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<mods:title id="D50005A4AD8C1677C41AC392A1DD6F60">Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key</mods:title>
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<paragraph id="8BD636BAFF87FF9FAB84FA89FD78CD51" blockId="5.[151,654,1303,1329]" box="[151,654,1303,1329]" pageId="5" pageNumber="6">
<heading id="D09E81D6FF87FF9FAB84FA89FD78CD51" bold="true" box="[151,654,1303,1329]" fontSize="11" level="1" pageId="5" pageNumber="6" reason="1">
<taxonomicName id="4C694D39FF87FF9FAB84FA89FD78CD51" ID-CoL="Y45N" authority="Boulenger, 1898" authorityName="Boulenger" authorityYear="1898" box="[151,654,1303,1329]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF87FF9FAB84FA89FD78CD51" bold="true" box="[151,654,1303,1329]" pageId="5" pageNumber="6">
<emphasis id="B91DEAA8FF87FF9FAB84FA89FE5BCD51" bold="true" box="[151,429,1303,1329]" italics="true" pageId="5" pageNumber="6">Cophixalus verrucosus</emphasis>
(Boulenger, 1898)
</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph id="8BD636BAFF87FF9FAB84FAC0FDD2CEBE" blockId="5.[151,1436,1374,2011]" pageId="5" pageNumber="6">
Boulenger (1898) also described
<taxonomicName id="4C694D39FF87FF9FA90EFAC0FCD5CD17" box="[541,803,1374,1399]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF87FF9FA90EFAC0FCD5CD17" box="[541,803,1374,1399]" italics="true" pageId="5" pageNumber="6">Cophixalus verrucosus</emphasis>
</taxonomicName>
on the basis of two specimens from Moroka and six specimens from Mt Victoria. Both localities are within approximately
<quantity id="4C919B5FFF87FF9FA8D4FA1CFBE7CDFA" box="[967,1041,1410,1434]" metricMagnitude="4" metricUnit="m" metricValue="7.0" pageId="5" pageNumber="6" unit="km" value="70.0">70 km</quantity>
of each other and outside of Port Moresby in the central Owen Stanley Range, Central Province,
<collectingCountry id="F37E762AFF87FF9FA880FA39FB8DCDDE" box="[915,1147,1446,1470]" name="Papua New Guinea" pageId="5" pageNumber="6">Papua New Guinea</collectingCountry>
. The diagnostic features mentioned by Boulenger are a sharp canthus, vertical lores, projecting snout, head as long as broad, large finger discs, tuberculate skin, and hidden surfaces of thighs and groin with a pattern of large white spots on a black ground.
<bibRefCitation id="EFF84B4BFF87FF9FABE6F98CFE5CCE4A" author="Zweifel" box="[245,426,1554,1578]" pageId="5" pageNumber="6" refString="Zweifel, R. G. (1956 a) Notes on microhylid frogs, genus Cophixalus, from New Guinea. American Museum Novitates, 1785, 1 - 8." type="journal article" year="1956" yearSuffix="a">Zweifel (1956a)</bibRefCitation>
emphasized the color pattern on the thighs as uniquely diagnostic within
<taxonomicName id="4C694D39FF87FF9FAFF6F98DFA92CE4B" box="[1253,1380,1555,1579]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B91DEAA8FF87FF9FAFF6F98DFA92CE4B" box="[1253,1380,1555,1579]" italics="true" pageId="5" pageNumber="6">Cophixalus</emphasis>
</taxonomicName>
, and
<bibRefCitation id="EFF84B4BFF87FF9FAB84F9A8FEB7CE2E" author="Zweifel" box="[151,321,1590,1614]" pageId="5" pageNumber="6" refString="Zweifel, R. G. (1979) A new cryptic species of microhylid frog (genus Cophixalus) from Papua New Guinea, with notes on related forms. American Museum Novitates, 2678, 1 - 14." type="journal article" year="1979">Zweifel (1979)</bibRefCitation>
and
<bibRefCitation id="EFF84B4BFF87FF9FAA68F9A8FDDBCE2E" author="Menzies" box="[379,557,1590,1614]" pageId="5" pageNumber="6" refString="Menzies, J. I. (2006) The frogs of New Guinea and the Solomon Islands. Pensoft, Sofia-Moscow." type="book" year="2006">Menzies (2006)</bibRefCitation>
added the detail that these colors are yellow on a dark-brown ground in life.
