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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.555.6522" ID-GBIF-Dataset="78858f45-6a20-4417-811f-17185cd352ab" ID-PMC="PMC4740822" ID-Pensoft-Pub="1313-2970-555-57" ID-PubMed="26877687" ID-ZBK="980FD5B24399479F848A16BD8F926405" ModsDocAuthor="" ModsDocDate="2016" ModsDocID="1313-2970-555-57" ModsDocOrigin="ZooKeys 555" ModsDocTitle="A new genus and species of arboreal toad with phytotelmonous larvae, from the Andaman Islands, India (Lissamphibia, Anura, Bufonidae)" checkinTime="1453318051294" checkinUser="pensoft" docAuthor="Chandramouli, S. R., Vasudevan, Karthikeyan, Harikrishnan, S., Dutta, Sushil Kumar, Janani, S. Jegath, Sharma, Richa, Das, Indraneil &amp; Aggarwal, Ramesh K." docDate="2016" docId="0D25AB69B598C01509BB7FB867A3FBE0" docLanguage="en" docName="ZooKeys 555: 57-90" docOrigin="ZooKeys 555" docSource="http://dx.doi.org/10.3897/zookeys.555.6522" docTitle="Blythophryne Chandramouli, Vasudevan, Harikrishnan, Dutta, Janani, Sharma, Das &amp; Aggarwal, 2016, gen. n." docType="treatment" docUuid="2BAB0154-53B8-43E3-BB14-F36F12FDD8DE" docUuidSource="ZooBank" docVersion="5" lastPageNumber="71" masterDocId="FFA8FF9ED4394663FF8A9C72FFBF057E" masterDocTitle="A new genus and species of arboreal toad with phytotelmonous larvae, from the Andaman Islands, India (Lissamphibia, Anura, Bufonidae)" masterLastPageNumber="90" masterPageNumber="57" pageNumber="60" updateTime="1668162501040" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>A new genus and species of arboreal toad with phytotelmonous larvae, from the Andaman Islands, India (Lissamphibia, Anura, Bufonidae)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Chandramouli, S. R.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Vasudevan, Karthikeyan</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Harikrishnan, S.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Dutta, Sushil Kumar</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Janani, S. Jegath</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Sharma, Richa</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Das, Indraneil</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Aggarwal, Ramesh K.</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
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<mods:part>
<mods:date>2016</mods:date>
<mods:detail type="volume">
<mods:number>555</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>57</mods:start>
<mods:end>90</mods:end>
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<mods:url>http://dx.doi.org/10.3897/zookeys.555.6522</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.555.6522</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-555-57</mods:identifier>
<mods:identifier type="ZBK">980FD5B24399479F848A16BD8F926405</mods:identifier>
<mods:identifier type="ZooBank">980FD5B24399479F848A16BD8F926405</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="127896854" LSID="urn:lsid:zoobank.org:act:2BAB0154-53B8-43E3-BB14-F36F12FDD8DE" httpUri="http://treatment.plazi.org/id/0D25AB69B598C01509BB7FB867A3FBE0" lastPageId="14" lastPageNumber="71" pageId="3" pageNumber="60">
<subSubSection pageId="3" pageNumber="60" type="multiple">
<paragraph pageId="3" pageNumber="60">Taxon classification Animalia ORDO FAMILIA</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="60" type="nomenclature">
<paragraph pageId="3" pageNumber="60">
<taxonomicName LSID="http://zoobank.org/2BAB0154-53B8-43E3-BB14-F36F12FDD8DE" class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne" order="Anura" pageId="3" pageNumber="60" phylum="Chordata" rank="genus">Blythophryne</taxonomicName>
<taxonomicNameLabel pageId="3" pageNumber="60">gen. n.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="60" type="type species">
<paragraph pageId="3" pageNumber="60">Type species.</paragraph>
<paragraph pageId="3" pageNumber="60">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="3" pageNumber="60" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. by monotypy (Fig. 1, Table 4).
</paragraph>
<caption pageId="3" pageNumber="60">
<paragraph pageId="3" pageNumber="60">
Table 4. Morphometric measurements of the holotype and paratype series of adult and two gravid (g) individuals of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="3" pageNumber="60" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n.
</paragraph>
</caption>
<paragraph pageId="3" pageNumber="60">
<table pageId="3" pageNumber="60">
<tr pageId="3" pageNumber="60">
<th colspan="1" pageId="3" pageNumber="60" rowspan="1">ZSI A-12521</th>
<th colspan="1" pageId="3" pageNumber="60" rowspan="1">ZSI A-12524</th>
<th colspan="1" pageId="3" pageNumber="60" rowspan="1">ZSI A-12522</th>
<th colspan="1" pageId="3" pageNumber="60" rowspan="1">ZSI A-12523</th>
<th colspan="1" pageId="3" pageNumber="60" rowspan="1">ZSI A-12526</th>
<th colspan="1" pageId="3" pageNumber="60" rowspan="1">ZSI A-12529</th>
<th colspan="1" pageId="3" pageNumber="60" rowspan="1">ZSI A-12527</th>
<th colspan="1" pageId="3" pageNumber="60" rowspan="1">ZSI A-12530</th>
<th colspan="1" pageId="3" pageNumber="60" rowspan="1">ZSI A-12528</th>
<th colspan="1" pageId="3" pageNumber="60" rowspan="1">ZSI A-12525</th>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">Sex</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">SVL</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">AG</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">HL</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">HW</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">HD</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">BW</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">EN</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">ES</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">ETY</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">UEW</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">IO</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">IN</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">TYH</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">TYV</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">UAL</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">LAL</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">PAL</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">FEL</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">TBL</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">FOL</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">ED</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">DL fold</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">PL</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">PW</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">f1</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">f2</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">f3</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">f4</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">t1</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">t2</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">t3</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">t4</td>
</tr>
<tr pageId="3" pageNumber="60">
<td colspan="1" pageId="3" pageNumber="60" rowspan="1">t5</td>
</tr>
</table>
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="60" type="content">
<paragraph pageId="3" pageNumber="60">Content.</paragraph>
<paragraph pageId="3" pageNumber="60">A single species is currently known.</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="60" type="type material">
<paragraph pageId="3" pageNumber="60">Type material.</paragraph>
<paragraph pageId="3" pageNumber="60">
Holotype ♀: ZSI_A-12521(Fig. 1), (SVL 27.4 mm) leg. S. R. Chandramouli and S. Harikrishnan on 12 December 2010 near Mt. Harriet National Park (ca.
<geoCoordinate direction="north" orientation="latitude" precision="925" value="11.7">11°42'N</geoCoordinate>
,
<geoCoordinate direction="east" orientation="longitude" precision="925" value="92.73333">92°44'E</geoCoordinate>
, 175 m asl.) within evergreen forests at ca. 2130 hours. Paratypes (paratopotypes): ZSI_A-12522 to ZSI_A-12530 (three ♀ and six ♂; Fig. 1g); leg. S. R. Chandramouli and S. Harikrishnan during 22 - 25 June 2010 from the above location but at an altitude range of ~150-330 m asl. Other paratypes (larvae): seven tadpoles (WII-115) collected from a phytotelm on Rutland Island. Referred material: WII-113, an adult topotype with six toes on both the feet.
</paragraph>
<caption pageId="3" pageNumber="60">
<paragraph pageId="3" pageNumber="60">
Figure 1. Morphological characters of the
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="3" pageNumber="60" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n.: a dorso-lateral view b dorsal view c ventral view d ventral view of left palm e ventral view of left foot of the adult female holotype (ZSI_A-12521) in life f adult female holotype in preservation g dorsal view of the male paratype (ZSI_A-12529) in life showing inverted-V shaped markings and the inter-ocular band on the dorsum.
</paragraph>
</caption>
</subSubSection>
<subSubSection lastPageId="6" lastPageNumber="63" pageId="3" pageNumber="60" type="etymology">
<paragraph pageId="3" pageNumber="60">Etymology.</paragraph>
<paragraph lastPageId="6" lastPageNumber="63" pageId="3" pageNumber="60">
The generic name is a patronym, coined in appreciation of Edward Blyth (1810-1873), the first curator of the Asiatic Society of Bengal, who initiated
<pageBreakToken pageId="4" pageNumber="61" start="start">herpetological</pageBreakToken>
studies in the Andaman and Nicobar Islands, through his phenomenal, pioneering paper &quot;Notes on the fauna of the Nicobar islands&quot; (Blyth 1846).
<bibRefCitation author="Das, I" editor="Ota, H" journalOrPublisher="Elsevier Science B. V. Amsterdam" pageId="19" pageNumber="76" pagination="43 - 77" title="Biogeography of the amphibians and reptiles of the Andaman and Nicobar Islands." volumeTitle="Tropical island herpetofauna. Origin, current diversity and conservation" year="1999">Das (1999)</bibRefCitation>
remarked, &quot;Blyth is to be credited for the description of a large number of species from the Andaman and Nicobar Islands that are still valid. Blyth (1846) wrote the
<pageBreakToken pageId="5" pageNumber="62" start="start">first</pageBreakToken>
account on the vertebrate fauna of these islands, and in 1863, compiled the first check-list&quot;. Further details of Edward Blyth and his contributions to studies on Indian natural history are in
<bibRefCitation author="Das, I" journalOrPublisher="Bonner Zoologische Beitraege" pageId="19" pageNumber="76" pagination="215 - 229" title="Herpetology of an antique land: the history of herpetological explorations and knowledge in India and south Asia." volume="52" year="2004">Das (2004)</bibRefCitation>
and Sridharan (2013). The specific epithet '
<taxonomicName lsidName="beryet" pageId="5" pageNumber="62" rank="species" species="beryet">beryet</taxonomicName>
' (in Great Andamanese language; http://www.andamanese.net/Great_Andamanese_Lexicon_English.pdf) refers to 'small
<normalizedToken originalValue="frog">frog'</normalizedToken>
. We believe that the Great Andamanese knew of the existence of this small arboreal anuran that is here described as new species to
<pageBreakToken pageId="6" pageNumber="63" start="start">science</pageBreakToken>
. We hope the name given here will also raise awareness about the dwindling, indigenous tribal populations in the Andamans, their culture and extinction of their tribal languages.
