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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.979.56863" ID-GBIF-Dataset="d78779d5-178d-4ec3-83ca-1ff64cde3f14" ID-PMC="PMC7609483" ID-Pensoft-Pub="1313-2970-979-133" ID-Pensoft-UUID="C9CBE921FA5E570CB82AD1BCFE3620C2" ID-PubMed="33192134" ID-ZooBank="2043280A15914D51ACE3F9015F170890" ModsDocID="1313-2970-979-133" checkinTime="1603838534300" checkinUser="pensoft" docAuthor="Nielsen, Stuart V., Conradie, Werner, Ceriaco, Luis M. P., Bauer, Aaron M., Heinicke, Matthew P., Stanley, Edward L. &amp; Blackburn, David C." docDate="2020" docId="978CF7A21BAB528191337943C913494E" docLanguage="en" docName="ZooKeys 979: 133-160" docOrigin="ZooKeys 979" docSource="http://dx.doi.org/10.3897/zookeys.979.56863" docTitle="Breviceps ombelanonga Nielsen &amp; Conradie &amp; Ceríaco &amp; Bauer &amp; Heinicke &amp; Stanley &amp; Blackburn 2020, sp. nov." docType="treatment" docUuid="E3815018-4176-4073-92B8-E65274D354FB" docUuidSource="ZooBank" docVersion="5" id="C9CBE921FA5E570CB82AD1BCFE3620C2" lastPageNumber="133" masterDocId="C9CBE921FA5E570CB82AD1BCFE3620C2" masterDocTitle="A new species of Rain Frog (Brevicipitidae, Breviceps) endemic to Angola" masterLastPageNumber="160" masterPageNumber="133" pageNumber="133" updateTime="1668169647161" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>A new species of Rain Frog (Brevicipitidae, Breviceps) endemic to Angola</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Nielsen, Stuart V.</mods:namePart>
<mods:affiliation>University of Michigan-Dearborn, Dearborn, Michigan, USA &amp; Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA &amp; Marquette University, Milwaukee, Wisconsin, USA</mods:affiliation>
<mods:nameIdentifier type="orcid">https://orcid.org/0000-0002-3114-1469</mods:nameIdentifier>
<mods:nameIdentifier type="email">stunie@gmail.com</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Conradie, Werner</mods:namePart>
<mods:affiliation>Port Elizabeth Museum (Bayworld), Humewood, Republic of South Africa &amp; Nelson Mandela University, George, Republic of South Africa &amp; National Geographic Okavango Wilderness Project, Wild Bird Trust, Republic of South Africa</mods:affiliation>
<mods:nameIdentifier type="orcid">https://orcid.org/0000-0003-0805-9683</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ceriaco, Luis M. P.</mods:namePart>
<mods:affiliation>Museu de Historia Natural e da Ciencia da Universidade do Porto, Porto, Portugal &amp; Museu Nacional de Historia Natural e da Ciencia, Lisboa, Portugal</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bauer, Aaron M.</mods:namePart>
<mods:affiliation>Villanova University, Villanova, Pennsylvania, USA</mods:affiliation>
<mods:nameIdentifier type="orcid">https://orcid.org/0000-0001-6839-8025</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Heinicke, Matthew P.</mods:namePart>
<mods:affiliation>University of Michigan-Dearborn, Dearborn, Michigan, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Stanley, Edward L.</mods:namePart>
<mods:affiliation>Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Blackburn, David C.</mods:namePart>
<mods:affiliation>Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA</mods:affiliation>
<mods:nameIdentifier type="orcid">https://orcid.org/0000-0002-1810-9886</mods:nameIdentifier>
</mods:name>
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<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2020</mods:date>
<mods:detail type="volume">
<mods:number>979</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>133</mods:start>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.979.56863</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.979.56863</mods:identifier>
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<mods:identifier type="Pensoft-UUID">C9CBE921FA5E570CB82AD1BCFE3620C2</mods:identifier>
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<treatment ID-GBIF-Taxon="168650533" LSID="urn:lsid:zoobank.org:act:E3815018-4176-4073-92B8-E65274D354FB" httpUri="http://treatment.plazi.org/id/978CF7A21BAB528191337943C913494E" lastPageNumber="133" pageId="0" pageNumber="133">
<subSubSection pageId="0" pageNumber="133" type="nomenclature">
<paragraph pageId="0" pageNumber="133">
<taxonomicName LSID="978CF7A2-1BAB-5281-9133-7943C913494E" authority="Nielsen &amp; Conradie &amp; Ceríaco &amp; Bauer &amp; Heinicke &amp; Stanley &amp; Blackburn, 2020" authorityName="Nielsen &amp; Conradie &amp; Ceríaco &amp; Bauer &amp; Heinicke &amp; Stanley &amp; Blackburn" authorityYear="2020" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps ombelanonga" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="ombelanonga" status="sp. nov.">Breviceps ombelanonga</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="133">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Breviceps ombelanonga sp. nov. holotype male (UF Herp 187172): A in life photo B dorsal and ventral aspects C additional views of the holotype, including the left pes, frontal, right lateral, and left manus and mental. Scale bar: 10 mm. Photographs by J. Cavagnaro (A) and SVN (B, C)." figureDoi="10.3897/zookeys.979.56863.figure4" httpUri="https://binary.pensoft.net/fig/467832" pageId="0" pageNumber="133">Figs 4</figureCitation>
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Variation in color and pattern within living paratypes of B. ombelanonga sp. nov.: A, B sub-adult (of unknown sex) from Embala Seque (14 km N of Cassumbi village), Bie Province (MHNCUP _ ANF 0320) C juvenile male, Cuito River source lake, Moxico Province (PEM A 12537) D adult female, Cuando River source, Moxico Province (PEM A 12770) E adult male, Quembo River source lake, Moxico Province (PEM A 12787) F adult male, Cuanavale River source lake, Moxico Province (PEM A 12800). Photographs by LMPC (A, B) and WC (C-F)." figureDoi="10.3897/zookeys.979.56863.figure5" httpUri="https://binary.pensoft.net/fig/467833" pageId="0" pageNumber="133">, 5</figureCitation>
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Variation in color and pattern within preserved paratypes of B. ombelanonga sp. nov.: A an adult male from Kawa Camp Headquarters, Luanda Province (UF Herp 187173) B sub-adult (of unknown sex) from Embala Seque, Bie Province (MHNCUPANF 320) C adult male from Cuanavale River source lake, Moxico Province (PEM A 12800) D juvenile male from Cuito River source lake, Moxico Province (PEM A 12537) E adult male from Quembo River source lake, Moxico Province (PEM A 12787) F adult female from Cuando River source, Moxico Province (PEM A 12770); and G adult male from Quembo River source lake, Moxico Province (SAIAB 204537). Photographs by SVN (A), LMPC (B), and WC (C-G)." figureDoi="10.3897/zookeys.979.56863.figure6" httpUri="https://binary.pensoft.net/fig/467834" pageId="0" pageNumber="133">, 6</figureCitation>
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Photos of typical habitat of B. ombelanonga sp. nov.: A a view of the Kwanza River and bordering savannah, near the type locality, in Kissama National Park, Luanda Province B savannah near Embala Seque (14 km N of Cassumbi village), Bie Province C Cuanavale River source lake and associated miombo savannah woodland. Photographs by LMPC (A, B) and WC (C)." figureDoi="10.3897/zookeys.979.56863.figure7" httpUri="https://binary.pensoft.net/fig/467835" pageId="0" pageNumber="133">
, 7 Suggested common names: Angolan Rain Frog (English), Sapinho das Chuvas de Angola (
<normalizedToken originalValue="Português">Portugues</normalizedToken>
).
</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="chresonymy">
<paragraph pageId="0" pageNumber="133">Chresonymy.1</paragraph>
<paragraph pageId="0" pageNumber="133">
<taxonomicName baseAuthorityName="Linnaeus" baseAuthorityYear="1758" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps gibbosus" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="gibbosus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps gibbosus</emphasis>
</taxonomicName>
:
<bibRefCitation author="Bocage, JVB" journalOrPublisher="Jornal de Sciencias Mathematicas, Physicas e Naturaes" pageId="0" pageNumber="133" pagination="66 - 68" refId="B4" refString="Bocage, JVB, 1870. Description de un &quot;saurien&quot; nouveau de l'Afrique occidentale. Jornal de Sciencias Mathematicas, Physicas e Naturaes 3: 66 - 68" title="Description de un &quot; saurien &quot; nouveau de l'Afrique occidentale." volume="3" year="1870">Bocage (1870</bibRefCitation>
: 68).
</paragraph>
<paragraph pageId="0" pageNumber="133">
<taxonomicName baseAuthorityName="Linnaeus" baseAuthorityYear="1758" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps gibbosus" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="gibbosus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps gibbosus</emphasis>
</taxonomicName>
:
<bibRefCitation author="Bocage, JVB" journalOrPublisher="Jornal de Sciencias Mathematicas, Physicas e Naturaes" pageId="0" pageNumber="133" pagination="209 - 227" refId="B5" refString="Bocage, JVB, 1873. Melanges erpetologiques. II. Sur quelques reptiles et batraciens nouveaux, rares ou peu connus d'Afrique occidental. Jornal de Sciencias Mathematicas, Physicas e Naturaes 4: 209 - 227" title="Melanges erpetologiques. II. Sur quelques reptiles et batraciens nouveaux, rares ou peu connus d'Afrique occidental." volume="4" year="1873">Bocage (1873</bibRefCitation>
: 227).
