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<document id="A02C8941017410D2742CFEF4574C8D09" ID-CLB-Dataset="7844" ID-DOI="10.1093/mspecies/sex001" ID-GBIF-Dataset="90dc3348-c6bc-4ecf-87bd-49d17e3168fd" ID-ISSN="1545-1410" ID-Zenodo-Dep="4573410" ID-ZooBank="10C0593B-41AC-49CD-90AD-EC53D38A9FEB" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1614694303045" checkinUser="felipe" docAuthor="Rose, Robert K., Pemberton, David A., Mooney, Nick J. &amp; Jones, Menna E." docDate="2017" docId="9F09878FFFEFE7143DE7F9911B36F9EB" docLanguage="en" docName="MammalSpecies.49.942.1-17.pdf" docOrigin="Mammalian Species 49 (942)" docSource="http://zoobank.org/10c0593b-41ac-49cd-90ad-ec53d38a9feb" docStyle="DocumentStyle:E260F2FEBB038053A7447521347BED36.4:MammalSpecies.2016-.journal_article" docStyleId="E260F2FEBB038053A7447521347BED36" docStyleName="MammalSpecies.2016-.journal_article" docStyleVersion="4" docTitle="Sarcophilus harrisii" docType="treatment" docVersion="6" lastPageNumber="13" masterDocId="6330FFF7FFEFE7183D0CFFC81E70FFC9" masterDocTitle="Sarcophilus harrisii (Dasyuromorphia: Dasyuridae)" masterLastPageNumber="17" masterPageNumber="1" pageNumber="1" updateTime="1698931748156" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:title id="AB1E29E0102DEF30BB8B9CACB0173DF7">Sarcophilus harrisii (Dasyuromorphia: Dasyuridae)</mods:title>
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<mods:namePart id="4E93C0E902192FDB09F6BB7149C3628A">Jones, Menna E.</mods:namePart>
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<treatment id="9F09878FFFEFE7143DE7F9911B36F9EB" ID-DOI="http://doi.org/10.5281/zenodo.4589273" ID-GBIF-Taxon="180300423" ID-Zenodo-Dep="4589273" LSID="urn:lsid:plazi:treatment:9F09878FFFEFE7143DE7F9911B36F9EB" httpUri="http://treatment.plazi.org/id/9F09878FFFEFE7143DE7F9911B36F9EB" lastPageId="12" lastPageNumber="13" pageId="0" pageNumber="1">
<subSubSection id="5FBA6512FFEFE7183DE7F9911CEAF9BA" box="[235,666,1625,1651]" pageId="0" pageNumber="1" type="nomenclature">
<paragraph id="171F3699FFEFE7183DE7F9911CEAF9BA" blockId="0.[109,792,1625,1831]" box="[235,666,1625,1651]" pageId="0" pageNumber="1">
<heading id="4C5781F5FFEFE7183DE7F9911CEAF9BA" box="[235,666,1625,1651]" centered="true" fontSize="11" level="2" pageId="0" pageNumber="1" reason="2">
<taxonomicName id="D0A04D1AFFEFE7183DE7F9911CEAF9BA" ID-CoL="6XMTZ" authority="(Boitard, 1842)" baseAuthorityName="Boitard" baseAuthorityYear="1842" box="[235,666,1625,1651]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEFE7183DE7F9911FA6F9BA" box="[235,470,1625,1651]" italics="true" pageId="0" pageNumber="1">Sarcophilus harrisii</emphasis>
<emphasis id="25D4EA8BFFEFE7183CD1F9911CEAF9BA" bold="true" box="[477,666,1625,1651]" pageId="0" pageNumber="1">
(
<bibRefCitation id="73314B68FFEFE7183CEBF9911CE1F9BA" author="BOITARD, P." box="[487,657,1625,1651]" pageId="0" pageNumber="1" refId="ref15036" refString="BOITARD, P. 1842. Le Jardin des Plantes description et moeurs des mammiferes de la Menagerie et du museum d'Histoire Naturelle precede d'une introduction historique, descriptive et pittoresque par M. J. Janin. Gustave Barba, Paris, France." type="book" year="1842">Boitard, 1842</bibRefCitation>
)
</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="5FBA6512FFEFE7183C72F94E1C42F954" box="[382,562,1670,1693]" pageId="0" pageNumber="1" type="description">
<paragraph id="171F3699FFEFE7183C72F94E1C42F954" blockId="0.[109,792,1625,1831]" box="[382,562,1670,1693]" pageId="0" pageNumber="1">Tasmanian Devil</paragraph>
</subSubSection>
<subSubSection id="5FBA6512FFEFE7193D61F9781E9BFDCB" lastPageId="1" lastPageNumber="2" pageId="0" pageNumber="1" type="reference_group">
<paragraph id="171F3699FFEFE7183D61F9781F15F92E" blockId="0.[109,792,1625,1831]" pageId="0" pageNumber="1">
<taxonomicName id="D0A04D1AFFEFE7183D61F9781FACF90E" authority="Harris, 1808: 176" authorityName="Harris" authorityPageNumber="176" authorityYear="1808" box="[109,476,1712,1735]" class="Mammalia" family="Didelphidae" genus="Didelphis" kingdom="Animalia" order="Didelphimorphia" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="ursina">
<emphasis id="25D4EA8BFFEFE7183D61F9781F6FF90E" box="[109,287,1712,1735]" italics="true" pageId="0" pageNumber="1">Didelphis ursina</emphasis>
<bibRefCitation id="73314B68FFEFE7183C29F9781FACF90E" author="HARRIS, G. P." box="[293,476,1712,1735]" pageId="0" pageNumber="1" pagination="174 - 178" refId="ref17365" refString="HARRIS, G. P. 1808. Description of two new species of Didelphis from Van Diemen's Land. Transactions of the Linnean Society of London 9: 174 - 178." type="journal article" year="1808">Harris, 1808:176</bibRefCitation>
</taxonomicName>
.
<typeStatus id="C81B883BFFEFE7183CEAF9781C6BF90E" box="[486,539,1712,1735]" pageId="0" pageNumber="1">Type</typeStatus>
locality
<quote id="655FC18BFFEFE7183F7AF9781E90F92E" pageId="0" pageNumber="1">“Van Diemens Land”</quote>
(
<collectingRegion id="D564F87BFFEFE7183DE2F9181F2AF92E" box="[238,346,1744,1767]" country="Australia" name="Tasmania" pageId="0" pageNumber="1">Tasmania</collectingRegion>
).
</paragraph>
<caption id="43DF6611FFEFE7183E44F9111B83F8E0" ID-DOI="http://doi.org/10.5281/zenodo.4573412" ID-Zenodo-Dep="4573412" httpUri="https://zenodo.org/record/4573412/files/figure.png" pageId="0" pageNumber="1" startId="0.[840,875,1753,1774]" targetBox="[877,1485,1098,1739]" targetPageId="0">
<paragraph id="171F3699FFEFE7183E44F9111B83F8E0" blockId="0.[840,1523,1753,1833]" pageId="0" pageNumber="1">
<emphasis id="25D4EA8BFFEFE7183E44F9111DF9F927" bold="true" box="[840,905,1753,1774]" pageId="0" pageNumber="1">Fig. 1.</emphasis>
—An alert young
<taxonomicName id="D0A04D1AFFEFE7183937F9111A8DF927" ID-CoL="6XMTZ" baseAuthorityName="Boitard" baseAuthorityYear="1842" box="[1083,1277,1753,1774]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEFE7183937F9111A8DF927" box="[1083,1277,1753,1774]" italics="true" pageId="0" pageNumber="1">Sarcophilus harrisii</emphasis>
</taxonomicName>
showing black coat with white marking on chest, prominent vibrissae, wet nose, and nearly naked ears. About 13% are all black. The sexes are similar except that males
</paragraph>
</caption>
<paragraph id="171F3699FFEFE7183D61F9381D68F8EE" blockId="0.[109,792,1625,1831]" pageId="0" pageNumber="1">
<taxonomicName id="D0A04D1AFFEFE7183D61F9381FF6F8CE" authority="Owen" authorityName="Owen" authorityYear="1838" box="[109,390,1776,1799]" class="Mammalia" family="Dasyuridae" genus="Dasyurus" kingdom="Animalia" order="Dasyuromorphia" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="laniarius">
<emphasis id="25D4EA8BFFEFE7183D61F9381F4DF8CE" box="[109,317,1776,1799]" italics="true" pageId="0" pageNumber="1">Dasyurus laniarius</emphasis>
Owen
</taxonomicName>
in Mitchell, 1838:363, plate 49, figures 3 and 4.
<typeStatus id="C81B883BFFEFE7183C3CF8D81F15F8EE" box="[304,357,1808,1831]" pageId="0" pageNumber="1">Type</typeStatus>
locality “
<collectingRegion id="D564F87BFFEFE7183CC1F8D81C35F8EE" box="[461,581,1808,1831]" country="New Zealand" name="Wellington" pageId="0" pageNumber="1">Wellington</collectingRegion>
Valley
<insertion id="8A255158FFEFE7183F9BF8D81D68F8EE" box="[663,792,1808,1831]" pageId="0" pageNumber="1">
[
<collectingRegion id="D564F87BFFEFE7183F92F8D81D68F8EE" box="[670,792,1808,1831]" country="Australia" name="New South Wales" pageId="0" pageNumber="1">New South</collectingRegion>
</insertion>
</paragraph>
<paragraph id="171F3699FFEFE7183D91F8F81BBFF88F" blockId="0.[157,716,1840,1863]" box="[157,1487,1840,1863]" lastBlockId="0.[840,1487,1841,1862]" pageId="0" pageNumber="1">
<collectingCountry id="6FB77609FFEFE7183D91F8F81E93F88E" box="[157,227,1840,1863]" name="United Kingdom" pageId="0" pageNumber="1">Wales</collectingCountry>
],
<collectingCountry id="6FB77609FFEFE7183DFFF8F81F2AF88E" box="[243,346,1840,1863]" name="Australia" pageId="0" pageNumber="1">Australia</collectingCountry>
;” fossil form from the Pleistocene. are 56% larger. Photograph by Christo Baars used with permission.
</paragraph>
<paragraph id="171F3699FFEEE7193D61FF031F8FFEEB" blockId="1.[109,792,203,589]" pageId="1" pageNumber="2">
<taxonomicName id="D0A04D1AFFEEE7193D61FF031F84FF2B" authority="Harrisii Boitard, 1842: 290" authorityName="Harrisii Boitard" authorityPageNumber="290" authorityYear="1842" box="[109,500,203,226]" class="Mammalia" family="Dasyuridae" genus="Ursinus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis id="25D4EA8BFFEEE7193D61FF031F52FF2B" box="[109,290,203,226]" italics="true" pageId="1" pageNumber="2">Ursinus Harrisii</emphasis>
<bibRefCitation id="73314B68FFEEE7193C20FF031F84FF2B" author="BOITARD, P." box="[300,500,203,226]" pageId="1" pageNumber="2" refId="ref15036" refString="BOITARD, P. 1842. Le Jardin des Plantes description et moeurs des mammiferes de la Menagerie et du museum d'Histoire Naturelle precede d'une introduction historique, descriptive et pittoresque par M. J. Janin. Gustave Barba, Paris, France." type="book" year="1842">Boitard, 1842:290</bibRefCitation>
</taxonomicName>
.
<typeStatus id="C81B883BFFEEE7193F0EFF031C47FF2B" box="[514,567,203,226]" pageId="1" pageNumber="2">Type</typeStatus>
locality
<quote id="655FC18BFFEEE7193F97FF031F47FECB" pageId="1" pageNumber="2">“la terre de Van-Diemen;”</quote>
replacement name for
<taxonomicName id="D0A04D1AFFEEE7193F56FF231F52FEEB" authority="Harris, 1808" authorityName="Harris" authorityYear="1808" class="Mammalia" family="Didelphidae" genus="Didelphis" kingdom="Animalia" order="Didelphimorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="ursina">
<emphasis id="25D4EA8BFFEEE7193F56FF231D68FECB" box="[602,792,235,258]" italics="true" pageId="1" pageNumber="2">Didelphis ursina</emphasis>
<bibRefCitation id="73314B68FFEEE7193D91FEC31F52FEEB" author="HARRIS, G. P." box="[157,290,267,290]" pageId="1" pageNumber="2" pagination="174 - 178" refId="ref17365" refString="HARRIS, G. P. 1808. Description of two new species of Didelphis from Van Diemen's Land. Transactions of the Linnean Society of London 9: 174 - 178." type="journal article" year="1808">Harris, 1808</bibRefCitation>
</taxonomicName>
vide
<bibRefCitation id="73314B68FFEEE7193C52FEC31F88FEEB" author="GROVES, C. P." box="[350,504,267,290]" pageId="1" pageNumber="2" refId="ref16815" refString="GROVES, C. P. 2005. Order Dasyuromorphia. Pp. 23 - 37 in Mammal species of the world: a taxonomic and geographic reference (D. E. Wilson and D. A. M. Reeder, eds.). 3 rd ed. Smithsonian Institution Press, Washington, D. C." type="book" year="2005">Groves (2005)</bibRefCitation>
.
</paragraph>
<paragraph id="171F3699FFEEE7193D61FEE31F50FEAB" blockId="1.[109,792,203,589]" pageId="1" pageNumber="2">
<taxonomicName id="D0A04D1AFFEEE7193D61FEE31C2FFE8B" authority="Thomas, 1903: 289" authorityName="Thomas" authorityPageNumber="289" authorityYear="1903" box="[109,607,299,322]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="satanicus">
<emphasis id="25D4EA8BFFEEE7193D61FEE31F1AFE8B" box="[109,362,299,322]" italics="true" pageId="1" pageNumber="2">Sarcophilus satanicus</emphasis>
<bibRefCitation id="73314B68FFEEE7193C8EFEE31C2FFE8B" author="THOMAS, O." box="[386,607,299,322]" pageId="1" pageNumber="2" pagination="289" refId="ref21523" refString="THOMAS, O. 1903. Note on the technical name of the Tasmanian devil. Annals and Magazine of Natural History 11: 289." type="journal article" year="1903">Thomas, 1903:289</bibRefCitation>
</taxonomicName>
.
<typeStatus id="C81B883BFFEEE7193F77FEE31CC0FE8B" box="[635,688,299,322]" pageId="1" pageNumber="2">Type</typeStatus>
locality
<quote id="655FC18BFFEEE7193D91FE831F50FEAB" box="[157,288,331,354]" pageId="1" pageNumber="2">
<collectingRegion id="D564F87BFFEEE7193DABFE831F63FEAB" box="[167,275,331,354]" country="Australia" name="Tasmania" pageId="1" pageNumber="2">Tasmania</collectingRegion>
.”
</quote>
</paragraph>
<paragraph id="171F3699FFEEE7193D61FEA31F1BFE6B" blockId="1.[109,792,203,589]" pageId="1" pageNumber="2">
<taxonomicName id="D0A04D1AFFEEE7193D61FEA31C58FE4B" ID-CoL="6XMTZ" authority="Thomas, 1912: 116" authorityName="Thomas" authorityPageNumber="116" authorityYear="1912" box="[109,552,363,386]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193D61FEA31F39FE4B" box="[109,329,363,386]" italics="true" pageId="1" pageNumber="2">Sarcophilus harrisii</emphasis>
<bibRefCitation id="73314B68FFEEE7193C5AFEA31C58FE4B" author="THOMAS, O." box="[342,552,363,386]" pageId="1" pageNumber="2" pagination="116" refId="ref21549" refString="THOMAS, O. 1912. The technical name of the Tasmanian devil. Proceedings of the Biological Society of Washington 25: 116." type="journal article" year="1912">Thomas, 1912:116</bibRefCitation>
</taxonomicName>
. First use of current name combination.
</paragraph>
<paragraph id="171F3699FFEEE7193D61FE631E9BFDCB" blockId="1.[109,792,203,589]" pageId="1" pageNumber="2">
<emphasis id="25D4EA8BFFEEE7193D61FE631FC0FE0B" box="[109,432,427,450]" italics="true" pageId="1" pageNumber="2">
<taxonomicName id="D0A04D1AFFEEE7193D61FE631F3FFE0B" ID-CoL="6XMTZ" authorityName="F. Cuvier" authorityYear="1837" box="[109,335,427,450]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">Sarcophilus. harrisii</taxonomicName>
. dixonae
</emphasis>
,
<bibRefCitation id="73314B68FFEEE7193CCCFE631CE0FE0B" author="WERDELIN, L." box="[448,656,427,450]" pageId="1" pageNumber="2" pagination="1 - 27" refId="ref21822" refString="WERDELIN, L. 1987 a. Some observations on Sarcophilus laniarius and the evolution of Sarcophilus. Records of the Queen Victoria Museum, Launceston 90: 1 - 27." type="journal article" year="1987">Werdelin 1987a:10</bibRefCitation>
.
<typeStatus id="C81B883BFFEEE7193F91FE631CA2FE0B" box="[669,722,427,450]" pageId="1" pageNumber="2">Type</typeStatus>
locality
<quote id="655FC18BFFEEE7193DC8FE031F07FE2B" box="[196,375,459,482]" pageId="1" pageNumber="2">“Mt. Hamilton;”</quote>
Victorian subfossil form vide
<bibRefCitation id="73314B68FFEEE7193FC0FE031E95FDCB" author="GROVES, C. P." pageId="1" pageNumber="2" refId="ref16815" refString="GROVES, C. P. 2005. Order Dasyuromorphia. Pp. 23 - 37 in Mammal species of the world: a taxonomic and geographic reference (D. E. Wilson and D. A. M. Reeder, eds.). 3 rd ed. Smithsonian Institution Press, Washington, D. C." type="book" year="2005">Groves (2005)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="5FBA6512FFEEE7193D61FDDE1F7EFC34" pageId="1" pageNumber="2" type="description">
<paragraph id="171F3699FFEEE7193D61FDDE1C8FFD84" blockId="1.[109,792,203,589]" pageId="1" pageNumber="2">
CONTEXT AND CONTENT. Context as for genus.
<emphasis id="25D4EA8BFFEEE7193F6AFDDE1EE8FD84" italics="true" pageId="1" pageNumber="2">
<taxonomicName id="D0A04D1AFFEEE7193F6AFDDE1C94FDE4" authorityName="F. Cuvier" authorityYear="1837" box="[614,740,534,557]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">Sarcophilus</taxonomicName>
<emphasis id="25D4EA8BFFEEE7193FE7FDDE1EE8FD84" italics="true" pageId="1" pageNumber="2">har-risii</emphasis>
</emphasis>
has no living subspecies (
<bibRefCitation id="73314B68FFEEE7193CBCFDFE1C2AFD84" author="WERDELIN, L." box="[432,602,566,589]" pageId="1" pageNumber="2" pagination="1 - 27" refId="ref21822" refString="WERDELIN, L. 1987 a. Some observations on Sarcophilus laniarius and the evolution of Sarcophilus. Records of the Queen Victoria Museum, Launceston 90: 1 - 27." type="journal article" year="1987">Werdelin 1987a</bibRefCitation>
;
<bibRefCitation id="73314B68FFEEE7193F6BFDFE1C9FFD84" author="GROVES, C. P." box="[615,751,566,589]" pageId="1" pageNumber="2" refId="ref16815" refString="GROVES, C. P. 2005. Order Dasyuromorphia. Pp. 23 - 37 in Mammal species of the world: a taxonomic and geographic reference (D. E. Wilson and D. A. M. Reeder, eds.). 3 rd ed. Smithsonian Institution Press, Washington, D. C." type="book" year="2005">Groves 2005</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFEEE7193D61FDAE1F7EFC34" blockId="1.[109,793,614,1021]" pageId="1" pageNumber="2">
NOMENCLATURAL NOTES.
<taxonomicName id="D0A04D1AFFEEE7193C88FDAE1C72FDB4" authorityName="F. Cuvier" authorityYear="1837" box="[388,514,614,637]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis id="25D4EA8BFFEEE7193C88FDAE1C72FDB4" box="[388,514,614,637]" italics="true" pageId="1" pageNumber="2">Sarcophilus</emphasis>
</taxonomicName>
is from the Greek words
<emphasis id="25D4EA8BFFEEE7193D61FD4E1EEBFD54" box="[109,155,646,669]" italics="true" pageId="1" pageNumber="2">sarx</emphasis>
, meaning flesh, and
<emphasis id="25D4EA8BFFEEE7193C9EFD4E1FA3FD54" box="[402,467,646,669]" italics="true" pageId="1" pageNumber="2">philos</emphasis>
, meaning loving or fond of (
<bibRefCitation id="73314B68FFEEE7193D7AFD6E1F75FD74" author="STRAHAN, R." box="[118,261,678,701]" pageId="1" pageNumber="2" refId="ref21423" refString="STRAHAN, R. 1981. A dictionary of Australian mammal names. Angus and Robertson Publishers, Sydney, Australia." type="book" year="1981">Strahan 1981</bibRefCitation>
). The specific name
<taxonomicName id="D0A04D1AFFEEE7193CE6FD6E1C4AFD74" authorityName="Thomas" authorityYear="1912" box="[490,570,678,701]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193CE6FD6E1C4AFD74" box="[490,570,678,701]" italics="true" pageId="1" pageNumber="2">harrisii</emphasis>
</taxonomicName>
is the Latin form of “harris.”
<bibRefCitation id="73314B68FFEEE7193DDDFD0E1F16FD14" author="HARRIS, G. P." box="[209,358,710,733]" pageId="1" pageNumber="2" pagination="174 - 178" refId="ref17365" refString="HARRIS, G. P. 1808. Description of two new species of Didelphis from Van Diemen's Land. Transactions of the Linnean Society of London 9: 174 - 178." type="journal article" year="1808">Harris (1808)</bibRefCitation>
, in his description of 2 new mammals from Van Diemens Land (at Australian independence in 1901, the state name was changed to “
<collectingRegion id="D564F87BFFEEE7193CC4FCCE1C46FCD4" box="[456,566,774,797]" country="Australia" name="Tasmania" pageId="1" pageNumber="2">Tasmania</collectingRegion>
”), gives the common name as “native devil.” Aboriginal names as recorded by early European settlers include “tarrabah,” “poirinnah,” and “parloo-mer-rer.”
<bibRefCitation id="73314B68FFEEE7193DF1FCAE1FC9FCB4" author="WERDELIN, L." box="[253,441,870,893]" pageId="1" pageNumber="2" pagination="1 - 27" refId="ref21822" refString="WERDELIN, L. 1987 a. Some observations on Sarcophilus laniarius and the evolution of Sarcophilus. Records of the Queen Victoria Museum, Launceston 90: 1 - 27." type="journal article" year="1987">Werdelin (1987a)</bibRefCitation>
, on the basis of priority and size comparisons, proposed the change in name from
<taxonomicName id="D0A04D1AFFEEE7193F96FC4E1ECDFC74" authorityName="Thomas" authorityYear="1912" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193F96FC4E1ECDFC74" italics="true" pageId="1" pageNumber="2">Sarcophilus harrisii</emphasis>
</taxonomicName>
to
<taxonomicName id="D0A04D1AFFEEE7193DE2FC6E1FABFC74" baseAuthorityName="Owen" baseAuthorityYear="1838" box="[238,475,934,957]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="laniarius">
<emphasis id="25D4EA8BFFEEE7193DE2FC6E1FABFC74" box="[238,475,934,957]" italics="true" pageId="1" pageNumber="2">Sarcophilus laniarius</emphasis>
</taxonomicName>
(the butcherer); acceptance was brief (
<bibRefCitation id="73314B68FFEEE7193DEDFC0E1F1AFC14" author="GROVES, C. P." box="[225,362,966,989]" pageId="1" pageNumber="2" refId="ref16754" refString="GROVES, C. P. 1993. Order Dasyuromorphia. Pp. 29 - 37 in Mammal species of the world: a taxonomic and geographic reference (D. E. Wilson and D. A. M. Reeder, eds.). 2 nd ed. Smithsonian Institution Press, Washington, D. C." type="book" year="1993">Groves 1993</bibRefCitation>
) and
<taxonomicName id="D0A04D1AFFEEE7193CA5FC0E1C64FC14" authorityName="Thomas" authorityYear="1912" box="[425,532,966,989]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193CA5FC0E1C64FC14" box="[425,532,966,989]" italics="true" pageId="1" pageNumber="2">S. harrisii</emphasis>
</taxonomicName>
is now widely accepted (
<bibRefCitation id="73314B68FFEEE7193D7AFC2E1E8EFC34" author="GROVES, C. P." box="[118,254,998,1021]" pageId="1" pageNumber="2" refId="ref16815" refString="GROVES, C. P. 2005. Order Dasyuromorphia. Pp. 23 - 37 in Mammal species of the world: a taxonomic and geographic reference (D. E. Wilson and D. A. M. Reeder, eds.). 3 rd ed. Smithsonian Institution Press, Washington, D. C." type="book" year="2005">Groves 2005</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSection id="7E2F2F72FFEEE7193C7DFBFC1C17FAE7" pageId="1" pageNumber="2" type="multiple">
<subSubSection id="5FBA6512FFEEE7193C7DFBFC1C17FAE7" pageId="1" pageNumber="2" type="diagnosis">
<paragraph id="171F3699FFEEE7193C7DFBFC1C64FB87" blockId="1.[369,532,1076,1102]" box="[369,532,1076,1102]" pageId="1" pageNumber="2">
<heading id="4C5781F5FFEEE7193C7DFBFC1C64FB87" allCaps="true" bold="true" box="[369,532,1076,1102]" centered="true" fontSize="11" level="1" pageId="1" pageNumber="2" reason="6">
<emphasis id="25D4EA8BFFEEE7193C7DFBFC1C64FB87" bold="true" box="[369,532,1076,1102]" pageId="1" pageNumber="2">DIAGNOSIS</emphasis>
</heading>
</paragraph>
<paragraph id="171F3699FFEEE7193D94FBBF1C17FAE7" blockId="1.[109,792,1143,1326]" pageId="1" pageNumber="2">
Restricted to
<collectingRegion id="D564F87BFFEEE7193C24FBBF1FFEFB47" box="[296,398,1143,1166]" country="Australia" name="Tasmania" pageId="1" pageNumber="2">Tasmania</collectingRegion>
in recent times,
<taxonomicName id="D0A04D1AFFEEE7193F4EFBBF1D68FB47" authorityName="Thomas" authorityYear="1912" box="[578,792,1143,1166]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193F4EFBBF1D68FB47" box="[578,792,1143,1166]" italics="true" pageId="1" pageNumber="2">Sarcophilus harrisii</emphasis>
</taxonomicName>
is distinctive among dasyurids by its large size (adults&gt;
<quantity id="D0589B7CFFEEE7193FD4FB5F1D7BFB67" box="[728,779,1175,1198]" metricMagnitude="0" metricUnit="kg" metricValue="5.0" pageId="1" pageNumber="2" unit="kg" value="5.0">5 kg</quantity>
); black coat, usually with diamond-shaped markings on the chest, shoulders, or rump; massive neck and head; short broad snout; and robust bone-crunching dentition. No other living dasyurid can be confused with it on any of these criteria.
</paragraph>
</subSubSection>
</subSection>
<subSection id="7E2F2F72FFEEE71A3C1DFAA11CB5F94C" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2" type="multiple">
<subSubSection id="5FBA6512FFEEE71A3C1DFAA11CB5F94C" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2" type="description">
<paragraph id="171F3699FFEEE7193C1DFAA11C05FA4A" blockId="1.[273,629,1385,1411]" box="[273,629,1385,1411]" pageId="1" pageNumber="2">
<heading id="4C5781F5FFEEE7193C1DFAA11C05FA4A" allCaps="true" bold="true" box="[273,629,1385,1411]" centered="true" fontSize="11" level="1" pageId="1" pageNumber="2" reason="6">
<emphasis id="25D4EA8BFFEEE7193C1DFAA11C05FA4A" bold="true" box="[273,629,1385,1411]" pageId="1" pageNumber="2">GENERAL CHARACTERS</emphasis>
</heading>
</paragraph>
<paragraph id="171F3699FFEEE7193D94FA641C40F88A" blockId="1.[109,793,1452,1955]" pageId="1" pageNumber="2">
<taxonomicName id="D0A04D1AFFEEE7193D94FA641F1EFA0A" authorityName="Thomas" authorityYear="1912" box="[152,366,1452,1475]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193D94FA641F1EFA0A" box="[152,366,1452,1475]" italics="true" pageId="1" pageNumber="2">Sarcophilus harrisii</emphasis>
</taxonomicName>
is the largest living carnivorous marsupial, under the assumption that the thylacine,
<taxonomicName id="D0A04D1AFFEEE7193F53FA041EBCF9CA" baseAuthorityName="Harris" baseAuthorityYear="1808" class="Mammalia" family="Thylacinidae" genus="Thylacinus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="cynocephalus">
<emphasis id="25D4EA8BFFEEE7193F53FA041EBCF9CA" italics="true" pageId="1" pageNumber="2">Thylacinus cynocephalus</emphasis>
</taxonomicName>
, has been extinct since 1936.
<taxonomicName id="D0A04D1AFFEEE7193F05FA241C03F9CA" authorityName="Thomas" authorityYear="1912" box="[521,627,1516,1539]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193F05FA241C03F9CA" box="[521,627,1516,1539]" italics="true" pageId="1" pageNumber="2">S. harrisii</emphasis>
</taxonomicName>
has short shiny black fur (
<quantity id="D0589B7CFFEEE7193DEEF9C41F40F9EA" box="[226,304,1548,1571]" metricMagnitude="-2" metricUnit="m" metricValue="3.0" pageId="1" pageNumber="2" unit="mm" value="30.0">30 mm</quantity>
long), with soft guard hairs and sparse soft underfur. Fur in the axillary regions is thin and that on the ears, if present, is short and fine. The hairs on the tail are about
<quantity id="D0589B7CFFEEE7193FC2F9841D68F9AA" box="[718,792,1612,1635]" metricMagnitude="-2" metricUnit="m" metricValue="6.0" pageId="1" pageNumber="2" unit="mm" value="60.0">60 mm</quantity>
long. In older animals, particularly males, the fur thins out on the face, tail, and rump. Some individuals have a distinct reddish tinge to their coat color, which can be almost ginger on the tail and around the ears. Diamond-shaped white marks, found most frequently on the chest, then rump, and least on shoulders and ribs, are highly variable and individually unique. About 13% of
<taxonomicName id="D0A04D1AFFEEE7193D61F8E41EA8F88A" authorityName="Thomas" authorityYear="1912" box="[109,216,1836,1859]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193D61F8E41EA8F88A" box="[109,216,1836,1859]" italics="true" pageId="1" pageNumber="2">S. harrisii</emphasis>
</taxonomicName>
are all black (
<bibRefCitation id="73314B68FFEEE7193C7DF8E41C50F88A" author="PEMBERTON, D." box="[369,544,1836,1859]" pageId="1" pageNumber="2" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFEEE7193D94F8841BD8FE0B" blockId="1.[109,793,1452,1955]" lastBlockId="1.[840,1524,203,1954]" pageId="1" pageNumber="2">
The size of a small stocky dog,
<taxonomicName id="D0A04D1AFFEEE7193CE4F8841C22F8AA" authorityName="Thomas" authorityYear="1912" box="[488,594,1868,1891]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193CE4F8841C22F8AA" box="[488,594,1868,1891]" italics="true" pageId="1" pageNumber="2">S. harrisii</emphasis>
</taxonomicName>
has short legs, tail one-half as long as the head and body, a wide head set on a thick neck, and roundish ears (
<figureCitation id="8F9B2A1CFFEEE7193C72F8441FCDF86A" box="[382,445,1932,1955]" captionStart="Fig" captionStartId="0.[840,875,1753,1774]" captionTargetBox="[877,1485,1098,1739]" captionTargetId="figure-463@0.[877,1485,1098,1739]" captionTargetPageId="0" captionText="Fig. 1.—An alert young Sarcophilus harrisii showing black coat with white marking on chest, prominent vibrissae, wet nose, and nearly naked ears. About 13% are all black. The sexes are similar except that males" figureDoi="http://doi.org/10.5281/zenodo.4573412" httpUri="https://zenodo.org/record/4573412/files/figure.png" pageId="1" pageNumber="2">Fig. 1</figureCitation>
). Fat is stored in the tail, which may be more than
<quantity id="D0589B7CFFEEE719391AFF031A12FF2B" box="[1046,1122,203,226]" metricMagnitude="-2" metricUnit="m" metricValue="5.0" pageId="1" pageNumber="2" unit="mm" value="50.0">50 mm</quantity>
wide at the base in large males, and usually is thickest about
<quantity id="D0589B7CFFEEE719395AFF231ABAFECB" box="[1110,1226,235,258]" metricMagnitude="-2" metricUnit="m" metricValue="5.5" metricValueMax="6.0" metricValueMin="5.0" pageId="1" pageNumber="2" unit="mm" value="55.0" valueMax="60.0" valueMin="50.0">5060 mm</quantity>
from its base. Wounds and scars are more prevalent in males than in females, probably the results of aggression in competition for food and when in conflict both with mates and other competing males during copulation. Muzzle wounds can be life-threatening, with jaw muscles ripped off and teeth exposed. The tail of older animals often is heavily scarred and sometimes shortened, as if bitten off.
</paragraph>
<paragraph id="171F3699FFEEE7193E7FFE031BFCFD2B" blockId="1.[840,1524,203,1954]" pageId="1" pageNumber="2">
The large and numerous backwardly swept black vibrissae of
<taxonomicName id="D0A04D1AFFEEE7193E64FE231DA6FDCB" authorityName="Thomas" authorityYear="1912" box="[872,982,491,514]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193E64FE231DA6FDCB" box="[872,982,491,514]" italics="true" pageId="1" pageNumber="2">S. harrisii</emphasis>
</taxonomicName>
include some from the lower jaw and eyebrows. The genial vibrissae, usually about 12, arise from a nearly circular area and extend past the ears to the neck (
<bibRefCitation id="73314B68FFEEE7193825FDE31BC3FD8B" author="POCOCK, R. I." box="[1321,1459,555,578]" pageId="1" pageNumber="2" pagination="1037 - 1084" refId="ref20750" refString="POCOCK, R. I. 1926. The external characters of Thylacinus, Sarcophilus, and some related marsupials. Proceedings of the Zoology Society of London 68: 1037 - 1084." type="journal article" year="1926">Pocock 1926</bibRefCitation>
). The mystacials are equally long and rigid, and sometimes sinuous. The rhinarium is large, wide, grooved, naked, and usually wet. The ear, higher than wide and attached by a broad base to the head, has numerous ridges and folds able to cover the auditory orifice, which lies at the base of the ventral-most cleft.
</paragraph>
<paragraph id="171F3699FFEEE7193E7FFD231D8EFB6B" blockId="1.[840,1524,203,1954]" pageId="1" pageNumber="2">
The entire plantar pad of the forefoot, covered with a coarse squamous tessellation, rests on the ground (
<bibRefCitation id="73314B68FFEEE719381FFCC31BECFCEB" author="POCOCK, R. I." box="[1299,1436,779,802]" pageId="1" pageNumber="2" pagination="1037 - 1084" refId="ref20750" refString="POCOCK, R. I. 1926. The external characters of Thylacinus, Sarcophilus, and some related marsupials. Proceedings of the Zoology Society of London 68: 1037 - 1084." type="journal article" year="1926">Pocock 1926</bibRefCitation>
). Digits 2, 3, and 4 of front feet are longer than 1 and 5; all digits are relatively long with long sharp claws that are not strongly recurved. Further, the forefoot is supple and capable of grasping. The hind foot has a less flexible plantar pad that extends to the heel. The heel, an important contact point when climbing, is sometimes placed on the ground while standing but when walking or loping
<taxonomicName id="D0A04D1AFFEEE7193E78FC231D93FBCB" authorityName="Thomas" authorityYear="1912" box="[884,995,1003,1026]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193E78FC231D93FBCB" box="[884,995,1003,1026]" italics="true" pageId="1" pageNumber="2">S. harrisii</emphasis>
</taxonomicName>
is digitigrade. The hallux is extremely reduced and if visible is 0.51.0 mm long. The other digits of hind feet are long, strongly clawed, and of equal size, creating a distinctive square hind footprint. Foot pads and claws are commonly black, but are brown or even pink in those individuals with reddish “henna” fur.
</paragraph>
<paragraph id="171F3699FFEEE7193E7FFB631BD8F90B" blockId="1.[840,1524,203,1954]" pageId="1" pageNumber="2">
Sexual size dimorphism in
<taxonomicName id="D0A04D1AFFEEE7193981FB631A86FB0B" authorityName="Thomas" authorityYear="1912" box="[1165,1270,1195,1218]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193981FB631A86FB0B" box="[1165,1270,1195,1218]" italics="true" pageId="1" pageNumber="2">S. harrisii</emphasis>
</taxonomicName>
is moderate, with males 56% larger than females by weight (Jones 1997). Mean mass of
<specimenCount id="01A6FD10FFEEE7193E44FB231DD1FACB" box="[840,929,1259,1282]" pageId="1" pageNumber="2" type="adult">50 adult</specimenCount>
males was
<quantity id="D0589B7CFFEEE719392BFB231A1DFACB" box="[1063,1133,1259,1282]" metricMagnitude="0" metricUnit="kg" metricValue="8.2" pageId="1" pageNumber="2" unit="kg" value="8.2">8.2 kg</quantity>
and that of
<specimenCount id="01A6FD10FFEEE71939F5FB231B00FACB" box="[1273,1392,1259,1282]" pageId="1" pageNumber="2" type="female">50 females</specimenCount>
was
<quantity id="D0589B7CFFEEE71938A1FB231B83FACB" box="[1453,1523,1259,1282]" metricMagnitude="0" metricUnit="kg" metricValue="6.1" pageId="1" pageNumber="2" unit="kg" value="6.1">6.1 kg</quantity>
obtained from a 10-year study on the west coast of
<collectingRegion id="D564F87BFFEEE7193881FAC31B83FAEB" box="[1421,1523,1291,1314]" country="Australia" name="Tasmania" pageId="1" pageNumber="2">Tasmania</collectingRegion>
(
<bibRefCitation id="73314B68FFEEE7193E5DFAE31DA1FA8B" author="GUILER, E. R." box="[849,977,1323,1346]" pageId="1" pageNumber="2" pagination="161 - 178" refId="ref16996" refString="GUILER, E. R. 1978. Observations on the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Mammalia) at Granville Harbour. Papers and Proceedings of the Royal Society of Tasmania 112: 161 - 178." type="journal article" year="1978">Guiler 1978</bibRefCitation>
), although this sample of males probably included a mixture of fully mature and growing (2- to 3-year-old) males. Pemberton and Renouf (1993—
<emphasis id="25D4EA8BFFEEE7193995FAA31AD6FA4B" box="[1177,1190,1387,1410]" italics="true" pageId="1" pageNumber="2">n</emphasis>
≥ 100 of each sex) report males averaged
<quantity id="D0589B7CFFEEE7193EBDFA431A75FA6B" box="[945,1029,1419,1442]" metricMagnitude="1" metricUnit="kg" metricValue="1.02" pageId="1" pageNumber="2" unit="kg" value="10.2">10.2 kg</quantity>
and females
<quantity id="D0589B7CFFEEE7193990FA431A92FA6B" box="[1180,1250,1419,1442]" metricMagnitude="0" metricUnit="kg" metricValue="7.1" pageId="1" pageNumber="2" unit="kg" value="7.1">7.1 kg</quantity>
from the drier northeast coast of
<collectingRegion id="D564F87BFFEEE7193EA8FA631A7DFA0B" box="[932,1037,1451,1474]" country="Australia" name="Tasmania" pageId="1" pageNumber="2">Tasmania</collectingRegion>
.
