamericidae</taxonomicName>
.
</paragraph>
<paragraph blockId="3.[816,1498,1071,1644]" pageId="3" pageNumber="4">
Previous phylogenetic analyses that include the recently discovered Early Cretaceous purported haramiyidan
<emphasis box="[1245,1358,1339,1358]" italics="true" pageId="3" pageNumber="4">Cifelliodon</emphasis>
<superScript attach="left" box="[1358,1392,1333,1346]" fontSize="5" pageId="3" pageNumber="4">
<bibRefCitation author="Huttenlocker, A. K. &amp; Grossnickle, D. M. &amp; Kirkland, J. I. &amp; Schultz, J. A. &amp; Luo, Z. - X." box="[1358,1373,1333,1346]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="108 - 112" part="558" refId="ref5276" refString="23. Huttenlocker, A. K., Grossnickle, D. M., Kirkland, J. I., Schultz, J. A. &amp; Luo, Z. - X. Late-surviving stem mammal links the lowermost Cretaceous of North America and Gondwana. Nature 558, 108 - 112 (2018)." title="Late-surviving stem mammal links the lowermost Cretaceous of North America and Gondwana" type="journal article" year="2018">23</bibRefCitation>
,
<bibRefCitation author="King, B. &amp; Beck, R." box="[1377,1392,1333,1346]" journalOrPublisher="stratigraphic fit and the early evolution of mammals. Preprint at bioRxiv" pageId="3" pageNumber="4" refId="ref5336" refString="24. King, B. &amp; Beck, R. Bayesian tip-dated phylogenetics: topological effects, stratigraphic fit and the early evolution of mammals. Preprint at bioRxiv https: // doi. org / 10.1101 / 533885 (2019)." title="Bayesian tip-dated phylogenetics: topological effects" type="book" year="2019">24</bibRefCitation>
</superScript>
advanced the idea that gondwanatherians are nested within Eleutherodontida, basal to the purported Early Cretaceous hahnodontids
<taxonomicName authorityName="Sigogneau-Russell" authorityYear="1991" box="[1375,1482,1396,1415]" class="Mammalia" family="Hahnodontidae" genus="Hahnodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[1375,1482,1396,1415]" italics="true" pageId="3" pageNumber="4">Hahnodon</emphasis>
</taxonomicName>
<bibRefCitation author="Sigogneau-Russell, D." box="[1482,1496,1390,1403]" journalOrPublisher="Neues Jahrb. Geol. Palaontol. Monatsh." pageId="3" pageNumber="4" pagination="119 - 125" part="2" refId="ref5380" refString="25. Sigogneau-Russell, D. First evidence of Multituberculata (Mammalia) in the Mesozoic of Africa. Neues Jahrb. Geol. Palaontol. Monatsh. 2, 119 - 125 (1991)." title="First evidence of Multituberculata (Mammalia) in the Mesozoic of Africa" type="journal article" year="1991">
<superScript attach="left" box="[1482,1496,1390,1403]" fontSize="5" pageId="3" pageNumber="4">25</superScript>
</bibRefCitation>
and
<emphasis box="[857,964,1425,1444]" italics="true" pageId="3" pageNumber="4">Cifelliodon</emphasis>
<bibRefCitation author="Huttenlocker, A. K. &amp; Grossnickle, D. M. &amp; Kirkland, J. I. &amp; Schultz, J. A. &amp; Luo, Z. - X." box="[964,978,1419,1432]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="108 - 112" part="558" refId="ref5276" refString="23. Huttenlocker, A. K., Grossnickle, D. M., Kirkland, J. I., Schultz, J. A. &amp; Luo, Z. - X. Late-surviving stem mammal links the lowermost Cretaceous of North America and Gondwana. Nature 558, 108 - 112 (2018)." title="Late-surviving stem mammal links the lowermost Cretaceous of North America and Gondwana" type="journal article" year="2018">
<superScript attach="left" box="[964,978,1419,1432]" fontSize="5" pageId="3" pageNumber="4">23</superScript>
</bibRefCitation>
. Although
<taxonomicName authorityName="Sigogneau-Russell" authorityYear="1991" box="[1083,1187,1425,1444]" class="Mammalia" family="Hahnodontidae" genus="Hahnodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[1083,1187,1425,1444]" italics="true" pageId="3" pageNumber="4">Hahnodon</emphasis>
</taxonomicName>
was not included in our analysis because it is represented by only one (or possibly two)isolated teeth
<superScript attach="left" box="[1460,1491,1448,1461]" fontSize="5" pageId="3" pageNumber="4">
<bibRefCitation author="Sigogneau-Russell, D." box="[1460,1473,1448,1461]" journalOrPublisher="Neues Jahrb. Geol. Palaontol. Monatsh." pageId="3" pageNumber="4" pagination="119 - 125" part="2" refId="ref5380" refString="25. Sigogneau-Russell, D. First evidence of Multituberculata (Mammalia) in the Mesozoic of Africa. Neues Jahrb. Geol. Palaontol. Monatsh. 2, 119 - 125 (1991)." title="First evidence of Multituberculata (Mammalia) in the Mesozoic of Africa" type="journal article" year="1991">25</bibRefCitation>
,
<bibRefCitation author="Butler, P. M. &amp; Hooker, J. J." box="[1477,1491,1448,1461]" journalOrPublisher="Acta Palaeontol. Pol." pageId="3" pageNumber="4" pagination="185 - 207" part="50" refId="ref5417" refString="26. Butler, P. M. &amp; Hooker, J. J. New teeth of allotherian mammals from the English Bathonian, including the earliest multituberculates. Acta Palaeontol. Pol. 50, 185 - 207 (2005)." title="New teeth of allotherian mammals from the English Bathonian, including the earliest multituberculates" type="journal article" year="2005">26</bibRefCitation>
</superScript>
,
<emphasis box="[816,925,1482,1501]" italics="true" pageId="3" pageNumber="4">Cifelliodon</emphasis>
is recovered at the base of Allotheria, which also includes Euharamiyida and