</paragraph>
<paragraph pageId="0" pageNumber="133">
<taxonomicName class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps mossambicus" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps mossambicus</emphasis>
</taxonomicName>
:
<bibRefCitation author="Bocage, JVB" journalOrPublisher="Ministerio da Marinha e das Colonias, Lisbonne, Portugal" pageId="0" pageNumber="133" refId="B6" refString="Bocage, JVB, 1895. Herpetologie d'Angola et du Congo. Ministerio da Marinha e das Colonias, Lisbonne, Portugal" title="Herpetologie d'Angola et du Congo." year="1895">Bocage (1895</bibRefCitation>
: 182);
<bibRefCitation author="Parker, HW" journalOrPublisher="Trustees of the British Museum, London" pageId="0" pageNumber="133" refId="B45" refString="Parker, HW, 1934. A monograph of the frogs of the family Microhylidae. Trustees of the British Museum, London" title="A monograph of the frogs of the family Microhylidae." year="1934">Parker (1934</bibRefCitation>
: 194); Monard (1937: 29, 1938: 56);
<bibRefCitation author="Laurent, RF" journalOrPublisher="Publicacoes Culturais da Companhia de Diamantes de Angola" pageId="0" pageNumber="133" pagination="1 - 165" refId="B33" refString="Laurent, RF, 1964. Reptiles et amphibiens de l'Angola. (Troiseme contribution). Publicacoes Culturais da Companhia de Diamantes de Angola 67: 1 - 165" title="Reptiles et amphibiens de l'Angola. (Troiseme contribution)." volume="67" year="1964">Laurent (1964</bibRefCitation>
: 156);
<bibRefCitation author="Cei, JM" journalOrPublisher="Boletim da Sociedade Portuguesa de Ciencias Naturais" pageId="0" pageNumber="133" pagination="5 - 26" refId="B9" refString="Cei, JM, 1977. Chaves para uma identificacao preliminar dos batraquios anuros da R. P. de Angola. Boletim da Sociedade Portuguesa de Ciencias Naturais 17: 5 - 26" title="Chaves para uma identificacao preliminar dos batraquios anuros da R. P. de Angola." volume="17" year="1977">Cei (1977</bibRefCitation>
: 17, 18);
<bibRefCitation author="Ruas, C" journalOrPublisher="Garcia de Orta, Seria de Zoologia" pageId="0" pageNumber="133" pagination="19 - 41" refId="B55" refString="Ruas, C, 1996. Contribuicao para o conhecimento da fauna de batraquios de Angola. Parte I: Familias Pipidae, Bufonidae, Microhylidae, Ranidae, Hemisidae e Arthroleptidae. Garcia de Orta, Seria de Zoologia 21 (1): 19 - 41" title="Contribuicao para o conhecimento da fauna de batraquios de Angola. Parte I: Familias Pipidae, Bufonidae, Microhylidae, Ranidae, Hemisidae e Arthroleptidae." volume="21" year="1996">Ruas (1996</bibRefCitation>
: 23).
</paragraph>
<paragraph pageId="0" pageNumber="133">
<taxonomicName class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rana mossambicus" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">Rana mossambicus</emphasis>
</taxonomicName>
: Hellmich (1957: 30).
</paragraph>
<paragraph pageId="0" pageNumber="133">
<taxonomicName class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps</emphasis>
</taxonomicName>
&quot;
<emphasis italics="true" pageId="0" pageNumber="133">
<taxonomicName class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="mossambicus" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="mossambicus">mossambicus</taxonomicName>
-
<taxonomicName authorityName="Peters" authorityYear="1882" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="adspersus" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="adspersus">adspersus</taxonomicName>
</emphasis>
&quot; complex:
<bibRefCitation DOI="https://doi.org/10.1080/02541858.1982.11447782" author="Poynton, JC" journalOrPublisher="African Journal of Zoology" pageId="0" pageNumber="133" pagination="67 - 74" refId="B52" refString="Poynton, JC, 1982. On species pairs among Southern African amphibians. African Journal of Zoology 17: 67 - 74, DOI: https://doi.org/10.1080/02541858.1982.11447782" title="On species pairs among Southern African amphibians." url="https://doi.org/10.1080/02541858.1982.11447782" volume="17" year="1982">Poynton (1982</bibRefCitation>
: 67);
<bibRefCitation author="Ruas, C" journalOrPublisher="Garcia de Orta, Serie de Zoologia" pageId="0" pageNumber="133" pagination="139 - 146" refId="B56" refString="Ruas, C, 2002. Batraquios de Angola em coleccao no Centro de Zoologia. Garcia de Orta, Serie de Zoologia 24 (1-2): 139 - 146" title="Batraquios de Angola em coleccao no Centro de Zoologia." volume="24" year="2002">Ruas (2002</bibRefCitation>
: 142).
</paragraph>
<paragraph pageId="0" pageNumber="133">
<taxonomicName authorityName="Peters" authorityYear="1882" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps adspersus" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="adspersus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps adspersus</emphasis>
</taxonomicName>
[part]:
<bibRefCitation author="Poynton, JC" journalOrPublisher="Annals of the Natal Museum" pageId="0" pageNumber="133" pagination="503 - 553" refId="B53" refString="Poynton, JC, Broadley, DG, 1985. Amphibia Zambesiaca 1. Scolecomorphidae, Pipidae, Microhylidae, Hemisidae, Arthroleptidae. Annals of the Natal Museum 26: 503 - 553" title="Amphibia Zambesiaca 1. Scolecomorphidae, Pipidae, Microhylidae, Hemisidae, Arthroleptidae." volume="26" year="1985">Poynton and Broadley (1985</bibRefCitation>
: 52).
</paragraph>
<paragraph pageId="0" pageNumber="133">
<taxonomicName class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps</emphasis>
</taxonomicName>
sp.:
<bibRefCitation author="Marques, MP" journalOrPublisher="Proceedings of the California Academy of Sciences Series 4 65, Supplement II" pageId="0" pageNumber="133" refId="B35" refString="Marques, MP, Ceríaco, LMP, Blackburn, DC, Bauer, AM, 2018. Diversity and distribution of the amphibians and terrestrial reptiles of Angola: Atlas of historical and bibliographic records (1840-2017). Proceedings of the California Academy of Sciences Series 4 65, Supplement II" title="Diversity and distribution of the amphibians and terrestrial reptiles of Angola: Atlas of historical and bibliographic records (1840 - 2017)." year="2018">Marques et al. (2018</bibRefCitation>
: 81);
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.4746.1.1" author="Ceriaco, LMP" journalOrPublisher="Zootaxa" pageId="0" pageNumber="133" pagination="1 - 71" refId="B14" refString="Ceriaco, LMP, Agarwal, I, Marques, MP, Bauer, AM, 2020. A review of the genus Hemidactylus Goldfuss, 1820 (Squamata: Gekkonidae) from Angola, with the description of two new species. Zootaxa 4746: 1 - 71, DOI: https://doi.org/10.11646/zootaxa.4746.1.1" title="A review of the genus Hemidactylus Goldfuss, 1820 (Squamata: Gekkonidae) from Angola, with the description of two new species." url="https://doi.org/10.11646/zootaxa.4746.1.1" volume="4746" year="2020">
<normalizedToken originalValue="Ceríaco">Ceriaco</normalizedToken>
et al. (2020
</bibRefCitation>
: 63).
</paragraph>
<paragraph pageId="0" pageNumber="133">
<taxonomicName authorityName="Peters" authorityYear="1882" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps adspersus" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="adspersus">Breviceps cf. adspersus</taxonomicName>
:
<bibRefCitation DOI="https://doi.org/10.1007/978-3-030-03083-4_12" author="Baptista, N" journalOrPublisher="Cham, Switzerland, Springer Open. XVIII" pageId="0" pageNumber="133" refId="B3" refString="Baptista, N, Conradie, W, Vaz Pinto, P, Branch, WR, 2019. The amphibians of Angola: Early studies and the current state of knowledge. In: Huntley BJ, Russo V, Lages F, Ferrand de Almeida N (Eds) Biodiversity in Angola. Science &amp; Conservation: A Modern Synthesis: 243-281. Cham, Switzerland, Springer Open. XVIII, DOI: https://doi.org/10.1007/978-3-030-03083-4_12" title="The amphibians of Angola: Early studies and the current state of knowledge. In: Huntley BJ, Russo V, Lages F, Ferrand de Almeida N (Eds) Biodiversity in Angola. Science &amp; Conservation: A Modern Synthesis: 243 - 281." url="https://doi.org/10.1007/978-3-030-03083-4_12" year="2019">Baptista et al. (2019</bibRefCitation>
: 270).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="materials_examined">
<paragraph pageId="0" pageNumber="133">Material examined.</paragraph>
<paragraph pageId="0" pageNumber="133">
<emphasis bold="true" pageId="0" pageNumber="133">
<emphasis italics="true" pageId="0" pageNumber="133">Holotype</emphasis>
.
