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<mods:title>An investigation into the taxonomy of Abavorana luctuosa (Peters, 1871) (Anura, Ranidae) and the resurrection of Rana decorata Mocquard, 1890 from Borneo</mods:title>
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<mods:namePart>Quah, Evan S. H.</mods:namePart>
<mods:affiliation>Institute of Tropical Biodiversity and Sustainable Development, Universiti Malaysia Terengganu, 21030 Kuala Nerus, Terengganu, Malaysia, https: // orcid. org / 0000 - 0002 - 5357 - 1953 &amp; Herpetology Laboratory, Department of Biology, La Sierra University, Riverside, California, USA, https: // orcid. org / 0000 - 0001 - 8422 - 3698 &amp; Lee Kong Chian Natural History Museum, National University of Singapore, 2 Conservatory Drive, 117377 Singapore, https: // orcid. org / 0000 - 0002 - 0638 - 9198</mods:affiliation>
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<mods:namePart>Grismer, L. Lee</mods:namePart>
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<mods:namePart>Wood Jr., Perry L.</mods:namePart>
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<mods:namePart>Lim, Kelvin K. P.</mods:namePart>
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<mods:namePart>Imbun, Paul Y.</mods:namePart>
<mods:affiliation>Museum of Zoology, Research &amp; Education Division, Sabah Parks, 10626 Kota Kinabalu, Sabah, Malaysia</mods:affiliation>
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<mods:namePart>Anuar, M. S. Shahrul</mods:namePart>
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<taxonomicName LSID="F4AB9D33-A382-5C03-8F5C-4566985CAF35" authority="(Peters, 1871)" baseAuthorityName="Peters" baseAuthorityYear="1871" class="Amphibia" family="Ranidae" genus="Abavorana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Abavorana luctuosa" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="luctuosa">Abavorana luctuosa (Peters, 1871)</taxonomicName>
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<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Holotype of Abavorana luctuosa; A: Original illustration of the holotype of Abavorana luctuosa from Peters (1872). B: Dorsum of the holotype of A. luctuosa (MSNG 29344). C: Venter of the holotype of A. luctuosa (MSNG 29344). Photographs by Giuliano Doria." figureDoi="10.3897/vz.71.e60921.figure3" httpUri="https://binary.pensoft.net/fig/518976" pageId="0" pageNumber="75">Common Mahogany Frog Figs 3</figureCitation>
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Bornean Abavorana luctuosa; Dorsal (A &amp; C) and ventral (B &amp; D) pattern of Abavorana luctuosa from Borneo. A &amp; B: Top row (from left to right); FMNH 234972, FMNH 234973, FMNH 234975 and FMNH 234977 (all specimens from Sipitang District, Sabah). Bottom row (from left to right); FMNH 234980 (Sipitang District, Sabah), FMNH 269142, FMNH 273216 and FMNH 273217 (specimens from Bintulu Division, Sarawak). C &amp; D (from left to right): ZRC 1.11938, ZRC 1.11943 (both from Kubah, Sarawak) and ZRC 1.2270 (unspecified location in Borneo)." figureDoi="10.3897/vz.71.e60921.figure4" httpUri="https://binary.pensoft.net/fig/518977" pageId="0" pageNumber="75">, 4</figureCitation>
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Living specimens of Abavorana decorata and Bornean A. luctuosa; A: Abavorana decorata from Tawau Hills, Sabah, East Malaysia. B: A. decorata from Maliau Basin, Sabah, East Malaysia. C: A. decorata from Kubah National Park, Sarawak, East Malaysia. D: A. decorata from Melawi Regency, West Kalimantan Province, Indonesia. E &amp; F: A. luctuosa from Kubah National Park, Sarawak, East Malaysia. G: A. luctuosa (FMNH 238720) from Sipitang, Sabah, East Malaysia. H: A. luctuosa (FMNH 242883) from Sipitang, Sabah, East Malaysia. (Photographs by Alexander Haas [A, C &amp; E], Paul Bertner [B], Muhammad Aini [D], Chien C. Lee [F], and Robert F. Inger [G &amp; H], Figs G &amp; H © Field Museum of Natural History. Created by Field Museum of Natural History, Amphibian and Reptile Collection and licensed under CC-BY-SA 4.0)." figureDoi="10.3897/vz.71.e60921.figure7" httpUri="https://binary.pensoft.net/fig/518980" pageId="0" pageNumber="75">, 7E-H</figureCitation>