<taxonomicName box="[1002,1176,1510,1530]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="order">Multituberculata</taxonomicName>
+ Gondwanatheria.Our analysis places the haramiyidans
<taxonomicName authorityName="Jenkins, Gatesy, Shubin &amp; Amaral" authorityYear="1997" box="[1062,1192,1540,1558]" class="Mammalia" family="Haramiyidae" genus="Haramiyavia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[1062,1192,1540,1558]" italics="true" pageId="3" pageNumber="4">Haramiyavia</emphasis>
</taxonomicName>
and
<taxonomicName box="[1239,1338,1539,1558]" class="Magnoliopsida" family="Malvaceae" genus="Thomasia" kingdom="Animalia" order="Malvales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
<emphasis box="[1239,1338,1539,1558]" italics="true" pageId="3" pageNumber="4">Thomasia</emphasis>
</taxonomicName>
—along with the poorly known taxon
<taxonomicName box="[1014,1124,1569,1587]" class="Mammalia" family="Eleutherodontidae" genus="Megaconus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haramiyida" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[1014,1124,1569,1587]" italics="true" pageId="3" pageNumber="4">Megaconus</emphasis>
</taxonomicName>
—outside of Mammaliaformes,with no close relations to allotherians. This finding is in contrast to previous analyses for
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[939,1019,1626,1644]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[939,1019,1626,1644]" italics="true" pageId="3" pageNumber="4">Vintana</emphasis>
</taxonomicName>
<bibRefCitation author="Krause, D. W." box="[1019,1026,1620,1633]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="512 - 517" part="515" refId="ref4435" refString="3. Krause, D. W. et al. First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism. Nature 515, 512 - 517 (2014)." title="First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism" type="journal article" year="2014">
<superScript attach="left" box="[1019,1026,1620,1633]" fontSize="5" pageId="3" pageNumber="4">3</superScript>
</bibRefCitation>
and for Jurassic euharamiyidans
<superScript attach="left" box="[1348,1382,1620,1633]" fontSize="5" pageId="3" pageNumber="4">
<bibRefCitation author="Han, G. &amp; Mao, F. &amp; Bi, S. &amp; Wang, Y. &amp; Meng, J." box="[1348,1361,1620,1633]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="451 - 456" part="551" refId="ref5461" refString="27. Han, G., Mao, F., Bi, S., Wang, Y. &amp; Meng, J. A Jurassic gliding euharamiyidan mammal with an ear of five auditory bones. Nature 551, 451 - 456 (2017)." title="A Jurassic gliding euharamiyidan mammal with an ear of five auditory bones" type="journal article" year="2017">27</bibRefCitation>
<bibRefCitation author="Bi, S. &amp; Wang, Y. &amp; Guan, J. &amp; Sheng, X. &amp; Meng, J." box="[1368,1382,1620,1633]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="579 - 584" part="514" refId="ref5556" refString="29. Bi, S., Wang, Y., Guan, J., Sheng, X. &amp; Meng, J. Three new Jurassic euharamiyidan species reinforce early divergence of mammals. Nature 514, 579 - 584 (2014)." title="Three new Jurassic euharamiyidan species reinforce early divergence of mammals" type="journal article" year="2014">29</bibRefCitation>
</superScript>
.
</paragraph>
<paragraph blockId="3.[816,1499,1701,1988]" box="[816,1083,1701,1726]" pageId="3" pageNumber="4">
<heading bold="true" box="[816,1083,1701,1726]" fontSize="10" level="4" pageId="3" pageNumber="4" reason="0">
<emphasis bold="true" box="[816,1083,1701,1726]" pageId="3" pageNumber="4">Evolution in isolation</emphasis>
</heading>
</paragraph>
<paragraph blockId="3.[816,1499,1701,1988]" lastBlockId="4.[106,787,1734,1988]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
Amongmammals,themostobviousandquantifiableinfluencesofevolving on islands are those related to body size.This observation has led to the articulation of the island rule,which states that—evolutionar- ily—small mammals on islands increase,and large mammals decrease,in size
<superScript attach="left" box="[853,882,1849,1862]" fontSize="5" pageId="3" pageNumber="4">
<bibRefCitation author="Lomolino, M. V." box="[853,863,1849,1862]" journalOrPublisher="J. Biogeogr." pageId="3" pageNumber="4" pagination="1427 - 1439" part="40" refId="ref4733" refString="11. Lomolino, M. V. Of mice and mammoths: generality and antiquity of the island rule. J. Biogeogr. 40, 1427 - 1439 (2013)." title="Of mice and mammoths: generality and antiquity of the island rule" type="journal article" year="2013">11</bibRefCitation>
,
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</superScript>
.Inaddition,evolutionininsularenvironmentsisthoughttoresult in changes in anatomy,physiology,behaviour and life-history strategies, and (at the faunal level)relatively low species richness,taxonomic imbalance,high endemism and a general level of primitiveness
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<bibRefCitation author="Sondaar, P. Y." box="[1373,1380,1935,1948]" editor="Hecht, M. K." journalOrPublisher="Plenum" pageId="3" pageNumber="4" pagination="671 - 707" refId="ref4551" refString="6. Sondaar, P. Y. in Major Patterns in Vertebrate Evolution (eds Hecht, M. K. et al.) 671 - 707 (Plenum, 1977)." type="book chapter" volumeTitle="Major Patterns in Vertebrate Evolution" year="1977">6</bibRefCitation>
