treatments-xml/data/03/82/87/038287F1FFB1FFEE0B847861FB584129.xml

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<mods:title id="E9BDECEC2AA300B1EC562976FE0BE226">A new species of Dainty Frog (Anura: Pyxicephalidae: Cacosternum) and the first endemic anuran to the Cederberg region of South Africa</mods:title>
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<mods:affiliation id="F0C853C1C1F2AC30FBE395FC200E4A9D">Ghent University, B- 9000, Sint-Pietersnieuwstraat 25, Belgium &amp; Norwegian University of Science and Technology, Postboks 8900, NO- 7491, Trondheim, Norway</mods:affiliation>
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<mods:namePart id="FECF10C0DF162CC40F8ED1E243F2C5D5">Ping, Tyrone</mods:namePart>
<mods:affiliation id="046A940A0E249C5CD37963EF7963E945">P. O. Box 800, Beach Road 37, Mouille Point, Cape Town, South Africa</mods:affiliation>
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<mods:affiliation id="81261D77497563EEEA5839967B767B91">Port Elizabeth Museum (Bayworld), P. O. Box 13147, Humewood 6013, Gqeberha, South Africa &amp; Department of Nature Conservation Management, Natural Resource Science and Management Cluster, Faculty of Science, George Campus, Nelson Mandela University, George, South Africa</mods:affiliation>
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<emphasis id="B95FEAF5FFB1FFE80B847861FE234641" bold="true" box="[151,493,1050,1077]" italics="true" pageId="8" pageNumber="9">Cacosternum cederbergense</emphasis>
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Cederberg Dainty Frog (
<figureCitation id="13102A62FFB1FFE80AB97819FE374609" box="[426,505,1123,1149]" captionStart="FIGURE 6" captionStartId="10.[151,250,1459,1483]" captionTargetBox="[206,1381,181,1434]" captionTargetId="figure-21@10.[206,1381,181,1435]" captionTargetPageId="10" captionText="FIGURE 6. Holotype of Cacosternum cederbergense sp. nov. (PEM A15391) from the eastern end of Pakhuis Pass in the northern Cederberg, Western Cape, South Africa: a) lateral, b) ventral, and a) dorsal view." figureDoi="http://doi.org/10.5281/zenodo.13848153" httpUri="https://zenodo.org/record/13848153/files/figure.png" pageId="8" pageNumber="9">Figs. 6</figureCitation>
, 
<figureCitation id="13102A62FFB1FFE809167819FDDB4609" box="[517,533,1123,1149]" captionStart="FIGURE 7" captionStartId="11.[151,250,928,952]" captionTargetBox="[159,1429,181,904]" captionTargetId="figure-18@11.[159,1429,181,904]" captionTargetPageId="11" captionText="FIGURE 7. Five specimens from the paratype series of Cacosternum cederbergense sp. nov.; a) PEM A15293 (male), b) PEM A15292 (male), c) PEM A15289 (female), and d) PEM A15287 (female)." figureDoi="http://doi.org/10.5281/zenodo.13848157" httpUri="https://zenodo.org/record/13848157/files/figure.png" pageId="8" pageNumber="9">7</figureCitation>
, 
<figureCitation id="13102A62FFB1FFE809327819FDF24609" box="[545,572,1123,1149]" captionStart="FIGURE 8" captionStartId="11.[151,250,1695,1719]" captionTargetBox="[174,1410,1075,1647]" captionTargetId="figure-52@11.[151,1436,1048,1671]" captionTargetPageId="11" captionText="FIGURE 8. Males in breeding condition: a) Cacosternum cederbergense sp. nov. (PEM A15293) from the northern Cederberg, b) C. capense from Hermon, Swartland District, c) C. namaquense from Kamieskroon, Namaqualand, and d) C. karooicum from Vrolijkheid Nature Reserve in the Breede River Valley. Only the C. cederbergense sp. nov. individual was collected as a voucher." figureDoi="http://doi.org/10.5281/zenodo.13848159" httpUri="https://zenodo.org/record/13848159/files/figure.png" pageId="8" pageNumber="9">8a</figureCitation>
)
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<materialsCitation id="3B433CBAFFB1FFE80B8478D1FBD24779" collectingDate="2023-06-14" collectionCode="PEM" country="South Africa" county="The" elevation="491" location="An" municipality="Oliver Angus" pageId="8" pageNumber="9" specimenCode="A15391" specimenCount="1" stateProvince="Western Cape" typeStatus="holotype">
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.
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adult male, 
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<specimenCode id="DB8D9E9CFFB1FFE80AF478D0FDF046B1" box="[487,574,1194,1221]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15391</specimenCode>
(
<figureCitation id="13102A62FFB1FFE8095878D1FD4446B1" box="[587,650,1195,1221]" captionStart="FIGURE 6" captionStartId="10.[151,250,1459,1483]" captionTargetBox="[206,1381,181,1434]" captionTargetId="figure-21@10.[206,1381,181,1435]" captionTargetPageId="10" captionText="FIGURE 6. Holotype of Cacosternum cederbergense sp. nov. (PEM A15391) from the eastern end of Pakhuis Pass in the northern Cederberg, Western Cape, South Africa: a) lateral, b) ventral, and a) dorsal view." figureDoi="http://doi.org/10.5281/zenodo.13848153" httpUri="https://zenodo.org/record/13848153/files/figure.png" pageId="8" pageNumber="9">Fig. 6</figureCitation>
), collected from the eastern end of 
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in the 
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, 
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, 
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(-32.10561˚ 19.06342˚, 
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above sea level
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) by 
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on 
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<collectingDate id="EFD1E9CFFFB1FFE80F6678B5FACC469D" box="[1141,1282,1231,1257]" pageId="8" pageNumber="9" value="2023-06-14">14 June 2023</collectingDate>
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. 
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specimen has a ventral incision from where liver tissue was removed for phylogenetic analyses.
</materialsCitation>
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<materialsCitation id="3B433CBAFFB1FFE80BD4796DFD19470D" collectionCode="PEM" collectorName="Kleinfontein" country="South Africa" county="Nicolas Telford" elevation="515" location="northern Cederberg" municipality="Tyrone Ping" pageId="8" pageNumber="9" specimenCode="A15287, A15288, A15289" specimenCount="16" specimenCount-adult="16" stateProvince="Western Cape" typeStatus="paratype">
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<typeStatus id="54908845FFB1FFE80BD4796DFE8F4745" box="[199,321,1303,1329]" pageId="8" pageNumber="9" type="paratype">Paratypes</typeStatus>
.
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<specimenCount id="9D2DFD6EFFB1FFE80A5C796DFE204745" box="[335,494,1303,1329]" count="16" pageId="8" pageNumber="9" type="adult">Sixteen adults</specimenCount>
collected from different localities in the 
<location id="8EF4603CFFB1FFE808D0796DFB6C4745" LSID="urn:lsid:plazi:treatment:038287F1FFB1FFEE0B847861FB584129:8EF4603CFFB1FFE808D0796DFB6C4745" box="[963,1186,1303,1329]" country="South Africa" county="Nicolas Telford" municipality="Tyrone Ping" name="northern Cederberg" pageId="8" pageNumber="9" stateProvince="Western Cape">northern Cederberg</location>
, 
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, 
<collectingCountry id="F33C7677FFB1FFE80E48796DFF114721" name="South Africa" pageId="8" pageNumber="9">South Africa</collectingCountry>
by 
<location id="8EF4603CFFB1FFE80A1A7941FE6C4721" LSID="urn:lsid:plazi:treatment:038287F1FFB1FFEE0B847861FB584129:8EF4603CFFB1FFE80A1A7941FE6C4721" box="[265,418,1339,1365]" country="South Africa" county="Nicolas Telford" municipality="Tyrone Ping" name="Oliver Angus" pageId="8" pageNumber="9" stateProvince="Western Cape">Oliver Angus</location>
, 
<collectingMunicipality id="6BF0AC9DFFB1FFE80ABE7941FDF84721" box="[429,566,1339,1365]" pageId="8" pageNumber="9">Tyrone Ping</collectingMunicipality>
and 
<collectingCounty id="62F54E6BFFB1FFE8097E7941FCD54721" box="[621,795,1339,1365]" pageId="8" pageNumber="9">Nicolas Telford</collectingCounty>
(
<figureCitation id="13102A62FFB1FFE808397941FCA14721" box="[810,879,1339,1365]" captionStart="FIGURE 7" captionStartId="11.[151,250,928,952]" captionTargetBox="[159,1429,181,904]" captionTargetId="figure-18@11.[159,1429,181,904]" captionTargetPageId="11" captionText="FIGURE 7. Five specimens from the paratype series of Cacosternum cederbergense sp. nov.; a) PEM A15293 (male), b) PEM A15292 (male), c) PEM A15289 (female), and d) PEM A15287 (female)." figureDoi="http://doi.org/10.5281/zenodo.13848157" httpUri="https://zenodo.org/record/13848157/files/figure.png" pageId="8" pageNumber="9">Fig. 7</figureCitation>
). 