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Peninsular Malaysian, Thai and Sumatran Abavorana luctuosa, and A. nazgul; A: A. luctuosa from Sungai Durian Forest Reserve, Kelantan, Peninsular Malaysia. B: A. luctuosa from Janda Baik, Pahang, Peninsular Malaysia. C: A. luctuosa from Semenyih, Selangor, Peninsular Malaysia. D: A. luctuosa from Fraser's Hill, Pahang, Peninsular Malaysia. E: A. luctuosa from Hala-Bala Wildlife Sanctuary, Thailand. F: A. luctuosa from Sumatra, Indonesia. G: A. nazgul (USMHC 2572) from Gunung Jerai, Kedah, Peninsular Malaysia. H. A. cf. nazgul from Thale Ban National Park, Satun Province, Thailand. (Photographs by Zaharil Dzulkafly [A], Steven Wong [B], Kurt H. P. Guek [C], Evan Quah [D &amp; G], Ian Dugdale [E], Hellen Kurniati [F] &amp; Bunyarit Dechochai [H])." figureDoi="10.3897/vz.71.e60921.figure8" httpUri="https://binary.pensoft.net/fig/518981" pageId="0" pageNumber="75">, 8A-F</figureCitation>
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Colour pattern comparison among species of the genus Abavorana; Dorsum (A, C &amp; E) and venter (B, D &amp; F) in life of A. luctuosa (USMHC 2576) (A &amp; B) from Penang Hill, Penang Island, Malaysia, A. decorata from Maliau Basin, Sabah, East Malaysia, Borneo (C &amp; D) and A. nazgul (USMHC 2233) (E &amp; F) from Gunung Jerai, Kedah, West Malaysia. (Photographs by Evan Quah [A, B, E &amp; F] and Stefan T. Hertwig [C &amp; D])." figureDoi="10.3897/vz.71.e60921.figure9" httpUri="https://binary.pensoft.net/fig/518982" pageId="0" pageNumber="75">, 9A-B</figureCitation>
</paragraph>
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<taxonomicName authorityName="Peters" authorityYear="1871" family="Ranidae" genus="Limnodytes" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Limnodytes luctuosus" order="Anura" pageId="0" pageNumber="75" rank="species" species="luctuosus">Limnodytes luctuosus</taxonomicName>
: 
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:579, 1872:43 &amp; pl. 6; 
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</paragraph>
<paragraph pageId="0" pageNumber="75">
<taxonomicName baseAuthorityName="Peters" baseAuthorityYear="1871" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rana luctuosa" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="luctuosa">Rana luctuosa</taxonomicName>
(in part): 
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<paragraph pageId="0" pageNumber="75">
<taxonomicName baseAuthorityName="Peters" baseAuthorityYear="1871" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rana (Limnodytes) luctuosa" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="luctuosa" subGenus="Limnodytes">Rana (Limnodytes) luctuosa</taxonomicName>
: 
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</paragraph>
<paragraph pageId="0" pageNumber="75">
<taxonomicName baseAuthorityName="Peters" baseAuthorityYear="1871" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rana (Hylorana) luctuosa" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="luctuosa" subGenus="Hylorana">Rana (Hylorana) luctuosa</taxonomicName>
(in part): 
<bibRefCitation DOI="https://doi.org/10.5962/bhl.title.12471" author="Boulenger, GA" journalOrPublisher="Records of the Indian Museum" pageId="0" pageNumber="75" pagination="1 - 226" refId="B6" refString="Boulenger, GA, 1920. A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana. Records of the Indian Museum 20: 1 - 226, DOI: https://doi.org/10.5962/bhl.title.12471" title="A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana." url="https://doi.org/10.5962/bhl.title.12471" volume="20" year="1920">Boulenger 1920</bibRefCitation>
:126.
