589 lines
58 KiB
XML
589 lines
58 KiB
XML
<document ID-DOI="http://dx.doi.org/10.3897/vz.72.e89647" ID-GBIF-Dataset="e83eba07-b611-4b55-b94a-96f2059d0efd" ID-Pensoft-Pub="2625-8498-72-997" ID-Pensoft-UUID="F478C7D1544650D99649E511AB6CED15" ID-ZooBank="0CA196A200E04E0AB21611AD5B3856EB" ModsDocID="2625-8498-72-997" checkinTime="1668159806196" checkinUser="pensoft" docAuthor="Kraus, Fred, Kaiser, Hinrich & O'Shea, Mark" docDate="2022" docId="BDA3BC47EB6C5C898B9757B513965967" docLanguage="en" docName="VertZool 72: 997-1034" docOrigin="Vertebrate Zoology 72" docPubDate="2022-11-10" docSource="http://dx.doi.org/10.3897/vz.72.e89647" docTitle="Toxicocalamus atratus Kraus & Kaiser & O’Shea 2022, sp. nov." docType="treatment" docUuid="2DED0238-6F17-414E-BEF3-801E852AB976" docUuidSource="ZooBank" docVersion="2" id="F478C7D1544650D99649E511AB6CED15" lastPageNumber="997" masterDocId="F478C7D1544650D99649E511AB6CED15" masterDocTitle="Hidden diversity in semi-fossorial Melanesian forest snakes: A revision of the Toxicocalamus loriae complex (Squamata, Elapidae) from New Guinea" masterLastPageNumber="1034" masterPageNumber="997" pageNumber="997" updateTime="1668247209537" updateUser="ExternalLinkService">
|
||
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
|
||
<mods:titleInfo>
|
||
<mods:title>Hidden diversity in semi-fossorial Melanesian forest snakes: A revision of the Toxicocalamus loriae complex (Squamata, Elapidae) from New Guinea</mods:title>
|
||
</mods:titleInfo>
|
||
<mods:name type="personal">
|
||
<mods:role>
|
||
<mods:roleTerm>Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart>Kraus, Fred</mods:namePart>
|
||
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0003-4194-4959</mods:nameIdentifier>
|
||
<mods:affiliation>Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, Michigan, USA</mods:affiliation>
|
||
</mods:name>
|
||
<mods:name type="personal">
|
||
<mods:role>
|
||
<mods:roleTerm>Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart>Kaiser, Hinrich</mods:namePart>
|
||
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-0001-9428</mods:nameIdentifier>
|
||
<mods:affiliation>Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA & Department of Vertebrate Zoology, Leibniz-Institut zur Analyse des Biodiversitaetswandels, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany</mods:affiliation>
|
||
<mods:nameIdentifier type="email">chalcopis@yahoo.com</mods:nameIdentifier>
|
||
</mods:name>
|
||
<mods:name type="personal">
|
||
<mods:role>
|
||
<mods:roleTerm>Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart>O'Shea, Mark</mods:namePart>
|
||
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-1566-7460</mods:nameIdentifier>
|
||
<mods:affiliation>Faculty of Science and Engineering, University of Wolverhampton, Wulfruna Street, Wolverhampton, West Midlands WV 1 1 LY, United Kingdom</mods:affiliation>
|
||
</mods:name>
|
||
<mods:typeOfResource>text</mods:typeOfResource>
|
||
<mods:relatedItem type="host">
|
||
<mods:titleInfo>
|
||
<mods:title>Vertebrate Zoology</mods:title>
|
||
</mods:titleInfo>
|
||
<mods:part>
|
||
<mods:date>2022</mods:date>
|
||
<mods:detail type="pubDate">
|
||
<mods:number>2022-11-10</mods:number>
|
||
</mods:detail>
|
||
<mods:detail type="volume">
|
||
<mods:number>72</mods:number>
|
||
</mods:detail>
|
||
<mods:extent unit="page">
|
||
<mods:start>997</mods:start>
|
||
<mods:end>1034</mods:end>
|
||
</mods:extent>
|
||
</mods:part>
|
||
</mods:relatedItem>
|
||
<mods:location>
|
||
<mods:url>http://dx.doi.org/10.3897/vz.72.e89647</mods:url>
|
||
</mods:location>
|
||
<mods:classification>journal article</mods:classification>
|
||
<mods:identifier type="DOI">http://dx.doi.org/10.3897/vz.72.e89647</mods:identifier>
|
||
<mods:identifier type="Pensoft-Pub">2625-8498-72-997</mods:identifier>
|
||
<mods:identifier type="ZooBank">0CA196A200E04E0AB21611AD5B3856EB</mods:identifier>
|
||
<mods:identifier type="Pensoft-UUID">F478C7D1544650D99649E511AB6CED15</mods:identifier>
|
||
</mods:mods>
|
||
<subSection pageId="0" pageNumber="997" type="taxonomic accounts">
|
||
<treatment ID-GBIF-Taxon="204928504" LSID="urn:lsid:zoobank.org:act:2DED0238-6F17-414E-BEF3-801E852AB976" httpUri="http://treatment.plazi.org/id/BDA3BC47EB6C5C898B9757B513965967" lastPageNumber="997" pageId="0" pageNumber="997">
|
||
<subSubSection pageId="0" pageNumber="997" type="nomenclature">
|
||
<paragraph pageId="0" pageNumber="997">
|
||
<taxonomicName LSID="https://zoobank.org/2DED0238-6F17-414E-BEF3-801E852AB976" authority="Kraus & Kaiser & O’Shea, 2022" authorityName="Kraus & Kaiser & O’Shea" authorityYear="2022" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus atratus" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="atratus" status="sp. nov.">Toxicocalamus atratus</taxonomicName>
|
||
<taxonomicNameLabel pageId="0" pageNumber="997">sp. nov.</taxonomicNameLabel>
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="description">
|
||
<paragraph pageId="0" pageNumber="997">
|
||
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Dorsal and right lateral views of heads of juveniles of Toxicocalamus loriae Group snakes. A, B T. loriae (BPBM 10966), Agaun, Milne Bay Province, PNG. C, D T. nymani comb. nov., spotted form (BPBM 5442), Kalolo, Morobe Province, PNG. E, F T. nymani, dark form (BPBM 23699), Wau, Morobe Province, PNG. G, H T. lamingtoni comb. nov. (AMS R 9352), Mt. Lamington, Oro Province, PNG. I, J T. vertebralis sp. nov. (KU 129086), Wau, Morobe Province, PNG. K, L T. spilorhynchus sp. nov. (AMNH R- 107204), Garaina, Morobe Province, PNG. M, N T. atratus sp. nov. (CAS 118958), Mintima, Chimbu Province, PNG." figureDoi="10.3897/vz.72.e89647.figure4" httpUri="https://binary.pensoft.net/fig/768170" pageId="0" pageNumber="997">Figs 4M, N</figureCitation>
|
||
<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Toxicocalamus loriae Group species in live and perimortem photographs to show colour in life. A, B T. vertebralis sp. nov. (CAS 140042) from Kol, Jimi Valley, Jiwaka Province, PNG. C, D T. spilorhynchus sp. nov. (MCZ R- 152431) from Garaina, Morobe Province, PNG. E T. atratus sp. nov. (MCZ R- 84026) from Kundiawa, Wahgi Valley, Chimbu Province, PNG. F T. atratus (MCZ R- 111767) from Kundiawa, Wahgi Valley, Chimbu Province, PNG. Photographs by Fred Parker." figureDoi="10.3897/vz.72.e89647.figure11" httpUri="https://binary.pensoft.net/fig/768178" pageId="0" pageNumber="997">, 11E, F</figureCitation>
