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<document ID-DOI="10.1206/0003-0090.457.1.1" ID-ISSN="0003-0090" ID-Zenodo-Dep="6971356" approvalRequired="59" approvalRequired_for_taxonomicNames="59" checkinTime="1659882917898" checkinUser="felipe" docAuthor="Beck, Robin M. D., Voss, Robert S. &amp; Jansa, Sharon A." docDate="2022" docId="03EFDD5DF77D696ED972FCC81EFFF9E3" docLanguage="en" docName="BulAmeMusNatHist.2022.457.1-350.pdf" docOrigin="Bulletin of the American Museum of Natural History 2022 (457)" docSource="https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-457/issue-1/0003-0090.457.1.1/Craniodental-Morphology-and-Phylogeny-of-Marsupials/10.1206/0003-0090.457.1.1.full" docStyle="DocumentStyle:915933466F796C9C739DF4DB6B8DCFA6.8:BulAmeMusNatHis.2011-.journal_article.1cover.type1" docStyleId="915933466F796C9C739DF4DB6B8DCFA6" docStyleName="BulAmeMusNatHis.2011-.journal_article.1cover.type1" docStyleVersion="8" docTitle="Litokoala undetermined" docType="treatment" docVersion="5" lastPageNumber="327" masterDocId="FFD6A525F63B6829DA0BFFAE1A62FFC7" masterDocTitle="Craniodental Morphology And Phylogeny Of Marsupials" masterLastPageNumber="353" masterPageNumber="1" pageNumber="326" updateTime="1659987007554" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title>Craniodental Morphology And Phylogeny Of Marsupials</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Beck, Robin M. D.</mods:namePart>
<mods:affiliation>School of Science, Engineering and Environment University of Salford, U. K. &amp; School of Biological, Earth &amp; Environmental Sciences University of New South Wales, Australia &amp; Division of Vertebrate Zoology (Mammalogy) American Museum of Natural History</mods:affiliation>
</mods:name>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Voss, Robert S.</mods:namePart>
<mods:affiliation>Division of Vertebrate Zoology (Mammalogy) American Museum of Natural History</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Jansa, Sharon A.</mods:namePart>
<mods:affiliation>Bell Museum and Department of Ecology, Evolution, and Behavior University of Minnesota</mods:affiliation>
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<mods:date>2022</mods:date>
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<mods:number>2022-06-28</mods:number>
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<mods:number>2022</mods:number>
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<mods:identifier type="DOI">10.1206/0003-0090.457.1.1</mods:identifier>
<mods:identifier type="ISSN">0003-0090</mods:identifier>
<mods:identifier type="Zenodo-Dep">6971356</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.6974476" ID-Zenodo-Dep="6974476" LSID="urn:lsid:plazi:treatment:03EFDD5DF77D696ED972FCC81EFFF9E3" httpUri="http://treatment.plazi.org/id/03EFDD5DF77D696ED972FCC81EFFF9E3" lastPageId="327" lastPageNumber="327" pageId="326" pageNumber="326">
<subSubSection box="[889,1008,870,894]" pageId="326" pageNumber="326" type="nomenclature">
<paragraph blockId="326.[889,1008,870,894]" box="[889,1008,870,894]" pageId="326" pageNumber="326">
<heading box="[889,1008,870,894]" centered="true" fontSize="10" level="2" pageId="326" pageNumber="326" reason="4">
<emphasis bold="true" box="[889,1008,870,894]" pageId="326" pageNumber="326">
<taxonomicName box="[902,1008,870,894]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" kingdom="Animalia" order="Diprotodontia" pageId="326" pageNumber="326" phylum="Chordata" rank="species">
<emphasis bold="true" box="[902,1008,870,894]" italics="true" pageId="326" pageNumber="326">Litokoala</emphasis>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection pageId="326" pageNumber="326" type="synonymic_list">