<bibRefCitation id="EFF84B4BFF87FF9FAB84F9C4FEBECE12" author="Menzies" box="[151,328,1626,1650]" pageId="5" pageNumber="6" refString="Menzies, J. I. (2006) The frogs of New Guinea and the Solomon Islands. Pensoft, Sofia-Moscow." type="book" year="2006">Menzies (2006)</bibRefCitation>
further noted that the venter is gray anteriorly and bright lemon yellow posteriorly and on the hind limbs. In my experience, this too appears unique among the known species of the genus. This species ranges from sea level to
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elevation throughout most of the Papuan Peninsula and on several of the offshore islands of Milne Bay Province (unpubl. data).
</paragraph>
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<bibRefCitation id="EFF84B4BFF87FF9FABD4F974FEACCF62" author="Hiaso" box="[199,346,1770,1794]" pageId="5" pageNumber="6" refString="Hiaso, J. (2002) A new species of Cophixalus (Anura: Microhylidae) from Tagula Island, New Guinea. Science in New Guinea, 27, 96 - 100." type="journal article" year="2002">Hiaso (2002)</bibRefCitation>
described
<taxonomicName id="4C694D39FF87FF9FAAC0F975FD3FCF63" box="[467,713,1771,1795]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="aimbensis">
<emphasis id="B91DEAA8FF87FF9FAAC0F975FD3FCF63" box="[467,713,1771,1795]" italics="true" pageId="5" pageNumber="6">Cophixalus aimbensis</emphasis>
</taxonomicName>
on the basis of
<specimenCount id="9D6FFD33FF87FF9FA869F974FBE7CF62" box="[890,1041,1770,1794]" pageId="5" pageNumber="6" type="generic">11 specimens</specimenCount>
(one cleared and stained) collected from along the Aimba River, Sudest Island, Milne Bay Province,
<collectingCountry id="F37E762AFF87FF9FA8AAF891FB55CF46" box="[953,1187,1806,1830]" name="Papua New Guinea" pageId="5" pageNumber="6">Papua New Guinea</collectingCountry>
. He claimed that all specimens were male, but in fact three of them (UPNG 8464, 8466, 8469) are female. Hiaso diagnosed his new species from its congeners with the characters: (1) head as long as broad, (2) snout rounded dorsally and projecting in profile, (3) fingers and toes with small terminal discs with marginal grooves, and (4) width of finger discs slightly larger than width of toe discs. The first two features were both mentioned by Boulenger (1898) in his description of
<taxonomicName id="4C694D39FF87FF9FAA2BF85DFE25CFBB" box="[312,467,1987,2011]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF87FF9FAA2BF85DFE25CFBB" box="[312,467,1987,2011]" italics="true" pageId="5" pageNumber="6">C. verrucosus</emphasis>
</taxonomicName>
. Furthermore, the first does not serve to distinguish
<taxonomicName id="4C694D39FF87FF9FAF0FF85DFB5ACFBB" box="[1052,1196,1987,2011]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="aimbensis">
<emphasis id="B91DEAA8FF87FF9FAF0FF85DFB5ACFBB" box="[1052,1196,1987,2011]" italics="true" pageId="5" pageNumber="6">C. aimbensis</emphasis>
</taxonomicName>
from most species of
<taxonomicName id="4C694D39FF84FF9CAB84FF06FEE1C8D0" box="[151,279,152,176]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B91DEAA8FF84FF9CAB84FF06FEE1C8D0" box="[151,279,152,176]" italics="true" pageId="6" pageNumber="7">Cophixalus</emphasis>
</taxonomicName>
. My remeasurements of the nine remaining
<typeStatus id="54D28818FF84FF9CA814FF06FCC1C8D0" box="[775,823,152,176]" pageId="6" pageNumber="7">type</typeStatus>
specimens of
<taxonomicName id="4C694D39FF84FF9CA8CBFF06FB9FC8D0" box="[984,1129,152,176]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="aimbensis">
<emphasis id="B91DEAA8FF84FF9CA8CBFF06FB9FC8D0" box="[984,1129,152,176]" italics="true" pageId="6" pageNumber="7">C. aimbensis</emphasis>
</taxonomicName>
(UPNG 8468 and 8470 are missing and presumably destroyed, cf.