|
||
<figureCitation captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Head of the Toxicocalamus atratus sp. nov. holotype (MCZ R- 84144) from Kundiawa, Chimbu Province, PNG, in both photographic and line-drawing illustrations. A, A ' dorsal, B, B ' right lateral, C, C ' left lateral, and D, D ' ventral views." figureDoi="10.3897/vz.72.e89647.figure14" httpUri="https://binary.pensoft.net/fig/768174" pageId="0" pageNumber="997">, 14</figureCitation>
|
||
<figureCitation captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. A Dorsum and B venter of the Toxicocalamus atratus sp. nov. holotype (MCZ R- 84144) from Kundiawa, Chimbu Province, PNG." figureDoi="10.3897/vz.72.e89647.figure15" httpUri="https://binary.pensoft.net/fig/768181" pageId="0" pageNumber="997">, 15</figureCitation>
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="reference_group">
|
||
<paragraph pageId="0" pageNumber="997">
|
||
<taxonomicName authority="Kraus & Kaiser & O’Shea, 2022" authorityName="Kraus & Kaiser & O’Shea" authorityYear="2022" baseAuthorityName="Kraus & Kaiser & O’Shea" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus (Apistocalamus) loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae" status="sp. nov." subGenus="Apistocalamus">Toxicocalamus (Apistocalamus) loriae</taxonomicName>
|
||
(part) - McDowell, 1969: 485.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="holotype">
|
||
<paragraph pageId="0" pageNumber="997">Holotype.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
MCZ R-84144, an adult female collected by Fred Parker at Kundiawa,
|
||
<geoCoordinate degrees="6.02" direction="south" orientation="latitude" precision="555" value="-6.02">6.02°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.97" direction="east" orientation="longitude" precision="555" value="144.97">144.97°E</geoCoordinate>
|
||
, elevation 1585 m, Chimbu Province, Papua New Guinea on 25 May 1964.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="paratypes (n = 90)">
|
||
<paragraph pageId="0" pageNumber="997">
|
||
Paratypes (
|
||
<emphasis italics="true" pageId="0" pageNumber="997">n</emphasis>
|
||
= 90).
|
||
</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
Papua New Guinea: Chimbu Prov-ince: along Wahgi River,
|
||
<geoCoordinate degrees="5.94" direction="south" orientation="latitude" precision="555" value="-5.94">5.94°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.80" direction="east" orientation="longitude" precision="555" value="144.8">144.80°E</geoCoordinate>
|
||
, elev. 1470 m, CAS 113665, 139564; Karimui,
|
||
<geoCoordinate degrees="6.50" direction="south" orientation="latitude" precision="555" value="-6.5">6.50°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.83" direction="east" orientation="longitude" precision="555" value="144.83">144.83°E</geoCoordinate>
|
||
, elev. 1170 m, CAS 118961-62; Kondiu,
|
||
<geoCoordinate degrees="5.98" direction="south" orientation="latitude" precision="555" value="-5.98">5.98°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.87" direction="east" orientation="longitude" precision="555" value="144.87">144.87°E</geoCoordinate>
|
||
, elev. 1600 m, AMNH R-75336-57; Kundiawa,
|
||
<geoCoordinate degrees="6.02" direction="south" orientation="latitude" precision="555" value="-6.02">6.02°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.97" direction="east" orientation="longitude" precision="555" value="144.97">144.97°E</geoCoordinate>
|
||
, elev. 1585 m, AMNH R-98495, R-98497, CAS 99916, 100069, 113670, 115986, 118948, 118960, 139584, 140043, MCZ R-83218, R-84026, R-111767, R-111788, R-115586, R-116774, R-116788, R-140818, R-145923, NMV 13421, USNM 166280; Kup,
|
||
<geoCoordinate degrees="5.95" direction="south" orientation="latitude" precision="555" value="-5.95">5.95°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.80" direction="east" orientation="longitude" precision="555" value="144.8">144.80°E</geoCoordinate>
|
||
, elev. 1500 m, AMNH R-72780-81; Mintima,
|
||
<geoCoordinate degrees="5.98" direction="south" orientation="latitude" precision="555" value="-5.98">5.98°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.91" direction="east" orientation="longitude" precision="555" value="144.91">144.91°E</geoCoordinate>
|
||
, elev. 1770 m, CAS 103374-75, 113663, 118957-59, MCZ R-116789-90, R-121547-48; Noru,
|
||
<geoCoordinate degrees="6.60" direction="south" orientation="latitude" precision="555" value="-6.6">6.60°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.63" direction="east" orientation="longitude" precision="555" value="144.63">144.63°E</geoCoordinate>
|
||
, elev. 1770 m, AMS R115365; Eastern Highlands Province: Agakamatasa,
|
||
<geoCoordinate degrees="6.72" direction="south" orientation="latitude" precision="555" value="-6.72">6.72°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="145.62" direction="east" orientation="longitude" precision="555" value="145.62">145.62°E</geoCoordinate>
|
||
, elev. 1720 m, MCZ R-121545; Lida Patrol Post,
|
||
<geoCoordinate degrees="6.32" direction="south" orientation="latitude" precision="555" value="-6.32">6.32°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="145.40" direction="east" orientation="longitude" precision="555" value="145.4">145.40°E</geoCoordinate>
|
||
, elev. 1800 m, CAS 139575; Lufa, Mt. Michael,
|
||
<geoCoordinate degrees="6.33" direction="south" orientation="latitude" precision="555" value="-6.33">6.33°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="145.25" direction="east" orientation="longitude" precision="555" value="145.25">145.25°E</geoCoordinate>
|
||
, elev. 1120 m, CAS 113666-67, MCZ R-121546; Nivi Unggai,
|
||
<geoCoordinate degrees="6.21" direction="south" orientation="latitude" precision="555" value="-6.21">6.21°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="145.31" direction="east" orientation="longitude" precision="555" value="145.31">145.31°E</geoCoordinate>
|
||
, elev. 2030 m, MCZ R-84142, R-116791 -92; Hela Province: Bobole,
|
||
<geoCoordinate degrees="6.23" direction="south" orientation="latitude" precision="555" value="-6.23">6.23°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="142.77" direction="east" orientation="longitude" precision="555" value="142.77">142.77°E</geoCoordinate>
|
||