</emphasis>
UF Herp 187172 (field number MCZ A-36476), an adult male, Kawa Camp Headquarters, 1 km south of the Kwanza River, Kissama National Park (
<geoCoordinate degrees="9.183068" direction="south" orientation="latitude" precision="1" value="-9.183068">-9.183068</geoCoordinate>
,
<geoCoordinate degrees="13.369314" direction="east" orientation="longitude" precision="1" value="13.369314">13.369314</geoCoordinate>
, WGS-84, elevation 151 m above sea level), Luanda Province, Angola, collected by LMPC, Mariana P. Marques, Philip Pastor, and John Cavagnaro on 2 June 2016 at approx. 22:00.
<emphasis bold="true" italics="true" pageId="0" pageNumber="133">Paratypes</emphasis>
(5 males, 1 female, 1 sex unknown) UF Herp 187173 (field number MCZ A-36495), an adult male, Kawa Camp Headquarters, 1 km south of the Kwanza River, Kissama National Park (
<geoCoordinate degrees="9.183068" direction="south" orientation="latitude" precision="1" value="-9.183068">-9.183068</geoCoordinate>
,
<geoCoordinate degrees="13.369314" direction="east" orientation="longitude" precision="1" value="13.369314">13.369314</geoCoordinate>
, WGS-84, elevation 151 m above sea level), Luanda Province, Angola, collected by LMPC, Mariana P. Marques, Philip Pastor, and John Cavagnaro on 8 June 2016; MHNCUP/ANF 320 (field number AMB 11736), sub-adult (sex unknown), Embala Seque, 14 km N of Cassumbi village (
<geoCoordinate degrees="11.083845" direction="south" orientation="latitude" precision="1" value="-11.083845">-11.083845</geoCoordinate>
,
<geoCoordinate degrees="16.66741" direction="east" orientation="longitude" precision="1" value="16.66741">16.66741</geoCoordinate>
),
<normalizedToken originalValue="Bié">Bie</normalizedToken>
Province, Angola, collected by LMPC, Mariana P. Marques, and Adam Ferguson on 16 June 2019; PEM A12800 (field number WC-4591), adult male, Cuanavale River source lake (
<geoCoordinate degrees="13.089343" direction="south" orientation="latitude" precision="1" value="-13.089343">-13.089343</geoCoordinate>
,
<geoCoordinate degrees="18.89485" direction="east" orientation="longitude" precision="1" value="18.89485">18.89485</geoCoordinate>
, 1396 m above sea level), Moxico Province, Angola, collected by Werner Conradie and Luke Verburgt on 24 October 2016; PEM A12537 (field number WC-3924), juvenile male, Cuito River source lake (
<geoCoordinate degrees="12.68935" direction="south" orientation="latitude" precision="1" value="-12.68935">-12.68935</geoCoordinate>
,
<geoCoordinate degrees="18.36012" direction="east" orientation="longitude" precision="1" value="18.36012">18.36012</geoCoordinate>
, 1435 m above sea level), Moxico Province, Angola, collected by Werner Conradie and Ninda Baptista on 18 February 2016 October; PEM A12787 (field number WC-4756), adult male, Quembo River source lake (
<geoCoordinate degrees="13.13544" direction="south" orientation="latitude" precision="1" value="-13.13544">-13.13544</geoCoordinate>
,
<geoCoordinate degrees="19.04397" direction="east" orientation="longitude" precision="1" value="19.04397">19.04397</geoCoordinate>
, 1375 m above sea level), Moxico Province, Angola collected by Werner Conradie on 11 November 2016; PEM A12770 (field number WC-4827), adult female, Cuando River source (
<geoCoordinate degrees="13.00334" direction="south" orientation="latitude" precision="1" value="-13.00334">-13.00334</geoCoordinate>
,
<geoCoordinate degrees="19.13564" direction="east" orientation="longitude" precision="1" value="19.13564">19.13564</geoCoordinate>
, 1364 m above sea level), Moxico Province, Angola, collected by Werner Conradie and James Harvey on 22 November 2016; SAIAB 204537 (field number Ang16-RB12), adult male, Quembo River source lake (
<geoCoordinate degrees="13.13583" direction="south" orientation="latitude" precision="1" value="-13.13583">-13.13583</geoCoordinate>
,
<geoCoordinate degrees="19.04528" direction="east" orientation="longitude" precision="1" value="19.04528">19.04528</geoCoordinate>
), Moxico Province, Angola, collected by Roger Bills on 9 November 2016.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="diagnosis">
<paragraph pageId="0" pageNumber="133">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="133">
A species referable to
<taxonomicName class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps</emphasis>
</taxonomicName>
due to the following characteristics (
<bibRefCitation author="Poynton, JC" journalOrPublisher="Annals of the Natal Museum" pageId="0" pageNumber="133" pagination="1 - 334" refId="B51" refString="Poynton, JC, 1964. The Amphibia of Southern Africa: a faunal study. Annals of the Natal Museum 17: 1 - 334" title="The Amphibia of Southern Africa: a faunal study." volume="17" year="1964">Poynton 1964</bibRefCitation>
;
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.4300.2.3" author="Minter, LR" journalOrPublisher="Zootaxa" pageId="0" pageNumber="133" pagination="195 - 216" refId="B41" refString="Minter, LR, Netherlands, EC, Du, Preez LH, 2017. Uncovering a hidden diversity: two new species of Breviceps (Anura: Brevicipitidae) from northern KwaZulu-Natal, South Africa. Zootaxa 4300: 195 - 216, DOI: https://doi.org/10.11646/zootaxa.4300.2.3" title="Uncovering a hidden diversity: two new species of Breviceps (Anura: Brevicipitidae) from northern KwaZulu-Natal, South Africa." url="https://doi.org/10.11646/zootaxa.4300.2.3" volume="4300" year="2017">Minter et al. 2017</bibRefCitation>
): snout extremely abbreviated; mouth narrow and downturned near jaw joint; short limbs which, at rest in life, are held close to the body, not projecting beyond the body outline; digits I and V short or rudimentary; inner metatarsal tubercle well developed and notably longer than pedal digit III, narrowly separated from a prominent conical outer metatarsal tubercle. Additionally, the results of the molecular phylogenetic analyses support this species as embedded within the diversity of
<taxonomicName class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps</emphasis>
</taxonomicName>
, specifically within the
<taxonomicName lsidName="B. mossambicus" pageId="0" pageNumber="133" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">B. mossambicus</emphasis>
</taxonomicName>
group (Fig.
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Geographic distribution and phylogenetic relationships of Breviceps spp. included in this study. A Map of Angola and surrounding countries with all known Breviceps spp. sampling localities indicated on legend. The proposed distributions of B. adspersus and B. poweri (blue and red polygons, respectively) are from IUCN (2013 a, b), but should be considered tentative and worthy of reevaluation in light of recent studies. Furthermore, B. mossambicus is not mapped as no samples of certain identification occur west of Malawi (see Nielsen et al. 2018). B Multi-locus phylogeny of Breviceps, with select clades collapsed that are not relevant directly to the B. mossambicus group. The backbone is from the likelihood analysis, although Bayesian analyses produced a nearly identical topology (with any topological differences subtended by poor support). A black dot at each node indicates high support (e. g., Bayesian posterior probability&gt; 0.95, Maximum Likelihood bootstrap&gt; 90), while values below that cutoff are indicated for deep nodes only. Tapered bars to the right of voucher IDs indicate from which Angolan locality they were collected. C Median-joining networks for the two nuclear loci indicating a lack of shared haplotypes between candidate and recognized species. Hash marks indicate unique sequence differences between lineages, and black circles are hypothetical intermediate haplotypes." figureDoi="10.3897/zookeys.979.56863.figure1" httpUri="https://binary.pensoft.net/fig/467829" pageId="0" pageNumber="133">1B</figureCitation>
).
<taxonomicName authorityName="Nielsen &amp; Conradie &amp; Ceríaco &amp; Bauer &amp; Heinicke &amp; Stanley &amp; Blackburn" authorityYear="2020" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps ombelanonga" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="ombelanonga">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps ombelanonga</emphasis>
</taxonomicName>
can be diagnosed from other species of
<taxonomicName class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps</emphasis>
</taxonomicName>
and especially those in the
<taxonomicName lsidName="B. mossambicus" pageId="0" pageNumber="133" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">B. mossambicus</emphasis>
</taxonomicName>
group by the combination of lacking a visible tympanum, males having a single, uniformly dark gular patch that is continuous with the mask extending from the eye, having generally smooth dorsal skin, lacking many small tubercles on the palmar surfaces (as in, e.g.,
<taxonomicName lsidName="B. branchi" pageId="0" pageNumber="133" rank="species" species="branchi">
<emphasis italics="true" pageId="0" pageNumber="133">B. branchi</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="B. sylvestris" pageId="0" pageNumber="133" rank="species" species="sylvestris">
<emphasis italics="true" pageId="0" pageNumber="133">B. sylvestris</emphasis>
</taxonomicName>
; FitzSimons 1930;
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.3381.1.4" author="Channing, A" journalOrPublisher="Zootaxa" pageId="0" pageNumber="133" pagination="62 - 68" refId="B15" refString="Channing, A, 2012. A new species of Rain Frog from Namaqualand, South Africa (Anura: Brevicipitidae: Breviceps). Zootaxa 3381: 62 - 68, DOI: https://doi.org/10.11646/zootaxa.3381.1.4" title="A new species of Rain Frog from Namaqualand, South Africa (Anura: Brevicipitidae: Breviceps)." url="https://doi.org/10.11646/zootaxa.3381.1.4" volume="3381" year="2012">Channing 2012</bibRefCitation>
), lacking pale spots along flanks and a pale patch above the vent (both present in
<taxonomicName lsidName="B. poweri" pageId="0" pageNumber="133" rank="species" species="poweri">
<emphasis italics="true" pageId="0" pageNumber="133">B. poweri</emphasis>
</taxonomicName>
;
<bibRefCitation author="Parker, HW" journalOrPublisher="Trustees of the British Museum, London" pageId="0" pageNumber="133" refId="B45" refString="Parker, HW, 1934. A monograph of the frogs of the family Microhylidae. Trustees of the British Museum, London" title="A monograph of the frogs of the family Microhylidae." year="1934">Parker 1934</bibRefCitation>
;
<bibRefCitation author="Du Preez, L" journalOrPublisher="Struik Nature, Cape Town" pageId="0" pageNumber="133" refId="B21" refString="Du Preez, L, Carruthers, V, 2017. Frogs of Southern Africa. A Complete Guide. Struik Nature, Cape Town" title="Frogs of Southern Africa. A Complete Guide." year="2017">du Preez and Carruthers 2017</bibRefCitation>
), lacking short dark band below nares (as in
<taxonomicName lsidName="B. poweri" pageId="0" pageNumber="133" rank="species" species="poweri">
<emphasis italics="true" pageId="0" pageNumber="133">B. poweri</emphasis>
</taxonomicName>
;
<bibRefCitation author="Du Preez, L" journalOrPublisher="Struik Nature, Cape Town" pageId="0" pageNumber="133" refId="B21" refString="Du Preez, L, Carruthers, V, 2017. Frogs of Southern Africa. A Complete Guide. Struik Nature, Cape Town" title="Frogs of Southern Africa. A Complete Guide." year="2017">du Preez and Carruthers 2017</bibRefCitation>
), lacking confluent inner and outer metatarsal tubercles, having a relatively narrower head, shorter thigh, and shorter manual digit III (Fig.