<bibRefCitation id="EFF84B4BFF84FF9CA979FF22FC85C8B4" author="Shea" box="[618,883,188,212]" pageId="6" pageNumber="7" refString="Shea, G., &amp; Kraus, F. (2007) A list of herpetological type specimens in the collections of the National Museum of Papua New Guinea and the University of Papua New Guinea. Zootaxa, 1514, 37 - 60." type="journal article" year="2007">Shea and Kraus, 2007</bibRefCitation>
) show that HL/HW varies in that series from 0.880.99, following the mode shown by most members of the genus, which tend to have the head slightly shorter than wide. These figures differ from those provided by Hiaso (HL/HW = 0.991.14), but he didnt define the landmarks used for his measurements, so the source of this difference is uncertain. The second feature is distinctive within Papuan
<taxonomicName id="4C694D39FF84FF9CAA53FED2FE49C904" box="[320,447,332,356]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B91DEAA8FF84FF9CAA53FED2FE49C904" box="[320,447,332,356]" italics="true" pageId="6" pageNumber="7">Cophixalus</emphasis>
</taxonomicName>
, but it is also shared with
<taxonomicName id="4C694D39FF84FF9CA9FAFED3FBDFC904" authority="Parker, 1934" authorityName="Parker" authorityYear="1934" box="[745,1065,332,356]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="cheesmanae">
<emphasis id="B91DEAA8FF84FF9CA9FAFED3FC62C904" box="[745,916,332,356]" italics="true" pageId="6" pageNumber="7">C. cheesmanae</emphasis>
<bibRefCitation id="EFF84B4BFF84FF9CA888FED2FBDFC904" author="Parker" box="[923,1065,332,356]" pageId="6" pageNumber="7" refString="Parker, H. W. (1934) A Monograph of the Frogs of the Family Microhylidae. British Museum (Natural History), London, U. K." type="book" year="1934">Parker, 1934</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C694D39FF84FF9CAF72FED3FB0BC904" box="[1121,1277,333,356]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CAF72FED3FB0BC904" box="[1121,1277,333,356]" italics="true" pageId="6" pageNumber="7">C. verrucosus</emphasis>
</taxonomicName>
. Hiasos third diagnostic feature characterizes almost all members of
<taxonomicName id="4C694D39FF84FF9CA812FEEEFC76C9E8" box="[769,896,368,392]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B91DEAA8FF84FF9CA812FEEEFC76C9E8" box="[769,896,368,392]" italics="true" pageId="6" pageNumber="7">Cophixalus</emphasis>
</taxonomicName>
and is, hence, uninformative. His fourth feature characterizes almost half the Papuan species of
<taxonomicName id="4C694D39FF84FF9CA9A1FE0AFCC7C9CC" box="[690,817,404,428]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B91DEAA8FF84FF9CA9A1FE0AFCC7C9CC" box="[690,817,404,428]" italics="true" pageId="6" pageNumber="7">Cophixalus</emphasis>
</taxonomicName>
. Indeed,
<bibRefCitation id="EFF84B4BFF84FF9CA888FE0AFB49C9CC" author="Menzies" box="[923,1215,404,428]" pageId="6" pageNumber="7" refString="Menzies, J. I. &amp; Tyler, M. J. (1977) The systematics and adaptations of some Papuan microhylid frogs which live underground. Journal of Zoology, London, 183, 431 - 464." type="journal article" year="1977">Menzies and Tyler (1977)</bibRefCitation>
thought the greater breadth of the finger discs relative to the toe discs was a diagnostic feature of
<taxonomicName id="4C694D39FF84FF9CA8E7FE26FB85C9B0" box="[1012,1139,440,464]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B91DEAA8FF84FF9CA8E7FE26FB85C9B0" box="[1012,1139,440,464]" italics="true" pageId="6" pageNumber="7">Cophixalus</emphasis>
</taxonomicName>
vis-à-vis
<taxonomicName id="4C694D39FF84FF9CAFF2FE26FF27C994" authority="Mehely, 1901" authorityName="Mehely" authorityYear="1901" class="Amphibia" family="Microhylidae" genus="Copiula" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B91DEAA8FF84FF9CAFF2FE26FACDC9B0" box="[1249,1339,440,464]" italics="true" pageId="6" pageNumber="7">Copiula</emphasis>
Méhely, 1901
</taxonomicName>
, but there is more variation in that feature within
<taxonomicName id="4C694D39FF84FF9CA9EFFE42FC8AC994" box="[764,892,476,500]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B91DEAA8FF84FF9CA9EFFE42FC8AC994" box="[764,892,476,500]" italics="true" pageId="6" pageNumber="7">Cophixalus</emphasis>
</taxonomicName>
(as currently constituted) than they thought, and a large number of species do not fit this description (
<bibRefCitation id="EFF84B4BFF84FF9CA9F0FE61FC0ACA77" author="Kraus" box="[739,1020,511,535]" pageId="6" pageNumber="7" refString="Kraus, F. &amp; Allison, A. (2009) New species of Cophixalus (Anura: Microhylidae) from Papua New Guinea. Zootaxa, 2128, 1 - 38." type="journal article" year="2009">Kraus and Allison, 2009</bibRefCitation>
).