</paragraph>
</subSubSection>
<subSubSection pageId="6" pageNumber="63" type="diagnosis">
<paragraph pageId="6" pageNumber="63">Diagnosis.</paragraph>
<paragraph pageId="6" pageNumber="63">This currently monotypic genus and species is diagnosed by the following suite of external morphological and osteological characters: small adult size (mean SVL 24.0 mm; range 21.8-27.4 mm); distinct tympanum, slightly smaller than eye; absence of cephalic ridges; absence of vomerine teeth; presence of a single, median, external vocal sac in males; presence of elongated pair of parotoid glands; absence of enlarged, keratinised tubercles on dorsum; presence of well developed, sheath-like webbing on fingers and on toes; digit tips dilated to discs, lacking circum-marginal grooves; presence of six presacral vertebrae; urostyle lacking lateral dilations; absence of omosternum and presence of arciferal pectoral girdle. Mature ova small (0.62 mm mean diameter), yolky and unpigmented; tadpoles lacking keratodont.</paragraph>
</subSubSection>
<subSubSection pageId="6" pageNumber="63" type="description of the holotype">
<paragraph pageId="6" pageNumber="63">Description of the holotype.</paragraph>
<paragraph pageId="6" pageNumber="63">
A small bufonid (mean SVL 24.2
<normalizedToken originalValue="±">+/-</normalizedToken>
0.6 mm), with depressed, moderately robust (AG:BW 1.0) habitus (Fig. 1
<normalizedToken originalValue="ac">a-c</normalizedToken>
). Head almost as long as broad (HL:HW 0.97), devoid of cephalic ridges, with a single, median internal vocal sac in males. Snout obtusely pointed in dorsal view, projecting beyond mandibles; nostrils oriented laterally, situated on lateral fold closer to tip of snout than to eye (EN:ES 0.7), loreal region mildly concave, canthal ridge well defined between nostril and the eye, distance between orbit and nostril greater than internarial distance (IN:EN 0.96), upper eyelid rough, densely covered with minute warts, eyes large (ED:HL 0.4), about twice length of tympanum (TYH:ED 0.6), separated from each other by twice internarial distance (IN:IO 0.6), and over twice width of upper eyelid (IO:UEW 1.9), pineal ocellus absent; vomerine teeth absent, tongue elongate, slender and oval, free posteriorly, not bifid, lacking lingual papilla; dorsolateral fold conspicuous, almost up to 48% SVL, beyond which it becomes indistinct and disappears; parotoid glands slender and elongate (PL:PW 4.3), as well-defined postorbital ridge. Limbs slender, upper arm short, 18.7% of SVL, lower arm longer than the upper arm (21% SVL), fingers basally webbed, webbing between Fingers II and III not exceeding penultimate subarticular tubercle (webbing formula I0-1II1-2III2-1IV; Fig. 1d); an enlarged, prominent outer metacarpal tubercle at palmar base (subequal to disc on Finger I), nuptial pad absent, subarticular tubercles prominent on fingers and toes, finger tips dilated to discs lacking circummarginal grooves that are much broader than long, and are less discernible in the first and second fingers; relative length of fingers 3&gt; 4&gt; 2&gt; 1; thigh 33.7% SVL, subequal to shank (38.6% SVL); toes partially webbed, webbing between Toes III and IV extending to penultimate subarticular tubercle (webbing formula I0-1II0-1III1-2IV
<normalizedToken originalValue="2½-½">21/2-1/2</normalizedToken>
V; Fig. 1e); tarsal ridge absent, inner meta-tarsal tubercle larger than outer. Relative length of toes 4&gt; 5&gt; 3&gt; 2&gt; 1. Skin rough dorsally and granular ventrally; lower abdomen with free, loose skin flap. Tubercles or granules absent on dorsum, scattered over venter, under surface of thighs less granular; throat and limb-insertions with dense granules, tibia with enlarged granular tubercles.
</paragraph>
</subSubSection>
<subSubSection lastPageId="7" lastPageNumber="64" pageId="6" pageNumber="63" type="colouration in life">
<paragraph pageId="6" pageNumber="63">Colouration in life.</paragraph>
<paragraph lastPageId="7" lastPageNumber="64" pageId="6" pageNumber="63">
Dorsum reddish-brown, with two feeble dark brown inverted
<normalizedToken originalValue="V">'V'</normalizedToken>
shaped markings which fail to reach flanks, interorbital band indistinct, canthus
<pageBreakToken pageId="7" pageNumber="64" start="start">dark</pageBreakToken>
chocolate brown, colour extending a little beyond tympanum, subequal to half-length of parotoid gland; forearm and hind limbs barred, one each on thigh, shank and tarsus. Venter heavily speckled with dark brown spots, throat dark brown, lower lip spotted with white and brown, pupil large, horizontally elliptical.
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="64" type="colouration in alcohol">
<paragraph pageId="7" pageNumber="64">Colouration in alcohol.</paragraph>
<paragraph pageId="7" pageNumber="64">
Dorsum drab brown with indistinct
<normalizedToken originalValue="inverted-V">'inverted-V'</normalizedToken>
shaped pattern, darker bands on limbs, venter cream, with black mottled pattern, throat black throughout (Fig. 1f).
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="64" type="osteology">
<paragraph pageId="7" pageNumber="64">Osteology</paragraph>
<paragraph pageId="7" pageNumber="64">
(based on paratype ZSI_A12527). Axial and appendicular skeleton composed primarily of bony elements; cartilaginous elements not observed. Atlas (the first vertebra) with rudimentary hypapophysis and not fused to axis, presacral vertebrae six in number, Vertebrae
<normalizedToken originalValue="IIV">II-V</normalizedToken>
bearing horizontally elongate hypapophyses, those on Vertebrae II and V oriented anteriorly; Vertebrae
<normalizedToken originalValue="IIIIV">III-IV</normalizedToken>
oriented horizontally; sacral diapophysis laterally dilated; coccyx not fused to sacrum; articulating with former by a double condyle and lacking lateral expansions, omosternum absent, pectoral girdle arciferal, with epicoracoids united to each other anteriorly and overlapping posteriorly (Fig. 2). Phalangeal formula of fingers 2-2-3-3; toes 2-2-3-4-3, terminal phalange obtusely curved, not truncate. Nasal bones of the skull large, about 1/3rd of frontoparietals and 1.25 times as large as orbital cavity. Maxillary and vomerine teeth absent.
</paragraph>
<caption pageId="7" pageNumber="64">
<paragraph pageId="7" pageNumber="64">
Figure 2. Skeletal characters of paratypes (ZSI_A-12527) of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="7" pageNumber="64" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. a complete dry structure
<normalizedToken originalValue="bf">b-f</normalizedToken>
various characters visible after staining/clearing of the skeleton. FP - frontoparietal; N - nasal.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="7" pageNumber="64" type="morphological variations">
<paragraph pageId="7" pageNumber="64">Morphological variations.</paragraph>
<paragraph pageId="7" pageNumber="64">
Adult females and males range between 25.2-27.4 mm and 21.8-25.5 mm, respectively. Measurements of paratypes are provided in Table 4. Dorsal colour in different shades of brown or reddish-brown. Intensity of inverted
<normalizedToken originalValue="V">'V'</normalizedToken>
-shaped pattern on dorsum variable. On one occasion, an abnormal specimen (WII-113) with a deformity was observed, with six digits, the first toe being preceded by a small additional toe on both feet. Fingers showed no such anomalies.
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="64" type="description of calls">
<paragraph pageId="7" pageNumber="64">Description of calls.</paragraph>
<paragraph pageId="7" pageNumber="64">
(Macaulay Library, Cornell Lab of Ornithology; voucher no: ML 174095). A calling male was observed on 24 November 2010 on the surface of leaves within bushes. Calls were composed of continuous syllables of
<normalizedToken originalValue="“pip-pip-pip-pip-pip-”">&quot;pip-pip-pip-pip-pip-&quot;</normalizedToken>
at a constant frequency of 8 kHz, without pause, lasting for 23 seconds, with mean amplitude of -3 db / 20 kU (Fig. 3). The call was composed of 198 pulses uttered within duration of 23 s, at a rate of 8 to 9 (mean = 8.6) pulses per second. Each pulse lasted for duration of 0.3 s (n = 198) with an interval of 8.5 s (n = 197) between two consecutive pulses.
</paragraph>
<caption pageId="7" pageNumber="64">
<paragraph pageId="7" pageNumber="64">
Figure 3. Sound spectrogram a and oscillogram b of a 23 second clip of a call of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="7" pageNumber="64" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n.. Detailed view of c frequency and d amplitude modulations of a one second long clip of the call e power spectrum of the call of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="7" pageNumber="64" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="7" pageNumber="64" type="distribution">
<paragraph pageId="7" pageNumber="64">Distribution.</paragraph>
<paragraph pageId="7" pageNumber="64">
This species has been documented from five islands of the Andaman archipelago, namely, the South Andaman (Mt. Harriet), Rutland, Little Andaman, Havelock Island in the
<normalizedToken originalValue="Ritchies">Ritchie's</normalizedToken>
Archipelago and North Andaman (Saddle Peak) (Fig. 4).