,
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</superScript>
. Although somewhat controversial and clearly not ubiquitous
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</bibRefCitation>
, the island rule is generally established as a pervasive pattern
<superScript attach="left" box="[541,580,1734,1747]" fontSize="5" pageId="4" pageNumber="5">
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,
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,
<bibRefCitation author="McClain, C. R. &amp; Durst, P. A. P. &amp; Boyer, A. G. &amp; Francis, C. D." box="[566,580,1734,1747]" journalOrPublisher="Biol. Lett." pageId="4" pageNumber="5" pagination="20120989" part="9" refId="ref5672" refString="32. McClain, C. R., Durst, P. A. P., Boyer, A. G. &amp; Francis, C. D. Unravelling the determinants of insular body size shifts. Biol. Lett. 9, 20120989 (2013)." title="Unravelling the determinants of insular body size shifts" type="journal article" year="2013">32</bibRefCitation>
</superScript>
. Examples of insular dwarfism and gigantism from Pliocene, Pleistocene and Holocene epochs abound
<superScript attach="left" box="[260,304,1792,1805]" fontSize="5" pageId="4" pageNumber="5">
<bibRefCitation author="Van der Geer, A. &amp; Lyras, G. &amp; de Vos, J. &amp; Dermitzakis, M." box="[260,274,1792,1805]" journalOrPublisher="Wiley-Blackwell" pageId="4" pageNumber="5" refId="ref4690" refString="10. Van der Geer, A., Lyras, G., de Vos, J. &amp; Dermitzakis, M. Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands (Wiley-Blackwell, 2010)." title="Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands" type="book" year="2010">10</bibRefCitation>
,
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,
<bibRefCitation author="De Vos, J. &amp; van den Hoek Ostende, L. W. &amp; van den Bergh, G. D." box="[292,304,1792,1805]" editor="Renema, W." journalOrPublisher="Springer" pageId="4" pageNumber="5" pagination="315 - 346" refId="ref4874" refString="15. De Vos, J., van den Hoek Ostende, L. W. &amp; van den Bergh, G. D. in Biogeography, Time and Place: Distributions, Barriers and Islands (ed. Renema, W.) 315 - 346 (Springer, 2007)." title="Biogeography, Time and Place: Distributions, Barriers and Islands" type="book chapter" year="2007">15</bibRefCitation>
</superScript>
, including from Madagascar (pygmy hippopotamuses
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andgiantlemurs
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<superScript attach="left" box="[339,353,1820,1833]" fontSize="5" pageId="4" pageNumber="5">34</superScript>
</bibRefCitation>
). ExamplesfromearlierintheCenozoiceraare relatively sparse
<superScript attach="left" box="[260,303,1849,1862]" fontSize="5" pageId="4" pageNumber="5">
<bibRefCitation author="Unay, E. &amp; De Bruijn, H. &amp; Sarac, G. The" box="[260,272,1849,1862]" journalOrPublisher="Deinsea" pageId="4" pageNumber="5" pagination="531 - 537" part="10" refId="ref4837" refString="14. Unay, E., De Bruijn, H. &amp; Sarac, G. The Oligocene rodent record of Anatolia: a review. Deinsea 10, 531 - 537 (2003)." title="Oligocene rodent record of Anatolia: a review" type="journal article" year="2003">14</bibRefCitation>
,
<bibRefCitation author="Worthy, T. H." box="[276,288,1849,1862]" journalOrPublisher="Proc. Natl Acad. Sci. USA" pageId="4" pageNumber="5" pagination="19419 - 19423" part="103" refId="ref4929" refString="16. Worthy, T. H. et al. Miocene mammal reveals a Mesozoic ghost lineage on insular New Zealand, southwest Pacific. Proc. Natl Acad. Sci. USA 103, 19419 - 19423 (2006)." title="Miocene mammal reveals a Mesozoic ghost lineage on insular New Zealand, southwest Pacific" type="journal article" year="2006">16</bibRefCitation>
,
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</superScript>
and the effects of long-term isolation are extremely poorly documented for Mesozoic mammaliaforms.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3784159" ID-Zenodo-Dep="3784159" httpUri="https://zenodo.org/record/3784159/files/figure.png" pageId="4" pageNumber="5" startId="4.[106,137,1572,1590]" targetBox="[138,1476,134,1542]" targetPageId="4">
<paragraph blockId="4.[106,786,1571,1696]" lastBlockId="4.[816,1492,1572,1670]" pageId="4" pageNumber="5">
<emphasis bold="true" box="[106,543,1571,1590]" pageId="4" pageNumber="5">
Fig.3 |Skeleton of
<taxonomicName authorityName="Krause, Hoffmann, Wible, and Rougier" authorityYear="2020" box="[270,321,1572,1590]" genus="Adalatherium" pageId="4" pageNumber="5" rank="species" species="hui">
<emphasis bold="true" box="[270,321,1572,1590]" italics="true" pageId="4" pageNumber="5">A.hui</emphasis>
</taxonomicName>
holotype (UA 9030). ag
</emphasis>
, Digitally reconstructed skeleton in left lateral view,highlighting the left scapulocoracoid in lateral view (
<emphasis bold="true" box="[112,122,1625,1643]" pageId="4" pageNumber="5">a</emphasis>
), thoracic vertebra 6 and lumbar vertebra 7 in anterior and dorsal views (missing parts mirrored and rendered as semi-transparent) (
<emphasis bold="true" box="[623,635,1652,1670]" pageId="4" pageNumber="5">b</emphasis>
