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<mods:title id="3EA7D4B5AC85D75F9122781761145F7E">A remarkable new species of Callulina (Amphibia: Anura: Brevicipitidae) with massive, boldly coloured limb glands</mods:title>
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<mods:namePart id="B633DC2F1FC353B17D5D903989D9C754">Gower, David J.</mods:namePart>
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<mods:namePart id="FF871B3B73455E7BF47D37A914F1E54B">Loader, Simon P.</mods:namePart>
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<mods:date id="589564F8DAF284DD22E2724323CBB2B4">2011</mods:date>
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<treatment id="870D0F513A60FFF572FAC1D14F7018D4" ID-DOI="http://doi.org/10.5281/zenodo.5667402" ID-GBIF-Taxon="119422497" ID-Zenodo-Dep="5667402" LSID="urn:lsid:plazi:treatment:870D0F513A60FFF572FAC1D14F7018D4" httpUri="http://treatment.plazi.org/id/870D0F513A60FFF572FAC1D14F7018D4" lastPageId="7" lastPageNumber="22" pageId="1" pageNumber="16">
<subSubSection id="47BEEDCC3A60FFF372FAC1D14D661A49" pageId="1" pageNumber="16" type="nomenclature">
<paragraph id="0F1BBE473A60FFF372FAC1D14FE219E1" blockId="1.[151,470,1495,1554]" box="[151,470,1495,1521]" pageId="1" pageNumber="16">
<heading id="5453092B3A60FFF372FAC1D14FE219E1" bold="true" box="[151,470,1495,1521]" fontSize="11" level="1" pageId="1" pageNumber="16" reason="1">
<emphasis id="3DD062553A60FFF372FAC1D14FE219E1" bold="true" box="[151,470,1495,1521]" pageId="1" pageNumber="16">
<taxonomicName id="C8A4C5C43A60FFF372FAC1D14F4619E1" ID-CoL="697CF" box="[151,370,1495,1521]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="1" pageNumber="16" phylum="Chordata" rank="species" species="meteora" status="sp. nov.">
<emphasis id="3DD062553A60FFF372FAC1D14F4619E1" bold="true" box="[151,370,1495,1521]" italics="true" pageId="1" pageNumber="16">Callulina meteora</emphasis>
</taxonomicName>
<taxonomicNameLabel id="26E3DF2E3A60FFF37314C1D14FE219E1" box="[377,470,1495,1521]" pageId="1" pageNumber="16" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="0F1BBE473A60FFF372FAC1FC4F9F1A02" blockId="1.[151,470,1495,1554]" box="[151,427,1530,1554]" pageId="1" pageNumber="16">
(
<figureCitation id="979FA2C23A60FFF372F2C1FC4F3E1A02" box="[159,266,1530,1554]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURE 3" captionStartId-0="2.[151,250,1991,2014]" captionStartId-1="3.[151,250,1370,1393]" captionStartId-2="4.[151,250,1918,1941]" captionTargetBox-0="[294,1287,553,1956]" captionTargetBox-1="[230,1345,207,1336]" captionTargetBox-2="[233,1356,413,1883]" captionTargetId-0="figure@2.[294,1287,553,1957]" captionTargetId-1="figure@3.[222,1356,193,1351]" captionTargetId-2="figure@4.[224,1363,409,1897]" captionTargetPageId-0="2" captionTargetPageId-1="3" captionTargetPageId-2="4" captionText-0="FIGURE 1. Dorsal, ventral and lateral views of the female holotype (BMNH 2008.450) of Callulina meteora sp. nov." captionText-1="FIGURE 2. Callulina meteora sp. nov. in life from the type locality, showing some of the species variability." captionText-2="FIGURE 3. Sonograms of Callulina species." httpUri-0="https://zenodo.org/record/202449/files/figure.png" httpUri-1="https://zenodo.org/record/202450/files/figure.png" httpUri-2="https://zenodo.org/record/202451/files/figure.png" pageId="1" pageNumber="16">Figs. 13</figureCitation>
,
<figureCitation id="979FA2C23A60FFF3737AC1FC4F771A02" box="[279,323,1530,1554]" captionStart-0="FIGURE 5" captionStart-1="FIGURE 6" captionStart-2="FIGURE 7" captionStartId-0="6.[151,250,1529,1552]" captionStartId-1="8.[151,250,1024,1047]" captionStartId-2="9.[151,250,1290,1313]" captionTargetBox-0="[150,1429,528,1460]" captionTargetBox-1="[231,1354,199,1000]" captionTargetBox-2="[184,1399,303,1259]" captionTargetId-0="figure@6.[144,1429,517,1496]" captionTargetId-1="figure@8.[226,1360,195,1003]" captionTargetId-2="figure@9.[184,1399,303,1259]" captionTargetPageId-0="6" captionTargetPageId-1="8" captionTargetPageId-2="9" captionText-0="FIGURE 5. Phylogenetic tree of all nominal species of Callulina, using a) Maximum likelihood, with bootstrap results given above and symbols below show the results of Kishino Hasegawa tests of differences in log likelihoods between optimal and the best suboptimal trees obtained from converse topological constraints. Presence (+) or lack (-) of support at the p ≤ 0.05 level, b) Maximum Parsimony, numbers above branches are bootstrap proportions, numbers below internal branches are decay indices. The symbols following the decay index values show the results of Templeton tests of differences in length between the MPTs and the best suboptimal trees obtained from converse topological constraints. Presence (+) or lack (-) of support at the p ≤ 0.05 level." captionText-1="FIGURE 6. Map of the Eastern Arc Mountains of Tanzania showing the distribution of the species in the genus Callulina and distribution of C. meteora sp. nov. (red outlined shaded area) in the Nguru Mountains." captionText-2="FIGURE 7. Similarity in enlarged limb glands in Callulina meteora sp. nov. (above) and Nectophrynoides viviparus (below)." httpUri-0="https://zenodo.org/record/202453/files/figure.png" httpUri-1="https://zenodo.org/record/202454/files/figure.png" httpUri-2="https://zenodo.org/record/202455/files/figure.png" pageId="1" pageNumber="16">57</figureCitation>
;
<tableCitation id="42268BFC3A60FFF37322C1FC4F941A02" box="[335,416,1530,1554]" captionStart="TABLE 1" captionStartId="1.[151,235,764,785]" captionTargetBox="[151,1436,844,994]" captionTargetPageId="1" captionText="TABLE 1. Morphometric data for all Callulina meteora sp. nov. specimens (n = 22), all measurements in mm. See Materials and Methods for explanation of abbreviations." httpUri="http://table.plazi.org/id/5BDBEECF3A60FFF372FAC6FA4C581F20" pageId="1" pageNumber="16" tableUuid="5BDBEECF3A60FFF372FAC6FA4C581F20">Table 1</tableCitation>
)
</paragraph>
<paragraph id="0F1BBE473A60FFF372FAC2444D661A49" blockId="1.[151,850,1602,1625]" box="[151,850,1602,1625]" pageId="1" pageNumber="16">
<taxonomicName id="C8A4C5C43A60FFF372FAC2444EC31A48" box="[151,247,1602,1624]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="1" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A60FFF372FAC2444EC31A48" box="[151,247,1602,1624]" italics="true" pageId="1" pageNumber="16">Callulina</emphasis>
</taxonomicName>
sp. 1 Menegon
<emphasis id="3DD062553A60FFF373F4C2454FF11A48" box="[409,453,1602,1624]" italics="true" pageId="1" pageNumber="16">et al</emphasis>
. (2008: p.114, appendix 1, tables 3, 4).
</paragraph>
</subSubSection>
<subSubSection id="47BEEDCC3A60FFF372FAC2834AE31B65" pageId="1" pageNumber="16" type="materials_examined">
<paragraph id="0F1BBE473A60FFF372FAC2834AA81AF5" blockId="1.[151,1436,1669,2018]" pageId="1" pageNumber="16">
<emphasis id="3DD062553A60FFF372FAC2834F3F1A8D" bold="true" box="[151,267,1669,1693]" pageId="1" pageNumber="16">
<typeStatus id="D01F00E53A60FFF372FAC2834F321A8D" box="[151,262,1669,1693]" pageId="1" pageNumber="16" type="holotype">Holotype</typeStatus>
.
</emphasis>
<collectionCode id="69B526823A60FFF3737FC2804F521A8D" box="[274,358,1670,1693]" pageId="1" pageNumber="16">BMNH</collectionCode>
2008.450 (Field tag MW 6825) a mature female (
<figureCitation id="979FA2C23A60FFF371F9C2834DE11A8D" box="[916,981,1669,1693]" captionStart="FIGURE 1" captionStartId="2.[151,250,1991,2014]" captionTargetBox="[294,1287,553,1956]" captionTargetId="figure@2.[294,1287,553,1957]" captionTargetPageId="2" captionText="FIGURE 1. Dorsal, ventral and lateral views of the female holotype (BMNH 2008.450) of Callulina meteora sp. nov." httpUri="https://zenodo.org/record/202449/files/figure.png" pageId="1" pageNumber="16">Fig. 1</figureCitation>
). Collected from the Maskati side of the Nguru South Forest Reserve,
<collectingCountry id="77B3FED73A60FFF37387C2AF4C651AD1" box="[490,593,1705,1729]" name="Tanzania" pageId="1" pageNumber="16">Tanzania</collectingCountry>
,
<geoCoordinate id="6A90D8803A60FFF37032C2AF4D3F1AD1" box="[607,779,1705,1729]" direction="south" orientation="latitude" pageId="1" pageNumber="16" precision="1" value="-6.069028">6.069027778 S</geoCoordinate>
-
<geoCoordinate id="6A90D8803A60FFF37148C2AF4DE11AD1" box="[805,981,1705,1729]" direction="east" orientation="longitude" pageId="1" pageNumber="16" precision="1" value="37.500668">37.50066667 E</geoCoordinate>
,
<quantity id="C85C13A23A60FFF3718FC2AF4A0D1AD1" box="[994,1081,1705,1729]" metricMagnitude="3" metricUnit="m" metricValue="1.98" pageId="1" pageNumber="16" unit="m" value="1980.0">1980 m</quantity>
(
<figureCitation id="979FA2C23A60FFF37627C2AF4ABB1AD1" box="[1098,1167,1705,1729]" captionStart="FIGURE 6" captionStartId="8.[151,250,1024,1047]" captionTargetBox="[231,1354,199,1000]" captionTargetId="figure@8.[226,1360,195,1003]" captionTargetPageId="8" captionText="FIGURE 6. Map of the Eastern Arc Mountains of Tanzania showing the distribution of the species in the genus Callulina and distribution of C. meteora sp. nov. (red outlined shaded area) in the Nguru Mountains." httpUri="https://zenodo.org/record/202454/files/figure.png" pageId="1" pageNumber="16">Fig. 6</figureCitation>
) by David Gower, Roy Hinde, Simon Loader, Hendrik Müller, Maria Müller, and Mark Wilkinson in
<date id="7B1A98873A60FFF37192C2C84AAD1AF5" box="[1023,1177,1741,1766]" pageId="1" pageNumber="16" value="2008-01">January 2008</date>
.