<taxonomicName id="D0A04D1AFFEEE7193914FA631AF3FA0B" authorityName="Thomas" authorityYear="1912" box="[1048,1155,1451,1474]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193914FA631AF3FA0B" box="[1048,1155,1451,1474]" italics="true" pageId="1" pageNumber="2">S. harrisii</emphasis>
</taxonomicName>
living at high elevation or in subalpine areas is smaller; males are
<quantity id="D0589B7CFFEEE71939BBFA031B78FA2B" box="[1207,1288,1483,1506]" metricMagnitude="0" metricUnit="kg" metricValue="8.43" pageId="1" pageNumber="2" unit="kg" value="8.43">8.43 kg</quantity>
(
<emphasis id="25D4EA8BFFEEE7193815FA031B56FA2B" box="[1305,1318,1483,1506]" italics="true" pageId="1" pageNumber="2">n</emphasis>
= 56) and females are
<quantity id="D0589B7CFFEEE7193E7DFA231DC6F9CB" box="[881,950,1515,1538]" metricMagnitude="0" metricUnit="kg" metricValue="5.4" pageId="1" pageNumber="2" unit="kg" value="5.4">5.4 kg</quantity>
(
<emphasis id="25D4EA8BFFEEE7193ECBFA231DA4F9CB" box="[967,980,1515,1538]" italics="true" pageId="1" pageNumber="2">n</emphasis>
= 130—Jones 1997). Healthy animals often have substantial amounts of body fat, especially in the skin, abdomen, and tail. Average and ranges of external measurements (mm) of
<specimenCount id="01A6FD10FFEEE7193E64F9831DB1F9AB" box="[872,961,1611,1634]" pageId="1" pageNumber="2" type="male">37 male</specimenCount>
and
<specimenCount id="01A6FD10FFEEE7193EF6F9831A18F9AB" box="[1018,1128,1611,1634]" pageId="1" pageNumber="2" type="female">32 female</specimenCount>
<taxonomicName id="D0A04D1AFFEEE719397DF9831AAFF9AB" authorityName="Thomas" authorityYear="1912" box="[1137,1247,1611,1634]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE719397DF9831AAFF9AB" box="[1137,1247,1611,1634]" italics="true" pageId="1" pageNumber="2">S. harrisii</emphasis>
</taxonomicName>
, respectively, were: total length, 824.1 (750975), 780.7 (710840); length of tail, 256.2 (220285), 253.0 (215265); length of hind foot, 91.2 (82108), 86.7 (77102); length of ear, 64.2 (4780), 56.2 (4473).
</paragraph>
<paragraph id="171F3699FFEEE71A3E7FF9031BD4FB4C" blockId="1.[840,1524,203,1954]" lastBlockId="2.[840,1523,942,1157]" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">
The skull is distinguished from that of other dasyurids by its massive bone thickness, relatively short rostrum, broad zygomatic arch, pronounced sagittal crest, and expansive zygoma to accommodate powerful masseter muscles (
<figureCitation id="8F9B2A1CFFEEE7193817F8E31B2BF88B" box="[1307,1371,1835,1858]" captionStart="Fig" captionStartId="2.[109,144,1373,1394]" captionTargetBox="[211,691,200,1360]" captionTargetId="figure-381@2.[211,691,200,1360]" captionTargetPageId="2" captionText="Fig. 2.—Dorsal, ventral, and lateral views of skull and lateral view of mandible of Sarcophilus harrisii (adult male, United States National Museum, USNM 252733) collected 4 June 1928; locality unknown.The specimen exhibits the tiny cranium, typical of marsupials. Photograph by Sam Hughes used with permission." figureDoi="http://doi.org/10.5281/zenodo.4573416" httpUri="https://zenodo.org/record/4573416/files/figure.png" pageId="1" pageNumber="2">Fig. 2</figureCitation>
). Canines are massive and molars are carnassial-like. The palatine is fenestrated. Average skull measurements (mm) for
<specimenCount id="01A6FD10FFEEE7193849F8A31BEEF84B" box="[1349,1438,1899,1922]" pageId="1" pageNumber="2" type="male">18 male</specimenCount>
and
<specimenCount id="01A6FD10FFEEE71938D4F8A31DE1F86B" pageId="1" pageNumber="2" type="female">15 female</specimenCount>
<taxonomicName id="D0A04D1AFFEEE7193E97F8431A79F86B" authorityName="Thomas" authorityYear="1912" box="[923,1033,1931,1954]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEEE7193E97F8431A79F86B" box="[923,1033,1931,1954]" italics="true" pageId="1" pageNumber="2">S. harrisii</emphasis>
</taxonomicName>
, respectively, were: greatest length of skull,
<taxonomicName id="D0A04D1AFFEDE71A3E44FC661DCBFC0C" authorityName="Thomas" authorityYear="1912" box="[840,955,942,965]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEDE71A3E44FC661DCBFC0C" box="[840,955,942,965]" italics="true" pageId="2" pageNumber="3">S. harrisii</emphasis>
</taxonomicName>
disappeared from the Australian mainland about 3,000 years ago, probably due to a convergence of extreme climate events (Brüniche-Olsen et al. 2014), competition with the dingo,
<taxonomicName id="D0A04D1AFFEDE71A3E98FBC61AF9FBEC" authorityName="Meyer" authorityYear="1793" box="[916,1161,1038,1061]" class="Mammalia" family="Canidae" genus="Canis" kingdom="Animalia" order="Carnivora" pageId="2" pageNumber="3" phylum="Chordata" rank="subSpecies" species="familiaris" subSpecies="dingo">
<emphasis id="25D4EA8BFFEDE71A3E98FBC61AF9FBEC" box="[916,1161,1038,1061]" italics="true" pageId="2" pageNumber="3">Canis familiaris dingo</emphasis>
</taxonomicName>
(Jones et al. 2003a;
<bibRefCitation id="73314B68FFEDE71A3863FBC61D0DFB8C" author="LETNIC, M. &amp; M. FILLIOS &amp; M. S. CROWTHER" pageId="2" pageNumber="3" pagination="34877" refId="ref19337" refString="LETNIC, M., M. FILLIOS, AND M. S. CROWTHER. 2012. Could direct killing by larger dingoes have caused the extinction of the thylacine from mainland Australia? PLoS ONE 7: e 34877." type="journal article" year="2012">Letnic et al. 2012</bibRefCitation>
), and an increase in the population density (
<bibRefCitation id="73314B68FFEDE71A386AFBE61D0DFBAC" author="PROWSE, T. A. A. &amp; C. N. JOHNSON &amp; C. J. A. BRADSHAW &amp; B. W. BROOK" pageId="2" pageNumber="3" pagination="693 - 702" refId="ref20784" refString="PROWSE, T. A. A., C. N. JOHNSON, C. J. A. BRADSHAW, AND B. W. BROOK. 2014. An ecological regime shift resulting from disrupted predatorprey interactions in Holocene Australia. Ecology 95: 693 - 702." type="journal article" year="2014">Prowse et al. 2014</bibRefCitation>
) of Aboriginal people, who also developed new hunting technologies around this time (Johnson and Wroe 2003).
</paragraph>
<caption id="43DF6611FFEDE71A3E44FD251A8DFCBE" ID-DOI="http://doi.org/10.5281/zenodo.4573414" ID-Zenodo-Dep="4573414" httpUri="https://zenodo.org/record/4573414/files/figure.png" pageId="2" pageNumber="3" startId="2.[840,875,749,770]" targetBox="[893,1467,201,735]" targetPageId="2">
<paragraph id="171F3699FFEDE71A3E44FD251A8DFCBE" blockId="2.[840,1523,749,887]" pageId="2" pageNumber="3">
<emphasis id="25D4EA8BFFEDE71A3E44FD251DF8FCCB" bold="true" box="[840,904,749,770]" pageId="2" pageNumber="3">Fig. 3.</emphasis>
—The distribution of
<taxonomicName id="D0A04D1AFFEDE71A3951FD251B6DFCCB" authorityName="Thomas" authorityYear="1912" box="[1117,1309,749,770]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEDE71A3951FD251B6DFCCB" box="[1117,1309,749,770]" italics="true" pageId="2" pageNumber="3">Sarcophilus harrisii</emphasis>
</taxonomicName>
has been restricted to the island state of Tasmania, Australia, in historic times. The land area of the larger island of Tasmania, 64,519 km
<superScript id="E0D59BD1FFEDE71A39FBFCEE1A8EFCFA" attach="left" box="[1271,1278,806,819]" fontSize="5" pageId="2" pageNumber="3">2</superScript>
, is approximately equivalent to that of the states of West Virginia or South Carolina, United States. Note the temperate location, 4142°S.
</paragraph>
</caption>
<caption id="43DF6611FFEDE71A3D61FA951FAFFA21" ID-DOI="http://doi.org/10.5281/zenodo.4573416" ID-Zenodo-Dep="4573416" httpUri="https://zenodo.org/record/4573416/files/figure.png" pageId="2" pageNumber="3" startId="2.[109,144,1373,1394]" targetBox="[211,691,200,1360]" targetPageId="2">
<paragraph id="171F3699FFEDE71A3D61FA951FAFFA21" blockId="2.[109,792,1373,1512]" pageId="2" pageNumber="3">
<emphasis id="25D4EA8BFFEDE71A3D61FA951EDEFABB" bold="true" box="[109,174,1373,1394]" pageId="2" pageNumber="3">Fig. 2.</emphasis>
—Dorsal, ventral, and lateral views of skull and lateral view of mandible of
<taxonomicName id="D0A04D1AFFEDE71A3DE1FAB31FC0FA59" authorityName="Thomas" authorityYear="1912" box="[237,432,1403,1424]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEDE71A3DE1FAB31FC0FA59" box="[237,432,1403,1424]" italics="true" pageId="2" pageNumber="3">Sarcophilus harrisii</emphasis>
</taxonomicName>
(adult male, United States National Museum, USNM 252733) collected 4 June 1928; locality unknown. The specimen exhibits the tiny cranium, typical of marsupials. Photograph by Sam Hughes used with permission.
</paragraph>
</caption>
<paragraph id="171F3699FFEDE71A3D61F9C61CB5F94C" blockId="2.[109,792,1550,1669]" pageId="2" pageNumber="3">137.9, 130.3; cranial breadth, 20.0, 20.8; zygomatic breadth, 110.7, 98.0; length of maxillary toothrow, 48.2, 49.0; length of molariform teeth in mandibular toothrow, 51.8, 53.5; maximal anterior-posterior diameter of upper canine, 10.03, 8.92.</paragraph>
</subSubSection>
</subSection>
<subSection id="7E2F2F72FFEDE71A3C5AF9081D68F853" pageId="2" pageNumber="3" type="multiple">
<subSubSection id="5FBA6512FFEDE71A3C5AF9081D68F853" pageId="2" pageNumber="3" type="distribution">
<paragraph id="171F3699FFEDE71A3C5AF9081C5EF913" blockId="2.[342,558,1728,1754]" box="[342,558,1728,1754]" pageId="2" pageNumber="3">
<heading id="4C5781F5FFEDE71A3C5AF9081C5EF913" allCaps="true" bold="true" box="[342,558,1728,1754]" centered="true" fontSize="11" level="1" pageId="2" pageNumber="3" reason="6">
<emphasis id="25D4EA8BFFEDE71A3C5AF9081C5EF913" bold="true" box="[342,558,1728,1754]" pageId="2" pageNumber="3">DISTRIBUTION</emphasis>
</heading>
</paragraph>
<paragraph id="171F3699FFEDE71A3D94F8CB1D68F853" blockId="2.[109,792,1795,1946]" pageId="2" pageNumber="3">
Distribution is restricted to
<collectingRegion id="D564F87BFFEDE71A3CB7F8CB1C53F8D3" box="[443,547,1795,1818]" country="Australia" name="Tasmania" pageId="2" pageNumber="3">Tasmania</collectingRegion>
, the island state off the southeast coast of mainland
<collectingCountry id="6FB77609FFEDE71A3C95F8EB1F8BF8F3" box="[409,507,1827,1850]" name="Australia" pageId="2" pageNumber="3">Australia</collectingCountry>
(
<figureCitation id="8F9B2A1CFFEDE71A3F06F8EB1C37F8F3" box="[522,583,1827,1850]" captionStart="Fig" captionStartId="2.[840,875,749,770]" captionTargetBox="[893,1467,201,735]" captionTargetId="figure-429@2.[893,1469,200,736]" captionTargetPageId="2" captionText="Fig. 3.—The distribution of Sarcophilus harrisii has been restricted to the island state of Tasmania, Australia, in historic times. The land area of the larger island of Tasmania, 64,519 km2, is approximately equivalent to that of the states of West Virginia or South Carolina, United States. Note the temperate location, 4142°S." figureDoi="http://doi.org/10.5281/zenodo.4573414" httpUri="https://zenodo.org/record/4573414/files/figure.png" pageId="2" pageNumber="3">Fig. 3</figureCitation>
). Within
<collectingRegion id="D564F87BFFEDE71A3FA0F8EB1D64F8F3" box="[684,788,1827,1850]" country="Australia" name="Tasmania" pageId="2" pageNumber="3">Tasmania</collectingRegion>
,
<taxonomicName id="D0A04D1AFFEDE71A3D61F88C1F32F892" authorityName="Thomas" authorityYear="1912" box="[109,322,1860,1883]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEDE71A3D61F88C1F32F892" box="[109,322,1860,1883]" italics="true" pageId="2" pageNumber="3">Sarcophilus harrisii</emphasis>
</taxonomicName>
is among the most widely distributed mammals, 3rd behind 2 macropods (Macropodidae—Rounsevell et al. 1991). Formerly widespread across southern
<collectingCountry id="6FB77609FFEDE71A3FBCF84B1D65F853" box="[688,789,1923,1946]" name="Australia" pageId="2" pageNumber="3">Australia</collectingCountry>
,
</paragraph>
</subSubSection>
</subSection>
<subSubSection id="5FBA6512FFEDE71E392AFB771B06F9CB" lastPageId="6" lastPageNumber="7" pageId="2" pageNumber="3" type="description">
<subSection id="7E2F2F72FFEDE71B392AFB771F4CFD2B" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3" type="multiple">
<paragraph id="171F3699FFEDE71A392AFB771B65FB10" blockId="2.[1062,1301,1215,1241]" box="[1062,1301,1215,1241]" pageId="2" pageNumber="3">
<heading id="4C5781F5FFEDE71A392AFB771B65FB10" allCaps="true" bold="true" box="[1062,1301,1215,1241]" centered="true" fontSize="11" level="1" pageId="2" pageNumber="3" reason="6">
<emphasis id="25D4EA8BFFEDE71A392AFB771B65FB10" bold="true" box="[1062,1301,1215,1241]" pageId="2" pageNumber="3">FOSSIL RECORD</emphasis>
</heading>
</paragraph>
<paragraph id="171F3699FFEDE71B3E7FFACB1F97FE6B" blockId="2.[840,1523,1283,1946]" lastBlockId="3.[109,792,203,738]" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3">
The recent ancestry of
<taxonomicName id="D0A04D1AFFEDE71A3966FACB1B30FAD3" authorityName="Thomas" authorityYear="1912" box="[1130,1344,1283,1306]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEDE71A3966FACB1B30FAD3" box="[1130,1344,1283,1306]" italics="true" pageId="2" pageNumber="3">Sarcophilus harrisii</emphasis>
</taxonomicName>
can be traced to several locations of Pleistocene age (Archer and Hand 1984), the oldest of which is from the early Pleistocene of Nelson Bay,
<collectingRegion id="D564F87BFFEDE71A3E44FAAB1DEEFAB3" box="[840,926,1379,1402]" country="Australia" name="Victoria" pageId="2" pageNumber="3">Victoria</collectingRegion>
(Gerdtz and Archbold 2003). Local faunas of
<geoCoordinate id="7294505EFFEDE71A3890FAAB1B95FAB3" box="[1436,1509,1379,1402]" degrees="70,000" direction="north" orientation="latitude" pageId="2" pageNumber="3" precision="55" value="70.0">70,000</geoCoordinate>
<geoCoordinate id="7294505EFFEDE71A38E9FAAB1DE1FA53" degrees="30,000" direction="west" orientation="longitude" pageId="2" pageNumber="3" precision="55" value="-30.0"> 30,000</geoCoordinate>
years ago with
<taxonomicName id="D0A04D1AFFEDE71A396DFA4B1AAFFA53" authorityName="F. Cuvier" authorityYear="1837" box="[1121,1247,1411,1434]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="25D4EA8BFFEDE71A396DFA4B1AAFFA53" box="[1121,1247,1411,1434]" italics="true" pageId="2" pageNumber="3">Sarcophilus</emphasis>
</taxonomicName>
fossils include Eastern Darling Downs (southeastern
<collectingRegion id="D564F87BFFEDE71A39BDFA6B1B43FA73" box="[1201,1331,1443,1466]" country="Australia" name="Queensland" pageId="2" pageNumber="3">Queensland</collectingRegion>
),
<collectingRegion id="D564F87BFFEDE71A385EFA6B1BD8FA73" box="[1362,1448,1443,1466]" country="Australia" name="Victoria" pageId="2" pageNumber="3">Victoria</collectingRegion>
Cave (southeastern
<collectingRegion id="D564F87BFFEDE71A3EEDFA0B1AFFFA13" box="[993,1167,1475,1498]" country="Australia" name="South Australia" pageId="2" pageNumber="3">South Australia</collectingRegion>
), and Mammoth Cave (
<collectingRegion id="D564F87BFFEDE71A3896FA0B1DDFFA33" country="Australia" name="Western Australia" pageId="2" pageNumber="3">Western Australia</collectingRegion>
). A late Quarternary record of
<taxonomicName id="D0A04D1AFFEDE71A3807FA2B1BF9FA33" authorityName="F. Cuvier" authorityYear="1837" box="[1291,1417,1507,1530]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="25D4EA8BFFEDE71A3807FA2B1BF9FA33" box="[1291,1417,1507,1530]" italics="true" pageId="2" pageNumber="3">Sarcophilus</emphasis>
</taxonomicName>
from the Mygoora Local Fauna, from Mygoora Lake in southern
<collectingRegion id="D564F87BFFEDE71A389EF9CB1DD8F9F3" country="Australia" name="Northern Territory" pageId="2" pageNumber="3">Northern Territory</collectingRegion>
, is the 1st record from central
<collectingCountry id="6FB77609FFEDE71A39EBF9EB1B39F9F3" box="[1255,1353,1571,1594]" name="Australia" pageId="2" pageNumber="3">Australia</collectingCountry>
(
<bibRefCitation id="73314B68FFEDE71A3854F9EB1D0DF993" author="MEGIRIAN, D. &amp; P. F. MURRAY &amp; P. K. LATZ &amp; K. A. JOHNSON" pageId="2" pageNumber="3" pagination="77 - 93" refId="ref19862" refString="MEGIRIAN, D., P. F. MURRAY, P. K. LATZ, AND K. A. JOHNSON. 2002. The Mygoora Local Fauna: a late Quarternary vertebrate assemblage from central Australia. The Beagle (Records of the Museums and Art Galleries of the Northern Territory) 18: 77 - 93." type="journal article" year="2002">Megirian et al. 2002</bibRefCitation>
). More recent remains are from the Devils Lair Local Fauna (southwestern
<collectingRegion id="D564F87BFFEDE71A3EEAF9AB1ADAF9B3" box="[998,1194,1635,1658]" country="Australia" name="Western Australia" pageId="2" pageNumber="3">Western Australia</collectingRegion>
,
<geoCoordinate id="7294505EFFEDE71A39BAF9AB1A8FF9B3" box="[1206,1279,1635,1658]" degrees="35,000" direction="north" orientation="latitude" pageId="2" pageNumber="3" precision="55" value="35.0">35,000</geoCoordinate>
<geoCoordinate id="7294505EFFEDE71A39F3F9AB1B39F9B3" box="[1279,1353,1635,1658]" degrees="2,000" direction="west" orientation="longitude" pageId="2" pageNumber="3" precision="55" value="-2.0">
<emphasis id="25D4EA8BFFEDE71A39F3F9AB1B7CF9B3" box="[1279,1292,1635,1658]" italics="true" pageId="2" pageNumber="3"></emphasis>
2,000
</geoCoordinate>
years ago) and Fromms Landing Local Fauna (
<collectingRegion id="D564F87BFFEDE71A39ABF94B1B23F953" box="[1191,1363,1667,1690]" country="Australia" name="South Australia" pageId="2" pageNumber="3">South Australia</collectingRegion>
, &lt;4,000 years ago).
<taxonomicName id="D0A04D1AFFEDE71A3E89F96B1D9FF973" authorityName="Thomas" authorityYear="1912" box="[901,1007,1699,1722]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEDE71A3E89F96B1D9FF973" box="[901,1007,1699,1722]" italics="true" pageId="2" pageNumber="3">S. harrisii</emphasis>
</taxonomicName>
also is known (from one i3) from the upper layers of Pyramids Cave in eastern
<collectingRegion id="D564F87BFFEDE71A39AFF90B1A8CF913" box="[1187,1276,1731,1754]" country="Australia" name="Victoria" pageId="2" pageNumber="3">Victoria</collectingRegion>
, a deposit of Holocene age (
<bibRefCitation id="73314B68FFEDE71A3E8CF92B1A47F933" author="WAKEFIELD, N. A." box="[896,1079,1763,1786]" pageId="2" pageNumber="3" pagination="211 - 214" refId="ref21689" refString="WAKEFIELD, N. A. 1967 a. Mammal bones in the Buchan district. Victorian Naturalist 84: 211 - 214." type="journal article" year="1967">Wakefield 1967a</bibRefCitation>
). Another Holocene (3,120 ± 100 years ago) record is from the Padypadiy archeological site in
<collectingRegion id="D564F87BFFEDE71A389EF8CB1DD8F8F3" country="Australia" name="Northern Territory" pageId="2" pageNumber="3">Northern Territory</collectingRegion>
. A 2nd record from northern
<collectingCountry id="6FB77609FFEDE71A39F2F8EB1B10F8F3" box="[1278,1376,1827,1850]" name="Australia" pageId="2" pageNumber="3">Australia</collectingCountry>
(Calaby and White 1967) is a skull fragment from Cape York,
<collectingRegion id="D564F87BFFEDE71A3862F88B1B9FF893" box="[1390,1519,1859,1882]" country="Australia" name="Queensland" pageId="2" pageNumber="3">Queensland</collectingRegion>
, dated
<geoCoordinate id="7294505EFFEDE71A3E80F8AB1DA5F8B3" box="[908,981,1891,1914]" degrees="11,000" direction="north" orientation="latitude" pageId="2" pageNumber="3" precision="55" value="11.0">11,000</geoCoordinate>
<geoCoordinate id="7294505EFFEDE71A3ED9F8AB1A5CF8B3" box="[981,1068,1891,1914]" degrees="10,000" direction="west" orientation="longitude" pageId="2" pageNumber="3" precision="55" value="-10.0">10,000</geoCoordinate>
years ago (
<bibRefCitation id="73314B68FFEDE71A39B8F8AB1B4FF8B3" author="HORTON, D. R." box="[1204,1343,1891,1914]" pageId="2" pageNumber="3" pagination="374 - 375" refId="ref17831" refString="HORTON, D. R. 1977. A 10,000 - yr old Sarcophilus from Cape York. Search 8: 374 - 375." type="journal article" year="1977">Horton 1977</bibRefCitation>
). Subfossils are known from 4 sites on the Basalt Plains of western
<collectingRegion id="D564F87BFFEDE71A386AF84B1BCFF853" box="[1382,1471,1923,1946]" country="Australia" name="Victoria" pageId="2" pageNumber="3">Victoria</collectingRegion>
, 2 of which are from Aboriginal middens; another (cave) site yielded an estimated 60 individuals (
<bibRefCitation id="73314B68FFECE71B3C92FF231C34FECB" author="WAKEFIELD, N. A." box="[414,580,235,258]" pageId="3" pageNumber="4" pagination="419 - 425" refId="ref21658" refString="WAKEFIELD, N. A. 1964. Recent mammalian sub-fossils of the Basalt Plains of Victoria. Proceedings of the Royal Society of Victoria 77: 419 - 425." type="journal article" year="1964">Wakefield 1964</bibRefCitation>
). Still other subfossils were found in McEacherns Cave in extreme southwestern
<collectingRegion id="D564F87BFFECE71B3D61FEE31EB3FE8B" box="[109,195,299,322]" country="Australia" name="Victoria" pageId="3" pageNumber="4">Victoria</collectingRegion>
(
<bibRefCitation id="73314B68FFECE71B3DD9FEE31FFBFE8B" author="WAKEFIELD, N. A." box="[213,395,299,322]" pageId="3" pageNumber="4" pagination="363 - 383" refId="ref21713" refString="WAKEFIELD, N. A. 1967 b. Preliminary report on McEachern's Cave, S. W. Victoria. Victorian Naturalist 84: 363 - 383." type="journal article" year="1967">Wakefield 1967b</bibRefCitation>
). Although no
<taxonomicName id="D0A04D1AFFECE71B3F3EFEE31CEFFE8B" authorityName="Thomas" authorityYear="1912" box="[562,671,299,322]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B3F3EFEE31CEFFE8B" box="[562,671,299,322]" italics="true" pageId="3" pageNumber="4">S. harrisii</emphasis>
</taxonomicName>
survive on any island of Bass Strait north of
<collectingRegion id="D564F87BFFECE71B3CEAFE831C3EFEAB" box="[486,590,331,354]" country="Australia" name="Tasmania" pageId="3" pageNumber="4">Tasmania</collectingRegion>
, subfossil remains from the Palana sand dunes on Flinders Island reveal that it lived there in the Holocene (
<bibRefCitation id="73314B68FFECE71B3C6EFE431FA7FE6B" author="HOPE, J. L." box="[354,471,395,418]" pageId="3" pageNumber="4" pagination="163 - 196" refId="ref17803" refString="HOPE, J. L. 1973. Mammals of the Bass Strait Islands. Proceedings of the Royal Society of Victoria 85: 163 - 196." type="journal article" year="1973">Hope 1973</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFECE71B3D94FE631F4CFD2B" blockId="3.[109,792,203,738]" pageId="3" pageNumber="4">
The
<typeStatus id="C81B883BFFECE71B3DCBFE631E85FE0B" box="[199,245,427,450]" pageId="3" pageNumber="4">type</typeStatus>
specimens (2 broken left maxillae) of the fossil subspecies
<taxonomicName id="D0A04D1AFFECE71B3DC8FE031F54FE2B" baseAuthorityName="Owen" baseAuthorityYear="1838" box="[196,292,459,482]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="laniarius">
<emphasis id="25D4EA8BFFECE71B3DC8FE031F54FE2B" box="[196,292,459,482]" italics="true" pageId="3" pageNumber="4">laniarius</emphasis>
</taxonomicName>
, found among fossils from
<collectingRegion id="D564F87BFFECE71B3F54FE031CBEFE2B" box="[600,718,459,482]" country="New Zealand" name="Wellington" pageId="3" pageNumber="4">Wellington</collectingRegion>
Caves (
<bibRefCitation id="73314B68FFECE71B3D7AFE231E86FDCB" author="OWEN, R." box="[118,246,491,514]" pageId="3" pageNumber="4" refId="ref20285" refString="OWEN, R. 1838. in T. L. Mitchell, Three expeditions into the interior of eastern Australia, with descriptions of the recently explored region of Australia Felix and of the present colony of New South Wales. T &amp; W. Boone, London, United Kingdom." type="book" year="1838">Owen 1838</bibRefCitation>
;
<bibRefCitation id="73314B68FFECE71B3C04FE231FEFFDCB" author="DAWSON, L." box="[264,415,491,514]" pageId="3" pageNumber="4" refId="ref15511" refString="DAWSON, L. 1982. Taxonomic status of fossil devils (Sarcophilus, Dasyuridae, Marsupialia) from late Quaternary eastern Australia localities. Pp. 517 - 525 in Carnivorous marsupials (M. Archer, ed.). Surrey Beatty &amp; Sons Pty Limited, Chipping Norton, New South Wales, Australia." type="book" year="1982">Dawson 1982</bibRefCitation>
) and later from other Pleistocene sites (
<bibRefCitation id="73314B68FFECE71B3DA2FDC31F39FDEB" author="MARSHALL, L. G." box="[174,329,523,546]" pageId="3" pageNumber="4" pagination="151 - 172" refId="ref19587" refString="MARSHALL, L. G. 1973. Fossil vertebrate faunas from the Lake Victoria region, S. W. New South Wales, Australia. Memoirs of the National Museum of Victoria 34: 151 - 172." type="journal article" year="1973">Marshall 1973</bibRefCitation>
), were determined by
<bibRefCitation id="73314B68FFECE71B3F33FDC31C8BFDEB" author="WERDELIN, L." box="[575,763,523,546]" pageId="3" pageNumber="4" pagination="1 - 27" refId="ref21822" refString="WERDELIN, L. 1987 a. Some observations on Sarcophilus laniarius and the evolution of Sarcophilus. Records of the Queen Victoria Museum, Launceston 90: 1 - 27." type="journal article" year="1987">Werdelin (1987a)</bibRefCitation>
to be 16% larger than
<taxonomicName id="D0A04D1AFFECE71B3C4EFDE31FDDFD8B" authorityName="Thomas" authorityYear="1912" box="[322,429,555,578]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B3C4EFDE31FDDFD8B" box="[322,429,555,578]" italics="true" pageId="3" pageNumber="4">S. harrisii</emphasis>
</taxonomicName>
, but were not sufficiently distinct to merit a different species designation.
<taxonomicName id="D0A04D1AFFECE71B3F14FD831CF3FDAB" authorityName="Thomas" authorityYear="1912" box="[536,643,587,610]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B3F14FD831CF3FDAB" box="[536,643,587,610]" italics="true" pageId="3" pageNumber="4">S. harrisii</emphasis>
</taxonomicName>
survives only in
<collectingRegion id="D564F87BFFECE71B3D87FDA31E84FD4B" box="[139,244,619,642]" country="Australia" name="Tasmania" pageId="3" pageNumber="4">Tasmania</collectingRegion>
. Because subfossil specimens from Mt. Hamilton,
<collectingRegion id="D564F87BFFECE71B3D61FD431EB6FD6B" box="[109,198,651,674]" country="Australia" name="Victoria" pageId="3" pageNumber="4">Victoria</collectingRegion>
, differ significantly in a number of dental and skull characteristics from the modern form,
<bibRefCitation id="73314B68FFECE71B3F01FD631CB9FD0B" author="WERDELIN, L." box="[525,713,683,706]" pageId="3" pageNumber="4" pagination="1 - 27" refId="ref21822" refString="WERDELIN, L. 1987 a. Some observations on Sarcophilus laniarius and the evolution of Sarcophilus. Records of the Queen Victoria Museum, Launceston 90: 1 - 27." type="journal article" year="1987">Werdelin (1987a)</bibRefCitation>
named these
<emphasis id="25D4EA8BFFECE71B3DA7FD031F44FD2B" box="[171,308,715,738]" italics="true" pageId="3" pageNumber="4">S. h. dixonae</emphasis>
.
</paragraph>
</subSection>
<subSection id="7E2F2F72FFECE71D3C2DFCE11F9DF90B" lastPageId="5" lastPageNumber="6" pageId="3" pageNumber="4" type="multiple">
<paragraph id="171F3699FFECE71B3C2DFCE11C14FC8A" blockId="3.[289,612,809,835]" box="[289,612,809,835]" pageId="3" pageNumber="4">
<heading id="4C5781F5FFECE71B3C2DFCE11C14FC8A" allCaps="true" bold="true" box="[289,612,809,835]" centered="true" fontSize="11" level="1" pageId="3" pageNumber="4" reason="6">
<emphasis id="25D4EA8BFFECE71B3C2DFCE11C14FC8A" bold="true" box="[289,612,809,835]" pageId="3" pageNumber="4">FORM AND FUNCTION</emphasis>
</heading>
</paragraph>
<paragraph id="171F3699FFECE71B3D94FCA51C55F8AD" blockId="3.[109,793,877,1956]" pageId="3" pageNumber="4">
The dental formula is i 4/3, c 1/1, p 2/2, m 4/4, total 42 (Green and Rainbird 2015). The dentition reflects the carnivorous diet of
<taxonomicName id="D0A04D1AFFECE71B3DFEFC651FB8FC0D" authorityName="Thomas" authorityYear="1912" box="[242,456,941,964]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B3DFEFC651FB8FC0D" box="[242,456,941,964]" italics="true" pageId="3" pageNumber="4">Sarcophilus harrisii</emphasis>
</taxonomicName>
, but is fundamentally different from that of the Order
<taxonomicName id="D0A04D1AFFECE71B3C68FC051FA1FC2D" authorityName="Bowdich" authorityYear="1821" box="[356,465,973,996]" class="Mammalia" kingdom="Animalia" order="Carnivora" pageId="3" pageNumber="4" phylum="Chordata" rank="order">Carnivora</taxonomicName>
. The incisor row is transverse. The massive canine teeth are circular in cross-section, equivalent among the placental carnivores only to some ursids and large mustelids, and are immensely strong and well adapted for eating bone and perhaps grasping large prey (Jones 2003a). Premolar teeth are of modest size, but molar teeth are massive; tooth and cusp size increase from M1 to M4. Like all dasyuroids,
<taxonomicName id="D0A04D1AFFECE71B3FC3FB451EE8FB0D" authorityName="Thomas" authorityYear="1912" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B3FC3FB451EE8FB0D" italics="true" pageId="3" pageNumber="4">S. harrisii</emphasis>
</taxonomicName>
lacks the specialized carnassial pair of the
<taxonomicName id="D0A04D1AFFECE71B3F71FB651C9AFB0D" authorityName="Bowdich" authorityYear="1821" box="[637,746,1197,1220]" class="Mammalia" kingdom="Animalia" order="Carnivora" pageId="3" pageNumber="4" phylum="Chordata" rank="order">Carnivora</taxonomicName>
. All 4 molars are similar in structure, because of phylogenetic constraints on tooth eruption and the resultant differences from other carnivorous marsupials in jaw geometry (
<bibRefCitation id="73314B68FFECE71B3F33FAC51CB9FAED" author="ARCHER, M." box="[575,713,1293,1316]" pageId="3" pageNumber="4" pagination="1 - 34" refId="ref14652" refString="ARCHER, M. 1976. The dasyurid dentition and its relationships to that of didelphids, thylacinids, borhyaenids (Marsupicarnivora) and peramelids (Peramelina: Marsupialia). Australian Journal of Zoology, Supplementary Series 39: 1 - 34." type="book chapter" year="1976">Archer 1976</bibRefCitation>
,
<bibRefCitation id="73314B68FFECE71B3FD7FAC51D60FAED" author="ARCHER, M." box="[731,784,1293,1316]" pageId="3" pageNumber="4" pagination="157 - 164" refId="ref14696" refString="ARCHER, M. 1978. The nature of the molar-premolar boundary in marsupials and a reinterpretation of the homology of marsupial cheekteeth. Memoirs of the Queensland Museum 18: 157 - 164." type="journal article" year="1978">1978</bibRefCitation>
;
<bibRefCitation id="73314B68FFECE71B3D61FAE51F6AFA8D" author="WERDELIN, L." box="[109,282,1325,1348]" pageId="3" pageNumber="4" pagination="342 - 350" refId="ref21854" refString="WERDELIN, L. 1987 b. Jaw geometry and molar morphology in marsupial carnivores: analysis of a constraint and its macroevolutionary consequences. Paleobiology 13: 342 - 350." type="journal article" year="1987">Werdelin 1987b</bibRefCitation>
). All molar teeth have meat-slicing and grinding areas, functions performed by carnassial and post-carnassial molars, respectively, of the
<taxonomicName id="D0A04D1AFFECE71B3C9EFAA51F8FFA4D" authorityName="Bowdich" authorityYear="1821" box="[402,511,1389,1412]" class="Mammalia" kingdom="Animalia" order="Carnivora" pageId="3" pageNumber="4" phylum="Chordata" rank="order">Carnivora</taxonomicName>
. The functional equivalent of the carnassial blade and notch in molar teeth of
<taxonomicName id="D0A04D1AFFECE71B3F9FFA451C8FFA6D" authorityName="Thomas" authorityYear="1912" box="[659,767,1421,1444]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B3F9FFA451C8FFA6D" box="[659,767,1421,1444]" italics="true" pageId="3" pageNumber="4">S. harrisii</emphasis>
</taxonomicName>
is the paracristid crest, which is in a similar functional position, on the outer anterior face of the lower molar tooth, to the placental carnassial blade (
<bibRefCitation id="73314B68FFECE71B3C27FA251FC3F9CD" author="ARCHER, M." box="[299,435,1517,1540]" pageId="3" pageNumber="4" pagination="1 - 34" refId="ref14652" refString="ARCHER, M. 1976. The dasyurid dentition and its relationships to that of didelphids, thylacinids, borhyaenids (Marsupicarnivora) and peramelids (Peramelina: Marsupialia). Australian Journal of Zoology, Supplementary Series 39: 1 - 34." type="book chapter" year="1976">Archer 1976</bibRefCitation>
,
<bibRefCitation id="73314B68FFECE71B3CC8FA251F89F9CD" author="ARCHER, M." box="[452,505,1517,1540]" pageId="3" pageNumber="4" pagination="157 - 164" refId="ref14696" refString="ARCHER, M. 1978. The nature of the molar-premolar boundary in marsupials and a reinterpretation of the homology of marsupial cheekteeth. Memoirs of the Queensland Museum 18: 157 - 164." type="journal article" year="1978">1978</bibRefCitation>
). Although the elongation and longitudinal orientation of the paracristid result in one long “carnassial” shearing unit (
<bibRefCitation id="73314B68FFECE71B3C82F9E51C64F98D" author="ARCHER, M." box="[398,532,1581,1604]" pageId="3" pageNumber="4" pagination="1 - 34" refId="ref14652" refString="ARCHER, M. 1976. The dasyurid dentition and its relationships to that of didelphids, thylacinids, borhyaenids (Marsupicarnivora) and peramelids (Peramelina: Marsupialia). Australian Journal of Zoology, Supplementary Series 39: 1 - 34." type="book chapter" year="1976">Archer 1976</bibRefCitation>
), the equivalent tooth to the placental carnassial is M4, which is in a geometrically similar position (
<bibRefCitation id="73314B68FFECE71B3DFAF9A51FE3F94D" author="WERDELIN, L." box="[246,403,1645,1668]" pageId="3" pageNumber="4" pagination="109 - 117" refId="ref21796" refString="WERDELIN, L. 1986. Comparison of skull shape in marsupial and placental carnivores. Australian Journal of Zoology 34: 109 - 117." type="journal article" year="1986">Werdelin 1986</bibRefCitation>
).