</paragraph>
<paragraph id="8BD636BAFF84FF9CABD4FDBAFD0BCA90" blockId="6.[151,1436,151,1545]" pageId="6" pageNumber="7">
Although not used in his diagnosis,
<bibRefCitation id="EFF84B4BFF84FF9CA949FDBAFD18CA5C" author="Hiaso" box="[602,750,548,572]" pageId="6" pageNumber="7" refString="Hiaso, J. (2002) A new species of Cophixalus (Anura: Microhylidae) from Tagula Island, New Guinea. Science in New Guinea, 27, 96 - 100." type="journal article" year="2002">Hiaso (2002)</bibRefCitation>
also mentioned that his specimens were bright yellow on the posterior of the abdomen and on the hidden surfaces of the limbs, a feature noted above to be unique to
<taxonomicName id="4C694D39FF84FF9CAE91FDD6FEE4CAE4" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CAE91FDD6FEE4CAE4" italics="true" pageId="6" pageNumber="7">C. verrucosus</emphasis>
</taxonomicName>
. In his description, Hiaso (2006) did not explicitly compare his new species to any other member of the genus
<taxonomicName id="4C694D39FF84FF9CABF1FD0EFE97CAC8" box="[226,353,656,680]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B91DEAA8FF84FF9CABF1FD0EFE97CAC8" box="[226,353,656,680]" italics="true" pageId="6" pageNumber="7">Cophixalus</emphasis>
</taxonomicName>
, but, instead, concentrated on distinguishing it from members of the genus
<taxonomicName id="4C694D39FF84FF9CAFC0FD0EFADBCAC8" box="[1235,1325,656,680]" class="Amphibia" family="Microhylidae" genus="Copiula" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B91DEAA8FF84FF9CAFC0FD0EFADBCAC8" box="[1235,1325,656,680]" italics="true" pageId="6" pageNumber="7">Copiula</emphasis>
</taxonomicName>
, which it resembles only in having a projecting, though differently shaped, snout. From the original description, then,
<taxonomicName id="4C694D39FF84FF9CAE91FD2BFEF1CA90" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="aimbensis">
<emphasis id="B91DEAA8FF84FF9CAE91FD2BFEF1CA90" italics="true" pageId="6" pageNumber="7">C. aimbensis</emphasis>
</taxonomicName>
cannot be distinguished from
<taxonomicName id="4C694D39FF84FF9CA948FD46FD00CA90" box="[603,758,728,752]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CA948FD46FD00CA90" box="[603,758,728,752]" italics="true" pageId="6" pageNumber="7">C. verrucosus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BD636BAFF84FF9CABD4FD63FDE5CE68" blockId="6.[151,1436,151,1545]" pageId="6" pageNumber="7">
In
<date id="FFD7107AFF84FF9CABF6FD62FED8CB74" box="[229,302,764,788]" pageId="6" pageNumber="7" value="2004-01">2004 I</date>
obtained a large series of
<taxonomicName id="4C694D39FF84FF9CA945FD62FCAECB74" box="[598,856,764,788]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CA945FD62FCAECB74" box="[598,856,764,788]" italics="true" pageId="6" pageNumber="7">Cophixalus verrucosus</emphasis>
</taxonomicName>
on Sudest Island, the
<typeStatus id="54D28818FF84FF9CAF41FD63FB74CB75" box="[1106,1154,765,789]" pageId="6" pageNumber="7">type</typeStatus>
locality of
<taxonomicName id="4C694D39FF84FF9CAE17FD63FA63CB74" box="[1284,1429,764,788]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="aimbensis">
<emphasis id="B91DEAA8FF84FF9CAE17FD63FA63CB74" box="[1284,1429,764,788]" italics="true" pageId="6" pageNumber="7">C. aimbensis</emphasis>
</taxonomicName>
. There is nothing to distinguish these specimens from the
<typeStatus id="54D28818FF84FF9CA80DFCBEFCB8CB58" box="[798,846,800,824]" pageId="6" pageNumber="7">type</typeStatus>
series of
<taxonomicName id="4C694D39FF84FF9CA8AEFCBEFBB9CB58" box="[957,1103,800,824]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="aimbensis">
<emphasis id="B91DEAA8FF84FF9CA8AEFCBEFBB9CB58" box="[957,1103,800,824]" italics="true" pageId="6" pageNumber="7">C. aimbensis</emphasis>
</taxonomicName>