</paragraph>
<paragraph id="0F1BBE473A60FFF372AAC2F44AE31B65" blockId="1.[151,1436,1669,2018]" pageId="1" pageNumber="16">
<emphasis id="3DD062553A60FFF372AAC2F44F721B1A" bold="true" box="[199,326,1778,1802]" pageId="1" pageNumber="16">
<typeStatus id="D01F00E53A60FFF372AAC2F44F751B1A" box="[199,321,1778,1802]" pageId="1" pageNumber="16" type="paratype">Paratypes</typeStatus>
.
</emphasis>
<collectionCode id="69B526823A60FFF3733FC2F74F941B19" box="[338,416,1777,1801]" pageId="1" pageNumber="16">MTSN</collectionCode>
8129-8134 (
<collectionCode id="69B526823A60FFF37054C2F74CBD1B19" box="[569,649,1777,1801]" pageId="1" pageNumber="16">MTSN</collectionCode>
8134, 39 ova in vitellogenesis;
<collectionCode id="69B526823A60FFF37661C2F74A6E1B19" box="[1036,1114,1777,1801]" pageId="1" pageNumber="16">MTSN</collectionCode>
8130 cleared and stained),
<collectionCode id="69B526823A60FFF372FAC3134ED11B3D" box="[151,229,1813,1837]" pageId="1" pageNumber="16">MTSN</collectionCode>
8141, collected by Michele Menegon between
<date id="7B1A98873A60FFF3709AC3134D401B3D" box="[759,884,1813,1837]" pageId="1" pageNumber="16" value="1926-10">October 26</date>
and November 0 2, 2004 in the Nguru South Forest Reserve,
<geoCoordinate id="6A90D8803A60FFF37347C33F4FFC1B41" box="[298,456,1849,1873]" direction="south" orientation="latitude" pageId="1" pageNumber="16" precision="1" value="-6.066302">6.06630176 S</geoCoordinate>
-
<geoCoordinate id="6A90D8803A60FFF37388C33F4CA01B41" box="[485,660,1849,1873]" direction="east" orientation="longitude" pageId="1" pageNumber="16" precision="1" value="37.498028">37.49802743 E</geoCoordinate>
, Nguru Mountains, Morogoro Region.
<collectingCountry id="77B3FED73A60FFF37635C33F4AF41B41" box="[1112,1216,1849,1873]" name="Tanzania" pageId="1" pageNumber="16">Tanzania</collectingCountry>
.
<collectionCode id="69B526823A60FFF376A0C33C4B151B41" box="[1229,1313,1850,1873]" pageId="1" pageNumber="16">BMNH</collectionCode>
2008.118- 451-452-453-454-455-456-457-458-459-460-461-462-463-464 same collection data as
<typeStatus id="D01F00E53A60FFF37600C35B4AE71B65" box="[1133,1235,1885,1909]" pageId="1" pageNumber="16" type="holotype">holotype</typeStatus>
.
</paragraph>
</subSubSection>
<subSubSection id="47BEEDCC3A60FFF172AAC3874FBE1A55" lastPageId="3" lastPageNumber="18" pageId="1" pageNumber="16" type="diagnosis">
<paragraph id="0F1BBE473A60FFF072AAC3874EED1DE4" blockId="1.[151,1436,1669,2018]" lastBlockId="2.[151,1436,151,500]" lastPageId="2" lastPageNumber="17" pageId="1" pageNumber="16">
<emphasis id="3DD062553A60FFF372AAC3874F751B8A" bold="true" box="[199,321,1921,1946]" pageId="1" pageNumber="16">Diagnosis.</emphasis>
The species is assigned to
<taxonomicName id="C8A4C5C43A60FFF37010C3874CD31B89" box="[637,743,1921,1945]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="1" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A60FFF37010C3874CD31B89" box="[637,743,1921,1945]" italics="true" pageId="1" pageNumber="16">Callulina</emphasis>
</taxonomicName>
within
<taxonomicName id="C8A4C5C43A60FFF3712CC3874DEB1B89" box="[833,991,1921,1945]" class="Amphibia" family="Brevicipitidae" kingdom="Animalia" order="Anura" pageId="1" pageNumber="16" phylum="Chordata" rank="family">Brevicipitidae</taxonomicName>
based on the following morphological features: Moderately sized wedge-shaped lobes on the mentomecklian elements, posteroventrally directed (variably reduced/enlarged in
<emphasis id="3DD062553A60FFF373DFC3CF4CC61BF1" box="[434,754,1993,2017]" italics="true" pageId="1" pageNumber="16">
<taxonomicName id="C8A4C5C43A60FFF373DFC3CF4C7C1BF1" box="[434,584,1993,2017]" class="Amphibia" family="Brevicipitidae" genus="Probreviceps" kingdom="Animalia" order="Anura" pageId="1" pageNumber="16" phylum="Chordata" rank="genus">Probreviceps</taxonomicName>
,
<taxonomicName id="C8A4C5C43A60FFF37039C3CF4CC61BF1" box="[596,754,1993,2017]" class="Amphibia" family="Brevicipitidae" genus="Balebreviceps" kingdom="Animalia" order="Anura" pageId="1" pageNumber="16" phylum="Chordata" rank="genus">Balebreviceps</taxonomicName>
</emphasis>
, and
<taxonomicName id="C8A4C5C43A60FFF37142C3CC4DA81BF1" box="[815,924,1994,2017]" class="Amphibia" family="Brevicipitidae" genus="Breviceps" kingdom="Animalia" order="Anura" pageId="1" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A60FFF37142C3CC4DA81BF1" box="[815,924,1994,2017]" italics="true" pageId="1" pageNumber="16">Breviceps</emphasis>
</taxonomicName>
, see
<bibRefCitation id="6B35C3B63A60FFF371B8C3CC4AD51BF1" author="Largen" box="[981,1249,1993,2018]" pageId="1" pageNumber="16" refString="Largen, M. J. &amp; Drewes, R. C. (1989) A new genus and species of brevicipitine frog (Amphibia: Anura: Microhylidae) from high altitude in the mountains of Ethiopia. Tropical Zoology, 2, 13 - 30." type="journal article" year="1989">Largen &amp; Drewes 1989</bibRefCitation>
); cultriform process of the parasphenoid with broad base but narrow alary processes, tapering laterally (cultriform process of the parasphenoids widely variable in breviciptids,
<bibRefCitation id="6B35C3B63A63FFF070DDC4BB4DF11CC4" author="Largen" box="[688,965,188,213]" pageId="2" pageNumber="17" refString="Largen, M. J. &amp; Drewes, R. C. (1989) A new genus and species of brevicipitine frog (Amphibia: Anura: Microhylidae) from high altitude in the mountains of Ethiopia. Tropical Zoology, 2, 13 - 30." type="journal article" year="1989">Largen &amp; Drewes 1989</bibRefCitation>
); nasals almost meet at midline (broadly separated in
<taxonomicName id="C8A4C5C43A63FFF0734EC4E74FA51CE8" box="[291,401,225,248]" class="Amphibia" family="Brevicipitidae" genus="Breviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF0734EC4E74FA51CE8" box="[291,401,225,248]" italics="true" pageId="2" pageNumber="17">Breviceps</emphasis>
</taxonomicName>
and
<taxonomicName id="C8A4C5C43A63FFF073A8C4E64C571CE8" box="[453,611,224,248]" class="Amphibia" family="Brevicipitidae" genus="Balebreviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF073A8C4E64C571CE8" box="[453,611,224,248]" italics="true" pageId="2" pageNumber="17">Balebreviceps</emphasis>
</taxonomicName>
); clavicle well-developed and straight though slightly curved anteriorly at the point of contact between coracoid and scapulae (clavicle straight in
<taxonomicName id="C8A4C5C43A63FFF071A8C5034A061D0C" box="[965,1074,261,284]" class="Amphibia" family="Brevicipitidae" genus="Breviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF071A8C5034A061D0C" box="[965,1074,261,284]" italics="true" pageId="2" pageNumber="17">Breviceps</emphasis>
</taxonomicName>
,
<taxonomicName id="C8A4C5C43A63FFF0762FC5024AE11D0C" box="[1090,1237,260,284]" class="Amphibia" family="Brevicipitidae" genus="Probreviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF0762FC5024AE11D0C" box="[1090,1237,260,284]" italics="true" pageId="2" pageNumber="17">Probreviceps</emphasis>
</taxonomicName>
,
