treatments-xml/data/DF/1D/30/DF1D302EFF90FFA2FF33D017FB02FEEE.xml
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<mods:title id="D2535BC6DD85CD59301060155678B5A3">Taxonomic revision of south-eastern Australian giant burrowing frogs (Anura: Limnodynastidae: Heleioporus Gray)</mods:title>
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<mods:namePart id="7B2838084BEB3702B4A44C4C44FA4F3D">Mahony, Michael J.</mods:namePart>
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<mods:affiliation id="7D921E937B93947EF2F736C5ED74E0B5">Department of Biological Sciences, The University of Newcastle, University Drive, Callaghan, NSW 2308, Australia. Michael. mahony @ newcastle. edu. au; https: // orcid. org / 0000 - 0002 - 1042 - 0848</mods:affiliation>
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<mods:affiliation id="87EDA73551746FEB97B666AC8285E3D9">Research and Development Division, Forests New South Wales, P. O Box 100 Beecroft, NSW 2119, Australia. &amp; Current address: Department of Forest and Ecosystem Science, University of Melbourne, 4 Water Street, Creswick, Victoria 3363, Australia. https: // orcid. org / 0000 - 0002 - 5203 - 9818</mods:affiliation>
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<mods:affiliation id="E3B3578ACCAC8E7E69B83198FE757269">The South Australian Museum, North Terrace Adelaide, SA 5000, Australia. &amp; School of Biological Sciences, The University of Adelaide, Adelaide SA 5005, Australia.</mods:affiliation>
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<mods:affiliation id="FCF0DF77745CD72673F7E048388B4290">Australian Museum Research Institute, Australian Museum, 1 William Street Sydney NSW 2010, Australia; Eco Logical Australia, Level 7, 19 Bolton Street, Newcastle, NSW 2300, Australia.</mods:affiliation>
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<mods:namePart id="5E6821A1EEA1EE3556F11C2BEA5C55FF">Bilney, Rohan</mods:namePart>
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<mods:namePart id="2554FA705E4B97177BF442F971046388">Donnellan, Stephen C.</mods:namePart>
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<mods:affiliation id="B7D56B9857823B69A3CEAEA39918C75C">The South Australian Museum, North Terrace Adelaide, SA 5000, Australia. &amp; Steve. Donnellan @ samuseum. sa. gov. au; https: // orcid. org / 0000 - 0002 - 5448 - 3226</mods:affiliation>
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<mods:date id="CC02418F616B9ECDDBBC843A84C78868">2021</mods:date>
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<treatment id="DF1D302EFF90FFA2FF33D017FB02FEEE" ID-DOI="http://doi.org/10.5281/zenodo.5222429" ID-Zenodo-Dep="5222429" LSID="urn:lsid:plazi:treatment:DF1D302EFF90FFA2FF33D017FB02FEEE" httpUri="http://treatment.plazi.org/id/DF1D302EFF90FFA2FF33D017FB02FEEE" lastPageId="25" lastPageNumber="475" pageId="24" pageNumber="475">
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<paragraph id="570B8138FF90FFA3FF33D017FD08FA93" blockId="24.[151,755,1350,1377]" box="[151,755,1350,1377]" pageId="24" pageNumber="475">
<heading id="0C433654FF90FFA3FF33D017FD08FA93" bold="true" box="[151,755,1350,1377]" fontSize="11" level="1" pageId="24" pageNumber="475" reason="1">
<taxonomicName id="90B4FABBFF90FFA3FF33D017FD08FA93" ID-CoL="3K4FC" authority="(Shaw &amp; Nodder, 1795)" baseAuthorityName="Shaw &amp; Nodder" baseAuthorityYear="1795" box="[151,755,1350,1377]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="australiacus">
<emphasis id="65C05D2AFF90FFA3FF33D017FD08FA93" bold="true" box="[151,755,1350,1377]" pageId="24" pageNumber="475">