: they show complete overlap in bivariate morphometric ratios with the
<typeStatus id="54D28818FF84FF9CA978FCDBFD6DCB3D" box="[619,667,837,861]" pageId="6" pageNumber="7">type</typeStatus>
series of
<taxonomicName id="4C694D39FF84FF9CA81AFCDBFC6CCB3C" box="[777,922,836,860]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="aimbensis">
<emphasis id="B91DEAA8FF84FF9CA81AFCDBFC6CCB3C" box="[777,922,836,860]" italics="true" pageId="6" pageNumber="7">C. aimbensis</emphasis>
</taxonomicName>
and they match those specimens in all details of morphology and color pattern. They also closely match specimens of
<taxonomicName id="4C694D39FF84FF9CA8E7FCF6FB62CBE0" box="[1012,1172,872,896]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CA8E7FCF6FB62CBE0" box="[1012,1172,872,896]" italics="true" pageId="6" pageNumber="7">C. verrucosus</emphasis>
</taxonomicName>
I have collected from several localities throughout the Papuan Peninsula in everything but their larger body size (Sudest male SV = 24.531.0 mm vs.
<quantity id="4C919B5FFF84FF9CAA74FC31FDFFCBA8" box="[359,521,943,968]" metricMagnitude="-2" metricUnit="m" metricValue="2.35" metricValueMax="2.75" metricValueMin="1.95" pageId="6" pageNumber="7" unit="mm" value="23.5" valueMax="27.5" valueMin="19.5">19.527.5 mm</quantity>
elsewhere; Sudest female SV = 29.637.0 mm vs. 23.031.7 elsewhere,
<tableCitation id="C6EB0301FF84FF9CAE2AFC31FA78CBA7" box="[1337,1422,943,967]" captionStart="TABLE 2" captionStartId="6.[151,239,1591,1615]" captionText="TABLE 2. Body-size variation among populations of Cophixalus verrucosus, with populations listed from north to south." pageId="6" pageNumber="7">Table 2</tableCitation>
). In particular, they share the projecting snout, vertical lores, hidden surfaces of thighs and groin dark brown with large yellow spots, grey chin and throat, and lemon-yellow abdomen and undersurfaces of legs that are characteristic of
<taxonomicName id="4C694D39FF84FF9CAA43FB83FE1DCC54" box="[336,491,1053,1076]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CAA43FB83FE1DCC54" box="[336,491,1053,1076]" italics="true" pageId="6" pageNumber="7">C. verrucosus</emphasis>
</taxonomicName>
. That all of these specimens represent
<taxonomicName id="4C694D39FF84FF9CA888FB83FBC1CC54" box="[923,1079,1053,1076]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CA888FB83FBC1CC54" box="[923,1079,1053,1076]" italics="true" pageId="6" pageNumber="7">C. verrucosus</emphasis>
</taxonomicName>
is further indicated by principal components analysis of the Sudest specimens compared to
<taxonomicName id="4C694D39FF84FF9CA823FBDEFC3ACC38" box="[816,972,1088,1112]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CA823FBDEFC3ACC38" box="[816,972,1088,1112]" italics="true" pageId="6" pageNumber="7">C. verrucosus</emphasis>
</taxonomicName>
from several localities elsewhere in their range. In both bivariate (
<figureCitation id="13522A3FFF84FF9CAABAFBFAFE1DCC1C" box="[425,491,1124,1148]" captionStart="FIGURE 3" captionStartId="7.[151,250,1800,1823]" captionTargetBox="[218,1349,243,1764]" captionTargetId="figure@7.[218,1369,233,1779]" captionTargetPageId="7" captionText="FIGURE 3. Bivariate plot of (A) principal component 1 (eigenvalue = 0.168) versus principal component 2 (eigenvalue = 0.012), and (B) principal component 2 versus principal component 3 (eigenvalue = 0.008) for eight type specimens of Cophixalus aimbensis (closed circles), ten other specimens of C. verrucosus from Sudest Island (open circles), ten specimens of C. verrucosus from Mt. Victoria and Moroka, the localities from which the type series was obtained (diamonds), and 27 other specimens of C. verrucosus from Milne Bay Province in southeastern Papua New Guinea (squares). Component loadings presented in Table 2." httpUri="https://zenodo.org/record/282922/files/figure.png" pageId="6" pageNumber="7">Fig. 3</figureCitation>