</paragraph>
<paragraph pageId="0" pageNumber="75">
<taxonomicName baseAuthorityName="Peters" baseAuthorityYear="1871" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rana (Hylarana) luctuosa" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="luctuosa" subGenus="Hylarana">Rana (Hylarana) luctuosa</taxonomicName>
(in part): 
<bibRefCitation DOI="https://doi.org/10.5962/bhl.title.12187" author="Van Kampen, PN" journalOrPublisher="Brill Ltd., Leiden" pageId="0" pageNumber="75" refId="B92" refString="Van Kampen, PN, 1923. The Amphibia of the Indo-Australian Archipelago. E.J. Brill Ltd., Leiden, DOI: https://doi.org/10.5962/bhl.title.12187" title="The Amphibia of the Indo-Australian Archipelago. E. J." url="https://doi.org/10.5962/bhl.title.12187" year="1923">Van Kampen 1923</bibRefCitation>
:196 &amp; 197; 
<bibRefCitation author="Bourret, R" journalOrPublisher="lnstitut Oceanographique de l'lndochine, Hanoi" pageId="0" pageNumber="75" refId="B9" refString="Bourret, R, 1942. Les Batraciens de l'lndochine. lnstitut Oceanographique de l'lndochine, Hanoi" title="Les Batraciens de l'lndochine." year="1942">Bourret 1942</bibRefCitation>
:360; Dubois, 1987: 42.
</paragraph>
<paragraph pageId="0" pageNumber="75">
<taxonomicName baseAuthorityName="Peters" baseAuthorityYear="1871" family="Ranidae" genus="Hylorana" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Hylorana luctuosa" order="Anura" pageId="0" pageNumber="75" rank="species" species="luctuosa">Hylorana luctuosa</taxonomicName>
(in part): 
<bibRefCitation author="Deckert, K" journalOrPublisher="Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin" pageId="0" pageNumber="75" pagination="127 - 184" refId="B24" refString="Deckert, K, 1938. Beitraege zur Osteologie und Systematik ranider Froschlurche. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1938: 127 - 184" title="Beitraege zur Osteologie und Systematik ranider Froschlurche." volume="1938" year="1938">Deckert 1938</bibRefCitation>
:144.
</paragraph>
<paragraph pageId="0" pageNumber="75">
<taxonomicName baseAuthorityName="Peters" baseAuthorityYear="1871" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rana (Pulchrana) luctuosa" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="luctuosa" subGenus="Pulchrana">Rana (Pulchrana) luctuosa</taxonomicName>
(in part): 
<bibRefCitation author="Dubois, A" journalOrPublisher="Bulletin Mensuel de la Societe Linneenne de Lyon" pageId="0" pageNumber="75" pagination="305 - 352" refId="B26" refString="Dubois, A, 1992. Notes sur la classification des Ranidae (Amphibiens anoures). Bulletin Mensuel de la Societe Linneenne de Lyon 61: 305 - 352" title="Notes sur la classification des Ranidae (Amphibiens anoures)." volume="61" year="1992">Dubois 1992</bibRefCitation>
:326.
</paragraph>
<paragraph pageId="0" pageNumber="75">
<taxonomicName baseAuthorityName="Peters" baseAuthorityYear="1871" class="Amphibia" family="Ranidae" genus="Pulchrana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pulchrana luctuosa" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="luctuosa">Pulchrana luctuosa</taxonomicName>
(in part): 
<bibRefCitation DOI="https://doi.org/10.1206/0003-0090(2006)297[0001:tatol]2.0.co;2" author="Frost, DR" journalOrPublisher="Proceedings of the Zoological Society of London" pageId="0" pageNumber="75" refId="B31" refString="Frost, DR, Grant, T, Faivovich, J, Bain, RH, Haas, A, Haddad, CFB, de Sa, RO, Channing, A, Wilkinson, M, Donnellan, SC, Raxworthy, CJ, Campbell, JA, Blotto, BL, Moler, PE, Drewes, RC, Nussbaum, RA, Lynch, JD, Green, DM, Wheeler, WC, 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1-291. https://doi.org/10.1206/0003-0090(2006)297[0001:tatol]2.0.co;2" title="The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1 - 291." url="https://doi.org/10.1206/0003-0090(2006)297[0001:tatol]2.0.co;2" year="2006">Frost et al. 2006</bibRefCitation>
:369; 
<bibRefCitation author="Fei, L" journalOrPublisher="Herpetologica Sinica" pageId="0" pageNumber="75" pagination="1 - 43" refId="B28" refString="Fei, L, Ye, C, Jiang, J, 2010. Phylogenetic systematics of Ranidae. Herpetologica Sinica 12: 1 - 43" title="Phylogenetic systematics of Ranidae." volume="12" year="2010">Fei et al. 2010</bibRefCitation>
:33.