), the left femur in distal and anterior views(
<emphasis bold="true" box="[326,335,1678,1696]" pageId="4" pageNumber="5">c</emphasis>
),the left humerus in anterior and distal views (
<emphasis bold="true" box="[734,746,1678,1696]" pageId="4" pageNumber="5">d</emphasis>
), the left astragalus and navicular in anterior view,and left calcaneus in medial and dorsal views (
<emphasis bold="true" box="[968,978,1598,1616]" pageId="4" pageNumber="5">e</emphasis>
),left hind foot in dorsal view (
<emphasis bold="true" box="[1235,1242,1598,1616]" pageId="4" pageNumber="5">f</emphasis>
),and the left tibia in lateral and anterior views(
<emphasis bold="true" box="[986,997,1625,1643]" pageId="4" pageNumber="5">g</emphasis>
). L,lumbar vertebra;T,thoracic vertebra.Scale bars,5 cm (main skeleton),1cm (
<emphasis bold="true" box="[1004,1038,1652,1670]" pageId="4" pageNumber="5">ad</emphasis>
,
<emphasis bold="true" box="[1046,1053,1652,1670]" pageId="4" pageNumber="5">f</emphasis>
,
<emphasis bold="true" box="[1061,1072,1652,1670]" pageId="4" pageNumber="5">g</emphasis>
), 5mm (
<emphasis bold="true" box="[1143,1153,1652,1670]" pageId="4" pageNumber="5">e</emphasis>
).
</paragraph>
</caption>
<paragraph blockId="4.[106,787,1734,1988]" lastBlockId="4.[816,1499,1734,1988]" pageId="4" pageNumber="5">
Among Mesozoic mammaliaforms,adaptations related to evolution in isolation have been most notably claimed for two island environments,both from the latest part of the Cretaceous period(Maastrichtian age): (1) the gondwanatherian
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[1118,1198,1741,1759]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1118,1198,1741,1759]" italics="true" pageId="4" pageNumber="5">Vintana</emphasis>
</taxonomicName>
from Madagascar
<bibRefCitation author="Krause, D. W." box="[1378,1385,1734,1747]" journalOrPublisher="Nature" pageId="4" pageNumber="5" pagination="512 - 517" part="515" refId="ref4435" refString="3. Krause, D. W. et al. First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism. Nature 515, 512 - 517 (2014)." title="First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism" type="journal article" year="2014">
<superScript attach="left" box="[1378,1385,1734,1747]" fontSize="5" pageId="4" pageNumber="5">3</superScript>
</bibRefCitation>
and (2) the multituberculates
<taxonomicName box="[1008,1132,1769,1788]" class="Mammalia" family="Kogaionidae" genus="Barbatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1008,1132,1769,1788]" italics="true" pageId="4" pageNumber="5">Barbatodon</emphasis>
</taxonomicName>
and
<emphasis box="[1179,1250,1769,1787]" italics="true" pageId="4" pageNumber="5">Litovoi</emphasis>
from the archaic Hațeg Island
<superScript attach="left" box="[884,918,1792,1805]" fontSize="5" pageId="4" pageNumber="5">
35,
<bibRefCitation author="Csiki-Sava, Z. &amp; Vremir, M. &amp; Meng, J. &amp; Brusatte, S. L. &amp; Norell, M. A." box="[902,918,1792,1805]" journalOrPublisher="Proc. Natl Acad. Sci. USA" pageId="4" pageNumber="5" pagination="4857 - 4862" part="115" refId="ref5855" refString="36. Csiki-Sava, Z., Vremir, M., Meng, J., Brusatte, S. L. &amp; Norell, M. A. Dome-headed, small-brained island mammal from the Late Cretaceous of Romania. Proc. Natl Acad. Sci. USA 115, 4857 - 4862 (2018)." title="Dome-headed, small-brained island mammal from the Late Cretaceous of Romania" type="journal article" year="2018">36</bibRefCitation>
</superScript>
(now part of Romania). Whether
<taxonomicName box="[1256,1379,1797,1816]" class="Mammalia" family="Kogaionidae" genus="Barbatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1256,1379,1797,1816]" italics="true" pageId="4" pageNumber="5">Barbatodon</emphasis>
</taxonomicName>
and
<emphasis box="[1426,1497,1798,1816]" italics="true" pageId="4" pageNumber="5">Litovoi</emphasis>
were part of a fauna that developed unique adaptations attributable to evolution in an insular environment is questionable—as is whether Hațeg Island was even an island (Supplementary Information).Because of its completeness and undoubted existence in an insular environment, the skeleton of
<taxonomicName box="[975,1118,1941,1960]" genus="Adalatherium" pageId="4" pageNumber="5" rank="genus">
<emphasis box="[975,1118,1941,1960]" italics="true" pageId="4" pageNumber="5">Adalatherium</emphasis>
</taxonomicName>
provides an opportunity to examine evolution in isolation among Mesozoic mammaliaforms.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3784161" ID-Zenodo-Dep="3784161" httpUri="https://zenodo.org/record/3784161/files/figure.png" pageId="5" pageNumber="6" startId="5.[106,137,1456,1474]" subCaptionStartIDs="5.[1299,1346,1562,1580]" subCaptionStarts="Maps " targetBox="[119,1483,135,1429]" targetPageId="5">
<paragraph blockId="5.[106,782,1455,1634]" lastBlockId="5.[816,1467,1455,1607]" pageId="5" pageNumber="6">
<emphasis bold="true" box="[106,638,1456,1474]" pageId="5" pageNumber="6">Fig.4 |Key stages in plate tectonic history of Madagascar.a</emphasis>
, Position of Madagascar before rifting between West Gondwana (South America and Africa)and East Gondwana (Madagascar,Seychelles,Indian subcontinent, Sri Lanka,Antarctica and Australia) at183 Myr ago (Early Jurassic epoch).