<collectionCode id="ED3AAE22FFB1FFE808977941FC704721" box="[900,958,1339,1365]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE808D07940FBEF4721" box="[963,1057,1338,1365]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15287</specimenCode>
, 
<collectionCode id="ED3AAE22FFB1FFE80F3F7941FBA84721" box="[1068,1126,1339,1365]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE80F787940FB064721" box="[1131,1224,1338,1365]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15288</specimenCode>
, and 
<collectionCode id="ED3AAE22FFB1FFE80E107941FAF34721" box="[1283,1341,1339,1365]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE80E517940FA524721" box="[1346,1436,1338,1365]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15289</specimenCode>
from 
<collectorName id="26DE5331FFB1FFE80BC47925FEAB470D" box="[215,357,1375,1401]" pageId="8" pageNumber="9">Kleinfontein</collectorName>
farm (-32.095˚ 19.055˚, 
<quantity id="4CD39B02FFB1FFE809967924FD01470D" box="[645,719,1374,1401]" metricMagnitude="2" metricUnit="m" metricValue="5.15" pageId="8" pageNumber="9" unit="m" value="515.0">
<elevation id="0006D1D4FFB1FFE809967924FD01470D" box="[645,719,1374,1401]" metricMagnitude="2" metricUnit="m" metricValue="5.15" pageId="8" pageNumber="9" unit="m" value="515.0">515 m</elevation>
</quantity>
)
</materialsCitation>
<materialsCitation id="3B433CBAFFB1FFE809F37925FB5B47E9" collectingDate="2022-07-19" collectionCode="PEM" collectorName="Powerlines' &amp; Traveller's Rest" country="South Africa" elevation="487" pageId="8" pageNumber="9" specimenCode="A15290-1, A15292-3" specimenCount="1" stateProvince="Western Cape" typeStatus="paratype">
on 
<date id="FF951027FFB1FFE808167925FC50470D" box="[773,926,1375,1401]" pageId="8" pageNumber="9" value="2022-07-19">
<collectingDate id="EFD1E9CFFFB1FFE808167925FC50470D" box="[773,926,1375,1401]" pageId="8" pageNumber="9" value="2022-07-19">19 July 2022</collectingDate>
</date>
; 
<collectionCode id="ED3AAE22FFB1FFE808BF7925FC28470D" box="[940,998,1375,1401]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE808FE7924FBA8470D" box="[1005,1126,1374,1401]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15290–1</specimenCode>
, and 
<collectionCode id="ED3AAE22FFB1FFE80FB67925FB11470D" box="[1189,1247,1375,1401]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE80FF57924FA93470D" box="[1254,1373,1374,1401]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15292–3</specimenCode>
from the ‘ 
<collectorName id="26DE5331FFB1FFE80BD679F9FE8047E9" box="[197,334,1411,1437]" pageId="8" pageNumber="9">Powerlines’</collectorName>
rock climbing site on 
<collectorName id="26DE5331FFB1FFE8095E79F9FCCE47E9" box="[589,768,1411,1437]" pageId="8" pageNumber="9">Traveller’s Rest</collectorName>
property (-32.086˚ 19.062˚, 
<quantity id="4CD39B02FFB1FFE80F5779F9FB4347E9" box="[1092,1165,1411,1437]" metricMagnitude="2" metricUnit="m" metricValue="4.87" pageId="8" pageNumber="9" unit="m" value="487.0">
<elevation id="0006D1D4FFB1FFE80F5779F9FB4347E9" box="[1092,1165,1411,1437]" metricMagnitude="2" metricUnit="m" metricValue="4.87" pageId="8" pageNumber="9" unit="m" value="487.0">487 m</elevation>
</quantity>
)
</materialsCitation>
<materialsCitation id="3B433CBAFFB1FFE80F8E79F9FA9F47B5" collectingDate="2022-07-19" collectionCode="PEM" collectorName="Hollanders' &amp; Traveller's Rest" country="South Africa" elevation="451" pageId="8" pageNumber="9" specimenCode="A15294" specimenCount="1" stateProvince="Western Cape" typeStatus="paratype">
on 
<date id="FF951027FFB1FFE80FD279F9FA9947E9" box="[1217,1367,1411,1437]" pageId="8" pageNumber="9" value="2022-07-19">
<collectingDate id="EFD1E9CFFFB1FFE80FD279F9FA9947E9" box="[1217,1367,1411,1437]" pageId="8" pageNumber="9" value="2022-07-19">19 July 2022</collectingDate>
</date>
; 
<collectionCode id="ED3AAE22FFB1FFE80E7079F9FA5347E9" box="[1379,1437,1411,1437]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE80B8479DCFF3C47B5" box="[151,242,1446,1473]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15294</specimenCode>
from the ‘ 
<collectorName id="26DE5331FFB1FFE80A7F79DDFE3D47B5" box="[364,499,1447,1473]" pageId="8" pageNumber="9">Hollanders’</collectorName>
rock climbing site on 
<collectorName id="26DE5331FFB1FFE809EF79DDFC7F47B5" box="[764,945,1447,1473]" pageId="8" pageNumber="9">Traveller’s Rest</collectorName>
property (-32.066˚ 19.036˚, 
<quantity id="4CD39B02FFB1FFE80FED79DCFA8647B5" box="[1278,1352,1446,1473]" metricMagnitude="2" metricUnit="m" metricValue="4.51" pageId="8" pageNumber="9" unit="m" value="451.0">
<elevation id="0006D1D4FFB1FFE80FED79DCFA8647B5" box="[1278,1352,1446,1473]" metricMagnitude="2" metricUnit="m" metricValue="4.51" pageId="8" pageNumber="9" unit="m" value="451.0">451 m</elevation>
</quantity>
)
</materialsCitation>
<materialsCitation id="3B433CBAFFB1FFE80E4979DDFB5B4791" collectingDate="2022-07-20" collectionCode="PEM" country="South Africa" elevation="494" location="Bushman's Cave" pageId="8" pageNumber="9" specimenCode="A15295" specimenCount="1" stateProvince="Western Cape" typeStatus="paratype">
on 
<date id="FF951027FFB1FFE80E9379DDFEC44791" pageId="8" pageNumber="9" value="2022-07-20">
<collectingDate id="EFD1E9CFFFB1FFE80E9379DDFEC44791" pageId="8" pageNumber="9" value="2022-07-20">20 July 2022</collectingDate>
</date>
; 
<collectionCode id="ED3AAE22FFB1FFE80A0679B1FE814791" box="[277,335,1483,1509]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE80A4679B0FE7E4791" box="[341,432,1482,1509]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15295</specimenCode>
from near 
<location id="8EF4603CFFB1FFE8093E79B1FD254791" LSID="urn:lsid:plazi:treatment:038287F1FFB1FFEE0B847861FB584129:8EF4603CFFB1FFE8093E79B1FD254791" box="[557,747,1483,1509]" country="South Africa" name="Bushman's Cave" pageId="8" pageNumber="9" stateProvince="Western Cape">Bushman’s Cave</location>
Bergteater (-31.958˚ 18.885˚, 
<quantity id="4CD39B02FFB1FFE80F5779B1FB434791" box="[1092,1165,1483,1509]" metricMagnitude="2" metricUnit="m" metricValue="4.9399999999999995" pageId="8" pageNumber="9" unit="m" value="494.0">
<elevation id="0006D1D4FFB1FFE80F5779B1FB434791" box="[1092,1165,1483,1509]" metricMagnitude="2" metricUnit="m" metricValue="4.9399999999999995" pageId="8" pageNumber="9" unit="m" value="494.0">494 m</elevation>
</quantity>
)
</materialsCitation>
<materialsCitation id="3B433CBAFFB1FFE80F8E79B1FDE74459" collectingDate="2022-07-20" collectionCode="PEM" country="South Africa" elevation="565" location="Klein Bergie' on Alpha Excelsior" pageId="8" pageNumber="9" specimenCode="A15296, A15297, A15298, A15299, A15300-1" specimenCount="1" stateProvince="Western Cape" typeStatus="paratype">
on 
<date id="FF951027FFB1FFE80FD279B1FA994791" box="[1217,1367,1483,1509]" pageId="8" pageNumber="9" value="2022-07-20">
<collectingDate id="EFD1E9CFFFB1FFE80FD279B1FA994791" box="[1217,1367,1483,1509]" pageId="8" pageNumber="9" value="2022-07-20">20 July 2022</collectingDate>
</date>
; 
<collectionCode id="ED3AAE22FFB1FFE80E7079B1FA534791" box="[1379,1437,1483,1509]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE80B847994FF3A447D" box="[151,244,1518,1545]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15296</specimenCode>
, 
<collectionCode id="ED3AAE22FFB1FFE80BEC7995FEF7447D" box="[255,313,1519,1545]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE80A2D7994FE52447D" box="[318,412,1518,1545]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15297</specimenCode>
, 
<collectionCode id="ED3AAE22FFB1FFE80AB47995FE2F447D" box="[423,481,1519,1545]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE80AF57994FD8D447D" box="[486,579,1518,1545]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15298</specimenCode>
, 
<collectionCode id="ED3AAE22FFB1FFE8095D7995FD46447D" box="[590,648,1519,1545]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE8099E7994FD24447D" box="[653,746,1518,1545]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15299</specimenCode>
, and 
<collectionCode id="ED3AAE22FFB1FFE808367995FC91447D" box="[805,863,1519,1545]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE808777994FC14447D" box="[868,986,1518,1545]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15300–1</specimenCode>
from ‘ 
<location id="8EF4603CFFB1FFE80F307995FA52447D" LSID="urn:lsid:plazi:treatment:038287F1FFB1FFEE0B847861FB584129:8EF4603CFFB1FFE80F307995FA52447D" box="[1059,1436,1519,1545]" country="South Africa" name="Klein Bergie' on Alpha Excelsior" pageId="8" pageNumber="9" stateProvince="Western Cape">Klein Bergie’ on Alpha Excelsior</location>
property (-32.106˚ 19.062˚, 
<quantity id="4CD39B02FFB1FFE80AC47A68FDEF4459" box="[471,545,1554,1581]" metricMagnitude="2" metricUnit="m" metricValue="5.65" pageId="8" pageNumber="9" unit="m" value="565.0">
<elevation id="0006D1D4FFB1FFE80AC47A68FDEF4459" box="[471,545,1554,1581]" metricMagnitude="2" metricUnit="m" metricValue="5.65" pageId="8" pageNumber="9" unit="m" value="565.0">565 m</elevation>
</quantity>
)
</materialsCitation>
<materialsCitation id="3B433CBAFFB1FFE809217A69FD304425" collectingDate="2022-07-20" collectionCode="PEM" collectorName="Kleinfontein Campsite &amp; Cederberg" country="South Africa" elevation="515" location="Western Cape" pageId="8" pageNumber="9" specimenCode="A15302" specimenCount="1" stateProvince="Western Cape" typeStatus="paratype">
on 
<date id="FF951027FFB1FFE809447A69FD3E4459" box="[599,752,1555,1581]" pageId="8" pageNumber="9" value="2022-07-20">
<collectingDate id="EFD1E9CFFFB1FFE809447A69FD3E4459" box="[599,752,1555,1581]" pageId="8" pageNumber="9" value="2022-07-20">20 July 2022</collectingDate>
</date>
; and 
<collectionCode id="ED3AAE22FFB1FFE8083D7A69FCA64459" box="[814,872,1555,1581]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB1FFE8087C7A68FC074459" box="[879,969,1554,1581]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="8" pageNumber="9" type="Museum">A15302</specimenCode>
from 
<collectorName id="26DE5331FFB1FFE80F027A69FADA4459" box="[1041,1300,1555,1581]" pageId="8" pageNumber="9">Kleinfontein Campsite</collectorName>
, 
<collectorName id="26DE5331FFB1FFE80E327A69FA564459" box="[1313,1432,1555,1581]" pageId="8" pageNumber="9">Cederberg</collectorName>
, 
<collectingRegion id="49EFF805FFB1FFE80B847A4DFEFB4425" box="[151,309,1591,1617]" country="South Africa" name="Western Cape" pageId="8" pageNumber="9">Western Cape</collectingRegion>
, 
<collectingCountry id="F33C7677FFB1FFE80A527A4DFE1E4425" box="[321,464,1591,1617]" name="South Africa" pageId="8" pageNumber="9">South Africa</collectingCountry>
(-32.095˚ 19.055˚, 
<quantity id="4CD39B02FFB1FFE809B97A4CFD3B4425" box="[682,757,1590,1617]" metricMagnitude="2" metricUnit="m" metricValue="5.15" pageId="8" pageNumber="9" unit="m" value="515.0">
<elevation id="0006D1D4FFB1FFE809B97A4CFD3B4425" box="[682,757,1590,1617]" metricMagnitude="2" metricUnit="m" metricValue="5.15" pageId="8" pageNumber="9" unit="m" value="515.0">515 m</elevation>
</quantity>
)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C331656CFFB1FFE80BD47A21FCCA44ED" pageId="8" pageNumber="9" type="etymology">
<paragraph id="8B9436E7FFB1FFE80BD47A21FCCA44ED" blockId="8.[151,1437,1194,2013]" pageId="8" pageNumber="9">
<emphasis id="B95FEAF5FFB1FFE80BD47A21FE804401" bold="true" box="[199,334,1627,1653]" pageId="8" pageNumber="9">Etymology.</emphasis>
The name is derived from the Cederberg mountain range from where the species occurs. The toponym is formed in agreement with the neuter genus.