</paragraph>
<caption pageId="7" pageNumber="64">
<paragraph pageId="7" pageNumber="64">
Figure 4. Map showing distribution of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="7" pageNumber="64" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. in the Andaman Islands, Bay of Bengal, India. Holotype collected from Mt Harriet (indicated with a red triangle).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="7" pageNumber="64" type="vernacular name">
<paragraph pageId="7" pageNumber="64">Vernacular name.</paragraph>
<paragraph pageId="7" pageNumber="64">
'Andaman bush
<normalizedToken originalValue="toad">toad'</normalizedToken>
is proposed as the common English name for this new species, indicating its arboreal habit and restricted distribution as understood currently.
</paragraph>
</subSubSection>
<subSubSection lastPageId="9" lastPageNumber="66" pageId="7" pageNumber="64" type="ecological notes">
<paragraph pageId="7" pageNumber="64">Ecological notes.</paragraph>
<paragraph lastPageId="8" lastPageNumber="65" pageId="7" pageNumber="64">
The new species is often seen on surface of leaves of herbaceous bushes. It is nocturnal and regularly seen year round. It was the third most common anuran in the islands (
<bibRefCitation author="Harikrishnan, S" journalOrPublisher="BMC Ecology" pageId="20" pageNumber="77" title="The devil is in the detail: estimating species richness, density, and relative abundance of tropical island herpetofauna." url="10.1186/s12898-015-0049-5" volume="15" year="2015">Harikrishnan and Vasudevan 2015</bibRefCitation>
). The high abundance of this species seems to be the result of it occupying a narrow range of distribution and a
<pageBreakToken pageId="8" pageNumber="65" start="start">unique</pageBreakToken>
niche of frogs belonging to the Old World tree frog family (
<taxonomicName family="Rhacophoridae" lsidName="" pageId="8" pageNumber="65" rank="family">Rhacophoridae</taxonomicName>
), which are not known to occur on the Andaman Islands. All other anuran amphibians recorded from these islands are ground-dwelling, with the exception of
<taxonomicName class="Amphibia" family="Microhylidae" genus="Kaloula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaloula baleata" order="Anura" pageId="8" pageNumber="65" phylum="Chordata" rank="subspecies" species="baleata" subspecies="ghoshi">Kaloula baleata ghoshi</taxonomicName>
, which is semi-arboreal, and
<taxonomicName class="Amphibia" family="Dicroglossidae" genus="Ingerana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ingerana charlesdarwini" order="Anura" pageId="8" pageNumber="65" phylum="Chordata" rank="species" species="charlesdarwini">Ingerana charlesdarwini</taxonomicName>
, which is known to use phytotelms for breeding and oviposition (
<bibRefCitation author="Das, I" journalOrPublisher="Hamadryad" pageId="19" pageNumber="76" pagination="41 - 49" title="A remarkable new species of ranid (Anura: Ranidae) with phytotelmonous larvae, from Mount Harriet, Andaman Islands." volume="23" year="1998">Das 1998</bibRefCitation>
). During day time, bush toads were found under leaf litter on the forest floor.
</paragraph>
<paragraph pageId="8" pageNumber="65">
The Andaman bush toad emits a white, viscous, pungent smelling secretion from the parotoid glands when handled (Fig. 5a); the secretion seems to be toxic, as other frogs kept within the same bag as one of these toads suffered mortality. Breeding commences in June with the onset of the Southwest Monsoon. Males were observed to call from heights of ca. 1-1.5 m above ground while sitting on leaves of bushes. Amplexus is axillary (Fig. 5b), and females deposit ova in phytotelms, which are tree-holes at a height of about 1-1.5 m above the ground filled with rainwater. Tadpoles develop in these phytotelms. The shrub from which the tadpoles described here were collected, measured 19 cm diameter at breast height, and eggs were found in a depression of 6 cm depth, filled with water up to 3 cm. The tree hole was oval, measuring 5
<normalizedToken originalValue="×">x</normalizedToken>
3 cm across (Fig. 6a).
</paragraph>
<caption pageId="8" pageNumber="65">
<paragraph pageId="8" pageNumber="65">
Figure 5. a A live, uncollected specimen of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="8" pageNumber="65" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. showing milky white secretion from the parotoid gland b Amplecting pair (live, uncollected) of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="8" pageNumber="65" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. showing axillary amplexus.
</paragraph>
</caption>
<caption pageId="8" pageNumber="65">
<paragraph pageId="8" pageNumber="65">
Figure 6. a Eggs and hatchling tadpoles of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="8" pageNumber="65" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. b, c endotrophic larvae of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="8" pageNumber="65" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. showing pale white abdominal yolk d Lateral view of a Stage 43 tadpole of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="8" pageNumber="65" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. e Oral disc of a Stage 35 larva of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="8" pageNumber="65" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n., showing absence of keratodont and the presence of keratinised jaw sheaths f a metamorph of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="8" pageNumber="65" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. showing initiation of tail absorption.
</paragraph>
</caption>
<paragraph pageId="9" pageNumber="66">
<pageBreakToken pageId="9" pageNumber="66" start="start">The</pageBreakToken>
Andaman bush toad is widely distributed in islands where it occurs, and occupies forested habitats from 29-250 m asl, more common above 100 m asl and rarer at lower altitudes. The forest types in this elevation range include littoral, moist-deciduous, giant evergreen and montane stunted evergreen forests (
<bibRefCitation author="Champion, HG" journalOrPublisher="Forest Research Institute, Dehradun" pageId="18" pageNumber="75" title="A revised survey of the forest types of India." year="1968">Champion and Seth 1968</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="9" pageNumber="66" type="conservation status">
<paragraph pageId="9" pageNumber="66">Conservation status.</paragraph>
<paragraph pageId="9" pageNumber="66">
The Andaman bush toad is known from five islands: North Andaman (Saddle Peak National Park only), South Andaman, Rutland, Havelock (only in a small patch of wet forest towards the south of the island) and Little Andaman. Based on searches carried out using 21 bounded quadrats of 100 m2 each in these islands, the new species occurs at densities of 1.1
<normalizedToken originalValue="±">+/-</normalizedToken>
0.37 toads per 100 m2 of forest floor (unpublished data). It is considered
<normalizedToken originalValue="Endangered">'Endangered'</normalizedToken>
based on IUCN Ver. 3.1. Second Edition (
<bibRefCitation author="Inger, RF" journalOrPublisher="Natural History Publications (Borneo), Kota Kinabalu" pageId="20" pageNumber="77" title="IUCN Red List of Threatened Species. Version 2013.2" url="http://www.iucnredlist.org" year="2014">IUCN 2014</bibRefCitation>
): criteria B.1 - extent of occurrence &lt;20000 km2 and B.1.a - severely fragmented population and known to exist at no more than 10 locations. A large array of invasive fauna in these Islands threatens the population of this toad. Additionally, stochastic events and anthropogenic pressures are potential threats to the species and its habitat.
</paragraph>
</subSubSection>
<subSubSection pageId="9" pageNumber="66" type="notes on larval development">
<paragraph pageId="9" pageNumber="66">Notes on larval development.</paragraph>
<paragraph pageId="9" pageNumber="66">
(Fig. 6
<normalizedToken originalValue="bf">b-f</normalizedToken>
) The clutch of ova in the phytotelm located in May 2011 at Rutland Island was monitored continuously until complete tadpole transformation. Unpigmented, early-stage larvae were observed on 2nd May 2011. A total of 73 hatchlings presumably from a single clutch could be counted in the phytotelm. Following subsequent rain showers four days later on 6th May, only 25 tadpoles of Stage 20 could be observed, the rest presumably washed out by overflow. At this stage, the tadpoles were translucent and colourless, but speckled with black, with white abdominal yolk region, dorsally positioned eyes and labia visible. On 19th May, i.e., 13 days later, two samples of Stages 30 and 35 were collected and preserved in formalin. Tadpoles of these stages had exposed hind limbs, lacking forelimb buds and were dull purplish-brown in colour, without a dorsal pattern. A week later, on 25th May, the tadpoles that developed into Stages of 41 and 43, were preserved. At these advanced stages, the tadpoles showed developed forelimbs, with expanded discs of fingers, more intense pigmentation on skin, and feeble barred pattern on limbs. The Stage 43 larva is briefly described: mouth positioned anteriorly, with prominent, keratinised pair of jaw sheaths; keratodont absent, eyes and nostrils positioned dorso-laterally (IO 1.46 mm), nostrils much closer to eyes than snout tip. Body depressed, head-body 1.5 times as long as broad (HBL: HBW 1.53), tail almost twice as long as head-body (tL/HBL 1.95) with well-developed caudal musculature. Measurements of the tadpoles are in Table 5.
</paragraph>
<caption pageId="9" pageNumber="66">
<paragraph pageId="9" pageNumber="66">
Table 5. Morphometric measurements of tadpoles of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="9" pageNumber="66" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n.