<emphasis id="65C05D2AFF90FFA3FF33D017FE3CFA92" bold="true" box="[151,455,1350,1377]" italics="true" pageId="24" pageNumber="475">Heleioporus australiacus</emphasis>
(
<bibRefCitation id="3325FCC9FF90FFA3FE73D017FD11FA93" author="Shaw, G. &amp; Nodder, F." box="[471,746,1350,1377]" pageId="24" pageNumber="475" pagination="200" refId="ref25595" refString="Shaw, G. &amp; Nodder, F. (1795) Rana australiaca. Naturalist's Miscellany London, 6, 200." type="journal article" year="1795">Shaw &amp; Nodder, 1795</bibRefCitation>
)
</emphasis>
</taxonomicName>
</heading>
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<subSubSection id="1FAED2B3FF90FFA2FF33D0DFFB02FEEE" lastPageId="25" lastPageNumber="476" pageId="24" pageNumber="475" type="diagnosis">
<paragraph id="570B8138FF90FFA3FF33D0DFFDCAF8C5" blockId="24.[151,1437,1422,2025]" pageId="24" pageNumber="475">
<emphasis id="65C05D2AFF90FFA3FF33D0DFFEF1FA5B" bold="true" box="[151,266,1422,1448]" pageId="24" pageNumber="475">Diagnosis</emphasis>
. Assigned to
<taxonomicName id="90B4FABBFF90FFA3FE06D0DEFDD7FA5A" authorityName="Gray" authorityYear="1841" box="[418,556,1423,1449]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="genus">
<emphasis id="65C05D2AFF90FFA3FE06D0DEFDD7FA5A" box="[418,556,1423,1449]" italics="true" pageId="24" pageNumber="475">Heleioporus</emphasis>
</taxonomicName>
based on the generic definition of
<bibRefCitation id="3325FCC9FF90FFA3FC15D0DEFBD7FA59" author="Lee, A." box="[945,1068,1423,1450]" pageId="24" pageNumber="475" pagination="367 - 439" refId="ref23752" refString="Lee, A. (1967) Studies in Australian amphibia II. Taxonomy, ecology and evolution of the genus Heleioporus Gray (Anura: Leptodactylidae). Australian Journal of Zoology, 15, 367 - 439. https: // doi. org / 10.1071 / ZO 9670367" type="journal article" year="1967">Lee (1967)</bibRefCitation>
and molecular phylogenetic evidence (
<bibRefCitation id="3325FCC9FF90FFA3FF4CD0E2FE31FA3E" author="Morgan, M. J. &amp; Roberts, J. D. &amp; Keogh, J. S." box="[232,458,1459,1485]" pageId="24" pageNumber="475" pagination="371 - 385" refId="ref24538" refString="Morgan, M. J., Roberts, J. D. &amp; Keogh, J. S. (2007) Molecular phylogenetic dating supports an ancient endemic speciation model in Australia's biodiversity hotspot. Molecular Phylogenetics and Evolution, 44, 371 - 385. https: // doi. org / 10.1016 / j. ympev. 2006.12.009" type="journal article" year="2007">
Morgan
<emphasis id="65C05D2AFF90FFA3FEEFD0E2FE85FA3E" box="[331,382,1459,1485]" italics="true" pageId="24" pageNumber="475">et al</emphasis>
. 2007
</bibRefCitation>
). Eastern Australian
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<emphasis id="65C05D2AFF90FFA3FD12D0E2FCBBFA3E" box="[694,832,1459,1485]" italics="true" pageId="24" pageNumber="475">Heleioporus</emphasis>
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are phylogenetically quite distant from their western Australian congeners with divergence estimated in the late Oligocene to early Miocene (
<bibRefCitation id="3325FCC9FF90FFA3FBD7D086FAABFA02" author="Morgan, M. J. &amp; Roberts, J. D. &amp; Keogh, J. S." box="[1139,1360,1495,1521]" pageId="24" pageNumber="475" pagination="371 - 385" refId="ref24538" refString="Morgan, M. J., Roberts, J. D. &amp; Keogh, J. S. (2007) Molecular phylogenetic dating supports an ancient endemic speciation model in Australia's biodiversity hotspot. Molecular Phylogenetics and Evolution, 44, 371 - 385. https: // doi. org / 10.1016 / j. ympev. 2006.12.009" type="journal article" year="2007">
Morgan
<emphasis id="65C05D2AFF90FFA3FB70D086FAFDFA02" box="[1236,1286,1495,1521]" italics="true" pageId="24" pageNumber="475">et al</emphasis>