<taxonomicName id="C8A4C5C43A63FFF07689C5024BBE1D0C" box="[1252,1418,260,284]" class="Amphibia" family="Brevicipitidae" genus="Spelaeophryne" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF07689C5024BBE1D0C" box="[1252,1418,260,284]" italics="true" pageId="2" pageNumber="17">Spelaeophryne</emphasis>
</taxonomicName>
); omosternum large (rudimentary or small in
<emphasis id="3DD062553A63FFF070E9C52F4DA61D50" box="[644,914,296,320]" italics="true" pageId="2" pageNumber="17">
<taxonomicName id="C8A4C5C43A63FFF070E9C52F4CC01D50" box="[644,756,297,320]" class="Amphibia" family="Brevicipitidae" genus="Breviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">Breviceps</taxonomicName>
,
<taxonomicName id="C8A4C5C43A63FFF07092C52E4DA61D50" box="[767,914,296,320]" class="Amphibia" family="Brevicipitidae" genus="Probreviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">Probreviceps</taxonomicName>
</emphasis>
, moderate in
<taxonomicName id="C8A4C5C43A63FFF07643C52E4AFF1D50" box="[1070,1227,296,320]" class="Amphibia" family="Brevicipitidae" genus="Balebreviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF07643C52E4AFF1D50" box="[1070,1227,296,320]" italics="true" pageId="2" pageNumber="17">Balebreviceps</emphasis>
</taxonomicName>
); tympanum present and usually well-differentiated (absent in
<taxonomicName id="C8A4C5C43A63FFF070FDC54A4D191D74" box="[656,813,332,356]" class="Amphibia" family="Brevicipitidae" genus="Balebreviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF070FDC54A4D191D74" box="[656,813,332,356]" italics="true" pageId="2" pageNumber="17">Balebreviceps</emphasis>
</taxonomicName>
and
<taxonomicName id="C8A4C5C43A63FFF0710EC54A4ABA1D74" box="[867,1166,332,356]" class="Amphibia" family="Brevicipitidae" genus="Probreviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="species" species="uluguruensis">
<emphasis id="3DD062553A63FFF0710EC54A4ABA1D74" box="[867,1166,332,356]" italics="true" pageId="2" pageNumber="17">Probreviceps uluguruensis</emphasis>
</taxonomicName>
); double condylar articulation between the urostyle and the sacral vertebrae (fused in
<emphasis id="3DD062553A63FFF071E2C5764A811D98" box="[911,1205,368,392]" italics="true" pageId="2" pageNumber="17">
<taxonomicName id="C8A4C5C43A63FFF071E2C5764A1B1D98" box="[911,1071,368,392]" class="Amphibia" family="Brevicipitidae" genus="Balebreviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">Balebreviceps</taxonomicName>
,
<taxonomicName id="C8A4C5C43A63FFF0762CC5774A851D98" box="[1089,1201,369,392]" class="Amphibia" family="Brevicipitidae" genus="Breviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">Breviceps</taxonomicName>
,
</emphasis>
and
<taxonomicName id="C8A4C5C43A63FFF07695C5764BBF1D98" box="[1272,1419,368,392]" class="Amphibia" family="Brevicipitidae" genus="Probreviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF07695C5764BBF1D98" box="[1272,1419,368,392]" italics="true" pageId="2" pageNumber="17">Probreviceps</emphasis>
</taxonomicName>
); truncated terminal phalanges (simple in
<emphasis id="3DD062553A63FFF07035C5924A161DBC" box="[600,1058,404,428]" italics="true" pageId="2" pageNumber="17">
Spelaeophyrne,
<taxonomicName id="C8A4C5C43A63FFF07161C5924D961DBC" box="[780,930,404,428]" class="Amphibia" family="Brevicipitidae" genus="Probreviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">Probreviceps</taxonomicName>
,
<taxonomicName id="C8A4C5C43A63FFF071C3C5934A2A1DBC" box="[942,1054,405,428]" class="Amphibia" family="Brevicipitidae" genus="Breviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">Breviceps</taxonomicName>
,
</emphasis>
and
<taxonomicName id="C8A4C5C43A63FFF07635C5924AC11DBC" box="[1112,1269,404,428]" class="Amphibia" family="Brevicipitidae" genus="Balebreviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF07635C5924AC11DBC" box="[1112,1269,404,428]" italics="true" pageId="2" pageNumber="17">Balebreviceps</emphasis>
</taxonomicName>
); single posterior denticulated row in the palate of
<taxonomicName id="C8A4C5C43A63FFF07029C5BE4C9A1DC0" box="[580,686,440,464]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF07029C5BE4C9A1DC0" box="[580,686,440,464]" italics="true" pageId="2" pageNumber="17">Callulina</emphasis>
</taxonomicName>
(two denticulated rows in
<taxonomicName id="C8A4C5C43A63FFF0718FC5BE4A411DC0" box="[994,1141,440,464]" class="Amphibia" family="Brevicipitidae" genus="Probreviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF0718FC5BE4A411DC0" box="[994,1141,440,464]" italics="true" pageId="2" pageNumber="17">Probreviceps</emphasis>
</taxonomicName>
, glandular mass in
<taxonomicName id="C8A4C5C43A63FFF0773AC5BF4EFD1DE4" class="Amphibia" family="Brevicipitidae" genus="Breviceps" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A63FFF0773AC5BF4EFD1DE4" italics="true" pageId="2" pageNumber="17">Breviceps</emphasis>
</taxonomicName>
).
</paragraph>
<caption id="5BDBEECF3A63FFF072FAC3C14B721BCD" box="[151,1350,1991,2014]" httpUri="https://zenodo.org/record/202449/files/figure.png" pageId="2" pageNumber="17" targetBox="[294,1287,553,1956]" targetPageId="2">
<paragraph id="0F1BBE473A63FFF072FAC3C14B721BCD" blockId="2.[151,1350,1991,2014]" box="[151,1350,1991,2014]" pageId="2" pageNumber="17">
<emphasis id="3DD062553A63FFF072FAC3C14F271BCD" bold="true" box="[151,275,1991,2014]" pageId="2" pageNumber="17">FIGURE 1.</emphasis>
Dorsal, ventral and lateral views of the female holotype (BMNH 2008.450) of
<taxonomicName id="C8A4C5C43A63FFF0765BC3CE4ADB1BCE" box="[1078,1263,1992,2014]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="2" pageNumber="17" phylum="Chordata" rank="species" species="meteora" status="sp. nov.">
<emphasis id="3DD062553A63FFF0765BC3CE4ADB1BCE" box="[1078,1263,1992,2014]" italics="true" pageId="2" pageNumber="17">Callulina meteora</emphasis>
</taxonomicName>
<taxonomicNameLabel id="26E3DF2E3A63FFF07698C3CE4B721BCD" box="[1269,1350,1992,2013]" pageId="2" pageNumber="17" rank="species">sp. nov.</taxonomicNameLabel>
</paragraph>
</caption>
<caption id="5BDBEECF3A62FFF172FAC15C4AD91960" box="[151,1261,1370,1393]" httpUri="https://zenodo.org/record/202450/files/figure.png" pageId="3" pageNumber="18" targetBox="[230,1345,207,1336]" targetPageId="3">
<paragraph id="0F1BBE473A62FFF172FAC15C4AD91960" blockId="3.[151,1261,1370,1393]" box="[151,1261,1370,1393]" pageId="3" pageNumber="18">
<emphasis id="3DD062553A62FFF172FAC15C4F271960" bold="true" box="[151,275,1370,1393]" pageId="3" pageNumber="18">FIGURE 2.</emphasis>
<taxonomicName id="C8A4C5C43A62FFF17374C15D4FE61961" box="[281,466,1371,1393]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="meteora" status="sp. nov.">
<emphasis id="3DD062553A62FFF17374C15D4FE61961" box="[281,466,1371,1393]" italics="true" pageId="3" pageNumber="18">Callulina meteora</emphasis>
</taxonomicName>
<taxonomicNameLabel id="26E3DF2E3A62FFF173B5C15D4C1C1960" box="[472,552,1371,1392]" pageId="3" pageNumber="18" rank="species">
<emphasis id="3DD062553A62FFF173B5C15D4C1C1960" bold="true" box="[472,552,1371,1392]" pageId="3" pageNumber="18">sp. nov.</emphasis>
</taxonomicNameLabel>
in life from the type locality, showing some of the species variability.
