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<document id="61B03E79C56C2F3C362F21F9300DA188" ID-DOI="10.11646/zootaxa.3620.3.1" ID-GBIF-Dataset="a501bd5f-fdee-474a-95cc-f791b29f5155" ID-ISSN="1175-5326" ID-Zenodo-Dep="215983" ID-ZooBank="03B8D237-7C7D-4E79-A020-4305ACF119B7" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460579560794" checkinUser="plazi" docAuthor="Channing, A., Hillers, A., Lötters, S., Rödel, O., Schick, S., Conradie, W., Rödder, D., Wagner, P., Dehling, J. M., Du Preez, L. H., Kielgast, J. &amp; Burger, M." docDate="2013" docId="E5775E59FFDDFFB3F8856906FEFA348A" docLanguage="en" docName="zt03620p350.pdf" docOrigin="Zootaxa 3620 (3)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Hyperolius benguellensis Bocage 1893" docType="treatment" docVersion="8" lastPageNumber="318" masterDocId="194E2621FFCDFFA2F8126D1AFA4F3332" masterDocTitle="Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species" masterLastPageNumber="350" masterPageNumber="301" pageNumber="317" updateTime="1698324059666" updateUser="plazi">
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<mods:title id="44F9E891B8EECE78188006B80F0DE760">Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species</mods:title>
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<treatment id="E5775E59FFDDFFB3F8856906FEFA348A" ID-DOI="http://doi.org/10.5281/zenodo.6154922" ID-GBIF-Taxon="119560980" ID-Zenodo-Dep="6154922" LSID="urn:lsid:plazi:treatment:E5775E59FFDDFFB3F8856906FEFA348A" httpUri="http://treatment.plazi.org/id/E5775E59FFDDFFB3F8856906FEFA348A" lastPageId="17" lastPageNumber="318" pageId="16" pageNumber="317">
<subSubSection id="25C4BCC4FFDDFFB2F8856906F8F337F3" pageId="16" pageNumber="317" type="nomenclature">
<paragraph id="6D61EF4FFFDDFFB2F8856906F8CA3707" blockId="16.[151,645,1051,1146]" box="[151,645,1051,1078]" pageId="16" pageNumber="317">
<heading id="36295823FFDDFFB2F8856906F8CA3707" bold="true" box="[151,645,1051,1078]" fontSize="11" level="1" pageId="16" pageNumber="317" reason="1">
<taxonomicName id="AADE94CCFFDDFFB2F8856906F8CA3707" ID-CoL="3NQRC" authority="Bocage, 1893" authorityName="Bocage" authorityYear="1893" box="[151,645,1051,1078]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="benguellensis">
<emphasis id="5FAA335DFFDDFFB2F8856906F8CA3707" bold="true" box="[151,645,1051,1078]" pageId="16" pageNumber="317">
<emphasis id="5FAA335DFFDDFFB2F8856906FB873704" bold="true" box="[151,456,1052,1078]" italics="true" pageId="16" pageNumber="317">Hyperolius benguellensis</emphasis>
(Bocage, 1893)
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<paragraph id="6D61EF4FFFDDFFB2F8856924FAA43748" blockId="16.[151,645,1051,1146]" pageId="16" pageNumber="317">
Benguella Long Reed Frog (
<figureCitation id="F5E5F3CAFFDDFFB2F88D6978FAAC3748" box="[159,227,1122,1146]" captionStart="FIGURE 4" captionStartId="11.[151,250,1898,1921]" captionTargetBox="[153,1434,193,1876]" captionTargetId="figure@11.[153,1434,193,1877]" captionTargetPageId="11" captionText="FIGURE 4. Hyperolius acuticeps Chongoni, Malawi, photo V. Mercurio (A); H. adspersus Bateka Nature Reserve, Gabon, photo G. Jongsma (B); H. benguellensis, Humpata, Angola, photo A. Channing (C); H. friedemanni sp. nov., Karonga, Malawi, photo V. Mercurio (D); H. howelli sp. nov. holotype, SAIAB 118979, Arusha, Tanzania, photo A. Channing (E); H. howelli sp. nov. female paratype, SAIAB 118980 - 1, Arusha, Tanzania, photo A. Channing (F)." httpUri="https://zenodo.org/record/215987/files/figure.png" pageId="16" pageNumber="317">Fig. 4</figureCitation>
)
</paragraph>
<paragraph id="6D61EF4FFFDDFFB2F88569B0F8F337F3" blockId="16.[151,700,1194,1217]" box="[151,700,1194,1217]" pageId="16" pageNumber="317">
<emphasis id="5FAA335DFFDDFFB2F88569B0FAB337F3" bold="true" box="[151,252,1194,1217]" pageId="16" pageNumber="317">Synonym</emphasis>
.