. 2007
</bibRefCitation>
). Eastern Australian
<taxonomicName id="90B4FABBFF90FFA3FE98D0AAFE3DF9E6" authorityName="Gray" authorityYear="1841" box="[316,454,1531,1557]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="genus">
<emphasis id="65C05D2AFF90FFA3FE98D0AAFE3DF9E6" box="[316,454,1531,1557]" italics="true" pageId="24" pageNumber="475">Heleioporus</emphasis>
</taxonomicName>
are readily diagnosed morphologically from their western congeners by the presence of black spines capping the tubercles on the dorsum, sides, and throat, and the uniform chocolate dorsum with yellow or white spots restricted to the sides separating
<taxonomicName id="90B4FABBFF90FFA3FD1AD312FC96F9AE" baseAuthorityName="Shaw &amp; Nodder" baseAuthorityYear="1795" box="[702,877,1603,1629]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="australiacus">
<emphasis id="65C05D2AFF90FFA3FD1AD312FC96F9AE" box="[702,877,1603,1629]" italics="true" pageId="24" pageNumber="475">H. australiacus</emphasis>
</taxonomicName>
from all other species of
<taxonomicName id="90B4FABBFF90FFA3FB3ED312FADFF9AE" authorityName="Gray" authorityYear="1841" box="[1178,1316,1603,1629]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="genus">
<emphasis id="65C05D2AFF90FFA3FB3ED312FADFF9AE" box="[1178,1316,1603,1629]" italics="true" pageId="24" pageNumber="475">Heleioporus</emphasis>
</taxonomicName>
except
<taxonomicName id="90B4FABBFF90FFA3FA25D312FEECF972" authorityName="Lee" authorityYear="1967" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="barycragus">
<emphasis id="65C05D2AFF90FFA3FA25D312FEECF972" italics="true" pageId="24" pageNumber="475">H. barycragus</emphasis>
</taxonomicName>
. The length of the metatarsal tubercle distinguishes
<taxonomicName id="90B4FABBFF90FFA3FCFBD336FBF0F972" baseAuthorityName="Shaw &amp; Nodder" baseAuthorityYear="1795" box="[863,1035,1639,1665]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="australiacus">
<emphasis id="65C05D2AFF90FFA3FCFBD336FBF0F972" box="[863,1035,1639,1665]" italics="true" pageId="24" pageNumber="475">H. australiacus</emphasis>
</taxonomicName>
(less than one half the length of the fourth toe) from
<taxonomicName id="90B4FABBFF90FFA3FEF1D3DAFE03F956" authorityName="Lee" authorityYear="1967" box="[341,504,1675,1701]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="barycragus">
<emphasis id="65C05D2AFF90FFA3FEF1D3DAFE03F956" box="[341,504,1675,1701]" italics="true" pageId="24" pageNumber="475">H. barycragus</emphasis>
</taxonomicName>
(at least one-half the length of the fourth toe) (
<bibRefCitation id="3325FCC9FF90FFA3FBAAD3DAFB86F955" author="Lee, A." box="[1038,1149,1675,1702]" pageId="24" pageNumber="475" pagination="367 - 439" refId="ref23752" refString="Lee, A. (1967) Studies in Australian amphibia II. Taxonomy, ecology and evolution of the genus Heleioporus Gray (Anura: Leptodactylidae). Australian Journal of Zoology, 15, 367 - 439. https: // doi. org / 10.1071 / ZO 9670367" type="journal article" year="1967">Lee 1967</bibRefCitation>
). The advertisement calls of
<taxonomicName id="90B4FABBFF90FFA3FF12D3FEFEA3F93A" authorityName="Lee" authorityYear="1967" box="[182,344,1711,1737]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="barycragus">
<emphasis id="65C05D2AFF90FFA3FF12D3FEFEA3F93A" box="[182,344,1711,1737]" italics="true" pageId="24" pageNumber="475">H. barycragus</emphasis>
</taxonomicName>
consist of a single note compared with multi-pulses in eastern
<taxonomicName id="90B4FABBFF90FFA3FBBFD3FEFB5EF93A" authorityName="Gray" authorityYear="1841" box="[1051,1189,1711,1737]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="genus">