</paragraph>
</caption>
<paragraph pageId="9" pageNumber="66">
<table pageId="9" pageNumber="66">
<tr pageId="9" pageNumber="66">
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">Stage</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">IOL</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">IND</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">NED</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">NSD</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">SS</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">SV</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">BL</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">TL</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">MBW</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">MTH</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">MTMW</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">TMH</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">ODD</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">VTL</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">DFH</abbrev>
</th>
<th colspan="1" pageId="9" pageNumber="66" rowspan="1">
<abbrev pageId="9" pageNumber="66">VFH</abbrev>
</th>
</tr>
<tr pageId="9" pageNumber="66">
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">30</td>
</tr>
<tr pageId="9" pageNumber="66">
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">35</td>
</tr>
<tr pageId="9" pageNumber="66">
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">41</td>
</tr>
<tr pageId="9" pageNumber="66">
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">42</td>
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">NA</td>
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">NA</td>
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">NA</td>
</tr>
<tr pageId="9" pageNumber="66">
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">43</td>
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">NA</td>
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">NA</td>
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">NA</td>
<td colspan="1" pageId="9" pageNumber="66" rowspan="1">NA</td>
</tr>
</table>
</paragraph>
</subSubSection>
<subSubSection lastPageId="10" lastPageNumber="67" pageId="9" pageNumber="66" type="description of tadpole">
<paragraph pageId="9" pageNumber="66">Description of Tadpole</paragraph>
<paragraph lastPageId="10" lastPageNumber="67" pageId="9" pageNumber="66">
(Stage 35). Body tubular in dorsal and ovoid in lateral views, respectively (Fig. 6c). When viewed laterally, body dorsum is flattened and depressed medially; ventrally body slightly flattened at anterior end and convex towards posterior; body length 35% of total length; body attains maximum diameter in region immediately behind eyes. Snout broad and truncate in dorsal and pointed in lateral views, respectively. Eyes large; located and oriented dorso-laterally. Nostrils rounded with elevated rim, located almost midway but closer to eyes than snout, placed linear to eye in dorsal view; internarial distance subequal to interorbital distance. Spiracle
<pageBreakToken pageId="10" pageNumber="67" start="start">sinistral</pageBreakToken>
and long with no inner wall; spiracle opening large; tube orientated postero-laterally, opening located approximately at midbody. Distance between spiracle and snout about 60% of body length. Intestinal coils not visible through the belly wall; vent tube medial. Tail tip broadly rounded; musculature linear till 1/3rd length of tail, after which it tapers. Dorsal fin slightly wider than ventral fin, originates posterior to body - tail junction and ventral fin at ventral terminus; both fins run parallel to tail muscle parallel through entire length of tail. Maximum tail height attained at about mid-length. Lateral line faintly visible. No glands observed on outer integument.
</paragraph>
<paragraph pageId="10" pageNumber="67">Oral disc positioned at terminal portion of body opening antero-ventrally (Fig. 6e); Rostral width of oral disc 27% body width, non-emarginate; entire oral disc visible dorsally; single row of seven to eight large marginal papillae present on lower labium and two to three on lateral corners; none present on upper labium; a single submarginal papilla located at each lateral corners; lower labium larger than upper labium. Denticle rows absent. Jaw sheaths well developed, heavily keratinised. Jaw sheaths completely serrated with minute serrations on lower jaw than upper jaw; suprarostrodont convex medially, longer than wide and lateral process of subequal height through length; infrarostrodont U-shaped.</paragraph>
<paragraph pageId="10" pageNumber="67">Measurements (in mm; mean shown without parentheses and standard errors are shown in parentheses): Measurements of the seven tadpoles of various stage of development (Stages 30, 35, 41, 42 and 43) are presented in Table 5.</paragraph>
</subSubSection>
<subSubSection pageId="10" pageNumber="67" type="colour">
<paragraph pageId="10" pageNumber="67">Colour.</paragraph>
<paragraph pageId="10" pageNumber="67">
In life, dorsally, outer integument brown, with no melanopores. Ventrally, integument translucent but the gut was not visible; throat speckled. Both tail fins transparent with few melanophores. Laterally, tail muscle white with a few brown spots spread mainly at anterior region of tail. A completely transformed metamorph (SVL 10.6 mm; HL 4.23 mm) resembles adult in morphology, with an evident inverted
<normalizedToken originalValue="V">'V'</normalizedToken>
mark on dorsum and transverse crossbars on limbs.
</paragraph>
</subSubSection>
<subSubSection lastPageId="13" lastPageNumber="70" pageId="10" pageNumber="67" type="morphological comparisons">
<paragraph pageId="10" pageNumber="67">Morphological comparisons.</paragraph>
<paragraph pageId="10" pageNumber="67">Morphological and osteological characteristics of this new taxon are compared with members of other known Oriental bufonid genera below. The new taxon described here differs from the following known genera thus (only opposing character states in the genera being compared are mentioned):</paragraph>
<paragraph pageId="10" pageNumber="67">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Parapelophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Parapelophryne" order="Anura" pageId="10" pageNumber="67" phylum="Chordata" rank="genus">Parapelophryne</taxonomicName>
Fei, Ye &amp; Jiang, 2003: type
<normalizedToken originalValue="species">species-</normalizedToken>
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophryne scalptus" order="Anura" pageId="10" pageNumber="67" phylum="Chordata" rank="species" species="scalptus">Nectophryne scalptus</taxonomicName>
[current name combination:
<taxonomicName class="Amphibia" family="Bufonidae" genus="Parapelophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Parapelophryne scalpta" order="Anura" pageId="10" pageNumber="67" phylum="Chordata" rank="species" species="scalpta">Parapelophryne scalpta</taxonomicName>
(
<bibRefCitation author="Liu, CC" journalOrPublisher="Acta Zoologica Sinica / Dong Wu Xue Bao" pageId="21" pageNumber="78" pagination="385 - 404" title="On collections of amphibians from Hainan Island." volume="19" year="1973">Liu and Hu 1973</bibRefCitation>
)]: Presence of eight presacral vertebrae and absence of parotoid glands (
<bibRefCitation author="Fei, L" journalOrPublisher="Acta Zootaxonomica Sinica" pageId="19" pageNumber="76" pagination="762 - 766" title="A new bufonid genus Parapelophryne from China (Amphibia, Anura)." volume="28" year="2003">Fei et al. 2003</bibRefCitation>
). The phylogenetic position of this taxon was assessed by
<bibRefCitation author="Matsui, M" journalOrPublisher="Current Herpetology" pageId="21" pageNumber="78" pagination="182 - 187" title="Unexpected phylogenetic position of Parapelophryne among Southeast Asian bufonids as revealed by mitochondrial DNA sequence (Amphibia, Anura, Bufonidae)." url="10.5358/hsj.34.182" volume="34" year="2015">Matsui et al. (2015)</bibRefCitation>
, who found it to be sister taxon to
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo japonicus" order="Anura" pageId="10" pageNumber="67" phylum="Chordata" rank="species" species="japonicus">Bufo japonicus</taxonomicName>
, thereby providing additional evidence for its distinctness from
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne" order="Anura" pageId="10" pageNumber="67" phylum="Chordata" rank="genus">Blythophryne</taxonomicName>
gen. n. described here. Distribution: Hainan, eastern China.
</paragraph>
<paragraph lastPageId="11" lastPageNumber="68" pageId="10" pageNumber="67">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes" order="Anura" pageId="10" pageNumber="67" phylum="Chordata" rank="genus">Pedostibes</taxonomicName>
<normalizedToken originalValue="Günther">Guenther</normalizedToken>
, 1875: type species -
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes tuberculosus" order="Anura" pageId="10" pageNumber="67" phylum="Chordata" rank="species" species="tuberculosus">Pedostibes tuberculosus</taxonomicName>
<normalizedToken originalValue="Günther">Guenther</normalizedToken>
, 1875: Larger adult size (SVL 36.6-38.5 mm), presence of eight presacral vertebrae; short, rounded parotoid glands; tips of fingers dilated into truncated discs; small, numerous pigmented ova laid in strings, as in members of the genus
<taxonomicName class="Amphibia" family="Bufonidae" genus="Duttaphrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Duttaphrynus" order="Anura" pageId="10" pageNumber="67" phylum="Chordata" rank="genus">Duttaphrynus</taxonomicName>
and exotrophic larvae (
<normalizedToken originalValue="Günther">Guenther</normalizedToken>
1875,
<bibRefCitation author="Inger, RF" journalOrPublisher="Journal of the Bombay Natural History Society" pageId="20" pageNumber="77" pagination="406 - 427" title="A report on a collection of amphibians and reptiles from the Ponmudi, Kerala, South India." volume="81" year="1984">Inger et al. 1984</bibRefCitation>
,
<bibRefCitation author="Fei, L" journalOrPublisher="Acta Zootaxonomica Sinica" pageId="19" pageNumber="76" pagination="762 - 766" title="A new bufonid genus Parapelophryne from China (Amphibia, Anura)." volume="28" year="2003">Fei et al. 2003</bibRefCitation>
,
<bibRefCitation author="Matsui, M" journalOrPublisher="Zoological Science" pageId="21" pageNumber="78" pagination="1159 - 1166" title="Taxonomic relationships of Ansoniaanotis Inger, Tan, Yambun, 2001 and Pedostibesmaculatus (Mocquard, 1890), with a description of a new genus (Amphibia, Bufonidae)." url="10.2108/zsj.24.1159" volume="24" year="2007">Matsui et al. 2007</bibRefCitation>
). Currently, the genus
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes" order="Anura" pageId="10" pageNumber="67" phylum="Chordata" rank="genus">Pedostibes</taxonomicName>
is represented by five nominal species, which show a disjointed distribution pattern. The westernmost of all,
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes tuberculosus" order="Anura" pageId="10" pageNumber="67" phylum="Chordata" rank="species" species="tuberculosus">Pedostibes tuberculosus</taxonomicName>
, is the type species associated
<pageBreakToken pageId="11" pageNumber="68" start="start">to</pageBreakToken>
the generic name (
<normalizedToken originalValue="Günther">Guenther</normalizedToken>
1875).