</paragraph>
</subSubSection>
<subSubSection id="C331656CFFB1FFE90BD47AD8FB2A46D5" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9" type="diagnosis">
<paragraph id="8B9436E7FFB1FFE80BD47AD8FDAD45E1" blockId="8.[151,1437,1194,2013]" pageId="8" pageNumber="9">
<emphasis id="B95FEAF5FFB1FFE80BD47AD8FE8C44C8" bold="true" box="[199,322,1698,1724]" pageId="8" pageNumber="9">Diagnosis.</emphasis>
Referable to the genus 
<taxonomicName id="4C2B4D64FFB1FFE809467AD9FD2444C9" authorityName="Boulenger" authorityYear="1887" box="[597,746,1699,1725]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B95FEAF5FFB1FFE809467AD9FD2444C9" box="[597,746,1699,1725]" italics="true" pageId="8" pageNumber="9">Cacosternum</emphasis>
</taxonomicName>
based on the absence of webbing between fingers and toes, concealed tympanum and smooth ventrum with brown mottling on white background, and molecular monophyly of the 
<emphasis id="B95FEAF5FFB1FFE80BD27A90FF254570" box="[193,235,1770,1796]" italics="true" pageId="8" pageNumber="9">16S</emphasis>
mitochondrial marker within 
<taxonomicName id="4C2B4D64FFB1FFE8092F7A91FD1F4571" authorityName="Boulenger" authorityYear="1887" box="[572,721,1771,1797]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B95FEAF5FFB1FFE8092F7A91FD1F4571" box="[572,721,1771,1797]" italics="true" pageId="8" pageNumber="9">Cacosternum</emphasis>
</taxonomicName>
. This new species differs from other 
<taxonomicName id="4C2B4D64FFB1FFE80F657A91FAAD4571" box="[1142,1379,1771,1797]" pageId="8" pageNumber="9">
<emphasis id="B95FEAF5FFB1FFE80F657A91FAC54571" box="[1142,1291,1771,1797]" italics="true" pageId="8" pageNumber="9">Cacosternum</emphasis>
species
</taxonomicName>
with regard to the 
<emphasis id="B95FEAF5FFB1FFE80A3B7B74FE9C455C" box="[296,338,1806,1832]" italics="true" pageId="8" pageNumber="9">16S</emphasis>
fragment, by a net uncorrected p-distance value of 1.9–7.1% (Table 1). This new species forms part of the morphologically and phylogenetically distinct 
<taxonomicName id="4C2B4D64FFB1FFE809FF7B49FCA94538" authorityName="Hewitt" authorityYear="1925" box="[748,871,1843,1869]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB1FFE809FF7B49FCA94538" box="[748,871,1843,1869]" italics="true" pageId="8" pageNumber="9">C. capense</emphasis>
</taxonomicName>
group, which can be separated from other species of 
<taxonomicName id="4C2B4D64FFB1FFE80BAB7B2DFE834505" authorityName="Boulenger" authorityYear="1887" box="[184,333,1879,1905]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis id="B95FEAF5FFB1FFE80BAB7B2DFE834505" box="[184,333,1879,1905]" italics="true" pageId="8" pageNumber="9">Cacosternum</emphasis>
</taxonomicName>
on the basis of their large size, pulse count of their ‘creak’ call, and possession of dorsolateral glands that produce defensive secretions.
</paragraph>
<paragraph id="8B9436E7FFB1FFE90BD47BE5FBE14665" blockId="8.[151,1437,1194,2013]" lastBlockId="9.[151,1437,151,1977]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
Morphologically, 
<taxonomicName id="4C2B4D64FFB1FFE80A9C7BE5FD9C45CD" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[399,594,1951,1977]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB1FFE80A9C7BE5FD9C45CD" box="[399,594,1951,1977]" italics="true" pageId="8" pageNumber="9">C. cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB1FFE8094F7BE5FD7945CD" bold="true" box="[604,695,1951,1977]" pageId="8" pageNumber="9">
<taxonomicNameLabel id="A26C578EFFB1FFE8094F7BE5FD7945CD" box="[604,695,1951,1977]" pageId="8" pageNumber="9" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is intermediate between 
<taxonomicName id="4C2B4D64FFB1FFE808CB7BE5FB9A45CC" authorityName="Hewitt" authorityYear="1925" box="[984,1108,1951,1977]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB1FFE808CB7BE5FB9A45CC" box="[984,1108,1951,1977]" italics="true" pageId="8" pageNumber="9">C. capense</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C2B4D64FFB1FFE80F827BE5FAF345CC" box="[1169,1341,1951,1977]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB1FFE80F827BE5FAF345CC" box="[1169,1341,1951,1977]" italics="true" pageId="8" pageNumber="9">C. namaquense</emphasis>
</taxonomicName>
but can be diagnosed based on the following character states: (1) rictal glands: enlarged, forming an uninterrupted arc with the angle of the jaw (interrupted in 
<taxonomicName id="4C2B4D64FFB0FFE909337CEDFD5742C4" authorityName="Hewitt" authorityYear="1925" box="[544,665,151,177]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE909337CEDFD5742C4" box="[544,665,151,177]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
; continuous with the upper lip and reaching the base of the arm in 
<taxonomicName id="4C2B4D64FFB0FFE90E907CEDFEEE42A0" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE90E907CEDFEEE42A0" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
), (2) supratympanic fold: continues posteriorly as a thickened fold (similar to 
<taxonomicName id="4C2B4D64FFB0FFE90F6D7CC1FAE642A0" box="[1150,1320,187,213]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE90F6D7CC1FAE642A0" box="[1150,1320,187,213]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
; indistinct in 
<taxonomicName id="4C2B4D64FFB0FFE90BA77CA5FEE3428C" authorityName="Hewitt" authorityYear="1925" box="[180,301,223,249]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE90BA77CA5FEE3428C" box="[180,301,223,249]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
), (3) nostrils: when viewed from above they are within one nostril diameter of the anterior snout edge (similar in 
<emphasis id="B95FEAF5FFB0FFE90A1D7D79FE424368" box="[270,396,259,285]" italics="true" pageId="9" pageNumber="10">
<taxonomicName id="4C2B4D64FFB0FFE90A1D7D79FE464368" authorityName="Hewitt" authorityYear="1925" box="[270,392,259,285]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">C. capense</taxonomicName>
,
</emphasis>
but located on the anterior margin of the snout in 
<taxonomicName id="4C2B4D64FFB0FFE908B97D79FB9C4368" box="[938,1106,259,285]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE908B97D79FB9C4368" box="[938,1106,259,285]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
), (4) pupil of the eye: absence of a dark streak of variable length running downward, from the pupil into the iris (present in 
<taxonomicName id="4C2B4D64FFB0FFE90FD87D5DFA884334" authorityName="Hewitt" authorityYear="1925" box="[1227,1350,295,321]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE90FD87D5DFA884334" box="[1227,1350,295,321]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C2B4D64FFB0FFE90E907D5DFEEE4310" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE90E907D5DFEEE4310" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
), (5) dorsum: smooth, with a diffusion of small dark protuberances concentrated between the scapular and sacral regions (smooth to granular in 
<taxonomicName id="4C2B4D64FFB0FFE909637D15FCD543FC" box="[624,795,367,393]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE909637D15FCD543FC" box="[624,795,367,393]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
, usually free of warts; covered with large rounded warts in 
<taxonomicName id="4C2B4D64FFB0FFE90BA57DE9FEFF43D8" authorityName="Hewitt" authorityYear="1925" box="[182,305,403,429]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE90BA57DE9FEFF43D8" box="[182,305,403,429]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
, each surrounded by dark flecks), (6) dorsal glands: a pair of dorsoventral subcutaneous glands run longitudinally from the vicinity of the urostyle to the scapula region (in 
<taxonomicName id="4C2B4D64FFB0FFE908AB7DCDFBFF43A4" authorityName="Hewitt" authorityYear="1925" box="[952,1073,439,465]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE908AB7DCDFBFF43A4" box="[952,1073,439,465]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
glands are bean-shaped and two pairs are present, one above the vent in a semi-fused condition, another anterior to the thighs; condition similar in 
<taxonomicName id="4C2B4D64FFB0FFE90E907DA1FEEE406C" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE90E907DA1FEEE406C" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
), (7) dorsal colouration: orange and olive-green tinges common (also present in 
<taxonomicName id="4C2B4D64FFB0FFE90F9D7D85FAC8406C" authorityName="Hewitt" authorityYear="1925" box="[1166,1286,511,537]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE90F9D7D85FAC8406C" box="[1166,1286,511,537]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
but rare in 
<taxonomicName id="4C2B4D64FFB0FFE90E907D85FEEE4048" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE90E907D85FEEE4048" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
