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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.109.1439" ID-PMC="PMC3118819" ID-Pensoft-Pub="1313-2970-109-19" ID-Pensoft-UUID="FFDE6B4A96644D30FFD8FFEA7F28FFF8" ID-PubMed="21852932" ID-Zenodo-Dep="577024" ModsDocID="1313-2970-109-19" checkinTime="1451250264676" checkinUser="pensoft" docAuthor="Kaiser, Hinrich, Carvalho, Venancio Lopes, Ceballos, Jester, Freed, Paul, Heacox, Scott, Lester, Barbara, Richards, Stephen J., Trainor, Colin R., Sanchez, Caitlin &amp; O'Shea, Mark" docDate="2011" docId="CA119BC729FEFFAEC39252B82BF98D8F" docLanguage="en" docName="ZooKeys 109: 19-86" docOrigin="ZooKeys 109" docPubDate="2011-06-20" docSource="http://dx.doi.org/10.3897/zookeys.109.1439" docTitle="Fejervarya Bolkay 1915" docType="treatment" docVersion="5" id="FFDE6B4A96644D30FFD8FFEA7F28FFF8" lastPageNumber="29" masterDocId="FFDE6B4A96644D30FFD8FFEA7F28FFF8" masterDocTitle="The herpetofauna of Timor-Leste: a first report" masterLastPageNumber="86" masterPageNumber="19" pageNumber="28" updateTime="1668151585581" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>The herpetofauna of Timor-Leste: a first report</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Kaiser, Hinrich</mods:namePart>
<mods:affiliation>Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA; and The Foundation for Post-Conflict Development, 245 Park Avenue, 24 th Floor, New York, New York 10167, USA</mods:affiliation>
<mods:nameIdentifier type="email">chalcopis@yahoo.com</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Carvalho, Venancio Lopes</mods:namePart>
<mods:affiliation>Universidade National Timor-Lorosa'e, Faculdade de Ciencias da Educacao, Departamentu da Biologia, Avenida Cidade de Lisboa, Liceu Dr. Francisco Machado, Dili, Timor-Leste</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ceballos, Jester</mods:namePart>
<mods:affiliation>Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA; and The Foundation for Post-Conflict Development, 245 Park Avenue, 24 th Floor, New York, New York 10167, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Freed, Paul</mods:namePart>
<mods:affiliation>14149 S. Butte Creek Road, Scotts Mills, Oregon 97375, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Heacox, Scott</mods:namePart>
<mods:affiliation>Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA; and The Foundation for Post-Conflict Development, 245 Park Avenue, 24 th Floor, New York, New York 10167, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Lester, Barbara</mods:namePart>
<mods:affiliation>14149 S. Butte Creek Road, Scotts Mills, Oregon 97375, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Richards, Stephen J.</mods:namePart>
<mods:affiliation>Conservation International, PO Box 1024, Atherton, Queensland 4883, Australia; and Herpetology Department, South Australian Museum, North Terrace, Adelaide, South Australia 5000, Australia</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Trainor, Colin R.</mods:namePart>
<mods:affiliation>School of Environmental and Life Sciences, Charles Darwin University, Darwin, Northern Territory 0909, Australia</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Sanchez, Caitlin</mods:namePart>
<mods:affiliation>Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA; and The Foundation for Post-Conflict Development, 245 Park Avenue, 24 th Floor, New York, New York 10167, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>O'Shea, Mark</mods:namePart>
<mods:affiliation>West Midland Safari Park, Bewdley, Worcestershire DY 12 1 LF, United Kingdom; and Australian Venom Research Unit, Department of Pharmacology, University of Melbourne, Victoria 3010, Australia</mods:affiliation>
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<mods:date>2011</mods:date>
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<mods:number>2011-06-20</mods:number>
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<mods:number>109</mods:number>