<taxonomicName id="D0A04D1AFFECE71B3CA6F9A51C65F94D" authorityName="Thomas" authorityYear="1912" box="[426,533,1645,1668]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B3CA6F9A51C65F94D" box="[426,533,1645,1668]" italics="true" pageId="3" pageNumber="4">S. harrisii</emphasis>
</taxonomicName>
is equivalent to the thylacines in its adaptation to carnivory, and the grinding area of the tooth is almost absent in M4, which remains sharp throughout life (Jones 1997). Processing of bones in
<taxonomicName id="D0A04D1AFFECE71B3F3FF9051CD0F92D" authorityName="Thomas" authorityYear="1912" box="[563,672,1741,1764]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B3F3FF9051CD0F92D" box="[563,672,1741,1764]" italics="true" pageId="3" pageNumber="4">S. harrisii</emphasis>
</taxonomicName>
, for which hyenas (
<taxonomicName id="D0A04D1AFFECE71B3DCBF9251F4CF8CD" authorityName="Gray" authorityYear="1821" box="[199,316,1773,1796]" class="Mammalia" family="Hyaenidae" kingdom="Animalia" order="Carnivora" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Hyaenidae</taxonomicName>
) have a specialized conical 3rd premolar, is carried out on M2, which is biomechanically in an equivalent part of the jaw (Jones 2003a) and which becomes extremely worn in older animals (
<bibRefCitation id="73314B68FFECE71B3C6AF8851C65F8AD" author="PEMBERTON, D." box="[358,533,1869,1892]" pageId="3" pageNumber="4" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFECE71B3D94F8A51A0EFDCB" blockId="3.[109,793,877,1956]" lastBlockId="3.[840,1523,203,1954]" pageId="3" pageNumber="4">
All teeth have closed roots, and hence animals can be aged by degree of tooth wear, generally into yearling, 2, 3, 4, 5 and more than 5-year-old age classes. Canine teeth overerupt and this feature can also be used to age animals. Broken incisor and canine teeth are common in older animals, occurring twice as frequently as expected if all teeth have an equal chance of fracture (Jones and Stoddart 1998). Supernumerary teeth, including incisors, canines, and molars, occur occasionally: a 5th upper right molar in 4 of 33 skulls (
<bibRefCitation id="73314B68FFECE71B399EFE431B61FE6B" author="GREEN, R. H." box="[1170,1297,395,418]" pageId="3" pageNumber="4" pagination="1 - 12" refId="ref16344" refString="GREEN, R. H. 1967. Notes on the devil (Sarcophilus harrisii) and the quoll (Dasyurus viverrinus) in north-eastern Tasmania. Records of the Queen Victoria Museum 27: 1 - 12." type="journal article" year="1967">Green 1967</bibRefCitation>
) and supernumerary partially erupted lower molars (1 on each side) in a young female (
<bibRefCitation id="73314B68FFECE71B3E5DFE031DBEFE2B" author="DORAN, A." box="[849,974,459,482]" pageId="3" pageNumber="4" pagination="379 - 381" refId="ref15732" refString="DORAN, A. 1975. A note on tooth anomalies in the Tasmanian devil, Sarcophilus harrisii. Australian Mammalogy 1: 379 - 381." type="journal article" year="1975">Doran 1975</bibRefCitation>
). Guiler and Heddle (1974b) describe the pattern of eruption and growth of teeth.
</paragraph>
<paragraph id="171F3699FFECE71B3E7FFDC31A16FC4B" blockId="3.[840,1523,203,1954]" pageId="3" pageNumber="4">
<taxonomicName id="D0A04D1AFFECE71B3E7FFDC31A39FDEB" authorityName="Thomas" authorityYear="1912" box="[883,1097,523,546]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B3E7FFDC31A39FDEB" box="[883,1097,523,546]" italics="true" pageId="3" pageNumber="4">Sarcophilus harrisii</emphasis>
</taxonomicName>
has a narrow, flattened skull, like other dasyurids (
<bibRefCitation id="73314B68FFECE71B3ECEFDE31A12FD8B" author="WERDELIN, L." box="[962,1122,555,578]" pageId="3" pageNumber="4" pagination="109 - 117" refId="ref21796" refString="WERDELIN, L. 1986. Comparison of skull shape in marsupial and placental carnivores. Australian Journal of Zoology 34: 109 - 117." type="journal article" year="1986">Werdelin 1986</bibRefCitation>
), but is remarkable for its tiny cranium (McNab and Eisenberg 1989) and short rostrum (
<bibRefCitation id="73314B68FFECE71B389EFD831D0DFD4B" author="WERDELIN, L." pageId="3" pageNumber="4" pagination="109 - 117" refId="ref21796" refString="WERDELIN, L. 1986. Comparison of skull shape in marsupial and placental carnivores. Australian Journal of Zoology 34: 109 - 117." type="journal article" year="1986">Werdelin 1986</bibRefCitation>
). Brain size is less than one-half (43%) that predicted by body mass allometry (McNab and Eisenberg 1989). The short rostrum combined with pronounced sagittal crest and expansive zygoma accommodating powerful masseter muscles result in a powerful bite force at the canines, which suits its crushing killing bite on large prey (Jones 1997). Also unusual in
<taxonomicName id="D0A04D1AFFECE71B388CFCC31B9CFCEB" authorityName="Thomas" authorityYear="1912" box="[1408,1516,779,802]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B388CFCC31B9CFCEB" box="[1408,1516,779,802]" italics="true" pageId="3" pageNumber="4">S. harrisii</emphasis>
</taxonomicName>
, although its functional significance is not known, is the postorbital process of the frontoparietal bone, which almost separates the orbit from the zygoma.
</paragraph>
<paragraph id="171F3699FFECE71B3E7FFC431BD6FB4B" blockId="3.[840,1523,203,1954]" pageId="3" pageNumber="4">
<taxonomicName id="D0A04D1AFFECE71B3E7FFC431A3BFC6B" authorityName="Thomas" authorityYear="1912" box="[883,1099,907,930]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B3E7FFC431A3BFC6B" box="[883,1099,907,930]" italics="true" pageId="3" pageNumber="4">Sarcophilus harrisii</emphasis>
</taxonomicName>
has short metatarsals compared to the length of the femur, indicating it is not a particularly fast runner, and so is unlikely to indulge in long, fast pursuits of prey (Jones 2003a). The metacarpal-to-phalanx length ratio (associated with running speed), the ratio of olecranon process to ulna length (which affects rotational and thus tree-climbing ability), and the flattish profile of the claws (associated with tree-climbing ability) are all typical of terrestrial carnivores (Jones 2003a).
</paragraph>
<paragraph id="171F3699FFECE71B3E7FFB431B3AF98B" blockId="3.[840,1523,203,1954]" pageId="3" pageNumber="4">
Females have a backward-directed abdominal pouch, formed when birth is imminent, with 2 pairs of teats surrounded in front and laterally by a horseshoe-shaped flap of skin. In males, the darkly pigmented scrotum is located in front of the penis in the inguinal region. The scrotum can be withdrawn by the action of cremaster muscles into a pseudo-pouch consisting of ventrolateral folds of abdominal skin (
<bibRefCitation id="73314B68FFECE71B39BCFA831B30FAAB" author="GUILER, E. R." box="[1200,1344,1355,1378]" pageId="3" pageNumber="4" pagination="63 - 70" refId="ref16876" refString="GUILER, E. R. 1970 a. Observations on the Tasmanian devil, Sarcophilus harrisii (Marsupialia: Dasyuridae). II. Reproduction, breeding, and growth of pouch young. Australian Journal of Zoology 18: 63 - 70." type="journal article" year="1970">Guiler 1970a</bibRefCitation>
;
<bibRefCitation id="73314B68FFECE71B385EFA831B93FAAB" author="HUGHES, R. L." box="[1362,1507,1355,1378]" pageId="3" pageNumber="4" refId="ref17904" refString="HUGHES, R. L. 1982. Reproduction in the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Marsupialia). Pp. 49 - 63 in Carnivorous marsupials (M. Archer, ed.). Royal Zoological Society of New South Wales, Sydney, Australia." type="book" year="1982">Hughes 1982</bibRefCitation>
). On relaxation of the cremaster muscles, such as when the male is resting or hot, the pseudo-pouch becomes shallow and scrotum pendulous. The penis normally is retracted within a pocket located ventro-anteriorly in the cloaca and the glans, although slightly furrowed, is not forked or bifid (MacKenzie and Owen 1919). Apart from their larger size and primary sexual characteristics, males are indistinguishable from females.
</paragraph>
<paragraph id="171F3699FFECE71B3E7FF9831A1FF92B" blockId="3.[840,1523,203,1954]" pageId="3" pageNumber="4">
The wild
<taxonomicName id="D0A04D1AFFECE71B3ED3F9831A3DF9AB" authorityName="Thomas" authorityYear="1912" box="[991,1101,1611,1634]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFECE71B3ED3F9831A3DF9AB" box="[991,1101,1611,1634]" italics="true" pageId="3" pageNumber="4">S. harrisii</emphasis>
</taxonomicName>
often has a subcutaneous fat layer up to
<quantity id="D0589B7CFFECE71B3E69F9A31DECF94B" box="[869,924,1643,1666]" metricMagnitude="-2" metricUnit="m" metricValue="3.0" pageId="3" pageNumber="4" unit="cm" value="3.0">3 cm</quantity>
thick (
<bibRefCitation id="73314B68FFECE71B3EE6F9A31A18F94B" author="GREEN, R. H." box="[1002,1128,1643,1666]" pageId="3" pageNumber="4" pagination="1 - 12" refId="ref16344" refString="GREEN, R. H. 1967. Notes on the devil (Sarcophilus harrisii) and the quoll (Dasyurus viverrinus) in north-eastern Tasmania. Records of the Queen Victoria Museum 27: 1 - 12." type="journal article" year="1967">Green 1967</bibRefCitation>
) and one of us (RKR) has observed visceral adipose that sometimes completely covers the intestines and kidneys with tallow-like deposits. The interscapular region is another area of fat stores.
</paragraph>
<paragraph id="171F3699FFECE71C3E7FF9231C74FE4B" blockId="3.[840,1523,203,1954]" lastBlockId="4.[109,793,203,1954]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
The short digestive tract is typical of carnivores, as are the high apparent dry matter (79%) and apparent energy (87%) digestibilities (Green and Eberhard 1979). The digestible energy required for maintenance is 545 W (kg)
<superScript id="E0D59BD1FFECE71B3831F8811BD0F89E" attach="left" box="[1341,1440,1865,1890]" fontSize="6" pageId="3" pageNumber="4">0.75 day1</superScript>
or
<geoCoordinate id="7294505EFFECE71B38C8F8831D11F84B" degrees="6.30" direction="west" orientation="longitude" pageId="3" pageNumber="4" precision="555" value="-6.3">6.30 W</geoCoordinate>
(kg)
<superScript id="E0D59BD1FFECE71B3E94F8A11DCDF8BE" attach="none" box="[920,957,1897,1911]" fontSize="6" pageId="3" pageNumber="4">0.75</superScript>
, or about 2.5 times the calculated basal metabolic rate (Green and Eberhard 1979). Mean water turnover rates of
<taxonomicName id="D0A04D1AFFEBE71C3D61FF031EA7FF2B" authorityName="Thomas" authorityYear="1912" box="[109,215,203,226]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3D61FF031EA7FF2B" box="[109,215,203,226]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
, measured by tritiated water, were considerably higher than those predicted from other studies of marsupials (
<bibRefCitation id="73314B68FFEBE71C3FD1FF231ED2FEEB" author="NICOL, S. C." pageId="4" pageNumber="5" pagination="465 - 473" refId="ref20037" refString="NICOL, S. C. 1978. Rates of water turnover in marsupials and eutherians: a comparative review, with new data on the Tasmanian devil. Australian Journal of Zoology 26: 465 - 473." type="journal article" year="1978">Nicol 1978</bibRefCitation>
). Four adults averaging 5.25 ± 1.21 (
<emphasis id="25D4EA8BFFEBE71C3F47FEC31C19FEEB" box="[587,617,267,290]" italics="true" pageId="4" pageNumber="5">SE</emphasis>
) kg had a total body water of 3.48 ± 0.41 l, with turnover of 1011% per day. Thus, the mean water turnover per day was 393 ± 98 ml/day, or 74.6 ± 2.6 ml
<superScript id="E0D59BD1FFEBE71C3C0FFEA31F1DFEBE" attach="left" box="[259,365,361,386]" fontSize="6" pageId="4" pageNumber="5">kg1 day1</superScript>
(
<bibRefCitation id="73314B68FFEBE71C3C71FEA31F84FE4B" author="NICOL, S. C." box="[381,500,363,386]" pageId="4" pageNumber="5" pagination="465 - 473" refId="ref20037" refString="NICOL, S. C. 1978. Rates of water turnover in marsupials and eutherians: a comparative review, with new data on the Tasmanian devil. Australian Journal of Zoology 26: 465 - 473." type="journal article" year="1978">Nicol 1978</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFEBE71C3D94FE431C01FC0B" blockId="4.[109,793,203,1954]" pageId="4" pageNumber="5">
Mean basal metabolic rate of
<taxonomicName id="D0A04D1AFFEBE71C3CE1FE431C2DFE6B" authorityName="Thomas" authorityYear="1912" box="[493,605,395,418]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3CE1FE431C2DFE6B" box="[493,605,395,418]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
, as measured in thermoneutrality, was 0.28 ml O 2
<superScript id="E0D59BD1FFEBE71C3CDEFE631C64FE7E" attach="left" box="[466,532,425,450]" fontSize="6" pageId="4" pageNumber="5">g1 h1</superScript>
(MacMillen and
<collectingRegion id="D564F87BFFEBE71C3FC0FE631D68FE0B" box="[716,792,427,450]" country="New Zealand" name="Nelson" pageId="4" pageNumber="5">Nelson</collectingRegion>
1969). Whereas its metabolic rate falls nearly on the line (0.90) predicted by body mass, its basal metabolic rate value lies on the line for 12 dasyurids as described by the equation: basal metabolic rate = 2.45 · W (kg)
<superScript id="E0D59BD1FFEBE71C3C82FDE11FCBFDFE" attach="left" box="[398,443,553,567]" fontSize="6" pageId="4" pageNumber="5">0.261</superScript>
(MacMillen and
<collectingRegion id="D564F87BFFEBE71C3F72FDE31CBAFD8B" box="[638,714,555,578]" country="New Zealand" name="Nelson" pageId="4" pageNumber="5">Nelson</collectingRegion>
1969). These basal metabolic rate values are 32.1% below those of placental mammals of comparable sizes, and close to the mean values for marsupials (Dawson and Hulbert 1970). The metabolic rate of free-living
<taxonomicName id="D0A04D1AFFEBE71C3C3EFD631FEDFD0B" authorityName="Thomas" authorityYear="1912" box="[306,413,683,706]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3C3EFD631FEDFD0B" box="[306,413,683,706]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
, determined in the field using doubly labeled water, was 2,591 kJ/day for a 7.9-kg summer animal and 2,890 kJ/day for a 7.1-kg winter animal (
<bibRefCitation id="73314B68FFEBE71C3F44FD231CB4FCCB" author="GREEN, B." box="[584,708,747,770]" pageId="4" pageNumber="5" refId="ref16440" refString="GREEN, B. 1997. Field energetics and water fluxes in marsupials. Pp. 143 - 162 in Marsupial biology: recent research, new perspectives (N. R. Saunders and L. A. Hinds, eds.). University of New South Wales Press, Sydney, Australia." type="book" year="1997">Green 1997</bibRefCitation>
). Water fluxes for these 2 animals were 724 and 743 ml/day, respectively. Both field metabolic rate and water fluxes are independent of season for
<taxonomicName id="D0A04D1AFFEBE71C3DE6FC831F29FCAB" authorityName="Thomas" authorityYear="1912" box="[234,345,843,866]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3DE6FC831F29FCAB" box="[234,345,843,866]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
. Water fluxes of lactating females were comparable to those of 4 other dasyurids of varying sizes, and field metabolic rate was significantly raised during late-stage lactation but not in early lactation (
<bibRefCitation id="73314B68FFEBE71C3CE8FC631C10FC0B" author="GREEN, B." box="[484,608,939,962]" pageId="4" pageNumber="5" refId="ref16440" refString="GREEN, B. 1997. Field energetics and water fluxes in marsupials. Pp. 143 - 162 in Marsupial biology: recent research, new perspectives (N. R. Saunders and L. A. Hinds, eds.). University of New South Wales Press, Sydney, Australia." type="book" year="1997">Green 1997</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFEBE71C3D94FC031C35FA2B" blockId="4.[109,793,203,1954]" pageId="4" pageNumber="5">
Body temperature in free-ranging
<taxonomicName id="D0A04D1AFFEBE71C3F3FFC031CD9FC2B" authorityName="Thomas" authorityYear="1912" box="[563,681,971,994]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3F3FFC031CD9FC2B" box="[563,681,971,994]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
averages 36.5°C, with a daily range of 0.6°C that tracks the diel cycle (
<bibRefCitation id="73314B68FFEBE71C3D7AFBC31F40FBEB" author="JONES, M. E. &amp; G. C. GRIGG &amp; L. A. BEARD" box="[118,304,1035,1058]" pageId="4" pageNumber="5" pagination="53 - 60" refId="ref18417" refString="JONES, M. E., G. C. GRIGG, AND L. A. BEARD. 1997. Body temperatures and activity patterns of Tasmanian devils (Sarcophilus harrisii) and eastern quolls (Dasyurus viverrinus) through a subalpine winter. Physiological Zoology 70: 53 - 60." type="journal article" year="1997">Jones et al. 1997</bibRefCitation>
). Body temperature rises at dark as animals become active, remains high all night, and then is lowered during the day when
<taxonomicName id="D0A04D1AFFEBE71C3C25FB831FE3FBAB" authorityName="Thomas" authorityYear="1912" box="[297,403,1099,1122]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3C25FB831FE3FBAB" box="[297,403,1099,1122]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
is resting in an underground burrow.
<taxonomicName id="D0A04D1AFFEBE71C3D61FBA31EABFB4B" authorityName="Thomas" authorityYear="1912" box="[109,219,1131,1154]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3D61FBA31EABFB4B" box="[109,219,1131,1154]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
sometimes suns itself, with young animals basking with hind legs stretched backwards. Radiotracked animals have been observed to wade into water after long foraging trips, probably to cool down. Nighttime body temperature of 36.1°C and midday maximum body temperature of 37.7°C are reported in the laboratory for a single captive animal (
<bibRefCitation id="73314B68FFEBE71C3F4AFAC31C98FAEB" author="MORRISON, P. R." box="[582,744,1291,1314]" pageId="4" pageNumber="5" pagination="173 - 187" refId="ref19979" refString="MORRISON, P. R. 1965. Body temperatures in some Australian mammals. Australian Journal of Zoology 13: 173 - 187." type="journal article" year="1965">Morrison 1965</bibRefCitation>
); of the 7 dasyurids examined in captivity, only the 15-g
<taxonomicName id="D0A04D1AFFEBE71C3F9BFAE31F7DFAAB" class="Mammalia" family="Dasyuridae" genus="Sminthopsis" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="crassicaudatus">
<emphasis id="25D4EA8BFFEBE71C3F9BFAE31F7DFAAB" italics="true" pageId="4" pageNumber="5">Sminthopsis crassicaudatus</emphasis>
</taxonomicName>
had a diel amplitude lower than the 1.6°C of
<taxonomicName id="D0A04D1AFFEBE71C3D61FAA31EAAFA4B" authorityName="Thomas" authorityYear="1912" box="[109,218,1387,1410]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3D61FAA31EAAFA4B" box="[109,218,1387,1410]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
. Body temperatures of 3435.5°C were reported for 10 experimental animals (Guiler and Heddle 1970), but were 23°C higher at night, when animals are most active, than during daytime rest (Guiler and Heddle 1974a).
</paragraph>
<paragraph id="171F3699FFEBE71C3D94FA231B01FE2B" blockId="4.[109,793,203,1954]" lastBlockId="4.[840,1524,203,1954]" pageId="4" pageNumber="5">
When held for periods in constant temperatures ranging from 5°C to +35°C,
<taxonomicName id="D0A04D1AFFEBE71C3C54F9C31FB3F9EB" authorityName="Thomas" authorityYear="1912" box="[344,451,1547,1570]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3C54F9C31FB3F9EB" box="[344,451,1547,1570]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
is a good thermoregulator, usually at 35 ± 1°C. At ambient temperatures outside the thermoneutral zone of 28.532°C (Nicol and Maskrey 1980),
<taxonomicName id="D0A04D1AFFEBE71C3FA2F9831D68F9AB" authorityName="Thomas" authorityYear="1912" box="[686,792,1611,1634]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3FA2F9831D68F9AB" box="[686,792,1611,1634]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
produces heat by shivering or loses heat by panting, the usual mammalian responses. When held in the cold well below its thermoneutral zone,
<taxonomicName id="D0A04D1AFFEBE71C3C41F9631FC9F90B" authorityName="Thomas" authorityYear="1912" box="[333,441,1707,1730]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3C41F9631FC9F90B" box="[333,441,1707,1730]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
is homeothermic, maintaining a body temperature of 36.0°C rather than the 36.8°C in thermoneutrality (MacMillen and
<collectingRegion id="D564F87BFFEBE71C3C9CF9231FACF8CB" box="[400,476,1771,1794]" country="New Zealand" name="Nelson" pageId="4" pageNumber="5">Nelson</collectingRegion>
1969). Despite snowy conditions through an alpine winter, 3 free-ranging male and female
<taxonomicName id="D0A04D1AFFEBE71C3D61F8E31EA9F88B" authorityName="Thomas" authorityYear="1912" box="[109,217,1835,1858]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3D61F8E31EA9F88B" box="[109,217,1835,1858]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
did not enter daily torpor (
<bibRefCitation id="73314B68FFEBE71C3F0CF8E31CC5F88B" author="JONES, M. E. &amp; G. C. GRIGG &amp; L. A. BEARD" box="[512,693,1835,1858]" pageId="4" pageNumber="5" pagination="53 - 60" refId="ref18417" refString="JONES, M. E., G. C. GRIGG, AND L. A. BEARD. 1997. Body temperatures and activity patterns of Tasmanian devils (Sarcophilus harrisii) and eastern quolls (Dasyurus viverrinus) through a subalpine winter. Physiological Zoology 70: 53 - 60." type="journal article" year="1997">Jones et al. 1997</bibRefCitation>
). After 2 weeks of acclimation to 23°C temperatures,
<taxonomicName id="D0A04D1AFFEBE71C3F53F8831CBCF8AB" authorityName="Thomas" authorityYear="1912" box="[607,716,1867,1890]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3F53F8831CBCF8AB" box="[607,716,1867,1890]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
exhibits a significant increase in metabolic rate after injection of norepinephrine compared to controls injected with isotonic saline (
<bibRefCitation id="73314B68FFEBE71C3E5DFF031A77FF2B" author="KABAT, A. P. &amp; R. W. ROSE &amp; AND A. K." box="[849,1031,203,226]" pageId="4" pageNumber="5" pagination="413 - 420" refId="ref18879" refString="KABAT, A. P., R. W. ROSE, AND A. K. W EST. 2003. Non-shivering thermogenesis in a carnivorous marsupial, Sarcophilus harrisii, in the absence of UCP 1. Journal of Thermal Biology 28: 413 - 420." type="journal article" year="2003">Kabat et al. 2003</bibRefCitation>
). These authors interpret this response as evidence that
<taxonomicName id="D0A04D1AFFEBE71C3ECFFF231A41FECB" authorityName="Thomas" authorityYear="1912" box="[963,1073,235,258]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3ECFFF231A41FECB" box="[963,1073,235,258]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
produces heat by non-shivering thermogenesis, despite a presumed lack of brown adipose tissue and the absence of uncoupling protein-1, both prerequisites for nonshivering thermogenesis in eutherian mammals. A further study (
<bibRefCitation id="73314B68FFEBE71C3E5DFEA31A64FE4B" author="KABAT, A. P. &amp; R. W. ROSE &amp; AND A. K." box="[849,1044,363,386]" pageId="4" pageNumber="5" pagination="109 - 115" refId="ref18929" refString="KABAT, A. P., R. W. ROSE, AND A. K. W EST. 2004. Molecular identification of uncoupling proteins 2 and 3 in a carnivorous marsupial, the Tasmanian devil (Sarcophilus harrisii). Physiological and Biochemical Zoology 77: 109 - 115." type="journal article" year="2004">Kabat et al. 2004</bibRefCitation>
), which examined the presence of 2 other members of the mitochondrial anion-carrier protein superfamily, revealed the presence of uncoupling protein-2 but not uncoupling protein-
<quantity id="D0589B7CFFEBE71C3ED0FE031A73FE2B" box="[988,1027,459,482]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="4" pageNumber="5" unit="in" value="3.0">3 in</quantity>
muscle and white adipose tissues.
</paragraph>
<paragraph id="171F3699FFEBE71C3E7FFE231DD3FBAB" blockId="4.[840,1524,203,1954]" pageId="4" pageNumber="5">
Hulbert and Rose (1972) concluded that
<taxonomicName id="D0A04D1AFFEBE71C382BFE231BE2FDCB" authorityName="Thomas" authorityYear="1912" box="[1319,1426,491,514]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C382BFE231BE2FDCB" box="[1319,1426,491,514]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
does not sweat (confirmed by
<bibRefCitation id="73314B68FFEBE71C393CFDC31AAAFDEB" author="BELL, C. J. &amp; R. V. BAUDINETTE &amp; S. C. NICOL" box="[1072,1242,523,546]" pageId="4" pageNumber="5" pagination="93 - 99" refId="ref14836" refString="BELL, C. J., R. V. BAUDINETTE, AND S. C. NICOL. 1983. Cutaneous water loss at rest and exercise in two species of marsupials. Australian Journal of Zoology 31: 93 - 99." type="journal article" year="1983">Bell et al. 1983</bibRefCitation>
), but loses heat via panting, when it raises its breathing rate 7-fold. At an ambient temperature of 40°C, body temperature is kept at 38.5°C but under exertion (running, up to
<quantity id="D0589B7CFFEBE71C3945FDA31AE3FD4B" box="[1097,1171,619,642]" metricMagnitude="3" metricUnit="m" metricValue="5.6" pageId="4" pageNumber="5" unit="km" value="5.6">5.6 km</quantity>
/h) or application of external heat load (Hulbert and Rose 1972), body temperature may rise to 39°C or 40°C. In contrast to eutherians in which less than 10% of heat is stored and most heat is lost by convection, conduction, or radiation,
<taxonomicName id="D0A04D1AFFEBE71C3EDEFD231A4CFCCB" authorityName="Thomas" authorityYear="1912" box="[978,1084,747,770]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3EDEFD231A4CFCCB" box="[978,1084,747,770]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
stores about 40% of produced heat (
<bibRefCitation id="73314B68FFEBE71C38CAFD231DC6FCEB" author="BELL, C. J. &amp; R. V. BAUDINETTE &amp; S. C. NICOL" pageId="4" pageNumber="5" pagination="93 - 99" refId="ref14836" refString="BELL, C. J., R. V. BAUDINETTE, AND S. C. NICOL. 1983. Cutaneous water loss at rest and exercise in two species of marsupials. Australian Journal of Zoology 31: 93 - 99." type="journal article" year="1983">Bell et al. 1983</bibRefCitation>
) and dissipates about one-half the heat by evaporative means, the latter almost entirely through breathing rather than the skin. These features are consistent with the relatively low body temperature and high rates of water turnover in
<taxonomicName id="D0A04D1AFFEBE71C388BFCA31A7CFC6B" authority="(Baudinette 1982)" baseAuthorityName="Baudinette" baseAuthorityYear="1982" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C388BFCA31B83FC4B" box="[1415,1523,875,898]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
(
<bibRefCitation id="73314B68FFEBE71C3E5DFC431A71FC6B" author="BAUDINETTE, R. V." box="[849,1025,907,930]" pageId="4" pageNumber="5" refId="ref14787" refString="BAUDINETTE, R. V. 1982. The energetics of locomotion in dasyurid marsupials. Pp. 261 - 265 in Carnivorous marsupials (M. Archer, ed.). Surrey Beatty &amp; Sons Pty Limited, Chipping Norton, New South Wales, Australia." type="book" year="1982">Baudinette 1982</bibRefCitation>
)
</taxonomicName>
. Although panting is the usual mechanism to dump heat in marsupials,
<taxonomicName id="D0A04D1AFFEBE71C3953FC631ABAFC0B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[1119,1226,939,962]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3953FC631ABAFC0B" box="[1119,1226,939,962]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
may possess a carotid rete, a network to send cooler blood to the brain (
<bibRefCitation id="73314B68FFEBE71C383EFC031B93FC2B" author="SHAH, S. K. H. &amp; S. C. NICOL &amp; R. SWAIN" box="[1330,1507,971,994]" pageId="4" pageNumber="5" pagination="125 - 133" refId="ref21139" refString="SHAH, S. K. H., S. C. NICOL, AND R. SWAIN. 1986. Functional morphology of the cranial vasculature and the nasal passage in the Tasmanian devil, Sarcophilus harrisii (Marsupialia: Dasyuridae): a marsupial carotid rete? Australian Journal of Zoology 34: 125 - 133." type="journal article" year="1986">Shah et al. 1986</bibRefCitation>
). Taken together, these studies indicate that
<taxonomicName id="D0A04D1AFFEBE71C3805FC231B03FBCB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[1289,1395,1003,1026]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3805FC231B03FBCB" box="[1289,1395,1003,1026]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
has a metabolic rate typical of marsupials, is a good thermoregulator, and often has a body temperature 23°C below the eutherian mean of 37°C.
</paragraph>
<paragraph id="171F3699FFEBE71C3E7FFBA31A4BFA8B" blockId="4.[840,1524,203,1954]" pageId="4" pageNumber="5">
Heart rate is somewhat variable, but averaged 102 ± 17 (
<emphasis id="25D4EA8BFFEBE71C38C0FBA31B9AFB4B" box="[1484,1514,1131,1154]" italics="true" pageId="4" pageNumber="5">SE</emphasis>
) beats per min for 20 animals at thermoneutrality, whereas blood pressure readings of unanesthetized animals at thermoneutrality averaged 131 ±
<quantity id="D0589B7CFFEBE71C3928FB031A02FB2B" box="[1060,1138,1227,1250]" metricMagnitude="-2" metricUnit="m" metricValue="2.6" pageId="4" pageNumber="5" unit="mm" value="26.0">26 mm</quantity>
Hg/98 ±
<quantity id="D0589B7CFFEBE71C39E8FB031B42FB2B" box="[1252,1330,1227,1250]" metricMagnitude="-2" metricUnit="m" metricValue="2.3" pageId="4" pageNumber="5" unit="mm" value="23.0">23 mm</quantity>
Hg (
<bibRefCitation id="73314B68FFEBE71C387DFB031B9CFB2B" author="NICOL, S. C." box="[1393,1516,1227,1250]" pageId="4" pageNumber="5" refId="ref20076" refString="NICOL, S. C. 1982. Physiology of the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Marsupialia). Pp. 273 - 278 in Carnivorous marsupials (M. Archer, ed.). Surrey Beatty &amp; Sons Pty Limited, Chipping Norton, New South Wales, Australia." type="book" year="1982">Nicol 1982</bibRefCitation>
, who also summarizes hematological data).
<bibRefCitation id="73314B68FFEBE71C381BFB231B83FACB" author="PARSONS, R. S. &amp; EDDLE, W. G. &amp; LUX, AND E. R." box="[1303,1523,1259,1282]" pageId="4" pageNumber="5" pagination="345 - 351" refId="ref20428" refString="PARSONS, R. S., R. W. L. H EDDLE, W. G. F LUX, AND E. R. G UILER. 1970. Studies of the blood of the Tasmanian devil. Comparative Biochemistry and Physiology 32: 345 - 351." type="journal article" year="1970">Parsons et al. (1970)</bibRefCitation>
recorded a mean aortic blood pressure (under anesthesia) of
<quantity id="D0589B7CFFEBE71C3E44FAE31DE2FA8B" box="[840,914,1323,1346]" metricMagnitude="-2" metricUnit="m" metricValue="9.4" pageId="4" pageNumber="5" unit="mm" value="94.0">94 mm</quantity>
Hg/
<quantity id="D0589B7CFFEBE71C3ECEFAE31A7CFA8B" box="[962,1036,1323,1346]" metricMagnitude="-2" metricUnit="m" metricValue="4.2" pageId="4" pageNumber="5" unit="mm" value="42.0">42 mm</quantity>
Hg.
</paragraph>
<paragraph id="171F3699FFEBE71D3E7FFA831F32FE2B" blockId="4.[840,1524,203,1954]" lastBlockId="5.[109,793,203,1730]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
Numerous studies have been conducted on the blood, starting with
<bibRefCitation id="73314B68FFEBE71C3EBCFAA31AE9FA4B" author="PARSONS, R. S. &amp; EDDLE, W. G. &amp; LUX, AND E. R." box="[944,1177,1387,1410]" pageId="4" pageNumber="5" pagination="345 - 351" refId="ref20428" refString="PARSONS, R. S., R. W. L. H EDDLE, W. G. F LUX, AND E. R. G UILER. 1970. Studies of the blood of the Tasmanian devil. Comparative Biochemistry and Physiology 32: 345 - 351." type="journal article" year="1970">Parsons et al. (1970)</bibRefCitation>
, who found, among other features, high levels of acid phosphatase. Both serum and tissue levels (16 μg) are exceedingly high (
<bibRefCitation id="73314B68FFEBE71C39D1FA631BC9FA0B" author="PARSONS, R. S. &amp; R. W. L. HEDDLE &amp; E. R. GUILER" box="[1245,1465,1451,1474]" pageId="4" pageNumber="5" pagination="219 - 226" refId="ref20480" refString="PARSONS, R. S., R. W. L. HEDDLE, AND E. R. GUILER. 1971 a. The distribution of acid phosphatase in the Tasmanian devil, Sarcophilus harrisii (Marsupialia: Dasyuridae). Comparative Biochemistry and Physiology, B. Comparative Biochemistry 39: 219 - 226." type="journal article" year="1971">Parsons et al. 1971a</bibRefCitation>
); the enzyme has a molecular weight of 85,000 (
<bibRefCitation id="73314B68FFEBE71C3825FA031B93FA2B" author="SALLIS, J. D. &amp; R. S. PARSONS &amp; E. R. GUILER" box="[1321,1507,1483,1506]" pageId="4" pageNumber="5" pagination="971 - 980" refId="ref21031" refString="SALLIS, J. D., R. S. PARSONS, AND E. R. GUILER. 1973. Isolation and some properties of an acid phosphatase from the plasma of the Tasmanian devil, Sarcophilus harrisii (Marsupialia: Dasyuridae). Comparative Biochemistry and Physiology, B. Comparative Biochemistry 44: 971 - 980." type="journal article" year="1973">Sallis et al. 1973</bibRefCitation>
).
<taxonomicName id="D0A04D1AFFEBE71C3E44FA231DC8F9CB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[840,952,1515,1538]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3E44FA231DC8F9CB" box="[840,952,1515,1538]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
and the next larger dasyurid,
<taxonomicName id="D0A04D1AFFEBE71C381FFA231B83F9CB" baseAuthorityName="Kerr" baseAuthorityYear="1792" box="[1299,1523,1515,1538]" class="Mammalia" family="Dasyuridae" genus="Dasyurus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="maculatus">
<emphasis id="25D4EA8BFFEBE71C381FFA231B83F9CB" box="[1299,1523,1515,1538]" italics="true" pageId="4" pageNumber="5">Dasyurus maculatus</emphasis>
</taxonomicName>
(spotted-tailed quoll), differ from other mammals in their high levels of acid phosphatase (Sallis and Guiler 1977). The values of 32.50 and
<quantity id="D0589B7CFFEBE71C3ED4F9831A5AF9AA" box="[984,1066,1611,1635]" metricMagnitude="1" metricUnit="m" metricValue="2.014" pageId="4" pageNumber="5" unit="m" value="20.14">20.14 µ</quantity>
mol-p-nitrophenol formed per ml per min in plasma, respectively, are the highest recorded for any mammalian species, and are 12 orders of magnitude greater than those of other marsupials and of humans.