), (8) a pale interorbital patch present but diffuse (in 
<taxonomicName id="4C2B4D64FFB0FFE908417E59FC344048" box="[850,1018,547,573]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE908417E59FC344048" box="[850,1018,547,573]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
the patch is free of mottling and forms an uninterrupted interorbital bar; absent in 
<taxonomicName id="4C2B4D64FFB0FFE909657E3DFD3E4014" authorityName="Hewitt" authorityYear="1925" box="[630,752,583,609]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE909657E3DFD3E4014" box="[630,752,583,609]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
, replaced by a dark triangle with a posterior vertex), (9) pale patch present between the scapular and sacral regions, diffuse (in 
<taxonomicName id="4C2B4D64FFB0FFE9087B7E11FBDF40F0" box="[872,1041,619,645]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE9087B7E11FBDF40F0" box="[872,1041,619,645]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
the patch on the back often extends to the flanks and is usually free of mottling; absent in 
<taxonomicName id="4C2B4D64FFB0FFE909F07EF5FC9540DC" authorityName="Hewitt" authorityYear="1925" box="[739,859,655,681]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE909F07EF5FC9540DC" box="[739,859,655,681]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
), (10) ventrum: breeding males with a rim of diffuse grey colouration around the lower jaw, brown spots underneath, extending to the base of the forearms (blotches rapidly coalesce towards the throat in 
<taxonomicName id="4C2B4D64FFB0FFE90AF67EADFD404084" box="[485,654,727,753]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE90AF67EADFD404084" box="[485,654,727,753]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
(
<figureCitation id="13102A62FFB0FFE9098D7EADFD234085" box="[670,749,727,753]" captionStart="FIGURE 8" captionStartId="11.[151,250,1695,1719]" captionTargetBox="[174,1410,1075,1647]" captionTargetId="figure-52@11.[151,1436,1048,1671]" captionTargetPageId="11" captionText="FIGURE 8. Males in breeding condition: a) Cacosternum cederbergense sp. nov. (PEM A15293) from the northern Cederberg, b) C. capense from Hermon, Swartland District, c) C. namaquense from Kamieskroon, Namaqualand, and d) C. karooicum from Vrolijkheid Nature Reserve in the Breede River Valley. Only the C. cederbergense sp. nov. individual was collected as a voucher." figureDoi="http://doi.org/10.5281/zenodo.13848159" httpUri="https://zenodo.org/record/13848159/files/figure.png" pageId="9" pageNumber="10">Fig. 8c</figureCitation>
), becoming paler and smaller; throat black in 
<taxonomicName id="4C2B4D64FFB0FFE90FFB7EADFAAF4084" authorityName="Hewitt" authorityYear="1925" box="[1256,1377,727,753]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE90FFB7EADFAAF4084" box="[1256,1377,727,753]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
(
<figureCitation id="13102A62FFB0FFE90E637EADFF7D4161" captionStart="FIGURE 8" captionStartId="11.[151,250,1695,1719]" captionTargetBox="[174,1410,1075,1647]" captionTargetId="figure-52@11.[151,1436,1048,1671]" captionTargetPageId="11" captionText="FIGURE 8. Males in breeding condition: a) Cacosternum cederbergense sp. nov. (PEM A15293) from the northern Cederberg, b) C. capense from Hermon, Swartland District, c) C. namaquense from Kamieskroon, Namaqualand, and d) C. karooicum from Vrolijkheid Nature Reserve in the Breede River Valley. Only the C. cederbergense sp. nov. individual was collected as a voucher." figureDoi="http://doi.org/10.5281/zenodo.13848159" httpUri="https://zenodo.org/record/13848159/files/figure.png" pageId="9" pageNumber="10">Fig. 8b</figureCitation>
)), (11) belly white with dark blotches often fusing together but never forming stripes (broad stripes often present in 
<taxonomicName id="4C2B4D64FFB0FFE90BA07F65FE92414C" box="[179,348,799,825]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE90BA07F65FE92414C" box="[179,348,799,825]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
and sometimes in 
<taxonomicName id="4C2B4D64FFB0FFE909397F65FD6C414C" authorityName="Hewitt" authorityYear="1925" box="[554,674,799,825]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE909397F65FD6C414C" box="[554,674,799,825]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
), (12) supernumerary tubercles: count of three to five, rounded, outer row closer to palmar tubercle than to proximal subarticular tubercles (same in 
<taxonomicName id="4C2B4D64FFB0FFE908E07F39FBA54128" authorityName="Hewitt" authorityYear="1925" box="[1011,1131,835,861]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE908E07F39FBA54128" box="[1011,1131,835,861]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
, in 
<taxonomicName id="4C2B4D64FFB0FFE90F857F39FAF14128" box="[1174,1343,835,861]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE90F857F39FAF14128" box="[1174,1343,835,861]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
conical, count of four to five, outer row closer to proximal subarticular tubercles than to palmar tubercles), and (13) palmar tubercle: heart-shaped (bifid in 
<taxonomicName id="4C2B4D64FFB0FFE90AEB7FF1FD6D41D0" box="[504,675,907,933]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE90AEB7FF1FD6D41D0" box="[504,675,907,933]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
; depressed in 
<taxonomicName id="4C2B4D64FFB0FFE908557FF1FC0E41D0" authorityName="Hewitt" authorityYear="1925" box="[838,960,907,933]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE908557FF1FC0E41D0" box="[838,960,907,933]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
). Meristic differences indicate the head of 
<taxonomicName id="4C2B4D64FFB0FFE90B847FD5FE9741BD" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[151,345,943,969]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB0FFE90B847FD5FE9741BD" box="[151,345,943,969]" italics="true" pageId="9" pageNumber="10">C. cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB0FFE90A717FD5FE7241BD" bold="true" box="[354,444,943,969]" pageId="9" pageNumber="10">
<taxonomicNameLabel id="A26C578EFFB0FFE90A717FD5FE7241BD" box="[354,444,943,969]" pageId="9" pageNumber="10" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is proportionately wider than 
<taxonomicName id="4C2B4D64FFB0FFE908007FD5FC7041BC" box="[787,958,943,969]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE908007FD5FC7041BC" box="[787,958,943,969]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
, but roughly equal to 
<taxonomicName id="4C2B4D64FFB0FFE90FAE7FD5FAF941BC" authorityName="Hewitt" authorityYear="1925" box="[1213,1335,943,969]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE90FAE7FD5FAF941BC" box="[1213,1335,943,969]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
(
<figureCitation id="13102A62FFB0FFE90E5A7FD5FA4341BD" box="[1353,1421,943,969]" captionStart="FIGURE 3" captionStartId="4.[151,250,1944,1968]" captionTargetBox="[372,1213,1640,1917]" captionTargetId="figure-97@4.[370,1217,1640,1920]" captionTargetPageId="4" captionText="FIGURE 3. Boxplots representing several informative meristic measurements for distinguishing each of the four Cacosternum species assessed in this study." figureDoi="http://doi.org/10.5281/zenodo.13848147" httpUri="https://zenodo.org/record/13848147/files/figure.png" pageId="9" pageNumber="10">Fig. 3</figureCitation>
). 
<taxonomicName id="4C2B4D64FFB0FFE90B847FA9FE014199" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[151,463,979,1005]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB0FFE90B847FA9FE014199" box="[151,463,979,1005]" italics="true" pageId="9" pageNumber="10">Cacosternum cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB0FFE90AC57FA9FDE04199" bold="true" box="[470,558,979,1005]" pageId="9" pageNumber="10">
<taxonomicNameLabel id="A26C578EFFB0FFE90AC57FA9FDE04199" box="[470,558,979,1005]" pageId="9" pageNumber="10" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
also has proportionately larger eyes than 
<taxonomicName id="4C2B4D64FFB0FFE908E47FA9FBA14198" authorityName="Hewitt" authorityYear="1925" box="[1015,1135,979,1005]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE908E47FA9FBA14198" box="[1015,1135,979,1005]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
and 
<emphasis id="B95FEAF5FFB0FFE90FB67FA9FA984198" box="[1189,1366,979,1005]" italics="true" pageId="9" pageNumber="10">
<taxonomicName id="4C2B4D64FFB0FFE90FB67FA9FA9C4198" box="[1189,1362,979,1005]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">C. namaquense</taxonomicName>
.