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<mods:url>http://dx.doi.org/10.3897/zookeys.109.1439</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.109.1439</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-109-19</mods:identifier>
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<treatment ID-GBIF-Taxon="152029820" LSID="urn:lsid:plazi:treatment:CA119BC729FEFFAEC39252B82BF98D8F" httpUri="http://treatment.plazi.org/id/CA119BC729FEFFAEC39252B82BF98D8F" lastPageId="10" lastPageNumber="29" pageId="9" pageNumber="28">
<subSubSection pageId="9" pageNumber="28" type="nomenclature">
<paragraph pageId="9" pageNumber="28">
Genus
<taxonomicName LSID="CA119BC7-29FE-FFAE-C392-52B82BF98D8F" authorityName="Bolkay" authorityYear="1915" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="Fejervarya" order="Anura" pageId="9" pageNumber="28" phylum="Chordata" rank="genus">Fejervarya</taxonomicName>
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Rice paddy frogs, genus Fejervarya. A Fejervarya sp. 1 from the Baucau area (SVL 58 mm) B Fejervarya sp. 2 from the Viqueque area (SVL 46 mm) C Fejervarya sp. 3 from the Viqueque area (SVL 38 mm) D All three species of rice paddy frogs found in Timor-Leste may show varying degrees of green coloration on the dorsal and lateral parts of the body. This specimen (SVL 56 mm) from the Viqueque area represents the most extreme green coloration we observed, in terms of both brightness and coverage. Photos by Mark O'Shea." httpUri="https://binary.pensoft.net/fig/15834" pageId="9" pageNumber="28">Fig. 4</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="9" pageNumber="28" type="common names">
<paragraph pageId="9" pageNumber="28">Common names.</paragraph>
<paragraph pageId="9" pageNumber="28">(E) Rice Paddy Frogs. (T) Manduku natar (manduku = frog, natar = rice paddy).</paragraph>
</subSubSection>
<subSubSection pageId="9" pageNumber="28" type="identification">
<paragraph pageId="9" pageNumber="28">Identification.</paragraph>
<paragraph pageId="9" pageNumber="28">
Rice paddy frogs (
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Rice paddy frogs, genus Fejervarya. A Fejervarya sp. 1 from the Baucau area (SVL 58 mm) B Fejervarya sp. 2 from the Viqueque area (SVL 46 mm) C Fejervarya sp. 3 from the Viqueque area (SVL 38 mm) D All three species of rice paddy frogs found in Timor-Leste may show varying degrees of green coloration on the dorsal and lateral parts of the body. This specimen (SVL 56 mm) from the Viqueque area represents the most extreme green coloration we observed, in terms of both brightness and coverage. Photos by Mark O'Shea." httpUri="https://binary.pensoft.net/fig/15834" pageId="9" pageNumber="28">Fig. 4</figureCitation>
) are the most common amphibians found in regions with rice agriculture. They may grow to over 60 mm in snout-vent length and can usually be recognized by their fairly stout body shape, brownish to gray-green coloration, shiny moist skin with ill-defined dorsal and lateral patterns, and warts scattered irregularly or in rows along the back (e.g.,
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Rice paddy frogs, genus Fejervarya. A Fejervarya sp. 1 from the Baucau area (SVL 58 mm) B Fejervarya sp. 2 from the Viqueque area (SVL 46 mm) C Fejervarya sp. 3 from the Viqueque area (SVL 38 mm) D All three species of rice paddy frogs found in Timor-Leste may show varying degrees of green coloration on the dorsal and lateral parts of the body. This specimen (SVL 56 mm) from the Viqueque area represents the most extreme green coloration we observed, in terms of both brightness and coverage. Photos by Mark O'Shea." httpUri="https://binary.pensoft.net/fig/15834" pageId="9" pageNumber="28">Fig. 4B</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="10" lastPageNumber="29" pageId="9" pageNumber="28" type="collection and natural history">
<paragraph pageId="9" pageNumber="28">Collection and natural history.</paragraph>
<paragraph pageId="9" pageNumber="28">
As the name suggests, rice paddy frogs are commonly found in rice paddies where they perch at the
<normalizedToken originalValue="waters">water's</normalizedToken>