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. PCA plots of ten size-corrected morphological characters from specimens of B. adspersus (n = 24), B. mossambicus (n = 9), B. poweri (n = 8), and the putative new Angolan species (n = 6) (Suppl. material 1: Table S 1), illustrating the PC 1 and PC 2 (A) and PC 2 and PC 3 (B) axes of variation, which combined represent ~ 70 % of the total variation (Table 4)." figureDoi="10.3897/zookeys.979.56863.figure2" httpUri="https://binary.pensoft.net/fig/467830" pageId="0" pageNumber="133">2</figureCitation>
; Table
<tableCitation captionStart="Table 4" captionStartId="T4" captionText="Table 4. Principal components analysis (PCA) loadings based on 10 size corrected morphological characters (head length, HL; head width, HW; eye diameter, ED; snout length, ES; interorbital distance, IOD; internarial distance, IND; thigh length, THL; crus length, CL; foot length, FL; and length of manual digit III, F 3 L)." httpUri="http://table.plazi.org/id/5E5D54EB9FBDD88A926AE7AA3D5ED9EE" pageId="0" pageNumber="133" tableUuid="5E5D54EB9FBDD88A926AE7AA3D5ED9EE">4</tableCitation>
), and having an advertisement call with both a longer interval between consecutive calls and a higher average dominant frequency (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Spectrograms and oscillograms showing a series of notes of the putatively novel Angolan Breviceps taxon compared to three closely related congeners." figureDoi="10.3897/zookeys.979.56863.figure3" httpUri="https://binary.pensoft.net/fig/467831" pageId="0" pageNumber="133">3</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="133">
The new species can be distinguished from other species of
<taxonomicName class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps</emphasis>
</taxonomicName>
occurring in the region by the following: pale paravertebral and dorsolateral patches are lacking, although a fine dorsolateral band may be present (versus no pale paravertebral or dorsolateral spots or patches in
<taxonomicName lsidName="B. mossambicus" pageId="0" pageNumber="133" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">B. mossambicus</emphasis>
</taxonomicName>
; series of both paravertebral and dorsolateral pale spots and patches present in
<taxonomicName lsidName="B. adspersus" pageId="0" pageNumber="133" rank="species" species="adspersus">
<emphasis italics="true" pageId="0" pageNumber="133">B. adspersus</emphasis>
</taxonomicName>
, a series of pale dorsolateral spots or patches present in
<taxonomicName lsidName="B. poweri" pageId="0" pageNumber="133" rank="species" species="poweri">
<emphasis italics="true" pageId="0" pageNumber="133">B. poweri</emphasis>
</taxonomicName>
); no conspicuous light patch above vent (present in
<taxonomicName lsidName="B. poweri" pageId="0" pageNumber="133" rank="species" species="poweri">
<emphasis italics="true" pageId="0" pageNumber="133">B. poweri</emphasis>
</taxonomicName>
); manual digit IV reaching approximately midway between the proximal and distal subarticular tubercles of manual digit III (versus nearly reaching distal subarticular tubercle of manual digit III in
<taxonomicName lsidName="B. mossambicus" pageId="0" pageNumber="133" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">B. mossambicus</emphasis>
</taxonomicName>
; not reaching or barely passing the proximal subarticular tubercle of the manual digit III in
<taxonomicName lsidName="B. poweri" pageId="0" pageNumber="133" rank="species" species="poweri">
<emphasis italics="true" pageId="0" pageNumber="133">B. poweri</emphasis>
</taxonomicName>
; similar to
<taxonomicName lsidName="B. adspersus" pageId="0" pageNumber="133" rank="species" species="adspersus">
<emphasis italics="true" pageId="0" pageNumber="133">B. adspersus</emphasis>
</taxonomicName>
in usually not reaching the distal subarticular of manual digit III); gular region with a single uniformly dark patch (versus a pair of marbled to freckled patches in
<taxonomicName lsidName="B. adspersus" pageId="0" pageNumber="133" rank="species" species="adspersus">
<emphasis italics="true" pageId="0" pageNumber="133">B. adspersus</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph pageId="0" pageNumber="133">
The advertisement call of the new species (Table
<tableCitation captionStart="Table 5" captionStartId="T5" captionText="Table 5. Comparison of the main variables for the advertisement calls of Breviceps ombelanonga sp. nov., Breviceps mossambicus, Breviceps adspersus and Breviceps poweri. Comparative data taken from Minter (1997, 2003)." httpUri="http://table.plazi.org/id/489D30395E35F00C8780E7CFC715E20F" pageId="0" pageNumber="133" tableUuid="489D30395E35F00C8780E7CFC715E20F">5</tableCitation>
, Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Spectrograms and oscillograms showing a series of notes of the putatively novel Angolan Breviceps taxon compared to three closely related congeners." figureDoi="10.3897/zookeys.979.56863.figure3" httpUri="https://binary.pensoft.net/fig/467831" pageId="0" pageNumber="133">3</figureCitation>
) can be differentiated from other potential Angolan congeners by its duration (0.175
<normalizedToken originalValue="±">+/-</normalizedToken>
0.083 s; shorter than in
<taxonomicName lsidName="B. adspersus" pageId="0" pageNumber="133" rank="species" species="adspersus">
<emphasis italics="true" pageId="0" pageNumber="133">B. adspersus</emphasis>
</taxonomicName>
0.196
<normalizedToken originalValue="±">+/-</normalizedToken>
0.047 s and
<taxonomicName lsidName="B. mossambicus" pageId="0" pageNumber="133" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">B. mossambicus</emphasis>
</taxonomicName>
, 0.500
<normalizedToken originalValue="±">+/-</normalizedToken>
0.070 s, and longer than in
<taxonomicName lsidName="B. poweri" pageId="0" pageNumber="133" rank="species" species="poweri">
<emphasis italics="true" pageId="0" pageNumber="133">B. poweri</emphasis>
</taxonomicName>
, 0.140
<normalizedToken originalValue="±">+/-</normalizedToken>
0.012 s), longer interval between consecutive calls (0.996
<normalizedToken originalValue="±">+/-</normalizedToken>
0.133 s;
<taxonomicName lsidName="B. adspersus" pageId="0" pageNumber="133" rank="species" species="adspersus">
<emphasis italics="true" pageId="0" pageNumber="133">B. adspersus</emphasis>
</taxonomicName>
, 0.745
<normalizedToken originalValue="±">+/-</normalizedToken>
0.636 s;
<taxonomicName lsidName="B. mossambicus" pageId="0" pageNumber="133" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">B. mossambicus</emphasis>
</taxonomicName>
, 0.710
<normalizedToken originalValue="±">+/-</normalizedToken>
0.168 s;
<taxonomicName lsidName="B. poweri" pageId="0" pageNumber="133" rank="species" species="poweri">
<emphasis italics="true" pageId="0" pageNumber="133">B. poweri</emphasis>
</taxonomicName>
, 0.743
<normalizedToken originalValue="±">+/-</normalizedToken>
0.166 s), and a higher dominant frequency (2156 Hz;
<taxonomicName lsidName="B. adspersus" pageId="0" pageNumber="133" rank="species" species="adspersus">
<emphasis italics="true" pageId="0" pageNumber="133">B. adspersus</emphasis>
</taxonomicName>
, 1742
<normalizedToken originalValue="±">+/-</normalizedToken>
100 Hz;
<taxonomicName lsidName="B. mossambicus" pageId="0" pageNumber="133" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">B. mossambicus</emphasis>
</taxonomicName>
, 1835
<normalizedToken originalValue="±">+/-</normalizedToken>
107 Hz;
<taxonomicName lsidName="B. poweri" pageId="0" pageNumber="133" rank="species" species="poweri">
<emphasis italics="true" pageId="0" pageNumber="133">B. poweri</emphasis>
</taxonomicName>
, 1728
<normalizedToken originalValue="±">+/-</normalizedToken>
83 Hz). The number of pulses per call (28-34) are similar to
<taxonomicName lsidName="B. adspersus" pageId="0" pageNumber="133" rank="species" species="adspersus">
<emphasis italics="true" pageId="0" pageNumber="133">B. adspersus</emphasis>
</taxonomicName>
(14-31),
<taxonomicName lsidName="B. mossambicus" pageId="0" pageNumber="133" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">B. mossambicus</emphasis>
</taxonomicName>
(7-31), and
<taxonomicName lsidName="B. poweri" pageId="0" pageNumber="133" rank="species" species="poweri">
<emphasis italics="true" pageId="0" pageNumber="133">B. poweri</emphasis>
</taxonomicName>
(10-74).