</paragraph>
</caption>
<paragraph id="0F1BBE473A62FFF172AAC19B4FBE1A55" blockId="3.[151,1437,1437,2037]" pageId="3" pageNumber="18">
<taxonomicName id="C8A4C5C43A62FFF172AAC19B4FA019A6" box="[199,404,1437,1462]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A62FFF172AAC19B4FA019A6" box="[199,404,1437,1462]" italics="true" pageId="3" pageNumber="18">Callulina meteora</emphasis>
</taxonomicName>
is morphologically distinct from most other species of
<taxonomicName id="C8A4C5C43A62FFF17669C19B4A5A19A5" box="[1028,1134,1437,1461]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A62FFF17669C19B4A5A19A5" box="[1028,1134,1437,1461]" italics="true" pageId="3" pageNumber="18">Callulina</emphasis>
</taxonomicName>
(
<emphasis id="3DD062553A62FFF17613C1984C3B19CA" italics="true" pageId="3" pageNumber="18">
<taxonomicName id="C8A4C5C43A62FFF17613C1984AD019A5" box="[1150,1252,1437,1461]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="kreffti">C. kreffti</taxonomicName>
,
<taxonomicName id="C8A4C5C43A62FFF1769DC1984BAC19A5" box="[1264,1432,1437,1461]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="kisiwamsitu">C. kisiwamsitu</taxonomicName>
,
<taxonomicName id="C8A4C5C43A62FFF172FAC1C44F3A19CA" box="[151,270,1474,1498]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="dawida">C. dawida</taxonomicName>
,
<taxonomicName id="C8A4C5C43A62FFF17376C1C44FB019CA" box="[283,388,1474,1498]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="kanga">C. kanga</taxonomicName>
,
<taxonomicName id="C8A4C5C43A62FFF173FCC1C44C3B19CA" box="[401,527,1474,1498]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="laphami">C. laphami</taxonomicName>
</emphasis>
and
<taxonomicName id="C8A4C5C43A62FFF17025C1C44CF619CA" box="[584,706,1474,1498]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="stanleyi">
<emphasis id="3DD062553A62FFF17025C1C44CF619CA" box="[584,706,1474,1498]" italics="true" pageId="3" pageNumber="18">C. stanleyi</emphasis>
</taxonomicName>
) in having large glands on the limbs. The new species is distinguished from
<taxonomicName id="C8A4C5C43A62FFF1735EC1E04F8A19EE" box="[307,446,1510,1534]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="shengena">
<emphasis id="3DD062553A62FFF1735EC1E04F8A19EE" box="[307,446,1510,1534]" italics="true" pageId="3" pageNumber="18">C. shengena</emphasis>
</taxonomicName>
and
<taxonomicName id="C8A4C5C43A62FFF17399C1E04C5819EE" box="[500,620,1510,1534]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="hanseni">
<emphasis id="3DD062553A62FFF17399C1E04C5819EE" box="[500,620,1510,1534]" italics="true" pageId="3" pageNumber="18">C. hanseni</emphasis>
</taxonomicName>
(also with enlarged limb glands) by the presence of a tympanum (absent in
<taxonomicName id="C8A4C5C43A62FFF172DEC20C4F0A1A32" box="[179,318,1546,1570]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="shengena">
<emphasis id="3DD062553A62FFF172DEC20C4F0A1A32" box="[179,318,1546,1570]" italics="true" pageId="3" pageNumber="18">C. shengena</emphasis>
</taxonomicName>
) and limb glands that are distinctly differently coloured to the rest of the limbs (no distinctive gland colour in
<taxonomicName id="C8A4C5C43A62FFF1736FC2284F4E1A56" box="[258,378,1582,1606]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="3" pageNumber="18" phylum="Chordata" rank="species" species="hanseni">
<emphasis id="3DD062553A62FFF1736FC2284F4E1A56" box="[258,378,1582,1606]" italics="true" pageId="3" pageNumber="18">C. hanseni</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="47BEEDCC3A62FFF572AAC2574AC41D74" lastPageId="7" lastPageNumber="22" pageId="3" pageNumber="18" type="description">
<paragraph id="0F1BBE473A62FFF672AAC2574F591D74" blockId="3.[151,1437,1437,2037]" lastBlockId="4.[151,1436,151,356]" lastPageId="4" lastPageNumber="19" pageId="3" pageNumber="18">
<emphasis id="3DD062553A62FFF172AAC2574FD31A7A" bold="true" box="[199,487,1617,1642]" pageId="3" pageNumber="18">
Description of
<typeStatus id="D01F00E53A62FFF17314C2544FD61A7A" box="[377,482,1618,1642]" pageId="3" pageNumber="18" type="holotype">holotype</typeStatus>
.
</emphasis>
Female. Body stout, head short but as wide as body. Snout truncate in lateral view, snout-tip extending slightly beyond upper and lower jaws. Snout tip rounded at edges, flattened not pointed at apex.
<emphasis id="3DD062553A62FFF172B7C29C4FA91AA2" box="[218,413,1690,1714]" italics="true" pageId="3" pageNumber="18">Canthus rostralis</emphasis>
rounded. Dorsal aspect of head covered by small, rounded, irregular-shaped warts. Ventral region with larger, granular like warts on chin and underside. Eyelids smooth with very small irregular shaped warts. Pupil was horizontal before preservation. Tympanum distinct, suboval (not as tall as long), smooth, with granular warts on slightly raised rim around edge of disc. Dorsum of body with small, irregular glandular masses giving warty appearance. Ventral surface with larger irregular shaped glandular masses, slightly larger and more granular on flanks. Forelimb slender. Massive continuous glands covering dorsal and ventral aspects of forearm. Surface of massive arm gland with smaller, irregular, slightly smoother bumps. Webbing almost absent on hand, only marginal rudimentary skin joining each finger. Distal phalanges moderately long, thick, truncate, expanded only slightly, rounded at edges. Inner tubercle smaller than outer tubercle, separated by a mid-palmar tubercle. First finger shortest, followed by second, fourth, third. Tubercles darker than silvery/metallic background of hand.
<collectingCountry id="77B3FED73A62FFF17709C3DB4BA81BE5" box="[1380,1436,2013,2037]" name="India" pageId="3" pageNumber="18">Hind</collectingCountry>
limbs stout, tibia, metatarsus and carpal area of tarsus covered by an enlarged glandular mass. Distal phalanges of feet moderately long, thick, truncate, expanded only slightly, rounded at edges. First toe marginally the shortest, followed by second, third, fifth, fourth. On toes and fingers, terminal phalanx darker, with a fold of skin, marked by a white line at the dorsal junction between the penultimate and ultimate phalanges. Webbing almost absent on foot, only marginal rudimentary skin joining each toe. Inner and outer tubercle in contact, equal in size. Vent ventro-posteriorly positioned.
</paragraph>
<caption id="5BDBEECF3A65FFF672FAC3784C561B85" box="[151,610,1918,1941]" httpUri="https://zenodo.org/record/202451/files/figure.png" pageId="4" pageNumber="19" targetBox="[233,1356,413,1883]" targetPageId="4">
<paragraph id="0F1BBE473A65FFF672FAC3784C561B85" blockId="4.[151,610,1918,1941]" box="[151,610,1918,1941]" pageId="4" pageNumber="19">
<emphasis id="3DD062553A65FFF672FAC3784F261B84" bold="true" box="[151,274,1918,1941]" pageId="4" pageNumber="19">FIGURE 3.</emphasis>
Sonograms of
<taxonomicName id="C8A4C5C43A65FFF673C1C3794C381B85" box="[428,524,1919,1941]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="4" pageNumber="19" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A65FFF673C1C3794C381B85" box="[428,524,1919,1941]" italics="true" pageId="4" pageNumber="19">Callulina</emphasis>
</taxonomicName>
species.
</paragraph>
</caption>
<paragraph id="0F1BBE473A64FFF772AAC49E4B1D1CC4" blockId="5.[151,1437,151,1293]" pageId="5" pageNumber="20">
<emphasis id="3DD062553A64FFF772AAC49E4FC91CA0" bold="true" box="[199,509,151,176]" pageId="5" pageNumber="20">
Measurement of
<typeStatus id="D01F00E53A64FFF773E2C49E4FCC1CA0" box="[399,504,152,176]" pageId="5" pageNumber="20" type="holotype">holotype</typeStatus>
.
</emphasis>
SUL = 38.2; TL = 13.5; ED = 4.2; TD = 2.0; ETD = 2.2; ND = 2.5; NED =2.8; JW = 13.8; LF3 = 4.4; LT4 = 5.7, TSL = 11; HL = 12.1; NLD =1.7; WDF3 = 1.3; WDTF3 = 1.2; IOD = 6.1.
</paragraph>
<paragraph id="0F1BBE473A64FFF772AAC4D94C351DBC" blockId="5.[151,1437,151,1293]" pageId="5" pageNumber="20">
<emphasis id="3DD062553A64FFF772AAC4D94F141CE8" bold="true" box="[199,288,223,248]" pageId="5" pageNumber="20">Colour.</emphasis>
In preservative, the
<typeStatus id="D01F00E53A64FFF7706EC4D94C511CE7" box="[515,613,223,247]" pageId="5" pageNumber="20" type="holotype">holotype</typeStatus>
dorsal ground colour is pale grey/brown, with darker brown patterning as irregular lateral dorsal markings. The flanks are a paler grey/brown. The dorsal and ventral surfaces of the thighs are dark brown, contrasting strongly with the silvery/metallic colouration of the glands. The ventral surface of the body is cream, with a dark brown colouration on the lateral edges. In life, the
<typeStatus id="D01F00E53A64FFF77199C54A4A621D74" box="[1012,1110,332,356]" pageId="5" pageNumber="20" type="holotype">holotype</typeStatus>
has the same patterning as in preservative but with more vivid colours. The massive glands on the limbs are silvery, giving a striking metallic sheen to the surface (see
<figureCitation id="979FA2C23A64FFF773DDC5924FC71DBC" box="[432,499,404,428]" captionStart="FIGURE 2" captionStartId="3.[151,250,1370,1393]" captionTargetBox="[230,1345,207,1336]" captionTargetId="figure@3.[222,1356,193,1351]" captionTargetPageId="3" captionText="FIGURE 2. Callulina meteora sp. nov. in life from the type locality, showing some of the species variability." httpUri="https://zenodo.org/record/202450/files/figure.png" pageId="5" pageNumber="20">Fig. 2</figureCitation>
).