edge, on tufts of vegetation, or even on cow patties. Our survey documented at least three species of rice paddy frogs in Timor-Leste, with two or more often occurring in the same suitable habitat. Species can be distinguished by the size of mature males, which can be clearly separated into three groups by their size, morphology of the tympanum and its associated structures, and the patterning of the throat in males. One of the species, designated as
<taxonomicName authorityName="Bolkay" authorityYear="1915" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Anura" pageId="9" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="9" pageNumber="28">Fejervarya</emphasis>
</taxonomicName>
sp. 1 [Ca CMD 431] is a lowland form and most similar to
<taxonomicName baseAuthorityName="Roux" baseAuthorityYear="1911" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Anura" pageId="9" pageNumber="28" phylum="Chordata" rank="species" species="verruculosa">
<emphasis italics="true" pageId="9" pageNumber="28">Fejervarya verruculosa</emphasis>
</taxonomicName>
(
<bibRefCitation author="Roux, J" journalOrPublisher="Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft" pageId="34" pageNumber="53" pagination="211 - 247" refId="B78" refString="Roux, J, 1910. Reptilien und Amphibien der Aru- und Kei-Inseln. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 33: 211 - 247" title="Reptilien und Amphibien der Aru- und Kei-Inseln." volume="33" year="1910">Roux 1910</bibRefCitation>
). This candidate species differs from
<taxonomicName baseAuthorityName="Roux" baseAuthorityYear="1911" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Anura" pageId="9" pageNumber="28" phylum="Chordata" rank="species" species="verruculosa">
<emphasis italics="true" pageId="9" pageNumber="28">Fejervarya verruculosa</emphasis>
</taxonomicName>
by the shape of the supratympanic fold, the size of the tympanum relative to the eye, patterning on the hidden surfaces of the legs, and number and position of maxillary teeth and the shape of the alary process of the premaxilla, among other characters. The other two candidate species can also be differentiated by these and other characters, and they have been designated
<taxonomicName authorityName="Bolkay" authorityYear="1915" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Anura" pageId="9" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="9" pageNumber="28">Fejervarya</emphasis>
</taxonomicName>
sp. 2 [Ca CMD 508] and
<taxonomicName authorityName="Bolkay" authorityYear="1915" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Anura" pageId="9" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="9" pageNumber="28">Fejervarya</emphasis>
</taxonomicName>
sp. 3 [Ca CMD 355].
</paragraph>
<paragraph lastPageId="10" lastPageNumber="29" pageId="9" pageNumber="28">
Among the specific habitats where we encountered these frogs were active rice paddies, roadside puddles, coffee plantations, and coastal forests at altitudes between 4 m and 1187 m. Our observations are consistent with those of
<bibRefCitation author="Menzies, JI" journalOrPublisher="Australian Journal of Zoology" pageId="34" pageNumber="53" pagination="373 - 418" publicationUrl="doi:10.1071/ZO9870373" refId="B68" refString="Menzies, JI, 1987. A taxonomic revision of Papuan Rana (Amphibia: Ranidae). Australian Journal of Zoology 35: 373 - 418, doi:10.1071/ZO9870373" title="A taxonomic revision of Papuan Rana (Amphibia: Ranidae)." url="doi:10.1071/ZO9870373" volume="35" year="1987">Menzies (1987)</bibRefCitation>
but we believe that not all three presumptive species are capable of such habitat plasticity.
<pageBreakToken pageId="10" pageNumber="29" start="start">More</pageBreakToken>
detailed investigation is needed to clarify the habitat requirements of these three candidate
<taxonomicName authorityName="Bolkay" authorityYear="1915" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Anura" pageId="10" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="29">Fejervarya</emphasis>
</taxonomicName>
species.