) and 3-dimensional (not shown) plots, the Sudest specimens cannot be distinguished from
<taxonomicName id="4C694D39FF84FF9CABC6FB16FE87CCC0" box="[213,369,1160,1184]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CABC6FB16FE87CCC0" box="[213,369,1160,1184]" italics="true" pageId="6" pageNumber="7">C. verrucosus</emphasis>
</taxonomicName>
, although they lie to the extreme of that species along PC1. However, this axis largely reflects specimen size, judging by the uniformly positive loadings of the constituent variables (
<tableCitation id="C6EB0301FF84FF9CAF68FB32FB25CCA4" box="[1147,1235,1196,1220]" captionStart="TABLE 3" captionStartId="8.[151,239,151,175]" captionTargetBox="[165,1351,273,910]" captionTargetPageId="8" captionText="TABLE 3. Loadings for log-transformed characters used in principal components analysis of covariance matrix for adults of Cophixalus aimbensis (type series) and C. verrucosus from populations throughout its range. Included specimens are listed in the Appendix." httpUri="http://table.plazi.org/id/DF166632FF8AFF92AB84FF09FDDEC88F" pageId="6" pageNumber="7" tableUuid="DF166632FF8AFF92AB84FF09FDDEC88F">Table 3</tableCitation>
), and this merely confirms that Sudest animals are on average larger than those from other populations of
<taxonomicName id="4C694D39FF84FF9CAF93FB4EFAEBCC88" box="[1152,1309,1232,1256]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CAF93FB4EFAEBCC88" box="[1152,1309,1232,1256]" italics="true" pageId="6" pageNumber="7">C. verrucosus</emphasis>
</taxonomicName>
. As can be seen from the loadings and from examination of scatter plots, there is no information expressed in PC2 or PC3 that serves to distinguish among populations (
<figureCitation id="13522A3FFF84FF9CA9B9FA86FD0CCD50" box="[682,762,1304,1328]" captionStart="FIGURE 3" captionStartId="7.[151,250,1800,1823]" captionTargetBox="[218,1349,243,1764]" captionTargetId="figure@7.[218,1369,233,1779]" captionTargetPageId="7" captionText="FIGURE 3. Bivariate plot of (A) principal component 1 (eigenvalue = 0.168) versus principal component 2 (eigenvalue = 0.012), and (B) principal component 2 versus principal component 3 (eigenvalue = 0.008) for eight type specimens of Cophixalus aimbensis (closed circles), ten other specimens of C. verrucosus from Sudest Island (open circles), ten specimens of C. verrucosus from Mt. Victoria and Moroka, the localities from which the type series was obtained (diamonds), and 27 other specimens of C. verrucosus from Milne Bay Province in southeastern Papua New Guinea (squares). Component loadings presented in Table 2." httpUri="https://zenodo.org/record/282922/files/figure.png" pageId="6" pageNumber="7">Fig. 3</figureCitation>
B). Smaller-sized specimens from Sudest overlap the morphological distribution of specimens from Moroka and approach those from Mt. Victoria, the
<typeStatus id="54D28818FF84FF9CAFCAFAA3FAFFCD35" box="[1241,1289,1341,1365]" pageId="6" pageNumber="7">type</typeStatus>
localities for
<taxonomicName id="4C694D39FF84FF9CAB84FAFEFECCCD18" box="[151,314,1376,1400]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CAB84FAFEFECCCD18" box="[151,314,1376,1400]" italics="true" pageId="6" pageNumber="7">C. verrucosus</emphasis>
</taxonomicName>
. More importantly, there is no obvious break in morphotype between mainland and Sudest populations. This holds as well if the mainland localities are each distinguished separately on the plot (graph not shown), in which case the Sudest specimens adjoin and overlap those from the Cloudy Mts., the geographically most proximate population on the mainland. That population, in turn, overlaps populations from elsewhere in Milne Bay and Central provinces.