<bibRefCitation id="73314B68FFEBE71C3808F9631B9CF90B" author="PARSONS, R. S. &amp; E. R. GUILER &amp; R. W. L. HEDDLE" box="[1284,1516,1707,1730]" pageId="4" pageNumber="5" pagination="209 - 217" refId="ref20367" refString="PARSONS, R. S., E. R. GUILER, AND R. W. L. HEDDLE. 1971 b. Comparative studies on the blood of monotremes and marsupials-II. Electrolyte organic constituents, proteins, gas analysis and enzymes. Comparative Biochemistry and Physiology, B. Comparative Biochemistry 39: 209 - 217." type="journal article" year="1971">Parsons et al. (1971b)</bibRefCitation>
, who determined levels of serum and red blood cell electrolytes as well as those of several organic constituents and serum proteins, recorded pH and levels of gases in blood for
<taxonomicName id="D0A04D1AFFEBE71C3889F8C31B83F8EB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[1413,1523,1803,1826]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEBE71C3889F8C31B83F8EB" box="[1413,1523,1803,1826]" italics="true" pageId="4" pageNumber="5">S. harrisii</emphasis>
</taxonomicName>
below those of 5 marsupials and of humans for CO 2, HCO 3, and both percentage of O 2 and O 2 as a percentage of saturation.
<bibRefCitation id="73314B68FFEBE71C3E44F8A31A6AF84B" author="BENGA, G. &amp; B. E. CHAPMAN &amp; C. H. GALLAGHER &amp; N. S. AGAR &amp; AND P. W." box="[840,1050,1899,1922]" pageId="4" pageNumber="5" pagination="515 - 518" refId="ref14881" refString="BENGA, G., B. E. CHAPMAN, C. H. GALLAGHER, N. S. AGAR, AND P. W. K UCHEL. 1993. NMR studies of diffusional water permeability of erythrocytes from eight species of marsupials. Comparative Biochemistry and Physiology, A. Comparative Physiology 106: 515 - 518." type="journal article" year="1993">Benga et al. (1993)</bibRefCitation>
measured membrane permeability to water and other features of red blood cells. More detailed accounts of blood components (
<bibRefCitation id="73314B68FFEAE71D3C4CFF031FC6FF2B" author="CLARK, P." box="[320,438,203,226]" pageId="5" pageNumber="6" refId="ref15439" refString="CLARK, P. 2004. Haematology of Australian mammals. CSIRO Publishing, Collingwood, Victoria, Australia." type="book" year="2004">Clark 2004</bibRefCitation>
) describe erythrocytes as eosinophilic discocytes with slight central pallor. Neutrophils have 3- to 7-lobed nuclei of clumped chromatin and colorless cytoplasm. Small- to medium-sized lymphocytes have round nuclei packed with dense chromatin and surrounded by little basophilic cytoplasm. The nucleus of monocytes is irregularly shaped and composed of lacy chromatin with a granular gray cytoplasm and each eosinophil has a 2- to 5-lobed nucleus and a pale basophilic granular cytoplasm.
</paragraph>
<paragraph id="171F3699FFEAE71D3D94FE231CAFFACB" blockId="5.[109,793,203,1730]" pageId="5" pageNumber="6">
The adrenal gland of
<taxonomicName id="D0A04D1AFFEAE71D3C84FE231F86FDCB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[392,502,491,514]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEAE71D3C84FE231F86FDCB" box="[392,502,491,514]" italics="true" pageId="5" pageNumber="6">S. harrisii</emphasis>
</taxonomicName>
is similar in size for both sexes,
<quantity id="D0589B7CFFEAE71D3DB9FDC31F78FDEB" box="[181,264,523,546]" metricMagnitude="-4" metricUnit="kg" metricValue="1.71" pageId="5" pageNumber="6" unit="mg" value="171.0">171 mg</quantity>
/kg in males and
<quantity id="D0589B7CFFEAE71D3CCAFDC31C68FDEB" box="[454,536,523,546]" metricMagnitude="-4" metricUnit="kg" metricValue="1.63" pageId="5" pageNumber="6" unit="mg" value="163.0">163 mg</quantity>
/kg in females. Cortisol is the major corticosteroid in the peripheral blood plasma (
<bibRefCitation id="73314B68FFEAE71D3D7AFD831F5BFDAB" author="MCDONALD, I. R." box="[118,299,587,610]" pageId="5" pageNumber="6" refId="ref19727" refString="MCDONALD, I. R. 1977. Adrenocortical functions in marsupials. Pp. 345 - 377 in The biology of marsupials (B. Stonehouse and D. Gilmore, eds.). The Macmillan Press Ltd., London, United Kingdom." type="book" year="1977">
<collectingCountry id="6FB77609FFEAE71D3D7AFD831E98FDAB" box="[118,232,587,610]" name="Heard Island and McDonald Islands" pageId="5" pageNumber="6">McDonald</collectingCountry>
1977
</bibRefCitation>
), which also contains a steroid tentatively identified as corticosterone (Weiss and Richards 1971). Plasmafree corticosteroid levels in free-ranging
<taxonomicName id="D0A04D1AFFEAE71D3F33FD431CC0FD6B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[575,688,651,674]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEAE71D3F33FD431CC0FD6B" box="[575,688,651,674]" italics="true" pageId="5" pageNumber="6">S. harrisii</emphasis>
</taxonomicName>
, 5.6 and 11.9 nM for adult males and females, were similar to values, 10.0 and 12.4 nM, for juvenile males and females, respectively (
<bibRefCitation id="73314B68FFEAE71D3D7AFD231F56FCCB" author="PEMBERTON, D." box="[118,294,747,770]" pageId="5" pageNumber="6" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
). Females have high corticosteroid levels during the breeding season and when young are 1st deposited in nests, whereas males have constant levels, including those of testosterone, throughout the year (
<bibRefCitation id="73314B68FFEAE71D3CC8FC831C05FCAB" author="PEMBERTON, D." box="[452,629,843,866]" pageId="5" pageNumber="6" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
). Males, when restrained for 510 min and lightly anesthetized, have peripheral plasma cortisol concentrations of
<quantity id="D0589B7CFFEAE71D3CE8FC431C29FC6B" box="[484,601,907,931]" metricMagnitude="-6" metricUnit="kg" metricValue="3.25" metricValueMax="3.7" metricValueMin="2.8" pageId="5" pageNumber="6" unit="mg" value="3.25" valueMax="3.7" valueMin="2.8">2.83.7 µg</quantity>
/100 ml, whereas females in the same conditions have a mean value of 7.5 (Weiss and Richards 1971). In males taken into captivity, plasma cortisol concentrations fall to basal levels after only 48 h in captivity (
<bibRefCitation id="73314B68FFEAE71D3DBBFBC31F26FBEB" author="LOCKHART, T. J." box="[183,342,1035,1058]" pageId="5" pageNumber="6" refId="ref19379" refString="LOCKHART, T. J. 2000. Effects of captivity on the reproduction of the Tasmanian devil, Sarcophilus harrisii. B. Sc. Honours thesis, University of Tasmania, Hobart, Australia." type="book" year="2000">Lockhart 2000</bibRefCitation>
). However, females have higher stressed secretion rates (
<quantity id="D0589B7CFFEAE71D3C2CFBE31F01FB8B" box="[288,369,1067,1091]" metricMagnitude="-4" metricUnit="kg" metricValue="1.9" pageId="5" pageNumber="6" unit="mg" value="190.0">190 µg</quantity>
<superScript id="E0D59BD1FFEAE71D3C71FBE31FA3FBFE" attach="left" box="[381,467,1065,1090]" fontSize="6" pageId="5" pageNumber="6">kg1 h1</superScript>
) and higher stressed plasma concentrations (
<quantity id="D0589B7CFFEAE71D3C15FB831F39FBAB" box="[281,329,1099,1123]" metricMagnitude="-6" metricUnit="kg" metricValue="8.0" pageId="5" pageNumber="6" unit="mg" value="8.0">8 µg</quantity>
/100 ml) than males (111 and
<quantity id="D0589B7CFFEAE71D3F81FB831CCFFBAB" box="[653,703,1099,1123]" metricMagnitude="-6" metricUnit="kg" metricValue="3.0" pageId="5" pageNumber="6" unit="mg" value="3.0">3 µg</quantity>
, respec-tively—McDonald 1977). Neither sex shows seasonal patterns of corticosteroid levels nor are seasonal peaks of testosterone detected during the breeding season, as is typical of some smaller semelparous dasyurids. Testosterone levels in free-ranging adult and juvenile males were 0.66 and
<quantity id="D0589B7CFFEAE71D3CD5FB231C59FACB" box="[473,553,1259,1282]" metricMagnitude="-13" metricUnit="kg" metricValue="7.8" pageId="5" pageNumber="6" unit="ng" value="0.78">0.78 ng</quantity>
/ml, respectively.
</paragraph>
<paragraph id="171F3699FFEAE71D3D94FAC31F7DFA6B" blockId="5.[109,793,203,1730]" pageId="5" pageNumber="6">
Both sexes show a neutrophil-to-lymphocyte shift over the mating season indicative of reproductive stress (
<bibRefCitation id="73314B68FFEAE71D3F8CFAE31D78FA8B" author="BURTON, B." box="[640,776,1323,1346]" pageId="5" pageNumber="6" refId="ref15237" refString="BURTON, B. 2002. The effects of the breeding season on immune responses in the Tasmanian devil, Sarcophilus laniarius. B. Sc. Honours thesis, University of Tasmania, Hobart, Australia." type="book" year="2002">Burton 2002</bibRefCitation>
). An increase in the total number of neutrophils in males may relate to the extensive injuries they sometimes sustain during the mating season.
</paragraph>
<paragraph id="171F3699FFEAE71D3D94FA631F9DF90B" blockId="5.[109,793,203,1730]" pageId="5" pageNumber="6">
Like other dasyurids,
<taxonomicName id="D0A04D1AFFEAE71D3C88FA631F9FFA0B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[388,495,1451,1474]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEAE71D3C88FA631F9FFA0B" box="[388,495,1451,1474]" italics="true" pageId="5" pageNumber="6">S. harrisii</emphasis>
</taxonomicName>
is more tolerant to the poison Compound 1080, used to control some pest species, than are eutherian carnivores, likely because similar defensive molecules are found in a native plant,
<taxonomicName id="D0A04D1AFFEAE71D3CA0F9C31C4FF9EB" box="[428,575,1547,1570]" class="Magnoliopsida" family="Fabaceae" genus="Gastrolobium" kingdom="Plantae" order="Fabales" pageId="5" pageNumber="6" phylum="Tracheophyta" rank="genus">
<emphasis id="25D4EA8BFFEAE71D3CA0F9C31C4FF9EB" box="[428,575,1547,1570]" italics="true" pageId="5" pageNumber="6">Gastrolobium</emphasis>
</taxonomicName>
. (Compound 1080, sodium monofluoroacetate, blocks enzymes that catalyze citrate and succinate metabolism, and thus inhibits the Krebs cycle, causing cardiac arrest.) On a mg per kg of body mass basis,
<taxonomicName id="D0A04D1AFFEAE71D3D61F9431EA8F96B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[109,216,1675,1698]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEAE71D3D61F9431EA8F96B" box="[109,216,1675,1698]" italics="true" pageId="5" pageNumber="6">S. harrisii</emphasis>
</taxonomicName>
is 38 times more tolerant of 1080 than is the dingo, a eutherian carnivore (
<bibRefCitation id="73314B68FFEAE71D3C46F9631FADF90B" author="MCILROY, J. C." box="[330,477,1707,1730]" pageId="5" pageNumber="6" refId="ref19774" refString="MCILROY, J. C. 1982. The sensitivity of Australian carnivorous mammals to 1080 poison. Pp. 267 - 271 in Carnivorous marsupials (M. Archer, ed.). Surrey Beatty &amp; Sons Pty Limited, Chipping Norton, New South Wales, Australia." type="book" year="1982">McIlroy 1982</bibRefCitation>
).
</paragraph>
</subSection>
<subSection id="7E2F2F72FFEAE71E3DC3F8CE1B06F9CB" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6" type="multiple">
<paragraph id="171F3699FFEAE71D3DC3F8CE1CC6F8E9" blockId="5.[207,694,1798,1824]" box="[207,694,1798,1824]" pageId="5" pageNumber="6">
<heading id="4C5781F5FFEAE71D3DC3F8CE1CC6F8E9" allCaps="true" bold="true" box="[207,694,1798,1824]" centered="true" fontSize="11" level="1" pageId="5" pageNumber="6" reason="6">
<emphasis id="25D4EA8BFFEAE71D3DC3F8CE1CC6F8E9" bold="true" box="[207,694,1798,1824]" pageId="5" pageNumber="6">ONTOGENY AND REPRODUCTION</emphasis>
</heading>
</paragraph>
<paragraph id="171F3699FFEAE71D3D94F8821BE3FD8B" blockId="5.[109,792,1866,1953]" lastBlockId="5.[840,1523,203,1954]" pageId="5" pageNumber="6">
<emphasis id="25D4EA8BFFEAE71D3D94F8821F7AF8AB" bold="true" box="[152,266,1866,1890]" italics="true" pageId="5" pageNumber="6">Ontogeny.</emphasis>
—Neonates, like other marsupial young, must clamber from the birth canal into the pouch and find an unoccupied teat or die. Because neonates are numerous and the number of teats only 4, natural selection (and chance) determine which survive. Neonates weigh about
<quantity id="D0589B7CFFEAE71D382DFF231B27FECB" box="[1313,1367,235,258]" metricMagnitude="-6" metricUnit="kg" metricValue="5.0" pageId="5" pageNumber="6" unit="mg" value="5.0">5 mg</quantity>
with a crownrump length of
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, spend the first 90105 days attached to teats, and a nearly equal time in the nest. Although weaned at 150240 days (
<bibRefCitation id="73314B68FFEAE71D3EFEFE831AF3FEAB" author="GUILER, E. R." box="[1010,1155,331,354]" pageId="5" pageNumber="6" pagination="63 - 70" refId="ref16876" refString="GUILER, E. R. 1970 a. Observations on the Tasmanian devil, Sarcophilus harrisii (Marsupialia: Dasyuridae). II. Reproduction, breeding, and growth of pouch young. Australian Journal of Zoology 18: 63 - 70." type="journal article" year="1970">Guiler 1970a</bibRefCitation>
reports 7 months, in November), or about 34 months later than other large dasyurids, maturity is not reached until nearly 2 years in most females. However, a fraction of females does breed as yearlings (immaturity confirmed by unfused tibial epiphyses) if they have achieved weights of
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(
<bibRefCitation id="73314B68FFEAE71D393FFE231AB4FDCB" author="HUGHES, R. L." box="[1075,1220,491,514]" pageId="5" pageNumber="6" refId="ref17904" refString="HUGHES, R. L. 1982. Reproduction in the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Marsupialia). Pp. 49 - 63 in Carnivorous marsupials (M. Archer, ed.). Royal Zoological Society of New South Wales, Sydney, Australia." type="book" year="1982">Hughes 1982</bibRefCitation>
). After the loss of adults in a diseased population, 1-year-old females with a mass of
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entered the breeding population (
<bibRefCitation id="73314B68FFEAE71D39BFFDE31BF2FD8B" author="LACHISH, S. &amp; H. MCCALLUM &amp; M. JONES" box="[1203,1410,555,578]" pageId="5" pageNumber="6" pagination="427 - 436" refId="ref19198" refString="LACHISH, S., H. MCCALLUM, AND M. JONES. 2009. Demography, disease, and the devil: life-history changes in a disease-affected population of the Tasmanian devil (Sarcophilus harrisii). Journal of Animal Ecology 78: 427 - 436." type="journal article" year="2009">Lachish et al. 2009</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFEAE71D3E7FFD831B12FCAB" blockId="5.[840,1523,203,1954]" pageId="5" pageNumber="6">
During early weeks of life, females are heavier than males, but by 18 months this pattern is reversed, and thereafter males can increase rapidly in mass (
<bibRefCitation id="73314B68FFEAE71D39AEFD431B58FD6B" author="GUILER, E. R." box="[1186,1320,651,674]" pageId="5" pageNumber="6" pagination="161 - 178" refId="ref16996" refString="GUILER, E. R. 1978. Observations on the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Mammalia) at Granville Harbour. Papers and Proceedings of the Royal Society of Tasmania 112: 161 - 178." type="journal article" year="1978">Guiler 1978</bibRefCitation>
). Although males can be sexually mature at
<quantity id="D0589B7CFFEAE71D3968FD631AD9FD0B" box="[1124,1193,683,706]" metricMagnitude="0" metricUnit="kg" metricValue="3.5" pageId="5" pageNumber="6" unit="kg" value="3.5">3.5 kg</quantity>
and most complete growth by 24 months of age (
<bibRefCitation id="73314B68FFEAE71D3913FD031AA2FD2B" author="PEMBERTON, D." box="[1055,1234,715,738]" pageId="5" pageNumber="6" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
), some males continue to grow until they are 34 years old, by which age they may reach their final adult masses of
<quantity id="D0589B7CFFEAE71D396FFCC31ACEFCEB" box="[1123,1214,779,802]" metricMagnitude="1" metricUnit="kg" metricValue="1.1" metricValueMax="1.4" metricValueMin="0.8" pageId="5" pageNumber="6" unit="kg" value="11.0" valueMax="14.0" valueMin="8.0">814 kg</quantity>
. Mass increased in 6 months from
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to
<quantity id="D0589B7CFFEAE71D3937FCE31AC0FC8B" box="[1083,1200,811,834]" metricMagnitude="0" metricUnit="kg" metricValue="6.45" metricValueMax="9.4" metricValueMin="3.5" pageId="5" pageNumber="6" unit="kg" value="6.45" valueMax="9.4" valueMin="3.5">3.59.4 kg</quantity>
for
<specimenCount id="01A6FD10FFEAE71D39EDFCE31B49FC8B" box="[1249,1337,811,834]" pageId="5" pageNumber="6" type="male">2 males</specimenCount>
, an indication of growth potential in ideal conditions (
<bibRefCitation id="73314B68FFEAE71D39DEFC831B22FCAB" author="GUILER, E. R." box="[1234,1362,843,866]" pageId="5" pageNumber="6" pagination="161 - 178" refId="ref16996" refString="GUILER, E. R. 1978. Observations on the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Mammalia) at Granville Harbour. Papers and Proceedings of the Royal Society of Tasmania 112: 161 - 178." type="journal article" year="1978">Guiler 1978</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFEAE71D3E7FFCA31DCDFACB" blockId="5.[840,1523,203,1954]" pageId="5" pageNumber="6">
<taxonomicName id="D0A04D1AFFEAE71D3E7FFCA31A38FC4B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[883,1096,875,898]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEAE71D3E7FFCA31A38FC4B" box="[883,1096,875,898]" italics="true" pageId="5" pageNumber="6">Sarcophilus harrisii</emphasis>
</taxonomicName>
can live up to 8 years in the wild but in some populations few live past 3 years, an indication of variation in life span of recruits (
<bibRefCitation id="73314B68FFEAE71D394BFC631AB8FC0B" author="GUILER, E. R." box="[1095,1224,939,962]" pageId="5" pageNumber="6" pagination="161 - 178" refId="ref16996" refString="GUILER, E. R. 1978. Observations on the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Mammalia) at Granville Harbour. Papers and Proceedings of the Royal Society of Tasmania 112: 161 - 178." type="journal article" year="1978">Guiler 1978</bibRefCitation>
). Juveniles have up to 80% mortality during their 1st year of independent life, with males outliving females, on average. Before weaned young enter the population, 61.591.0% (
<emphasis id="25D4EA8BFFEAE71D3955FBC01A1DFBD4" box="[1113,1133,1032,1053]" italics="true" pageId="5" pageNumber="6">X</emphasis>
= 76.2%) of the population is adult, a high value for carnivores (
<bibRefCitation id="73314B68FFEAE71D3971FBE31A8EFB8B" author="GUILER, E. R." box="[1149,1278,1067,1090]" pageId="5" pageNumber="6" pagination="161 - 178" refId="ref16996" refString="GUILER, E. R. 1978. Observations on the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Mammalia) at Granville Harbour. Papers and Proceedings of the Royal Society of Tasmania 112: 161 - 178." type="journal article" year="1978">Guiler 1978</bibRefCitation>
). Symptoms of physical deterioration due to age 1st appeared in zoo-held
<taxonomicName id="D0A04D1AFFEAE71D388BFB831B83FBAB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[1415,1523,1099,1122]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEAE71D388BFB831B83FBAB" box="[1415,1523,1099,1122]" italics="true" pageId="5" pageNumber="6">S. harrisii</emphasis>
</taxonomicName>
after 50 months of age (Holz and Little 1995). Gradual loss of balance leading to total hindquarter paralysis, interpreted as agerelated degeneration, is common; necropsies of aged animals revealed degenerative neurological disease consistent with these symptoms.
</paragraph>
<paragraph id="171F3699FFEAE71D3E7FFAC31B7CFA6B" blockId="5.[840,1523,203,1954]" pageId="5" pageNumber="6">
Although young are prey of wedge-tailed eagles (
<taxonomicName id="D0A04D1AFFEAE71D38A0FAC31DF9FA8B" baseAuthorityName="Latham" baseAuthorityYear="1802" class="Aves" family="Accipitridae" genus="Aquila" kingdom="Animalia" order="Accipitriformes" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="audax">
<emphasis id="25D4EA8BFFEAE71D38A0FAC31DF9FA8B" italics="true" pageId="5" pageNumber="6">Aquila audax</emphasis>
</taxonomicName>
) or adult spotted-tailed quolls, adults have no predators, except dogs (
<taxonomicName id="D0A04D1AFFEAE71D3ED1FA831AFEFAAB" authorityName="Linnaeus" authorityYear="1758" box="[989,1166,1355,1378]" class="Mammalia" family="Canidae" genus="Canis" kingdom="Animalia" order="Carnivora" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="familiaris">
<emphasis id="25D4EA8BFFEAE71D3ED1FA831AFEFAAB" box="[989,1166,1355,1378]" italics="true" pageId="5" pageNumber="6">Canis familiaris</emphasis>
</taxonomicName>
). Vehicular traffic can be a significant source of mortality for animals scavenging for roadkill (Jones 2000; Hobday and Minstrell 2008).
</paragraph>
<paragraph id="171F3699FFEAE71D3E7FFA631D83F9AB" blockId="5.[840,1523,203,1954]" pageId="5" pageNumber="6">
<emphasis id="25D4EA8BFFEAE71D3E7FFA631A62FA0A" bold="true" box="[883,1042,1451,1475]" italics="true" pageId="5" pageNumber="6">Reproduction.</emphasis>
—Almost all dasyurids are seasonal breeders, with the period of pouch emergence and late lactation coinciding with spring and early summer in temperate
<collectingCountry id="6FB77609FFEAE71D3E44F9C31DDCF9EB" box="[840,940,1547,1570]" name="Australia" pageId="5" pageNumber="6">Australia</collectingCountry>
(Tyndale-Biscoe and Renfree 1987). Copulation in
<taxonomicName id="D0A04D1AFFEAE71D3E44F9E31A56F98B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[840,1062,1579,1602]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFEAE71D3E44F9E31A56F98B" box="[840,1062,1579,1602]" italics="true" pageId="5" pageNumber="6">Sarcophilus harrisii</emphasis>
</taxonomicName>
is most common between mid-February and late March.
</paragraph>
<paragraph id="171F3699FFEAE71D3E7FF9A31B19F8EB" blockId="5.[840,1523,203,1954]" pageId="5" pageNumber="6">
Gestation is 17.9 ± 1.0 days (
<emphasis id="25D4EA8BFFEAE71D39A9F9A31AB3F94B" box="[1189,1219,1643,1666]" italics="true" pageId="5" pageNumber="6">SE</emphasis>
: range 1422 days—Hester-man et al. 2008a), with most births in April but some as late as July. Young are carried in the pouch for about 1415 weeks, after which they are deposited in a den. They do not leave dens on their own until November and are weaned from mid-December to early February (
<bibRefCitation id="73314B68FFEAE71D3903F8C31AECF8EB" author="GUILER, E. R." box="[1039,1180,1803,1826]" pageId="5" pageNumber="6" pagination="63 - 70" refId="ref16876" refString="GUILER, E. R. 1970 a. Observations on the Tasmanian devil, Sarcophilus harrisii (Marsupialia: Dasyuridae). II. Reproduction, breeding, and growth of pouch young. Australian Journal of Zoology 18: 63 - 70." type="journal article" year="1970">Guiler 1970a</bibRefCitation>
;
<bibRefCitation id="73314B68FFEAE71D39A5F8C31B28F8EB" author="PEMBERTON, D." box="[1193,1368,1803,1826]" pageId="5" pageNumber="6" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFEAE71E3E7FF8E31C1EFEEB" blockId="5.[840,1523,203,1954]" lastBlockId="6.[109,793,203,1954]" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
Based on [22Na] and [3H] isotope-dilution techniques, females in late lactation have turnover rates per kg of body mass that were 90% and 60% (respectively) greater than those of males and non-lactating females, or of females carrying small young (Green and Eberhard 1979). The milk of
<taxonomicName id="D0A04D1AFFE9E71E3F81FF031C8BFF2B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[653,763,203,226]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE9E71E3F81FF031C8BFF2B" box="[653,763,203,226]" italics="true" pageId="6" pageNumber="7">S. harrisii</emphasis>
</taxonomicName>
is similar in 6 minerals to that of eutherian mammals, except for substantially higher levels of iron (
<bibRefCitation id="73314B68FFE9E71E3CEDFEC31C2EFEEB" author="GREEN, B." box="[481,606,267,290]" pageId="6" pageNumber="7" pagination="369 - 387" refId="ref16383" refString="GREEN, B. 1984. Composition of milk and energetics of growth in marsupials. Symposia of the Zoological Society of London 51: 369 - 387." type="journal article" year="1984">Green 1984</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE9E71E3D94FEE31D68FD6B" blockId="6.[109,793,203,1954]" pageId="6" pageNumber="7">
All
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males from April, May, and August exhibited spermatogenesis, and all males greater than 5.0 kg were judged to be fertile (
<bibRefCitation id="73314B68FFE9E71E3DF0FEA31FFBFE4B" author="HUGHES, R. L." box="[252,395,363,386]" pageId="6" pageNumber="7" refId="ref17904" refString="HUGHES, R. L. 1982. Reproduction in the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Marsupialia). Pp. 49 - 63 in Carnivorous marsupials (M. Archer, ed.). Royal Zoological Society of New South Wales, Sydney, Australia." type="book" year="1982">Hughes 1982</bibRefCitation>
). As in other marsupials, there is no evidence of either seminal vesicles or ampullary enlargement of the vas deferens, but a carrot-shaped prostate gland and 2 pairs of Cowpers glands are present (MacKenzie and Owen 1919). The prostate gland contains glucose as a minor constituent but has substantial glycogen (
<quantity id="D0589B7CFFE9E71E3C58FDC31FACFDEB" box="[340,476,523,546]" metricMagnitude="-4" metricUnit="kg" metricValue="4.5" metricValueMax="5.0" metricValueMin="4.0" pageId="6" pageNumber="7" unit="mg" value="450.0" valueMax="500.0" valueMin="400.0">400500 mg</quantity>
%—Rodger and White 1977). Although the form in which the prostatic glycogen reaches the semen is unknown, glycogen is found in extremely high concentrations (
<quantity id="D0589B7CFFE9E71E3DC0FDA31F45FD4B" box="[204,309,619,642]" metricMagnitude="-3" metricUnit="kg" metricValue="4.0" pageId="6" pageNumber="7" unit="mg" value="4000.0">4,000 mg</quantity>
%) in secretions recovered from the prostatic urethra and thus probably enters the semen intact (
<bibRefCitation id="73314B68FFE9E71E3F73FD431D78FD6B" author="RODGER, J. C." box="[639,776,651,674]" pageId="6" pageNumber="7" pagination="73 - 80" refId="ref20869" refString="RODGER, J. C. 1978. Male reproduction - its usefulness in discussions of Macropodidae evolution. Australian Mammalogy 2: 73 - 80." type="journal article" year="1978">Rodger 1978</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE9E71E3D94FD631F6BFC6B" blockId="6.[109,793,203,1954]" pageId="6" pageNumber="7">
The scrotum, which is evident at birth (
<bibRefCitation id="73314B68FFE9E71E3F33FD631CBBFD0B" author="GUILER, E. R." box="[575,715,683,706]" pageId="6" pageNumber="7" pagination="63 - 70" refId="ref16876" refString="GUILER, E. R. 1970 a. Observations on the Tasmanian devil, Sarcophilus harrisii (Marsupialia: Dasyuridae). II. Reproduction, breeding, and growth of pouch young. Australian Journal of Zoology 18: 63 - 70." type="journal article" year="1970">Guiler 1970a</bibRefCitation>
), has a temperature at maturity of about 27.7°C, or 5.6°C lower than the body temperature of anesthetized animals (
<bibRefCitation id="73314B68FFE9E71E3F46FD231CAFFCCB" author="SETCHELL, B. P." box="[586,735,747,770]" pageId="6" pageNumber="7" refId="ref21092" refString="SETCHELL, B. P. 1977. Reproduction in male marsupials. Pp. 411 - 457 in The biology of marsupials (B. Stonehouse and D. Gilmore, eds.). The Macmillan Press Ltd., London, United Kingdom." type="book" year="1977">Setchell 1977</bibRefCitation>
). By contrast, Guiler and Heddle (1970), using thermistors inside the scrotum, found testicular and body temperatures to be similar (34.5 ± 1°C). The diameter of the seminiferous tubules in the testes of
<taxonomicName id="D0A04D1AFFE9E71E3DDEFCA31F4FFC4B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[210,319,875,898]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE9E71E3DDEFCA31F4FFC4B" box="[210,319,875,898]" italics="true" pageId="6" pageNumber="7">S. harrisii</emphasis>
</taxonomicName>
is
<quantity id="D0589B7CFFE9E71E3C68FCA31FCBFC4B" box="[356,443,875,899]" metricMagnitude="-1" metricUnit="m" metricValue="3.6" pageId="6" pageNumber="7" unit="mm" value="360.0">360 µm</quantity>
, an average value for dasyurids (
<bibRefCitation id="73314B68FFE9E71E3D7AFC431F7BFC6B" author="WOOLLEY, P." box="[118,267,907,930]" pageId="6" pageNumber="7" pagination="255 - 261" refId="ref21946" refString="WOOLLEY, P. 1975. The seminiferous tubules in dasyurid marsupials. Journal of Reproduction and Fertility 45: 255 - 261." type="journal article" year="1975">Woolley 1975</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE9E71E3D94FC631E82FA8B" blockId="6.[109,793,203,1954]" pageId="6" pageNumber="7">
The sperm of
<taxonomicName id="D0A04D1AFFE9E71E3C30FC631FDEFC0B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[316,430,939,962]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE9E71E3C30FC631FDEFC0B" box="[316,430,939,962]" italics="true" pageId="6" pageNumber="7">S. harrisii</emphasis>
</taxonomicName>
, at
<quantity id="D0589B7CFFE9E71E3CD2FC631C48FC0B" box="[478,568,939,963]" metricMagnitude="-1" metricUnit="m" metricValue="2.4" pageId="6" pageNumber="7" unit="mm" value="240.0">240 µm</quantity>
, is among the largest in mammals. Mature sperm in the proximal one-third of the epididymis show an unusual feature: the long axis of the head is held at an angle of 6090° to the axis of the tail, supported by a conical cytoplasmic droplet (
<bibRefCitation id="73314B68FFE9E71E3F3CFBE31C96FB8B" author="HUGHES, R. L." box="[560,742,1067,1090]" pageId="6" pageNumber="7" refId="ref17904" refString="HUGHES, R. L. 1982. Reproduction in the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Marsupialia). Pp. 49 - 63 in Carnivorous marsupials (M. Archer, ed.). Royal Zoological Society of New South Wales, Sydney, Australia." type="book" year="1982">Hughes 1982:60</bibRefCitation>
, figure 28). Identical in gross morphology and in ultrastructure to those of
<taxonomicName id="D0A04D1AFFE9E71E3DC1FBA31FDEFB4B" baseAuthorityName="Shaw" baseAuthorityYear="1800" box="[205,430,1131,1154]" class="Mammalia" family="Dasyuridae" genus="Dasyurus" kingdom="Animalia" order="Dasyuromorphia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="viverrinus">
<emphasis id="25D4EA8BFFE9E71E3DC1FBA31FDEFB4B" box="[205,430,1131,1154]" italics="true" pageId="6" pageNumber="7">Dasyurus viverrinus</emphasis>
</taxonomicName>
(eastern quoll), a closely related dasyurid, the sperm also share some unusual features with the bandicoots (Peramelidae—Hughes 1982). Whether spermiogenesis is suspended during part of the year in adult males is moot. Among large dasyurids, only
<taxonomicName id="D0A04D1AFFE9E71E3CF6FB231C19FACB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[506,617,1259,1282]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE9E71E3CF6FB231C19FACB" box="[506,617,1259,1282]" italics="true" pageId="6" pageNumber="7">S. harrisii</emphasis>
</taxonomicName>
has sufficiently large testes to be a candidate for sperm competition (
<bibRefCitation id="73314B68FFE9E71E3FC9FAC31E92FA8B" author="TAGGART, D. A. &amp; G. A. SHIMMIN &amp; C. R. DICKMAN &amp; W. G. BREED" pageId="6" pageNumber="7" refId="ref21445" refString="TAGGART, D. A., G. A. SHIMMIN, C. R. DICKMAN, AND W. G. BREED. 2003. Reproductive biology of carnivorous marsupials: clues to the likelihood of sperm competition. Pp. 358 - 375 in Predators with pouches (M. Jones, C. Dickman, and M. Archer, eds.). CSIRO Publishing, Collingwood, Victoria, Australia." type="book" year="2003">Taggart et al. 2003</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE9E71E3D94FA831E96F86B" blockId="6.[109,793,203,1954]" pageId="6" pageNumber="7">
The onset of breeding in females is marked by a conspicuous enlargement of the pouch to a moist, fist-sized structure with 4 well-developed mammary glands (
<bibRefCitation id="73314B68FFE9E71E3CD1FA431CA3FA6B" author="HESTERMAN, H. &amp; S. M. JONES &amp; F. SCHWARZENBERGER" box="[477,723,1419,1442]" pageId="6" pageNumber="7" pagination="130 - 138" refId="ref17493" refString="HESTERMAN, H., S. M. JONES, AND F. SCHWARZENBERGER. 2008 b. Pouch appearance is a reliable indicator of the reproductive status in the Tasmanian devil and the spotted-tailed quoll. Journal of Zoology (London) 275: 130 - 138." type="journal article" year="2008">Hesterman et al. 2008b</bibRefCitation>
), with the uterus showing hypertrophy in association with preovulatory enlargement of Graafian follicles (
<bibRefCitation id="73314B68FFE9E71E3CE6FA031C0AFA2B" author="HUGHES, R. L." box="[490,634,1483,1506]" pageId="6" pageNumber="7" refId="ref17904" refString="HUGHES, R. L. 1982. Reproduction in the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Marsupialia). Pp. 49 - 63 in Carnivorous marsupials (M. Archer, ed.). Royal Zoological Society of New South Wales, Sydney, Australia." type="book" year="1982">Hughes 1982</bibRefCitation>
;
<bibRefCitation id="73314B68FFE9E71E3F80FA031ED2F9CB" author="KEELEY, T. &amp; J. K. O'BRIEN &amp; B. G. FANSON &amp; K. MASTERS &amp; P. D. MCGREEVY" pageId="6" pageNumber="7" pagination="182 - 191" refId="ref18983" refString="KEELEY, T., J. K. O'BRIEN, B. G. FANSON, K. MASTERS, AND P. D. MCGREEVY. 2012. The reproductive cycle of the Tasmanian devil (Sarcophilus harrisii) and factors associated with reproductive success in captivity. General and Comparative Endocrinology 176: 182 - 191." type="journal article" year="2012">Keeley et al. 2012</bibRefCitation>
). Behavioral predictors of estrus in captive females include development of a fluid-filled neck roll, increased nesting behavior, loss of appetite, and lethargy (
<bibRefCitation id="73314B68FFE9E71E3CD4F9E31CE7F98B" author="KEELEY, T. &amp; J. K. O'BRIEN &amp; B. G. FANSON &amp; K. MASTERS &amp; P. D. MCGREEVY" box="[472,663,1579,1602]" pageId="6" pageNumber="7" pagination="182 - 191" refId="ref18983" refString="KEELEY, T., J. K. O'BRIEN, B. G. FANSON, K. MASTERS, AND P. D. MCGREEVY. 2012. The reproductive cycle of the Tasmanian devil (Sarcophilus harrisii) and factors associated with reproductive success in captivity. General and Comparative Endocrinology 176: 182 - 191." type="journal article" year="2012">Keeley et al. 2012</bibRefCitation>
). Ovulation is spontaneous, with large numbers of eggs released. Like other dasyurids,
<taxonomicName id="D0A04D1AFFE9E71E3DEFF9A31F20F94B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[227,336,1643,1666]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE9E71E3DEFF9A31F20F94B" box="[227,336,1643,1666]" italics="true" pageId="6" pageNumber="7">S. harrisii</emphasis>
</taxonomicName>
females produce more eggs and embryos (21 eggs—Flynn 1922; 15 embryos—Guiler 1970a; 11
<specimenCount id="01A6FD10FFE9E71E3FC9F9431D68F96B" box="[709,792,1675,1698]" pageId="6" pageNumber="7" type="egg">56 eggs</specimenCount>
and embryos in 6 females—Hughes 1982) than can be accommodated by the 4 teats. Although 4 is the maximum litter, usually only 2 or 3 survive to weaning age.
<bibRefCitation id="73314B68FFE9E71E3F36F9231CA0F8CB" author="GUILER, E. R." box="[570,720,1771,1794]" pageId="6" pageNumber="7" pagination="161 - 178" refId="ref16996" refString="GUILER, E. R. 1978. Observations on the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Mammalia) at Granville Harbour. Papers and Proceedings of the Royal Society of Tasmania 112: 161 - 178." type="journal article" year="1978">Guiler (1978)</bibRefCitation>
found up to 80% of eligible females carrying pouch young; across his 10-year study 58.1% of females had young (
<emphasis id="25D4EA8BFFE9E71E3F5BF8E01C1BF8F4" box="[599,619,1832,1853]" italics="true" pageId="6" pageNumber="7">X</emphasis>
= 2.67 young). By contrast,
<bibRefCitation id="73314B68FFE9E71E3DFEF8831FDAF8AB" author="PEMBERTON, D." box="[242,426,1867,1890]" pageId="6" pageNumber="7" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton (1990</bibRefCitation>
—2.30) and
<bibRefCitation id="73314B68FFE9E71E3F21F8831C8DF8AB" author="LACHISH, S. &amp; H. MCCALLUM &amp; M. JONES" box="[557,765,1867,1890]" pageId="6" pageNumber="7" pagination="427 - 436" refId="ref19198" refString="LACHISH, S., H. MCCALLUM, AND M. JONES. 2009. Demography, disease, and the devil: life-history changes in a disease-affected population of the Tasmanian devil (Sarcophilus harrisii). Journal of Animal Ecology 78: 427 - 436." type="journal article" year="2009">Lachish et al. (2009</bibRefCitation>
— 3.40) recorded lower and greater mean numbers of pouch young per female.