<emphasis bold="true" box="[106,118,1562,1580]" pageId="5" pageNumber="6">b</emphasis>
,Separation of Indo-Madagascar from Antarctica and Australia at 124Myr ago (mid-Early Cretaceous epoch).
<emphasis bold="true" box="[371,381,1589,1607]" pageId="5" pageNumber="6">c</emphasis>
, Separation of Indian subcontinent from Madagascar at 88 Myr ago (mid-Late Cretaceous epoch).
<emphasis bold="true" box="[592,604,1616,1634]" pageId="5" pageNumber="6">d</emphasis>
, Approximate time of deposition of Maevarano Formation at 66 Myr ago (latest Cretaceous period).Solid black lines indicate current coastlines of Madagascar and east Africa;brown represents Precambrian terranes;and yellow indicates sedimentary basins along west coast of Madagascar.The discovery site of UA 9030 is indicated by red star in
<emphasis bold="true" box="[1110,1122,1562,1580]" pageId="5" pageNumber="6">d</emphasis>
.Scale bars,500 km.Maps adapted from Earthworks (www.reeves.nl/gond.com).
</paragraph>
</caption>
<paragraph blockId="5.[106,787,1681,1988]" pageId="5" pageNumber="6">
Madagascar separated from the Indian subcontinent and the Seychelles about 88 Myr ago
<superScript attach="left" box="[355,386,1706,1719]" fontSize="5" pageId="5" pageNumber="6">
<bibRefCitation author="Reeves, C." box="[355,367,1706,1719]" journalOrPublisher="J. Afr. Earth Sci." pageId="5" pageNumber="6" pagination="45 - 57" part="94" refId="ref5914" refString="37. Reeves, C. The position of Madagascar within Gondwana and its movements during Gondwana dispersal. J. Afr. Earth Sci. 94, 45 - 57 (2014)." title="The position of Madagascar within Gondwana and its movements during Gondwana dispersal" type="journal article" year="2014">37</bibRefCitation>
,
<bibRefCitation author="Krause, D. W. &amp; Sertich, J. J. W. &amp; O'Connor, P. M. &amp; Curry Rogers, K. &amp; Rogers, R. R. The" box="[371,386,1706,1719]" journalOrPublisher="Annu. Rev. Earth Planet. Sci." pageId="5" pageNumber="6" pagination="519 - 553" part="47" refId="ref5949" refString="38. Krause, D. W., Sertich, J. J. W., O'Connor, P. M., Curry Rogers, K. &amp; Rogers, R. R. The Mesozoic biogeographic history of Gondwanan terrestrial vertebrates: insights from Madagascar's fossil record. Annu. Rev. Earth Planet. Sci. 47, 519 - 553 (2019)." title="Mesozoic biogeographic history of Gondwanan terrestrial vertebrates: insights from Madagascar's fossil record" type="journal article" year="2019">38</bibRefCitation>
</superScript>
(
<figureCitation box="[397,452,1710,1730]" captionStart="Fig" captionStartId="5.[106,137,1456,1474]" captionTargetBox="[119,1483,135,1429]" captionTargetId="graphics@5.[170,1432,811,1406]" captionTargetPageId="5" captionText="Fig.4 |Keystages in platetectonic historyof Madagascar.a, Position of Madagascar before rifting betweenWestGondwana (South America and Africa)and EastGondwana (Madagascar,Seychelles,Indiansubcontinent, Sri Lanka,Antarctica and Australia) at183 Myr ago (Early Jurassic epoch). b,Separation of Indo-Madagascarfrom Antarctica andAustralia at 124Myr ago (mid-Early Cretaceous epoch).c, Separation of Indiansubcontinent from Madagascar at 88 Myr ago (mid-Late Cretaceous epoch).d, Approximate time of deposition of MaevaranoFormation at 66 Myrago (latest Cretaceous period).Solid blacklines indicate current coastlines of Madagascar andeast Africa;brown representsPrecambrian terranes;andyellow indicates sedimentary basins along west coast of Madagascar.The discovery siteof UA 9030 isindicated by red star ind.Scale bars,500 km.Maps adapted from Earthworks (www.reeves.nl/gond.com)." figureDoi="http://doi.org/10.5281/zenodo.3784161" httpUri="https://zenodo.org/record/3784161/files/figure.png" pageId="5" pageNumber="6">Fig. 4</figureCitation>
). As a result,after separation, the obligate terrestrial taxa of Madagascar evolved in complete isolation and the only taxa that gained access to the island subsequently were flying, swimming or rafting forms that were able to disperse across considerable marine barriers
<superScript attach="left" box="[396,432,1820,1833]" fontSize="5" pageId="5" pageNumber="6">
<bibRefCitation author="Krause, D. W. &amp; Sertich, J. J. W. &amp; O'Connor, P. M. &amp; Curry Rogers, K. &amp; Rogers, R. R. The" box="[396,411,1820,1833]" journalOrPublisher="Annu. Rev. Earth Planet. Sci." pageId="5" pageNumber="6" pagination="519 - 553" part="47" refId="ref5949" refString="38. Krause, D. W., Sertich, J. J. W., O'Connor, P. M., Curry Rogers, K. &amp; Rogers, R. R. The Mesozoic biogeographic history of Gondwanan terrestrial vertebrates: insights from Madagascar's fossil record. Annu. Rev. Earth Planet. Sci. 47, 519 - 553 (2019)." title="Mesozoic biogeographic history of Gondwanan terrestrial vertebrates: insights from Madagascar's fossil record" type="journal article" year="2019">38</bibRefCitation>
<bibRefCitation author="Samonds, K. E." box="[419,432,1820,1833]" journalOrPublisher="PLoS ONE" pageId="5" pageNumber="6" pagination="62086" part="8" refId="ref6093" refString="41. Samonds, K. E. et al. Imperfect isolation: factors and filters shaping Madagascar's extant vertebrate fauna. PLoS ONE 8, e 62086 (2013)." title="Imperfect isolation: factors and filters shaping Madagascar's extant vertebrate fauna" type="journal article" year="2013">41</bibRefCitation>
</superScript>