</paragraph>
<caption ID-Table-UUID="489D30395E35F00C8780E7CFC715E20F" httpUri="http://table.plazi.org/id/489D30395E35F00C8780E7CFC715E20F" pageId="0" pageNumber="133" start="Table 5" startId="T5">
<paragraph pageId="0" pageNumber="133">
<emphasis bold="true" pageId="0" pageNumber="133">Table 5.</emphasis>
Comparison of the main variables for the advertisement calls of
<taxonomicName authorityName="Nielsen &amp; Conradie &amp; Ceríaco &amp; Bauer &amp; Heinicke &amp; Stanley &amp; Blackburn" authorityYear="2020" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps ombelanonga" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="ombelanonga">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps ombelanonga</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps mossambicus" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps mossambicus</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="Peters" authorityYear="1882" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps adspersus" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="adspersus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps adspersus</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Parker" authorityYear="1934" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps poweri" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="poweri">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps poweri</emphasis>
</taxonomicName>
. Comparative data taken from
<bibRefCitation author="Minter, LR" journalOrPublisher="Annals of the Natal Museum" pageId="0" pageNumber="133" pagination="5 - 19" refId="B39" refString="Minter, LR, 1997. Advertisement call structure and morphology of Breviceps mossambicus Peters and B. poweri Parker (Anura: Microhylidae) from northern Mozambique. Annals of the Natal Museum 38: 5 - 19" title="Advertisement call structure and morphology of Breviceps mossambicus Peters and B. poweri Parker (Anura: Microhylidae) from northern Mozambique." volume="38" year="1997">Minter (1997</bibRefCitation>
,
<bibRefCitation DOI="https://doi.org/10.1080/21564574.2003.9635473" author="Minter, LR" journalOrPublisher="African Journal of Herpetology" pageId="0" pageNumber="133" pagination="9 - 21" refId="B40" refString="Minter, LR, 2003. Two new cryptic species of Breviceps (Anura: Microhylidae) from southern Africa. African Journal of Herpetology 52 (1): 9 - 21, DOI: https://doi.org/10.1080/21564574.2003.9635473" title="Two new cryptic species of Breviceps (Anura: Microhylidae) from southern Africa." url="https://doi.org/10.1080/21564574.2003.9635473" volume="52" year="2003">2003</bibRefCitation>
).
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="133">
<table pageId="0" pageNumber="133">
<tr pageId="0" pageNumber="133">
<th colspan="1" isEmpty="true" pageId="0" pageNumber="133" rowspan="1">-</th>
<th colspan="2" pageId="0" pageNumber="133" rowspan="1">
<taxonomicName lsidName="B. ombelanonga" pageId="0" pageNumber="133" rank="species" species="ombelanonga">
<emphasis italics="true" pageId="0" pageNumber="133">B. ombelanonga</emphasis>
</taxonomicName>
sp. nov.
</th>
<th colspan="2" pageId="0" pageNumber="133" rowspan="1">
<taxonomicName lsidName="B. adspersus" pageId="0" pageNumber="133" rank="species" species="adspersus">
<emphasis italics="true" pageId="0" pageNumber="133">B. adspersus</emphasis>
</taxonomicName>
</th>
<th colspan="2" pageId="0" pageNumber="133" rowspan="1">
<taxonomicName lsidName="B. mossambicus" pageId="0" pageNumber="133" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">B. mossambicus</emphasis>
</taxonomicName>
</th>
<th colspan="2" pageId="0" pageNumber="133" rowspan="1">
<taxonomicName lsidName="B. poweri" pageId="0" pageNumber="133" rank="species" species="poweri">
<emphasis italics="true" pageId="0" pageNumber="133">B. poweri</emphasis>
</taxonomicName>
</th>
</tr>
<tr pageId="0" pageNumber="133">
<td colspan="1" isEmpty="true" pageId="0" pageNumber="133" rowspan="1">-</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
avg
<normalizedToken originalValue="±">+/-</normalizedToken>
sd
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">range</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
avg
<normalizedToken originalValue="±">+/-</normalizedToken>
sd
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">range</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
avg
<normalizedToken originalValue="±">+/-</normalizedToken>
sd
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">range</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
avg
<normalizedToken originalValue="±">+/-</normalizedToken>
sd
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">range</td>
</tr>
<tr pageId="0" pageNumber="133">
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
<emphasis bold="true" pageId="0" pageNumber="133">Call duration (s)</emphasis>
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
0.175
<normalizedToken originalValue="±">+/-</normalizedToken>
0.083
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">0.064-0.342</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
0.196
<normalizedToken originalValue="±">+/-</normalizedToken>
0.047
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">0.077-0.293</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
0.500
<normalizedToken originalValue="±">+/-</normalizedToken>
0.070
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">0.036-0.079</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
0.140
<normalizedToken originalValue="±">+/-</normalizedToken>
0.012
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">0.111-0.160</td>
</tr>
<tr pageId="0" pageNumber="133">
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
<emphasis bold="true" pageId="0" pageNumber="133">Call interval (s)</emphasis>
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
0.996
<normalizedToken originalValue="±">+/-</normalizedToken>
0.133
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">0.742-1.190</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
0.745
<normalizedToken originalValue="±">+/-</normalizedToken>
0.636
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">0.363-0.745*</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
0.710
<normalizedToken originalValue="±">+/-</normalizedToken>
0.168
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">0.396-1.17</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
0.743
<normalizedToken originalValue="±">+/-</normalizedToken>
0.166
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">0.500-1.100</td>
</tr>
<tr pageId="0" pageNumber="133">
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
<emphasis bold="true" pageId="0" pageNumber="133">No. of pulses/call</emphasis>
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
30
<normalizedToken originalValue="±">+/-</normalizedToken>
2.6
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">28-34</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
23
<normalizedToken originalValue="±">+/-</normalizedToken>
3.3
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">14-31</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
9
<normalizedToken originalValue="±">+/-</normalizedToken>
1.2
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">7-13</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
30
<normalizedToken originalValue="±">+/-</normalizedToken>
16.3
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">10-74</td>
</tr>
<tr pageId="0" pageNumber="133">
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
<emphasis bold="true" pageId="0" pageNumber="133">Dominant frequency (Hz)</emphasis>
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">2156</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">na</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
1742
<normalizedToken originalValue="±">+/-</normalizedToken>
100
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">1482-2179</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
1835
<normalizedToken originalValue="±">+/-</normalizedToken>
107
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">1600-2193</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">
1728
<normalizedToken originalValue="±">+/-</normalizedToken>
83
</td>
<td colspan="1" pageId="0" pageNumber="133" rowspan="1">1557-1903</td>
</tr>
</table>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="description">
<paragraph pageId="0" pageNumber="133">Description of the holotype.</paragraph>
<paragraph pageId="0" pageNumber="133">
Adult male (SUL 30.5 mm), with globular body and well-developed short limbs with medialmost and lateralmost digits reduced (Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Breviceps ombelanonga sp. nov. holotype male (UF Herp 187172): A in life photo B dorsal and ventral aspects C additional views of the holotype, including the left pes, frontal, right lateral, and left manus and mental. Scale bar: 10 mm. Photographs by J. Cavagnaro (A) and SVN (B, C)." figureDoi="10.3897/zookeys.979.56863.figure4" httpUri="https://binary.pensoft.net/fig/467832" pageId="0" pageNumber="133">4</figureCitation>
; Table
<tableCitation captionStart="Table 3" captionStartId="T3" captionText="Table 3. Measurements (mm) of type series." httpUri="http://table.plazi.org/id/50583A53262E79D975ABC6B44E1C423F" pageId="0" pageNumber="133" tableUuid="50583A53262E79D975ABC6B44E1C423F">3</tableCitation>
); snout abbreviated, protruding and angular in lateral profile, blunt and rectangular in dorsal view; eyes projecting beyond profile of head in both dorsal and ventral views; pupils horizontally elliptical; nares small oval slits, directed horizontally and visible in dorsal and lateral views; mouth narrow and directed ventrally near jaw joint; choana largely obscured by maxillae in ventral view; well-developed gland at midline of palate between choana; tongue ovoid and filling floor of mouth, and lacking median papilla; single medial bony point on lower jaw at symphysis; tympana not distinguishable; teeth absent on premaxilla, maxilla, and vomer.