<emphasis id="65C05D2AFF90FFA3FBBFD3FEFB5EF93A" box="[1051,1189,1711,1737]" italics="true" pageId="24" pageNumber="475">Heleioporus</emphasis>
</taxonomicName>
(FrogID application). Contra to
<bibRefCitation id="3325FCC9FF90FFA3FEA2D382FE79F91D" author="Lee, A." box="[262,386,1747,1774]" pageId="24" pageNumber="475" pagination="367 - 439" refId="ref23752" refString="Lee, A. (1967) Studies in Australian amphibia II. Taxonomy, ecology and evolution of the genus Heleioporus Gray (Anura: Leptodactylidae). Australian Journal of Zoology, 15, 367 - 439. https: // doi. org / 10.1071 / ZO 9670367" type="journal article" year="1967">Lee (1967)</bibRefCitation>
, the number and distribution of the pre-orbital papillae in the anterior corner of the eye is quite variable in
<taxonomicName id="90B4FABBFF90FFA3FEBDD3A6FE3CF8E2" baseAuthorityName="Shaw &amp; Nodder" baseAuthorityYear="1795" box="[281,455,1783,1809]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="australiacus">
<emphasis id="65C05D2AFF90FFA3FEBDD3A6FE3CF8E2" box="[281,455,1783,1809]" italics="true" pageId="24" pageNumber="475">H. australiacus</emphasis>
</taxonomicName>
(
<figureCitation id="CF8F9DBDFF90FFA3FE7DD3A6FDE6F8E2" box="[473,541,1783,1809]" captionStart="FIGURE 8" captionStartId="19.[151,250,1901,1925]" captionTargetBox="[235,1353,181,1877]" captionTargetId="figure-17@19.[235,1353,181,1877]" captionTargetPageId="19" captionText="FIGURE 8. Variation in the pre-orbital papillae in anterior corner of eye of eastern Australian Heleioporus. Heleioporus a. australiacus—A) AMS R104952 (neotype) Maroota State Forest, B) AMS R5958 North Sydney, C) AMS R9470 Wollemi National Park, D) AMS R78853 Commodore Heights. H. a. flavopunctatus subsp. nov.—E) AMS R136719 Coolangubra State Forest, F) ANWC A0357 Wadbilliga, G) AMS R136730 Coolangubra State Forest, H) ANWC A1504 Bondi State Forest." figureDoi="http://doi.org/10.5281/zenodo.5222427" httpUri="https://zenodo.org/record/5222427/files/figure.png" pageId="24" pageNumber="475">Fig. 8</figureCitation>
) and includes exemplars that closely resemble the condition in
<taxonomicName id="90B4FABBFF90FFA3FB5DD3A6FA66F8E2" authorityName="Lee" authorityYear="1967" box="[1273,1437,1783,1809]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="barycragus">
<emphasis id="65C05D2AFF90FFA3FB5DD3A6FA66F8E2" box="[1273,1437,1783,1809]" italics="true" pageId="24" pageNumber="475">H. barycragus</emphasis>
</taxonomicName>
illustrated in
<figureCitation id="CF8F9DBDFF90FFA3FE8FD24AFE72F8C6" box="[299,393,1818,1845]" captionStart="FIGURE 5" captionStartId="13.[151,250,1959,1983]" captionTargetBox="[151,1435,633,1936]" captionTargetId="figure-244@13.[151,1435,633,1936]" captionTargetPageId="13" captionText="FIGURE 5. Histograms of linear discriminant (LD) for analyses of morphological variation on transformed morphological data for female and male H. a. australiacus (red), and H. a. flavopunctatus subsp. nov. (blue)." figureDoi="http://doi.org/10.5281/zenodo.5222421" httpUri="https://zenodo.org/record/5222421/files/figure.png" pageId="24" pageNumber="475">Figure 5</figureCitation>
of
<bibRefCitation id="3325FCC9FF90FFA3FE0AD24AFDD6F8C5" author="Lee, A." box="[430,557,1819,1846]" pageId="24" pageNumber="475" pagination="367 - 439" refId="ref23752" refString="Lee, A. (1967) Studies in Australian amphibia II. Taxonomy, ecology and evolution of the genus Heleioporus Gray (Anura: Leptodactylidae). Australian Journal of Zoology, 15, 367 - 439. https: // doi. org / 10.1071 / ZO 9670367" type="journal article" year="1967">Lee (1967)</bibRefCitation>
.