, elev. 1145 m, AMS R122803, R122806; Halalinja,
|
||
<geoCoordinate degrees="6.15" direction="south" orientation="latitude" precision="555" value="-6.15">6.15°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="143.37" direction="east" orientation="longitude" precision="555" value="143.37">143.37°E</geoCoordinate>
|
||
, elev. 2140 m, BMNH 1976.92; Nipa,
|
||
<geoCoordinate degrees="6.10" direction="south" orientation="latitude" precision="555" value="-6.1">6.10°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="143.25" direction="east" orientation="longitude" precision="555" value="143.25">143.25°E</geoCoordinate>
|
||
, elev. 2070 m, UPNG 5811; Jiwaka Province: Banz,
|
||
<geoCoordinate degrees="5.78" direction="south" orientation="latitude" precision="555" value="-5.78">5.78°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.62" direction="east" orientation="longitude" precision="555" value="144.62">144.62°E</geoCoordinate>
|
||
, elev. 1650 m, AMNH R-85743, R-88060; Minj,
|
||
<geoCoordinate degrees="5.86" direction="south" orientation="latitude" precision="555" value="-5.86">5.86°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.87" direction="east" orientation="longitude" precision="555" value="144.87">144.87°E</geoCoordinate>
|
||
, elev. 1560 m, MCZ R-141849; Morobe Province: Kwaplalim, 12-13 km W Menyamya,
|
||
<geoCoordinate degrees="7.19" direction="south" orientation="latitude" precision="555" value="-7.19">7.19°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="145.97" direction="east" orientation="longitude" precision="555" value="145.97">145.97°E</geoCoordinate>
|
||
, elev. 1490 m, NMW 37670, UCM 51552-53; Southern Highlands Province: Mendi,
|
||
<geoCoordinate degrees="6.14" direction="south" orientation="latitude" precision="555" value="-6.14">6.14°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="143.66" direction="east" orientation="longitude" precision="555" value="143.66">143.66°E</geoCoordinate>
|
||
, elev. 1750 m, CAS 113664, MCZ R-121543-44; Moro Camp, Lake Kutubu,
|
||
<geoCoordinate degrees="6.36" direction="south" orientation="latitude" precision="555" value="-6.36">6.36°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="143.23" direction="east" orientation="longitude" precision="555" value="143.23">143.23°E</geoCoordinate>
|
||
, elev. 840 m, SAMA R69950; Western Highlands Province: Igindi,
|
||
<geoCoordinate degrees="6.19" direction="south" orientation="latitude" precision="555" value="-6.19">6.19°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.98" direction="east" orientation="longitude" precision="555" value="144.98">144.98°E</geoCoordinate>
|
||
, elev. 1630 m, AMNH R-98134; Kimil,
|
||
<geoCoordinate degrees="5.72" direction="south" orientation="latitude" precision="555" value="-5.72">5.72°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.53" direction="east" orientation="longitude" precision="555" value="144.53">144.53°E</geoCoordinate>
|
||
, elev. 1700 m, AMNH R-14783; Korn, Mt. Hagen,
|
||
<geoCoordinate degrees="5.84" direction="south" orientation="latitude" precision="555" value="-5.84">5.84°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="144.31" direction="east" orientation="longitude" precision="555" value="144.31">144.31°E</geoCoordinate>
|
||
, elev. 1630 m, AMNH R-14771, R-14773, R-14785-88.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="referred specimen">
|
||
<paragraph pageId="0" pageNumber="997">Referred specimen.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
Papua New Guinea: Morobe Prov-ince: S side Ekuti Divide,
|
||
<geoCoordinate degrees="7.42" direction="south" orientation="latitude" precision="555" value="-7.42">7.42°S</geoCoordinate>
|
||
,
|
||
<geoCoordinate degrees="146.43" direction="east" orientation="longitude" precision="555" value="146.43">146.43°E</geoCoordinate>
|
||
, elev. 1050 m, BPBM 17423.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="etymology">
|
||
<paragraph pageId="0" pageNumber="997">Etymology.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">The species epithet is a masculine Latin adjective meaning "dressed in black," in recognition of the dark dorsal and ventral colouration of adults of this species.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="diagnosis">
|
||
<paragraph pageId="0" pageNumber="997">Diagnosis.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
A large member of the
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
|
||
</taxonomicName>
|
||
Group (male SVL up to 655 mm, female SVL up to 682 mm) with the following unique combination of characters: sexual size dimorphism in SVL present (
|
||
<emphasis italics="true" pageId="0" pageNumber="997">t</emphasis>
|
||
71 = 2.5689,
|
||
<emphasis italics="true" pageId="0" pageNumber="997">p</emphasis>
|
||
= 0.0062); two scales covering vent; posterior genials usually entirely separated (80%) but may be in anterior contact (20%) with each other; intergenial usually widest posteriorly (89%) or centrally (11%); preocular usually contacting nasal (77%), not contacting internasal; preocular rather short, less than twice as long as deep; postoculars two (one in 26%); posterior temporals two (58%) or three (42%); ventrals sexually dimorphic (
|
||
<emphasis italics="true" pageId="0" pageNumber="997">t</emphasis>
|
||
85 = 7.400,
|
||
<emphasis italics="true" pageId="0" pageNumber="997">p</emphasis>
|
||
<0.00001), 177-206 in males, 187-218 in females; subcaudals sexually dimorphic with overlap (
|
||
<emphasis italics="true" pageId="0" pageNumber="997">t</emphasis>
|
||
87 = -24.8814,
|
||
<emphasis italics="true" pageId="0" pageNumber="997">p</emphasis>
|
||
<0.00001), 40-47 in males, 26-41 in females; two scales covering vent; yellow nuchal collar and yellow markings on prefrontals present in juveniles, usually absent (but sometimes merely faded) in adults; tail spine paler than remainder of tail; and venter uniformly dark brown or dark brown with the posterior of each ventral paler brown or yellowish brown in adults, giving a banded appearance, black in life; venter yellow with a black spot on lateral margins of each ventral in juveniles.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="comparisons">
|
||
<paragraph pageId="0" pageNumber="997">Comparisons with other species.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
Adult
|
||