</paragraph>
<paragraph id="171F3699FFE9E71E3E7FFF031DFDFCEB" blockId="6.[840,1523,203,1538]" pageId="6" pageNumber="7">
The 14-day follicular phase of the estrous cycle is followed by an 18-day luteal phase leading to ovulation (
<bibRefCitation id="73314B68FFE9E71E3849FF231DF9FEEB" author="HESTERMAN, H. &amp; S. M. JONES &amp; F. SCHWARZENBERGER" pageId="6" pageNumber="7" pagination="234 - 244" refId="ref17435" refString="HESTERMAN, H., S. M. JONES, AND F. SCHWARZENBERGER. 2008 a. Reproductive endocrinology of the large dasyurids: characterization of ovarian cycles by plasma and fecal steroid monitoring. Part I. The Tasmanian devil (Sarcophilus harrisii). General and Comparative Endocrinology 155: 234 - 244." type="journal article" year="2008">Hesterman et al. 2008a</bibRefCitation>
). In a zoo-study sample with 10 times more females,
<bibRefCitation id="73314B68FFE9E71E3E44FEE31A67FE8B" author="KEELEY, T. &amp; J. K. O'BRIEN &amp; B. G. FANSON &amp; K. MASTERS &amp; P. D. MCGREEVY" box="[840,1047,299,322]" pageId="6" pageNumber="7" pagination="182 - 191" refId="ref18983" refString="KEELEY, T., J. K. O'BRIEN, B. G. FANSON, K. MASTERS, AND P. D. MCGREEVY. 2012. The reproductive cycle of the Tasmanian devil (Sarcophilus harrisii) and factors associated with reproductive success in captivity. General and Comparative Endocrinology 176: 182 - 191." type="journal article" year="2012">Keeley et al. (2012)</bibRefCitation>
report similar lengths of the luteal phases for both unsuccessful and successful matings (12.5 ± 1.4 (
<emphasis id="25D4EA8BFFE9E71E3898FE831BC2FEAB" box="[1428,1458,331,354]" italics="true" pageId="6" pageNumber="7">SE</emphasis>
) days for the latter group). Pregnancy is accommodated within the luteal phase, and regression of the corpus luteum coincides with parturition. If lactation follows, the next follicular phase is suppressed and the corpus luteum of pregnancy slowly disappears or shrinks to a corpus albicans (Tyndale-Biscoe and Renfree 1987). An infertile cycle or loss of a litter is followed by another estrus after an interval of 33.7 ± 5.9 (
<emphasis id="25D4EA8BFFE9E71E398AFDE31AD4FD8B" box="[1158,1188,555,578]" italics="true" pageId="6" pageNumber="7">SE</emphasis>
) days (
<bibRefCitation id="73314B68FFE9E71E39FDFDE31B93FD8B" author="HESTERMAN, H. &amp; S. M. JONES &amp; F. SCHWARZENBERGER" box="[1265,1507,555,578]" pageId="6" pageNumber="7" pagination="234 - 244" refId="ref17435" refString="HESTERMAN, H., S. M. JONES, AND F. SCHWARZENBERGER. 2008 a. Reproductive endocrinology of the large dasyurids: characterization of ovarian cycles by plasma and fecal steroid monitoring. Part I. The Tasmanian devil (Sarcophilus harrisii). General and Comparative Endocrinology 155: 234 - 244." type="journal article" year="2008">Hesterman et al. 2008a</bibRefCitation>
). As many as 3 estrous cycles in a single year among females with unsuccessful matings are known (
<bibRefCitation id="73314B68FFE9E71E39A0FDA31B1AFD4B" author="KEELEY, T. &amp; J. K. O'BRIEN &amp; B. G. FANSON &amp; K. MASTERS &amp; P. D. MCGREEVY" box="[1196,1386,619,642]" pageId="6" pageNumber="7" pagination="182 - 191" refId="ref18983" refString="KEELEY, T., J. K. O'BRIEN, B. G. FANSON, K. MASTERS, AND P. D. MCGREEVY. 2012. The reproductive cycle of the Tasmanian devil (Sarcophilus harrisii) and factors associated with reproductive success in captivity. General and Comparative Endocrinology 176: 182 - 191." type="journal article" year="2012">Keeley et al. 2012</bibRefCitation>
).
<bibRefCitation id="73314B68FFE9E71E3873FDA31DABFD6B" author="HESTERMAN, H. &amp; S. M. JONES &amp; F. SCHWARZENBERGER" pageId="6" pageNumber="7" pagination="234 - 244" refId="ref17435" refString="HESTERMAN, H., S. M. JONES, AND F. SCHWARZENBERGER. 2008 a. Reproductive endocrinology of the large dasyurids: characterization of ovarian cycles by plasma and fecal steroid monitoring. Part I. The Tasmanian devil (Sarcophilus harrisii). General and Comparative Endocrinology 155: 234 - 244." type="journal article" year="2008">Hesterman et al. (2008a)</bibRefCitation>
documented female reproductive cycles between January and June, thus explaining the occasional presence of pouch young in late winter (JulySeptember—Green 1967;
<bibRefCitation id="73314B68FFE9E71E3E44FD231DA9FCCB" author="GUILER, E. R." box="[840,985,747,770]" pageId="6" pageNumber="7" pagination="63 - 70" refId="ref16876" refString="GUILER, E. R. 1970 a. Observations on the Tasmanian devil, Sarcophilus harrisii (Marsupialia: Dasyuridae). II. Reproduction, breeding, and growth of pouch young. Australian Journal of Zoology 18: 63 - 70." type="journal article" year="1970">Guiler 1970a</bibRefCitation>
) and 1-month-old young in August (
<bibRefCitation id="73314B68FFE9E71E3873FD231D0DFCEB" author="PEMBERTON, D." pageId="6" pageNumber="7" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE9E71E3E7FFCE31A5CFBEB" blockId="6.[840,1523,203,1538]" pageId="6" pageNumber="7">
Eggs are about
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in diameter (
<bibRefCitation id="73314B68FFE9E71E39E1FCE31B09FC8B" author="HUGHES, R. L." box="[1261,1401,811,834]" pageId="6" pageNumber="7" refId="ref17904" refString="HUGHES, R. L. 1982. Reproduction in the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Marsupialia). Pp. 49 - 63 in Carnivorous marsupials (M. Archer, ed.). Royal Zoological Society of New South Wales, Sydney, Australia." type="book" year="1982">Hughes 1982</bibRefCitation>
), including
<specimenCount id="01A6FD10FFE9E71E3E44FC831DF2FCAB" box="[840,898,843,866]" pageId="6" pageNumber="7" type="egg">3 egg</specimenCount>
membranes: zona pellucida (
<quantity id="D0589B7CFFE9E71E39B1FC831A88FCAB" box="[1213,1272,843,867]" metricMagnitude="-3" metricUnit="m" metricValue="4.0" pageId="6" pageNumber="7" unit="mm" value="4.0">4 µm</quantity>
), mucoid coat (
<quantity id="D0589B7CFFE9E71E3892FC831B96FCAB" box="[1438,1510,843,867]" metricMagnitude="-2" metricUnit="m" metricValue="2.8" pageId="6" pageNumber="7" unit="mm" value="28.0">28 µm</quantity>
), and shell membrane (
<quantity id="D0589B7CFFE9E71E394FFCA31AF4FC4B" box="[1091,1156,875,899]" metricMagnitude="-3" metricUnit="m" metricValue="5.0" pageId="6" pageNumber="7" unit="mm" value="5.0">5 µm</quantity>
). All membranes and numerous trapped sperm remain intact while cleavages produce the cells of a unilaminar blastocyst (bypassing the morula stage of development). At least 20% of uterine zygotes fail to cleave and in some females less than one-half develop to the expanded blastocyst stage (
<bibRefCitation id="73314B68FFE9E71E3E82FBC31A6BFBEB" author="HUGHES, R. L." box="[910,1051,1035,1058]" pageId="6" pageNumber="7" refId="ref17904" refString="HUGHES, R. L. 1982. Reproduction in the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Marsupialia). Pp. 49 - 63 in Carnivorous marsupials (M. Archer, ed.). Royal Zoological Society of New South Wales, Sydney, Australia." type="book" year="1982">Hughes 1982</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE9E71E3E7FFBE31D88FAEB" blockId="6.[840,1523,203,1538]" pageId="6" pageNumber="7">
The pouch, which develops from a circular fold of skin in the lower abdomen, expands as if to accommodate the growing young, opening backwards and downwards. The sex ratio of pouch young was unity in one study (
<bibRefCitation id="73314B68FFE9E71E3800FB431BCEFB6B" author="PEMBERTON, D." box="[1292,1470,1163,1186]" pageId="6" pageNumber="7" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
) but favored females (nearly double) in another (
<bibRefCitation id="73314B68FFE9E71E3814FB631BD3FB0B" author="GUILER, E. R." box="[1304,1443,1195,1218]" pageId="6" pageNumber="7" pagination="63 - 70" refId="ref16876" refString="GUILER, E. R. 1970 a. Observations on the Tasmanian devil, Sarcophilus harrisii (Marsupialia: Dasyuridae). II. Reproduction, breeding, and growth of pouch young. Australian Journal of Zoology 18: 63 - 70." type="journal article" year="1970">Guiler 1970a</bibRefCitation>
), a pattern seen in another population (
<bibRefCitation id="73314B68FFE9E71E39A7FB031B0CFB2B" author="LACHISH, S. &amp; H. MCCALLUM &amp; M. JONES" box="[1195,1404,1227,1250]" pageId="6" pageNumber="7" pagination="427 - 436" refId="ref19198" refString="LACHISH, S., H. MCCALLUM, AND M. JONES. 2009. Demography, disease, and the devil: life-history changes in a disease-affected population of the Tasmanian devil (Sarcophilus harrisii). Journal of Animal Ecology 78: 427 - 436." type="journal article" year="2009">Lachish et al. 2009</bibRefCitation>
), in which litters in diseased females had twice as many female pups as healthy females.
</paragraph>
<paragraph id="171F3699FFE9E71E3E7FFAE31B06F9CB" blockId="6.[840,1523,203,1538]" pageId="6" pageNumber="7">
Young females average 2.0 young in the 1st litters (
<bibRefCitation id="73314B68FFE9E71E38A3FAE31D0DFAAB" author="GUILER, E. R." pageId="6" pageNumber="7" pagination="161 - 178" refId="ref16996" refString="GUILER, E. R. 1978. Observations on the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Mammalia) at Granville Harbour. Papers and Proceedings of the Royal Society of Tasmania 112: 161 - 178." type="journal article" year="1978">Guiler 1978</bibRefCitation>
), but have 3.6 young per year during their 2nd4th seasons, and 2.0 young in their last season, making a total of 14.8 young for the female that lives the longest in the wild.
<bibRefCitation id="73314B68FFE9E71E3873FA431DFFFA0B" author="PEMBERTON, D." pageId="6" pageNumber="7" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton (1990)</bibRefCitation>
also found that fertility decreased in older females, as indicated by increases in reproductive failure (no pouch young) as well as reductions in litter size from earlier years.
</paragraph>
</subSection>
</subSubSection>
<subSection id="7E2F2F72FFE9E710395EF98E1A2FFC6B" lastPageId="8" lastPageNumber="9" pageId="6" pageNumber="7" type="multiple">
<subSubSection id="5FBA6512FFE9E710395EF98E1A2FFC6B" lastPageId="8" lastPageNumber="9" pageId="6" pageNumber="7" type="biology_ecology">
<paragraph id="171F3699FFE9E71E395EF98E1A98F9A9" blockId="6.[1106,1256,1606,1632]" box="[1106,1256,1606,1632]" pageId="6" pageNumber="7">
<heading id="4C5781F5FFE9E71E395EF98E1A98F9A9" allCaps="true" bold="true" box="[1106,1256,1606,1632]" centered="true" fontSize="11" level="1" pageId="6" pageNumber="7" reason="6">
<emphasis id="25D4EA8BFFE9E71E395EF98E1A98F9A9" bold="true" box="[1106,1256,1606,1632]" pageId="6" pageNumber="7">ECOLOGY</emphasis>
</heading>
</paragraph>
<paragraph id="171F3699FFE9E71F3E7FF9421F59FD6B" blockId="6.[840,1523,1674,1953]" lastBlockId="7.[109,793,203,1954]" lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7">
<emphasis id="25D4EA8BFFE9E71E3E7FF9421AEDF96B" bold="true" box="[883,1181,1674,1698]" italics="true" pageId="6" pageNumber="7">Population characteristics.</emphasis>
—Both distribution and density of
<taxonomicName id="D0A04D1AFFE9E71E3E60F9621A3CF908" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[876,1100,1706,1729]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE9E71E3E60F9621A3CF908" box="[876,1100,1706,1729]" italics="true" pageId="6" pageNumber="7">Sarcophilus harrisii</emphasis>
</taxonomicName>
correlate with low-to-medium mean annual rainfall (Jones and Rose 1996). Densities sometimes exceed 1 animal per km
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in
</quantity>
the eastern one-half of
<collectingRegion id="D564F87BFFE9E71E3888F9221B9EF8C8" box="[1412,1518,1770,1793]" country="Australia" name="Tasmania" pageId="6" pageNumber="7">Tasmania</collectingRegion>
, across the northern coastal region, and in a narrow strip down the west coast (Jones and Rose 1996). Population density varies across the landscape at a scale of tens of kilometers.
<taxonomicName id="D0A04D1AFFE9E71E3E44F8A21DC5F848" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[840,949,1898,1921]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE9E71E3E44F8A21DC5F848" box="[840,949,1898,1921]" italics="true" pageId="6" pageNumber="7">S. harrisii</emphasis>
</taxonomicName>
was not always common, for
<bibRefCitation id="73314B68FFE9E71E39F4F8A21BF7F848" author="FLYNN, T. T." box="[1272,1415,1898,1921]" pageId="6" pageNumber="7" pagination="16 - 19" refId="ref16185" refString="FLYNN, T. T. 1939. Note on the egg of the rare marsupial Sarcophilus. Proceedings of the Royal Irish Academy 1938 - 1939 45: 16 - 19." type="journal article" year="1939">Flynn (1939)</bibRefCitation>
describes the species as rare in his title. Earlier,
<bibRefCitation id="73314B68FFE9E71E382AF8421BCDF868" author="FLYNN, T. T." box="[1318,1469,1930,1953]" pageId="6" pageNumber="7" pagination="225 - 231" refId="ref16154" refString="FLYNN, T. T. 1922. Notes on certain reproductive phenomena in some Tasmanian marsupials. Annals of the Magazine of Natural History 9: 225 - 231." type="journal article" year="1922">Flynn (1922)</bibRefCitation>
had difficulty obtaining sufficient animals over 18 years of effort to study its reproduction. Numbers were thought to be low also in the 1860s, the early1900s, and the 1940s (
<bibRefCitation id="73314B68FFE8E71F3F3AFEC31CC8FEEB" author="GUILER, E. R." box="[566,696,267,290]" pageId="7" pageNumber="8" refId="ref17036" refString="GUILER, E. R. 1992. The Tasmanian devil. St. David's Park Publishing, Hobart, Australia." type="book" year="1992">Guiler 1992</bibRefCitation>
).
<taxonomicName id="D0A04D1AFFE8E71F3FC3FEC31EE9FE8B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3FC3FEC31EE9FE8B" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
may have been most abundant in the 1970s1990s, with estimates of 130,000 (Jones and Rose 1996). However, since the mid-1990s, an aggressive, rapidly progressing, and fatal disease, characterized by facial tumors, has resulted in massive population declines. The disease, 1st detected in
<quantity id="D0589B7CFFE8E71F3F9BFE631C9DFE0B" box="[663,749,427,450]" metricMagnitude="1" metricUnit="m" metricValue="5.06984" pageId="7" pageNumber="8" unit="in" value="1996.0">1996 in</quantity>
the dense populations of the northeast coast, has spread westward across the state at the rate of about
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per year, resulting in an estimated loss of one-third to one-half of the population by the end of 2003 (Jones 2003b) and perhaps 80% declines later (
<bibRefCitation id="73314B68FFE8E71F3D7AFD831F17FDAB" author="LACHISH, S. &amp; M. JONES &amp; H. MCCALLUM" box="[118,359,587,610]" pageId="7" pageNumber="8" pagination="926 - 936" refId="ref19156" refString="LACHISH, S., M. JONES, AND H. MCCALLUM. 2007. The impact of disease on the survival and population growth rate of the Tasmanian devil. Journal of Animal Ecology 76: 926 - 936." type="journal article" year="2007">McCallum et al. 2007</bibRefCitation>
). In 2016, the disease still was moving westward unabated but disease-free populations still remain in parts of the state.
</paragraph>
<paragraph id="171F3699FFE8E71F3D94FD631F05FC4B" blockId="7.[109,793,203,1954]" pageId="7" pageNumber="8">
Numbers also vary from year to year. A study population on the west coast ranged from 20 to 106 animals over 10 years (
<bibRefCitation id="73314B68FFE8E71F3D7AFD231E86FCCB" author="GUILER, E. R." box="[118,246,747,770]" pageId="7" pageNumber="8" pagination="161 - 178" refId="ref16996" refString="GUILER, E. R. 1978. Observations on the Tasmanian devil, Sarcophilus harrisii (Dasyuridae, Mammalia) at Granville Harbour. Papers and Proceedings of the Royal Society of Tasmania 112: 161 - 178." type="journal article" year="1978">Guiler 1978</bibRefCitation>
).
<bibRefCitation id="73314B68FFE8E71F3C00FD231FBEFCCB" author="PEMBERTON, D." box="[268,462,747,770]" pageId="7" pageNumber="8" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton (1990)</bibRefCitation>
, who trapped and radiotracked
<taxonomicName id="D0A04D1AFFE8E71F3D61FCC31EA9FCEB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[109,217,779,802]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3D61FCC31EA9FCEB" box="[109,217,779,802]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
in the prime habitat of Mt. William National Park in northeastern
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, estimated a maximum of 200 animals on his 16-km
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study area, with mark-recapture estimates being nearly double that value.
</paragraph>
<paragraph id="171F3699FFE8E71F3D94FC431CA1FBAB" blockId="7.[109,793,203,1954]" pageId="7" pageNumber="8">
After the large influx of weaned young into the population during January and February, the population was 72% young of the year, 7% juveniles (1224 months of age), and 21% adults (
<bibRefCitation id="73314B68FFE8E71F3DB2FC231F1EFBCB" author="PEMBERTON, D." box="[190,366,1003,1026]" pageId="7" pageNumber="8" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
). Young females tend to be philopatric, whereas most young males disperse. In healthy populations, the number of transient
<taxonomicName id="D0A04D1AFFE8E71F3C4EFBE31FDBFB8B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[322,427,1067,1090]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3C4EFBE31FDBFB8B" box="[322,427,1067,1090]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
decreases throughout the year, but the number of adult residents remains relatively constant.
</paragraph>
<paragraph id="171F3699FFE8E71F3D94FBA31FCEFB2B" blockId="7.[109,793,203,1954]" pageId="7" pageNumber="8">
Preweaning mortality was 27%, but far greater mortality (80%) was suffered by juveniles during their 1st year of independence (
<bibRefCitation id="73314B68FFE8E71F3DB1FB631F1EFB0B" author="PEMBERTON, D." box="[189,366,1195,1218]" pageId="7" pageNumber="8" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
). Annual adult mortality of 20% probably increases in older animals.
</paragraph>
<paragraph id="171F3699FFE8E71F3D94FB231E8DFA6B" blockId="7.[109,793,203,1954]" pageId="7" pageNumber="8">
<emphasis id="25D4EA8BFFE8E71F3D94FB231C66FACB" box="[152,534,1259,1283]" italics="true" pageId="7" pageNumber="8">
<emphasis id="25D4EA8BFFE8E71F3D94FB231F68FACA" bold="true" box="[152,280,1259,1283]" italics="true" pageId="7" pageNumber="8">Space use.</emphasis>
<taxonomicName id="D0A04D1AFFE8E71F3C3FFB231C66FACB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[307,534,1259,1282]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">Sarcophilus harrisii</taxonomicName>
</emphasis>
occupies overlapping home ranges that average
<quantity id="D0589B7CFFE8E71F3CAEFAC31C70FAEB" box="[418,512,1291,1314]" metricMagnitude="4" metricUnit="m" metricValue="1.33" pageId="7" pageNumber="8" unit="km" value="13.3">13.3 km</quantity>
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(range
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for both sexes); the smallest are those of females with young in dens (
<bibRefCitation id="73314B68FFE8E71F3DBFFA831F19FAAB" author="PEMBERTON, D." box="[179,361,1355,1378]" pageId="7" pageNumber="8" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
). Females attend denned young every night at 1st, but more sporadically later as the young approach weaning age.
</paragraph>
<paragraph id="171F3699FFE8E71F3D94FA631EC3F88B" blockId="7.[109,793,203,1954]" pageId="7" pageNumber="8">
Its widespread distribution within
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suggests that
<taxonomicName id="D0A04D1AFFE8E71F3D61FA031EAAFA2B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[109,218,1483,1506]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3D61FA031EAAFA2B" box="[109,218,1483,1506]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
has wide habitat tolerances. In general,
<taxonomicName id="D0A04D1AFFE8E71F3F9EFA031C8EFA2B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[658,766,1483,1506]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3F9EFA031C8EFA2B" box="[658,766,1483,1506]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
is found in open forest and woodlands that are not too shrubby at ground level and so offer clear runs for hunting. It avoids open habitats, steep or rocky areas, and tall or dense wet forests, including rainforest (Jones and Rose 1996; Jones and Barmuta 2000), but is attracted to predictable point sources of food, such as roadkills (Jones 2000), garbage pits, “devil restaurants” for tourism, and carcass dumps on farms.
<taxonomicName id="D0A04D1AFFE8E71F3CF1F9631C17F90B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[509,615,1707,1730]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3CF1F9631C17F90B" box="[509,615,1707,1730]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
reaches its highest population densities where prey is abundant, such as the short green sward of a sheep pasture that is attractive to macropods, its primary native prey in some locations (Jones and Barmuta 2000).
</paragraph>
<paragraph id="171F3699FFE8E71F3D94F8831AB9FDEB" blockId="7.[109,793,203,1954]" lastBlockId="7.[840,1523,203,1954]" pageId="7" pageNumber="8">
<emphasis id="25D4EA8BFFE8E71F3D94F8831FB3F8AB" box="[152,451,1867,1891]" italics="true" pageId="7" pageNumber="8">
<emphasis id="25D4EA8BFFE8E71F3D94F8831EBDF8AA" bold="true" box="[152,205,1867,1891]" italics="true" pageId="7" pageNumber="8">Diet.</emphasis>
<taxonomicName id="D0A04D1AFFE8E71F3DE5F8831FB3F8AB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[233,451,1867,1890]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">Sarcophilus harrisii</taxonomicName>
</emphasis>
, a dweller of forest, woodland, and coastal scrub, is a generalist predator and scavenger, eating a variety of vertebrates, but mostly medium-to-large mammals.
<taxonomicName id="D0A04D1AFFE8E71F3E44FF031DC7FF2B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[840,951,203,226]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3E44FF031DC7FF2B" box="[840,951,203,226]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
undoubtedly has benefited from the introduction of sheep in 1804;
<collectingRegion id="D564F87BFFE8E71F3EFCFF231A27FECB" box="[1008,1111,235,258]" country="Australia" name="Tasmania" pageId="7" pageNumber="8">Tasmania</collectingRegion>
had 2.4 million sheep in 2015, many of them under low-intensity “rangeland” management which results in many mortalities.
<taxonomicName id="D0A04D1AFFE8E71F397FFEE31AAFFE8B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[1139,1247,299,322]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F397FFEE31AAFFE8B" box="[1139,1247,299,322]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
can be a significant predator on young and moribund sheep if livestock husbandry practices are poor. Some breeds are better than others at protecting their lambs, but 1st borns of multiple births are particularly vulnerable while the 2nd is being born. The frequent presence of wool in feces indicates that sheep are often part of the diet of
<taxonomicName id="D0A04D1AFFE8E71F3E64FE231DA7FDCB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[872,983,491,514]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3E64FE231DA7FDCB" box="[872,983,491,514]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
, especially in the drier eastern half of
<collectingRegion id="D564F87BFFE8E71F3887FE231B82FDCB" box="[1419,1522,491,514]" country="Australia" name="Tasmania" pageId="7" pageNumber="8">Tasmania</collectingRegion>
where most of the sheep are raised.
</paragraph>
<paragraph id="171F3699FFE8E71F3E7FFDE31B7AFCEB" blockId="7.[840,1523,203,1954]" pageId="7" pageNumber="8">
<taxonomicName id="D0A04D1AFFE8E71F3E7FFDE31A39FD8B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[883,1097,555,578]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3E7FFDE31A39FD8B" box="[883,1097,555,578]" italics="true" pageId="7" pageNumber="8">Sarcophilus harrisii</emphasis>
</taxonomicName>
can consume food equal to as much as 40% of its body mass, making them noticeably plump (
<bibRefCitation id="73314B68FFE8E71F3873FD831D0DFD4B" author="PEMBERTON, D." pageId="7" pageNumber="8" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
).
<taxonomicName id="D0A04D1AFFE8E71F3E9EFDA31D8CFD4B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[914,1020,619,642]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3E9EFDA31D8CFD4B" box="[914,1020,619,642]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
consumes hair and fragmented bones as well as the flesh of prey, leaving only parts of large skulls and the often still-filled colon. The scraps are strewn over an area of as much as
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. Dried chalky feces, often 20 by
<quantity id="D0589B7CFFE8E71F381BFD031B1FFD2B" box="[1303,1391,715,738]" metricMagnitude="-1" metricUnit="m" metricValue="1.5" pageId="7" pageNumber="8" unit="mm" value="150.0">150 mm</quantity>
and grayish from the remains of digested bone (
<bibRefCitation id="73314B68FFE8E71F39C8FD231B4AFCCB" author="JONES, M. E." box="[1220,1338,747,770]" pageId="7" pageNumber="8" refId="ref18123" refString="JONES, M. E. 1995. Guild structure of the large marsupial carnivores in Tasmania. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1995">Jones 1995</bibRefCitation>
), usually contain bone fragments and large amounts of hair.
</paragraph>
<paragraph id="171F3699FFE8E71F3E7FFCE31D80F94B" blockId="7.[840,1523,203,1954]" pageId="7" pageNumber="8">
In the central highlands of
<collectingRegion id="D564F87BFFE8E71F399BFCE31A8FFC8B" box="[1175,1279,811,834]" country="Australia" name="Tasmania" pageId="7" pageNumber="8">Tasmania</collectingRegion>
, lacking domestic animals, fecal analysis indicates adult
<taxonomicName id="D0A04D1AFFE8E71F39D4FC831B38FCAB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[1240,1352,843,866]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F39D4FC831B38FCAB" box="[1240,1352,843,866]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
consume large prey, such as common wombats,
<taxonomicName id="D0A04D1AFFE8E71F39B0FCA31B0CFC4B" baseAuthorityName="Shaw" baseAuthorityYear="1800" box="[1212,1404,875,898]" class="Mammalia" family="Vombatidae" genus="Vombatus" kingdom="Animalia" order="Diprotodontia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="ursinus">
<emphasis id="25D4EA8BFFE8E71F39B0FCA31B0CFC4B" box="[1212,1404,875,898]" italics="true" pageId="7" pageNumber="8">Vombatus ursinus</emphasis>
</taxonomicName>
, Bennetts wallabies,
<taxonomicName id="D0A04D1AFFE8E71F3EB3FC431AD8FC6B" box="[959,1192,907,930]" class="Mammalia" family="Macropodidae" genus="Macropus" kingdom="Animalia" order="Diprotodontia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="rufigriseus">
<emphasis id="25D4EA8BFFE8E71F3EB3FC431AD8FC6B" box="[959,1192,907,930]" italics="true" pageId="7" pageNumber="8">Macropus rufigriseus</emphasis>
</taxonomicName>
, and Tasmanian pademelons,
<taxonomicName id="D0A04D1AFFE8E71F3E44FC631B2AFC0B" authority="(Jones and Barmuta 1998)" baseAuthorityName="Jones and Barmuta" baseAuthorityYear="1998" box="[840,1370,939,962]" class="Mammalia" family="Macropodidae" genus="Thylogale" kingdom="Animalia" order="Diprotodontia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="billardierii">
<emphasis id="25D4EA8BFFE8E71F3E44FC631A5FFC0B" box="[840,1071,939,962]" italics="true" pageId="7" pageNumber="8">Thylogale billardierii</emphasis>
(Jones and Barmuta 1998)
</taxonomicName>
, and rely less on smaller mammals and birds, the primary prey of subadults and juveniles. Diet overlap between adults and juveniles is less in summer, when adults shift toward the larger end of the prey spectrum, possibly because juveniles of their larger prey species are available then (Jones and Barmuta 1998). This dependence on large prey is reflected in stomach and fecal analyses from other parts of
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, but a wide range of foods is eaten, including birds, rabbits, and sheep (review—Pemberton et al. 2008). In a study of 3 coastal and 3 inland sites from the western onehalf of
<collectingRegion id="D564F87BFFE8E71F3E95FB231D8FFACB" box="[921,1023,1259,1282]" country="Australia" name="Tasmania" pageId="7" pageNumber="8">Tasmania</collectingRegion>
and using microscopic analysis of hairs from scats,
<bibRefCitation id="73314B68FFE8E71F3E86FAC31AF7FAEB" author="PEMBERTON, D. &amp; S. GALES &amp; B. BAUER &amp; R. GALES &amp; B. LAZENBY &amp; K. MEDLOCK" box="[906,1159,1291,1314]" pageId="7" pageNumber="8" pagination="13 - 21" refId="ref20687" refString="PEMBERTON, D., S. GALES, B. BAUER, R. GALES, B. LAZENBY, AND K. MEDLOCK. 2008. The diet of the Tasmanian devil, Sarcophilus harrisii, as determined from analysis of scat and stomach contents. Papers and Proceedings of the Royal Society of Tasmania 142: 13 - 21." type="journal article" year="2008">Pemberton et al. (2008)</bibRefCitation>
report a remarkably catholic diet: scats had remains of birds (50%), wallaby or pademelon (36%), common ringtail possum (
<taxonomicName id="D0A04D1AFFE8E71F396BFA831B0DFAAB" baseAuthorityName="Boddaert" baseAuthorityYear="1785" box="[1127,1405,1355,1378]" class="Mammalia" family="Pseudocheiridae" genus="Pseudocheirus" kingdom="Animalia" order="Diprotodontia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="peregrinus">
<emphasis id="25D4EA8BFFE8E71F396BFA831B0DFAAB" box="[1127,1405,1355,1378]" italics="true" pageId="7" pageNumber="8">Pseudocheirus peregrinus</emphasis>
</taxonomicName>
) or brushtailed possum (
<taxonomicName id="D0A04D1AFFE8E71F3EE5FAA31AA6FA4B" baseAuthorityName="Kerr" baseAuthorityYear="1792" box="[1001,1238,1387,1410]" class="Mammalia" family="Phalangeridae" genus="Trichosurus" kingdom="Animalia" order="Diprotodontia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="vulpecula">
<emphasis id="25D4EA8BFFE8E71F3EE5FAA31AA6FA4B" box="[1001,1238,1387,1410]" italics="true" pageId="7" pageNumber="8">Trichosurus vulpecula</emphasis>
</taxonomicName>
—30%). Both possums are semiarboreal. Four of 44 scats had remains of the Tasmanian monotremes: platypus (
<taxonomicName id="D0A04D1AFFE8E71F3941FA631B1AFA0B" baseAuthorityName="Shaw" baseAuthorityYear="1799" box="[1101,1386,1451,1474]" class="Mammalia" family="Ornithorhynchidae" genus="Ornithorhynchus" kingdom="Animalia" order="Monotremata" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="anatinus">
<emphasis id="25D4EA8BFFE8E71F3941FA631B1AFA0B" box="[1101,1386,1451,1474]" italics="true" pageId="7" pageNumber="8">Ornithorhynchus anatinus</emphasis>
</taxonomicName>
) or echidna (
<taxonomicName id="D0A04D1AFFE8E71F3E5DFA031A3BFA2B" baseAuthorityName="Shaw" baseAuthorityYear="1792" box="[849,1099,1483,1506]" class="Mammalia" family="Tachyglossidae" genus="Tachyglossus" kingdom="Animalia" order="Monotremata" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="aculeatus">
<emphasis id="25D4EA8BFFE8E71F3E5DFA031A3BFA2B" box="[849,1099,1483,1506]" italics="true" pageId="7" pageNumber="8">Tachyglossus aculeatus</emphasis>
</taxonomicName>
). Some of the identifiable bird remains were those of little penguin (
<taxonomicName id="D0A04D1AFFE8E71F3988FA231B49F9CB" baseAuthorityName="J.R.Forster" baseAuthorityYear="1781" box="[1156,1337,1515,1538]" class="Aves" family="Spheniscidae" genus="Eudyptula" kingdom="Animalia" order="Sphenisciformes" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="minor">
<emphasis id="25D4EA8BFFE8E71F3988FA231B49F9CB" box="[1156,1337,1515,1538]" italics="true" pageId="7" pageNumber="8">Eudyptula minor</emphasis>
</taxonomicName>
) and short-tailed shearwater (
<taxonomicName id="D0A04D1AFFE8E71F3EC2F9C31ADAF9EB" baseAuthorityName="Temminck" baseAuthorityYear="1836" box="[974,1194,1547,1570]" class="Aves" family="Procellariidae" genus="Puffinus" kingdom="Animalia" order="Procellariiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="tenuirostris">
<emphasis id="25D4EA8BFFE8E71F3EC2F9C31ADAF9EB" box="[974,1194,1547,1570]" italics="true" pageId="7" pageNumber="8">Puffinus tenuirostris</emphasis>
</taxonomicName>
), both of which nest in cavities in coastal dunes.
<taxonomicName id="D0A04D1AFFE8E71F3939F9E31AD2F98B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[1077,1186,1579,1602]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3939F9E31AD2F98B" box="[1077,1186,1579,1602]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
is the most carnivorous of the large dasyurids, consuming almost no plant material (Jones and Barmuta 1998).
</paragraph>
<paragraph id="171F3699FFE8E7103E7FF9431C28FEEB" blockId="7.[840,1523,203,1954]" lastBlockId="8.[109,793,203,1954]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
All foods have a high content of protein, water, vitamins, and minerals, variable amounts of fat, but with small amounts of carbohydrates. Thus,
<taxonomicName id="D0A04D1AFFE8E71F3921F9031AE8F92B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[1069,1176,1739,1762]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3921F9031AE8F92B" box="[1069,1176,1739,1762]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
obtains its hexoses largely from amino acids. With dentition and bite force to deal with every part of a carcass,
<taxonomicName id="D0A04D1AFFE8E71F3EDEF8C31A4DF8EB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[978,1085,1803,1826]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3EDEF8C31A4DF8EB" box="[978,1085,1803,1826]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
scavenges more than the syntopic quolls (
<bibRefCitation id="73314B68FFE8E71F3E5DF8E31D8FF88B" author="PEMBERTON, D." box="[849,1023,1835,1858]" pageId="7" pageNumber="8" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
;
<bibRefCitation id="73314B68FFE8E71F3901F8E31AF3F88B" author="JONES, M. E." box="[1037,1155,1835,1858]" pageId="7" pageNumber="8" refId="ref18123" refString="JONES, M. E. 1995. Guild structure of the large marsupial carnivores in Tasmania. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1995">Jones 1995</bibRefCitation>
). When placed in small enclosures with
<quantity id="D0589B7CFFE8E71F3E71F8831DB0F8AB" box="[893,960,1867,1890]" metricMagnitude="-1" metricUnit="kg" metricValue="5.0" pageId="7" pageNumber="8" unit="kg" value="0.5">0.5 kg</quantity>
brown rats,
<taxonomicName id="D0A04D1AFFE8E71F394EF8831B70F8AB" baseAuthorityName="Berkenhout" baseAuthorityYear="1769" box="[1090,1280,1867,1890]" class="Mammalia" family="Muridae" genus="Rattus" kingdom="Animalia" order="Rodentia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="norvegicus">
<emphasis id="25D4EA8BFFE8E71F394EF8831B70F8AB" box="[1090,1280,1867,1890]" italics="true" pageId="7" pageNumber="8">Rattus norvegicus</emphasis>
</taxonomicName>
,
<taxonomicName id="D0A04D1AFFE8E71F3800F8831B06F8AB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[1292,1398,1867,1890]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE8E71F3800F8831B06F8AB" box="[1292,1398,1867,1890]" italics="true" pageId="7" pageNumber="8">S. harrisii</emphasis>
</taxonomicName>
is a clumsy killer (
<bibRefCitation id="73314B68FFE8E71F3E9FF8A31A79F84B" author="EWER, R. F." box="[915,1033,1899,1922]" pageId="7" pageNumber="8" pagination="23 - 38" refId="ref15882" refString="EWER, R. F. 1969. Some observations on the killing and eating of prey by two dasyurid marsupials: the mulgara, Dasycercus cristicauda, and the Tasmanian devil, Sarcophilus harrisii. Zeitschrift fur Tierpsychologie 26: 23 - 38." type="journal article" year="1969">Ewer 1969</bibRefCitation>
; Buchmann and Guiler 1977). Nevertheless, incidental (but no direct) observations show that they can be successful predators of 8 or
<quantity id="D0589B7CFFE7E7103C95FF031FA5FF2B" box="[409,469,203,226]" metricMagnitude="1" metricUnit="kg" metricValue="3.0" pageId="8" pageNumber="9" unit="kg" value="30.0">30 kg</quantity>
macropods. Whether they can attack and kill a common wombat, adults of which are twice the mass of an adult male
<taxonomicName id="D0A04D1AFFE7E7103C51FEC31FB7FEEB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[349,455,267,290]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103C51FEC31FB7FEEB" box="[349,455,267,290]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
, is uncertain.