</emphasis>
Snout length is longer in 
<taxonomicName id="4C2B4D64FFB0FFE90A777F8DFE154664" authorityName="Hewitt" authorityYear="1925" box="[356,475,1015,1041]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE90A777F8DFE154664" box="[356,475,1015,1041]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
than in 
<taxonomicName id="4C2B4D64FFB0FFE909277F8DFD134664" box="[564,733,1015,1041]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE909277F8DFD134664" box="[564,733,1015,1041]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C2B4D64FFB0FFE908017F8DFC1F4665" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[786,977,1015,1041]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB0FFE908017F8DFC1F4665" box="[786,977,1015,1041]" italics="true" pageId="9" pageNumber="10">C. cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB0FFE908C47F8DFBE14665" bold="true" box="[983,1071,1015,1041]" pageId="9" pageNumber="10">
<taxonomicNameLabel id="A26C578EFFB0FFE908C47F8DFBE14665" box="[983,1071,1015,1041]" pageId="9" pageNumber="10" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</paragraph>
<paragraph id="8B9436E7FFB0FFE90BD47861FB2A46D5" blockId="9.[151,1437,151,1977]" pageId="9" pageNumber="10">
The advertisement call consists of a brief pulsed creak emitted at long intervals (
<figureCitation id="13102A62FFB0FFE90F797861FB734641" box="[1130,1213,1051,1077]" captionStart="FIGURE 9" captionStartId="12.[151,250,1442,1466]" captionTargetBox="[191,1396,562,1414]" captionTargetId="figure-219@12.[186,1402,561,1419]" captionTargetPageId="12" captionText="FIGURE 9. Spectrograms of advertisement calls for a) Cacosternum cederbergense sp. nov., b) C. capense, c) C. namaquense, and d) C. karooicum produced using the seewave package in R statistical environment." figureDoi="http://doi.org/10.5281/zenodo.13848161" httpUri="https://zenodo.org/record/13848161/files/figure.png" pageId="9" pageNumber="10">Fig. 9a</figureCitation>
, 
<tableCitation id="C6A9035CFFB0FFE90FD87861FAEC4641" box="[1227,1314,1051,1077]" captionStart="TABLE 3" captionStartId="7.[151,239,151,175]" captionTargetPageId="7" captionText="TABLE 3. Mean values for each of the six acoustic parameters used to characterise each species’ call. See Methods and Materials for an explanation of abbreviations. SD, standard deviation." pageId="9" pageNumber="10">Table 3</tableCitation>
). It differs from the harsh creaks produced by 
<taxonomicName id="4C2B4D64FFB0FFE909307845FD53462C" authorityName="Hewitt" authorityYear="1925" box="[547,669,1087,1113]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB0FFE909307845FD53462C" box="[547,669,1087,1113]" italics="true" pageId="9" pageNumber="10">C. capense</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C2B4D64FFB0FFE909C77845FC4F462C" box="[724,897,1087,1113]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE909C77845FC4F462C" box="[724,897,1087,1113]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
which always have a mean dominant frequency greater than 2 kHz. The call of 
<taxonomicName id="4C2B4D64FFB0FFE90AE77819FD6E4608" box="[500,672,1123,1149]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB0FFE90AE77819FD6E4608" box="[500,672,1123,1149]" italics="true" pageId="9" pageNumber="10">C. namaquense</emphasis>
</taxonomicName>
can have more than 26 pulses and often a duration greater than 
<geoCoordinate id="EE1F5020FFB0FFE90E787819FF6D46D5" degrees="0.30" direction="south" orientation="latitude" pageId="9" pageNumber="10" precision="555" value="-0.3">0.30 s</geoCoordinate>
, whereas that of 
<taxonomicName id="4C2B4D64FFB0FFE90A7178FDFDEB46D5" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[354,549,1159,1185]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB0FFE90A7178FDFDEB46D5" box="[354,549,1159,1185]" italics="true" pageId="9" pageNumber="10">C. cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB0FFE9093F78FDFD4846D5" bold="true" box="[556,646,1159,1185]" pageId="9" pageNumber="10">
<taxonomicNameLabel id="A26C578EFFB0FFE9093F78FDFD4846D5" box="[556,646,1159,1185]" pageId="9" pageNumber="10" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
ranges from 18 to 26 pulses and lasts less than 
<geoCoordinate id="EE1F5020FFB0FFE90F8978FDFB1146D5" box="[1178,1247,1159,1185]" degrees="0.30" direction="south" orientation="latitude" pageId="9" pageNumber="10" precision="555" value="-0.3">0.30 s</geoCoordinate>
.
</paragraph>
</subSubSection>
<subSubSection id="C331656CFFB0FFEE0BD478D0FD9C4049" lastPageId="14" lastPageNumber="15" pageId="9" pageNumber="10" type="description">
<paragraph id="8B9436E7FFB0FFE90BD478D0FD9C4791" blockId="9.[151,1437,151,1977]" pageId="9" pageNumber="10">
<emphasis id="B95FEAF5FFB0FFE90BD478D0FDD346B1" bold="true" box="[199,541,1194,1221]" pageId="9" pageNumber="10">
Description of the 
<typeStatus id="54908845FFB0FFE90ABC78D1FDD646B1" box="[431,536,1195,1221]" pageId="9" pageNumber="10" type="holotype">holotype</typeStatus>
.
</emphasis>
An adult male in breeding condition, SVL 
<quantity id="4CD39B02FFB0FFE90F3278D1FB4746B1" box="[1057,1161,1195,1221]" metricMagnitude="-2" metricUnit="m" metricValue="2.32" pageId="9" pageNumber="10" unit="mm" value="23.2">23.2 mm</quantity>
(
<figureCitation id="13102A62FFB0FFE90F8F78D1FB2B46B1" box="[1180,1253,1195,1221]" captionStart="FIGURE 6" captionStartId="10.[151,250,1459,1483]" captionTargetBox="[206,1381,181,1434]" captionTargetId="figure-21@10.[206,1381,181,1435]" captionTargetPageId="10" captionText="FIGURE 6. Holotype of Cacosternum cederbergense sp. nov. (PEM A15391) from the eastern end of Pakhuis Pass in the northern Cederberg, Western Cape, South Africa: a) lateral, b) ventral, and a) dorsal view." figureDoi="http://doi.org/10.5281/zenodo.13848153" httpUri="https://zenodo.org/record/13848153/files/figure.png" pageId="9" pageNumber="10">Fig. 6</figureCitation>
). Body slender, urostyle posterior to vent, widest around the angle of the jaw (HW = 
<quantity id="4CD39B02FFB0FFE908BD78B5FBC4469D" box="[942,1034,1231,1257]" metricMagnitude="-3" metricUnit="m" metricValue="9.7" pageId="9" pageNumber="10" unit="mm" value="9.7">9.7 mm</quantity>
). The head is bluntly rounded from above, truncated in profile. Head length is about one quarter of the body length (HL/SVL 0.26). 
<emphasis id="B95FEAF5FFB0FFE90FCA7889FA524779" box="[1241,1436,1267,1293]" italics="true" pageId="9" pageNumber="10">Canthus rostralis</emphasis>
rounded, straight from eye to nostril, loreal region slightly concave; nostrils small, rounded, directed anterolaterally, raised marginally. The nostrils are placed closer to the snout than to the eye (EN/SL 0.66). Internarial distance is slightly less than distance between eye and nostril (NN/EN 0.94). Eyes directed anterolaterally, the eyes protrude, and barely visible from below, relatively large (ED/SL 1.02). Distance between anterior corners of eyes more than double the internarial distance (NN/EE 0.45). The angle of the jaw is posterior to a line drawn vertically from the back of the eye. Tympanum not visible.
</paragraph>
<paragraph id="8B9436E7FFB0FFE90BD47995FBD145CD" blockId="9.[151,1437,151,1977]" pageId="9" pageNumber="10">The dorsal surfaces of the head, trunk and limbs are smooth, with small indistinct protuberances; the rictal gland is unbroken, continuing around the posterior angle of the jaw, expanded posteriorly. The supratympanic fold continues posteriorly, thick and glandular to the arm insertion, forming a thickened saddle. Subcutaneous dorsolateral glands running from urostyle along the flanks, small, do not reach the insertion of the forelimbs. The forelimbs are slender, hands small (HAN/SVL 0.26), fingertips bluntly rounded without discs. Relative finger lengths I&lt;II&lt;IV&lt;III; subarticular tubercles distinct, rounded, with one on fingers I and II, two on fingers III and IV. No webbing between fingers, although each digit tapers laterally to a thin margin. Thenar tubercle moderate size, partially obscured by nuptial pad that reaches the distal phalanx of the first finger; palmar tubercles large, heart shaped. Four supernumerary tubercles on palm. Hind limbs moderately long (TIB/SVL 0.42; FOT/SVL 0.45), foot slightly longer than tibia (TIB/ FOT 0.93); thighs are poorly developed, granular on the posterior faces with a diffusion of small glands; relative toe lengths are I&lt;II&lt;V&lt;III&lt;IV. The toe tips are not expanded; subarticular tubercles: one on toes I and II, two on toes III and V, and IV. No webbing between the toes. The inner metatarsal tubercle is large and conical, protruding, equal in width to the tip of the first toe. Outer metatarsal tubercle present as a small bump.</paragraph>
<caption id="DF54666FFFB3FFEA0B8479C9FBEB479B" ID-DOI="http://doi.org/10.5281/zenodo.13848153" ID-Zenodo-Dep="13848153" httpUri="https://zenodo.org/record/13848153/files/figure.png" pageId="10" pageNumber="11" startId="10.[151,250,1459,1483]" targetBox="[206,1381,181,1434]" targetPageId="10" targetType="figure">
<paragraph id="8B9436E7FFB3FFEA0B8479C9FBEB479B" blockId="10.[151,1436,1459,1519]" pageId="10" pageNumber="11">
<emphasis id="B95FEAF5FFB3FFEA0B8479C9FEDB47BF" bold="true" box="[151,277,1459,1483]" pageId="10" pageNumber="11">FIGURE 6.</emphasis>
Holotype of 
<taxonomicName id="4C2B4D64FFB3FFEA0AB779C9FD0947BF" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[420,711,1459,1483]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB3FFEA0AB779C9FD0947BF" box="[420,711,1459,1483]" italics="true" pageId="10" pageNumber="11">Cacosternum cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB3FFEA09C379C9FCED47BF" bold="true" box="[720,803,1459,1483]" pageId="10" pageNumber="11">
<taxonomicNameLabel id="A26C578EFFB3FFEA09C379C9FCED47BF" box="[720,803,1459,1483]" pageId="10" pageNumber="11" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(PEM A15391) from the eastern end of Pakhuis Pass in the northern Cederberg, Western Cape, South Africa: a) lateral, b) ventral, and a) dorsal view.
</paragraph>
</caption>
<paragraph id="8B9436E7FFB3FFEA0BD47A3BFBAC4547" blockId="10.[151,1436,1601,2024]" pageId="10" pageNumber="11">
<emphasis id="B95FEAF5FFB3FFEA0BD47A3BFEA9442F" bold="true" box="[199,359,1601,1627]" pageId="10" pageNumber="11">Colour in life</emphasis>
(
<figureCitation id="13102A62FFB3FFEA0A6B7A3BFE70442F" box="[376,446,1601,1627]" captionStart="FIGURE 6" captionStartId="10.[151,250,1459,1483]" captionTargetBox="[206,1381,181,1434]" captionTargetId="figure-21@10.[206,1381,181,1435]" captionTargetPageId="10" captionText="FIGURE 6. Holotype of Cacosternum cederbergense sp. nov. (PEM A15391) from the eastern end of Pakhuis Pass in the northern Cederberg, Western Cape, South Africa: a) lateral, b) ventral, and a) dorsal view." figureDoi="http://doi.org/10.5281/zenodo.13848153" httpUri="https://zenodo.org/record/13848153/files/figure.png" pageId="10" pageNumber="11">Fig. 6</figureCitation>
). Dorsum with olive mottling over paler beige patches; small protuberances covered in black blotches; pale beige patch between eyes and in the area between the scapula and sacral regions; limbs with transverse dark and pale bands, the darker bands contain olive-green centres; iris hazel with a pale rim around the pupil. Ventrum white with large black blotches that fuse together and are tinged with orange; throat with smaller black spots anteriorly; diffuse grey beneath lower lip that extends to the insertion of the forelimbs; palms, feet and underside of limbs orange with many dark reticulations. Light intensity has a substantial effect on colour shade, where adults become dark in the absence of light and turn paler when exposed to light.