</paragraph>
</subSubSection>
<subSubSection pageId="10" pageNumber="29" type="biogeography">
<paragraph pageId="10" pageNumber="29">Biogeography.</paragraph>
<paragraph pageId="10" pageNumber="29">
The presence in Timor-Leste of three morphologically similar and seemingly endemic dicroglossid frogs, with an evolutionary origin in Asia, raises interesting biogeographic questions. The simplest explanation would be a single or a series of introductions in modern times, with the influx of peoples and cargo from points all across the Indonesian Archipelago. However, a more ancient, classic island biogeography scenario is also feasible. Based on the two main concepts of speciation, sympatric speciation among amphibians is possible but presumed rare among dicroglossid frogs. The concept of allopatric speciation is the alternative, by which the three species may represent descendants of three separate introductions that occurred as early farmers brought rice plants to the island. It is generally accepted that rice agriculture originally spread from China into South and Southeast Asia (
<bibRefCitation author="Crawford, GW" journalOrPublisher="Antiquity" pageId="34" pageNumber="53" pagination="858 - 866" refId="B23" refString="Crawford, GW, Chen, S, 1998. The origins of rice agriculture: recent progress in East Asia. Antiquity 72: 858 - 866" title="The origins of rice agriculture: recent progress in East Asia." volume="72" year="1998">Crawford and Chen 1998</bibRefCitation>
) and reached the Lesser Sunda Islands in waves after spreading throughout the Greater Sunda Islands during the Neolithic Period (
<bibRefCitation author="Chi, Z" journalOrPublisher="Asian Perspectives" pageId="34" pageNumber="53" pagination="299 - 329" refId="B20" refString="Chi, Z, Hung, H-C, 2008. The Neolithic of southern China-origin, development, and dispersal. Asian Perspectives 47: 299 - 329" title="The Neolithic of southern China-origin, development, and dispersal." volume="47" year="2008">Chi and Hung 2008</bibRefCitation>
). During this time, it is quite possible that stowaway frogs arrived on Timor with rice plants. The occurrence of several similar rice paddy frog species in sympatry is not unique to Timor-Leste (e.g., Burma; G. Zug, in litt.). Molecular studies to obtain some insights into this conundrum are progressing. The hypothesis that multiple human-mitigated introductions of
<taxonomicName authorityName="Bolkay" authorityYear="1915" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Anura" pageId="10" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="29">Fejervarya</emphasis>
</taxonomicName>
populations occurred mirroring the development of rice cultivation is a plausible explanation for the many species of this genus listed as
<emphasis italics="true" pageId="10" pageNumber="29">incertae sedis</emphasis>
with respect to their intrageneric relationships.
</paragraph>
<caption httpUri="https://binary.pensoft.net/fig/15834" pageId="10" pageNumber="29" start="Figure 4" startId="F4">
<paragraph pageId="10" pageNumber="29">
<emphasis bold="true" pageId="10" pageNumber="29">Figure 4.</emphasis>
Rice paddy frogs, genus
<taxonomicName authorityName="Bolkay" authorityYear="1915" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Anura" pageId="10" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="29">Fejervarya</emphasis>
</taxonomicName>
.
<emphasis bold="true" pageId="10" pageNumber="29">A</emphasis>
<taxonomicName authorityName="Bolkay" authorityYear="1915" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Anura" pageId="10" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="29">Fejervarya</emphasis>
</taxonomicName>
sp. 1 from the Baucau area (SVL 58 mm)
<emphasis bold="true" pageId="10" pageNumber="29">B</emphasis>
<taxonomicName authorityName="Bolkay" authorityYear="1915" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Anura" pageId="10" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="29">Fejervarya</emphasis>
</taxonomicName>
sp. 2 from the Viqueque area (SVL 46 mm)
<emphasis bold="true" pageId="10" pageNumber="29">C</emphasis>
<taxonomicName authorityName="Bolkay" authorityYear="1915" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Anura" pageId="10" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="29">Fejervarya</emphasis>
</taxonomicName>
sp. 3 from the Viqueque area (SVL 38 mm)
<emphasis bold="true" pageId="10" pageNumber="29">D</emphasis>
All three species of rice paddy frogs found in Timor-Leste may show varying degrees of green coloration on the dorsal and lateral parts of the body. This specimen (SVL 56 mm) from the Viqueque area represents the most extreme green coloration we observed, in terms of both brightness and coverage. Photos by Mark
<normalizedToken originalValue="OShea">O'Shea</normalizedToken>
.
</paragraph>
</caption>
</subSubSection>
</treatment>
</document>