</paragraph>
<caption doi="10.3897/zookeys.979.56863.figure4" httpUri="https://binary.pensoft.net/fig/467832" pageId="0" pageNumber="133" start="Figure 4" startId="F4">
<paragraph pageId="0" pageNumber="133">
<emphasis bold="true" pageId="0" pageNumber="133">Figure 4.</emphasis>
<taxonomicName authorityName="Nielsen &amp; Conradie &amp; Ceríaco &amp; Bauer &amp; Heinicke &amp; Stanley &amp; Blackburn" authorityYear="2020" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps ombelanonga" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="ombelanonga">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps ombelanonga</emphasis>
</taxonomicName>
sp. nov. holotype male (UF Herp 187172):
<emphasis bold="true" pageId="0" pageNumber="133">A</emphasis>
in life photo
<emphasis bold="true" pageId="0" pageNumber="133">B</emphasis>
dorsal and ventral aspects
<emphasis bold="true" pageId="0" pageNumber="133">C</emphasis>
additional views of the holotype, including the left pes, frontal, right lateral, and left manus and mental. Scale bar: 10 mm. Photographs by J. Cavagnaro (
<emphasis bold="true" pageId="0" pageNumber="133">A</emphasis>
) and SVN (
<emphasis bold="true" pageId="0" pageNumber="133">B, C</emphasis>
).
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="133">Skin of dorsum and head smooth, and weakly glandular with irregular folds; skin of ventrum smooth; skin folds overlying vent creating triangular shape.</paragraph>
<paragraph pageId="0" pageNumber="133">Limbs short with digits I and V short or rudimentary; webbing absent on manus and pes; nuptial pads absent and adhesive glands not discernable; relative manual digit lengths when adpressed: III&gt;II&gt;I&gt;IV; only tip of first pedal digit extending beyond fleshy webbing and sole; fourth (outer) manual digit reaches midway between the large tubercle at metacarpophalangeal joint and subarticular tubercle at most proximal interphalangeal joint; finger tips conical, not expanded; several small globular palmar tubercles; single subarticular tubercles present on pedal digits II, III, and IV; pedal digit V very short, falling short of most proximal subarticular tubercle of pedal digit IV; well-developed (though not keratinized) inner metatarsal tubercle visibly longer than pedal digit III, separated from conical outer metatarsal tubercle by deep cleft.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="coloration">
<paragraph pageId="0" pageNumber="133">Coloration.</paragraph>
<paragraph pageId="0" pageNumber="133">
In life, dorsum of body mottled dark brown on pale tan base, transitioning to golden yellow on the lateral aspects, before stark transition to solid dark brown flanks with a dark boundary becoming paler ventrally (Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Breviceps ombelanonga sp. nov. holotype male (UF Herp 187172): A in life photo B dorsal and ventral aspects C additional views of the holotype, including the left pes, frontal, right lateral, and left manus and mental. Scale bar: 10 mm. Photographs by J. Cavagnaro (A) and SVN (B, C)." figureDoi="10.3897/zookeys.979.56863.figure4" httpUri="https://binary.pensoft.net/fig/467832" pageId="0" pageNumber="133">4</figureCitation>
); limbs dark grayish brown dorsally; plantar and palmar surfaces pale grayish brown; subarticular, palmar, and inner and outer metatarsal tubercles pale gray; posterior dorsum dark gray-brown with scattered pale gray spots; bold facial mask composed of broad dark brown stripe running obliquely downwards, from margin of lower eyelid towards base of arm (but not attaining it) and joining dorsolateral aspect of gular patch, giving appearance of a large dark bib; region below nares generally same coloration as dorsal and lateral rostrum, and not more darkly pigmented; lower eyelid with white opaque patch at anterior margin; margins of mouth and lateral angle of mouth off-white to cream; gular patch uniformly dark anteriorly, becoming mottled posteriorly and merging with ventral coloration; pectoral region and ventrum creamy pale gray with scattered punctate gray dots sometimes coalescing into larger spots in the gular region and laterally; iris bright orange, scattered with dark brown flecks (dark brown in preservative), with black pupil (pale gray in preservative; no mid-vertebral line; faint pale line extending across posterior hindlimbs extending between heels.
</paragraph>
<paragraph pageId="0" pageNumber="133">
In preservative, coloration is largely similar but more muted and overall darker (Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Breviceps ombelanonga sp. nov. holotype male (UF Herp 187172): A in life photo B dorsal and ventral aspects C additional views of the holotype, including the left pes, frontal, right lateral, and left manus and mental. Scale bar: 10 mm. Photographs by J. Cavagnaro (A) and SVN (B, C)." figureDoi="10.3897/zookeys.979.56863.figure4" httpUri="https://binary.pensoft.net/fig/467832" pageId="0" pageNumber="133">4</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="description">
<paragraph pageId="0" pageNumber="133">Measurements.</paragraph>
<paragraph pageId="0" pageNumber="133">
Measurements of the type series are shown in Table
<tableCitation captionStart="Table 3" captionStartId="T3" captionText="Table 3. Measurements (mm) of type series." httpUri="http://table.plazi.org/id/50583A53262E79D975ABC6B44E1C423F" pageId="0" pageNumber="133" tableUuid="50583A53262E79D975ABC6B44E1C423F">3</tableCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="variation">
<paragraph pageId="0" pageNumber="133">Variations.</paragraph>
<paragraph pageId="0" pageNumber="133">
All specimens resemble the holotype in the absence of a visible tympanum, and skin that is densely granular dorsally and laterally and smooth ventrally (Figs
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Variation in color and pattern within living paratypes of B. ombelanonga sp. nov.: A, B sub-adult (of unknown sex) from Embala Seque (14 km N of Cassumbi village), Bie Province (MHNCUP _ ANF 0320) C juvenile male, Cuito River source lake, Moxico Province (PEM A 12537) D adult female, Cuando River source, Moxico Province (PEM A 12770) E adult male, Quembo River source lake, Moxico Province (PEM A 12787) F adult male, Cuanavale River source lake, Moxico Province (PEM A 12800). Photographs by LMPC (A, B) and WC (C-F)." figureDoi="10.3897/zookeys.979.56863.figure5" httpUri="https://binary.pensoft.net/fig/467833" pageId="0" pageNumber="133">5</figureCitation>
-
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Variation in color and pattern within preserved paratypes of B. ombelanonga sp. nov.: A an adult male from Kawa Camp Headquarters, Luanda Province (UF Herp 187173) B sub-adult (of unknown sex) from Embala Seque, Bie Province (MHNCUPANF 320) C adult male from Cuanavale River source lake, Moxico Province (PEM A 12800) D juvenile male from Cuito River source lake, Moxico Province (PEM A 12537) E adult male from Quembo River source lake, Moxico Province (PEM A 12787) F adult female from Cuando River source, Moxico Province (PEM A 12770); and G adult male from Quembo River source lake, Moxico Province (SAIAB 204537). Photographs by SVN (A), LMPC (B), and WC (C-G)." figureDoi="10.3897/zookeys.979.56863.figure6" httpUri="https://binary.pensoft.net/fig/467834" pageId="0" pageNumber="133">6</figureCitation>
). The distal tip of manual digit IV reaches well past the proximal subarticular tubercle of manual digit III in all specimens. PEM A12770 have both manual digit II and III proximal subarticular tubercles divided. Inner and outer metatarsal tubercles not separated by a deep cleft in paratypes PEM A12800, PEM A12537, PEM A12787, PEM A12770 and SAIAB 204537.
</paragraph>
<caption doi="10.3897/zookeys.979.56863.figure5" httpUri="https://binary.pensoft.net/fig/467833" pageId="0" pageNumber="133" start="Figure 5" startId="F5">
<paragraph pageId="0" pageNumber="133">
<emphasis bold="true" pageId="0" pageNumber="133">Figure 5.</emphasis>
Variation in color and pattern within living paratypes of
<taxonomicName lsidName="B. ombelanonga" pageId="0" pageNumber="133" rank="species" species="ombelanonga">
<emphasis italics="true" pageId="0" pageNumber="133">B. ombelanonga</emphasis>
</taxonomicName>
sp. nov.:
<emphasis bold="true" pageId="0" pageNumber="133">A, B</emphasis>
sub-adult (of unknown sex) from Embala Seque (14 km N of Cassumbi village),
<normalizedToken originalValue="Bié">Bie</normalizedToken>
Province (MHNCUP_ANF 0320)
<emphasis bold="true" pageId="0" pageNumber="133">C</emphasis>
juvenile male, Cuito River source lake, Moxico Province (PEM A12537)
<emphasis bold="true" pageId="0" pageNumber="133">D</emphasis>
adult female, Cuando River source, Moxico Province (PEM A12770)
<emphasis bold="true" pageId="0" pageNumber="133">E</emphasis>
adult male, Quembo River source lake, Moxico Province (PEM A12787)
<emphasis bold="true" pageId="0" pageNumber="133">F</emphasis>
adult male, Cuanavale River source lake, Moxico Province (PEM A12800). Photographs by LMPC (
<emphasis bold="true" pageId="0" pageNumber="133">A, B</emphasis>
) and WC (
<emphasis bold="true" pageId="0" pageNumber="133">C-F</emphasis>
).