</paragraph>
<paragraph id="8B9436E7FFB3FFEA0BD47B47FBF74593" blockId="10.[151,1436,1601,2024]" pageId="10" pageNumber="11">
<emphasis id="B95FEAF5FFB3FFEA0BD47B47FE164523" bold="true" box="[199,472,1853,1879]" pageId="10" pageNumber="11">Colour in preservative.</emphasis>
Dorsum dark grey with pale grey mottling; pale grey crossbar anterior to the eyes, with anterior pale grey line to snout; supratympanic and subcutaneous dorsolateral glands orange-mustard colouration, with olive darker blotches; pale grey and dark grey transverse bands on limbs. Ventrum white with grey blotches that fuse together; gular grey anterior and white with grey blotches posteriorly; palm of hands dark grey, with fingers and tubercles white; sole of feet uniform dark grey, except subarticular tubercles white.
</paragraph>
<caption id="DF54666FFFB2FFEB0B847FDAFC4E41A8" ID-DOI="http://doi.org/10.5281/zenodo.13848157" ID-Zenodo-Dep="13848157" httpUri="https://zenodo.org/record/13848157/files/figure.png" pageId="11" pageNumber="12" startId="11.[151,250,928,952]" targetBox="[159,1429,181,904]" targetPageId="11" targetType="figure">
<paragraph id="8B9436E7FFB2FFEB0B847FDAFC4E41A8" blockId="11.[151,1436,928,988]" pageId="11" pageNumber="12">
<emphasis id="B95FEAF5FFB2FFEB0B847FDAFEDD41CC" bold="true" box="[151,275,928,952]" pageId="11" pageNumber="12">FIGURE 7.</emphasis>
Five specimens from the paratype series of 
<taxonomicName id="4C2B4D64FFB2FFEB09C37FDBFC3E41CC" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[720,1008,928,952]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB2FFEB09C37FDBFC3E41CC" box="[720,1008,928,952]" italics="true" pageId="11" pageNumber="12">Cacosternum cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB2FFEB08E57FDAFB8941CC" bold="true" box="[1014,1095,928,952]" pageId="11" pageNumber="12">
<taxonomicNameLabel id="A26C578EFFB2FFEB08E57FDAFB8941CC" box="[1014,1095,928,952]" pageId="11" pageNumber="12" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
; a) PEM A15293 (male), b) PEM A15292 (male), c) PEM A15289 (female), and d) PEM A15287 (female).
</paragraph>
</caption>
<caption id="DF54666FFFB2FFEB0B847AE5FF214557" ID-DOI="http://doi.org/10.5281/zenodo.13848159" ID-Zenodo-Dep="13848159" httpUri="https://zenodo.org/record/13848159/files/figure.png" pageId="11" pageNumber="12" startId="11.[151,250,1695,1719]" targetBox="[174,1410,1075,1647]" targetPageId="11" targetType="figure">
<paragraph id="8B9436E7FFB2FFEB0B847AE5FF214557" blockId="11.[151,1437,1695,1827]" pageId="11" pageNumber="12">
<emphasis id="B95FEAF5FFB2FFEB0B847AE5FEDC44C3" bold="true" box="[151,274,1695,1719]" pageId="11" pageNumber="12">FIGURE 8.</emphasis>
Males in breeding condition: a) 
<taxonomicName id="4C2B4D64FFB2FFEB09497ADAFCB744C3" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[602,889,1695,1719]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB2FFEB09497ADAFCB744C3" box="[602,889,1695,1719]" italics="true" pageId="11" pageNumber="12">Cacosternum cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB2FFEB086C7ADAFC0144CC" bold="true" box="[895,975,1696,1720]" pageId="11" pageNumber="12">
<taxonomicNameLabel id="A26C578EFFB2FFEB086C7ADAFC0144CC" box="[895,975,1696,1720]" pageId="11" pageNumber="12" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(PEM A15293) from the northern Cederberg, b) 
<taxonomicName id="4C2B4D64FFB2FFEB0BA67ABEFEE944AF" authorityName="Hewitt" authorityYear="1925" box="[181,295,1732,1755]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB2FFEB0BA67ABEFEE944AF" box="[181,295,1732,1755]" italics="true" pageId="11" pageNumber="12">C. capense</emphasis>
</taxonomicName>
from Hermon, Swartland District, c) 
<taxonomicName id="4C2B4D64FFB2FFEB09A17ABEFC9F44AF" box="[690,849,1732,1755]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB2FFEB09A17ABEFC9F44AF" box="[690,849,1732,1755]" italics="true" pageId="11" pageNumber="12">C. namaquense</emphasis>
</taxonomicName>
from Kamieskroon, Namaqualand, and d) 
<taxonomicName id="4C2B4D64FFB2FFEB0E027ABEFA5244AF" box="[1297,1436,1731,1755]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="karooicum">
<emphasis id="B95FEAF5FFB2FFEB0E027ABEFA5244AF" box="[1297,1436,1731,1755]" italics="true" pageId="11" pageNumber="12">C. karooicum</emphasis>
</taxonomicName>
from Vrolijkheid Nature Reserve in the Breede River Valley. Only the 
<taxonomicName id="4C2B4D64FFB2FFEB08747A92FBD6448B" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[871,1048,1767,1791]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB2FFEB08747A92FBD6448B" box="[871,1048,1767,1791]" italics="true" pageId="11" pageNumber="12">C. cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB2FFEB0F0D7A92FBA14574" bold="true" box="[1054,1135,1768,1792]" pageId="11" pageNumber="12">
<taxonomicNameLabel id="A26C578EFFB2FFEB0F0D7A92FBA14574" box="[1054,1135,1768,1792]" pageId="11" pageNumber="12" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
individual wa 
<emphasis id="B95FEAF5FFB2FFEB0E117A92FAC2448B" box="[1282,1292,1768,1791]" italics="true" pageId="11" pageNumber="12">s</emphasis>
collected as a voucher.
</paragraph>
</caption>
<paragraph id="8B9436E7FFB2FFEC0BD47B0CFD704335" blockId="11.[151,1436,1909,2008]" lastBlockId="12.[151,1436,151,537]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
<emphasis id="B95FEAF5FFB2FFEB0BD47B0CFE7E45FB" bold="true" box="[199,432,1909,1936]" pageId="11" pageNumber="12">
<typeStatus id="54908845FFB2FFEB0BD47B0CFEFA45E4" box="[199,308,1910,1936]" pageId="11" pageNumber="12" type="paratype">Paratype</typeStatus>
variation.
</emphasis>
SVL in females ranges from 23.0– 
<quantity id="4CD39B02FFB2FFEB08527B0CFC6645E4" box="[833,936,1910,1936]" metricMagnitude="-2" metricUnit="m" metricValue="2.9699999999999998" pageId="11" pageNumber="12" unit="mm" value="29.7">29.7 mm</quantity>
, and in males 
<quantity id="4CD39B02FFB2FFEB0F5F7B0CFB3A45E4" box="[1100,1268,1910,1936]" metricMagnitude="-2" metricUnit="m" metricValue="2.3649999999999998" metricValueMax="2.66" metricValueMin="2.07" pageId="11" pageNumber="12" unit="mm" value="23.65" valueMax="26.6" valueMin="20.7">20.7–26.6 mm</quantity>
. The condition of the subcutaneous dorsolateral glands varies in size, correlating with SVL, from prominent in large individuals to indistinct in smaller ones. The density and intensity of dark mottling on the ventrum, the size of individual blotches, and the degree to which they fuse differs from specimen to specimen (
<figureCitation id="13102A62FFB5FFEC08D27CEDFBC642C5" box="[961,1032,151,177]" captionStart="FIGURE 7" captionStartId="11.[151,250,928,952]" captionTargetBox="[159,1429,181,904]" captionTargetId="figure-18@11.[159,1429,181,904]" captionTargetPageId="11" captionText="FIGURE 7. Five specimens from the paratype series of Cacosternum cederbergense sp. nov.; a) PEM A15293 (male), b) PEM A15292 (male), c) PEM A15289 (female), and d) PEM A15287 (female)." figureDoi="http://doi.org/10.5281/zenodo.13848157" httpUri="https://zenodo.org/record/13848157/files/figure.png" pageId="12" pageNumber="13">Fig. 7</figureCitation>
). Males have grey throats (
<figureCitation id="13102A62FFB5FFEC0E517CEDFA4842C5" box="[1346,1414,151,177]" captionStart="FIGURE 7" captionStartId="11.[151,250,928,952]" captionTargetBox="[159,1429,181,904]" captionTargetId="figure-18@11.[159,1429,181,904]" captionTargetPageId="11" captionText="FIGURE 7. Five specimens from the paratype series of Cacosternum cederbergense sp. nov.; a) PEM A15293 (male), b) PEM A15292 (male), c) PEM A15289 (female), and d) PEM A15287 (female)." figureDoi="http://doi.org/10.5281/zenodo.13848157" httpUri="https://zenodo.org/record/13848157/files/figure.png" pageId="12" pageNumber="13">Fig. 7</figureCitation>
ab, 8a), and nuptial pads that reach the distal phalanx of the first finger (
<figureCitation id="13102A62FFB5FFEC08C37CC1FBFC42A1" box="[976,1074,187,213]" captionStart="FIGURE 10" captionStartId="12.[151,250,1929,1953]" captionTargetBox="[379,1208,1563,1903]" captionTargetId="figure-256@12.[376,1211,1562,1906]" captionTargetPageId="12" captionText="FIGURE 10. Diagram of a) the right hand of C. cederbergense sp. nov. holotype male (PEM A15391) in breeding condition, and b) the right hand of a paratype female (PEM A15288). Abbreviations: P = palmar tubercle, T = thenar tubercle, np = nuptial pad, sn = supernumerary tubercles, st = subarticular tubercle." figureDoi="http://doi.org/10.5281/zenodo.13848163" httpUri="https://zenodo.org/record/13848163/files/figure.png" pageId="12" pageNumber="13">Fig. 10a</figureCitation>
). Supernumerary tubercles may be prominent to indistinct, and vary in number, from three to five. Those located beneath the basal subarticular tubercle on the first finger are often obscured by the thenar tubercle. Meristic differences between the 
<typeStatus id="54908845FFB5FFEC0E127D79FAA04369" box="[1281,1390,259,285]" pageId="12" pageNumber="13" type="holotype">holotypes</typeStatus>
and 
<typeStatus id="54908845FFB5FFEC0B847D5DFECB4335" box="[151,261,295,321]" pageId="12" pageNumber="13" type="paratype">paratypes</typeStatus>
, and males and females, are negligible.