<paragraph blockId="326.[683,1213,920,1738]" pageId="326" pageNumber="326">
SPECIES SCORED: †
<emphasis box="[933,1212,920,944]" italics="true" pageId="326" pageNumber="326">
<taxonomicName box="[933,1035,920,944]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="326" pageNumber="326" phylum="Chordata" rank="genus">Litokoala</taxonomicName>
kutjamarpensis
</emphasis>
(
<typeStatus box="[692,741,953,977]" pageId="326" pageNumber="326">type</typeStatus>
species), †
<emphasis box="[860,1005,953,977]" italics="true" pageId="326" pageNumber="326">L. dicksmithi.</emphasis>
</paragraph>
</subSubSection>
<subSubSection pageId="326" pageNumber="326" type="materials_examined">
<paragraph blockId="326.[683,1213,920,1738]" pageId="326" pageNumber="326">
GEOLOGICAL PROVENANCE OF SCORED
<collectionCode box="[684,742,1019,1043]" pageId="326" pageNumber="326">SPECIMENS</collectionCode>
: UCR Locality RV-8453 (Kanunka North Local Fauna), Etadunna Formation, Lake Kanunka,
<collectingRegion box="[683,857,1085,1109]" country="Australia" name="South Australia" pageId="326" pageNumber="326">South Australia</collectingRegion>
,
<collectingCountry box="[873,970,1085,1109]" name="Australia" pageId="326" pageNumber="326">Australia</collectingCountry>
(†
<emphasis box="[1004,1196,1085,1109]" italics="true" pageId="326" pageNumber="326">L. kutjamarpensis</emphasis>
); Jims Carousel, Henks Hollow, Dwornamor, and Gotham sites (Riversleigh Faunal Zone C), Riversleigh World Heritage Area,
<collectingRegion box="[981,1108,1184,1208]" country="Australia" name="Queensland" pageId="326" pageNumber="326">Queensland</collectingRegion>
,
<collectingCountry box="[1118,1213,1184,1208]" name="Australia" pageId="326" pageNumber="326">Australia</collectingCountry>
(†
<emphasis box="[706,894,1218,1242]" italics="true" pageId="326" pageNumber="326">L. kutjamarpensis</emphasis>
); Ross Scott-Orr Site (Riversleigh Faunal Zone B), Riversleigh World Heritage Area,
<collectingRegion box="[747,874,1284,1308]" country="Australia" name="Queensland" pageId="326" pageNumber="326">Queensland</collectingRegion>
,
<collectingCountry box="[885,979,1284,1308]" name="Australia" pageId="326" pageNumber="326">Australia</collectingCountry>
(†
<emphasis box="[1006,1137,1284,1308]" italics="true" pageId="326" pageNumber="326">L. dicksmithi</emphasis>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="327" lastPageNumber="327" pageId="326" pageNumber="326" type="description">
<paragraph blockId="326.[683,1213,920,1738]" pageId="326" pageNumber="326">AGE OF SCORED SPECIMENS: Based on the paleomagnetic data of Woodburne et al. (1994), Metzger and Retallack (2010) estimated the Etadunna Formation to span 26.123.6 Mya. Riversleigh Faunal Zone B and C are interpreted to be early and middle Miocene, respectively, based on biostratigraphy (see above), but only Ross ScottOrr (RSO) Site has a radiometric date, of 16.24 16.86 Mya (Woodhead et al., 2014). We conservatively assign an age range of the entire late Oligocene to middle Miocene (Chattian to Serravalian; Cohen et al., 2013 [updated]) for this taxon.</paragraph>
<paragraph blockId="327.[108,638,226,1738]" box="[140,621,226,250]" pageId="327" pageNumber="327">
ASSIGNED AGE
<collectionCode box="[311,388,226,250]" pageId="327" pageNumber="327">RANGE</collectionCode>
:
<geoCoordinate box="[402,475,226,250]" degrees="27.820" direction="north" orientation="latitude" pageId="327" pageNumber="327" precision="55" value="27.82">27.820</geoCoordinate>
<geoCoordinate box="[474,559,226,250]" degrees="11.630" direction="west" orientation="longitude" pageId="327" pageNumber="327" precision="55" value="-11.63">11.630</geoCoordinate>
Mya.