<taxonomicName authorityName="Kraus & Kaiser & O’Shea" authorityYear="2022" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus atratus" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus atratus</emphasis>
|
||
</taxonomicName>
|
||
is easily distinguished from all other members of the
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
|
||
</taxonomicName>
|
||
species group except
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
in having a dark venter that is black or very dark brown in life and either uniformly brown or with each ventral banded dark brown in preservative. Other members of this complex have venters that are yellow (
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. lamingtoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="lamingtoni">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. lamingtoni</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loennbergii" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loennbergii">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. loennbergii</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. spilorhynchus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="spilorhynchus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. spilorhynchus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. vertebralis" order="Squamata" pageId="0" pageNumber="997" rank="species" species="vertebralis">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. vertebralis</emphasis>
|
||
</taxonomicName>
|
||
), grey (
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nigrescens" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nigrescens">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nigrescens</emphasis>
|
||
</taxonomicName>
|
||
), or yellow or pale grey with grey bands across each ventral (
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. mattisoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="mattisoni">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. mattisoni</emphasis>
|
||
</taxonomicName>
|
||
).
|
||
<taxonomicName authorityName="Kraus & Kaiser & O’Shea" authorityYear="2022" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus atratus" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus atratus</emphasis>
|
||
</taxonomicName>
|
||
can be distinguished from
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
by its juvenile ventral colour pattern (pale yellow with a black spot on the lateral edges of each ventral vs. uniformly black or dark brown but banded with dark brown in preservative in
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
), larger size (males to 655 mm SVL, females to 682 mm SVL vs. 422 mm and 540 mm, respectively, in
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
), and the greater number of ventrals with overlap in both sexes (males 177-206 vs. 178-198 in
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
;
|
||
<emphasis italics="true" pageId="0" pageNumber="997">t</emphasis>
|
||
65 = 5.0261,
|
||
<emphasis italics="true" pageId="0" pageNumber="997">p</emphasis>
|
||
<0.00001; females: 187-218 vs. 191-210 in
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<emphasis italics="true" pageId="0" pageNumber="997">t</emphasis>
|
||
62 = 4.3472,
|
||
<emphasis italics="true" pageId="0" pageNumber="997">p</emphasis>
|
||
= 0.000026). Although the venter of
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. atratus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. atratus</emphasis>
|
||
</taxonomicName>
|
||
is much darker, its banded pattern in most preserved specimens could perhaps be mistaken for the paler banded pattern seen in preserved
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nigrescens" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nigrescens">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nigrescens</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. mattisoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="mattisoni">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. mattisoni</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
<taxonomicName authorityName="Kraus & Kaiser & O’Shea" authorityYear="2022" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus atratus" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus atratus</emphasis>
|
||
</taxonomicName>
|
||
can be further distinguished from
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nigrescens" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nigrescens">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nigrescens</emphasis>
|
||
</taxonomicName>
|
||
in having a shorter preocular (preocular approximately squarish, slightly longer than tall in
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. atratus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. atratus</emphasis>
|
||
</taxonomicName>
|
||
vs. more than twice as long as tall in
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nigrescens" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nigrescens">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nigrescens</emphasis>
|
||
</taxonomicName>
|
||
), and from
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. mattisoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="mattisoni">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. mattisoni</emphasis>
|
||
</taxonomicName>
|
||
in having the preocular usually contacting nasal (vs. never in
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. mattisoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="mattisoni">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. mattisoni</emphasis>
|
||
</taxonomicName>
|
||
) and usually two postoculars (vs. one in
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. mattisoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="mattisoni">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. mattisoni</emphasis>
|
||
</taxonomicName>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="description">
|
||
<paragraph pageId="0" pageNumber="997">Description of the holotype.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
Adult female, 590 mm SVL + 72 mm TL = 662 mm TTL. Rostral broader than high, notched ventromedially; internasals angulate, semi-triangular, wider than long; prefrontals distinct from preoculars, approximately square but angled laterally and posteriorly, slightly wider than long (Fig.