. Madagascar remains today a large, isolated continental block that is topographically high, geotectonically stable and at a minimum distance of 430 km from the closest mainland (Africa).
</paragraph>
<paragraph blockId="5.[106,787,1681,1988]" lastBlockId="6.[106,787,134,1358]" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
The postcranial skeleton of UA9030indicates that
<taxonomicName box="[607,740,1941,1960]" genus="Adalatherium" pageId="5" pageNumber="6" rank="genus">
<emphasis box="[607,740,1941,1960]" italics="true" pageId="5" pageNumber="6">Adalatherium</emphasis>
</taxonomicName>
(and perhaps other gondwanatherians) was neither volant nor aquatic;it was an obligate terrestrial form that was relatively less capable of dispersal across marine barriers and more likely to have evolved on Madagascar. There are at least two other gondwanatherians that lived on Madagascar contemporaneously with
<taxonomicName box="[1112,1252,1769,1788]" genus="Adalatherium" pageId="5" pageNumber="6" rank="genus">
<emphasis box="[1112,1252,1769,1788]" italics="true" pageId="5" pageNumber="6">Adalatherium</emphasis>
</taxonomicName>
:
<taxonomicName box="[1262,1350,1769,1788]" class="Mammalia" family="Sudamericidae" genus="Lavanify" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1262,1350,1769,1788]" italics="true" pageId="5" pageNumber="6">Lavanify</emphasis>
</taxonomicName>
(based on two fragmentary isolated teeth
<bibRefCitation author="Krause, D. W. &amp; Prasad, G. V. R. &amp; Koenigswald, W. V. &amp; Sahni, A. &amp; Grine, F. E." box="[1084,1098,1792,1805]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="504 - 507" part="390" refId="ref6126" refString="42. Krause, D. W., Prasad, G. V. R., Koenigswald, W. V., Sahni, A. &amp; Grine, F. E. Cosmopolitanism among Gondwanan Late Cretaceous mammals. Nature 390, 504 - 507 (1997)." title="Cosmopolitanism among Gondwanan Late Cretaceous mammals" type="journal article" year="1997">
<superScript attach="left" box="[1084,1098,1792,1805]" fontSize="5" pageId="5" pageNumber="6">42</superScript>
</bibRefCitation>
that are insufficient to be informative in the current context) and
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[1070,1152,1827,1845]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1070,1152,1827,1845]" italics="true" pageId="5" pageNumber="6">Vintana</emphasis>
</taxonomicName>
(based on a complete cranium
<superScript attach="left" box="[1464,1483,1820,1833]" fontSize="5" pageId="5" pageNumber="6">
<bibRefCitation author="Krause, D. W." box="[1464,1471,1820,1833]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="512 - 517" part="515" refId="ref4435" refString="3. Krause, D. W. et al. First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism. Nature 515, 512 - 517 (2014)." title="First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism" type="journal article" year="2014">3</bibRefCitation>
,
<bibRefCitation author="Krause, D. W." box="[1475,1483,1820,1833]" journalOrPublisher="J. Vertebr. Paleontol." pageId="5" pageNumber="6" pagination="6" part="34" refId="ref4467" refString="4. Krause, D. W. (ed.) Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar. J. Vertebr. Paleontol. 34, suppl. to issue 6 (2014)." title="Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar" type="journal article" year="2014">4</bibRefCitation>
</superScript>
). The cranium and upper postcanine dentition of
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[1297,1378,1855,1873]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1297,1378,1855,1873]" italics="true" pageId="5" pageNumber="6">Vintana</emphasis>
</taxonomicName>
exhibit several features that are unknown among Mesozoic mammaliaforms,but the number of such features in
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[1133,1216,1913,1931]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1133,1216,1913,1931]" italics="true" pageId="5" pageNumber="6">Vintana</emphasis>
</taxonomicName>
are far fewer than those for
<taxonomicName box="[816,954,1941,1960]" genus="Adalatherium" pageId="5" pageNumber="6" rank="genus">
<emphasis box="[816,954,1941,1960]" italics="true" pageId="5" pageNumber="6">Adalatherium</emphasis>
</taxonomicName>
(based on a complete skeleton).Furthermore,given its deeply nested phylogenetic position within Allotheria (Extended Data
<figureCitation box="[106,177,134,154]" captionStart="Extended" captionStartId="18.[106,195,1761,1779]" captionTargetBox="[193,1407,135,1734]" captionTargetId="figure@18.[192,1411,131,1737]" captionText="ExtendedData Fig.10 | Phylogenetic relationshipsof A. huiand selected mammaliaforms.Strict consensus treeof 16 equallyparsimonious trees(tree length = 2,315,consistencyindex =0.3015 and retentionindex = 0.7001) derived from analysis of 84 cynodont taxaand 530characters,withmultistate characters unorderedand unweighted.Bremer values arelisted next tothe nodes.Adalatheriumis highlighted inred.Allotheria—consisting of Cifelliodon, Euharamiyida,Gondwanatheria (including Adalatherium)and Multituberculata—is highlighted in blue.Taxon andcharacter lists,the data matrix,limitations andassumptions,phylogeneticmethods and amore detailed explanation of theresults are providedin theSupplementary Information." figureDoi="http://doi.org/10.5281/zenodo.3784181" httpUri="https://zenodo.org/record/3784181/files/figure.png" pageId="6" pageNumber="7">Fig. 10</figureCitation>