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes kempi" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="kempi">Pedostibes kempi</taxonomicName>
is known from the Garo Hills in Meghalaya, north-east India. Presently,
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes kempi" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="kempi">Pedostibes kempi</taxonomicName>
is considered congeneric, but differs in having a concealed tympanum. The remaining species, namely,
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes rugosus" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="rugosus">Pedostibes rugosus</taxonomicName>
,
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes hosii" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="hosii">Pedostibes hosii</taxonomicName>
and
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes everetti" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="everetti">Pedostibes everetti</taxonomicName>
occur in the Indo-Chinese and Indo-Malayan regions (
<bibRefCitation author="Frost, DR" journalOrPublisher="Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenchaftliche Classe" pageId="19" pageNumber="76" title="Amphibian Species of the World: an Online Reference. Version 6" url="http://research.amnh.org/vz/herpetology/amphibia" year="2014">Frost 2014</bibRefCitation>
).
<bibRefCitation author="Bocxlaer, IV" journalOrPublisher="BMC Evolutionary Biology" pageId="18" pageNumber="75" title="Toad radiation reveals into-India dispersal as a source of endemism in the Western Ghats-Sri Lanka biodiversity hotspot." url="10.1186/1471-2148-9-131" volume="9" year="2009">Bocxlaer et al. (2009)</bibRefCitation>
, and more recently
<bibRefCitation author="Ron, SR" journalOrPublisher="Zootaxa" pageId="22" pageNumber="79" pagination="347 - 366" title="Systematics of the endangered toad genus Andinophryne (Anura: Bufonidae): phylogenetic position and synonymy under the genus Rhaebo." url="10.11646/zootaxa.3947.3.3" volume="3" year="2015">Ron et al. (2015)</bibRefCitation>
, in their phylogenetic studies, showed that the genus
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="genus">Pedostibes</taxonomicName>
, as currently defined, does not constitute a monophyletic group. According to their study, the type species,
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes tuberculosus" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="tuberculosus">Pedostibes tuberculosus</taxonomicName>
does not show a close relationship with the south-east Asian
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes hosii" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="hosii">Pedostibes hosii</taxonomicName>
. On the other hand, they demonstrated that
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes hosii" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="hosii">Pedostibes hosii</taxonomicName>
is the sister taxon to
<taxonomicName class="Amphibia" family="Bufonidae" genus="Phrynoidis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynoidis juxtasper" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="juxtasper">Phrynoidis juxtasper</taxonomicName>
. In addition, the generic placement of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes kempi" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="kempi">Pedostibes kempi</taxonomicName>
is also uncertain owing to the inconsistencies in morphological characters associated with this taxon. Hence, resolving the higher level systematic status of the south-east Asian taxa currently allocated to the genus
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="genus">Pedostibes</taxonomicName>
will require further study. Distribution: Western Ghats, Indochina, Malay Peninsula.
</paragraph>
<paragraph pageId="11" pageNumber="68">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufoides" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="genus">Bufoides</taxonomicName>
Pillai &amp; Yazdani, 1973: type
<normalizedToken originalValue="species">species-</normalizedToken>
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ansonia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ansonia meghalayana" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="meghalayana">Ansonia meghalayana</taxonomicName>
[current name combination:
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufoides meghalayanus" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="meghalayanus">Bufoides meghalayanus</taxonomicName>
(Yazdani &amp; Chanda, 1971); currently monotypic, but additional, unnamed species recognised;
<bibRefCitation author="Das, I" journalOrPublisher="Froglog" pageId="19" pageNumber="76" pagination="1 - 4" title="Rediscovery of the endangered Khasi Hills rock toad, Bufoidesmeghalayana in Meghalaya, Northeastern India." volume="92" year="2009">Das et al. 2009</bibRefCitation>
]: Larger adult size (mean 42.9 mm, range 37-47 mm), absence of webbing and expanded discs in fingers, hidden tympanum, presence of cranial ridges and large, pigmented ova laid in strings, as in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Duttaphrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Duttaphrynus" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="genus">Duttaphrynus</taxonomicName>
(Yazdni and Chanda 1972,
<bibRefCitation author="Pillai, RS" journalOrPublisher="Journal of the Zoological Society of India" pageId="21" pageNumber="78" pagination="65 - 70" title="Bufoides, a new genus for the rock-toad, Ansoniameghalayana Yazdani &amp; Chanda, with notes on its ecology and breeding habits." volume="25" year="1973">Pillai and Yazdani 1973</bibRefCitation>
,
<bibRefCitation author="Fei, L" journalOrPublisher="Acta Zootaxonomica Sinica" pageId="19" pageNumber="76" pagination="762 - 766" title="A new bufonid genus Parapelophryne from China (Amphibia, Anura)." volume="28" year="2003">Fei et al. 2003</bibRefCitation>
), presence of seven presacral vertebrae, distinguish this taxon from the newly described genus. Distribution: Khasi Hills, Meghalaya, north-east India (
<bibRefCitation author="Frost, DR" journalOrPublisher="Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenchaftliche Classe" pageId="19" pageNumber="76" title="Amphibian Species of the World: an Online Reference. Version 6" url="http://research.amnh.org/vz/herpetology/amphibia" year="2014">Frost 2014</bibRefCitation>
).
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pelophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelophryne" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="genus">Pelophryne</taxonomicName>
Barbour, 1938: type
<normalizedToken originalValue="species">species-</normalizedToken>
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pelophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelophryne albotaeniata" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="albotaeniata">Pelophryne albotaeniata</taxonomicName>
Barbour, 1938: Presence of coccygeal expansions, absence of parotoid glands; fleshy manus with one phalange free of web and presence of seven (occasionally six) presacral vertebrae, urostyle fused to the sacrum and less number (n ≤ 30) of larger sized yolky eggs (
<bibRefCitation author="Barbour, T" journalOrPublisher="Proceedings of the Biological Society of Washington" pageId="18" pageNumber="75" pagination="191 - 196" title="Notes on the &quot; Nectophryne &quot;." volume="51" year="1938">Barbour 1938</bibRefCitation>
,
<bibRefCitation author="Inger, RF" journalOrPublisher="Fieldiana Zoology" pageId="20" pageNumber="77" pagination="183 - 531" title="Systematics and zoogeography of Philippine Amphibia." url="10.5962/bhl.title.5571" volume="33" year="1954">Inger 1954</bibRefCitation>
,
<bibRefCitation author="Inger, RF" journalOrPublisher="Fieldiana Zoology" pageId="20" pageNumber="77" pagination="1 - 402" title="The systematics and zoogeography of the Amphibia of Borneo." volume="52" year="1966">1966</bibRefCitation>
;
<bibRefCitation author="Matsui, M" journalOrPublisher="Zoological Science" pageId="21" pageNumber="78" pagination="1159 - 1166" title="Taxonomic relationships of Ansoniaanotis Inger, Tan, Yambun, 2001 and Pedostibesmaculatus (Mocquard, 1890), with a description of a new genus (Amphibia, Bufonidae)." url="10.2108/zsj.24.1159" volume="24" year="2007">Matsui et al. 2007</bibRefCitation>
). Distribution: eastern Asia, Sundaland and the Philippines (Frost 2011).
</paragraph>
<paragraph pageId="11" pageNumber="68">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Sabahphrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sabahphrynus" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="genus">Sabahphrynus</taxonomicName>
Matsui, Yambun &amp; Sudin, 2007: type
<normalizedToken originalValue="species">species-</normalizedToken>
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophryne maculata" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="maculata">Nectophryne maculata</taxonomicName>
[current name combination:
<taxonomicName class="Amphibia" family="Bufonidae" genus="Sabahphrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sabahphrynus maculatus" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="maculatus">Sabahphrynus maculatus</taxonomicName>
(Mocquard, 1890)]: Larger adult size (41.21
<normalizedToken originalValue="±">+/-</normalizedToken>
2.5, 30.4-52.6), presence of eight presacral vertebrae, absence of tympanum and parotoid glands, absence of webbing between the fingers, over 50 eggs/ovary and absence of an external vocal sac in males (
<bibRefCitation author="Matsui, M" journalOrPublisher="Zoological Science" pageId="21" pageNumber="78" pagination="1159 - 1166" title="Taxonomic relationships of Ansoniaanotis Inger, Tan, Yambun, 2001 and Pedostibesmaculatus (Mocquard, 1890), with a description of a new genus (Amphibia, Bufonidae)." url="10.2108/zsj.24.1159" volume="24" year="2007">Matsui et al. 2007</bibRefCitation>
). Distribution: endemic to Borneo (
<bibRefCitation author="Frost, DR" journalOrPublisher="Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenchaftliche Classe" pageId="19" pageNumber="76" title="Amphibian Species of the World: an Online Reference. Version 6" url="http://research.amnh.org/vz/herpetology/amphibia" year="2014">Frost 2014</bibRefCitation>
).