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="133">Color and pattern in UF Herp 187173 is very similar to the holotype. Dorsum gray with scattered black spots (MHNCUP/ANF 320); red with scattered black blotches in two specimens (PEM A12537 and PEM A12770), dark brown to black with red spots and markings (PEM A12787 and PEM A12800), light brown with red spots and darker black blotches (SAIAB 204537). Interocular bar visible in all paratypes, except PEM A12537, PEM A12770 and SAIAB 204537. Light dorsolateral patches present in PEM A12878, absent in PEM A12770, dark black band present in PEM A12537. Mid-vertebral line present in most paratypes, but very faint in PEM A12537 and PEM A12770, and absent in SAIAB 204537 and MHNCUP/ANF 320. Heel-to-heel line present in all specimens, but faint in UF Herp 187173, PEM A12537, and PEM A12770. A broad, black stripe runs obliquely downwards from margin of lower eyelid towards base of arm, not reaching the shoulder in all specimens. Dark orbital band partly reaching the gular patch in all specimens, falling short in PEM A12770 (female). Anterior to the orbital bar, a broad white stripe runs down to angle of mouth and onto upper and lower lips in all individuals. Gular patch uniform dark brown to black in all paratypes, except PEM A12537 in which it is dark brown with scattered darker blotches. Pectoral region white, with scattered spots in all specimens. Ventrum white with scattered darker spots in all paratypes.</paragraph>
<caption doi="10.3897/zookeys.979.56863.figure6" httpUri="https://binary.pensoft.net/fig/467834" pageId="0" pageNumber="133" start="Figure 6" startId="F6">
<paragraph pageId="0" pageNumber="133">
<emphasis bold="true" pageId="0" pageNumber="133">Figure 6.</emphasis>
Variation in color and pattern within preserved paratypes of
<taxonomicName lsidName="B. ombelanonga" pageId="0" pageNumber="133" rank="species" species="ombelanonga">
<emphasis italics="true" pageId="0" pageNumber="133">B. ombelanonga</emphasis>
</taxonomicName>
sp. nov.:
<emphasis bold="true" pageId="0" pageNumber="133">A</emphasis>
an adult male from Kawa Camp Headquarters, Luanda Province (UF Herp 187173)
<emphasis bold="true" pageId="0" pageNumber="133">B</emphasis>
sub-adult (of unknown sex) from Embala Seque,
<normalizedToken originalValue="Bié">Bie</normalizedToken>
Province (MHNCUPANF 320)
<emphasis bold="true" pageId="0" pageNumber="133">C</emphasis>
adult male from Cuanavale River source lake, Moxico Province (PEM A12800)
<emphasis bold="true" pageId="0" pageNumber="133">D</emphasis>
juvenile male from Cuito River source lake, Moxico Province (PEM A12537)
<emphasis bold="true" pageId="0" pageNumber="133">E</emphasis>
adult male from Quembo River source lake, Moxico Province (PEM A12787)
<emphasis bold="true" pageId="0" pageNumber="133">F</emphasis>
adult female from Cuando River source, Moxico Province (PEM A12770); and
<emphasis bold="true" pageId="0" pageNumber="133">G</emphasis>
adult male from Quembo River source lake, Moxico Province (SAIAB 204537). Photographs by SVN (
<emphasis bold="true" pageId="0" pageNumber="133">A</emphasis>
), LMPC (
<emphasis bold="true" pageId="0" pageNumber="133">B</emphasis>
), and WC (
<emphasis bold="true" pageId="0" pageNumber="133">C-G</emphasis>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="advertisement call">
<paragraph pageId="0" pageNumber="133">Advertisement call.</paragraph>
<paragraph pageId="0" pageNumber="133">
The following call description is based on a recording of a paratype male (PEM A12800) from the source lake of the Cuanavale River recorded on 24 October 2016 at 8:50 in the morning. Ambient temperature was not recorded. Frogs began calling during the daytime following heavy rains, and stopped after sunset. Call sites were among leaf litter in dense miombo woodland. The call can be described as a short whistle with a call duration of 0.064-0.342 seconds and call interval of 0.742-1.190 seconds. Each call consists of about 28-34 pulses and a dominant frequency of 2156 Hz (Table
<tableCitation captionStart="Table 4" captionStartId="T4" captionText="Table 4. Principal components analysis (PCA) loadings based on 10 size corrected morphological characters (head length, HL; head width, HW; eye diameter, ED; snout length, ES; interorbital distance, IOD; internarial distance, IND; thigh length, THL; crus length, CL; foot length, FL; and length of manual digit III, F 3 L)." httpUri="http://table.plazi.org/id/5E5D54EB9FBDD88A926AE7AA3D5ED9EE" pageId="0" pageNumber="133" tableUuid="5E5D54EB9FBDD88A926AE7AA3D5ED9EE">4</tableCitation>
, Fig.
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. PCA plots of ten size-corrected morphological characters from specimens of B. adspersus (n = 24), B. mossambicus (n = 9), B. poweri (n = 8), and the putative new Angolan species (n = 6) (Suppl. material 1: Table S 1), illustrating the PC 1 and PC 2 (A) and PC 2 and PC 3 (B) axes of variation, which combined represent ~ 70 % of the total variation (Table 4)." figureDoi="10.3897/zookeys.979.56863.figure2" httpUri="https://binary.pensoft.net/fig/467830" pageId="0" pageNumber="133">2</figureCitation>
). The small number of calls from a geographically restricted sample does not allow for further statistical analysis.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="distribution">
<paragraph pageId="0" pageNumber="133">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="133">
Based on our phylogenetic analysis, this species is currently confirmed from three widely separated localities and elevations ranging from near sea level to&gt; 1400 m: i) Kissama National Park, on the outskirts of
<normalizedToken originalValue="Angolas">Angola's</normalizedToken>
capital city, Luanda, in coastal western Angola (Luanda Province); ii) central Angola (
<normalizedToken originalValue="Bié">Bie</normalizedToken>
Province); and iii) the source of the Cuanavale, Cuito, Cuando and Quembo rivers (Moxico Province). The identity of other known Angolan localities for
<taxonomicName class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps</emphasis>
</taxonomicName>
(black diamonds) remain uncertain without additional sampling and genetic data (Fig.
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Geographic distribution and phylogenetic relationships of Breviceps spp. included in this study. A Map of Angola and surrounding countries with all known Breviceps spp. sampling localities indicated on legend. The proposed distributions of B. adspersus and B. poweri (blue and red polygons, respectively) are from IUCN (2013 a, b), but should be considered tentative and worthy of reevaluation in light of recent studies. Furthermore, B. mossambicus is not mapped as no samples of certain identification occur west of Malawi (see Nielsen et al. 2018). B Multi-locus phylogeny of Breviceps, with select clades collapsed that are not relevant directly to the B. mossambicus group. The backbone is from the likelihood analysis, although Bayesian analyses produced a nearly identical topology (with any topological differences subtended by poor support). A black dot at each node indicates high support (e. g., Bayesian posterior probability&gt; 0.95, Maximum Likelihood bootstrap&gt; 90), while values below that cutoff are indicated for deep nodes only. Tapered bars to the right of voucher IDs indicate from which Angolan locality they were collected. C Median-joining networks for the two nuclear loci indicating a lack of shared haplotypes between candidate and recognized species. Hash marks indicate unique sequence differences between lineages, and black circles are hypothetical intermediate haplotypes." figureDoi="10.3897/zookeys.979.56863.figure1" httpUri="https://binary.pensoft.net/fig/467829" pageId="0" pageNumber="133">1</figureCitation>
, Appendix 1; see
<bibRefCitation author="Marques, MP" journalOrPublisher="Proceedings of the California Academy of Sciences Series 4 65, Supplement II" pageId="0" pageNumber="133" refId="B35" refString="Marques, MP, Ceríaco, LMP, Blackburn, DC, Bauer, AM, 2018. Diversity and distribution of the amphibians and terrestrial reptiles of Angola: Atlas of historical and bibliographic records (1840-2017). Proceedings of the California Academy of Sciences Series 4 65, Supplement II" title="Diversity and distribution of the amphibians and terrestrial reptiles of Angola: Atlas of historical and bibliographic records (1840 - 2017)." year="2018">Marques et al. 2018</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="genetic divergence">
<paragraph pageId="0" pageNumber="133">Genetic divergence.</paragraph>
<paragraph pageId="0" pageNumber="133">
<taxonomicName authorityName="Nielsen &amp; Conradie &amp; Ceríaco &amp; Bauer &amp; Heinicke &amp; Stanley &amp; Blackburn" authorityYear="2020" class="Amphibia" family="Brevicipitidae" genus="Breviceps" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Breviceps ombelanonga" order="Anura" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="ombelanonga">
<emphasis italics="true" pageId="0" pageNumber="133">Breviceps ombelanonga</emphasis>
</taxonomicName>
differs from other species within the
<taxonomicName lsidName="B. mossambicus" pageId="0" pageNumber="133" rank="species" species="mossambicus">
<emphasis italics="true" pageId="0" pageNumber="133">B. mossambicus</emphasis>
</taxonomicName>
group by net uncorrected mitochondrial
<emphasis italics="true" pageId="0" pageNumber="133">p</emphasis>
-distances of at least 9% (12S) and 11% (16S; Table
<tableCitation captionStart="Table 2" captionStartId="T2" captionText="Table 2. Uncorrected mean pairwise 12 S and 16 S mitochondrial sequence differences between ingroup Breviceps sequence pairs (above / below the diagonal, respectively) and within species (along the diagonal) conducted in MEGA." httpUri="http://table.plazi.org/id/463DF8D27796180B7D7A76622BE22FEC" pageId="0" pageNumber="133" tableUuid="463DF8D27796180B7D7A76622BE22FEC">2</tableCitation>
), as well as unique nuclear haplotypes for both RAG1 and BDNF (Fig.