</paragraph>
<paragraph id="171F3699FFE7E7103D94FEE31C9AFD6B" blockId="8.[109,793,203,1954]" pageId="8" pageNumber="9">
<emphasis id="25D4EA8BFFE7E7103D94FEE31FDBFE8A" bold="true" box="[152,427,299,323]" italics="true" pageId="8" pageNumber="9">Diseases and parasites.</emphasis>
—An analysis of 100 years of disease and mortality for Australasian marsupials held at the London Zoo (Canfield and Cunningham 1993:165) revealed that “ill-defined dermatopathies” on the faces of
<taxonomicName id="D0A04D1AFFE7E7103F93FE431D61FE6B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[671,785,395,418]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103F93FE431D61FE6B" box="[671,785,395,418]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
, usually attributed to injuries related to fights, may have been “erosive squamous cell carcinomas.” The origin of these multiple proliferative lesions, previously reported for
<taxonomicName id="D0A04D1AFFE7E7103FABFE231D68FDCB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[679,792,491,514]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103FABFE231D68FDCB" box="[679,792,491,514]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
(
<bibRefCitation id="73314B68FFE7E7103D7AFDC31E89FDEB" author="GRINER, L. A." box="[118,249,523,546]" pageId="8" pageNumber="9" pagination="589 - 595" refId="ref16729" refString="GRINER, L. A. 1979. Neoplasms in Tasmanian devils. Journal of the National Cancer Institute 62: 589 - 595." type="journal article" year="1979">Griner 1979</bibRefCitation>
;
<bibRefCitation id="73314B68FFE7E7103C0BFDC31FA8FDEB" author="CANFIELD, P. J. &amp; W. J. HARTLEY &amp; G. L. REDDACLIFF" box="[263,472,523,546]" pageId="8" pageNumber="9" pagination="147 - 158" refId="ref15356" refString="CANFIELD, P. J., W. J. HARTLEY, AND G. L. REDDACLIFF. 1990. Spontaneous proliferations in Australian marsupials-a survey and review. 2. Dasyurids and bandicoots. Journal of Comparative Pathology 103: 147 - 158." type="journal article" year="1990">Canfield et al. 1990</bibRefCitation>
), may not be due to traumatic wounds but to a genetic predisposition for tumor development in
<taxonomicName id="D0A04D1AFFE7E7103D86FD831E88FDAB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[138,248,587,610]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103D86FD831E88FDAB" box="[138,248,587,610]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
, as suggested by Griner. That work was prescient in light of the appearance in the 1990s of a facial cancer transmitted by bites (see details in the “Conservation” section).
</paragraph>
<paragraph id="171F3699FFE7E7103D94FD631F23FBCB" blockId="8.[109,793,203,1954]" pageId="8" pageNumber="9">
<taxonomicName id="D0A04D1AFFE7E7103D94FD631FDFFD0B" authorityName="Garkavi" authorityYear="1972" box="[152,431,683,706]" class="Adenophorea" family="Trichinellidae" genus="Trichinella" kingdom="Animalia" order="Trichocephalida" pageId="8" pageNumber="9" phylum="Nematoda" rank="species" species="pseudospiralis">
<emphasis id="25D4EA8BFFE7E7103D94FD631FDFFD0B" box="[152,431,683,706]" italics="true" pageId="8" pageNumber="9">Trichinella pseudospiralis</emphasis>
</taxonomicName>
, a nematode whose larvae encyst in muscle tissue, was detected in
<taxonomicName id="D0A04D1AFFE7E7103CD6FD031C36FD2B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[474,582,715,738]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103CD6FD031C36FD2B" box="[474,582,715,738]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
in 1988 (
<bibRefCitation id="73314B68FFE7E7103FBEFD031E92FCCB" author="OBENDORF, D. L." pageId="8" pageNumber="9" pagination="108 - 110" refId="ref20257" refString="OBENDORF, D. L., ET AL. 1990. Trichinella pseudospiralis infection in Tasmanian wildlife. Australian Veterinary Journal 67: 108 - 110." type="journal article" year="1990">Obendorf et al. 1990</bibRefCitation>
). A survey of 9 locations across
<collectingRegion id="D564F87BFFE7E7103F5CFD231CC6FCCB" box="[592,694,747,770]" country="Australia" name="Tasmania" pageId="8" pageNumber="9">Tasmania</collectingRegion>
revealed infected
<taxonomicName id="D0A04D1AFFE7E7103DC6FCC31F45FCEB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[202,309,779,802]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103DC6FCC31F45FCEB" box="[202,309,779,802]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
at all sites, and an overall prevalence of 30% in 153 animals. At least 3 other Tasmanian mammals eaten by
<taxonomicName id="D0A04D1AFFE7E7103D61FC831EAFFCAB" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[109,223,843,866]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103D61FC831EAFFCAB" box="[109,223,843,866]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
harbor this parasite, and cannibalism likely contributes to maintenance of the parasite in wildlife populations. When fed muscle from infected
<taxonomicName id="D0A04D1AFFE7E7103CCBFC431C42FC6B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[455,562,907,930]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103CCBFC431C42FC6B" box="[455,562,907,930]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
, laboratory rats, cats, 2 native raptorial birds, brush-tailed possum, eastern quoll, and other
<taxonomicName id="D0A04D1AFFE7E7103DA1FC031F69FC2B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[173,281,971,994]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103DA1FC031F69FC2B" box="[173,281,971,994]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
all became infected with
<taxonomicName id="D0A04D1AFFE7E7103F38FC031C9BFC2B" authorityName="Garkavi" authorityYear="1972" box="[564,747,971,994]" class="Adenophorea" family="Trichinellidae" genus="Trichinella" kingdom="Animalia" order="Trichocephalida" pageId="8" pageNumber="9" phylum="Nematoda" rank="species" species="pseudospiralis">
<emphasis id="25D4EA8BFFE7E7103F38FC031C9BFC2B" box="[564,747,971,994]" italics="true" pageId="8" pageNumber="9">T. pseudospiralis</emphasis>
</taxonomicName>
larvae (
<bibRefCitation id="73314B68FFE7E7103DADFC231F33FBCB" author="OBENDORF, D. L." box="[161,323,1003,1026]" pageId="8" pageNumber="9" pagination="266 - 268" refId="ref20221" refString="OBENDORF, D. L. 1993. Experimental infections with the Tasmanian isolate of Trichinella pseudospiralis using a non-enzymatic recovery technique. Journal of the Helminthological Society of Washington 60: 266 - 268." type="journal article" year="1993">Obendorf 1993</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE7E7103D94FBC31F59FA2B" blockId="8.[109,793,203,1954]" pageId="8" pageNumber="9">
The cestodes
<taxonomicName id="D0A04D1AFFE7E7103C3FFBC31C52FBEB" box="[307,546,1035,1058]" class="Cestoda" family="Taeniidae" genus="Anoplotaenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyclophyllidea" pageId="8" pageNumber="9" phylum="Platyhelminthes" rank="species" species="dasyure">
<emphasis id="25D4EA8BFFE7E7103C3FFBC31C52FBEB" box="[307,546,1035,1058]" italics="true" pageId="8" pageNumber="9">Anoplotaenia dasyure</emphasis>
</taxonomicName>
, up to 15,000, in the small intestine (
<bibRefCitation id="73314B68FFE7E7103C14FBE31F97FB8B" author="GREGORY, G. G. &amp; B. L. MUNDAY &amp; I. BEVERIDGE &amp; M. D. RICKARD" box="[280,487,1067,1090]" pageId="8" pageNumber="9" pagination="187 - 191" refId="ref16591" refString="GREGORY, G. G., B. L. MUNDAY, I. BEVERIDGE, AND M. D. RICKARD. 1975. Studies on Anoplotaenia dasyuri Beddard, 1911 (Cestoda: Taeniidae), a parasite of the Tasmanian devil: life cycle and epidemiology. International Journal of Parasitology 5: 187 - 191." type="journal article" year="1975">Gregory et al. 1975</bibRefCitation>
) and
<taxonomicName id="D0A04D1AFFE7E7103F28FBE31F5BFBAB" authority="(Beveridge 1984)" baseAuthorityName="Beveridge" baseAuthorityYear="1984" class="Cestoda" family="Dilepididae" genus="Dasyurotaenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyclophyllidea" pageId="8" pageNumber="9" phylum="Platyhelminthes" rank="species" species="robusta">
<emphasis id="25D4EA8BFFE7E7103F28FBE31D68FB8B" box="[548,792,1067,1090]" italics="true" pageId="8" pageNumber="9">Dasyurotaenia robusta</emphasis>
(
<bibRefCitation id="73314B68FFE7E7103D7AFB831F51FBAB" author="BEVERIDGE, I." box="[118,289,1099,1122]" pageId="8" pageNumber="9" pagination="185 - 195" refId="ref14943" refString="BEVERIDGE, I. 1984. Dasyurotaenia robusta Beddard, 1912 and D. dasyuri sp. nov. from carnivorous Australian marsupials. Transactions of the Royal Society of South Australia 108: 185 - 195." type="journal article" year="1984">Beveridge 1984</bibRefCitation>
)
</taxonomicName>
, and a stomach nematode,
<taxonomicName id="D0A04D1AFFE7E7103F5BFB831F88FB4B" authority="(Beveridge and Spratt 2003)" baseAuthorityName="Beveridge and Spratt" baseAuthorityYear="2003" class="Secernentea" family="Physalopteridae" genus="Physaloptera" kingdom="Animalia" order="Spirurida" pageId="8" pageNumber="9" phylum="Nematoda" rank="species" species="sarcophili">
<emphasis id="25D4EA8BFFE7E7103F5BFB831EC7FB4B" italics="true" pageId="8" pageNumber="9">Physaloptera sarcophili</emphasis>
(Beveridge and Spratt 2003)
</taxonomicName>
, all infect
<taxonomicName id="D0A04D1AFFE7E7103F7DFBA31CAFFB4B" baseAuthorityName="Baudinette" baseAuthorityYear="1982" box="[625,735,1131,1154]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103F7DFBA31CAFFB4B" box="[625,735,1131,1154]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
. The filarioid nematode,
<taxonomicName id="D0A04D1AFFE7E7103C4BFB431C18FB6B" box="[327,616,1163,1186]" class="Chromadorea" family="Onchocercidae" genus="Cercopithifilaria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Spirurida" pageId="8" pageNumber="9" phylum="Nematoda" rank="species" species="johnstoni">
<emphasis id="25D4EA8BFFE7E7103C4BFB431C18FB6B" box="[327,616,1163,1186]" italics="true" pageId="8" pageNumber="9">Cercopithifilaria johnstoni</emphasis>
</taxonomicName>
, transmitted by the tick
<taxonomicName id="D0A04D1AFFE7E7103DC7FB631FF4FB0B" box="[203,388,1195,1218]" class="Arachnida" family="Ixodidae" genus="Ixodes" kingdom="Animalia" order="Ixodida" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="trichosuri">
<emphasis id="25D4EA8BFFE7E7103DC7FB631FF4FB0B" box="[203,388,1195,1218]" italics="true" pageId="8" pageNumber="9">Ixodes trichosuri</emphasis>
</taxonomicName>
, sometimes is present in the subcutis of
<taxonomicName id="D0A04D1AFFE7E7103DA2FB031C3BFB2B" authority="(Spratt and Haycock 1988)" baseAuthorityName="Spratt and Haycock" baseAuthorityYear="1988" box="[174,587,1227,1250]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103DA2FB031F6AFB2B" box="[174,282,1227,1250]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
(Spratt and Haycock 1988)
</taxonomicName>
.
<taxonomicName id="D0A04D1AFFE7E7103F54FB031EA8FACB" authorityName="Sprent" authorityYear="1970" class="Chromadorea" family="Ascarididae" genus="Baylisascaris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rhabditida" pageId="8" pageNumber="9" phylum="Nematoda" rank="species" species="tasmaniensis">
<emphasis id="25D4EA8BFFE7E7103F54FB031EA8FACB" italics="true" pageId="8" pageNumber="9">Baylisascaris tasmaniensis</emphasis>
</taxonomicName>
is an ascaridoid nematode sometimes found in the intestines of
<taxonomicName id="D0A04D1AFFE7E7103C0CFAC31C67FAEB" authority="(Sprent 1970)" baseAuthorityName="Sprent" baseAuthorityYear="1970" box="[256,535,1291,1314]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103C0CFAC31F00FAEB" box="[256,368,1291,1314]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
(
<bibRefCitation id="73314B68FFE7E7103C89FAC31C7BFAEB" author="SPRENT, J. F. A." box="[389,523,1291,1314]" pageId="8" pageNumber="9" pagination="75 - 86" refId="ref21396" refString="SPRENT, J. F. A. 1970. Baylisascaris tasmaniensis sp. nov. in marsupial carnivores. Parasitology 61: 75 - 86." type="journal article" year="1970">Sprent 1970</bibRefCitation>
)
</taxonomicName>
. A new genus of mite that produced mange in a
<taxonomicName id="D0A04D1AFFE7E7103C98FAE31C73FA8B" baseAuthorityName="Sprent" baseAuthorityYear="1970" box="[404,515,1323,1346]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103C98FAE31C73FA8B" box="[404,515,1323,1346]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
held in the London Zoo was reported by Fain and Laurance (1975) and the following 4 species of mites of 3 families were taken from
<taxonomicName id="D0A04D1AFFE7E7103F81FAA31C8AFA4B" baseAuthorityName="Sprent" baseAuthorityYear="1970" box="[653,762,1387,1410]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103F81FAA31C8AFA4B" box="[653,762,1387,1410]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
in
<collectingRegion id="D564F87BFFE7E7103D61FA431EA7FA6B" box="[109,215,1419,1442]" country="Australia" name="Tasmania" pageId="8" pageNumber="9">Tasmania</collectingRegion>
:
<taxonomicName id="D0A04D1AFFE7E7103DECFA431F86FA6B" box="[224,502,1419,1442]" class="Arachnida" family="Sarcoptidae" genus="Diabolicoptes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astigmata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="sarcophilus">
<emphasis id="25D4EA8BFFE7E7103DECFA431F86FA6B" box="[224,502,1419,1442]" italics="true" pageId="8" pageNumber="9">Diabolicoptes sarcophilus</emphasis>
</taxonomicName>
,
<taxonomicName id="D0A04D1AFFE7E7103F0FFA431D62FA6B" box="[515,786,1419,1442]" class="Arachnida" family="Laelapidae" genus="Haemolaelaps" higherTaxonomySource="GBIF" kingdom="Animalia" order="Mesostigmata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="flagellatus">
<emphasis id="25D4EA8BFFE7E7103F0FFA431D62FA6B" box="[515,786,1419,1442]" italics="true" pageId="8" pageNumber="9">Haemolaelaps flagellatus</emphasis>
</taxonomicName>
,
<taxonomicName id="D0A04D1AFFE7E7103D61FA631C76FA0B" box="[109,518,1451,1474]" class="Arachnida" family="Macronyssidae" genus="Ornithonyssus" kingdom="Animalia" order="Mesostigmata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="dasyuri" subGenus="Trichonyssus">
<emphasis id="25D4EA8BFFE7E7103D61FA631F77FA0B" box="[109,263,1451,1474]" italics="true" pageId="8" pageNumber="9">Ornithonyssus</emphasis>
(
<emphasis id="25D4EA8BFFE7E7103C15FA631FD4FA0B" box="[281,420,1451,1474]" italics="true" pageId="8" pageNumber="9">Trichonyssus</emphasis>
)
<emphasis id="25D4EA8BFFE7E7103CBAFA631C76FA0B" box="[438,518,1451,1474]" italics="true" pageId="8" pageNumber="9">dasyuri</emphasis>
</taxonomicName>
, and
<taxonomicName id="D0A04D1AFFE7E7103F4AFA631F54FA2B" authority="(Green 1989)" baseAuthorityName="Green" baseAuthorityYear="1989" class="Arachnida" family="Sarcoptidae" genus="Satanicoptes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Astigmata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="armatus">
<emphasis id="25D4EA8BFFE7E7103F4AFA631EFCFA2B" italics="true" pageId="8" pageNumber="9">Satanicoptes armatus</emphasis>
(
<bibRefCitation id="73314B68FFE7E7103D90FA031F69FA2B" author="GREEN, R. H." box="[156,281,1483,1506]" pageId="8" pageNumber="9" pagination="1 - 25" refId="ref16412" refString="GREEN, R. H. 1989. The ectoparasitic mites of Tasmanian vertebrate animals. Records of the Queen Victoria Museum 98: 1 - 25." type="journal article" year="1989">Green 1989</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="171F3699FFE7E7103D94FA231EC3F86B" blockId="8.[109,793,203,1954]" pageId="8" pageNumber="9">
<emphasis id="25D4EA8BFFE7E7103D94FA231FC6F9CA" bold="true" box="[152,438,1515,1539]" italics="true" pageId="8" pageNumber="9">Interspecific interactions.</emphasis>
—The 3 largest (&gt;
<quantity id="D0589B7CFFE7E7103F80FA231CBCF9CB" box="[652,716,1515,1538]" metricMagnitude="-1" metricUnit="kg" metricValue="5.0" pageId="8" pageNumber="9" unit="g" value="500.0">500 g</quantity>
) dasyurid carnivores,
<taxonomicName id="D0A04D1AFFE7E7103C2EF9C31F8CF9EB" baseAuthorityName="Sprent" baseAuthorityYear="1970" box="[290,508,1547,1570]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103C2EF9C31F8CF9EB" box="[290,508,1547,1570]" italics="true" pageId="8" pageNumber="9">Sarcophilus harrisii</emphasis>
</taxonomicName>
, spotted-tailed quoll, and eastern quoll, are sympatric only in
<collectingRegion id="D564F87BFFE7E7103F0EF9E31C1CF98B" box="[514,620,1579,1602]" country="Australia" name="Tasmania" pageId="8" pageNumber="9">Tasmania</collectingRegion>
. As the largest,
<taxonomicName id="D0A04D1AFFE7E7103D61F9831EADF9AB" baseAuthorityName="Sprent" baseAuthorityYear="1970" box="[109,221,1611,1634]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103D61F9831EADF9AB" box="[109,221,1611,1634]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
dominates at a carcass, although an adult spottedtailed quoll can chase off a young
<taxonomicName id="D0A04D1AFFE7E7103F08F9A31D63F94B" authority="(Jones 1995)" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[516,787,1643,1666]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103F08F9A31C05F94B" box="[516,629,1643,1666]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
(
<bibRefCitation id="73314B68FFE7E7103F87F9A31D78F94B" author="JONES, M. E." box="[651,776,1643,1666]" pageId="8" pageNumber="9" refId="ref18123" refString="JONES, M. E. 1995. Guild structure of the large marsupial carnivores in Tasmania. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1995">Jones 1995</bibRefCitation>
)
</taxonomicName>
.
<taxonomicName id="D0A04D1AFFE7E7103D61F9431EAFF96B" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[109,223,1675,1698]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103D61F9431EAFF96B" box="[109,223,1675,1698]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
overlaps in diet with spotted-tailed quolls, which in turn overlaps with and dominates eastern quolls (Jones and Barmuta 1998). Morphological patterning in trophic structures (canine tooth strength and masseter muscle strength) among the species and sexes of quolls in
<collectingRegion id="D564F87BFFE7E7103CB0F8C31C55F8EB" box="[444,549,1803,1826]" country="Australia" name="Tasmania" pageId="8" pageNumber="9">Tasmania</collectingRegion>
, and character release among the same quolls on the adjacent Australian mainland where
<taxonomicName id="D0A04D1AFFE7E7103DB7F8831F5CF8AB" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[187,300,1867,1890]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103DB7F8831F5CF8AB" box="[187,300,1867,1890]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
is absent, constitute evidence for competition among these carnivores at an evolutionary timescale (Jones 1997).
</paragraph>
<paragraph id="171F3699FFE7E7103E7FFF031A2FFC6B" blockId="8.[840,1523,203,930]" pageId="8" pageNumber="9">
The role of
<taxonomicName id="D0A04D1AFFE7E7103EF8FF031A11FF2B" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[1012,1121,203,226]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103EF8FF031A11FF2B" box="[1012,1121,203,226]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
in the community structure of carnivores is complex (
<bibRefCitation id="73314B68FFE7E710391DFF231A9EFECB" author="DICKMAN, C. R. &amp; GLEN, M. E. &amp; JONES, M. E. &amp; E. G. RICHIE &amp; A. D. WALLACH" box="[1041,1262,235,258]" pageId="8" pageNumber="9" refId="ref15639" refString="DICKMAN, C. R., A. S. GLEN, M. E. JONES, M. E. SOULe, E. G. RICHIE, AND A. D. WALLACH. 2014. Strongly interactive carnivore species: maintaining and restoring ecosystem function. Pp. 301 - 322 in Carnivores of Australia: past, present, and future (A. S. Glen and C. R. Dickman, eds.). CSIRO Publishing, Collingwood, Victoria, Australia." type="book" year="2014">Dickman et al. 2014</bibRefCitation>
). For example, spottedtailed quolls adjust their activity to avoid
<taxonomicName id="D0A04D1AFFE7E7103817FEC31BFAFEEB" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[1307,1418,267,290]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103817FEC31BFAFEEB" box="[1307,1418,267,290]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
where
<taxonomicName id="D0A04D1AFFE7E71038D3FEC31DE9FE8B" baseAuthorityName="Jones" baseAuthorityYear="1995" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E71038D3FEC31B9CFEEB" box="[1503,1516,267,290]" italics="true" pageId="8" pageNumber="9">S</emphasis>
.
<emphasis id="25D4EA8BFFE7E7103E44FEE31DE9FE8B" box="[840,921,299,322]" italics="true" pageId="8" pageNumber="9">harrisii</emphasis>
</taxonomicName>
is at high density (
<bibRefCitation id="73314B68FFE7E710397EFEE31B7EFE8B" author="HOLLINGS, T. A." box="[1138,1294,299,322]" pageId="8" pageNumber="9" refId="ref17655" refString="HOLLINGS, T. A. 2013. Ecological effects of disease-induced apex predator decline. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="2013">Hollings 2013</bibRefCitation>
). Further,
<taxonomicName id="D0A04D1AFFE7E7103889FEE31B83FE8B" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[1413,1523,299,322]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103889FEE31B83FE8B" box="[1413,1523,299,322]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
seemingly suppresses the behavior and probably the abundance of feral cats (
<taxonomicName id="D0A04D1AFFE7E7103EEDFEA31A27FE4B" authorityName="Linnaeus" authorityYear="1758" box="[993,1111,363,386]" class="Mammalia" family="Felidae" genus="Felis" kingdom="Animalia" order="Carnivora" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="catus">
<emphasis id="25D4EA8BFFE7E7103EEDFEA31A27FE4B" box="[993,1111,363,386]" italics="true" pageId="8" pageNumber="9">Felis catus</emphasis>
</taxonomicName>
), with increased detections observed in areas where the number of
<taxonomicName id="D0A04D1AFFE7E7103980FE431A89FE6B" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[1164,1273,395,418]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103980FE431A89FE6B" box="[1164,1273,395,418]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
has declined from disease; 2 studies using different methods support this conjecture (
<bibRefCitation id="73314B68FFE7E7103E5DFE031A59FE2B" author="HOLLINGS, T. &amp; M. JONES &amp; N. MOONEY &amp; H. MCCALLUM" box="[849,1065,459,482]" pageId="8" pageNumber="9" pagination="63 - 75" refId="ref17685" refString="HOLLINGS, T., M. JONES, N. MOONEY, AND H. MCCALLUM. 2014. Trophic cascades following the disease-induced decline of an apex predator, the Tasmanian devil. Conservation Biology 28: 63 - 75." type="journal article" year="2014">Hollings et al. 2014</bibRefCitation>
;
<bibRefCitation id="73314B68FFE7E7103934FE031B61FE2B" author="HOLLINGS, T. A. &amp; ONES, N. &amp; MOONEY, AND &amp; H. I. MCCALLUM" box="[1080,1297,459,482]" pageId="8" pageNumber="9" refId="ref17728" refString="HOLLINGS, T. A., M. J ONES, N. MOONEY, AND H. I. MCCALLUM. 2015. Diseaseinduced decline of an apex predator drives invasive dominated states and threatens biodiversity. Ecology Online." type="book" year="2015">Hollings et al. 2015</bibRefCitation>
). Even when
<taxonomicName id="D0A04D1AFFE7E71038A5FE031D04FDCB" baseAuthorityName="Jones" baseAuthorityYear="1995" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E71038A5FE031D04FDCB" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
is at low densities, cats still avoid them, both spatially (Lazenby and Dickman 2013) and temporally (
<bibRefCitation id="73314B68FFE7E710385FFDC31DFAFD8B" author="FANCOURT, B. A. &amp; C. E. HAWKINS &amp; E. Z. CAMERON &amp; M. E. JONES &amp; S. C. NICOL" pageId="8" pageNumber="9" pagination="119303" refId="ref15984" refString="FANCOURT, B. A., C. E. HAWKINS, E. Z. CAMERON, M. E. JONES, AND S. C. NICOL. 2015 a. Devil declines and catastrophic cascades: is mesopredator release of feral cats inhibiting recovery of the eastern quoll? PLoS ONE 10: e 119303." type="journal article" year="2015">Fancourt et al. 2015a</bibRefCitation>
), indicating their strong suppressive influence. Also,
<taxonomicName id="D0A04D1AFFE7E7103E44FD831DC3FDAB" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[840,947,587,610]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103E44FD831DC3FDAB" box="[840,947,587,610]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
may protect smaller carnivores such as eastern quolls through controlling cat numbers. Although severe weather may have contributed to the recent decline in eastern quoll populations (
<bibRefCitation id="73314B68FFE7E7103EAFFD631AF2FD0B" author="FANCOURT, B. A. &amp; C. E. HAWKINS &amp; S. C. NICOL" box="[931,1154,683,706]" pageId="8" pageNumber="9" pagination="195 - 205" refId="ref16045" refString="FANCOURT, B. A., C. E. HAWKINS, AND S. C. NICOL. 2013. Evidence of rapid decline of the eastern quoll (Dasyurus viverrinus) in Tasmania. Australian Mammalogy 35: 195 - 205." type="journal article" year="2013">Fancourt et al. 2013</bibRefCitation>
;
<bibRefCitation id="73314B68FFE7E710399FFD631BF0FD0B" author="FANCOURT, B. A." box="[1171,1408,683,706]" pageId="8" pageNumber="9" pagination="0129420" refId="ref16090" refString="FANCOURT, B. A., ET AL. 2015 b. Testing the role of climate change in species decline: is the eastern quoll a victim of a change in the weather. PLoS ONE 10: e 0129420." type="journal article" year="2015">Fancourt et al. 2015b</bibRefCitation>
), high cat densities now threaten the species (
<bibRefCitation id="73314B68FFE7E71039C5FD031BD0FD2B" author="HOLLINGS, T. A. &amp; ONES, N. &amp; MOONEY, AND &amp; H. I. MCCALLUM" box="[1225,1440,715,738]" pageId="8" pageNumber="9" refId="ref17728" refString="HOLLINGS, T. A., M. J ONES, N. MOONEY, AND H. I. MCCALLUM. 2015. Diseaseinduced decline of an apex predator drives invasive dominated states and threatens biodiversity. Ecology Online." type="book" year="2015">Hollings et al. 2015</bibRefCitation>
), a further indication of the complex role of
<taxonomicName id="D0A04D1AFFE7E71039FFFD231B12FCCB" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[1267,1378,747,770]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E71039FFFD231B12FCCB" box="[1267,1378,747,770]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
in the carnivore community in
<collectingRegion id="D564F87BFFE7E7103916FCC31AF4FCEB" box="[1050,1156,779,802]" country="Australia" name="Tasmania" pageId="8" pageNumber="9">Tasmania</collectingRegion>
. The disease-induced reduction in
<taxonomicName id="D0A04D1AFFE7E7103E44FCE31DC6FC8B" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[840,950,811,834]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103E44FCE31DC6FC8B" box="[840,950,811,834]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
densities likely is triggering a complex trophic cascade, including an increase in invasive rodents and a decline in native small mammals in areas where cat detections are increasing (
<bibRefCitation id="73314B68FFE7E7103E76FC431A3FFC6B" author="HOLLINGS, T. A. &amp; ONES, N. &amp; MOONEY, AND &amp; H. I. MCCALLUM" box="[890,1103,907,930]" pageId="8" pageNumber="9" refId="ref17728" refString="HOLLINGS, T. A., M. J ONES, N. MOONEY, AND H. I. MCCALLUM. 2015. Diseaseinduced decline of an apex predator drives invasive dominated states and threatens biodiversity. Ecology Online." type="book" year="2015">Hollings et al. 2015</bibRefCitation>
).
</paragraph>
</subSubSection>
</subSection>
<subSection id="7E2F2F72FFE7E711394DFC2E1CA5FECB" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9" type="multiple">
<subSubSection id="5FBA6512FFE7E711394DFC2E1CA5FECB" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9" type="description">
<paragraph id="171F3699FFE7E710394DFC2E1A89FBC9" blockId="8.[1089,1273,998,1024]" box="[1089,1273,998,1024]" pageId="8" pageNumber="9">
<heading id="4C5781F5FFE7E710394DFC2E1A89FBC9" allCaps="true" bold="true" box="[1089,1273,998,1024]" centered="true" fontSize="11" level="1" pageId="8" pageNumber="9" reason="6">
<emphasis id="25D4EA8BFFE7E710394DFC2E1A89FBC9" bold="true" box="[1089,1273,998,1024]" pageId="8" pageNumber="9">HUSBANDRY</emphasis>
</heading>
</paragraph>
<paragraph id="171F3699FFE7E7103E7FFBE21B2CFA28" blockId="8.[840,1524,1066,1953]" pageId="8" pageNumber="9">
<taxonomicName id="D0A04D1AFFE7E7103E7FFBE21A20FB88" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[883,1104,1066,1089]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103E7FFBE21A20FB88" box="[883,1104,1066,1089]" italics="true" pageId="8" pageNumber="9">Sarcophilus harrisii</emphasis>
</taxonomicName>
has been reared successfully for a hundred years in
<collectingRegion id="D564F87BFFE7E7103906FB821A02FBA8" box="[1034,1138,1098,1121]" country="Australia" name="Tasmania" pageId="8" pageNumber="9">Tasmania</collectingRegion>
, initially using methods developed by Mary Roberts of Tasmanias Beaumarais Zoo. In response to the threat of what is now called Devil Facial Tumor Disease (DFTD), quarantine protocols and breeding programs incorporating genetic variability have been developed to ensure substantial captive “insurance” populations. The pooled experience of zoo and wildlife park cooperators and of wildlife carers has led to standard guidelines, under the Australian Zoo and Aquarium Association, for use in the Tasmanian governments Save the Tasmanian Devil Program (Hogg and Hockley 2013). In 2015, about 700
<taxonomicName id="D0A04D1AFFE7E7103EB4FA421A53FA68" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[952,1059,1418,1441]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103EB4FA421A53FA68" box="[952,1059,1418,1441]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
were held in captivity, including more than
<quantity id="D0589B7CFFE7E7103E44FA621DFDFA08" box="[840,909,1450,1473]" metricMagnitude="1" metricUnit="m" metricValue="1.27" pageId="8" pageNumber="9" unit="in" value="500.0">500 in</quantity>
40 institutions in
<collectingCountry id="6FB77609FFE7E710395CFA621AC5FA08" box="[1104,1205,1450,1473]" name="Australia" pageId="8" pageNumber="9">Australia</collectingCountry>
; others were in 3 locations in
<collectingCountry id="6FB77609FFE7E7103E44FA021DA6FA28" box="[840,982,1482,1505]" name="New Zealand" pageId="8" pageNumber="9">New Zealand</collectingCountry>
and in 6 places in the
<collectingCountry id="6FB77609FFE7E71039C9FA021B28FA28" box="[1221,1368,1482,1505]" name="United States of America" pageId="8" pageNumber="9">United States</collectingCountry>
.
</paragraph>
<paragraph id="171F3699FFE7E7113E7FFA221CA5FECB" blockId="8.[840,1524,1066,1953]" lastBlockId="9.[109,792,203,258]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
Facilities to house
<taxonomicName id="D0A04D1AFFE7E7103948FA221AC2F9C8" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[1092,1202,1514,1537]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103948FA221AC2F9C8" box="[1092,1202,1514,1537]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
under intensive management must be kept separated from other captive wildlife, have sufficient area for several animals, and be enclosed with non-climbable fencing. Environmental enrichment is vital, and includes hidden food, scats from others, natural vegetation including hollow logs, and substrate for digging. Less intensive facilities, known as “Free Range Enclosures,” in which other wildlife can mix with
<taxonomicName id="D0A04D1AFFE7E7103EBDF9021A6EF928" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[945,1054,1738,1761]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E7103EBDF9021A6EF928" box="[945,1054,1738,1761]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
, also are permitted. Socialization of young animals is important both for developing competitive feeding behaviors at carcasses and for mate selection. Diet is almost entirely whole animal food including fur, feather, and bone, and occasionally a large carcass with offal and gut contents is presented. Water is provided for drinking, playing, bathing and, as needed, cooling; indeed,
<taxonomicName id="D0A04D1AFFE7E710398AF8421A86F868" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[1158,1270,1930,1953]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE7E710398AF8421A86F868" box="[1158,1270,1930,1953]" italics="true" pageId="8" pageNumber="9">S. harrisii</emphasis>
</taxonomicName>
enjoys shallow water. The diet of captive females is more closely adjusted in preparation for the breeding season in February and early March.
</paragraph>
</subSubSection>
</subSection>
<subSection id="7E2F2F72FFE6E7123C79FEF51A8CFCCB" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10" type="multiple">
<subSubSection id="5FBA6512FFE6E7123C79FEF51A8CFCCB" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10" type="biology_ecology">
<paragraph id="171F3699FFE6E7113C79FEF51C60FE9E" blockId="9.[373,528,317,343]" box="[373,528,317,343]" pageId="9" pageNumber="10">
<heading id="4C5781F5FFE6E7113C79FEF51C60FE9E" allCaps="true" bold="true" box="[373,528,317,343]" centered="true" fontSize="11" level="1" pageId="9" pageNumber="10" reason="6">
<emphasis id="25D4EA8BFFE6E7113C79FEF51C60FE9E" bold="true" box="[373,528,317,343]" pageId="9" pageNumber="10">BEHAVIOR</emphasis>
</heading>
</paragraph>
<paragraph id="171F3699FFE6E7113D94FE491CE7FE31" blockId="9.[109,793,385,1944]" pageId="9" pageNumber="10">
The behavior of
<taxonomicName id="D0A04D1AFFE6E7113C5BFE491C41FE51" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[343,561,385,408]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113C5BFE491C41FE51" box="[343,561,385,408]" italics="true" pageId="9" pageNumber="10">Sarcophilus harrisii</emphasis>
</taxonomicName>
has been studied as much as that of any marsupial (
<bibRefCitation id="73314B68FFE6E7113CBEFE691C54FE71" author="CROFT, D. B." box="[434,548,417,440]" pageId="9" pageNumber="10" refId="ref15459" refString="CROFT, D. B. 1982. Communication in the Dasyuridae (Marsupialia): a review. Pp. 291 - 309 in Carnivorous marsupials (M. Archer, ed.). Surrey Beatty &amp; Sons Pty Limited, Chipping Norton, New South Wales, Australia." type="book" year="1982">Croft 1982</bibRefCitation>
). In the wild, the major studies of behavior are those of
<bibRefCitation id="73314B68FFE6E7113CDDFE091CE5FE11" author="PEMBERTON, D." box="[465,661,449,472]" pageId="9" pageNumber="10" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton (1990)</bibRefCitation>
, Pemberton and Renouf (1993), and Jones (
<bibRefCitation id="73314B68FFE6E7113CB0FE291CF7FE31" author="LACHISH, S. &amp; M. JONES &amp; H. MCCALLUM" box="[444,647,481,504]" pageId="9" pageNumber="10" pagination="926 - 936" refId="ref19156" refString="LACHISH, S., M. JONES, AND H. MCCALLUM. 2007. The impact of disease on the survival and population growth rate of the Tasmanian devil. Journal of Animal Ecology 76: 926 - 936." type="journal article" year="2007">Lachish et al. 2007</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE6E7113D94FDC91C1FFD11" blockId="9.[109,793,385,1944]" pageId="9" pageNumber="10">
<taxonomicName id="D0A04D1AFFE6E7113D94FDC91F1DFDD1" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[152,365,513,536]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113D94FDC91F1DFDD1" box="[152,365,513,536]" italics="true" pageId="9" pageNumber="10">Sarcophilus harrisii</emphasis>
</taxonomicName>
spends the day in one of its well-hidden dens (
<emphasis id="25D4EA8BFFE6E7113DBCFDD51EB4FDFB" box="[176,196,541,562]" italics="true" pageId="9" pageNumber="10">X</emphasis>
= 3.8 dens per adult—Pemberton 1990), often located in a wombat burrow or on the surface under a tussock. When the young become too large to be carried in the pouch, they are deposited in a den for a period of weeks (
<bibRefCitation id="73314B68FFE6E7113F3AFD491C97FD51" author="PEMBERTON, D." box="[566,743,641,664]" pageId="9" pageNumber="10" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
). At least in captivity, females build a nest of dry vegetation in the dens, chewing and digging to create a thick bed.