</paragraph>
<paragraph id="0F1BBE473A64FFF772AAC5B14B321E94" blockId="5.[151,1437,151,1293]" pageId="5" pageNumber="20">
<emphasis id="3DD062553A64FFF772AAC5B14F0F1DC0" bold="true" box="[199,315,439,464]" pageId="5" pageNumber="20">Variation.</emphasis>
The tympanum is usually distinct in
<taxonomicName id="C8A4C5C43A64FFF77085C5BE4D821DC0" box="[744,950,440,464]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="5" pageNumber="20" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A64FFF77085C5BE4D821DC0" box="[744,950,440,464]" italics="true" pageId="5" pageNumber="20">Callulina meteora</emphasis>
</taxonomicName>
, but in some
<typeStatus id="D01F00E53A64FFF7763FC5BE4A891DC0" box="[1106,1213,440,464]" pageId="5" pageNumber="20" type="paratype">paratypes</typeStatus>
(BMNH 2008.464, BMNH 2008.453-455, BMNH 2008.461, BMNH 2008.451) it is poorly demarcated and was measured by dissecting the skin around the region. Otherwise there is little notable (non-colour) morphological variation among individuals apart from between the sexes. Males are significantly smaller in body length (T-test: &lt;0.05, females, SUL=
<quantity id="C85C13A23A64FFF772FAC6414F081E4F" box="[151,316,583,607]" metricMagnitude="-2" metricUnit="m" metricValue="3.7700000000000005" metricValueMax="4.0600000000000005" metricValueMin="3.4799999999999995" pageId="5" pageNumber="20" unit="mm" value="37.7" valueMax="40.6" valueMin="34.8">34.840.6 mm</quantity>
, x=
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, number= 8; males,
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, x=
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, number= 9), and head width (T-test: &lt;0.05). All other morphological characters analysed are not significantly different between the sexes.
</paragraph>
<paragraph id="0F1BBE473A64FFF772AAC6894ABB1F4D" blockId="5.[151,1437,151,1293]" pageId="5" pageNumber="20">
<emphasis id="3DD062553A64FFF772AAC6894FA31EB8" bold="true" box="[199,407,655,680]" pageId="5" pageNumber="20">Colour variation.</emphasis>
<taxonomicName id="C8A4C5C43A64FFF773F2C6964C581EB8" box="[415,620,656,680]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="5" pageNumber="20" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A64FFF773F2C6964C581EB8" box="[415,620,656,680]" italics="true" pageId="5" pageNumber="20">Callulina meteora</emphasis>
</taxonomicName>
individuals show wide colour variability with some rather constant patterns. In all examined individuals, the massive limb glands are paler than surrounding areas, often whitish or silvery with, as in the rest of the body, a metallic sheen (see
<figureCitation id="979FA2C23A64FFF77151C6D14DB71EFF" box="[828,899,727,751]" captionStart="FIGURE 2" captionStartId="3.[151,250,1370,1393]" captionTargetBox="[230,1345,207,1336]" captionTargetId="figure@3.[222,1356,193,1351]" captionTargetPageId="3" captionText="FIGURE 2. Callulina meteora sp. nov. in life from the type locality, showing some of the species variability." httpUri="https://zenodo.org/record/202450/files/figure.png" pageId="5" pageNumber="20">Fig. 2</figureCitation>
). The body can be almost completely white or coppery brown, with darker areas on the dorsum, inguinal, axillary and tympanic zones and on limbs. In some specimens (e.g., MTSN 8129), darker areas on the dorsum are extensive and almost black. The entire animal has a metallic sheen that persists after preservation. Eye colour in life is bright yellow to orange.
</paragraph>
<paragraph id="0F1BBE473A64FFF772AAC7614DD5191C" blockId="5.[151,1437,151,1293]" pageId="5" pageNumber="20">
<emphasis id="3DD062553A64FFF772AAC7614F341F90" bold="true" box="[199,256,871,896]" pageId="5" pageNumber="20">Call.</emphasis>
Advertisement calls of two males
<taxonomicName id="C8A4C5C43A64FFF770C8C76E4D4C1F90" box="[677,888,872,896]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="5" pageNumber="20" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A64FFF770C8C76E4D4C1F90" box="[677,888,872,896]" italics="true" pageId="5" pageNumber="20">Callulina meteora</emphasis>
</taxonomicName>
were recorded at the collecting site by M.M. between
<date id="7B1A98873A64FFF77291C78A4C491FB4" box="[252,637,908,932]" pageId="5" pageNumber="20" value="2004-10-26" valueMax="2004-11-02" valueMin="2004-10-26">October 26 and November 2 2004</date>
both during the day and the night. Calling males were seen at the base of trees. The call is composed of a single periodic pulse train, introductory notes, and repeated notes. These three elements are sometimes arranged in different ways: a single periodic pulse train could be heard with or without introductory notes. Full calls in rainy weather were heard to comprise three or four modules of introductory notes followed by a single pulse train and a final introductory note followed by a group of similar pulse trains. The presence of the introductory note was previously known for the recently described
<taxonomicName id="C8A4C5C43A64FFF7762CC0464A9E1848" box="[1089,1194,1088,1112]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="5" pageNumber="20" phylum="Chordata" rank="species" species="kanga">
<emphasis id="3DD062553A64FFF7762CC0464A9E1848" box="[1089,1194,1088,1112]" italics="true" pageId="5" pageNumber="20">C. kanga</emphasis>
</taxonomicName>
only (
<bibRefCitation id="6B35C3B63A64FFF77696C0464ED5186C" author="Loader" pageId="5" pageNumber="20" refString="Loader, S. P., Muller, H., Gower, D. J., &amp; Menegon, M. (2010 b) Two new species of Callulina (Amphibia: Anura: Brevicipitidae) from the Nguru Mountains, Tanzania. Zootaxa, 2694, 26 - 42." type="journal article" year="2010" yearSuffix="b">
Loader
<emphasis id="3DD062553A64FFF77735C0474BBA1848" box="[1368,1422,1088,1112]" italics="true" pageId="5" pageNumber="20">et al</emphasis>
., 2010b
</bibRefCitation>
). The last call element of
<taxonomicName id="C8A4C5C43A64FFF77067C0634CBC186C" box="[522,648,1125,1148]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="5" pageNumber="20" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A64FFF77067C0634CBC186C" box="[522,648,1125,1148]" italics="true" pageId="5" pageNumber="20">C. meteora</emphasis>
</taxonomicName>
considered alone shows temporal properties similar to the call of the other
<taxonomicName id="C8A4C5C43A64FFF772B4C08E4F7718B0" box="[217,323,1160,1184]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="5" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A64FFF772B4C08E4F7718B0" box="[217,323,1160,1184]" italics="true" pageId="5" pageNumber="20">Callulina</emphasis>
</taxonomicName>
species. The single periodic pulse train is composed of six pulses and has a length ranging from 0.067 to 0.078 sec, but the temporal properties of repeated note remains the same. The introductory notes usually rising in intensity are composed of 16 to 21 pulses and have a length ranging from 0.203 to 0.238 sec. All sound emissions have an average intensity maximum around 1.6 kHz (see
<figureCitation id="979FA2C23A64FFF771FDC0F24DE7191C" box="[912,979,1268,1292]" captionStart="FIGURE 3" captionStartId="4.[151,250,1918,1941]" captionTargetBox="[233,1356,413,1883]" captionTargetId="figure@4.[224,1363,409,1897]" captionTargetPageId="4" captionText="FIGURE 3. Sonograms of Callulina species." httpUri="https://zenodo.org/record/202451/files/figure.png" pageId="5" pageNumber="20">Fig. 3</figureCitation>
).