</paragraph>
<paragraph pageId="0" pageNumber="75">
<taxonomicName baseAuthorityName="Peters" baseAuthorityYear="1871" class="Amphibia" family="Ranidae" genus="Hylarana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hylarana luctuosa" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="luctuosa">Hylarana luctuosa</taxonomicName>
(in part): 
<bibRefCitation author="Manthey, U" journalOrPublisher="- herpetofauna, Weinstadt" pageId="0" pageNumber="75" pagination="20 - 31" refId="B62" refString="Manthey, U, 1983. Exkursion am Mt. Kinabalu 4100 m, Nord - Borneo. Teil 3: Checkliste der Herpetofauna oberhalb 600m ueNN. - herpetofauna, Weinstadt 5: 20 - 31" title="Exkursion am Mt. Kinabalu 4100 m, Nord - Borneo. Teil 3: Checkliste der Herpetofauna oberhalb 600 m ueNN." volume="5" year="1983">Manthey 1983</bibRefCitation>
:22; 
<bibRefCitation DOI="https://doi.org/10.1016/j.ympev.2006.11.032" author="Che, J" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="75" pagination="1 - 13" refId="B17" refString="Che, J, Pang, J, Zhao, H, Wu, GF, Zhao, EM, Zhang, YP, 2007. Phylogeny of Raninae (Anura: Ranidae) inferred from mitochondrial and nuclear sequences. Molecular Phylogenetics and Evolution 43: 1 - 13, DOI: https://doi.org/10.1016/j.ympev.2006.11.032" title="Phylogeny of Raninae (Anura: Ranidae) inferred from mitochondrial and nuclear sequences." url="https://doi.org/10.1016/j.ympev.2006.11.032" volume="43" year="2007">Che et al. 2007</bibRefCitation>
: 1-13 (by implication); 
<bibRefCitation author="Das, I" journalOrPublisher="Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut" pageId="0" pageNumber="75" pagination="141 - 174" refId="B23" refString="Das, I, Jankowski, A, Makmor, MIB, Haas, A, 2007. Species diversity, elevational distribution and reproductive modes in an amphibian community at the Matang Range, Sarawak (Borneo). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 104: 141 - 174" title="Species diversity, elevational distribution and reproductive modes in an amphibian community at the Matang Range, Sarawak (Borneo)." volume="104" year="2007">Das et al. 2007</bibRefCitation>
:159, 160, 167, 170, fig. 7d; 
<bibRefCitation author="Imbun, PY" journalOrPublisher="Dewan Bahasa dan Pustaka, Kuala Lumpur" pageId="0" pageNumber="75" refId="B41" refString="Imbun, PY, 2014. Amfibia Sabah. Dewan Bahasa dan Pustaka, Kuala Lumpur" title="Amfibia Sabah." year="2014">Imbun 2014</bibRefCitation>
:99.
</paragraph>
<paragraph pageId="0" pageNumber="75">
<taxonomicName baseAuthorityName="Peters" baseAuthorityYear="1871" class="Amphibia" family="Ranidae" genus="Abavorana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Abavorana luctuosa" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="luctuosa">Abavorana luctuosa</taxonomicName>
: 
<bibRefCitation DOI="https://doi.org/10.1016/j.ympev.2015.05.001" author="Oliver, LA" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="75" pagination="176 - 192" refId="B71" refString="Oliver, LA, Prendini, E, Kraus, F, Raxworthy, CJ, 2015. Systematics and biogeography of the Hylarana frog (Anura: Ranidae) radiation across tropical Australasia, Southeast Asia, and Africa. Molecular Phylogenetics and Evolution 90: 176 - 192, DOI: https://doi.org/10.1016/j.ympev.2015.05.001" title="Systematics and biogeography of the Hylarana frog (Anura: Ranidae) radiation across tropical Australasia, Southeast Asia, and Africa." url="https://doi.org/10.1016/j.ympev.2015.05.001" volume="90" year="2015">Oliver et al. 2015</bibRefCitation>
:186 (in part); 