</paragraph>
<paragraph id="171F3699FFE6E7113D94FD291D7DFB71" blockId="9.[109,793,385,1944]" pageId="9" pageNumber="10">
A tireless runner,
<taxonomicName id="D0A04D1AFFE6E7113C5EFD291FCCFD31" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[338,444,737,760]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113C5EFD291FCCFD31" box="[338,444,737,760]" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
often runs a predictable transect, up to
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, to places where food was found in a recent night (Pemberton and Renouf 1993), the long-distance gait being a characteristic 1-1-2 pattern of right hindleft foreleft hind and right forefoot together has been observed by one of us (NJM). An average
<taxonomicName id="D0A04D1AFFE6E7113DFEFC491F2FFC51" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[242,351,897,920]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113DFEFC491F2FFC51" box="[242,351,897,920]" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
spends 7.7 h traveling
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per night while hunting (
<bibRefCitation id="73314B68FFE6E7113C1DFC691FB0FC71" author="PEMBERTON, D." box="[273,448,929,952]" pageId="9" pageNumber="10" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
). Animals from an alpine population (
<bibRefCitation id="73314B68FFE6E7113DB0FC091F03FC11" author="JONES, M. E. &amp; G. C. GRIGG &amp; L. A. BEARD" box="[188,371,961,984]" pageId="9" pageNumber="10" pagination="53 - 60" refId="ref18417" refString="JONES, M. E., G. C. GRIGG, AND L. A. BEARD. 1997. Body temperatures and activity patterns of Tasmanian devils (Sarcophilus harrisii) and eastern quolls (Dasyurus viverrinus) through a subalpine winter. Physiological Zoology 70: 53 - 60." type="journal article" year="1997">Jones et al. 1997</bibRefCitation>
) spent an average of 8 active hours in both summer and winter (in snow). Continuous radiotracking revealed that travel was intermittent, perhaps indicating a dual foraging tactic, i.e., that of ambush predator (waiting to rush and overpower prey) while also covering large distances to maximize chances of finding a carcass. Nicol and Maskrey (1986) determined that a 5.2-kg
<taxonomicName id="D0A04D1AFFE6E7113C41FB491FCAFB51" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[333,442,1153,1176]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113C41FB491F2AFB51" box="[333,346,1153,1176]" italics="true" pageId="9" pageNumber="10">S</emphasis>
.
<emphasis id="25D4EA8BFFE6E7113C66FB491FCAFB51" box="[362,442,1153,1176]" italics="true" pageId="9" pageNumber="10">harrisii</emphasis>
</taxonomicName>
had a maximal aerobic running speed of about
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/h, making such a dual strategy plausible.
</paragraph>
<paragraph id="171F3699FFE6E7113D94FB091F65FA31" blockId="9.[109,793,385,1944]" pageId="9" pageNumber="10">
As is typical of most dasyurids,
<taxonomicName id="D0A04D1AFFE6E7113CE0FB091C26FB11" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[492,598,1217,1240]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113CE0FB091C26FB11" box="[492,598,1217,1240]" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
catches and holds small prey with its forepaws. Dead prey is examined tactilely with nose, lips, and the numerous facial vibrissae.
<taxonomicName id="D0A04D1AFFE6E7113FAFFAC91D61FAD1" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[675,785,1281,1304]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113FAFFAC91D61FAD1" box="[675,785,1281,1304]" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
, which eats its own dead as willingly as those of other animals, consumes small prey head 1st but devours larger prey by working against the lie of the hairs toward the head (Eisenberg and Leyhausen 1972). With wallabies, the fatty tail often is eaten 1st (N. J. Mooney, pers. obs.). Isotope studies (
<bibRefCitation id="73314B68FFE6E7113F57FA691D7FFA71" author="PEMBERTON, D." box="[603,783,1441,1464]" pageId="9" pageNumber="10" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
) show that
<taxonomicName id="D0A04D1AFFE6E7113DD6FA091F35FA11" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[218,325,1473,1496]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113DD6FA091F35FA11" box="[218,325,1473,1496]" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
, as gorge feeders, consumes one large meal every 48 days.
</paragraph>
<paragraph id="171F3699FFE6E7113D94F9C91A1FFEEB" blockId="9.[109,793,385,1944]" lastBlockId="9.[840,1524,203,1282]" pageId="9" pageNumber="10">
Once a carcass is located, consumption begins immediately unless another
<taxonomicName id="D0A04D1AFFE6E7113C03F9E91F0BF9F1" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[271,379,1569,1592]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113C03F9E91F0BF9F1" box="[271,379,1569,1592]" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
already has possession of the carcass; entry usually is through a wound, the cloaca, or the thin skin near the pouch or scrotum (Pemberton and Renouf 1993). Eating and tearing quickly,
<taxonomicName id="D0A04D1AFFE6E7113C4AF9491FC1F951" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[326,433,1665,1688]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113C4AF9491FC1F951" box="[326,433,1665,1688]" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
consumes the softer organs it can easily reach and then moves to limbs, completely consuming the bones as it does. The sounds of breaking bones, and the snuffling and other noises made by a feeding animal, usually are heard by others in the vicinity, and then the aggressive interactions at the carcass begin. Increasing amounts of time must be spent driving off intruders. Based on 47 nights (482 h) of direct observation at standardized carcasses at 7 sites of differing densities (
<bibRefCitation id="73314B68FFE6E7113FB3F8A91EACF851" author="HAMEDE, R. K. &amp; H. MCCALLUM &amp; M. E. JONES" pageId="9" pageNumber="10" pagination="614 - 622" refId="ref17203" refString="HAMEDE, R. K., H. MCCALLUM, AND M. E. JONES. 2008. Seasonal, demographic and density-related patterns of contact between Tasmanian devils (Sarcophilus harrisii): implications for transmission of devil facial tumour disease. Austral Ecology 33: 614 - 622." type="journal article" year="2008">Hamede et al. 2008</bibRefCitation>
), the proportion of more than 1
<taxonomicName id="D0A04D1AFFE6E7113F3CF8491CEAF851" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[560,666,1921,1944]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113F3CF8491CEAF851" box="[560,666,1921,1944]" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
at a carcass increased with density and, more importantly, so did the rate at which bites were inflicted. Nearly 90% of bites, and most puncture bites, were to the head.
</paragraph>
<paragraph id="171F3699FFE6E7113E7FFEE31A42FCEB" blockId="9.[840,1524,203,1282]" pageId="9" pageNumber="10">
Lone animals feeding on a carcass spend more time being vigilant than multiple feeders (Jones 1998). If the carcass is large, such as adult common wombat or sheep, feeding
<taxonomicName id="D0A04D1AFFE6E71138A5FEA31D03FE6B" baseAuthorityName="Jones" baseAuthorityYear="1995" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E71138A5FEA31D03FE6B" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
can take up positions at the ends and sides, but they usually do not feed shoulder to shoulder. The successful outcome of agonistic interactions (
<figureCitation id="8F9B2A1CFFE6E7113953FE031AEFFE2B" box="[1119,1183,459,482]" captionStart="Fig" captionStartId="9.[840,875,1773,1794]" captionTargetBox="[861,1501,1324,1760]" captionTargetId="figure-919@9.[861,1501,1324,1760]" captionTargetPageId="9" captionText="Fig. 4.—Typical aggression between adult Sarcophilus harrisii, whether at a carcass or in contention for mates. Note the massive head, the postures during threat displays, the differential placement of white markings, and the pale cheeks of the older animal on the right, the result of scars from past conflicts. Photograph by Christo Baars used with permission." figureDoi="http://doi.org/10.5281/zenodo.4573418" httpUri="https://zenodo.org/record/4573418/files/figure.png" pageId="9" pageNumber="10">Fig. 4</figureCitation>
) at carcasses depends on body size (larger), age (older), and the degree of hunger (hungrier—
<bibRefCitation id="73314B68FFE6E7113E44FDC31DB1FDEB" author="JONES, M. E." box="[840,961,523,546]" pageId="9" pageNumber="10" refId="ref18123" refString="JONES, M. E. 1995. Guild structure of the large marsupial carnivores in Tasmania. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1995">Jones 1995</bibRefCitation>
). Such interactions sometimes result in injury, particularly on the jowls and face, and wounds are common on older animals, often lightening the color of the face from the scars. Lactating females with denned young are especially aggressive. Young animals do not fare well in such feeding competitions and often obtain scraps only after larger animals depart. Young show higher levels of vigilance behavior at carcasses than adults, consistent with their smaller size and risk of injury from older animals (Jones 1998).
</paragraph>
<paragraph id="171F3699FFE6E7113E7FFCE31AE2FC0B" blockId="9.[840,1524,203,1282]" pageId="9" pageNumber="10">
The eastern quoll, the smallest syntopic dasyurid, readily forages on mammal carcasses while maintaining higher levels of vigilance than
<taxonomicName id="D0A04D1AFFE6E7113EEAFCA31A21FC4B" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[998,1105,875,898]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113EEAFCA31A21FC4B" box="[998,1105,875,898]" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
does (Jones 1998). Once an
<taxonomicName id="D0A04D1AFFE6E711388BFCA31B83FC4B" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[1415,1523,875,898]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E711388BFCA31B83FC4B" box="[1415,1523,875,898]" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
arrives, the quoll moves to the sidelines and when a 2nd
<taxonomicName id="D0A04D1AFFE6E71138A6FC431D03FC0B" baseAuthorityName="Jones" baseAuthorityYear="1995" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E71138A6FC431D03FC0B" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
appears, the quoll departs.
</paragraph>
<paragraph id="171F3699FFE6E7123E7FFC031CEAFDEB" blockId="9.[840,1524,203,1282]" lastBlockId="10.[109,793,203,1922]" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
Pemberton and Renouf (1993) examined the role of visual communication at dimly illuminated carcasses and recorded 20 postures observed during social interactions. Although physical damage seldom resulted, agonistic interactions led to wounds on the muzzle and rump, as supported by heavy scarring in those regions, particularly in males. There is no hierarchical structure to the sequence in which
<taxonomicName id="D0A04D1AFFE6E7113950FB431AB7FB6B" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[1116,1223,1163,1186]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E7113950FB431AB7FB6B" box="[1116,1223,1163,1186]" italics="true" pageId="9" pageNumber="10">S. harrisii</emphasis>
</taxonomicName>
feeds, and feeding duration seems to be determined by satiety; feeding bouts of all individuals were similar in length except that some were longer because of interruptions caused by conflicts. Pemberton and Renouf (1993), who placed
<quantity id="D0589B7CFFE5E7123DF8FF031F43FF2B" box="[244,307,203,226]" metricMagnitude="1" metricUnit="kg" metricValue="2.0" pageId="10" pageNumber="11" unit="kg" value="20.0">20 kg</quantity>
of wombat and wallaby bait before making observations in controlled conditions, report that a mean of 7.5 devils consumed 90% of the carcasses (leaving vertebrae, skulls, and some long bones), each consuming about
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. After a cow was shot in the study area, 22
<taxonomicName id="D0A04D1AFFE5E7123CC8FE831C41FEAB" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[452,561,331,354]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E7123CC8FE831C41FEAB" box="[452,561,331,354]" italics="true" pageId="10" pageNumber="11">S. harrisii</emphasis>
</taxonomicName>
were observed feeding simultaneously, but overall, 46.8% of observations involved 25
<taxonomicName id="D0A04D1AFFE5E7123DA8FE431F65FE6B" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[164,277,395,418]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E7123DA8FE431F65FE6B" box="[164,277,395,418]" italics="true" pageId="10" pageNumber="11">S. harrisii</emphasis>
</taxonomicName>
feeding simultaneously on smaller carcasses (Pemberton and Renouf 1993). Feeding duration averaged 34 min for both sexes; however, adult males fed much longer, 57.1 min. At a study site with a low population density of
<taxonomicName id="D0A04D1AFFE5E7123F70FE231C98FDCB" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[636,744,491,514]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E7123F70FE231C98FDCB" box="[636,744,491,514]" italics="true" pageId="10" pageNumber="11">S. harrisii</emphasis>
</taxonomicName>
, 1.6 animals fed for an average of 33.4 min (
<bibRefCitation id="73314B68FFE5E7123F18FDC31CFAFDEB" author="JONES, M. E." box="[532,650,523,546]" pageId="10" pageNumber="11" refId="ref18123" refString="JONES, M. E. 1995. Guild structure of the large marsupial carnivores in Tasmania. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1995">Jones 1995</bibRefCitation>
).
</paragraph>
<caption id="43DF6611FFE6E7113E44F9251DC8F85C" ID-DOI="http://doi.org/10.5281/zenodo.4573418" ID-Zenodo-Dep="4573418" httpUri="https://zenodo.org/record/4573418/files/figure.png" pageId="9" pageNumber="10" startId="9.[840,875,1773,1794]" targetBox="[861,1501,1324,1760]" targetPageId="9">
<paragraph id="171F3699FFE6E7113E44F9251DC8F85C" blockId="9.[840,1523,1773,1941]" pageId="9" pageNumber="10">
<emphasis id="25D4EA8BFFE6E7113E44F9251DE3F8CB" bold="true" box="[840,915,1773,1794]" pageId="9" pageNumber="10">Fig. 4.</emphasis>
—Typical aggression between adult
<taxonomicName id="D0A04D1AFFE6E711382CF9251B9CF8CB" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[1312,1516,1773,1794]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE6E711382CF9251B9CF8CB" box="[1312,1516,1773,1794]" italics="true" pageId="9" pageNumber="10">Sarcophilus harrisii</emphasis>
</taxonomicName>
, whether at a carcass or in contention for mates. Note the massive head, the postures during threat displays, the differential placement of white markings, and the pale cheeks of the older animal on the right, the result of scars from past conflicts. Photograph by Christo Baars used with permission.
</paragraph>
</caption>
<paragraph id="171F3699FFE5E7123D94FDE31CBEFBCB" blockId="10.[109,793,203,1922]" pageId="10" pageNumber="11">
Social behavior is poorly developed in dasyurids, predation and scavenging being largely solitary activities. Pemberton and Renouf (1993), who observed more than 200
<taxonomicName id="D0A04D1AFFE5E7123F6DFDA31CBDFD4B" baseAuthorityName="Jones" baseAuthorityYear="1995" box="[609,717,619,642]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E7123F6DFDA31CBDFD4B" box="[609,717,619,642]" italics="true" pageId="10" pageNumber="11">S. harrisii</emphasis>
</taxonomicName>
across a 46-km
<superScript id="E0D59BD1FFE5E7123DC5FD411EA1FD5E" attach="left" box="[201,209,649,663]" fontSize="6" pageId="10" pageNumber="11">2</superScript>
area, never saw animals traveling in pairs or groups and never heard them vocalize except at carcasses. However, the overlapping home ranges indicate a tolerance among neighbors (a prerequisite to sociality), allowing Buchmann and Guiler (1977) to speculate that closed (family?) groups may feed communally on large carcasses. Play behaviors such as locomotory, mock attack, and wrestling are described for
<taxonomicName id="D0A04D1AFFE5E7123F0DFC831C88FCAB" authority="(Fleay 1935)" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[513,760,843,866]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E7123F0DFC831C19FCAB" box="[513,617,843,866]" italics="true" pageId="10" pageNumber="11">S. harrisii</emphasis>
(
<bibRefCitation id="73314B68FFE5E7123F74FC831C9DFCAB" author="FLEAY, D." box="[632,749,843,866]" pageId="10" pageNumber="11" pagination="100 - 105" refId="ref16133" refString="FLEAY, D. 1935. Notes on breeding of Tasmanian devils. Victorian Naturalist 52: 100 - 105." type="journal article" year="1935">Fleay 1935</bibRefCitation>
)
</taxonomicName>
. In captivity, males dominate females except during pregnancy and when females are guarding young (Buchmann and Guiler 1977). The male often confines the female to her den for up to 12 days after mating (
<bibRefCitation id="73314B68FFE5E7123C0DFC031F0AFC2B" author="FLEAY, D." box="[257,378,971,994]" pageId="10" pageNumber="11" pagination="100 - 105" refId="ref16133" refString="FLEAY, D. 1935. Notes on breeding of Tasmanian devils. Victorian Naturalist 52: 100 - 105." type="journal article" year="1935">Fleay 1935</bibRefCitation>
), a behavior confirmed in the wild by
<bibRefCitation id="73314B68FFE5E7123D61FC231F5DFBCB" author="PEMBERTON, D." box="[109,301,1003,1026]" pageId="10" pageNumber="11" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton (1990)</bibRefCitation>
and in zoo studies (
<bibRefCitation id="73314B68FFE5E7123F0CFC231CCFFBCB" author="KEELEY, T. &amp; J. K. O'BRIEN &amp; B. G. FANSON &amp; K. MASTERS &amp; P. D. MCGREEVY" box="[512,703,1003,1026]" pageId="10" pageNumber="11" pagination="182 - 191" refId="ref18983" refString="KEELEY, T., J. K. O'BRIEN, B. G. FANSON, K. MASTERS, AND P. D. MCGREEVY. 2012. The reproductive cycle of the Tasmanian devil (Sarcophilus harrisii) and factors associated with reproductive success in captivity. General and Comparative Endocrinology 176: 182 - 191." type="journal article" year="2012">Keeley et al. 2012</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE5E7123D94FBC31F88FACB" blockId="10.[109,793,203,1922]" pageId="10" pageNumber="11">
As in most nocturnal mammals, olfactory and auditory communication are most important, but
<taxonomicName id="D0A04D1AFFE5E7123F0BFBE31C07FB8B" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[519,631,1067,1090]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E7123F0BFBE31C07FB8B" box="[519,631,1067,1090]" italics="true" pageId="10" pageNumber="11">S. harrisii</emphasis>
</taxonomicName>
also has well developed open-mouth (visual and olfactory) behaviors that can be a preliminary to such agonistic behaviors as chases, bites, wrestling, and, rarely, locking of jaws (Buchmann and Guiler 1977; Eisenberg and Golani 1977).
<taxonomicName id="D0A04D1AFFE5E7123CE9FB631C3FFB0B" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[485,591,1195,1218]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E7123CE9FB631C3FFB0B" box="[485,591,1195,1218]" italics="true" pageId="10" pageNumber="11">S. harrisii</emphasis>
</taxonomicName>
also displays a lateral posture after “shouldering” against an opponent, including bumping and pushing with the rump.
</paragraph>
<paragraph id="171F3699FFE5E7123D94FAC31CDEFA2B" blockId="10.[109,793,203,1922]" pageId="10" pageNumber="11">
Visual and olfactory behaviors are usually accompanied by vocalizations; 8 vocalizations and 2 non-vocal sounds are made by
<taxonomicName id="D0A04D1AFFE5E7123D82FA831E87FAAB" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[142,247,1355,1378]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E7123D82FA831E87FAAB" box="[142,247,1355,1378]" italics="true" pageId="10" pageNumber="11">S. harrisii</emphasis>
</taxonomicName>
in captivity (
<bibRefCitation id="73314B68FFE5E7123C8EFA831C2FFAAB" author="EISENBERG, J. F. &amp; L. R. COLLINS &amp; C. WEMMER" box="[386,607,1355,1378]" pageId="10" pageNumber="11" pagination="379 - 399" refId="ref15759" refString="EISENBERG, J. F., L. R. COLLINS, AND C. WEMMER. 1975. Communication in the Tasmanian devil (Sarcophilus harrisii) and a survey of auditory communication in the Marsupialia. Zeitschrift fur Tierpsychologie 37: 379 - 399." type="journal article" year="1975">Eisenberg et al. 1975</bibRefCitation>
). The loud snorts and barks of a
<taxonomicName id="D0A04D1AFFE5E7123C1EFAA31F0FFA4B" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[274,383,1387,1410]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E7123C1EFAA31F0FFA4B" box="[274,383,1387,1410]" italics="true" pageId="10" pageNumber="11">S. harrisii</emphasis>
</taxonomicName>
defending a food source can explode with the arrival of intruders into a loud cacophony of agonistic vocalizations, including growls, grunts, hisses, moans, footstamping, whines, and shrieks (
<bibRefCitation id="73314B68FFE5E7123CB2FA031CEEFA2B" author="EISENBERG, J. F. &amp; L. R. COLLINS &amp; C. WEMMER" box="[446,670,1483,1506]" pageId="10" pageNumber="11" pagination="379 - 399" refId="ref15759" refString="EISENBERG, J. F., L. R. COLLINS, AND C. WEMMER. 1975. Communication in the Tasmanian devil (Sarcophilus harrisii) and a survey of auditory communication in the Marsupialia. Zeitschrift fur Tierpsychologie 37: 379 - 399." type="journal article" year="1975">Eisenberg et al. 1975</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE5E7123D94FA231CB0F9AB" blockId="10.[109,793,203,1922]" pageId="10" pageNumber="11">
Of 11 vocalizations identified in the wild by Pemberton and Renouf (1993), the briefest was a snort lasting 162 ms and the longest a growl emitted for nearly
<geoCoordinate id="7294505EFFE5E7123CD4F9E31C7CF98B" box="[472,524,1579,1602]" degrees="5.5" direction="south" orientation="latitude" pageId="10" pageNumber="11" precision="5555" value="-5.5">5.5 s</geoCoordinate>
. All were distinguishable by humans, and most had low frequencies of 912 kHz.
</paragraph>
<paragraph id="171F3699FFE5E7123D94F9A31FDEF84B" blockId="10.[109,793,203,1922]" pageId="10" pageNumber="11">During courtship, the male makes a “huff” or “clap” sound while pursuing the female. Prolonged chases and thwarted mounting attempts usually are part of courtship. In captivity, a male marks its space with scent, either from a sternal rub, urine dribble, or a cloacal drag (Eisenberg and Golani 1977). The cloacal drag seemingly is important early in an interaction but once an animal has established dominance the frequency of marking behavior diminishes progressively in the subordinate animal (Buchmann and Guiler 1977).</paragraph>
<paragraph id="171F3699FFE5E7123E7FFF031BDEFE0B" blockId="10.[840,1523,203,770]" pageId="10" pageNumber="11">
In copulation, the typical neck-grip behavior of dasyurids sometimes is augmented by the male dragging the female. During copulation, which can last for several hours, the male grasps the female with his forelimbs, sometimes palpating her abdomen, especially when thrusting (
<bibRefCitation id="73314B68FFE5E712397FFE831A9BFEAB" author="CROFT, D. B." box="[1139,1259,331,354]" pageId="10" pageNumber="11" refId="ref15459" refString="CROFT, D. B. 1982. Communication in the Dasyuridae (Marsupialia): a review. Pp. 291 - 309 in Carnivorous marsupials (M. Archer, ed.). Surrey Beatty &amp; Sons Pty Limited, Chipping Norton, New South Wales, Australia." type="book" year="1982">Croft 1982</bibRefCitation>
). Because males sometimes hold females in dens during a copulation period lasting more than 1 week,
<taxonomicName id="D0A04D1AFFE5E712391FFE431A0EFE6B" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[1043,1150,395,418]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E712391FFE431A0EFE6B" box="[1043,1150,395,418]" italics="true" pageId="10" pageNumber="11">S. harrisii</emphasis>
</taxonomicName>
may be facultatively monogamous in response to high population density (
<bibRefCitation id="73314B68FFE5E71239E2FE631BEDFE0B" author="PEMBERTON, D." box="[1262,1437,427,450]" pageId="10" pageNumber="11" refId="ref20617" refString="PEMBERTON, D. 1990. Social organisation and behaviour of the Tasmanian devil, Sarcophilus harrisii. Ph. D. dissertation, University of Tasmania, Hobart, Australia." type="book" year="1990">Pemberton 1990</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE5E7123E7FFE031A8CFCCB" blockId="10.[840,1523,203,770]" pageId="10" pageNumber="11">After the long period of pouch life, young are left behind in a vegetation-lined nesting chamber of a den while the mother goes out foraging. The female continues to nurse young for another 4 months and may bring food back to the den until the young are weaned, usually between mid-December and mid- January. About 2 months after permanent pouch vacation, usually in October, the young start to venture out from the den by themselves at night. The young are not taught to hunt nor do they accompany the mother at night. Although dispersal is malebiased, some young females do disperse.</paragraph>
</subSubSection>
</subSection>
<subSection id="7E2F2F72FFE5E712395EFC8E1A20F848" pageId="10" pageNumber="11" type="multiple">
<subSubSection id="5FBA6512FFE5E712395EFC8E1A20F848" pageId="10" pageNumber="11" type="description">
<paragraph id="171F3699FFE5E712395EFC8E1A9AFCA9" blockId="10.[1106,1258,838,864]" box="[1106,1258,838,864]" pageId="10" pageNumber="11">
<heading id="4C5781F5FFE5E712395EFC8E1A9AFCA9" allCaps="true" bold="true" box="[1106,1258,838,864]" centered="true" fontSize="11" level="1" pageId="10" pageNumber="11" reason="6">
<emphasis id="25D4EA8BFFE5E712395EFC8E1A9AFCA9" bold="true" box="[1106,1258,838,864]" pageId="10" pageNumber="11">GENETICS</emphasis>
</heading>
</paragraph>
<paragraph id="171F3699FFE5E7123E7FFC421A3AF988" blockId="10.[840,1523,906,1921]" pageId="10" pageNumber="11">
<taxonomicName id="D0A04D1AFFE5E7123E7FFC421A3AFC68" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[883,1098,906,929]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E7123E7FFC421A3AFC68" box="[883,1098,906,929]" italics="true" pageId="10" pageNumber="11">Sarcophilus harrisii</emphasis>
</taxonomicName>
has a diploid number (2n) of 14 chromosomes, with 12 metacentric autosomes, a small metacentric X chromosome, and a tiny Y chromosome (
<bibRefCitation id="73314B68FFE5E712384EFC021B93FC28" author="SHARMAN, G. B." box="[1346,1507,970,993]" pageId="10" pageNumber="11" pagination="38 - 60" refId="ref21199" refString="SHARMAN, G. B. 1961. The mitotic chromosomes of marsupials and their bearing on taxonomy and phylogeny. Australian Journal of Zoology 9: 38 - 60." type="journal article" year="1961">Sharman 1961</bibRefCitation>
). An analysis of population genetic structure using microsatellite (
<bibRefCitation id="73314B68FFE5E7123E5DFBC21A64FBE8" author="JONES, M. E. &amp; D. PAETKAU &amp; E. GEFFEN &amp; C. MORITZ" box="[849,1044,1034,1057]" pageId="10" pageNumber="11" pagination="277 - 279" refId="ref18524" refString="JONES, M. E., D. PAETKAU, E. GEFFEN, AND C. MORITZ. 2003 b. Microsatellites for the Tasmanian devil (Sarcophilus lanarius). Molecular Ecology Notes 3: 277 - 279." type="journal article" year="2003">Jones et al. 2003b</bibRefCitation>
;
<bibRefCitation id="73314B68FFE5E7123928FBC21AAAFBE8" author="JONES, M. E. &amp; D. PAETKAU &amp; E. GEFFEN &amp; C. MORITZ" box="[1060,1242,1034,1057]" pageId="10" pageNumber="11" pagination="2197 - 2209" refId="ref18566" refString="JONES, M. E., D. PAETKAU, E. GEFFEN, AND C. MORITZ. 2004. Genetic diversity and population structure of Tasmanian devils, the largest marsupial carnivore. Molecular Ecology 13: 2197 - 2209." type="journal article" year="2004">Jones et al. 2004</bibRefCitation>
), SNP, and mitochondrial (
<bibRefCitation id="73314B68FFE5E7123E5DFBE21A76FB88" author="LACHISH, S. &amp; K. J. MILLER &amp; A. STORFER &amp; A. W. GOLDIZEN &amp; M. E. JONES" box="[849,1030,1066,1089]" pageId="10" pageNumber="11" pagination="172 - 182" refId="ref19247" refString="LACHISH, S., K. J. MILLER, A. STORFER, A. W. GOLDIZEN, AND M. E. JONES. 2011. Evidence that disease-induced population decline changes genetic structure and alters dispersal patterns in the Tasmanian devil. Heredity 106: 172 - 182." type="journal article" year="2011">Miller et al. 2011</bibRefCitation>
) and immune (
<bibRefCitation id="73314B68FFE5E71239ABFBE21B2DFB88" author="SIDDLE, H. V. &amp; J. MARSEE &amp; Y. CHENG &amp; M. JONES &amp; K. BELOV" box="[1191,1373,1066,1089]" pageId="10" pageNumber="11" pagination="2001 - 2006" refId="ref21296" refString="SIDDLE, H. V., J. MARSEE, Y. CHENG, M. JONES, AND K. BELOV. 2010. MHC gene copy number variation in Tasmanian devils: implications for the spread of a contagious cancer. Proceedings of the Royal Society of London, B. Biological Sciences 277: 2001 - 2006." type="journal article" year="2010">Siddle et al. 2010</bibRefCitation>
) loci revealed moderately low genetic variability (allelic diversity = 2.73.3; heterozygosity = 0.390.47—
<bibRefCitation id="73314B68FFE5E712399DFBA21B2FFB48" author="JONES, M. E. &amp; D. PAETKAU &amp; E. GEFFEN &amp; C. MORITZ" box="[1169,1375,1130,1153]" pageId="10" pageNumber="11" pagination="277 - 279" refId="ref18524" refString="JONES, M. E., D. PAETKAU, E. GEFFEN, AND C. MORITZ. 2003 b. Microsatellites for the Tasmanian devil (Sarcophilus lanarius). Molecular Ecology Notes 3: 277 - 279." type="journal article" year="2003">Jones et al. 2003b</bibRefCitation>
;
<bibRefCitation id="73314B68FFE5E712387FFBA21D0DFB68" author="JONES, M. E. &amp; D. PAETKAU &amp; E. GEFFEN &amp; C. MORITZ" pageId="10" pageNumber="11" pagination="2197 - 2209" refId="ref18566" refString="JONES, M. E., D. PAETKAU, E. GEFFEN, AND C. MORITZ. 2004. Genetic diversity and population structure of Tasmanian devils, the largest marsupial carnivore. Molecular Ecology 13: 2197 - 2209." type="journal article" year="2004">Jones et al. 2004</bibRefCitation>
).
<taxonomicName id="D0A04D1AFFE5E7123E99FB421A72FB68" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[917,1026,1162,1185]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE5E7123E99FB421A72FB68" box="[917,1026,1162,1185]" italics="true" pageId="10" pageNumber="11">S. harrisii</emphasis>
</taxonomicName>
has lost genetic diversity perhaps twice in its history: at the last glacial maximum (about 20,000 years ago) and more significantly about 3,000 years ago, at the end of prolonged and severe ENSO climatic events (Brüniche-Olsen 2014). Neighborhood size is about
<quantity id="D0589B7CFFE5E712397FFAC21AB6FAE8" box="[1139,1222,1290,1313]" metricMagnitude="5" metricUnit="m" metricValue="1.0" pageId="10" pageNumber="11" unit="km" value="100.0">100 km</quantity>
, similar to the postnatal dispersal distance from capture-recapture data of animals marked as juveniles (
<bibRefCitation id="73314B68FFE5E7123ED0FA821AC1FAA8" author="LACHISH, S. &amp; K. J. MILLER &amp; A. STORFER &amp; A. W. GOLDIZEN &amp; M. E. JONES" box="[988,1201,1354,1377]" pageId="10" pageNumber="11" pagination="172 - 182" refId="ref19247" refString="LACHISH, S., K. J. MILLER, A. STORFER, A. W. GOLDIZEN, AND M. E. JONES. 2011. Evidence that disease-induced population decline changes genetic structure and alters dispersal patterns in the Tasmanian devil. Heredity 106: 172 - 182." type="journal article" year="2011">Lachish et al. 2011</bibRefCitation>
). This movement distance is reflected in population structure, with extensive gene flow, which reduces at scales above
<quantity id="D0589B7CFFE5E7123992FA421A85FA68" box="[1182,1269,1418,1441]" metricMagnitude="5" metricUnit="m" metricValue="1.5" pageId="10" pageNumber="11" unit="km" value="150.0">150 km</quantity>
, and with just 2 major genetic subpopulations in
<collectingRegion id="D564F87BFFE5E7123968FA621ABCFA08" box="[1124,1228,1450,1473]" country="Australia" name="Tasmania" pageId="10" pageNumber="11">Tasmania</collectingRegion>
, due to a band of less suitable natural and anthropogenic habitat (e.g., cleared land, urban development, tall wet forest or rainforest, and alpine habitat) that separates northwestern populations from central and eastern ones (
<bibRefCitation id="73314B68FFE5E7123E84F9E21A4AF988" author="JONES, M. E. &amp; D. PAETKAU &amp; E. GEFFEN &amp; C. MORITZ" box="[904,1082,1578,1601]" pageId="10" pageNumber="11" pagination="2197 - 2209" refId="ref18566" refString="JONES, M. E., D. PAETKAU, E. GEFFEN, AND C. MORITZ. 2004. Genetic diversity and population structure of Tasmanian devils, the largest marsupial carnivore. Molecular Ecology 13: 2197 - 2209." type="journal article" year="2004">Jones et al. 2004</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE5E7123E7FF9821A20F848" blockId="10.[840,1523,906,1921]" pageId="10" pageNumber="11">
The karyotype of cancer cells present in DFTD is distinctive and stable in diseased animals from geographic regions and has the following features (Pearse and Swift 2006): 13 autosomal chromosomes but no sex-determing chromosomes; no chromosome-2 pair; only 1 chromosome-6; the long arm of one chromosome-1 was deleted; and 4 unidentified marker chromosomes were present (designated as M 1 M 4). Importantly, these anomalies were the same in tumors of all animals (
<emphasis id="25D4EA8BFFE5E7123815F8E21B56F888" box="[1305,1318,1834,1857]" italics="true" pageId="10" pageNumber="11">n</emphasis>
= 11). Thus, tumor cells are karyotypically distinctive from those of their hosts (Pearse and Swift 2006).
</paragraph>
</subSubSection>
</subSection>
<subSection id="7E2F2F72FFE4E7143C43FF031B36F9EB" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12" type="multiple">
<subSubSection id="5FBA6512FFE4E7143C43FF031B36F9EB" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12" type="conservation">
<paragraph id="171F3699FFE4E7133C43FF031C46FF2C" blockId="11.[335,566,203,229]" box="[335,566,203,229]" pageId="11" pageNumber="12">
<heading id="4C5781F5FFE4E7133C43FF031C46FF2C" allCaps="true" bold="true" box="[335,566,203,229]" centered="true" fontSize="11" level="1" pageId="11" pageNumber="12" reason="6">
<emphasis id="25D4EA8BFFE4E7133C43FF031C46FF2C" bold="true" box="[335,566,203,229]" pageId="11" pageNumber="12">CONSERVATION</emphasis>
</heading>
</paragraph>
<paragraph id="171F3699FFE4E7133D94FEC71C78FCCF" blockId="11.[109,793,271,1350]" pageId="11" pageNumber="12">
Fully protected by law since 1941,
<taxonomicName id="D0A04D1AFFE4E7133F37FEC71D68FEEF" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[571,792,271,294]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE4E7133F37FEC71D68FEEF" box="[571,792,271,294]" italics="true" pageId="11" pageNumber="12">Sarcophilus harrisii</emphasis>
</taxonomicName>
received increased protection under the Wildlife Regulations of the Tasmanian National Parks and Wildlife Act of 1971. It is now listed under both Federal (Section 178 of the Environment Protection and Biodiversity Conservation Act of 1999 [40]) and Tasmanian (Threatened Species Protection Act of 1995) laws. Until about 2000, with estimated numbers as high as 130,000 (Jones and Rose 1996),
<taxonomicName id="D0A04D1AFFE4E7133C9BFE271C7CFDCF" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[407,524,495,518]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE4E7133C9BFE271C7CFDCF" box="[407,524,495,518]" italics="true" pageId="11" pageNumber="12">S. harrisii</emphasis>
</taxonomicName>
was classified by the International Union for Conservation of Nature and Natural Resources at the category of “Lower RiskLeast Concern.” This status changed after the discovery in
<quantity id="D0589B7CFFE4E7133F1CFD871C15FDAF" box="[528,613,591,614]" metricMagnitude="1" metricUnit="m" metricValue="5.06984" pageId="11" pageNumber="12" unit="in" value="1996.0">1996 in</quantity>
a population in northeastern
<collectingRegion id="D564F87BFFE4E7133DF5FDA71F2FFD4F" box="[249,351,623,646]" country="Australia" name="Tasmania" pageId="11" pageNumber="12">Tasmania</collectingRegion>
of DFTD, a new, infectious, and always fatal cancer that has reduced some local populations by up to 90% in about 10 years, while overspreading much of
<collectingRegion id="D564F87BFFE4E7133FBFFD671D69FD0F" box="[691,793,687,710]" country="Australia" name="Tasmania" pageId="11" pageNumber="12">Tasmania</collectingRegion>
(
<bibRefCitation id="73314B68FFE4E7133D7AFD071F7FFD2F" author="MOONEY, N." box="[118,271,719,742]" pageId="11" pageNumber="12" pagination="34 - 41" refId="ref19956" refString="MOONEY, N. 2004. The devil's new Hell. Nature Australia Summer 2004 - 2005 28: 34 - 41." type="journal article" year="2004">Mooney 2004</bibRefCitation>
;
<bibRefCitation id="73314B68FFE4E7133C2FFD071C76FD2F" author="HAWKINS, C. E." box="[291,518,719,742]" pageId="11" pageNumber="12" pagination="307 - 324" refId="ref17398" refString="HAWKINS, C. E., ET AL. 2006. Emerging disease and population decline of an island endemic, the Tasmanian devil, Sarcophilus harrisii. Biological Conservation 131: 307 - 324." type="journal article" year="2006">Hawkins et al. 2006</bibRefCitation>
;
<bibRefCitation id="73314B68FFE4E7133F15FD071D60FD2F" author="LACHISH, S. &amp; M. JONES &amp; H. MCCALLUM" box="[537,784,719,742]" pageId="11" pageNumber="12" pagination="926 - 936" refId="ref19156" refString="LACHISH, S., M. JONES, AND H. MCCALLUM. 2007. The impact of disease on the survival and population growth rate of the Tasmanian devil. Journal of Animal Ecology 76: 926 - 936." type="journal article" year="2007">McCallum et al. 2007</bibRefCitation>
;
<bibRefCitation id="73314B68FFE4E7133D61FD271F48FCCF" author="LACHISH, S. &amp; H. MCCALLUM &amp; M. JONES" box="[109,312,751,774]" pageId="11" pageNumber="12" pagination="427 - 436" refId="ref19198" refString="LACHISH, S., H. MCCALLUM, AND M. JONES. 2009. Demography, disease, and the devil: life-history changes in a disease-affected population of the Tasmanian devil (Sarcophilus harrisii). Journal of Animal Ecology 78: 427 - 436." type="journal article" year="2009">Lachish et al. 2009</bibRefCitation>
; Jones et al. 2014).