</paragraph>
<paragraph id="570B8138FF90FFA2FF63D26EFB02FEEE" blockId="24.[151,1437,1422,2025]" lastBlockId="25.[151,1437,151,285]" lastPageId="25" lastPageNumber="476" pageId="24" pageNumber="475">
<taxonomicName id="90B4FABBFF90FFA3FF63D26EFE1AF8AA" baseAuthorityName="Shaw &amp; Nodder" baseAuthorityYear="1795" box="[199,481,1855,1881]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="australiacus">
<emphasis id="65C05D2AFF90FFA3FF63D26EFE1AF8AA" box="[199,481,1855,1881]" italics="true" pageId="24" pageNumber="475">Heleioporus australiacus</emphasis>
</taxonomicName>
comprises two sub-species-
<emphasis id="65C05D2AFF90FFA3FCBAD26EFC1FF8AA" box="[798,996,1855,1881]" italics="true" pageId="24" pageNumber="475">
H. a.
<taxonomicName id="90B4FABBFF90FFA3FCFED26EFC1FF8AA" authorityName="Mahony &amp; Penman &amp; Bertozzi &amp; Lemckert &amp; Bilney &amp; Donnellan" authorityYear="2021" box="[858,996,1855,1881]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="australiacus" status="comb. nov.">australiacus</taxonomicName>
</emphasis>
<emphasis id="65C05D2AFF90FFA3FC4ED26EFB4EF8AB" bold="true" box="[1002,1205,1854,1881]" pageId="24" pageNumber="475">
<taxonomicNameLabel id="7EF3E051FF90FFA3FC4ED26EFB4EF8AB" box="[1002,1205,1854,1881]" pageId="24" pageNumber="475" rank="species">new combination</taxonomicNameLabel>
</emphasis>
and
<emphasis id="65C05D2AFF90FFA3FB4DD26EFF36F88F" italics="true" pageId="24" pageNumber="475">H. a. flavopunctatus</emphasis>
<emphasis id="65C05D2AFF90FFA3FF70D232FEADF88E" bold="true" box="[212,342,1891,1917]" pageId="24" pageNumber="475">subsp. nov.</emphasis>
which closely resemble each other in most morphological features but can be diagnosed from each other by two features in combination as follows. In
<emphasis id="65C05D2AFF90FFA3FD73D2D6FC64F852" box="[727,927,1927,1953]" italics="true" pageId="24" pageNumber="475">
H. a.
<taxonomicName id="90B4FABBFF90FFA3FCB1D2D6FC64F852" authorityName="Mahony &amp; Penman &amp; Bertozzi &amp; Lemckert &amp; Bilney &amp; Donnellan" authorityYear="2021" box="[789,927,1927,1953]" class="Amphibia" family="Limnodynastidae" genus="Heleioporus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="australiacus">australiacus</taxonomicName>
</emphasis>
the tubercles on the flanks have small central white- and yellow-coloured spots which rarely cover the entire tubercle or coalesce with adjacent spots; yellow or white spots occur on the raised tubercles surrounding the cloaca, but they do not coalesce to form a complete ring. In
<emphasis id="65C05D2AFF91FFA2FF13D5C6FE66FF42" box="[183,413,151,177]" italics="true" pageId="25" pageNumber="476">H. a. flavopunctatus</emphasis>
tubercles on the flanks have cream and yellow-coloured spots that coalesce with adjacent spots; yellow or white spots on raised tubercles coalesce to from an almost complete ring surrounding the cloaca. Higher mean number of pulses of the note in the advertisement call of
<emphasis id="65C05D2AFF91FFA2FC15D58EFB6FFF0A" box="[945,1172,223,249]" italics="true" pageId="25" pageNumber="476">H. a. flavopunctatus</emphasis>
. Apomorphic character states for at least 16 nucleotide sites in the mitochondrial
<emphasis id="65C05D2AFF91FFA2FCBFD452FCABFEEE" box="[795,848,259,285]" italics="true" pageId="25" pageNumber="476">ND4</emphasis>
gene diagnose the two taxa (
<tableCitation id="1A36B483FF91FFA2FB31D452FB17FEEE" box="[1173,1260,259,285]" captionStart="TABLE 2" captionStartId="9.[151,239,908,932]" captionText="TABLE 2. Net average sequence divergence between lineages (dA). Values in bold are sister taxon comparisons." pageId="25" pageNumber="476">Table 2</tableCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>