</paragraph>
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<paragraph id="0F1BBE473A64FFF772FAC3CE4DE31BCE" blockId="5.[151,983,1992,2015]" box="[151,983,1992,2015]" pageId="5" pageNumber="20">
<emphasis id="3DD062553A64FFF772FAC3CE4F261BCE" bold="true" box="[151,274,1992,2015]" pageId="5" pageNumber="20">FIGURE 4.</emphasis>
Views of the habitat at the type locality of
<taxonomicName id="C8A4C5C43A64FFF770A5C3CE4DB51BCF" box="[712,897,1992,2015]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="5" pageNumber="20" phylum="Chordata" rank="species" species="meteora" status="sp. nov.">
<emphasis id="3DD062553A64FFF770A5C3CE4DB51BCF" box="[712,897,1992,2015]" italics="true" pageId="5" pageNumber="20">Callulina meteora</emphasis>
</taxonomicName>
<taxonomicNameLabel id="26E3DF2E3A64FFF771EAC3CF4DE31BCE" box="[903,983,1993,2014]" pageId="5" pageNumber="20" rank="species">sp. nov.</taxonomicNameLabel>
</paragraph>
</caption>
<paragraph id="0F1BBE473A67FFF472AAC4914BBF1DDF" blockId="6.[151,1437,151,464]" pageId="6" pageNumber="21">
<emphasis id="3DD062553A67FFF472AAC4914C211CA0" bold="true" box="[199,533,151,176]" pageId="6" pageNumber="21">Habitat and natural history.</emphasis>
<taxonomicName id="C8A4C5C43A67FFF47071C49E4CDC1CA0" box="[540,744,152,176]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="6" pageNumber="21" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A67FFF47071C49E4CDC1CA0" box="[540,744,152,176]" italics="true" pageId="6" pageNumber="21">Callulina meteora</emphasis>
</taxonomicName>
is partly sympatric with
<taxonomicName id="C8A4C5C43A67FFF47191C49E4A471CA0" box="[1020,1139,152,176]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="6" pageNumber="21" phylum="Chordata" rank="species" species="hanseni">
<emphasis id="3DD062553A67FFF47191C49E4A471CA0" box="[1020,1139,152,176]" italics="true" pageId="6" pageNumber="21">C. hanseni</emphasis>
</taxonomicName>
(
<bibRefCitation id="6B35C3B63A67FFF476ECC4914B521CBF" author="Loader" box="[1153,1382,151,176]" pageId="6" pageNumber="21" refString="Loader, S. P., Muller, H., Gower, D. J., &amp; Menegon, M. (2010 b) Two new species of Callulina (Amphibia: Anura: Brevicipitidae) from the Nguru Mountains, Tanzania. Zootaxa, 2694, 26 - 42." type="journal article" year="2010" yearSuffix="b">
Loader
<emphasis id="3DD062553A67FFF476B4C49F4B3E1CA0" box="[1241,1290,152,176]" italics="true" pageId="6" pageNumber="21">et al</emphasis>
., 2010b
</bibRefCitation>
) and seems to be restricted to the montane and upper montane forest of the Nguru South Forest Reserve (Menegon
<emphasis id="3DD062553A67FFF47730C4BB4BBA1CC4" box="[1373,1422,188,212]" italics="true" pageId="6" pageNumber="21">et al</emphasis>
., 2008). All specimens were collected between
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asl but calls were heard up to
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. This distribution is similar to that reported for
<taxonomicName id="C8A4C5C43A67FFF47385C5024DCC1D0C" authority="Poynton et al., 2009" authorityName="Poynton et al." authorityYear="2009" box="[488,1016,260,284]" class="Amphibia" family="Arthroleptidae" genus="Arthroleptis" kingdom="Animalia" order="Anura" pageId="6" pageNumber="21" phylum="Chordata" rank="species" species="nguruensis">
<emphasis id="3DD062553A67FFF47385C5024CC61D0C" box="[488,754,260,284]" italics="true" pageId="6" pageNumber="21">Arthroleptis nguruensis</emphasis>
(
<bibRefCitation id="6B35C3B63A67FFF4716FC5024DC41D0C" author="Poynton" box="[770,1008,260,284]" pageId="6" pageNumber="21" refString="Poynton, J. C., Menegon, M. &amp; Loader, S. P. (2009) A new giant species of Arthroleptis (Amphibia: Anura) from the forests of the Nguru Mountains, Tanzania. African Journal of Herpetology, 57, 63 - 74." type="journal article" year="2009">
Poynton
<emphasis id="3DD062553A67FFF47107C5034DA91D0C" box="[874,925,260,284]" italics="true" pageId="6" pageNumber="21">et al</emphasis>
., 2009
</bibRefCitation>
)
</taxonomicName>
. Some specimens were found during the day by digging in soft soil and leaf litter accumulating at the base of large trees. This microhabitat together with the presence of strongly keratinized and well-raised metatarsal tubercles is suggestive of semifossoriality. On the basis of the presence of large eggs found in the oviduct of a dissected female, we presume that the species is oviparous with direct development. Although no direct evidence is known for the genus
<taxonomicName id="C8A4C5C43A67FFF47653C5924A9C1DBC" box="[1086,1192,404,428]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="6" pageNumber="21" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A67FFF47653C5924A9C1DBC" box="[1086,1192,404,428]" italics="true" pageId="6" pageNumber="21">Callulina</emphasis>
</taxonomicName>
, oviparity with direct development is regarded as the most likely reproductive mode for this genus of breviciptids (
<bibRefCitation id="6B35C3B63A67FFF476CAC5B14B4B1DDF" author="Muller" box="[1191,1407,439,464]" pageId="6" pageNumber="21" refString="Muller, H., Loader, S. P., Ngalason, W., Howell, K. M. &amp; Gower, D. J. (2007) Reproduction in brevicipitid frogs (Amphibia: Anura: Brevicipitinae) - evidence from Probreviceps m. macrodactylus. Copeia, 2007, 728 - 734." type="journal article" year="2007">
Müller
<emphasis id="3DD062553A67FFF47691C5BF4B191DC0" box="[1276,1325,440,464]" italics="true" pageId="6" pageNumber="21">et al</emphasis>
., 2007
</bibRefCitation>
).
</paragraph>
<caption id="5BDBEECF3A67FFF472FAC1FF4F221AD7" httpUri="https://zenodo.org/record/202453/files/figure.png" pageId="6" pageNumber="21" targetBox="[150,1429,528,1460]" targetPageId="6">
<paragraph id="0F1BBE473A67FFF472FAC1FF4F221AD7" blockId="6.[151,1437,1529,1735]" pageId="6" pageNumber="21">
<emphasis id="3DD062553A67FFF472FAC1FF4F271A1F" bold="true" box="[151,275,1529,1552]" pageId="6" pageNumber="21">FIGURE 5.</emphasis>
Phylogenetic tree of all nominal species of
<emphasis id="3DD062553A67FFF470BBC1FF4D081A1F" box="[726,828,1529,1551]" italics="true" pageId="6" pageNumber="21">
<taxonomicName id="C8A4C5C43A67FFF470BBC1FF4D0C1A1F" box="[726,824,1529,1551]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="6" pageNumber="21" phylum="Chordata" rank="genus">Callulina</taxonomicName>
,
</emphasis>
using a) Maximum likelihood, with bootstrap results given above and symbols below show the results of KishinoHasegawa tests of differences in log likelihoods between optimal and the best suboptimal trees obtained from converse topological constraints. Presence (+) or lack (-) of support at the p ≤ 0.05 level, b) Maximum Parsimony, numbers above branches are bootstrap proportions, numbers below internal branches are decay indices. The symbols following the decay index values show the results of Templeton tests of differences in length between the MPTs and the best suboptimal trees obtained from converse topological constraints. Presence (+) or lack (-) of support at the p ≤ 0.05 level.
</paragraph>
</caption>
<paragraph id="0F1BBE473A67FFF572AAC2F24AC41D74" blockId="6.[151,1436,1780,2029]" lastBlockId="7.[151,1436,151,1220]" lastPageId="7" lastPageNumber="22" pageId="6" pageNumber="21">
<emphasis id="3DD062553A67FFF472AAC2F24F8A1B1D" bold="true" box="[199,446,1780,1805]" pageId="6" pageNumber="21">Conservation status.</emphasis>
Based on current knowledge of the species distribution and habitat preference, the estimated extent of occurrence of
<taxonomicName id="C8A4C5C43A67FFF4738BC31D4C991B23" box="[486,685,1819,1843]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="6" pageNumber="21" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A67FFF4738BC31D4C991B23" box="[486,685,1819,1843]" italics="true" pageId="6" pageNumber="21">Callulina meteora</emphasis>
</taxonomicName>
is equal to or less than
<quantity id="C85C13A23A67FFF471DFC31D4DCC1B23" box="[946,1016,1819,1843]" metricMagnitude="4" metricUnit="m" metricValue="4.2" pageId="6" pageNumber="21" unit="km" value="42.0">42 km</quantity>
2, and the estimated area of occupancy not larger than
<quantity id="C85C13A23A67FFF47356C3444FB41B4A" box="[315,384,1858,1882]" metricMagnitude="4" metricUnit="m" metricValue="2.6" pageId="6" pageNumber="21" unit="km" value="26.0">26 km</quantity>
2; these are respectively the area including the elevational distribution (
<quantity id="C85C13A23A67FFF47611C3444B221B4A" box="[1148,1302,1858,1882]" metricMagnitude="3" metricUnit="m" metricValue="2.04" metricValueMax="2.1" metricValueMin="1.98" pageId="6" pageNumber="21" unit="m" value="2040.0" valueMax="2100.0" valueMin="1980.0">19802100 m</quantity>
) of this species in the Nguru South Forest Reserve and the area included in the polygon obtained by linking the localities where the presence of the species was recorded (
<figureCitation id="979FA2C23A67FFF47056C38C4C4B1BB2" box="[571,639,1930,1954]" captionStart="FIGURE 6" captionStartId="8.[151,250,1024,1047]" captionTargetBox="[231,1354,199,1000]" captionTargetId="figure@8.[226,1360,195,1003]" captionTargetPageId="8" captionText="FIGURE 6. Map of the Eastern Arc Mountains of Tanzania showing the distribution of the species in the genus Callulina and distribution of C. meteora sp. nov. (red outlined shaded area) in the Nguru Mountains." httpUri="https://zenodo.org/record/202454/files/figure.png" pageId="6" pageNumber="21">Fig. 6</figureCitation>
). Therefore, according to Red List (IUCN 2009) categories based on the criterion of an extent of occurrence estimated to be less than
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2, the presence of one population at only a single location, compounded with an observed decline in area, extent and quality of the habitat (IUCN, 2010), we suggest that
<taxonomicName id="C8A4C5C43A66FFF572FAC49E4F271CA0" box="[151,275,152,176]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A66FFF572FAC49E4F271CA0" box="[151,275,152,176]" italics="true" pageId="7" pageNumber="22">C. meteora</emphasis>
</taxonomicName>
qualifies as critically endangered or, more technically, CR B1b (iii). The proposed conservation status would need to be re-evaluated if specimens are recovered below
<quantity id="C85C13A23A66FFF5713FC4BA4D931CC4" box="[850,935,188,212]" metricMagnitude="3" metricUnit="m" metricValue="1.98" pageId="7" pageNumber="22" unit="m" value="1980.0">1980 m</quantity>
, but herpetological surveys examining species turnover between
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have been conducted in the area over the past seven years (S.P.L. and M.M. unpublished data; Owen
<emphasis id="3DD062553A66FFF57372C5034F641D0C" box="[287,336,260,284]" italics="true" pageId="7" pageNumber="22">et al</emphasis>
., 2008) and
<taxonomicName id="C8A4C5C43A66FFF573B2C5034C6E1D0C" box="[479,602,261,284]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A66FFF573B2C5034C6E1D0C" box="[479,602,261,284]" italics="true" pageId="7" pageNumber="22">C. meteora</emphasis>
</taxonomicName>
has never been recovered below
<quantity id="C85C13A23A66FFF571BCC5024A1E1D0C" box="[977,1066,260,284]" metricMagnitude="3" metricUnit="m" metricValue="1.98" pageId="7" pageNumber="22" unit="m" value="1980.0">1980 m</quantity>
. Currently the population density appears to be locally high but with increasing pressure due to land-use changes in the region (pers. obs.), and predicted climate-mediated changes that could affect the high montane zone, the species is clearly facing threats.