|
||
<figureCitation captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Head of the Toxicocalamus atratus sp. nov. holotype (MCZ R- 84144) from Kundiawa, Chimbu Province, PNG, in both photographic and line-drawing illustrations. A, A ' dorsal, B, B ' right lateral, C, C ' left lateral, and D, D ' ventral views." figureDoi="10.3897/vz.72.e89647.figure14" httpUri="https://binary.pensoft.net/fig/768174" pageId="0" pageNumber="997">14A, A</figureCitation>
|
||
'), bordered below by preocular and nasal; preoculars angulate, slightly longer than high, bordered anteriorly by nasal, below by second and third supralabials (Fig.
|
||
<figureCitation captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Head of the Toxicocalamus atratus sp. nov. holotype (MCZ R- 84144) from Kundiawa, Chimbu Province, PNG, in both photographic and line-drawing illustrations. A, A ' dorsal, B, B ' right lateral, C, C ' left lateral, and D, D ' ventral views." figureDoi="10.3897/vz.72.e89647.figure14" httpUri="https://binary.pensoft.net/fig/768174" pageId="0" pageNumber="997">14B, B</figureCitation>
|
||
', C,
|
||
<normalizedToken originalValue="C’">C'</normalizedToken>
|
||
); frontal shield-shaped, lateral margins straight, not fused with supraoculars, anterior margin extending slightly anterior to remainder of scale medially; parietals approximately twice as long as wide. Nasals divided by large nares, with a short groove above and below along the posterior of naris. Postoculars single on each side, irregularly pentagonal in shape; one elongate anterior temporal above fifth and sixth supralabials, separating latter from parietal; two posterior temporals, subequal in size, lower abutting posterodorsal margin of sixth supralabial. Supralabials six on each side, third and fourth entering eye; infralabials six, first three in contact with anterior genial. Mental small, shallow, triangular, wider than deep, bordered behind by first supralabials; anterior genials slightly larger than posterior genials, in medial contact along entire length; posterior genials longer on left than right, completely separated by single elongate intergenial, which is widest posteriorly; seven gulars separate intergenial from first ventral in the midline; first sublabial separates posterior genial from fifth infralabial (Fig.
|
||
<figureCitation captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Head of the Toxicocalamus atratus sp. nov. holotype (MCZ R- 84144) from Kundiawa, Chimbu Province, PNG, in both photographic and line-drawing illustrations. A, A ' dorsal, B, B ' right lateral, C, C ' left lateral, and D, D ' ventral views." figureDoi="10.3897/vz.72.e89647.figure14" httpUri="https://binary.pensoft.net/fig/768174" pageId="0" pageNumber="997">14D, D</figureCitation>
|
||
'). Eye relatively small; pupil round.
|
||
</paragraph>
|
||
<caption doi="10.3897/vz.72.e89647.figure14" httpUri="https://binary.pensoft.net/fig/768174" pageId="0" pageNumber="997" start="Figure 14" startId="F14">
|
||
<paragraph pageId="0" pageNumber="997">
|
||
<emphasis bold="true" pageId="0" pageNumber="997">Figure 14.</emphasis>
|
||
Head of the
|
||
<taxonomicName authorityName="Kraus & Kaiser & O’Shea" authorityYear="2022" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus atratus" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus atratus</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" pageId="0" pageNumber="997">sp. nov.</emphasis>
|
||
holotype (MCZ R-84144) from Kundiawa, Chimbu Province, PNG, in both photographic and line-drawing illustrations.
|
||
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
|
||
,
|
||
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
|
||
' dorsal,
|
||
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
|
||
,
|
||
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
|
||
' right lateral,
|
||
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
|
||
,
|
||
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
|
||
' left lateral, and
|
||
<emphasis bold="true" pageId="0" pageNumber="997">D, D</emphasis>
|
||
' ventral views.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="0" pageNumber="997">Dorsal scale rows 15-15-15, smooth, not notched posteriorly, without apical pits. Ventrals 207, each approximately four times wider than long; two scales covering vent; subcaudals 30, paired. Tail tipped by a pointed coni-cal spine.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
In preservative (50 years after collection), dorsum dark brown dorsally, slightly paler laterally (Fig.
|
||
<figureCitation captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. A Dorsum and B venter of the Toxicocalamus atratus sp. nov. holotype (MCZ R- 84144) from Kundiawa, Chimbu Province, PNG." figureDoi="10.3897/vz.72.e89647.figure15" httpUri="https://binary.pensoft.net/fig/768181" pageId="0" pageNumber="997">15A</figureCitation>
|
||
). Venter dark brown, anterior ventrals dark brown, narrowly margined with paler brown posteriorly (Fig.
|
||
<figureCitation captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. A Dorsum and B venter of the Toxicocalamus atratus sp. nov. holotype (MCZ R- 84144) from Kundiawa, Chimbu Province, PNG." figureDoi="10.3897/vz.72.e89647.figure15" httpUri="https://binary.pensoft.net/fig/768181" pageId="0" pageNumber="997">15B</figureCitation>
|
||
), subcaudals dark brown. Supralabials yellow, dark brown on approximately dorsal half of each scale. Head dark brown with vague yellow blotch on each prefrontal just extending onto posterolateral corner of each internasal. No pale nuchal collar present. Chin and throat with pale straw yellow ground heavily suffused with brown throughout, imparting a largely brown impression overall. Tail spine white. Iris black.
|
||
</paragraph>
|
||
<caption doi="10.3897/vz.72.e89647.figure15" httpUri="https://binary.pensoft.net/fig/768181" pageId="0" pageNumber="997" start="Figure 15" startId="F15">
|
||
<paragraph pageId="0" pageNumber="997">
|
||
<emphasis bold="true" pageId="0" pageNumber="997">Figure 15.</emphasis>
|
||
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
|
||
Dorsum and
|
||
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
|
||
venter of the
|
||
<taxonomicName authorityName="Kraus & Kaiser & O’Shea" authorityYear="2022" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus atratus" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus atratus</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" pageId="0" pageNumber="997">sp. nov.</emphasis>
|
||
holotype (MCZ R-84144) from Kundiawa, Chimbu Province, PNG.