</paragraph>
</subSubSection>
<subSubSection pageId="327" pageNumber="327" type="discussion">
<paragraph blockId="327.[108,638,226,1738]" pageId="327" pageNumber="327">
REMARKS: †
<taxonomicName box="[272,372,259,283]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">
<emphasis box="[272,372,259,283]" italics="true" pageId="327" pageNumber="327">Litokoala</emphasis>
</taxonomicName>
is one of only two fossil phascolarctids known from relatively complete craniodental material (Louys et al., 2009; Black et al., 2014a), but its taxonomic history is complex and remains controversial. †
<emphasis italics="true" pageId="327" pageNumber="327">
<taxonomicName box="[482,583,391,415]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">Litokoala</taxonomicName>
kutjamarpensis
</emphasis>
was described by Stirton et al. (1967a) based on a single M1 from the Kutjamarpu Local Fauna, Wipajiri Formation, Lake Ngapakaldi,
<collectingRegion box="[255,438,523,547]" country="Australia" name="South Australia" pageId="327" pageNumber="327">South Australia</collectingRegion>
. Springer (1987) then described a second species, †
<emphasis italics="true" pageId="327" pageNumber="327">L. kanunkaensis</emphasis>
, based on two isolated lower molars and two partial upper molars from the Kanunka North Local Fauna, Etadunna Formation, Lake Kanunka,
<collectingRegion box="[341,515,688,712]" country="Australia" name="South Australia" pageId="327" pageNumber="327">South Australia</collectingRegion>
. Black and Archer (1997b) subsequently referred several phascolarctid dental specimens from a number of Riversleigh Faunal Zone C sites to †
<emphasis italics="true" pageId="327" pageNumber="327">L. kanunkaensis</emphasis>
. However, based on study of the dentition of a new, well-preserved partial cranium (
<collectionCode box="[181,227,887,911]" country="Australia" lsid="urn:lsid:biocol.org:col:35019" name="Queensland Museum" pageId="327" pageNumber="327" type="Museum">QM</collectionCode>
<accessionNumber box="[234,317,887,911]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/F51382" pageId="327" pageNumber="327">F51382</accessionNumber>
) from the Riversleigh Faunal Zone C JC Site, Louys et al. (2007) concluded that †
<emphasis box="[171,343,953,977]" italics="true" pageId="327" pageNumber="327">L. kanunkaensis</emphasis>
is a junior synonym of †
<emphasis italics="true" pageId="327" pageNumber="327">L. kutjamarpensis</emphasis>
. Louys et al. (2007) also named a new species, †
<emphasis box="[268,445,1019,1043]" italics="true" pageId="327" pageNumber="327">L. garyjohnstoni</emphasis>
, based on a partial maxilla and isolated M4 from the Riversleigh Faunal Zone B Outasite Site. Louys et al. (2009) subsequently gave a detailed description of the cranial morphology of
<collectionCode box="[366,410,1152,1176]" country="Australia" lsid="urn:lsid:biocol.org:col:35019" name="Queensland Museum" pageId="327" pageNumber="327" type="Museum">QM</collectionCode>
<accessionNumber box="[418,501,1151,1175]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/F51382" pageId="327" pageNumber="327">F51382</accessionNumber>
, which they maintained represents †
<emphasis box="[373,387,1185,1208]" italics="true" pageId="327" pageNumber="327">L</emphasis>
.
<emphasis box="[401,565,1184,1208]" italics="true" pageId="327" pageNumber="327">kutjamarpensis</emphasis>
.
</paragraph>
<paragraph blockId="327.[108,638,226,1738]" lastBlockId="327.[684,1214,226,1573]" pageId="327" pageNumber="327">
Pledge (2010), however, retained †
<emphasis italics="true" pageId="327" pageNumber="327">L. kanunkaensis</emphasis>
as a separate species for the isolated molars from the Kanunka North Local Fauna described by Springer (1987), referred the Riversleigh Faunal Zone C specimens (including
<collectionCode box="[108,152,1383,1407]" country="Australia" lsid="urn:lsid:biocol.org:col:35019" name="Queensland Museum" pageId="327" pageNumber="327" type="Museum">QM</collectionCode>
<accessionNumber box="[162,245,1383,1407]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/F51382" pageId="327" pageNumber="327">F51382</accessionNumber>
) to a new species, †
<emphasis box="[473,630,1383,1407]" italics="true" pageId="327" pageNumber="327">L. dicktedfordi</emphasis>