</paragraph>
</subSubSection>
<subSubSection id="47BEEDCC3A66FFF572A8C5764F7018D4" pageId="7" pageNumber="22" type="etymology">
<paragraph id="0F1BBE473A66FFF572A8C5764BB41DBC" blockId="7.[151,1436,151,1220]" pageId="7" pageNumber="22">
<emphasis id="3DD062553A66FFF572A8C5764F7F1D98" bold="true" box="[197,331,368,392]" pageId="7" pageNumber="22">Etymology.</emphasis>
The specific epithet is used as an adjective and derives from the greek word
<emphasis id="3DD062553A66FFF576AEC5774B061D98" box="[1219,1330,369,392]" italics="true" pageId="7" pageNumber="22">meteoron,</emphasis>
meaning &quot;thing high up,&quot; in reference to the
<typeStatus id="D01F00E53A66FFF5704EC5934C671DBD" box="[547,595,405,429]" pageId="7" pageNumber="22">type</typeStatus>
locality of the species, situated close to the top of the Nguru Mountains.
</paragraph>
<paragraph id="0F1BBE473A66FFF572A8C5BE4ED81E94" blockId="7.[151,1436,151,1220]" pageId="7" pageNumber="22">
<emphasis id="3DD062553A66FFF572A8C5BE4F871DC0" bold="true" box="[197,435,440,464]" pageId="7" pageNumber="22">Molecular analyses.</emphasis>
To examine the distinctiveness of and relationships among
<taxonomicName id="C8A4C5C43A66FFF57636C5BE4AF11DC0" box="[1115,1221,440,464]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A66FFF57636C5BE4AF11DC0" box="[1115,1221,440,464]" italics="true" pageId="7" pageNumber="22">Callulina</emphasis>
</taxonomicName>
species, we analysed sequence data for 12S, 16S, and cytb mt genes for all nine nominal species, including
<taxonomicName id="C8A4C5C43A66FFF576FDC5DB4B381DE4" box="[1168,1292,477,500]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A66FFF576FDC5DB4B381DE4" box="[1168,1292,477,500]" italics="true" pageId="7" pageNumber="22">C. meteora</emphasis>
</taxonomicName>
. The dataset comprised 24 individuals and 1124 unambiguously aligned characters, of which 707 were constant, 140 variable and uninformative, and 277 informative under parsimony.
<taxonomicName id="C8A4C5C43A66FFF57146C6234A011E2C" box="[811,1077,548,572]" class="Amphibia" family="Brevicipitidae" genus="Breviceps" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="mossambicus">
<emphasis id="3DD062553A66FFF57146C6234A011E2C" box="[811,1077,548,572]" italics="true" pageId="7" pageNumber="22">Breviceps mossambicus</emphasis>
</taxonomicName>
was used to root trees and
<emphasis id="3DD062553A66FFF57707C6234FE31E70" italics="true" pageId="7" pageNumber="22">
<taxonomicName id="C8A4C5C43A66FFF57707C6234F351E70" class="Amphibia" family="Brevicipitidae" genus="Probreviceps" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="genus">Probreviceps</taxonomicName>
m. macrodactylus
</emphasis>
and
<taxonomicName id="C8A4C5C43A66FFF57078C64E4D1F1E70" box="[533,811,584,608]" class="Amphibia" family="Brevicipitidae" genus="Spelaeophryne" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="methneri">
<emphasis id="3DD062553A66FFF57078C64E4D1F1E70" box="[533,811,584,608]" italics="true" pageId="7" pageNumber="22">Spelaeophryne methneri</emphasis>
</taxonomicName>
were used as additional outgroups (see
<bibRefCitation id="6B35C3B63A66FFF5776AC6414ED41E94" author="Loader" pageId="7" pageNumber="22" refString="Loader, S. P., Gower, D. J., Ngalason, W. &amp; Menegon, M. (2010 a) Three new species of Callulina (Amphibia: Anura: Brevicipitidae) highlight local endemism and conservation plight of Africa's Eastern Arc forests. Zoological Journal of the Linnean Society, 160, 496 - 514." type="journal article" year="2010" yearSuffix="a">Loader et al. 2010a</bibRefCitation>
).
</paragraph>
<paragraph id="0F1BBE473A66FFF572AAC6894F7018D4" blockId="7.[151,1436,151,1220]" pageId="7" pageNumber="22">
Parsimony analysis yielded two most parsimonious (MP) trees of 915 steps (
<figureCitation id="979FA2C23A66FFF57640C6894A441EB7" box="[1069,1136,655,679]" captionStart="FIGURE 5" captionStartId="6.[151,250,1529,1552]" captionTargetBox="[150,1429,528,1460]" captionTargetId="figure@6.[144,1429,517,1496]" captionTargetPageId="6" captionText="FIGURE 5. Phylogenetic tree of all nominal species of Callulina, using a) Maximum likelihood, with bootstrap results given above and symbols below show the results of Kishino Hasegawa tests of differences in log likelihoods between optimal and the best suboptimal trees obtained from converse topological constraints. Presence (+) or lack (-) of support at the p ≤ 0.05 level, b) Maximum Parsimony, numbers above branches are bootstrap proportions, numbers below internal branches are decay indices. The symbols following the decay index values show the results of Templeton tests of differences in length between the MPTs and the best suboptimal trees obtained from converse topological constraints. Presence (+) or lack (-) of support at the p ≤ 0.05 level." httpUri="https://zenodo.org/record/202453/files/figure.png" pageId="7" pageNumber="22">Fig. 5</figureCitation>
b). These topologies differ from the optimal likelihood tree (
<figureCitation id="979FA2C23A66FFF57060C6B24C651EDC" box="[525,593,692,716]" captionStart="FIGURE 5" captionStartId="6.[151,250,1529,1552]" captionTargetBox="[150,1429,528,1460]" captionTargetId="figure@6.[144,1429,517,1496]" captionTargetPageId="6" captionText="FIGURE 5. Phylogenetic tree of all nominal species of Callulina, using a) Maximum likelihood, with bootstrap results given above and symbols below show the results of Kishino Hasegawa tests of differences in log likelihoods between optimal and the best suboptimal trees obtained from converse topological constraints. Presence (+) or lack (-) of support at the p ≤ 0.05 level, b) Maximum Parsimony, numbers above branches are bootstrap proportions, numbers below internal branches are decay indices. The symbols following the decay index values show the results of Templeton tests of differences in length between the MPTs and the best suboptimal trees obtained from converse topological constraints. Presence (+) or lack (-) of support at the p ≤ 0.05 level." httpUri="https://zenodo.org/record/202453/files/figure.png" pageId="7" pageNumber="22">Fig. 5</figureCitation>
a). The two MP trees differ in the position of
<taxonomicName id="C8A4C5C43A66FFF57622C6B34AED1EDC" box="[1103,1241,692,716]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="shengena">
<emphasis id="3DD062553A66FFF57622C6B34AED1EDC" box="[1103,1241,692,716]" italics="true" pageId="7" pageNumber="22">C. shengena</emphasis>
</taxonomicName>
and
<taxonomicName id="C8A4C5C43A66FFF5777CC6B34B421EDC" box="[1297,1398,692,716]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="kanga">
<emphasis id="3DD062553A66FFF5777CC6B34B421EDC" box="[1297,1398,692,716]" italics="true" pageId="7" pageNumber="22">C. kanga</emphasis>
</taxonomicName>
. In one MP tree,
<taxonomicName id="C8A4C5C43A66FFF57343C6DE4F8D1EE0" box="[302,441,728,752]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="shengena">
<emphasis id="3DD062553A66FFF57343C6DE4F8D1EE0" box="[302,441,728,752]" italics="true" pageId="7" pageNumber="22">C. shengena</emphasis>
</taxonomicName>
is sister to
<taxonomicName id="C8A4C5C43A66FFF57054C6DE4C811EE0" box="[569,693,728,752]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="laphami">
<emphasis id="3DD062553A66FFF57054C6DE4C811EE0" box="[569,693,728,752]" italics="true" pageId="7" pageNumber="22">C. laphami</emphasis>
</taxonomicName>
, and in the other it is sister to all other (except
<taxonomicName id="C8A4C5C43A66FFF576AAC6DE4B701EE0" box="[1223,1348,728,752]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="laphami">
<emphasis id="3DD062553A66FFF576AAC6DE4B701EE0" box="[1223,1348,728,752]" italics="true" pageId="7" pageNumber="22">C. laphami</emphasis>
</taxonomicName>
)
<taxonomicName id="C8A4C5C43A66FFF57739C6DE4EF71F04" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A66FFF57739C6DE4EF71F04" italics="true" pageId="7" pageNumber="22">Callulina</emphasis>
</taxonomicName>
species.