|
||
</paragraph>
|
||
</caption>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="variation">
|
||
<paragraph pageId="0" pageNumber="997">Variation.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">Preoculars contact nasals in most specimens but are separated by prefrontal contact with the second supralabial on both sides in 17 specimens and on one side in nine specimens; preoculars are invariably separated from contact with internasals. Postoculars one (25.5%) or two (74%), absent on left side of MCZ R-141849. Posterior temporals two (58%) or three (42%). Supralabials six, except five on right side of AMS R14788, CAS 118948, and MCZ R-84142, the left side of MCZ R-116790, and both sides of CAS 115986, 118962, and MCZ R-141849; third and fourth supralabials contacting eye, except third through fifth on right side of AMNH R-75344, and only third supralabial on right side of AMS R14788, CAS 118962, and MCZ R-84142. Infralabials invariably six. Posterior genials in small anterior contact (20% of specimens) or entirely separated (80%) by single intergenial (two small intergenials in AMNH R-72780-81 from Kup). Intergenials widest posteriorly (89% of specimens) but widest centrally in ten (11%).</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
Dorsal scale rows invariably 15-15-15. Ventrals 177-206 (196
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
5) in 49 males, 187-218 (206
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
7) in 43 females; subcaudals 40-47 (44
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
2) in 48 males, 26-41 (31
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
3) in 41 females. SCR 16.9-21.0% (18.3
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
0.8%) in 48 males, 11.2-17.8% (12.9
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
1.3%) in 41 females. Tail tipped by a blunt to pointed conical spine. Maximum male SVL = 655 mm, TLR in 38 adult males 12.9-19.4% (15.5
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
1.7%); maximum female SVL = 682 mm, TLR in 33 adult females 6.7-18.7% (10.2
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
2.2%); sexual size dimorphism present (
|
||
<emphasis italics="true" pageId="0" pageNumber="997">t</emphasis>
|
||
71 = 2.5689,
|
||
<emphasis italics="true" pageId="0" pageNumber="997">p</emphasis>
|
||
= 0.0062). There appears to be an ontogenetic effect of tail length in juvenile males, whose tails are relatively shorter (mean TLR 12.5
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
0.6;
|
||
<emphasis italics="true" pageId="0" pageNumber="997">n</emphasis>
|
||
= 10) than those of adult males (mean TLR 15.6
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
1.7;
|
||
<emphasis italics="true" pageId="0" pageNumber="997">n</emphasis>
|
||
= 39). This difference is not present in females (mean TLR in eight juveniles 10.4
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
1.4, in 32 adults 9.9
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
1.7). The vent is typically covered by two scales, but this covering is incompletely formed in AMNH R-75355 and covered by a single scale in CAS 103774-75 and 139584.
|
||
</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
Variation in colouration is largely ontogenetic, with adults being black or dark brown above and below, although these colours fade to brown in preservative, and often with each ventral appearing banded with dark and light brown as colouration fades in preservative. Small juveniles (SVL = 161-262 mm) are often paler than this, with a dark vertebral stripe often being present, but this seems to be lost as animals darken ontogenetically. Small juveniles also have a large yellow blotch on each prefrontal and an incomplete yellow nuchal collar whose sides are separated by two to four brown dorsal scale rows (Fig.
|
||
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Dorsal and right lateral views of heads of juveniles of Toxicocalamus loriae Group snakes. A, B T. loriae (BPBM 10966), Agaun, Milne Bay Province, PNG. C, D T. nymani comb. nov., spotted form (BPBM 5442), Kalolo, Morobe Province, PNG. E, F T. nymani, dark form (BPBM 23699), Wau, Morobe Province, PNG. G, H T. lamingtoni comb. nov. (AMS R 9352), Mt. Lamington, Oro Province, PNG. I, J T. vertebralis sp. nov. (KU 129086), Wau, Morobe Province, PNG. K, L T. spilorhynchus sp. nov. (AMNH R- 107204), Garaina, Morobe Province, PNG. M, N T. atratus sp. nov. (CAS 118958), Mintima, Chimbu Province, PNG." figureDoi="10.3897/vz.72.e89647.figure4" httpUri="https://binary.pensoft.net/fig/768170" pageId="0" pageNumber="997">4M, N</figureCitation>
|
||
,
|
||
<figureCitation captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Uncollected juvenile Toxicocalamus atratus sp. nov. from Kutubu area, 1760 m elevation, Southern Highlands Province, PNG, to show the colouration in life. Photographs by Nick Baker." figureDoi="10.3897/vz.72.e89647.figure16" httpUri="https://binary.pensoft.net/fig/768182" pageId="0" pageNumber="997">16</figureCitation>
|
||
). These markings usually disappear in adults but may be retained as vague markings. Furthermore, small juveniles have a yellow venter with each ventral marked on each lateral extremity with a black spot. Because the first row of dorsal scales is also yellow or pale brown, this imparts the impression of the venter having a row of black spots along each side. In a few small specimens (CAS 103374, 113666, MCZ R-116774), these yellow venters also have a few small brown flecks. The tail spine is white in most specimens but pale brown in two.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="colour in life">
|
||
<paragraph pageId="0" pageNumber="997">Colour in life.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
Photographs of MCZ R-84026 (540 mm SVL) and R-111767 (480 mm SVL), both adult males, show uniformly black animals, with the venter of the latter also being black (Fig.
|
||
<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Toxicocalamus loriae Group species in live and perimortem photographs to show colour in life. A, B T. vertebralis sp. nov. (CAS 140042) from Kol, Jimi Valley, Jiwaka Province, PNG. C, D T. spilorhynchus sp. nov. (MCZ R- 152431) from Garaina, Morobe Province, PNG. E T. atratus sp. nov. (MCZ R- 84026) from Kundiawa, Wahgi Valley, Chimbu Province, PNG. F T. atratus (MCZ R- 111767) from Kundiawa, Wahgi Valley, Chimbu Province, PNG. Photographs by Fred Parker." figureDoi="10.3897/vz.72.e89647.figure11" httpUri="https://binary.pensoft.net/fig/768178" pageId="0" pageNumber="997">11E, F</figureCitation>
|
||
). Photographs provided to us by Nick Baker of an uncollected juvenile from Southern Highlands Province, PNG, show an animal with a black dorsum (Fig.