</paragraph>
<paragraph id="8B9436E7FFB5FFEC0BD47D30FDC7406D" blockId="12.[151,1436,151,537]" pageId="12" pageNumber="13">
<emphasis id="B95FEAF5FFB5FFEC0BD47D30FE614311" bold="true" box="[199,431,330,357]" pageId="12" pageNumber="13">Advertisement call.</emphasis>
A total of 20 calls from the 
<typeStatus id="54908845FFB5FFEC09E37D31FC9C4311" box="[752,850,331,357]" pageId="12" pageNumber="13" type="holotype">holotype</typeStatus>
specimen (
<collectionCode id="ED3AAE22FFB5FFEC08C17D31FBC34311" box="[978,1037,331,357]" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="12" pageNumber="13" type="Museum">PEM</collectionCode>
<specimenCode id="DB8D9E9CFFB5FFEC0F077D30FBA04311" box="[1044,1134,330,357]" collectionCode="PEM" country="South Africa" httpUri="http://grbio.org/cool/xqn4-34sv" name="Port Elizabeth Museum" pageId="12" pageNumber="13" type="Museum">A15391</specimenCode>
<emphasis id="B95FEAF5FFB5FFEC0F7D7D30FBB94310" bold="true" box="[1134,1143,330,356]" pageId="12" pageNumber="13">)</emphasis>
were recorded at the type locality over a period of 18.5 minutes. 
<collectorName id="26DE5331FFB5FFEC09557D15FDA543FD" box="[582,619,367,393]" pageId="12" pageNumber="13">Air</collectorName>
temperature during this time was 10º 
<collectorName id="26DE5331FFB5FFEC0F007D15FB9043FD" box="[1043,1118,367,393]" pageId="12" pageNumber="13">C. The</collectorName>
call of 
<taxonomicName id="4C2B4D64FFB5FFEC0FA17D15FABA43FD" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[1202,1396,367,393]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB5FFEC0FA17D15FABA43FD" box="[1202,1396,367,393]" italics="true" pageId="12" pageNumber="13">C. cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB5FFEC0E687D15FF0643D9" bold="true" pageId="12" pageNumber="13">
<taxonomicNameLabel id="A26C578EFFB5FFEC0E687D15FF0643D9" pageId="12" pageNumber="13" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
can be described as a short, pulsed bray emitted singly at long intervals (58.3 ± 
<geoCoordinate id="EE1F5020FFB5FFEC0F2C7DE9FB5F43D9" box="[1087,1169,403,429]" degrees="97.23" direction="south" orientation="latitude" pageId="12" pageNumber="13" precision="555" value="-97.23">97.23 s</geoCoordinate>
, 
<figureCitation id="13102A62FFB5FFEC0F8E7DE9FB2243D9" box="[1181,1260,403,429]" captionStart="FIGURE 9" captionStartId="12.[151,250,1442,1466]" captionTargetBox="[191,1396,562,1414]" captionTargetId="figure-219@12.[186,1402,561,1419]" captionTargetPageId="12" captionText="FIGURE 9. Spectrograms of advertisement calls for a) Cacosternum cederbergense sp. nov., b) C. capense, c) C. namaquense, and d) C. karooicum produced using the seewave package in R statistical environment." figureDoi="http://doi.org/10.5281/zenodo.13848161" httpUri="https://zenodo.org/record/13848161/files/figure.png" pageId="12" pageNumber="13">Fig. 9a</figureCitation>
, 
<tableCitation id="C6A9035CFFB5FFEC0FE47DE9FA8543D9" box="[1271,1355,403,429]" captionStart="TABLE 3" captionStartId="7.[151,239,151,175]" captionTargetPageId="7" captionText="TABLE 3. Mean values for each of the six acoustic parameters used to characterise each species’ call. See Methods and Materials for an explanation of abbreviations. SD, standard deviation." pageId="12" pageNumber="13">Table 3</tableCitation>
). 
<collectorName id="26DE5331FFB5FFEC0E4C7DE9FABE43D9" box="[1375,1392,403,429]" pageId="12" pageNumber="13">It</collectorName>
has a mean dominant frequency of 1.8 ± 0.03 kHz, and a dominant frequency range of 0.92 kHz. 
<collectorName id="26DE5331FFB5FFEC0FD37DCDFB3443A5" box="[1216,1274,439,465]" pageId="12" pageNumber="13">Calls</collectorName>
have between 18 and 26 pulses at a rate of 90.6 ± 7.35 pulses per second. 
<collectorName id="26DE5331FFB5FFEC08267DA1FC994381" box="[821,855,475,501]" pageId="12" pageNumber="13">No</collectorName>
territorial call was heard, but no more than a single male was heard calling in a pool.
</paragraph>
<caption id="DF54666FFFB5FFEC0B8479D8FBC947AA" ID-DOI="http://doi.org/10.5281/zenodo.13848161" ID-Zenodo-Dep="13848161" httpUri="https://zenodo.org/record/13848161/files/figure.png" pageId="12" pageNumber="13" startId="12.[151,250,1442,1466]" targetBox="[191,1396,562,1414]" targetPageId="12" targetType="figure">
<paragraph id="8B9436E7FFB5FFEC0B8479D8FBC947AA" blockId="12.[151,1436,1442,1502]" pageId="12" pageNumber="13">
<emphasis id="B95FEAF5FFB5FFEC0B8479D8FEDC47CE" bold="true" box="[151,274,1442,1466]" pageId="12" pageNumber="13">FIGURE 9.</emphasis>
Spectrograms of advertisement calls for a) 
<taxonomicName id="4C2B4D64FFB5FFEC09DB79D8FC2647CE" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[712,1000,1442,1466]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB5FFEC09DB79D8FC2647CE" box="[712,1000,1442,1466]" italics="true" pageId="12" pageNumber="13">Cacosternum cederbergense</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A26C578EFFB5FFEC08FD79D8FBF147CE" box="[1006,1087,1442,1466]" pageId="12" pageNumber="13" rank="species">
<emphasis id="B95FEAF5FFB5FFEC08FD79D8FBF747CE" bold="true" box="[1006,1081,1442,1466]" pageId="12" pageNumber="13">sp. nov</emphasis>
.
</taxonomicNameLabel>
, b) 
<taxonomicName id="4C2B4D64FFB5FFEC0F7579D8FB1B47CE" authorityName="Hewitt" authorityYear="1925" box="[1126,1237,1442,1466]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB5FFEC0F7579D8FB1B47CE" box="[1126,1237,1442,1466]" italics="true" pageId="12" pageNumber="13">C. capense</emphasis>
</taxonomicName>
, c) 
<taxonomicName id="4C2B4D64FFB5FFEC0FE979D8FA5847CE" box="[1274,1430,1442,1466]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB5FFEC0FE979D8FA5847CE" box="[1274,1430,1442,1466]" italics="true" pageId="12" pageNumber="13">C. namaquense</emphasis>
</taxonomicName>
, and d) 
<taxonomicName id="4C2B4D64FFB5FFEC0BCD79BCFEA847AA" box="[222,358,1478,1502]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="karooicum">
<emphasis id="B95FEAF5FFB5FFEC0BCD79BCFEA847AA" box="[222,358,1478,1502]" italics="true" pageId="12" pageNumber="13">C. karooicum</emphasis>
</taxonomicName>
produced using the 
<emphasis id="B95FEAF5FFB5FFEC092779BDFD4647AA" box="[564,648,1479,1502]" italics="true" pageId="12" pageNumber="13">seewave</emphasis>
package in R statistical environment.
</paragraph>
</caption>
<caption id="DF54666FFFB5FFEC0B847BF3FCCD459E" ID-DOI="http://doi.org/10.5281/zenodo.13848163" ID-Zenodo-Dep="13848163" httpUri="https://zenodo.org/record/13848163/files/figure.png" pageId="12" pageNumber="13" startId="12.[151,250,1929,1953]" targetBox="[379,1208,1563,1903]" targetPageId="12" targetType="figure">
<paragraph id="8B9436E7FFB5FFEC0B847BF3FCCD459E" blockId="12.[151,1437,1929,2026]" pageId="12" pageNumber="13">
<emphasis id="B95FEAF5FFB5FFEC0B847BF3FEEE45D6" bold="true" box="[151,288,1929,1954]" pageId="12" pageNumber="13">FIGURE 10.</emphasis>
Diagram of a) the right hand of 
<taxonomicName id="4C2B4D64FFB5FFEC09637BF0FCEF45D6" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[624,801,1930,1954]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB5FFEC09637BF0FCEF45D6" box="[624,801,1930,1954]" italics="true" pageId="12" pageNumber="13">C. cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB5FFEC083B7BF0FCB745D6" bold="true" box="[808,889,1930,1954]" pageId="12" pageNumber="13">
<taxonomicNameLabel id="A26C578EFFB5FFEC083B7BF0FCB745D6" box="[808,889,1930,1954]" pageId="12" pageNumber="13" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
holotype male (PEM A15391) in breeding condition, and b) the right hand of a paratype female (PEM A15288). Abbreviations: P = palmar tubercle, T = thenar tubercle, np = nuptial pad, sn = supernumerary tubercles, st = subarticular tubercle.
</paragraph>
</caption>
<caption id="DF54666FFFB4FFED0B8479B6FDFE4424" ID-DOI="http://doi.org/10.5281/zenodo.13848165" ID-Zenodo-Dep="13848165" httpUri="https://zenodo.org/record/13848165/files/figure.png" pageId="13" pageNumber="14" startId="13.[151,250,1484,1508]" targetBox="[230,1359,181,1460]" targetPageId="13" targetType="figure">
<paragraph id="8B9436E7FFB4FFED0B8479B6FDFE4424" blockId="13.[151,1437,1484,1616]" pageId="13" pageNumber="14">
<emphasis id="B95FEAF5FFB4FFED0B8479B6FED14790" bold="true" box="[151,287,1484,1508]" pageId="13" pageNumber="14">FIGURE 11.</emphasis>
Habitat of 
<taxonomicName id="4C2B4D64FFB4FFED0A8279B7FD8C4790" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[401,578,1484,1508]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB4FFED0A8279B7FD8C4790" box="[401,578,1484,1508]" italics="true" pageId="13" pageNumber="14">C. cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB4FFED095B79B7FD574791" bold="true" box="[584,665,1485,1509]" pageId="13" pageNumber="14">
<taxonomicNameLabel id="A26C578EFFB4FFED095B79B7FD574791" box="[584,665,1485,1509]" pageId="13" pageNumber="14" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
a–b) typical breeding habitat consisting of small rock pools with high walls and overhangs, c) a collection of pools at the Hollanders locality, only certain pools are used for breeding, d) a large boulder complex near Bushman’s Cave Bergteater with several breeding pools on top, raised several meters above the ground and scalable only by climbing vertical faces.