</paragraph>
<caption id="DF166632FF84FF9CAB84F9A9FF2ECE0F" pageId="6" pageNumber="7">
<paragraph id="8BD636BAFF84FF9CAB84F9A9FF2ECE0F" blockId="6.[151,1436,1591,1647]" pageId="6" pageNumber="7">
<emphasis id="B91DEAA8FF84FF9CAB84F9A9FEF8CE2F" bold="true" box="[151,270,1591,1615]" pageId="6" pageNumber="7">TABLE 2.</emphasis>
Body-size variation among populations of
<taxonomicName id="4C694D39FF84FF9CA9E6F9A6FC18CE2F" box="[757,1006,1592,1615]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF84FF9CA9E6F9A6FC18CE2F" box="[757,1006,1592,1615]" italics="true" pageId="6" pageNumber="7">Cophixalus verrucosus</emphasis>
</taxonomicName>
, with populations listed from north to south.
</paragraph>
</caption>
<paragraph id="8BD636BAFF84FF9CABFBF90FFB33CEC6" blockId="6.[165,1358,1680,2010]" box="[232,1221,1680,1703]" pageId="6" pageNumber="7">SV (mm) Males Females</paragraph>
<paragraph id="8BD636BAFF84FF92AAF8F923FC5CCD14" blockId="6.[165,1358,1680,2010]" lastBlockId="8.[151,1436,976,1396]" lastPageId="8" lastPageNumber="9" pageId="6" pageNumber="7">
mean (n) range mean (n) range Mt Victoria 24.1 (2) 23.225.0 28.6 (4) 26.831.7 Moroka 22.8 (2) 22.323.3 26.2 (2) 26.026.4 Mt Simpson 20.6 (4) 19.522.5 24.7 (3) 24.125.1 Oya Tabu 25.1 (7) 23.327.5 28.1 (3) 23.931.5 Cloudy Mts ― 27.3 (10) 23.031.1 Sudest Island 28.6 (5) 24.531.0 33.0 (5) 29.637.0 It could perhaps be posited that the Sudest population in fact represents a distinct, separate species that is merely closely related to
<taxonomicName id="4C694D39FF8AFF92AAA7FC6BFDA6CC6C" box="[436,592,1013,1036]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF8AFF92AAA7FC6BFDA6CC6C" box="[436,592,1013,1036]" italics="true" pageId="8" pageNumber="9">C. verrucosus</emphasis>
</taxonomicName>
. If true, one might expect differences to appear in call parameters between the Sudest animals and representatives of
<taxonomicName id="4C694D39FF8AFF92A97DFB87FCFCCC50" box="[622,778,1049,1072]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF8AFF92A97DFB87FCFCCC50" box="[622,778,1049,1072]" italics="true" pageId="8" pageNumber="9">C. verrucosus</emphasis>
</taxonomicName>
from the mainland. This too is not the case. The call of
<taxonomicName id="4C694D39FF8AFF92AE91FB87FEE4CC34" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF8AFF92AE91FB87FEE4CC34" italics="true" pageId="8" pageNumber="9">C. verrucosus</emphasis>
</taxonomicName>
is a single pulsed note emitted at time intervals ranging from 1.552 s in animals recorded by me. To the human ear the call sounds raspy, similar to a thumb dragged over the teeth of a comb. No call parameter serves to distinguish the Sudest population from mainland populations of
<taxonomicName id="4C694D39FF8AFF92A86DFB1BFBECCCFC" box="[894,1050,1157,1180]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF8AFF92A86DFB1BFBECCCFC" box="[894,1050,1157,1180]" italics="true" pageId="8" pageNumber="9">C. verrucosus</emphasis>
</taxonomicName>
(
<tableCitation id="C6EB0301FF8AFF92AF34FB1AFB8BCCFC" box="[1063,1149,1156,1180]" captionStart="TABLE 4" captionStartId="8.[151,239,1439,1463]" captionTargetBox="[151,1435,1522,1904]" captionTargetPageId="8" captionText="TABLE 4. Call parameters for Cophixalus verrucosus from Sudest Island (type locality of C. aimbensis) and four mainland populations. Population numbers refer to sites in Fig. 2. Numbers for call parameters are mean ± SD (range)." httpUri="http://table.plazi.org/id/DF166632FF8AFF92AB84FA01FA7CCDB7" pageId="8" pageNumber="9" tableUuid="DF166632FF8AFF92AB84FA01FA7CCDB7">Table 4</tableCitation>
,