|
||
<figureCitation captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Uncollected juvenile Toxicocalamus atratus sp. nov. from Kutubu area, 1760 m elevation, Southern Highlands Province, PNG, to show the colouration in life. Photographs by Nick Baker." figureDoi="10.3897/vz.72.e89647.figure16" httpUri="https://binary.pensoft.net/fig/768182" pageId="0" pageNumber="997">16A</figureCitation>
|
||
), yellow venter having a mid-ventral row of brown spots (Fig.
|
||
<figureCitation captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Uncollected juvenile Toxicocalamus atratus sp. nov. from Kutubu area, 1760 m elevation, Southern Highlands Province, PNG, to show the colouration in life. Photographs by Nick Baker." figureDoi="10.3897/vz.72.e89647.figure16" httpUri="https://binary.pensoft.net/fig/768182" pageId="0" pageNumber="997">16B, C</figureCitation>
|
||
), and yellow supralabials, a yellow spot on each prefrontal, and a very incomplete nuchal collar consisting of only two small yellow lateral spots (Fig.
|
||
<figureCitation captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Uncollected juvenile Toxicocalamus atratus sp. nov. from Kutubu area, 1760 m elevation, Southern Highlands Province, PNG, to show the colouration in life. Photographs by Nick Baker." figureDoi="10.3897/vz.72.e89647.figure16" httpUri="https://binary.pensoft.net/fig/768182" pageId="0" pageNumber="997">16C, D</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption doi="10.3897/vz.72.e89647.figure16" httpUri="https://binary.pensoft.net/fig/768182" pageId="0" pageNumber="997" start="Figure 16" startId="F16">
|
||
<paragraph pageId="0" pageNumber="997">
|
||
<emphasis bold="true" pageId="0" pageNumber="997">Figure 16.</emphasis>
|
||
Uncollected juvenile
|
||
<taxonomicName authorityName="Kraus & Kaiser & O’Shea" authorityYear="2022" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus atratus" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus atratus</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" pageId="0" pageNumber="997">sp. nov.</emphasis>
|
||
from Kutubu area, 1760 m elevation, Southern Highlands Province, PNG, to show the colouration in life. Photographs by Nick Baker.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="0" pageNumber="997">Fred Parker, the collector, took numerous field notes on this species in life. Colour patterns for adults were described by him as follows: "Black with pair of yellow spots on the snout, partial bars laterally on back of head." (MCZ R-121547, SVL = 360 mm); "Part grown, black above and below." (CAS 139564, SVL = 455 mm); "Olive-black above, a narrow darker vertebral line from behind head to tip of tail. Lips translucent pinkish marked grey. Ventrals greyish, paler than dorsum." (MCZ R-84142, SVL = 470 mm); "Uniform iridescent black on dorsal surfaces. Ventrally and laterally uniform black with no markings. Lips faintly yellowish. Spike on tail yellow with black tip." (MCZ R-84026, SVL = 540 mm). "Black above, brown laterally. Grey-brown underneath. Lips paler." (CAS 99916, SVL = 610 mm). "Black above and below. Head slightly paler with olive tinge. Highly iridescent." (AMNH R-98495, SVL = 515 mm).</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">He described the colour pattern for two juveniles as follows: "A juvenile with bright yellow bar across snout and short similar bars laterally on back of head from behind corner of jaw to posterior corners of large head shields but not meeting dorsally. Ventrally, translucent yellow, with a black spot at side of each ventral [scale]. Tail with yellow spike at tip, the spike with small brown tip. Whole of upper lips yellow. Some greyish markings under the head. A narrow dark vertebral stripe. Flanks paler than dorsum. Pair of anals marked grey." (MCZ R-83218, SVL = 211 mm); "Grey-brown above, whitish below, yellow bar on snout, lips yellow, part bars laterally at back of head." (AMNH R-98498, SVL = 299 mm). Finally, a neonate that hatched on 23 December 1964 was recorded as black above, translucent below, and with paler lips; we have been unable to locate this specimen in any museum catalogue.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="range">
|
||
<paragraph pageId="0" pageNumber="997">Range.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
Known from the Central Highlands of Papua New Guinea and extending as far east as the southern side of the Ekuti Dividing Range, Morobe Province, at elevations of 840-2140 m (Fig.
|
||
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Type localities (stars) and other collection sites (circles) of Toxicocalamus loriae Group species. A Map of New Guinea and its satellite islands, showing the positions of the inset maps and the type locality of T. mintoni (magenta), an endemic of Sudest Island, Milne Bay Province, PNG. Scale = 500 km. B Papua New Guinea and adjacent areas of West New Guinea, Indonesia, with white lines demarcating provincial boundaries. Scale = 250 km. C West Papua Province, Indonesia (WP), showing the type locality of T. loennbergii comb. nov. (orange). Other species treated in this report are T. loriae (dark blue), T. lamingtoni comb. nov. (yellow), T. spilorhynchus sp. nov. (red), T. nymani comb. nov. (neon green), T. vertebralis sp. nov. (pink), and T. atratus sp. nov. (light blue). Additional related species in the T. loriae clade are T. goodenoughensis (brown), T. mattisoni (violet), T. nigrescens (black), T. pachysomus (dark green), and T. " Toxicocalamus loriae " Clade 4 (white). The type locality of the recently described dubious taxon T. longhagen (tan) is also indicated. Scale = 250 km. Provinces of PNG are Central (Ce), Chimbu (Ch; also spelled Simbu), Gulf (Gu), Hela (He; created in 2012), Eastern Highlands (EH), East Sepik (ES), Enga (En), Jiwaka (Ji; created in 2012), Madang (Ma), Milne Bay (MB), Morobe (Mo), National Capital District (NCD), Oro (Or; also known as Northern Province); Sandaun (Sa; formerly West Sepik), Southern Highlands (SH), Western (We), and Western Highlands (WH). Provinces of West New Guinea are Highland Papua (HP; created in 2022), Papua (Pa), and South Papua (SP; created in 2022). Sites of sympatry are Wau (W; neon green + pink) with two species and Garaina (G; neon green + red + yellow) with three species." figureDoi="10.3897/vz.72.e89647.figure6" httpUri="https://binary.pensoft.net/fig/768172" pageId="0" pageNumber="997">6B</figureCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="ecological notes">
|
||
<paragraph pageId="0" pageNumber="997">Ecological notes.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
Considered common at Kundiawa, Chimbu Province, PNG (Fred Parker, pers. comm.). For example, 88 animals were obtained in gardens on 3 December 1967. In general, most individuals of this species were collected during the day from under piles of vegetation in sweet potato gardens. In forested situations, only the occasional animal would be collected in or under logs during the day. Parker never observed the species active at night, which is consistent with the first
|
||
<normalizedToken originalValue="author’s">author's</normalizedToken>
|
||
experience with other
|
||
<taxonomicName authorityName="Boulenger" authorityYear="1896" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus</emphasis>
|
||
</taxonomicName>
|
||
species. One specimen (MCZ R-111769, SVL = 437 mm) contained a large earthworm in its stomach.