</paragraph>
<paragraph lastPageId="12" lastPageNumber="69" pageId="11" pageNumber="68">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Duttaphrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Duttaphrynus" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="genus">Duttaphrynus</taxonomicName>
Frost, Grant, Faivovich, Bain, Haas, Haddad, de
<normalizedToken originalValue="Sá">Sa</normalizedToken>
, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green &amp; Wheeler, 2006: type
<normalizedToken originalValue="species">species-</normalizedToken>
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo melanostictus" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="melanostictus">Bufo melanostictus</taxonomicName>
[current name combination:
<taxonomicName class="Amphibia" family="Bufonidae" genus="Duttaphrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Duttaphrynus melanostictus" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="melanostictus">Duttaphrynus melanostictus</taxonomicName>
(Schneider, 1799)]: Large adult size (mean SVL 43.7 mm), presence of eight presacral vertebrae, presence of keratinised cephalic ridges in some species, presence of large, keratinised warts on the dorsum, absence of expanded discs in finger and toe tips, absence of webbing between the fingers, numerous black pigmented ova laid in long, continuous strings, exotrophic larvae and terrestrial habit (
<bibRefCitation author="Dubois, A" journalOrPublisher="Journal of South Asian Natural History" pageId="19" pageNumber="76" pagination="133 - 180" title="Asian and Oriental toads of the Bufomelanostictus, Bufoscaber and Bufostejnegeri groups (Amphibia, Anura): a list of available names and redescription of some name-bearing types." volume="4" year="1999">Dubois and Ohler 1999</bibRefCitation>
,
<bibRefCitation author="Manamendra-Arachchi, K" journalOrPublisher="Journal of South Asian Natural History" pageId="21" pageNumber="78" pagination="213 - 246" title="A synopsis of the Sri Lankan Bufonidae (Amphibia: Anura) with description of two new species." volume="3" year="1998">Manamendra-Arachchi and Pethiyagoda 1998</bibRefCitation>
). Particularly, the nomen
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo camortensis" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="camortensis">Bufo camortensis</taxonomicName>
(holotype - ZSI A 6955) erected for a species that is currently considered to represent
<taxonomicName class="Amphibia" family="Bufonidae" genus="Duttaphrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Duttaphrynus melanostictus" order="Anura" pageId="11" pageNumber="68" phylum="Chordata" rank="species" species="melanostictus">Duttaphrynus melanostictus</taxonomicName>
differs from the new taxon described here by its
<pageBreakToken pageId="12" pageNumber="69" start="start">considerably</pageBreakToken>
large adult size [SVL - 67 mm (vs. much smaller mean adult size of 24 mm in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Blythophryne</taxonomicName>
gen. n.), presence of keratinised cephalic ridges and glandular tubercles on the body (vs. absent in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Blythophryne</taxonomicName>
gen. n.), absence of webbing between the fingers and dilated terminal discs in the digits (vs. present in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Blythophryne</taxonomicName>
gen. n.). Distribution: East Africa through the Middle East, India, Indochina, east to the Sundas till Bali (
<bibRefCitation author="Frost, DR" journalOrPublisher="Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenchaftliche Classe" pageId="19" pageNumber="76" title="Amphibian Species of the World: an Online Reference. Version 6" url="http://research.amnh.org/vz/herpetology/amphibia" year="2014">Frost 2014</bibRefCitation>
).
</paragraph>
<paragraph pageId="12" pageNumber="69">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ansonia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ansonia" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Ansonia</taxonomicName>
Stoliczka, 1870: type species -
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ansonia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ansonia penangensis" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="penangensis">Ansonia penangensis</taxonomicName>
Stoliczka, 1870: small to medium adult size (35-40 mm), absence of (or rudimentary) webbing between the fingers, presence of eight presacral vertebrae, absence of dilations in finger and toe tips, absence of parotoid glands, exotropic larvae with prominent oral discs and torrential stream dwelling habit (
<bibRefCitation author="Inger, RF" journalOrPublisher="Fieldiana Zoology" pageId="20" pageNumber="77" pagination="473 - 503" title="A review of the Oriental toads of the genus Ansonia Stoliczka." url="10.5962/bhl.title.2697" volume="39" year="1960">Inger 1960</bibRefCitation>
,
<bibRefCitation author="Matsui, M" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="21" pageNumber="78" pagination="561 - 570" title="Phylogenetic relationships of Ansonia from Southeast Asia inferred from mitochondrial DNA sequences: systematic and biogeographic implications (Anura: Bufonidae)." url="10.1016/j.ympev.2009.08.003" volume="54" year="2010">Matsui et al. 2010</bibRefCitation>
). Distribution: Indo-Malayan region and the Philippines (
<bibRefCitation author="Frost, DR" journalOrPublisher="Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenchaftliche Classe" pageId="19" pageNumber="76" title="Amphibian Species of the World: an Online Reference. Version 6" url="http://research.amnh.org/vz/herpetology/amphibia" year="2014">Frost 2014</bibRefCitation>
).
</paragraph>
<paragraph pageId="12" pageNumber="69">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Adenomus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Adenomus" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Adenomus</taxonomicName>
Cope, 1861: type
<normalizedToken originalValue="species">species-</normalizedToken>
<taxonomicName class="Amphibia" family="Bufonidae" genus="Adenomus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Adenomus badioflavus" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="badioflavus">Adenomus badioflavus</taxonomicName>
Cope, 1861, a junior synonym of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo kelaartii" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="kelaartii">Bufo kelaartii</taxonomicName>
[current name combination:
<taxonomicName class="Amphibia" family="Bufonidae" genus="Adenomus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Adenomus kelaarti" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="kelaarti">Adenomus kelaarti</taxonomicName>
(
<normalizedToken originalValue="Günther">Guenther</normalizedToken>
, 1858)]: The genus
<taxonomicName class="Amphibia" family="Bufonidae" genus="Adenomus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Adenomus" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Adenomus</taxonomicName>
was resurrected from the synonymy of '
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Bufo</taxonomicName>
' by
<bibRefCitation author="Manamendra-Arachchi, K" journalOrPublisher="Journal of South Asian Natural History" pageId="21" pageNumber="78" pagination="213 - 246" title="A synopsis of the Sri Lankan Bufonidae (Amphibia: Anura) with description of two new species." volume="3" year="1998">Manamendra-Arachchi and Pethiyagoda (1998)</bibRefCitation>
to accommodate members of the
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo kelaarti" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="kelaarti">' Bufo ' kelaarti</taxonomicName>
group, characterised by smooth finger edges; differing from the new taxon described here by its larger adult size (mean SVL 38.4 mm), presence of seven presacral vertebrae, absence of sheath-like webbing between fingers, absence of expanded discs at digit tips, presence of cranial ridges and indistinct tympanum (in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Adenomus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Adenomus kelaarti" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="kelaarti">Adenomus kelaarti</taxonomicName>
), terrestrial habit, pronounced sexual size dimorphism and unpigmented ova laid in long, continuous strings as in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Duttaphrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Duttaphrynus" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Duttaphrynus</taxonomicName>
(
<bibRefCitation author="Manamendra-Arachchi, K" journalOrPublisher="Journal of South Asian Natural History" pageId="21" pageNumber="78" pagination="213 - 246" title="A synopsis of the Sri Lankan Bufonidae (Amphibia: Anura) with description of two new species." volume="3" year="1998">Manamendra-Arachchi and Pethiyagoda 1998</bibRefCitation>
;
<bibRefCitation author="Haas, W" journalOrPublisher="Elaphe" pageId="20" pageNumber="77" pagination="16 - 19" title="Zur Biologie von Bufokelaartii Guenther, 1859." volume="7" year="1999">Haas 1999</bibRefCitation>
;
<bibRefCitation author="Meegaskumbura, M" journalOrPublisher="Zootaxa" pageId="21" pageNumber="78" pagination="245 - 261" title="The Sri Lankan torrent toads (Bufonidae: Adenominae: Adenomus): species boundaries assessed using multiple criteria." url="10.11646/zootaxa.3911.2.6" volume="3911" year="2015">Meegaskumbura et al. 2015</bibRefCitation>
). Distribution: endemic to Sri Lanka (
<bibRefCitation author="Frost, DR" journalOrPublisher="Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenchaftliche Classe" pageId="19" pageNumber="76" title="Amphibian Species of the World: an Online Reference. Version 6" url="http://research.amnh.org/vz/herpetology/amphibia" year="2014">Frost 2014</bibRefCitation>
).
</paragraph>
<paragraph pageId="12" pageNumber="69">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ghatophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ghatophryne" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Ghatophryne</taxonomicName>
Biju, Bocxlaer, Giri, Loader &amp; Bossuyt, 2009: type
<normalizedToken originalValue="species">species-</normalizedToken>
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ansonia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ansonia ornata" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="ornata">Ansonia ornata</taxonomicName>
[current name combination:
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ghatophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ghatophryne ornata" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="ornata">Ghatophryne ornata</taxonomicName>
(
<normalizedToken originalValue="Günther">Guenther</normalizedToken>
, 1876)]: larger adult size (up to 35 mm SVL), characteristic reddish dorsal and ventral colouration, absence of parotoid glands, absence of webbing between the fingers, finger tips not dilated to discs and torrential stream dwelling habit (
<bibRefCitation author="Biju, SD" journalOrPublisher="BMC Research Notes" pageId="18" pageNumber="75" title="Two new endemic genera and one new species of toad (Anura: Bufonidae) from the Western Ghats of India." url="10.1186/1756-0500-2-241" volume="2" year="2009">Biju et al. 2009</bibRefCitation>
). Distribution: Central Western Ghats in the states of Kerala and Karnataka (
<bibRefCitation author="Frost, DR" journalOrPublisher="Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenchaftliche Classe" pageId="19" pageNumber="76" title="Amphibian Species of the World: an Online Reference. Version 6" url="http://research.amnh.org/vz/herpetology/amphibia" year="2014">Frost 2014</bibRefCitation>
).