), several additional features of
<taxonomicName box="[489,629,135,154]" genus="Adalatherium" pageId="6" pageNumber="7" rank="genus">
<emphasis box="[489,629,135,154]" italics="true" pageId="6" pageNumber="7">Adalatherium</emphasis>
</taxonomicName>
clearly evolved convergently with those of non-allotherian mammals—particularly therians (Supplementary Information). Considered together, and in concert with the palaeogeographical history of Madagascar,we hypothesize that the unusual morphological attributes of these two gondwanatherians are due to long-term evolution in isolation in an island environment, paralleling the cases adduced for various mammals on Mediterranean islands during the Neogene subperiod
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<bibRefCitation author="Sondaar, P. Y." box="[739,747,330,343]" editor="Hecht, M. K." journalOrPublisher="Plenum" pageId="6" pageNumber="7" pagination="671 - 707" refId="ref4551" refString="6. Sondaar, P. Y. in Major Patterns in Vertebrate Evolution (eds Hecht, M. K. et al.) 671 - 707 (Plenum, 1977)." type="book chapter" volumeTitle="Major Patterns in Vertebrate Evolution" year="1977">6</bibRefCitation>
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,
<bibRefCitation author="Van der Geer, A. &amp; Lyras, G. &amp; de Vos, J. &amp; Dermitzakis, M." box="[766,780,330,343]" journalOrPublisher="Wiley-Blackwell" pageId="6" pageNumber="7" refId="ref4690" refString="10. Van der Geer, A., Lyras, G., de Vos, J. &amp; Dermitzakis, M. Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands (Wiley-Blackwell, 2010)." title="Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands" type="book" year="2010">10</bibRefCitation>
</superScript>
. Furthermore,both
<taxonomicName box="[296,434,364,383]" genus="Adalatherium" pageId="6" pageNumber="7" rank="genus">
<emphasis box="[296,434,364,383]" italics="true" pageId="6" pageNumber="7">Adalatherium</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[478,558,365,383]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[478,558,365,383]" italics="true" pageId="6" pageNumber="7">Vintana</emphasis>
</taxonomicName>
are large,being among the largest mammaliaforms known from the entire Mesozoic era of Gondwana.
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[228,311,422,440]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[228,311,422,440]" italics="true" pageId="6" pageNumber="7">Vintana</emphasis>
</taxonomicName>
, which is even larger than
<taxonomicName box="[579,721,421,440]" genus="Adalatherium" pageId="6" pageNumber="7" rank="genus">
<emphasis box="[579,721,421,440]" italics="true" pageId="6" pageNumber="7">Adalatherium</emphasis>
</taxonomicName>
, is the second-largest known Mesozoic mammaliaform globally and the largest from Gondwana
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<bibRefCitation author="Krause, D. W." box="[304,311,473,486]" journalOrPublisher="Nature" pageId="6" pageNumber="7" pagination="512 - 517" part="515" refId="ref4435" refString="3. Krause, D. W. et al. First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism. Nature 515, 512 - 517 (2014)." title="First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism" type="journal article" year="2014">3</bibRefCitation>
,
<bibRefCitation author="Krause, D. W." box="[315,323,473,486]" journalOrPublisher="J. Vertebr. Paleontol." pageId="6" pageNumber="7" pagination="6" part="34" refId="ref4467" refString="4. Krause, D. W. (ed.) Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar. J. Vertebr. Paleontol. 34, suppl. to issue 6 (2014)." title="Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar" type="journal article" year="2014">4</bibRefCitation>
</superScript>
. Although the fossil record of Mesozoic mammaliaforms (especially from Gondwana) is too poor to conclusively establish that the large size of
<taxonomicName authorityName="Krause, Hoffmann, Wible, Kirk, Schultz, von Koenigswald, Groenke, Rossie, O'Connor, Seiffert, Dumont, Holloway, Rogers, Rahantarisoa, Kemp &amp; Andriamialison" authorityYear="2014" box="[392,471,537,555]" class="Mammalia" family="Sudamericidae" genus="Vintana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Multituberculata" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[392,471,537,555]" italics="true" pageId="6" pageNumber="7">Vintana</emphasis>
</taxonomicName>
and
<taxonomicName box="[513,648,536,555]" genus="Adalatherium" pageId="6" pageNumber="7" rank="genus">
<emphasis box="[513,648,536,555]" italics="true" pageId="6" pageNumber="7">Adalatherium</emphasis>
</taxonomicName>
reflects island gigantism,such a conclusion is consistent with the island rule
<superScript attach="left" box="[714,761,559,572]" fontSize="5" pageId="6" pageNumber="7">
<bibRefCitation author="Sondaar, P. Y." box="[714,722,559,572]" editor="Hecht, M. K." journalOrPublisher="Plenum" pageId="6" pageNumber="7" pagination="671 - 707" refId="ref4551" refString="6. Sondaar, P. Y. in Major Patterns in Vertebrate Evolution (eds Hecht, M. K. et al.) 671 - 707 (Plenum, 1977)." type="book chapter" volumeTitle="Major Patterns in Vertebrate Evolution" year="1977">6</bibRefCitation>
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,
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</superScript>
.