|
||
</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
<normalizedToken originalValue="Parker’s">Parker's</normalizedToken>
|
||
field notes also recorded that MCZ R-84144 (SVL = 590 mm) laid two eggs on 25 May 1964 and one on the following day; one of these hatched on 23 December, indicating a 7-month incubation period. MCZ R-111790 (SVL = 440 mm) was captured on 3 December 1967 and "laid four eggs after capture," CAS 115987 (SVL = 375 mm) was captured on 29 February 1969 when gravid with three eggs, CAS 115992 (SVL = 435 mm) and CAS 115996 (SVL = 625 mm) were captured on 11 March 1968 and noted to be gravid, and CAS 115997 (SVL = 480 mm) was captured on 19 April 1968 gravid with six eggs. All of these snakes were from Kundiawa, Chimbu Province. Thus, gravid females were captured during the months of February to April and eggs were laid in May and December, suggesting that reproduction in this species occurs year-round. The observed eggs were cylindrical, bluntly rounded at each end, and measured 35
|
||
<normalizedToken originalValue="×">x</normalizedToken>
|
||
12 mm, 35
|
||
<normalizedToken originalValue="×">x</normalizedToken>
|
||
12.5 mm, and 39
|
||
<normalizedToken originalValue="×">x</normalizedToken>
|
||
14 mm.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="997" type="remarks">
|
||
<paragraph pageId="0" pageNumber="997">Remarks.</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
The primary feature distinguishing
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. atratus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. atratus</emphasis>
|
||
</taxonomicName>
|
||
from the very similar
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
is the ontogenetic change in ventral colour pattern, which is dramatic in
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. atratus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. atratus</emphasis>
|
||
</taxonomicName>
|
||
but absent in
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
. Some populations of
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
also retain the yellow nuchal collar and prefrontal spots into adulthood, which individuals of
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. atratus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. atratus</emphasis>
|
||
</taxonomicName>
|
||
never do. Furthermore,
|
||
<bibRefCitation DOI="https://doi.org/10.1080/00222930008678427" author="Loennberg, E" journalOrPublisher="Biodiversity and Conservation" pageId="0" pageNumber="997" refId="B25" refString="Loennberg, E, 1900. Reptiles and batrachians collected in German New Guinea by the late Dr Erik Nyman. Annals and Magazine of Natural History, Series 7 6: 574-582. https://doi.org/10.1080/-00-222930008678427" title="Reptiles and batrachians collected in German New Guinea by the late Dr Erik Nyman. Annals and Magazine of Natural History, Series 7 6: 574 - 582." url="https://doi.org/10.1080/00222930008678427" year="1900">
|
||
<normalizedToken originalValue="Lönnberg">Loennberg</normalizedToken>
|
||
(1900
|
||
</bibRefCitation>
|
||
: 579) described the dorsal colour of his freshly preserved specimens of
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
as "bronzy brown (almost blackish in the largest specimen)", whereas notes by Fred Parker recorded that a neonate and small juvenile of
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. atratus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. atratus</emphasis>
|
||
</taxonomicName>
|
||
were black above (see also Fig.
|
||
<figureCitation captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Uncollected juvenile Toxicocalamus atratus sp. nov. from Kutubu area, 1760 m elevation, Southern Highlands Province, PNG, to show the colouration in life. Photographs by Nick Baker." figureDoi="10.3897/vz.72.e89647.figure16" httpUri="https://binary.pensoft.net/fig/768182" pageId="0" pageNumber="997">16</figureCitation>
|
||
). As noted in the Comparisons section above,
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. atratus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. atratus</emphasis>
|
||
</taxonomicName>
|
||
also differs from
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
by its larger maximum size and, statistically, by its greater mean count of ventral scales in both males and females. These differences are confirmed by post-hoc Tukey Tests in an ANOVA (
|
||
<emphasis italics="true" pageId="0" pageNumber="997">p</emphasis>
|
||
= 0.00551 and
|
||
<emphasis italics="true" pageId="0" pageNumber="997">p</emphasis>
|
||
= 0.04195 for males and females, respectively). Its lesser mean SCR in males also differs significantly from that seen in
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
. Each of these differences suggests that
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. atratus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. atratus</emphasis>
|
||
</taxonomicName>
|
||
are separate species, but scalational differences that would be useful to field observation are not obvious at this point. It will be useful to investigate this issue in greater detail once fresh specimens with tissues and colour notes become available for both species.
|
||
</paragraph>
|
||
<paragraph pageId="0" pageNumber="997">
|
||
We refer the sole specimen from south of the Ekuti Divide (BPBM 17423) to this species on the basis of its ventral colour pattern, which indicates the lateral rows of brown spots on each ventral. However, the venter of this specimen is somewhat discoloured, and its collection locality is close to Wau, where
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
|
||
</taxonomicName>
|
||
primarily resides, so confirmation of this assignment to
|
||
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. atratus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="atratus">
|
||
<emphasis italics="true" pageId="0" pageNumber="997">T. atratus</emphasis>
|
||
</taxonomicName>
|
||
would be desirable once further specimens become available. This uncertainty leads us to exclude this specimen from the series of paratypes.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</subSection>
|
||
</document> |