</paragraph>
<paragraph pageId="12" pageNumber="69">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Xanthophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Xanthophryne" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Xanthophryne</taxonomicName>
Biju Bocxlaer, Giri, Loader &amp; Bossuyt, 2009: type
<normalizedToken originalValue="species">species-</normalizedToken>
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo koynaensis" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="koynaensis">Bufo koynaensis</taxonomicName>
[current name combination:
<taxonomicName class="Amphibia" family="Bufonidae" genus="Xanthophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Xanthophryne koynaensis" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="koynaensis">Xanthophryne koynaensis</taxonomicName>
(Soman, 1963)]: Larger adult size (up to 35.3 mm SVL), presence of characteristic chrome yellow patches along the flanks and sides of the abdomen, indistinct tympanum, weak, rounded parotoid glands, absence of webbing in fingers and discs in toes and fingers; large, pigmented ova laid in stagnant puddles on the ground (
<bibRefCitation author="Biju, SD" journalOrPublisher="BMC Research Notes" pageId="18" pageNumber="75" title="Two new endemic genera and one new species of toad (Anura: Bufonidae) from the Western Ghats of India." url="10.1186/1756-0500-2-241" volume="2" year="2009">Biju et al. 2009</bibRefCitation>
). Distribution: Known only from Northern Western Ghats in Maharashtra, India (
<bibRefCitation author="Frost, DR" journalOrPublisher="Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenchaftliche Classe" pageId="19" pageNumber="76" title="Amphibian Species of the World: an Online Reference. Version 6" url="http://research.amnh.org/vz/herpetology/amphibia" year="2014">Frost 2014</bibRefCitation>
).
</paragraph>
<paragraph pageId="12" pageNumber="69">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Leptophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptophryne" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Leptophryne</taxonomicName>
Fitzinger, 1843: type species -
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo cruentatus" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="cruentatus">Bufo cruentatus</taxonomicName>
[current name combination:
<taxonomicName class="Amphibia" family="Bufonidae" genus="Leptophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptophryne cruentata" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="cruentata">Leptophryne cruentata</taxonomicName>
(Tschudi, 1838)]:
<bibRefCitation author="Dubois, A" journalOrPublisher="Journal of Herpetology" pageId="19" pageNumber="76" pagination="173 - 174" title="Leptophryne Fitzinger, 1843, a senior synonym of Cacophryne Davis, 1935 (Bufonidae)." url="10.2307/1563811" volume="16" year="1982">Dubois (1982)</bibRefCitation>
resurrected the genus
<taxonomicName class="Amphibia" family="Bufonidae" genus="Leptophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptophryne" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Leptophryne</taxonomicName>
Fitzinger, 1843 as the senior synonym of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Cacophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cacophryne" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="genus">Cacophryne</taxonomicName>
Davis, 1935, which currently comprises two species -
<taxonomicName class="Amphibia" family="Bufonidae" genus="Leptophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptophryne borbonica" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="borbonica">Leptophryne borbonica</taxonomicName>
(Tschudi, 1838) and
<taxonomicName class="Amphibia" family="Bufonidae" genus="Leptophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptophryne cruentata" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="cruentata">Leptophryne cruentata</taxonomicName>
(Tschudi, 1838). Presence of eight presacral vertebrae; firmisternal pectoral girdle; elongate subarticular tubercles near the base of each toe, numerous pigmented eggs and exotrophic larvae (
<bibRefCitation author="Fei, L" journalOrPublisher="Acta Zootaxonomica Sinica" pageId="19" pageNumber="76" pagination="762 - 766" title="A new bufonid genus Parapelophryne from China (Amphibia, Anura)." volume="28" year="2003">Fei et al. 2003</bibRefCitation>
) distinguish it from
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="12" pageNumber="69" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. Distribution: Sundaland (
<bibRefCitation author="Frost, DR" journalOrPublisher="Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenchaftliche Classe" pageId="19" pageNumber="76" title="Amphibian Species of the World: an Online Reference. Version 6" url="http://research.amnh.org/vz/herpetology/amphibia" year="2014">Frost 2014</bibRefCitation>
).
</paragraph>
<paragraph pageId="13" pageNumber="70">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pseudobufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudobufo" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">
<pageBreakToken pageId="13" pageNumber="70" start="start">Pseudobufo</pageBreakToken>
</taxonomicName>
Tschudi, 1838: type species -
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pseudobufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudobufo subasper" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="subasper">Pseudobufo subasper</taxonomicName>
Tschudi, 1838: Large body size, stout habitus; presence of seven presacral vertebrae (vs. six in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Blythophryne</taxonomicName>
gen. n.) completely (to the tip of Toe IV) webbed feet (vs. incomplete toe webbing in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n.), fingers basally webbed; parotoid glands absent; dorsal, lateral and ventral skin surfaces with fine spinules, dorsoventrally depressed body with large, round warts and dorsally positioned nostrils (vs. lateral) distinguish it from the new genus described here (
<bibRefCitation author="Fei, L" journalOrPublisher="Acta Zootaxonomica Sinica" pageId="19" pageNumber="76" pagination="762 - 766" title="A new bufonid genus Parapelophryne from China (Amphibia, Anura)." volume="28" year="2003">Fei et al. 2003</bibRefCitation>
;
<bibRefCitation author="Inger, RF" journalOrPublisher="Natural History Publications (Borneo), Kota Kinabalu" pageId="20" pageNumber="77" pagination="1 - 201" title="A field guide to the frogs of Borneo" year="2005">Inger and Stuebing 2005</bibRefCitation>
). Distribution: Sundaland.
</paragraph>
<paragraph pageId="13" pageNumber="70">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ingerophrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ingerophrynus" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Ingerophrynus</taxonomicName>
Frost, Grant, Faivovich, Bain, Haas, Haddad, de
<normalizedToken originalValue="Sá">Sa</normalizedToken>
, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green &amp; Wheeler, 2006: type
<normalizedToken originalValue="species">species-</normalizedToken>
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo biporcatus" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="biporcatus">Bufo biporcatus</taxonomicName>
[current name combination:
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ingerophrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ingerophrynus biporcatus" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="biporcatus">Ingerophrynus biporcatus</taxonomicName>
(Gravenhorst, 1829)]: Presence of seven presacral vertebrae (vs. six); absence of lateral dilations in the digit tips (vs. present); absence of webbing between the fingers (vs. present) and endotrophic (vs. exotrophic) larvae distinguish
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. from
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ingerophrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ingerophrynus" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Ingerophrynus</taxonomicName>
. Distribution: Southern Yunnan, Indochina, the Malay Peninsula, the islands of Indo-Malaya, and Philippines.
</paragraph>
<paragraph pageId="13" pageNumber="70">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Phrynoidis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynoidis" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Phrynoidis</taxonomicName>
Fitzinger, 1843: type species -
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo asper" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="asper">Bufo asper</taxonomicName>
[current name combination:
<taxonomicName class="Amphibia" family="Bufonidae" genus="Phrynoidis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynoidis asper" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="asper">Phrynoidis asper</taxonomicName>
(Gravenhorst, 1829)]: Large adult size (up to 100 mm SVL) presence of an omosternum, (vs. absent); presence of seven presacral vertebrae (vs. six); absence of lateral dilations of digit tips (vs. present) and exotrophic (vs. endotrophic) larvae distinguish this genus from the new genus
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Blythophryne</taxonomicName>
gen. n. Distribution: Myanmar through western and peninsular Thailand, the Malay Peninsula, Sumatra, Borneo, and Java.
</paragraph>
<paragraph pageId="13" pageNumber="70">Apart from the above bufonid genera known from Oriental Asia, the new taxon described herein differs from the following central-west African genera:</paragraph>
<paragraph pageId="13" pageNumber="70">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophryne" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Nectophryne</taxonomicName>
Buchholz &amp; Peters, 1875: type species -
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophryne afra" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="afra">Nectophryne afra</taxonomicName>
Buchholz &amp; Peters, 1875 by the presence of eight presacral vertebrae (vs. six in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n.); presence of lamelliform subdigital pads - a character unique to
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophryne" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Nectophryne</taxonomicName>
which is absent in the new taxon described here. Oriental forms including members of the genera
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Pedostibes</taxonomicName>
and
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pelophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelophryne" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Pelophryne</taxonomicName>
were attributed to
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophryne" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Nectophryne</taxonomicName>
earlier (
<bibRefCitation author="Boulenger, GA" journalOrPublisher="Proceedings of the Zoological Society of London" pageId="18" pageNumber="75" pagination="505 - 508" title="An account of the reptiles and batrachians collected by Mr. C. Hose on Mt. Dulit, Borneo." volume="1892" year="1892">Boulenger 1892</bibRefCitation>
,
<bibRefCitation author="Boulenger, GA" journalOrPublisher="Annals and Magazine of Natural History (Series 6)" pageId="18" pageNumber="75" pagination="449 - 450" title="Descriptions of two new batrachians obtained by Mr. Everett on Mt. Kina Balu, North Borneo." volume="17" year="1896">1896</bibRefCitation>
,
<bibRefCitation author="Boulenger, GA" journalOrPublisher="Records of the Indian Museum" pageId="18" pageNumber="75" pagination="207 - 208" title="Descriptions of three new batrachians from the Garo Hills, Assam." volume="16" year="1919">1919</bibRefCitation>
), until
<bibRefCitation author="Barbour, T" journalOrPublisher="Proceedings of the Biological Society of Washington" pageId="18" pageNumber="75" pagination="191 - 196" title="Notes on the &quot; Nectophryne &quot;." volume="51" year="1938">Barbour (1938)</bibRefCitation>
redefined these genera.