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Geographic distribution and phylogenetic relationships of Breviceps spp. included in this study. A Map of Angola and surrounding countries with all known Breviceps spp. sampling localities indicated on legend. The proposed distributions of B. adspersus and B. poweri (blue and red polygons, respectively) are from IUCN (2013 a, b), but should be considered tentative and worthy of reevaluation in light of recent studies. Furthermore, B. mossambicus is not mapped as no samples of certain identification occur west of Malawi (see Nielsen et al. 2018). B Multi-locus phylogeny of Breviceps, with select clades collapsed that are not relevant directly to the B. mossambicus group. The backbone is from the likelihood analysis, although Bayesian analyses produced a nearly identical topology (with any topological differences subtended by poor support). A black dot at each node indicates high support (e. g., Bayesian posterior probability&gt; 0.95, Maximum Likelihood bootstrap&gt; 90), while values below that cutoff are indicated for deep nodes only. Tapered bars to the right of voucher IDs indicate from which Angolan locality they were collected. C Median-joining networks for the two nuclear loci indicating a lack of shared haplotypes between candidate and recognized species. Hash marks indicate unique sequence differences between lineages, and black circles are hypothetical intermediate haplotypes." figureDoi="10.3897/zookeys.979.56863.figure1" httpUri="https://binary.pensoft.net/fig/467829" pageId="0" pageNumber="133">1B</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="habitat">
<paragraph pageId="0" pageNumber="133">Habitat and natural history notes.</paragraph>
<paragraph pageId="0" pageNumber="133">
The preferred habitat for
<taxonomicName lsidName="B. ombelanonga" pageId="0" pageNumber="133" rank="species" species="ombelanonga">
<emphasis italics="true" pageId="0" pageNumber="133">B. ombelanonga</emphasis>
</taxonomicName>
ranges from typical western Angolan savannah, with sandy soils and vegetation dominated by
<taxonomicName class="Magnoliopsida" family="Malvaceae" genus="Adansonia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Adansonia digitata" order="Malvales" pageId="0" pageNumber="133" phylum="Tracheophyta" rank="species" species="digitata">
<emphasis italics="true" pageId="0" pageNumber="133">Adansonia digitata</emphasis>
</taxonomicName>
,
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Euphorbia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Euphorbia conspicua" order="Malpighiales" pageId="0" pageNumber="133" phylum="Tracheophyta" rank="species" species="conspicua">
<emphasis italics="true" pageId="0" pageNumber="133">Euphorbia conspicua</emphasis>
</taxonomicName>
,
<taxonomicName class="Magnoliopsida" family="Fabaceae" genus="Acacia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Acacia welwitschii" order="Fabales" pageId="0" pageNumber="133" phylum="Tracheophyta" rank="species" species="welwitschii">
<emphasis italics="true" pageId="0" pageNumber="133">Acacia welwitschii</emphasis>
</taxonomicName>
and
<taxonomicName class="Magnoliopsida" family="Combretaceae" genus="Combretum" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Combretum" order="Myrtales" pageId="0" pageNumber="133" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="133">Combretum</emphasis>
</taxonomicName>
sp., together with a good grass coverage (
<bibRefCitation author="Grandvaux-Barbosa, LA" journalOrPublisher="Instituto de Investigacao Cientifica de Angola, Luanda" pageId="0" pageNumber="133" refId="B24" refString="Grandvaux-Barbosa, LA, 1970. Carta Fitogeografica de Angola. Instituto de Investigacao Cientifica de Angola, Luanda" title="Carta Fitogeografica de Angola." year="1970">Grandvaux-Barbosa 1970</bibRefCitation>
), to dense Angolan wet miombo woodland in the east (Fig.
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Photos of typical habitat of B. ombelanonga sp. nov.: A a view of the Kwanza River and bordering savannah, near the type locality, in Kissama National Park, Luanda Province B savannah near Embala Seque (14 km N of Cassumbi village), Bie Province C Cuanavale River source lake and associated miombo savannah woodland. Photographs by LMPC (A, B) and WC (C)." figureDoi="10.3897/zookeys.979.56863.figure7" httpUri="https://binary.pensoft.net/fig/467835" pageId="0" pageNumber="133">7</figureCitation>
). The type series was collected after gentle rains, either by hand or in traps. The holotype was first observed feeding on small, unidentified ants (family
<taxonomicName family="Formicidae" lsidName="" pageId="0" pageNumber="133" rank="family">Formicidae</taxonomicName>
). No information is available on egg deposit sites and clutch sizes. One of us (WC) has discovered remains of
<taxonomicName lsidName="B. ombelanonga" pageId="0" pageNumber="133" rank="species" species="ombelanonga">
<emphasis italics="true" pageId="0" pageNumber="133">B. ombelanonga</emphasis>
</taxonomicName>
in the stomach contents of two snake species,
<taxonomicName class="Squamata" family="Psammophiidae" genus="Kladirostratus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kladirostratus acutus" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="acutus">
<emphasis italics="true" pageId="0" pageNumber="133">Kladirostratus acutus</emphasis>
</taxonomicName>
(
<taxonomicName family="Psammophiidae" lsidName="" pageId="0" pageNumber="133" rank="family">Psammophiidae</taxonomicName>
; PEM R23450) and
<taxonomicName authorityName="Boulenger" authorityYear="1905" class="Reptilia" family="Viperidae" genus="Causus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Causus bilineatus" order="Squamata" pageId="0" pageNumber="133" phylum="Chordata" rank="species" species="bilineatus">
<emphasis italics="true" pageId="0" pageNumber="133">Causus bilineatus</emphasis>
</taxonomicName>
(
<taxonomicName authorityName="Oppel" authorityYear="1811" family="Viperidae" lsidName="" pageId="0" pageNumber="133" rank="family">Viperidae</taxonomicName>
; PEM R23321) from the Cuando and Cuito River sources, respectively.
</paragraph>
<caption doi="10.3897/zookeys.979.56863.figure7" httpUri="https://binary.pensoft.net/fig/467835" pageId="0" pageNumber="133" start="Figure 7" startId="F7">
<paragraph pageId="0" pageNumber="133">
<emphasis bold="true" pageId="0" pageNumber="133">Figure 7.</emphasis>
Photos of typical habitat of
<taxonomicName lsidName="B. ombelanonga" pageId="0" pageNumber="133" rank="species" species="ombelanonga">
<emphasis italics="true" pageId="0" pageNumber="133">B. ombelanonga</emphasis>
</taxonomicName>
sp. nov.:
<emphasis bold="true" pageId="0" pageNumber="133">A</emphasis>
a view of the Kwanza River and bordering savannah, near the type locality, in Kissama National Park, Luanda Province
<emphasis bold="true" pageId="0" pageNumber="133">B</emphasis>
savannah near Embala Seque (14 km N of Cassumbi village),
<normalizedToken originalValue="Bié">Bie</normalizedToken>
Province
<emphasis bold="true" pageId="0" pageNumber="133">C</emphasis>
Cuanavale River source lake and associated miombo savannah woodland. Photographs by LMPC (
<emphasis bold="true" pageId="0" pageNumber="133">A, B</emphasis>
) and WC (
<emphasis bold="true" pageId="0" pageNumber="133">C</emphasis>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="etymology">
<paragraph pageId="0" pageNumber="133">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="133">
The name
<emphasis italics="true" pageId="0" pageNumber="133">ombelanonga</emphasis>
is a derived combination of two words in Umbundu, a native Angolan language, for rain (
<emphasis italics="true" pageId="0" pageNumber="133">ombela</emphasis>
) and frog (
<emphasis italics="true" pageId="0" pageNumber="133">anonga</emphasis>
). The species epithet is used as an invariable noun in apposition to the generic name.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="133" type="conservation status">
<paragraph pageId="0" pageNumber="133">Conservation status.</paragraph>
<paragraph pageId="0" pageNumber="133">
Given that it appears widely distributed, we suggest that
<taxonomicName lsidName="B. ombelanonga" pageId="0" pageNumber="133" rank="species" species="ombelanonga">
<emphasis italics="true" pageId="0" pageNumber="133">B. ombelanonga</emphasis>
</taxonomicName>
be included in the IUCN category of Least Concern. The type locality lies within Kissama National Park, which grants some legal protection from major habitat degradation and loss, though the park has recently experienced significant wildfires. Additionally, the paratype localities in southeastern Angola (visited during field activities related to the
<bibRefCitation author="National Geographic Okavango Wilderness Project" journalOrPublisher="Arquivos do Museu Bocage" pageId="0" pageNumber="133" refId="B43" refString="National Geographic Okavango Wilderness Project, 2017. Initial Findings from Exploration of the Upper Catchments of the Cuito, Cuanavale, and Cuando Rivers, May 2015 to December 2016. Unpublished report." title="Initial Findings from Exploration of the Upper Catchments of the Cuito, Cuanavale, and Cuando Rivers, May 2015 to December 2016. Unpublished report." year="2017">National Geographic Okavango Wilderness Project 2017</bibRefCitation>
) are relatively pristine and ecologically intact miombo savannah that comprise an area recently proposed for formal protection.
</paragraph>
</subSubSection>
</treatment>
</document>