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.4320.2.4" author="Quah, ESH" journalOrPublisher="Zootaxa" pageId="0" pageNumber="75" pagination="272 - 288" refId="B75" refString="Quah, ESH, Shahrul, AMS, Grismer, LL, Wood, JrPL, Siti, AMN, Muin, MA, 2017. A new species of frog of the genus Abavorana Oliver, Prendini, Kraus &amp; Raxworthy 2015 (Anura: Ranidae) from Gunung Jerai, Kedah, northwestern Peninsular Malaysia. Zootaxa 4320: 272 - 288, DOI: https://doi.org/10.11646/zootaxa.4320.2.4" title="A new species of frog of the genus Abavorana Oliver, Prendini, Kraus &amp; Raxworthy 2015 (Anura: Ranidae) from Gunung Jerai, Kedah, northwestern Peninsular Malaysia." url="https://doi.org/10.11646/zootaxa.4320.2.4" volume="4320" year="2017">Quah et al. 2017</bibRefCitation>
: 272-288 (in part); 
<bibRefCitation author="Zainudin, R" journalOrPublisher="Turkish Journal of Zoology" pageId="0" pageNumber="75" pagination="876 - 891" refId="B93" refString="Zainudin, R, Zain, BMM, Ahmad, N, Nor, SM, 2017. Microhabitat partitioning of closely related Sarawak (Malaysian Borneo) frog species previously assigned to the genus Hylarana (Amphibia: Anura). Turkish Journal of Zoology 41: 876 - 891" title="Microhabitat partitioning of closely related Sarawak (Malaysian Borneo) frog species previously assigned to the genus Hylarana (Amphibia: Anura)." volume="41" year="2017">Zainudin et al. 2017</bibRefCitation>
:876-891 (in part); 
<bibRefCitation DOI="https://doi.org/10.15560/15.6.1055" author="Chan, KO" journalOrPublisher="Check List" pageId="0" pageNumber="75" pagination="1055 - 1069" refId="B16" refString="Chan, KO, Muin, MA, Anuar, S, Andam, J, Razak, N, Aziz, MA, 2019. First checklist on the amphibians and reptiles of Mount Korbu, the second highest peak in Peninsular Malaysia. Check List 15: 1055 - 1069, DOI: https://doi.org/10.15560/15.6.1055" title="First checklist on the amphibians and reptiles of Mount Korbu, the second highest peak in Peninsular Malaysia." url="https://doi.org/10.15560/15.6.1055" volume="15" year="2019">Chan et al. 2019</bibRefCitation>
:1057, 1062.
</paragraph>
<paragraph pageId="0" pageNumber="75">
<taxonomicName baseAuthorityName="Peters" baseAuthorityYear="1871" class="Amphibia" family="Ranidae" genus="Hylarana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hylarana (Abavorana) luctuosa" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="luctuosa" subGenus="Abavorana">Hylarana (Abavorana) luctuosa</taxonomicName>
(in part): 
<bibRefCitation author="Inger, RF" journalOrPublisher=", Kota Kinabalu, Sabah" pageId="0" pageNumber="75" refId="B48" refString="Inger, RF, Stuebing, RB, Grafe, TU, Dehling, JM, 2017. A Field Guide to the Frogs of Borneo. 3rd Edition. Natural History Publications (Borneo) Sdn Bhd., Kota Kinabalu, Sabah" title="A Field Guide to the Frogs of Borneo. 3 rd Edition. Natural History Publications (Borneo) Sdn Bhd." year="2017">Inger et al. 2017</bibRefCitation>
:147 &amp; 148.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="75" type="holotype">
<paragraph pageId="0" pageNumber="75">Holotype.</paragraph>
<paragraph pageId="0" pageNumber="75">
Adult female (MSNG 29344) collected by Giacomo Doria and Odoardo Beccari from 
<normalizedToken originalValue="“Sarawak”">&quot;Sarawak&quot;</normalizedToken>
, (Borneo) Malaysia (
<bibRefCitation author="Capocaccia, L" journalOrPublisher="Annali del Museo Civico di Storia Naturale di Genova" pageId="0" pageNumber="75" pagination="208 - 222" refId="B12" refString="Capocaccia, L, 1957. Catalogo dei tipi di anfibi del Museo Civico di Storia Naturale di Genova. Annali del Museo Civico di Storia Naturale di Genova 3: 208 - 222" title="Catalogo dei tipi di anfibi del Museo Civico di Storia Naturale di Genova." volume="3" year="1957">Capocaccia 1957</bibRefCitation>
) (Fig. 