, and named a further species, †
<emphasis box="[481,629,1416,1440]" italics="true" pageId="327" pageNumber="327">L. thurmerae</emphasis>
, based on an isolated M3 from the?late Oligocene Ngama Local Fauna, Lake Palankarinna,
<collectingRegion box="[108,281,1515,1539]" country="Australia" name="South Australia" pageId="327" pageNumber="327">South Australia</collectingRegion>
. Thus, Pledge (2010) recognized five species: †
<emphasis box="[274,478,1548,1572]" italics="true" pageId="327" pageNumber="327">L. kutjamarpensis</emphasis>
(Kutjamarpu Local Fauna;?early or middle Miocene), †
<emphasis italics="true" pageId="327" pageNumber="327">L. kanunkaensis</emphasis>
(Kanunka North Local Fauna;?late Oligocene), †
<emphasis box="[258,436,1647,1671]" italics="true" pageId="327" pageNumber="327">L. garyjohnstoni</emphasis>
(Riversleigh Faunal Zone B;?early Miocene), †
<emphasis box="[474,636,1680,1704]" italics="true" pageId="327" pageNumber="327">L. dicktedfordi</emphasis>
(Riversleigh Faunal Zone C;?middle Miocene), and †
<emphasis box="[750,895,226,250]" italics="true" pageId="327" pageNumber="327">L. thurmerae</emphasis>
(Ngama Local Fauna;?late Oligocene). Most recently, Black et al. (2014a, 2014b) reaffirmed that †
<emphasis box="[947,1118,292,316]" italics="true" pageId="327" pageNumber="327">L. kanunkaensis</emphasis>
(including the Riversleigh Faunal Zone C species referred to †
<emphasis box="[822,977,358,382]" italics="true" pageId="327" pageNumber="327">L. dicktedfordi</emphasis>
by Pledge, 2010) is a junior synonym of †
<emphasis box="[918,1113,391,415]" italics="true" pageId="327" pageNumber="327">L. kutjamarpensis</emphasis>
, considered †
<emphasis box="[753,893,424,448]" italics="true" pageId="327" pageNumber="327">L. thurmerae</emphasis>
to be a nomen dubium, and named yet another new species, †
<emphasis box="[1065,1206,457,481]" italics="true" pageId="327" pageNumber="327">L. dicksmithi</emphasis>
, based on a well-preserved rostral fragment (
<collectionCode box="[1166,1212,490,514]" country="Australia" lsid="urn:lsid:biocol.org:col:35019" name="Queensland Museum" pageId="327" pageNumber="327" type="Museum">QM</collectionCode>
<accessionNumber box="[685,769,523,547]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/F54567" pageId="327" pageNumber="327">F54567</accessionNumber>
). Here we follow the taxonomy of Louys et al. (2007, 2009) and Black et al. (2014a, 2014b), who recognized only two species within the genus: the
<typeStatus box="[801,848,623,647]" pageId="327" pageNumber="327">type</typeStatus>
species, †
<emphasis italics="true" pageId="327" pageNumber="327">
<taxonomicName box="[958,1058,622,646]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">Litokoala</taxonomicName>
kutjamarpensis
</emphasis>
, and †
<emphasis box="[785,922,655,679]" italics="true" pageId="327" pageNumber="327">L. dicksmithi</emphasis>
.
</paragraph>
<paragraph blockId="327.[684,1214,226,1573]" pageId="327" pageNumber="327">
We scored our †
<taxonomicName box="[901,1001,688,712]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">
<emphasis box="[901,1001,688,712]" italics="true" pageId="327" pageNumber="327">Litokoala</emphasis>
</taxonomicName>
terminal based on specimens of †
<emphasis box="[851,1045,722,746]" italics="true" pageId="327" pageNumber="327">L. kutjamarpensis</emphasis>
from both the Kanunka North Local Fauna and Riversleigh Faunal Zone C sites, and also on the only known specimen (
<collectionCode box="[803,849,821,845]" country="Australia" lsid="urn:lsid:biocol.org:col:35019" name="Queensland Museum" pageId="327" pageNumber="327" type="Museum">QM</collectionCode>
<accessionNumber box="[857,938,821,845]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/F54567" pageId="327" pageNumber="327">F54567</accessionNumber>
) of †
<emphasis box="[996,1133,821,845]" italics="true" pageId="327" pageNumber="327">L. dicksmithi</emphasis>
. Based on this material, †
<taxonomicName box="[909,1012,854,878]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">
<emphasis box="[909,1012,854,878]" italics="true" pageId="327" pageNumber="327">Litokoala</emphasis>
</taxonomicName>
appears to have lacked masticatory specializations seen in