</paragraph>
</caption>
<paragraph id="8B9436E7FFB4FFEE0BD47AD8FE234369" blockId="13.[151,1437,1698,2013]" lastBlockId="14.[151,1437,151,861]" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14">
<emphasis id="B95FEAF5FFB4FFED0BD47AD8FE6444C8" bold="true" box="[199,426,1698,1725]" pageId="13" pageNumber="14">Habits and Habitat</emphasis>
(
<figureCitation id="13102A62FFB4FFED0AAA7AD9FDC744C9" box="[441,521,1699,1725]" captionStart="FIGURE 11" captionStartId="13.[151,250,1484,1508]" captionTargetBox="[230,1359,181,1460]" captionTargetId="figure-18@13.[228,1359,181,1460]" captionTargetPageId="13" captionText="FIGURE 11. Habitat of C. cederbergense sp. nov. a–b) typical breeding habitat consisting of small rock pools with high walls and overhangs, c) a collection of pools at the Hollanders locality, only certain pools are used for breeding, d) a large boulder complex near Bushman’s Cave Bergteater with several breeding pools on top, raised several meters above the ground and scalable only by climbing vertical faces." figureDoi="http://doi.org/10.5281/zenodo.13848165" httpUri="https://zenodo.org/record/13848165/files/figure.png" pageId="13" pageNumber="14">Fig. 11</figureCitation>
). All records come from Nardouw and Cederberg Sandstone Fynbos sensu 
<bibRefCitation id="EFBA4B16FFB4FFED0E5B7AD9FE404494" author="Mucina, L. &amp; Rutherford, M. C." pageId="13" pageNumber="14" refId="ref11286" refString="Mucina, L. &amp; Rutherford, M. C. (2006) The Vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria, 807 pp." type="book" year="2006">Mucina and Rutherford (2006)</bibRefCitation>
, who describe the landscape as flat to gently sloping tableland dominated by rocky outcrops and gullies that are surrounded by deep sandy flats. Restios are common in moist areas, and small shrubs in drier sections. Breeding takes place atop large sandstone boulders and rocky outcrops in rock pools that form after the first winter rains. 
<taxonomicName id="4C2B4D64FFB4FFED0BC97B49FDDD4539" authorityName="Angus &amp; Telford &amp; Ping &amp; Conradie" authorityYear="2024" box="[218,531,1843,1869]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="cederbergense" status="sp. nov.">
<emphasis id="B95FEAF5FFB4FFED0BC97B49FDDD4539" box="[218,531,1843,1869]" italics="true" pageId="13" pageNumber="14">Cacosternum cederbergense</emphasis>
</taxonomicName>
<emphasis id="B95FEAF5FFB4FFED09097B49FDBD4539" bold="true" box="[538,627,1843,1869]" pageId="13" pageNumber="14">
<taxonomicNameLabel id="A26C578EFFB4FFED09097B49FDBD4539" box="[538,627,1843,1869]" pageId="13" pageNumber="14" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
appears to favour highly heterogeneous habitat with rocky and vegetated sections, often on scree covered slopes. During the day, adult and juvenile frogs have been found sheltering under rocks, or in small cracks in the pool edge. At night, adults may be observed resting entirely submerged on the pool bottom, or partially submerged clinging to the rock. If disturbed, they will dart downwards and walk along the substrate away from the perceived threat. When picked up they show similarities to the death-feigning that has been observed in 
<taxonomicName id="4C2B4D64FFB7FFEE0BA77CEDFEE342C4" authorityName="Hewitt" authorityYear="1925" box="[180,301,151,177]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB7FFEE0BA77CEDFEE342C4" box="[180,301,151,177]" italics="true" pageId="14" pageNumber="15">C. capense</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C2B4D64FFB7FFEE0A767CEDFDC042C4" box="[357,526,151,177]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="namaquense">
<emphasis id="B95FEAF5FFB7FFEE0A767CEDFDC042C4" box="[357,526,151,177]" italics="true" pageId="14" pageNumber="15">C. namaquense</emphasis>
</taxonomicName>
, whereby they tuck their limbs and head inwards, stiffening up in the process (De Villiers 1929; OA personal observation). On occasion, individuals may secrete a white substance from their large dorsolateral glands. As is the case with 
<taxonomicName id="4C2B4D64FFB7FFEE09417CA5FD05428C" authorityName="Hewitt" authorityYear="1925" box="[594,715,223,249]" class="Amphibia" family="Pyxicephalidae" genus="Cacosternum" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="capense">
<emphasis id="B95FEAF5FFB7FFEE09417CA5FD05428C" box="[594,715,223,249]" italics="true" pageId="14" pageNumber="15">C. capense</emphasis>
</taxonomicName>
, this secretion is likely toxic, acting as a defensive adaptation to avoid predation (
<bibRefCitation id="EFBA4B16FFB7FFEE0A437D79FE2F4369" author="Rebelo, A. D." box="[336,481,259,285]" pageId="14" pageNumber="15" pagination="27 - 28" refId="ref11719" refString="Rebelo, A. D. (2021) Cacosternum karooicum (Boycott, de Villiers, Scott, 2002). Karoo Caco. Defensive Secretion. African Herp News, 76, 27 - 28." type="journal article" year="2021">Rebelo 2021</bibRefCitation>
).
</paragraph>
<paragraph id="8B9436E7FFB7FFEE0BD47D5DFD9C4049" blockId="14.[151,1437,151,861]" pageId="14" pageNumber="15">
<emphasis id="B95FEAF5FFB7FFEE0BD47D5DFE454334" bold="true" box="[199,395,294,321]" pageId="14" pageNumber="15">Natural History.</emphasis>
Calls have only been heard on two occasions despite multiple night excursions. In 
<date id="FF951027FFB7FFEE0E377D5DFA564335" box="[1316,1432,295,321]" pageId="14" pageNumber="15" value="2023-06">June 2023</date>
, on the same day as a significant rainfall event (
<quantity id="4CD39B02FFB7FFEE09A57D30FCC24311" box="[694,780,330,357]" metricMagnitude="-2" metricUnit="m" metricValue="5.4" pageId="14" pageNumber="15" unit="mm" value="54.0">54 mm</quantity>
), only 
<specimenCount id="9D2DFD6EFFB7FFEE084F7D31FC134311" box="[860,989,331,357]" count="3" pageId="14" pageNumber="15" type="male">three males</specimenCount>
were heard calling at the 
<typeStatus id="54908845FFB7FFEE0E197D31FAF44311" box="[1290,1338,331,357]" pageId="14" pageNumber="15">type</typeStatus>
locality. Males began calling one hour after darkness, uttering soft creaks at long intervals. Call sites were limited to the pool edge, whereby males call half submerged holding onto the pool edge with their forelimbs. No territorial call has been heard, although two or more males have never been observed in the same pool. Pairs in amplexus have not been observed. A single egg clump (
<specimenCount id="9D2DFD6EFFB7FFEE09497DA1FD7D4381" box="[602,691,475,501]" count="30" pageId="14" pageNumber="15" type="egg">30 eggs</specimenCount>
) was found resting on the bottom of a pool away from the water’s edge. Individual eggs have been found scattered across the rock pool, presumably as a result of wave action caused by wind that dissembles the egg clump.
</paragraph>
</subSubSection>
<subSubSection id="C331656CFFB7FFEE0BD47E3CFB584129" pageId="14" pageNumber="15" type="distribution">
<paragraph id="8B9436E7FFB7FFEE0BD47E3CFB584129" blockId="14.[151,1437,151,861]" pageId="14" pageNumber="15">
<emphasis id="B95FEAF5FFB7FFEE0BD47E3CFEAF4014" bold="true" box="[199,353,582,608]" pageId="14" pageNumber="15">Distribution.</emphasis>
All records come from the northern Cederberg between 
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and 
<geoCoordinate id="EE1F5020FFB7FFEE0F487E3DFB654015" box="[1115,1195,583,609]" degrees="31.9" direction="south" orientation="latitude" pageId="14" pageNumber="15" precision="5555" value="-31.9">31.9º S</geoCoordinate>
at elevation of 
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to 
<specimenCount id="9D2DFD6EFFB7FFEE0BA77E10FECC40F1" box="[180,258,618,645]" count="540" pageId="14" pageNumber="15" type="generic">
<quantity id="4CD39B02FFB7FFEE0BA77E10FF3040F1" box="[180,254,618,645]" metricMagnitude="2" metricUnit="m" metricValue="5.4" pageId="14" pageNumber="15" unit="m" value="540.0">540 m</quantity>
.
</specimenCount>
The distribution of this species is likely controlled by its strict habitat requirements, namely the presence of rocky outcrops and the small rock pools that form atop them. The apparent disuse of habitat associated with rocky flats is reflected in the SDM for the species, which does not predict their occurrence on much of the plateau that extends from the Gifberg to Nieuwoudtville, in the lowlands of the Olifants River valley, or at elevations above 
<specimenCount id="9D2DFD6EFFB7FFEE0E617EADFF7A4161" count="800" pageId="14" pageNumber="15" type="generic">
<quantity id="4CD39B02FFB7FFEE0E617EADFF7E4161" metricMagnitude="2" metricUnit="m" metricValue="8.0" pageId="14" pageNumber="15" unit="m" value="800.0">800 m</quantity>
.
</specimenCount>
Rather, it suggests a distribution intermediate between lowland and montane environments, from the vicinity of Wupperthal in the south to the Doringrivier in the north. It also identifies two narrow linear strips of suitable habitat, one north of the Doringrivier and the second immediately west of the Olifants River valley.
</paragraph>
</subSubSection>
</treatment>
</document>