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Holotype of Abavorana luctuosa; A: Original illustration of the holotype of Abavorana luctuosa from Peters (1872). B: Dorsum of the holotype of A. luctuosa (MSNG 29344). C: Venter of the holotype of A. luctuosa (MSNG 29344). Photographs by Giuliano Doria." figureDoi="10.3897/vz.71.e60921.figure3" httpUri="https://binary.pensoft.net/fig/518976" pageId="0" pageNumber="75">3</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="75" type="diagnosis">
<paragraph pageId="0" pageNumber="75">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="75">
Body robust, medium-sized; head moderate; snout short, rounded, canthus rostralis smoothly rounded; interorbital space broader than the upper eyelid; tympanum distinct, not quite two-thirds the size of the eye with no pale colouration on the margins of the tympanum; no vocal sacs in males; vomerine teeth in two small oblique patches on a level with the posterior edge of the choanae; length of 1st finger greater than 2nd finger; disc width to finger width ratios of finger 3 and toe 4 is 1-1.5; dorsolateral fold indistinct or absent; the humeral gland in males is prominent, raised and centrally positioned on the ventral surface of the upper arm; a weak or absent rictal ridge; outer metatarsal tubercle weak or absent; subarticular tubercles moderate; skin of dorsum smooth or finely shagreened; throat, abdomen, and flanks smooth; posterior section of venter and back of the thigh rugose; dorsum reddish-orange to chocolate-brown, encircled by a white or cream coloured dorsolateral line that encircles the snout, canthus rostralis, outer edge of the upper eyelids, and dorsum along the dorsolateral fold to the vent; lower flanks dark-brown or black below the dorsolateral line grading into a paler venter; dorsal colouration of the limbs same as the flanks with whitish or light-grey speckles or stripes. 
<taxonomicName baseAuthorityName="Peters" baseAuthorityYear="1871" class="Amphibia" family="Ranidae" genus="Abavorana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Abavorana luctuosa" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="luctuosa">
<emphasis italics="true" pageId="0" pageNumber="75">Abavorana luctuosa</emphasis>
</taxonomicName>
can be easily differentiated from its congeners on the basis of its ventral colour pattern which is usually immaculate or with only very faint, sparse, light speckling (Figs 
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Holotype of Abavorana luctuosa; A: Original illustration of the holotype of Abavorana luctuosa from Peters (1872). B: Dorsum of the holotype of A. luctuosa (MSNG 29344). C: Venter of the holotype of A. luctuosa (MSNG 29344). Photographs by Giuliano Doria." figureDoi="10.3897/vz.71.e60921.figure3" httpUri="https://binary.pensoft.net/fig/518976" pageId="0" pageNumber="75">3</figureCitation>
, 
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Bornean Abavorana luctuosa; Dorsal (A &amp; C) and ventral (B &amp; D) pattern of Abavorana luctuosa from Borneo. A &amp; B: Top row (from left to right); FMNH 234972, FMNH 234973, FMNH 234975 and FMNH 234977 (all specimens from Sipitang District, Sabah). Bottom row (from left to right); FMNH 234980 (Sipitang District, Sabah), FMNH 269142, FMNH 273216 and FMNH 273217 (specimens from Bintulu Division, Sarawak). C &amp; D (from left to right): ZRC 1.11938, ZRC 1.11943 (both from Kubah, Sarawak) and ZRC 1.2270 (unspecified location in Borneo)." figureDoi="10.3897/vz.71.e60921.figure4" httpUri="https://binary.pensoft.net/fig/518977" pageId="0" pageNumber="75">4</figureCitation>
, 
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Colour pattern comparison among species of the genus Abavorana; Dorsum (A, C &amp; E) and venter (B, D &amp; F) in life of A. luctuosa (USMHC 2576) (A &amp; B) from Penang Hill, Penang Island, Malaysia, A. decorata from Maliau Basin, Sabah, East Malaysia, Borneo (C &amp; D) and A. nazgul (USMHC 2233) (E &amp; F) from Gunung Jerai, Kedah, West Malaysia. (Photographs by Evan Quah [A, B, E &amp; F] and Stefan T. Hertwig [C &amp; D])." figureDoi="10.3897/vz.71.e60921.figure9" httpUri="https://binary.pensoft.net/fig/518982" pageId="0" pageNumber="75">9A-B</figureCitation>