</paragraph>
<paragraph id="171F3699FFE4E7133D94FCC71E86FBAF" blockId="11.[109,793,271,1350]" pageId="11" pageNumber="12">
Although population fluctuations have happened in the past (
<bibRefCitation id="73314B68FFE4E7133DA6FCE71F49FC8F" author="GUILER, E. R." box="[170,313,815,838]" pageId="11" pageNumber="12" pagination="134 - 137" refId="ref16969" refString="GUILER, E. R. 1970 c. Tasmanian devils and agriculture. Tasmanian Journal of Agriculture (1970) 41: 134 - 137." type="journal article" year="1970">Guiler 1970c</bibRefCitation>
; Bradshaw and Brook 2005), DFTD, if not controlled, clearly poses a threat of endangerment and even extinction in the wild (
<bibRefCitation id="73314B68FFE4E7133C50FCA71C54FC4F" author="LACHISH, S. &amp; M. JONES &amp; H. MCCALLUM" box="[348,548,879,902]" pageId="11" pageNumber="12" pagination="926 - 936" refId="ref19156" refString="LACHISH, S., M. JONES, AND H. MCCALLUM. 2007. The impact of disease on the survival and population growth rate of the Tasmanian devil. Journal of Animal Ecology 76: 926 - 936." type="journal article" year="2007">Lachish et al. 2007</bibRefCitation>
). Soon after the consequences of DFTD became known (McCallum and Jones 2006), state and federal agencies mounted broad-scale efforts to diagnose, describe, and curb the spread of the disease (Owen and Pemberton 2005). Information was needed from a range of scientists: ecologists, geneticists, immunologists, modelers, pathologists, and veterinarians; even the general public provided many new records.
</paragraph>
<paragraph id="171F3699FFE4E7133D94FBA71BC0FE2B" blockId="11.[109,793,271,1350]" lastBlockId="11.[840,1523,203,1186]" pageId="11" pageNumber="12">
The 1st disease symptoms appear as patches in the mouth or on the cheeks, lips, and neck (
<figureCitation id="8F9B2A1CFFE4E7133CBDFB471F9FFB6F" box="[433,495,1167,1190]" captionStart="Fig" captionStartId="11.[110,145,1837,1858]" captionTargetBox="[131,771,1398,1824]" captionTargetId="figure-692@11.[131,771,1398,1824]" captionTargetPageId="11" captionText="Fig. 5.—The 1st symptoms of the fatal disease, Devil Facial Tumor Disease, often are patches of redness on the palate, lips, and snout. Note the relatively unscarred cheeks of this young adult Sarcophilus harrisii. Photograph by Christo Baars used with permission." figureDoi="http://doi.org/10.5281/zenodo.4573422" httpUri="https://zenodo.org/record/4573422/files/figure.png" pageId="11" pageNumber="12">Fig. 5</figureCitation>
), then grow into large solid masses of soft tissue (
<figureCitation id="8F9B2A1CFFE4E7133C6BFB671FD8FB0F" box="[359,424,1199,1222]" captionStart="Fig" captionStartId="11.[840,875,1778,1799]" captionTargetBox="[861,1501,1247,1765]" captionTargetId="figure-735@11.[861,1501,1247,1765]" captionTargetPageId="11" captionText="Fig. 6.—Adult Sarcophilus harrisii (photographed prior to 1996) with what is now called Devil Facial Tumor Disease. Such gross deformities of the advanced stages of the disease prevent normal feeding behavior, leading to death by starvation or secondary infections. At the time this photograph was taken by Christo Baars, Devil Facial Tumor Disease was undocumented in the literature." figureDoi="http://doi.org/10.5281/zenodo.4573420" httpUri="https://zenodo.org/record/4573420/files/figure.png" pageId="11" pageNumber="12">Fig. 6</figureCitation>
), usually circumscribed and flattened, and later progressing to tumors with ulcerative and exudative surfaces (
<bibRefCitation id="73314B68FFE4E7133DF1FB271FD4FACF" author="LOH, R." box="[253,420,1263,1286]" pageId="11" pageNumber="12" pagination="890 - 895" refId="ref19416" refString="LOH, R., ET AL. 2006 a. The pathology of devil facial tumor disease (DFTD) in the Tasmanian devil (Sarcophilus harrisii). Veterinary Pathology 43: 890 - 895." type="journal article" year="2006">Loh et al. 2006a</bibRefCitation>
). The tumors, usually located in the dermis, are nodular aggregates of round- to spindle-shaped cells and often with a pseudocapsule divided by septae into lobules. Histologically, the cells are of one
<typeStatus id="C81B883BFFE4E71339D9FF031B73FF2B" box="[1237,1283,203,226]" pageId="11" pageNumber="12">type</typeStatus>
and characterized by large nuclei. Tumors are locally aggressive and metastasize in 65% of cases (
<bibRefCitation id="73314B68FFE4E7133EE8FEC31AE0FEEB" author="LOH, R." box="[996,1168,267,290]" pageId="11" pageNumber="12" pagination="890 - 895" refId="ref19416" refString="LOH, R., ET AL. 2006 a. The pathology of devil facial tumor disease (DFTD) in the Tasmanian devil (Sarcophilus harrisii). Veterinary Pathology 43: 890 - 895." type="journal article" year="2006">Loh et al. 2006a</bibRefCitation>
), typical of undifferentiated softtissue neoplasms. As tumors grow, body condition deteriorates until death, usually within 6 months; causes of death vary but likely include starvation and secondary infections. Importantly, histological sections of tumors and related lymph nodes revealed a general absence of infiltrating T-lymphocytes, indicative of a failed immune response by the host (
<bibRefCitation id="73314B68FFE4E71339DEFE031BD0FE2B" author="HOWSON, L. J." box="[1234,1440,459,482]" pageId="11" pageNumber="12" pagination="925 - 938" refId="ref17856" refString="HOWSON, L. J., ET AL. 2014. Identification of dendritic cells, B cell and T cell subsets in Tasmanian lymphoid tissue; evidence for poor immune cell infiltration into devil facial tumors. The Anatomical Record 297: 925 - 938." type="journal article" year="2014">Howson et al. 2014</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE4E7133E7FFE231B91FCAB" blockId="11.[840,1523,203,1186]" pageId="11" pageNumber="12">
Despite
<taxonomicName id="D0A04D1AFFE4E7133EC7FE231A46FDCB" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[971,1078,491,514]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE4E7133EC7FE231A46FDCB" box="[971,1078,491,514]" italics="true" pageId="11" pageNumber="12">S. harrisii</emphasis>
</taxonomicName>
having a functionally competent immune system (
<bibRefCitation id="73314B68FFE4E7133EAEFDC31A11FDEB" author="WOODS, G. M. &amp; A. KREISS &amp; K. BELOV &amp; H. V. SIDDLE &amp; D. L. OBENDORF &amp; H. K. MULLER" box="[930,1121,523,546]" pageId="11" pageNumber="12" pagination="338 - 345" refId="ref21886" refString="WOODS, G. M., A. KREISS, K. BELOV, H. V. SIDDLE, D. L. OBENDORF, AND H. K. MULLER. 2007. The immune response of the Tasmanian devil (Sarcophilus harrisii) and devil facial tumour disease. EcoHealth 4: 338 - 345." type="journal article" year="2007">Woods et al. 2007</bibRefCitation>
), the cells introduced by bites are not recognized as foreign, are not attacked by the immune system, and consequently are not rejected. The failure of the immune system to mount a proper response, while not well understood, is due in part to low genetic diversity, including in the major histocompatibility complex, the most variable part of the mammalian genome and important in recognition of self and non-self (
<bibRefCitation id="73314B68FFE4E7133E5DFD231A77FCCB" author="LOH, R. &amp; D. HAYES &amp; A. MAHJOOR &amp; A. O'HARA &amp; S. PYECROFT &amp; S. RAIDAL" box="[849,1031,747,770]" pageId="11" pageNumber="12" pagination="896 - 903" refId="ref19454" refString="LOH, R., D. HAYES, A. MAHJOOR, A. O'HARA, S. PYECROFT, AND S. RAIDAL. 2006 b. The immunohistochemical characterization of devil facial tumor disease (DFTD) in the Tasmanian devil (Sarcophilus harrisii). Veterinary Pathology 43: 896 - 903." type="journal article" year="2006">Loh et al. 2006b</bibRefCitation>
;
<bibRefCitation id="73314B68FFE4E713391BFD231AABFCCB" author="CHENG, Y. &amp; C. SANDERSON &amp; M. JONES" box="[1047,1243,747,770]" pageId="11" pageNumber="12" pagination="525 - 533" refId="ref15403" refString="CHENG, Y., C. SANDERSON, AND M. JONES. 2012. Low MHC class II diversity in the Tasmanian devil, Sarcophilus harrisii. Immunogenetics 64: 525 - 533." type="journal article" year="2012">Cheng et al. 2012</bibRefCitation>
). Recent genetic analysis of complete mitochondrial genomes from current and museum specimens indicates that low genetic diversity in extant populations predates DFTD by at least 100 years (
<bibRefCitation id="73314B68FFE4E7133814FC831BA0FCAB" author="LACHISH, S. &amp; K. J. MILLER &amp; A. STORFER &amp; A. W. GOLDIZEN &amp; M. E. JONES" box="[1304,1488,843,866]" pageId="11" pageNumber="12" pagination="172 - 182" refId="ref19247" refString="LACHISH, S., K. J. MILLER, A. STORFER, A. W. GOLDIZEN, AND M. E. JONES. 2011. Evidence that disease-induced population decline changes genetic structure and alters dispersal patterns in the Tasmanian devil. Heredity 106: 172 - 182." type="journal article" year="2011">Miller et al. 2011</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE4E7143E7FFCA31F29FE6B" blockId="11.[840,1523,203,1186]" lastBlockId="12.[109,793,203,1954]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
Evidence that DFTD is a transmissible neoplasm, transmitted as a clonal cell line by injurious bites between animals at carcasses or inflicted during courtship and mating (
<bibRefCitation id="73314B68FFE4E713386EFC631D0DFC2B" author="HAMEDE, R. K. &amp; H. MCCALLUM &amp; M. E. JONES" pageId="11" pageNumber="12" pagination="614 - 622" refId="ref17203" refString="HAMEDE, R. K., H. MCCALLUM, AND M. E. JONES. 2008. Seasonal, demographic and density-related patterns of contact between Tasmanian devils (Sarcophilus harrisii): implications for transmission of devil facial tumour disease. Austral Ecology 33: 614 - 622." type="journal article" year="2008">Hamede et al. 2008</bibRefCitation>
;
<bibRefCitation id="73314B68FFE4E7133E86FC031A25FC2B" author="HAMEDE, R. K. &amp; H. I. MCCALLUM &amp; M. E. JONES" box="[906,1109,971,994]" pageId="11" pageNumber="12" pagination="182 - 190" refId="ref17256" refString="HAMEDE, R. K., H. I. MCCALLUM, AND M. E. JONES. 2013. Biting injuries and transmission of devil facial tumour disease. Journal of Animal Ecology 82: 182 - 190." type="journal article" year="2013">Hamede et al. 2013</bibRefCitation>
), includes identical chromosomal rearrangements in all clones within each identified morphotype and the successful induction of the disease in healthy
<taxonomicName id="D0A04D1AFFE4E7133847FBC31BC4FBEB" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[1355,1460,1035,1058]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE4E7133847FBC31BC4FBEB" box="[1355,1460,1035,1058]" italics="true" pageId="11" pageNumber="12">S. harrisii</emphasis>
</taxonomicName>
using cells from both natural tumors and cultured cell lines (
<bibRefCitation id="73314B68FFE4E7133894FBE31DC5FBAB" author="PYECROFT, S. B." pageId="11" pageNumber="12" pagination="346 - 351" refId="ref20836" refString="PYECROFT, S. B., ET AL. 2007. Towards a case definition for devil facial tumour disease: what is it? EcoHealth 4: 346 - 351." type="journal article" year="2007">Pyecroft et al. 2007</bibRefCitation>
). DFTD arose in a female, as evidenced by X chromosome but no Y chromosome (
<bibRefCitation id="73314B68FFE4E7133970FBA31B5BFB4B" author="TOVAR, C. &amp; D. OBENDORF &amp; E. P. MURCHISON &amp; A. T. PAPENFUSS &amp; A. KREISS &amp; G. M. WOODS" box="[1148,1323,1131,1154]" pageId="11" pageNumber="12" pagination="1195 - 1203" refId="ref21574" refString="TOVAR, C., D. OBENDORF, E. P. MURCHISON, A. T. PAPENFUSS, A. KREISS, AND G. M. WOODS. 2011. A tumor-specific diagnostic marker for transmissible facial tumors of Tasmanian devils: immunohistochemistry studies. Veterinary Pathology 48: 1195 - 1203." type="journal article" year="2011">Tovar et al. 2011</bibRefCitation>
), in cells of neuroendocrine origin (
<bibRefCitation id="73314B68FFE4E7133905FB431ACBFB6B" author="LOH, R. &amp; D. HAYES &amp; A. MAHJOOR &amp; A. O'HARA &amp; S. PYECROFT &amp; S. RAIDAL" box="[1033,1211,1163,1186]" pageId="11" pageNumber="12" pagination="896 - 903" refId="ref19454" refString="LOH, R., D. HAYES, A. MAHJOOR, A. O'HARA, S. PYECROFT, AND S. RAIDAL. 2006 b. The immunohistochemical characterization of devil facial tumor disease (DFTD) in the Tasmanian devil (Sarcophilus harrisii). Veterinary Pathology 43: 896 - 903." type="journal article" year="2006">Loh et al. 2006b</bibRefCitation>
), more specifically Schwann cells (
<bibRefCitation id="73314B68FFE3E7143DBCFF031FD1FF2B" author="MURCHISON, E. P." box="[176,417,203,226]" pageId="12" pageNumber="13" pagination="84 - 87" refId="ref20004" refString="MURCHISON, E. P., ET AL. 2010. The Tasmanian devil transcriptome reveals Schwann cell origins of a clonally transmissible cancer. Science 327: 84 - 87." type="journal article" year="2010">Murchison et al. 2010</bibRefCitation>
). One Schwann cell protein, periaxin, was identified as being diagnostic for DFTD (
<bibRefCitation id="73314B68FFE3E7143F92FF231ED2FEEB" author="TOVAR, C. &amp; D. OBENDORF &amp; E. P. MURCHISON &amp; A. T. PAPENFUSS &amp; A. KREISS &amp; G. M. WOODS" pageId="12" pageNumber="13" pagination="1195 - 1203" refId="ref21574" refString="TOVAR, C., D. OBENDORF, E. P. MURCHISON, A. T. PAPENFUSS, A. KREISS, AND G. M. WOODS. 2011. A tumor-specific diagnostic marker for transmissible facial tumors of Tasmanian devils: immunohistochemistry studies. Veterinary Pathology 48: 1195 - 1203." type="journal article" year="2011">Tovar et al. 2011</bibRefCitation>
). This same research group (
<bibRefCitation id="73314B68FFE3E7143CD6FEC31CE5FEEB" author="KREISS, A. &amp; C. TOVAR &amp; D. L. OBENDORF &amp; K. DUN &amp; G. M. WOODS" box="[474,661,267,290]" pageId="12" pageNumber="13" pagination="475 - 481" refId="ref19108" refString="KREISS, A., C. TOVAR, D. L. OBENDORF, K. DUN, AND G. M. WOODS. 2011. A murine xenograft model for a transmissible cancer in Tasmanian devils. Veterinary Pathology 48: 475 - 481." type="journal article" year="2011">Kreiss et al. 2011</bibRefCitation>
) also developed a mouse model that reproduces DFTD with fidelity: mouse cells are similar histologically and karyotypically, and xenografted DFTD tumors expressed periaxin, the marker protein that diagnoses DFTD.
</paragraph>
<caption id="43DF6611FFE4E7133E44F93A1AD0F853" ID-DOI="http://doi.org/10.5281/zenodo.4573420" ID-Zenodo-Dep="4573420" httpUri="https://zenodo.org/record/4573420/files/figure.png" pageId="11" pageNumber="12" startId="11.[840,875,1778,1799]" targetBox="[861,1501,1247,1765]" targetPageId="11">
<paragraph id="171F3699FFE4E7133E44F93A1AD0F853" blockId="11.[840,1523,1778,1946]" pageId="11" pageNumber="12">
<emphasis id="25D4EA8BFFE4E7133E44F93A1DF8F8CE" bold="true" box="[840,904,1778,1799]" pageId="11" pageNumber="12">Fig. 6.</emphasis>
—Adult
<taxonomicName id="D0A04D1AFFE4E7133ED2F93B1AEFF8C1" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[990,1183,1779,1800]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE4E7133ED2F93B1AEFF8C1" box="[990,1183,1779,1800]" italics="true" pageId="11" pageNumber="12">Sarcophilus harrisii</emphasis>
</taxonomicName>
(photographed prior to 1996) with what is now called Devil Facial Tumor Disease. Such gross deformities of the advanced stages of the disease prevent normal feeding behavior, leading to death by starvation or secondary infections. At the time this photograph was taken by Christo Baars, Devil Facial Tumor Disease was undocumented in the literature.
</paragraph>
</caption>
<caption id="43DF6611FFE4E7133D62F8E51C2DF853" ID-DOI="http://doi.org/10.5281/zenodo.4573422" ID-Zenodo-Dep="4573422" httpUri="https://zenodo.org/record/4573422/files/figure.png" pageId="11" pageNumber="12" startId="11.[110,145,1837,1858]" targetBox="[131,771,1398,1824]" targetPageId="11">
<paragraph id="171F3699FFE4E7133D62F8E51C2DF853" blockId="11.[109,793,1837,1946]" pageId="11" pageNumber="12">
<emphasis id="25D4EA8BFFE4E7133D62F8E51EC1F88B" bold="true" box="[110,177,1837,1858]" pageId="11" pageNumber="12">Fig. 5.</emphasis>
—The 1st symptoms of the fatal disease, Devil Facial Tumor Disease, often are patches of redness on the palate, lips, and snout. Note the relatively unscarred cheeks of this young adult
<emphasis id="25D4EA8BFFE4E7133F5FF8A01D69F8B4" box="[595,793,1896,1917]" italics="true" pageId="11" pageNumber="12">
<taxonomicName id="D0A04D1AFFE4E7133F5FF8A01D65F8B4" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[595,789,1896,1917]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="harrisii">Sarcophilus harrisii</taxonomicName>
.
</emphasis>
Photograph by Christo Baars used with permission.
</paragraph>
</caption>
<paragraph id="171F3699FFE3E7143D94FE631F99FCEB" blockId="12.[109,793,203,1954]" pageId="12" pageNumber="13">
Cells of all tumors have identical genotypes at multiple microsatellite and histocompatibility complex loci (
<bibRefCitation id="73314B68FFE3E7143FD8FE031EAFFDCB" author="WOODS, G. M. &amp; A. KREISS &amp; K. BELOV &amp; H. V. SIDDLE &amp; D. L. OBENDORF &amp; H. K. MULLER" pageId="12" pageNumber="13" pagination="338 - 345" refId="ref21886" refString="WOODS, G. M., A. KREISS, K. BELOV, H. V. SIDDLE, D. L. OBENDORF, AND H. K. MULLER. 2007. The immune response of the Tasmanian devil (Sarcophilus harrisii) and devil facial tumour disease. EcoHealth 4: 338 - 345." type="journal article" year="2007">Siddle et al. 2007</bibRefCitation>
), further indication that all tumors are a single clone derived from one
<taxonomicName id="D0A04D1AFFE3E7143C4CFDC31FC2FDEB" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[320,434,523,546]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE3E7143C4CFDC31FC2FDEB" box="[320,434,523,546]" italics="true" pageId="12" pageNumber="13">S. harrisii</emphasis>
</taxonomicName>
, an opinion also supported by
<bibRefCitation id="73314B68FFE3E7143D61FDE31F17FD8B" author="MURCHISON, E. P." box="[109,359,555,578]" pageId="12" pageNumber="13" pagination="84 - 87" refId="ref20004" refString="MURCHISON, E. P., ET AL. 2010. The Tasmanian devil transcriptome reveals Schwann cell origins of a clonally transmissible cancer. Science 327: 84 - 87." type="journal article" year="2010">Murchison et al. (2010)</bibRefCitation>
. Thus, tumor cells are genetically different from host cells. Furthermore, the karyotypes of cells taken from tumors of animals of different sexes, ages, and regions are consistently similar in the high degree of aneuploidy and chromosomal rearrangement, indicating that cytogenetically, DFTD is relatively stable, leading Pearse and Swift (2006) to hypothesize that the cancer is acting as a transmissible allograft, with infectious cells introduced by bites.
</paragraph>
<paragraph id="171F3699FFE3E7143D94FCE31C9CFA8B" blockId="12.[109,793,203,1954]" pageId="12" pageNumber="13">
The best early evidence of the local impact of the disease is from a longitudinal study, initiated in 1999 by M. Jones, of a
<taxonomicName id="D0A04D1AFFE3E7143D8EFCA31E9FFC4B" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[130,239,875,898]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE3E7143D8EFCA31E9FFC4B" box="[130,239,875,898]" italics="true" pageId="12" pageNumber="13">S. harrisii</emphasis>
</taxonomicName>
population at Freycinet National Park, also on the east coast (
<bibRefCitation id="73314B68FFE3E7143DEFFC431FDDFC6B" author="LACHISH, S. &amp; M. JONES &amp; H. MCCALLUM" box="[227,429,907,930]" pageId="12" pageNumber="13" pagination="926 - 936" refId="ref19156" refString="LACHISH, S., M. JONES, AND H. MCCALLUM. 2007. The impact of disease on the survival and population growth rate of the Tasmanian devil. Journal of Animal Ecology 76: 926 - 936." type="journal article" year="2007">Lachish et al. 2007</bibRefCitation>
). Annual adult survival was about 0.5 and adults comprised 5153% of the population until 2001, when the 1st diseased animal was detected. As DFTD moved southward, adult survival dropped to near zero within 3 years. By contrast, apparent survival of subadults, although more variable among years, was relatively constant across the 7-year study, declining slightly at the end. During the 1st two years of the decline in adult numbers, population size of subadults increased substantially, as if in response to release from competition from adults (
<bibRefCitation id="73314B68FFE3E7143DCCFB631FE4FB0B" author="LACHISH, S. &amp; M. JONES &amp; H. MCCALLUM" box="[192,404,1195,1218]" pageId="12" pageNumber="13" pagination="926 - 936" refId="ref19156" refString="LACHISH, S., M. JONES, AND H. MCCALLUM. 2007. The impact of disease on the survival and population growth rate of the Tasmanian devil. Journal of Animal Ecology 76: 926 - 936." type="journal article" year="2007">Lachish et al. 2007</bibRefCitation>
;
<figureCitation id="8F9B2A1CFFE3E7143CA9FB631F96FB0B" box="[421,486,1195,1218]" captionStart="Fig" captionStartId="11.[110,145,1837,1858]" captionTargetBox="[131,771,1398,1824]" captionTargetId="figure-692@11.[131,771,1398,1824]" captionTargetPageId="11" captionText="Fig. 5.—The 1st symptoms of the fatal disease, Devil Facial Tumor Disease, often are patches of redness on the palate, lips, and snout. Note the relatively unscarred cheeks of this young adult Sarcophilus harrisii. Photograph by Christo Baars used with permission." figureDoi="http://doi.org/10.5281/zenodo.4573422" httpUri="https://zenodo.org/record/4573422/files/figure.png" pageId="12" pageNumber="13">Fig. 5</figureCitation>
). But this effect was shortlived and after adult population size fell to near zero, subadult numbers dipped to one-third of former levels. Documenting population declines is difficult but spotlight surveys of this population indicate reductions of 6990% (
<bibRefCitation id="73314B68FFE3E7143CF8FAE31CACFA8B" author="LACHISH, S. &amp; M. JONES &amp; H. MCCALLUM" box="[500,732,1323,1346]" pageId="12" pageNumber="13" pagination="926 - 936" refId="ref19156" refString="LACHISH, S., M. JONES, AND H. MCCALLUM. 2007. The impact of disease on the survival and population growth rate of the Tasmanian devil. Journal of Animal Ecology 76: 926 - 936." type="journal article" year="2007">McCallum et al. 2007</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE3E7143D94FA831C70F86B" blockId="12.[109,793,203,1954]" pageId="12" pageNumber="13">
The outbreak and spread of DFTD through a population is characterized by a change in age structure, due to the loss of breeding adults (≥ 2 years). The average age of diseased animals in the Freycinet population was 2.27 years, but after 6 years adults comprised only 20% of the population and only one older animal had survived (
<bibRefCitation id="73314B68FFE3E7143C5BFA231C54F9CB" author="LACHISH, S. &amp; H. MCCALLUM &amp; M. JONES" box="[343,548,1515,1538]" pageId="12" pageNumber="13" pagination="427 - 436" refId="ref19198" refString="LACHISH, S., H. MCCALLUM, AND M. JONES. 2009. Demography, disease, and the devil: life-history changes in a disease-affected population of the Tasmanian devil (Sarcophilus harrisii). Journal of Animal Ecology 78: 427 - 436." type="journal article" year="2009">Lachish et al. 2009</bibRefCitation>
). Death (or disappearance) usually follows within 6 months of the appearance of DFTD symptoms. The higher growth rates of young animals at low density are attributable to greater feeding success because of fewer adults at carcasses or less competition for resources in other ways. In response to higher growth rates, some large 1-year-old females have become precocial breeders; thresholds were related to greater head width and heavy (
<quantity id="D0589B7CFFE3E7143F68F9031CE6F92B" box="[612,662,1739,1762]" metricMagnitude="0" metricUnit="kg" metricValue="6.0" pageId="12" pageNumber="13" unit="kg" value="6.0">6 kg</quantity>
) body mass (
<bibRefCitation id="73314B68FFE3E7143D7AF9231F37F8CB" author="LACHISH, S. &amp; H. MCCALLUM &amp; M. JONES" box="[118,327,1771,1794]" pageId="12" pageNumber="13" pagination="427 - 436" refId="ref19198" refString="LACHISH, S., H. MCCALLUM, AND M. JONES. 2009. Demography, disease, and the devil: life-history changes in a disease-affected population of the Tasmanian devil (Sarcophilus harrisii). Journal of Animal Ecology 78: 427 - 436." type="journal article" year="2009">Lachish et al. 2009</bibRefCitation>
). Precocial breeding increased in diseased populations by an average of 16-fold (range 1383%—Jones et al. 2008). Another population response was that diseased females, although having similar litter sizes as healthy females (3.40 versus 3.42), had twice as many female pups compared to healthy females (
<bibRefCitation id="73314B68FFE3E7143C29F8431F9FF86B" author="LACHISH, S. &amp; H. MCCALLUM &amp; M. JONES" box="[293,495,1931,1954]" pageId="12" pageNumber="13" pagination="427 - 436" refId="ref19198" refString="LACHISH, S., H. MCCALLUM, AND M. JONES. 2009. Demography, disease, and the devil: life-history changes in a disease-affected population of the Tasmanian devil (Sarcophilus harrisii). Journal of Animal Ecology 78: 427 - 436." type="journal article" year="2009">Lachish et al. 2009</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE3E7143E7FFF031BB2FCCB" blockId="12.[840,1524,203,1570]" pageId="12" pageNumber="13">
Besides documenting the progression of DFTD across the island, researchers and agencies have formulated plans to preserve disease-free isolated populations, such as those that might be established on nearby islands or on the mainland, i.e., places where healthy
<taxonomicName id="D0A04D1AFFE3E7143EE4FE831A23FEAB" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[1000,1107,331,354]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE3E7143EE4FE831A23FEAB" box="[1000,1107,331,354]" italics="true" pageId="12" pageNumber="13">S. harrisii</emphasis>
</taxonomicName>
might be bred and held for later reintroduction after the disease has run its course (
<bibRefCitation id="73314B68FFE3E7143835FEA31B9BFE4B" author="JONES, M. E. &amp; P. J. JARMAN &amp; C. M. LEES &amp; H. HESTERMAN" box="[1337,1515,363,386]" pageId="12" pageNumber="13" pagination="326 - 337" refId="ref18471" refString="JONES, M. E., P. J. JARMAN, C. M. LEES, AND H. HESTERMAN. 2007. Conservation management of Tasmanian devils in the context of an emerging, extinction-threatening disease: devil facial tumor disease. EcoHealth 4: 326 - 337." type="journal article" year="2007">Jones et al. 2007</bibRefCitation>
; McCallum and Jones 2010). By 2015, captive breeding populations were being studied at 7 secure sites on mainland
<collectingCountry id="6FB77609FFE3E714389EFE631B83FE0B" box="[1426,1523,427,450]" name="Australia" pageId="12" pageNumber="13">Australia</collectingCountry>
and also on Maria Island,
<collectingRegion id="D564F87BFFE3E7143942FE031AC5FE2B" box="[1102,1205,459,482]" country="Australia" name="Tasmania" pageId="12" pageNumber="13">Tasmania</collectingRegion>
, and on the Forestier Peninsula through removal of the original population and introduction of healthy animals, at a time when disease-free populations were still present in the northwest and southwest of the state, and in pockets elsewhere (
<bibRefCitation id="73314B68FFE3E714392DFD831B75FDAB" author="HUXTABLE, S. J. &amp; D. V. LEE &amp; P. WISE &amp; S. T. T. D. PROGRAM" box="[1057,1285,587,610]" pageId="12" pageNumber="13" refId="ref17991" refString="HUXTABLE, S. J., D. V. LEE, P. WISE AND S. T. T. D. PROGRAM. 2015. Metapopulation management of an extreme disease scenario. Pp. 141 - 154 in Advances in reintroduction biology of Australian and New Zealand fauna (D. P. Armstrong, M. W. Hayward, D. Moro, and B. P. Seddon, eds.). CSIRO Publishing, Clayton, Victoria, Australia." type="book" year="2015">Huxtable et al. 2015</bibRefCitation>
). Also, in
<date id="631E1059FFE3E714388EFD831D0FFD4B" pageId="12" pageNumber="13" value="2015-09">September 2015</date>
, 20
<taxonomicName id="D0A04D1AFFE3E7143EBDFDA31A6DFD4B" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[945,1053,619,642]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE3E7143EBDFDA31A6DFD4B" box="[945,1053,619,642]" italics="true" pageId="12" pageNumber="13">S. harrisii</emphasis>
</taxonomicName>
injected with a vaccine derived from heatkilled tumor cells (
<bibRefCitation id="73314B68FFE3E7143902FD431AB4FD6B" author="KREISS, A. &amp; G. K. BROWN &amp; C. TOVAR &amp; A. B. LYONS &amp; G. M. WOODS" box="[1038,1220,651,674]" pageId="12" pageNumber="13" pagination="3016 - 3025" refId="ref19043" refString="KREISS, A., G. K. BROWN, C. TOVAR, A. B. LYONS, AND G. M. WOODS. 2015. Evidence for induction of humoral and cytotoxic immune responses against devil facial tumor disease cells in Tasmanian devils (Sarcophilus harrisii) immunized with killed cell preparations. Vaccine 33: 3016 - 3025." type="journal article" year="2015">Kreiss et al. 2015</bibRefCitation>
) were released in a trial conducted at Narawntapu National Park in northern
<collectingRegion id="D564F87BFFE3E7143854FD631BCFFD0B" box="[1368,1471,683,706]" country="Australia" name="Tasmania" pageId="12" pageNumber="13">Tasmania</collectingRegion>
, and a month later 40 unvaccinated animals raised in captivity were released on the Tasman Peninsula in southeastern
<collectingRegion id="D564F87BFFE3E714385AFD231BCEFCCB" box="[1366,1470,747,770]" country="Australia" name="Tasmania" pageId="12" pageNumber="13">Tasmania</collectingRegion>
.
</paragraph>
<paragraph id="171F3699FFE3E7143E7FFCC31BAAFB2B" blockId="12.[840,1524,203,1570]" pageId="12" pageNumber="13">
If
<taxonomicName id="D0A04D1AFFE3E7143E87FCC31D84FCEB" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[907,1012,779,802]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE3E7143E87FCC31D84FCEB" box="[907,1012,779,802]" italics="true" pageId="12" pageNumber="13">S. harrisii</emphasis>
</taxonomicName>
is reduced to extremely low densities as a result of DFTD, other sources of mortality, which otherwise are not significant, might become important to survival (McCallum and Jones 2006). In such circumstances, mortality factors such as roadkill and persecution for alleged and actual livestock depredations gain additional importance (
<bibRefCitation id="73314B68FFE3E71439AFFC631B5CFC0B" author="GUILER, E. R." box="[1187,1324,939,962]" pageId="12" pageNumber="13" pagination="49 - 62" refId="ref16921" refString="GUILER, E. R. 1970 b. Observations on the Tasmanian devil, Sarcophilus harrisii (Marsupialia: Dasyuridae). I. Numbers, home range, movements, and food in two populations. Australian Journal of Zoology 18: 49 - 62." type="journal article" year="1970">Guiler 1970b</bibRefCitation>
; Jones et al. 2014). Although
<taxonomicName id="D0A04D1AFFE3E7143EBAFC031A53FC2B" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[950,1059,971,994]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE3E7143EBAFC031A53FC2B" box="[950,1059,971,994]" italics="true" pageId="12" pageNumber="13">S. harrisii</emphasis>
</taxonomicName>
comprises only a small proportion of the overall roadkilled wildlife in
<collectingRegion id="D564F87BFFE3E7143973FC231A93FBCB" box="[1151,1251,1003,1026]" country="Australia" name="Tasmania" pageId="12" pageNumber="13">Tasmania</collectingRegion>
(&lt;1%—
<bibRefCitation id="73314B68FFE3E7143844FC231BCEFBCB" author="SHAW, R." box="[1352,1470,1003,1026]" pageId="12" pageNumber="13" refId="ref21230" refString="SHAW, R. 2003. Predicting wildlife roadkill in Tasmania. B. Sc. Honours thesis, University of Tasmania, Hobart, Australia." type="book" year="2003">Shaw 2003</bibRefCitation>
, and 8%—Hobday and Minstrell 2008), local mortality can be significant (e.g., the 50% population decline in 18 months after the paving of a road—Jones 2000).
<taxonomicName id="D0A04D1AFFE3E7143966FB831AA1FBAB" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[1130,1233,1099,1122]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE3E7143966FB831AA1FBAB" box="[1130,1233,1099,1122]" italics="true" pageId="12" pageNumber="13">S. harrisii</emphasis>
</taxonomicName>
, in its nightly forays, moves easily along or is attracted to the roadway to scavenge on roadkilled wildlife. When dazzled by headlights of oncoming vehicles,
<taxonomicName id="D0A04D1AFFE3E7143E44FB631DC3FB0B" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[840,947,1195,1218]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE3E7143E44FB631DC3FB0B" box="[840,947,1195,1218]" italics="true" pageId="12" pageNumber="13">S. harrisii</emphasis>
</taxonomicName>
often reacts unpredictably and so is among the most difficult wildlife for drivers to detect and avoid (
<bibRefCitation id="73314B68FFE3E7143831FB031BBAFB2B" author="HOBDAY, A. J." box="[1341,1482,1227,1250]" pageId="12" pageNumber="13" pagination="265 - 272" refId="ref17543" refString="HOBDAY, A. J. 2010. Night time driver detection distance for Tasmanian fauna: informing speed limits to reduce roadkill. Wildlife Research 37: 265 - 272." type="journal article" year="2010">Hobday 2010</bibRefCitation>
).
</paragraph>
<paragraph id="171F3699FFE3E7143E7FFB231B36F9EB" blockId="12.[840,1524,203,1570]" pageId="12" pageNumber="13">
With almost 100% adult mortality,
<taxonomicName id="D0A04D1AFFE3E714381FFB231BF6FACB" baseAuthorityName="Fleay" baseAuthorityYear="1935" box="[1299,1414,1259,1282]" class="Mammalia" family="Dasyuridae" genus="Sarcophilus" kingdom="Animalia" order="Dasyuromorphia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="harrisii">
<emphasis id="25D4EA8BFFE3E714381FFB231BF6FACB" box="[1299,1414,1259,1282]" italics="true" pageId="12" pageNumber="13">S. harrisii</emphasis>
</taxonomicName>
is under extreme selection pressure to develop resistance to DFTD, to reduce transmission through behavioral mechanisms (
<bibRefCitation id="73314B68FFE3E71438A1FAE31D0DFAAB" author="DEWAR, E." pageId="12" pageNumber="13" refId="ref15607" refString="DEWAR, E. 2013. Understanding behaviour, stress and disease in Tasmanian devils: implications for selective adaptations. Honours thesis, University of Tasmania, Hobart, Australia." type="book" year="2013">Dewar 2013</bibRefCitation>
), or to make such life-history adjustments as breeding at a younger age or producing more female than male offspring. The tumor also is evolving (
<bibRefCitation id="73314B68FFE3E714397BFA431B12FA6B" author="MURCHISON, E. P." box="[1143,1378,1419,1442]" pageId="12" pageNumber="13" pagination="84 - 87" refId="ref20004" refString="MURCHISON, E. P., ET AL. 2010. The Tasmanian devil transcriptome reveals Schwann cell origins of a clonally transmissible cancer. Science 327: 84 - 87." type="journal article" year="2010">Murchison et al. 2010</bibRefCitation>
;
<bibRefCitation id="73314B68FFE3E714387DFA431D0DFA0B" author="PEARSE, A. - M." pageId="12" pageNumber="13" pagination="101 - 112" refId="ref20567" refString="PEARSE, A. - M., ET AL. 2012. Evolution in a transmissible cancer: A study of the chromosomal changes in devil facial tumor (DFT) as it spreads through the wild Tasmanian devil population. Cancer Genetics 205: 101 - 112." type="journal article" year="2012">Pearse et al. 2012</bibRefCitation>
), with tumor lineages being associated with epidemic and demographic effects (
<bibRefCitation id="73314B68FFE3E7143939FA031B7DFA2B" author="HAMEDE, R. K. &amp; A. - M. PEARSE &amp; K. SWIFT &amp; L. A. BARMUTA &amp; E. P. MURCHISON &amp; M. E. JONES" box="[1077,1293,1483,1506]" pageId="12" pageNumber="13" refId="ref17298" refString="HAMEDE, R. K., A. - M. PEARSE, K. SWIFT, L. A. BARMUTA, E. P. MURCHISON, AND M. E. JONES. 2015. Transmissible cancer in Tasmanian devils: localized lineage replacement and host population response. Proceedings of the Royal Society of London, B. Biological Sciences 282.20151468." type="book" year="2015">Hamede et al. 2015</bibRefCitation>
). Rapid evolution of animal and tumor is expected, with DFTD eventually becoming an endemic disease and populations recovering.
</paragraph>
</subSubSection>
</subSection>
</treatment>
</document>