<taxonomicName authority="(Louys et al., 2009)" baseAuthorityName="Louys" baseAuthorityYear="2009" class="Mammalia" family="Phascolarctidae" genus="Phascolarctos" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="327" pageNumber="327">Phascolarctos</emphasis>
(Louys et al., 2009)
</taxonomicName>
. In addition, estimated body masses for †
<taxonomicName box="[1009,1212,953,977]" pageId="327" pageNumber="327">
<emphasis box="[1009,1114,953,977]" italics="true" pageId="327" pageNumber="327">Litokoala</emphasis>
species
</taxonomicName>
(
<quantity box="[691,809,986,1010]" metricMagnitude="0" metricUnit="kg" metricValue="3.5" metricValueMax="4.6" metricValueMin="2.4" pageId="327" pageNumber="327" unit="kg" value="3.5" valueMax="4.6" valueMin="2.4">2.44.6 kg</quantity>
; Black et al., 2014a; 2014b) are considerably less than those of
<taxonomicName baseAuthorityName="Goldfuss" baseAuthorityYear="1817" box="[992,1105,1020,1043]" class="Mammalia" family="Phascolarctidae" genus="Phascolarctos" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="species">
<emphasis box="[992,1105,1020,1043]" italics="true" pageId="327" pageNumber="327">P. cinereus</emphasis>
</taxonomicName>
(
<quantity metricMagnitude="0" metricUnit="kg" metricValue="8.8" metricValueMax="13.5" metricValueMin="4.1" pageId="327" pageNumber="327" unit="kg" value="8.8" valueMax="13.5" valueMin="4.1">4.113.5 kg</quantity>
; Martin et al., 2008). Nevertheless, phylogenetic analyses of vombatiform relationships based on 71 craniodental characters by Black et al. (2012a) supported a sister-group relationship between †
<taxonomicName box="[805,908,1184,1208]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">
<emphasis box="[805,908,1184,1208]" italics="true" pageId="327" pageNumber="327">Litokoala</emphasis>
</taxonomicName>
and
<taxonomicName box="[978,1126,1184,1208]" class="Mammalia" family="Phascolarctidae" genus="Phascolarctos" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">
<emphasis box="[978,1126,1184,1208]" italics="true" pageId="327" pageNumber="327">Phascolarctos</emphasis>
</taxonomicName>
to the exclusion of the other currently named fossil phascolarctid genera, and Beck et al. (2020 fig. 5) found †
<taxonomicName box="[769,869,1284,1308]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">
<emphasis box="[769,869,1284,1308]" italics="true" pageId="327" pageNumber="327">Litokoala</emphasis>
</taxonomicName>
to be paraphyletic with respect to
<taxonomicName box="[718,865,1317,1341]" class="Mammalia" family="Phascolarctidae" genus="Phascolarctos" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">
<emphasis box="[718,865,1317,1341]" italics="true" pageId="327" pageNumber="327">Phascolarctos</emphasis>
</taxonomicName>
, with †
<emphasis box="[958,1159,1317,1341]" italics="true" pageId="327" pageNumber="327">L. kutjamarpensis</emphasis>
and
<taxonomicName box="[684,828,1350,1374]" class="Mammalia" family="Phascolarctidae" genus="Phascolarctos" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">
<emphasis box="[684,828,1350,1374]" italics="true" pageId="327" pageNumber="327">Phascolarctos</emphasis>
</taxonomicName>
forming a clade to the exclusion of †
<emphasis box="[698,843,1383,1407]" italics="true" pageId="327" pageNumber="327">L. dicksmithi</emphasis>
. However, an undescribed new genus and species of phascolarctid from the Riversleigh Faunal Zone D Encore Site, previously identified as
<taxonomicName box="[836,1025,1482,1506]" class="Mammalia" family="Phascolarctidae" genus="Phascolarctos" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="species">
<emphasis box="[836,983,1482,1506]" italics="true" pageId="327" pageNumber="327">Phascolarctos</emphasis>
sp.
</taxonomicName>
by Myers et al. (2001), may be more closely related to the living genus than is †
<taxonomicName authority="(Black et al., 2014 b)" baseAuthorityName="Black" baseAuthorityYear="2014" box="[849,1177,1548,1572]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="327" pageNumber="327" phylum="Chordata" rank="genus">
<emphasis box="[849,949,1548,1572]" italics="true" pageId="327" pageNumber="327">Litokoala</emphasis>
(Black et al., 2014b)
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>