). Adult males with SVL 35.05-44.27 mm, adult females with SVL 37.05-51.06 mm; adult males with SW 7.11-8.62 mm, adult females with SW 7.65-9.38 mm; adult males with IOD 3.25-5.14 mm, adult females with IOD 3.80-5.64 mm; adult males with TD 2.50-3.83 mm, adult females with TD 2.95-4.24 mm; adult males with FL 17.81-22.04 mm, adult females with FL 19.13-26.37 mm (Table 
<tableCitation captionStart="Table 5" captionStartId="T5" captionText="Table 5. Summary statistics and ventral colour pattern among members of the genus Abavorana. Summary statistics and ventral colour pattern for Abavorana decorata, A. nazgul, and A. luctuosa populations from the Malay Peninsula and Borneo, SD = standard deviation, N = sample size, / = no data. Humeral gland only present in males and all measurements in mm." httpUri="http://table.plazi.org/id/B91F734E592A25063FCA3E8C57D388D3" pageId="0" pageNumber="75" tableUuid="B91F734E592A25063FCA3E8C57D388D3">5</tableCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="75" type="distribution">
<paragraph pageId="0" pageNumber="75">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="75">
Its distribution spans southern Peninsular Thailand, Peninsular Malaysia, Borneo, and Sumatra (Manthey &amp; Grossmann 1997; 
<bibRefCitation DOI="https://doi.org/10.1016/j.ympev.2015.05.001" author="Oliver, LA" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="75" pagination="176 - 192" refId="B71" refString="Oliver, LA, Prendini, E, Kraus, F, Raxworthy, CJ, 2015. Systematics and biogeography of the Hylarana frog (Anura: Ranidae) radiation across tropical Australasia, Southeast Asia, and Africa. Molecular Phylogenetics and Evolution 90: 176 - 192, DOI: https://doi.org/10.1016/j.ympev.2015.05.001" title="Systematics and biogeography of the Hylarana frog (Anura: Ranidae) radiation across tropical Australasia, Southeast Asia, and Africa." url="https://doi.org/10.1016/j.ympev.2015.05.001" volume="90" year="2015">Oliver et al. 2015</bibRefCitation>
; 
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.4320.2.4" author="Quah, ESH" journalOrPublisher="Zootaxa" pageId="0" pageNumber="75" pagination="272 - 288" refId="B75" refString="Quah, ESH, Shahrul, AMS, Grismer, LL, Wood, JrPL, Siti, AMN, Muin, MA, 2017. A new species of frog of the genus Abavorana Oliver, Prendini, Kraus &amp; Raxworthy 2015 (Anura: Ranidae) from Gunung Jerai, Kedah, northwestern Peninsular Malaysia. Zootaxa 4320: 272 - 288, DOI: https://doi.org/10.11646/zootaxa.4320.2.4" title="A new species of frog of the genus Abavorana Oliver, Prendini, Kraus &amp; Raxworthy 2015 (Anura: Ranidae) from Gunung Jerai, Kedah, northwestern Peninsular Malaysia." url="https://doi.org/10.11646/zootaxa.4320.2.4" volume="4320" year="2017">Quah et al. 2017</bibRefCitation>
). Within Borneo it is confirmed from Sabah state: Imbak Canyon and Sipitang; Sarawak state: Bukit Kana, Samarakan and Tubau in Bintulu Division and Kubah National Park where it is sympatric with 
<taxonomicName authorityName="comb. nov. (Mocquard" authorityYear="1890" class="Amphibia" family="Ranidae" genus="Abavorana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Abavorana decorata" order="Anura" pageId="0" pageNumber="75" phylum="Chordata" rank="species" species="decorata">
<emphasis italics="true" pageId="0" pageNumber="75">Abavorana decorata</emphasis>
</taxonomicName>
(Fig. 
<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Distribution map of Abavorana decorata and A. luctuosa in Borneo; Known distribution of Abavorana decorata (red) and A. luctuosa (blue) in Borneo. Star = type locality." figureDoi="10.3897/vz.71.e60921.figure10" httpUri="https://binary.pensoft.net/fig/518983" pageId="0" pageNumber="75">10</figureCitation>
). The species is expected to range across Borneo.
</paragraph>
</subSubSection>
</treatment>
</document>