<figureCitation id="13522A3FFF8AFF92AF9AFB1AFB20CCFC" box="[1161,1238,1156,1180]" captionStart="FIGURE 2" captionStartId="4.[151,250,1652,1675]" captionTargetBox="[281,1296,896,1631]" captionTargetId="figure@4.[280,1307,890,1631]" captionTargetPageId="4" captionText="FIGURE 2. Map of southeastern New Guinea and adjacent islands showing type localities for Cophixalus ateles (filled star), C. sisyphus and C. nexipus sp. nov. (filled circle), C. verrucosus (stars), C. albolineatus sp. nov. (filled square), C. clapporum and C. amabilis (filled triangle), and an additional locality for C. albolineatus sp. nov. (open square). Five additional numbered localities from which call samples of C. verrucosus were obtained for this study are: 1 = Sudest Island, type locality for C. aimbensis; 2 = Cloudy Mts.; 3 = N slope of Mt. Simpson; 4 = W slope of Mt. Dayman; and 5 = Fane. Elevational contours are at 1000 m (light gray) and 1500 m (dark gray)." httpUri="https://zenodo.org/record/282921/files/figure.png" pageId="8" pageNumber="9">Figs. 2</figureCitation>
,
<figureCitation id="13522A3FFF8AFF92AFF1FB1AFB05CCFC" box="[1250,1267,1156,1180]" captionStart="FIGURE 4" captionStartId="9.[151,250,1322,1345]" captionTargetBox="[180,1416,205,1289]" captionTargetId="figure@9.[151,1436,193,1301]" captionTargetPageId="9" captionText="FIGURE 4. (A) Waveform, (B) power spectrum, and (C) spectrogram of three calls of Cophixalus verrucosus (BPBM 15261) recorded in the Cloudy Mts., 10 April 2002, air temperature 24.0 C; and (D) waveform, (E) power spectrum, and (F) spectrogram of three calls of C. verrucosus (BPBM 20185) recorded on Sudest Island, 16 April 2004, 26.0 C." httpUri="https://zenodo.org/record/282923/files/figure.png" pageId="8" pageNumber="9">4</figureCitation>
); instead, there is considerable overlap with mainland populations in all these parameters. Of particular interest is that the range of variation in the call parameters from the Sudest sample, with the exception of pulse rate, are entirely contained within the range seen for those same parameters from the Cloudy Mts sample (
<tableCitation id="C6EB0301FF8AFF92AF03FB6EFB91CD68" box="[1040,1127,1264,1288]" captionStart="TABLE 4" captionStartId="8.[151,239,1439,1463]" captionTargetBox="[151,1435,1522,1904]" captionTargetPageId="8" captionText="TABLE 4. Call parameters for Cophixalus verrucosus from Sudest Island (type locality of C. aimbensis) and four mainland populations. Population numbers refer to sites in Fig. 2. Numbers for call parameters are mean ± SD (range)." httpUri="http://table.plazi.org/id/DF166632FF8AFF92AB84FA01FA7CCDB7" pageId="8" pageNumber="9" tableUuid="DF166632FF8AFF92AB84FA01FA7CCDB7">Table 4</tableCitation>
), its geographically nearest neighbor. Because no morphological, color-pattern, or call differences serve to distinguish
<taxonomicName id="4C694D39FF8AFF92AFB4FA8BFACCCD4C" box="[1191,1338,1300,1324]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="aimbensis">
<emphasis id="B91DEAA8FF8AFF92AFB4FA8BFACCCD4C" box="[1191,1338,1300,1324]" italics="true" pageId="8" pageNumber="9">C. aimbensis</emphasis>
</taxonomicName>
from
<taxonomicName id="4C694D39FF8AFF92AE91FA8BFEE4CD30" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91DEAA8FF8AFF92AE91FA8BFEE4CD30" italics="true" pageId="8" pageNumber="9">C. verrucosus</emphasis>
</taxonomicName>
, and both taxa share a number of color-pattern features unique within
<taxonomicName id="4C694D39FF8AFF92AF56FAA6FB32CD30" box="[1093,1220,1336,1360]" class="Amphibia" family="Microhylidae" genus="Cophixalus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B91DEAA8FF8AFF92AF56FAA6FB32CD30" box="[1093,1220,1336,1360]" italics="true" pageId="8" pageNumber="9">Cophixalus</emphasis>
</taxonomicName>
, I synonymize the former with the latter. The taxonomic history for the species becomes:
</paragraph>
</subSubSection>
</treatment>
</document>