</paragraph>
<paragraph blockId="6.[106,787,134,1358]" pageId="6" pageNumber="7">
The vertebrate fauna associated with
<taxonomicName box="[488,623,593,612]" genus="Adalatherium" pageId="6" pageNumber="7" rank="genus">
<emphasis box="[488,623,593,612]" italics="true" pageId="6" pageNumber="7">Adalatherium</emphasis>
</taxonomicName>
(Supplementary Table 2) also exhibits unique characteristics reflective of its relictual nature.Forinstance,relativelyhighnumbersofderivedfeaturesareseen in other terrestrial members of the latest Cretaceous fauna of Madagascar,including the ceratophryid frog
<taxonomicName box="[451,555,708,727]" class="Amphibia" family="Ceratophryidae" genus="Beelzebufo" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[451,555,708,727]" italics="true" pageId="6" pageNumber="7">Beelzebufo</emphasis>
</taxonomicName>
,the crocodyliform
<taxonomicName class="Reptilia" family="Chimaerasuchidae" genus="Simosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="6" pageNumber="7">Simosuchus</emphasis>
</taxonomicName>
and the theropods
<taxonomicName authorityName="Lavocat" authorityYear="1955" box="[374,529,737,755]" class="Reptilia" family="Abelisauridae" genus="Majungasaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[374,529,737,755]" italics="true" pageId="6" pageNumber="7">Majungasaurus</emphasis>
</taxonomicName>
and
<taxonomicName box="[573,724,737,756]" class="Reptilia" family="Noasauridae" genus="Masiakasaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[573,724,737,756]" italics="true" pageId="6" pageNumber="7">Masiakasaurus</emphasis>
</taxonomicName>
(ref.
<bibRefCitation author="Krause, D. W. &amp; Sertich, J. J. W. &amp; O'Connor, P. M. &amp; Curry Rogers, K. &amp; Rogers, R. R. The" box="[771,786,731,744]" journalOrPublisher="Annu. Rev. Earth Planet. Sci." pageId="6" pageNumber="7" pagination="519 - 553" part="47" refId="ref5949" refString="38. Krause, D. W., Sertich, J. J. W., O'Connor, P. M., Curry Rogers, K. &amp; Rogers, R. R. The Mesozoic biogeographic history of Gondwanan terrestrial vertebrates: insights from Madagascar's fossil record. Annu. Rev. Earth Planet. Sci. 47, 519 - 553 (2019)." title="Mesozoic biogeographic history of Gondwanan terrestrial vertebrates: insights from Madagascar's fossil record" type="journal article" year="2019">
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and references therein).None of these genera is known from any other landmass,which again attests to high endemicity (although the very poor Mesozoic fossil record from Gondwana must be acknowledged in this regard).Furthermore,many of these forms had ghost lineages that extend back to or before theEarlyLate Cretaceous boundary (100 Myr ago)
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, thus suggesting that their ancestors had probably arrived on Indo-Madagascar before its separation from other Gondwanan landmasses(that is,via a deep-time vicariance event rather than overwater dispersal).Despite originating in different ways,we conclude that the latest Cretaceous insular vertebrate fauna of Madagascar was probably as unique relative to mainland faunas as it is today.
</paragraph>
<paragraph blockId="6.[106,787,134,1358]" pageId="6" pageNumber="7">
The currently known latest Cretaceous (Maastrichtian) vertebrate fauna of Madagascar, assembled as an island fauna over the course of more than 20 Myr (around 8866 Myr ago),became completely or nearlycompletelyextinct—presumablytheresultoftheend-Cretaceous bolide impact and/or the penecontemporaneous nearby volcanic eruptions that resulted in the Deccan Traps of India
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<superScript attach="left" box="[555,570,1218,1231]" fontSize="5" pageId="6" pageNumber="7">44</superScript>
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. Thereafter,assembly of a Madagascan fauna began afresh.The establishment of a new vertebrate fauna required the arrival of transoceanic dispersers,primarily from Africa,that encountered habitats that were largely to completely devoid of Maastrichtian antecedents
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</superScript>
.
</paragraph>
</subSubSection>
</treatment>
</document>