<taxonomicName id="C8A4C5C43A66FFF57346C6FA4FD51F04" box="[299,481,764,788]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="kanga">
<emphasis id="3DD062553A66FFF57346C6FA4FD51F04" box="[299,481,764,788]" italics="true" pageId="7" pageNumber="22">Callulina kanga</emphasis>
</taxonomicName>
is recovered as sister to a clade comprising a Nguru radiation (
<emphasis id="3DD062553A66FFF576D4C6FB4EF01F28" italics="true" pageId="7" pageNumber="22">
<taxonomicName id="C8A4C5C43A66FFF576D4C6FB4B0E1F04" box="[1209,1338,765,788]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="meteora">C. meteora</taxonomicName>
,
<taxonomicName id="C8A4C5C43A66FFF5772AC6FB4EF01F28" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="hanseni">C. hanseni</taxonomicName>
</emphasis>
),
<emphasis id="3DD062553A66FFF572B6C7264FE61F28" box="[219,466,800,824]" italics="true" pageId="7" pageNumber="22">
<taxonomicName id="C8A4C5C43A66FFF572B6C7264F641F28" box="[219,336,800,824]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="dawida">C. dawida</taxonomicName>
,
<taxonomicName id="C8A4C5C43A66FFF57336C7264FE61F28" box="[347,466,800,824]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="stanleyi">C. stanleyi</taxonomicName>
</emphasis>
and
<taxonomicName id="C8A4C5C43A66FFF5706AC7264C981F28" box="[519,684,800,824]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="kisiwamsitu">
<emphasis id="3DD062553A66FFF5706AC7264C981F28" box="[519,684,800,824]" italics="true" pageId="7" pageNumber="22">C. kisiwamsitu</emphasis>
</taxonomicName>
in the former MP tree. In the second,
<taxonomicName id="C8A4C5C43A66FFF5763CC7264A811F28" box="[1105,1205,800,824]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="kanga">
<emphasis id="3DD062553A66FFF5763CC7264A811F28" box="[1105,1205,800,824]" italics="true" pageId="7" pageNumber="22">C. kanga</emphasis>
</taxonomicName>
is sister to
<taxonomicName id="C8A4C5C43A66FFF5775EC7264BA11F28" box="[1331,1429,800,824]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="kreffti">
<emphasis id="3DD062553A66FFF5775EC7264BA11F28" box="[1331,1429,800,824]" italics="true" pageId="7" pageNumber="22">C. kreffti</emphasis>
</taxonomicName>
. Neither of the alternative topologies is well supported. Most analyses supported the monophyly of
<taxonomicName id="C8A4C5C43A66FFF57699C7424B6A1F4C" box="[1268,1374,836,860]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A66FFF57699C7424B6A1F4C" box="[1268,1374,836,860]" italics="true" pageId="7" pageNumber="22">Callulina</emphasis>
</taxonomicName>
(best trees with a non-monophyletic
<taxonomicName id="C8A4C5C43A66FFF5706CC76E4C5F1F90" box="[513,619,872,896]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A66FFF5706CC76E4C5F1F90" box="[513,619,872,896]" italics="true" pageId="7" pageNumber="22">Callulina</emphasis>
</taxonomicName>
are significantly suboptimal in Templeton but not likelihood topology tests). All nominal species are robustly monophyletic (
<figureCitation id="979FA2C23A66FFF5716CC78A4D731FB4" box="[769,839,908,932]" captionStart="FIGURE 5" captionStartId="6.[151,250,1529,1552]" captionTargetBox="[150,1429,528,1460]" captionTargetId="figure@6.[144,1429,517,1496]" captionTargetPageId="6" captionText="FIGURE 5. Phylogenetic tree of all nominal species of Callulina, using a) Maximum likelihood, with bootstrap results given above and symbols below show the results of Kishino Hasegawa tests of differences in log likelihoods between optimal and the best suboptimal trees obtained from converse topological constraints. Presence (+) or lack (-) of support at the p ≤ 0.05 level, b) Maximum Parsimony, numbers above branches are bootstrap proportions, numbers below internal branches are decay indices. The symbols following the decay index values show the results of Templeton tests of differences in length between the MPTs and the best suboptimal trees obtained from converse topological constraints. Presence (+) or lack (-) of support at the p ≤ 0.05 level." httpUri="https://zenodo.org/record/202453/files/figure.png" pageId="7" pageNumber="22">Fig. 5</figureCitation>
). Likelihood analysis recovered a tree similar to that presented by
<bibRefCitation id="6B35C3B63A66FFF57341C7A94C261FD7" author="Loader" box="[300,530,943,968]" pageId="7" pageNumber="22" refString="Loader, S. P., Gower, D. J., Ngalason, W. &amp; Menegon, M. (2010 a) Three new species of Callulina (Amphibia: Anura: Brevicipitidae) highlight local endemism and conservation plight of Africa's Eastern Arc forests. Zoological Journal of the Linnean Society, 160, 496 - 514." type="journal article" year="2010" yearSuffix="a">
Loader
<emphasis id="3DD062553A66FFF573EFC7B74F871FD8" box="[386,435,944,968]" italics="true" pageId="7" pageNumber="22">et al</emphasis>
. (2010a
</bibRefCitation>
: fig. 8b) differing only in the position of
<taxonomicName id="C8A4C5C43A66FFF571B7C7B64A501FD8" box="[986,1124,944,968]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="shengena">
<emphasis id="3DD062553A66FFF571B7C7B64A501FD8" box="[986,1124,944,968]" italics="true" pageId="7" pageNumber="22">C. shengena</emphasis>
</taxonomicName>
. In
<bibRefCitation id="6B35C3B63A66FFF576FDC7A94B481FD7" box="[1168,1404,943,968]" pageId="7" pageNumber="22" refString="Loader, S. P., Gower, D. J., Ngalason, W. &amp; Menegon, M. (2010 a) Three new species of Callulina (Amphibia: Anura: Brevicipitidae) highlight local endemism and conservation plight of Africa's Eastern Arc forests. Zoological Journal of the Linnean Society, 160, 496 - 514." type="journal article">
Loader
<emphasis id="3DD062553A66FFF5768BC7B74B231FD8" box="[1254,1303,944,968]" italics="true" pageId="7" pageNumber="22">et al</emphasis>
. (2010a)
</bibRefCitation>
<taxonomicName id="C8A4C5C43A66FFF577EFC7B64F351FFC" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="shengena">
<emphasis id="3DD062553A66FFF577EFC7B64F351FFC" italics="true" pageId="7" pageNumber="22">C. shengena</emphasis>
</taxonomicName>
was sister to all other
<taxonomicName id="C8A4C5C43A66FFF57393C7D24C5C1FFC" box="[510,616,980,1004]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A66FFF57393C7D24C5C1FFC" box="[510,616,980,1004]" italics="true" pageId="7" pageNumber="22">Callulina</emphasis>
</taxonomicName>
species (but not well supported) whereas here
<taxonomicName id="C8A4C5C43A66FFF5761AC7D34B361FFC" box="[1143,1282,980,1004]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="shengena">
<emphasis id="3DD062553A66FFF5761AC7D34B361FFC" box="[1143,1282,980,1004]" italics="true" pageId="7" pageNumber="22">C. shengena</emphasis>
</taxonomicName>
is sister to
<taxonomicName id="C8A4C5C43A66FFF577EFC7D34EC71800" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="laphami">
<emphasis id="3DD062553A66FFF577EFC7D34EC71800" italics="true" pageId="7" pageNumber="22">C. laphami</emphasis>
</taxonomicName>
, also only very weakly supported. The new species,
<taxonomicName id="C8A4C5C43A66FFF5712EC7FE4DF41800" box="[835,960,1016,1040]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A66FFF5712EC7FE4DF41800" box="[835,960,1016,1040]" italics="true" pageId="7" pageNumber="22">C. meteora</emphasis>
</taxonomicName>
, forms a clade with
<taxonomicName id="C8A4C5C43A66FFF576C8C7FE4B281800" box="[1189,1308,1016,1040]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="hanseni">
<emphasis id="3DD062553A66FFF576C8C7FE4B281800" box="[1189,1308,1016,1040]" italics="true" pageId="7" pageNumber="22">C. hanseni</emphasis>
</taxonomicName>
in all analyses. Pairwise distances values highlight the genetic distinctiveness of all named
<taxonomicName id="C8A4C5C43A66FFF5765FC01A4AAF1824" box="[1074,1179,1052,1076]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="3DD062553A66FFF5765FC01A4AAF1824" box="[1074,1179,1052,1076]" italics="true" pageId="7" pageNumber="22">Callulina</emphasis>
</taxonomicName>
species (as previously shown in
<bibRefCitation id="6B35C3B63A66FFF57365C0464FC01848" author="Loader" box="[264,500,1088,1112]" pageId="7" pageNumber="22" refString="Loader, S. P., Muller, H., Gower, D. J., &amp; Menegon, M. (2010 b) Two new species of Callulina (Amphibia: Anura: Brevicipitidae) from the Nguru Mountains, Tanzania. Zootaxa, 2694, 26 - 42." type="journal article" year="2010" yearSuffix="b">
Loader
<emphasis id="3DD062553A66FFF5730DC0474FAF1848" box="[352,411,1088,1112]" italics="true" pageId="7" pageNumber="22">et al.</emphasis>
, 2010b
</bibRefCitation>
), including the new species
<taxonomicName id="C8A4C5C43A66FFF57157C0464D8C1848" box="[826,952,1088,1112]" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="meteora">
<emphasis id="3DD062553A66FFF57157C0464D8C1848" box="[826,952,1088,1112]" italics="true" pageId="7" pageNumber="22">C. meteora</emphasis>
</taxonomicName>
(3.84.0 % different to its sister group
<taxonomicName id="C8A4C5C43A66FFF577EFC0464EDA186C" class="Amphibia" family="Brevicipitidae" genus="Callulina" kingdom="Animalia" order="Anura" pageId="7" pageNumber="22" phylum="Chordata" rank="species" species="hanseni">
<emphasis id="3DD062553A66FFF577EFC0464EDA186C" italics="true" pageId="7" pageNumber="22">C. hanseni</emphasis>
</taxonomicName>
). Despite the genetic distinctiveness of the nominal species, the phylogenetic signal in the available mt sequence data is insufficient to provide a compelling resolution of their interrelationships using the methods employed here.
</paragraph>
</subSubSection>
</treatment>
</document>