</paragraph>
<paragraph pageId="13" pageNumber="70">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophrynoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophrynoides" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Nectophrynoides</taxonomicName>
Noble, 1926: type species -
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophryne tornieri" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="tornieri">Nectophryne tornieri</taxonomicName>
[current name combination:
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophrynoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophrynoides tornieri" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="tornieri">Nectophrynoides tornieri</taxonomicName>
(Roux, 1906)]: The comparisons made here are restricted to the type species of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophrynoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophrynoides" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Nectophrynoides</taxonomicName>
because the genus is poorly defined and is composed of representatives with a broad spectrum of morphological and developmental characteristics. Though unique among bufonids in possessing an omosternum and a direct developmental mode (in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophrynoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophrynoides viviparus" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="viviparus">Nectophrynoides viviparus</taxonomicName>
), members of this genus are poorly diagnosed with respect to other genera (
<bibRefCitation author="Menegon, M" journalOrPublisher="Tropical Zoology" pageId="21" pageNumber="78" pagination="97 - 121" title="Five new species of Nectophrynoides Noble, 1926 (Amphibia: Anura, Bufonidae) from the Eastern Arc Mountains, Tanzania." url="10.1080/03946975.2004.10531201" volume="17" year="2004">Menegon et al. 2004</bibRefCitation>
). Larger adult size (SVL 21-30 mm), presence of expanded, truncate fingertips (vs. expanded and curved in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n.), presence of eight presacral vertebrae (vs. 6 in
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n.) however, distinguish
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophrynoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophrynoides" order="Anura" pageId="13" pageNumber="70" phylum="Chordata" rank="genus">Nectophrynoides</taxonomicName>
from the new taxon described here (see
<bibRefCitation author="Tihen, JA" journalOrPublisher="Copeia" pageId="22" pageNumber="79" pagination="225 - 233" title="Two new genera of African bufonids, with remarks on the phylogeny of related genera." url="10.2307/1439662" volume="1960" year="1960">Tihen 1960</bibRefCitation>
;
<bibRefCitation author="Menegon, M" journalOrPublisher="Tropical Zoology" pageId="21" pageNumber="78" pagination="97 - 121" title="Five new species of Nectophrynoides Noble, 1926 (Amphibia: Anura, Bufonidae) from the Eastern Arc Mountains, Tanzania." url="10.1080/03946975.2004.10531201" volume="17" year="2004">Menegon et al. 2004</bibRefCitation>
;
<bibRefCitation author="Harper, EB" journalOrPublisher="Camerapix Publishers International, Nairobi" pageId="20" pageNumber="77" title="Field guide to amphibians of the Eastern Arc Mountains and coastal forests of Tanzania and Kenya." year="2010">Harper et al. 2010</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="14" lastPageNumber="71" pageId="13" pageNumber="70" type="molecular phylogeny">
<paragraph pageId="13" pageNumber="70">Molecular phylogeny.</paragraph>
<paragraph lastPageId="14" lastPageNumber="71" pageId="13" pageNumber="70">
Multiple sequence alignment of the 16S homologous regions resulted in 498 conserved sites and 246 parsimoniously informative sites. In the
<pageBreakToken pageId="14" pageNumber="71" start="start">phylogenetic</pageBreakToken>
analysis using both Maximum likelihood and Bayesian inference, the focal taxon showed a unique taxonomic position. The phylograms of both inference methods were similar (Fig. 7).
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufoides meghalayanus" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="meghalayanus">Bufoides meghalayanus</taxonomicName>
was found to be the closest taxon to the focal species,
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n. in the tree generated using 36 species from Asia and Africa but with relatively low support (Fig. 7a). However, when analysed with more of the Asian toads, it clearly separates out from species of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Duttaphrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Duttaphrynus" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Duttaphrynus</taxonomicName>
, as well as, those of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Xanthophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Xanthophryne" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Xanthophryne</taxonomicName>
and
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufoides" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Bufoides</taxonomicName>
(Fig. 7b, c). The average within-genus pairwise K2p distances at the partial 16S rRNA gene for all the described genera considered under this study was 0.0642, with 99% confidence interval (CI) of 0.0512-0.0687 (Table 6). The average pairwise k2p distance of the focal species with all other taxa at partial 16S rRNA gene considered here was 0.103, with a 99% CI of 0.096-0.113, strongly supporting its distinctiveness and unique phylogenetic position within the Bufonids. Similarly, for partial 12S rRNA gene, the average within-genus pairwise K2p distances for all described genera was 0.0495, with the 99% CI of 0.0387-0.0603. The average pairwise k2p distance of the focal species with all other taxa at partial 12S rRNA gene was 0.0783, with a 99% CI of 0.072-0.085. Both tree-based and distance-based analyses clearly indicate the uniqueness of its phylogenetic position. Thus, the rDNA typing strongly suggest the new taxon as a candidate to be named as a new genus/species.
</paragraph>
<caption pageId="14" pageNumber="71">
<paragraph pageId="14" pageNumber="71">
Figure 7. Phylogenetic position of
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n., inferred from concatenated partial 12S and 16S rDNA sequences. The posterior probabilities for Bayesian Inference (BI) and the bootstrap support values for the ML are given as (BI /ML) above and below the branch nodes. a The tree was generated using 36 species related to 21 genera, and was rooted using
<taxonomicName genus="Rhabeo" lsidName="Rhabeo gutattus" pageId="14" pageNumber="71" rank="species" species="gutattus">Rhabeo gutattus</taxonomicName>
as outgroup b the subclade containing the Indian and Sri Lankan toads (7 genera, 17 species) rooted using
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ghatophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ghatophryne ornata" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="ornata">Ghatophryne ornata</taxonomicName>
as outgroup; and c the subclade containing the Indian and Sri Lankan toads (5 genera, 15 species) rooted using
<taxonomicName class="Amphibia" family="Bufonidae" genus="Adenomus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Adenomus kelaartii" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="kelaartii">Adenomus kelaartii</taxonomicName>
as outgroup.
</paragraph>
</caption>
<caption pageId="14" pageNumber="71">
<paragraph pageId="14" pageNumber="71">
Table 6. The average uncorrected K2p distance estimates across genera of
<taxonomicName family="Bufonidae" lsidName="" pageId="14" pageNumber="71" rank="family">Bufonidae</taxonomicName>
based on partial 16S (lower triangular matrix) and 12S (upper triangular matrix) rDNA sequences, calculated using MEGA6.06; within genus K2p distances are given along the diagonal (left value for 16S and right value for 12S).
</paragraph>
</caption>
<paragraph pageId="14" pageNumber="71">
<table pageId="14" pageNumber="71">
<tr pageId="14" pageNumber="71">
<th colspan="21" pageId="14" pageNumber="71" rowspan="1">16S/12S k2p uncorrected pair-wise distance estimates</th>
</tr>
<tr pageId="14" pageNumber="71">
<th colspan="2" pageId="14" pageNumber="71" rowspan="1">Taxa (Genus*/Species)</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">1</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">2</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">3</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">4</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">5</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">6</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">7</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">8</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">9</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">10</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">11</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">12</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">13</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">14</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">15</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">16</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">17</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">18</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">19</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">20</th>
<th colspan="1" pageId="14" pageNumber="71" rowspan="1">21</th>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Amietophrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Amietophrynus" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Amietophrynus</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">0.068/0.044</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ansonia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ansonia" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Ansonia</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">0.066/0.060</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Duttaphrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Duttaphrynus" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Duttaphrynus</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">0.049/ 0.038</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ingerophrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ingerophrynus" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Ingerophrynus</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">0.056/0.077</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Mertensophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mertensophryne" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Mertensophryne</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">0.059/ 0.061</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophryne" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Nectophryne</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">0.089/0.049</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Nectophrynoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nectophrynoides" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Nectophrynoides</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">0.021/0.015</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pedostibes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pedostibes" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Pedostibes</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">0.076/0.065</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Phrynoidis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynoidis" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="genus">Phrynoidis</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">0.039/0.085</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Ghatophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ghatophryne ornata" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="ornata">Ghatophryne ornata</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Leptophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptophryne borbonica" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="borbonica">Leptophryne borbonica</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Vandijkophrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Vandijkophrynus robinsoni" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="robinsoni">Vandijkophrynus robinsoni</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Schismaderma" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Schismaderma carens" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="carens">Schismaderma carens</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo bufo" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="bufo">Bufo bufo</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Sabahphrynus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sabahphrynus maculatus" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="maculatus">Sabahphrynus maculatus</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Pelophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelophryne api" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="api">Pelophryne api</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufoides meghalayanus" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="meghalayanus">Bufoides meghalayanus</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Adenomus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Adenomus kelaartii" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="kelaartii">Adenomus kelaartii</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Xanthophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Xanthophryne koynayensis" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="koynayensis">Xanthophryne koynayensis</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Rhaebo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhaebo guttatus" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="guttatus">Rhaebo guttatus</taxonomicName>
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
<tr pageId="14" pageNumber="71">
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">
<taxonomicName class="Amphibia" family="Bufonidae" genus="Blythophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blythophryne beryet" order="Anura" pageId="14" pageNumber="71" phylum="Chordata" rank="species" species="beryet">Blythophryne beryet</taxonomicName>
gen. et sp. n.
</td>
<td colspan="1" pageId="14" pageNumber="71" rowspan="1">n/a</td>
</tr>
</table>
</paragraph>
</subSubSection>
</treatment>
</document>