<taxonomicName id="AADE94CCFFDDFFB2F91B69B0F8F337F3" authority="Boulenger, 1901" authorityName="Boulenger" authorityYear="1901" box="[265,700,1194,1217]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="oxyrhynchus">
<emphasis id="5FAA335DFFDDFFB2F91B69B0FBB137F2" box="[265,510,1194,1216]" italics="true" pageId="16" pageNumber="317">Hyperolius oxyrhynchus</emphasis>
(Boulenger, 1901)
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</paragraph>
</subSubSection>
<subSubSection id="25C4BCC4FFDDFFB2F88569F7F886367F" pageId="16" pageNumber="317" type="materials_examined">
<paragraph id="6D61EF4FFFDDFFB2F88569F7F886367F" blockId="16.[151,1437,1261,2006]" pageId="16" pageNumber="317">
<emphasis id="5FAA335DFFDDFFB2F88569F7FB293634" bold="true" box="[151,358,1261,1286]" pageId="16" pageNumber="317">Genetic material</emphasis>
. ZMB 772712, ZMB 77318 (Humpata,
<collectingCountry id="15C9AFDFFFDDFFB2FB7169F7F9F53637" box="[867,954,1261,1285]" name="Angola" pageId="16" pageNumber="317">Angola</collectingCountry>
); ZMB 772734 (Bicuar National Park,
<collectingCountry id="15C9AFDFFFDDFFB2F885680BFAA1361B" box="[151,238,1297,1321]" name="Angola" pageId="16" pageNumber="317">Angola</collectingCountry>
); ZMB 77275 (Zootecnica Plateau, Humpata,
<collectingCountry id="15C9AFDFFFDDFFB2FB17680BF913361B" box="[773,860,1297,1321]" name="Angola" pageId="16" pageNumber="317">Angola</collectingCountry>
); AACRG 1030 (Kaparotta,
<collectingCountry id="15C9AFDFFFDDFFB2FCBB680BFF52361B" box="[1193,1309,1297,1321]" name="Botswana" pageId="16" pageNumber="317">Botswana</collectingCountry>
); GenBank
<accessionNumber id="728D72ACFFDDFFB2F885682FFB5E367F" box="[151,273,1333,1357]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/AF215224" pageId="16" pageNumber="317">AF215224</accessionNumber>
,
<accessionNumber id="728D72ACFFDDFFB2F90E682FFBDB367F" box="[284,404,1333,1357]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/AF215442" pageId="16" pageNumber="317">AF215442</accessionNumber>
(Rundu,
<collectingCountry id="15C9AFDFFFDDFFB2F9EF682FF810367F" box="[509,607,1333,1357]" name="Namibia" pageId="16" pageNumber="317">Namibia</collectingCountry>
) (
<figureCitation id="F5E5F3CAFFDDFFB2FA64682FF8F8367F" box="[630,695,1333,1357]" captionStart="FIGURE 1" captionStartId="4.[151,250,1838,1861]" captionTargetBox="[170,1406,1021,1787]" captionTargetId="figure@4.[151,1436,985,1817]" captionTargetPageId="4" captionText="FIGURE 1. Map showing the distribution of the sequenced specimens of recognised species in the Hyperolius nasutus group plus H. lamottei. Species numbers represent the following: 1 — H. acuticeps, 2 — H. adspersus, 3 — H. benguellensis, 4 — H. friedemanni sp. nov., 5 — H. howelli sp. nov., 6 — H. igbettensis, 7 — H. inyangae sp. nov., 8 — H. jacobseni sp. nov., 9 — H. lamottei, 10 — H. lupiroensis sp. nov., 11 — H. nasutus, 12 — H. poweri, 13 — H. rwandae sp. nov., 14 — H. dartevellei, 15 — H. viridis, 16 — H. nasicus." httpUri="https://zenodo.org/record/215984/files/figure.png" pageId="16" pageNumber="317">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="25C4BCC4FFDDFFB2F8DB6843F83135EB" pageId="16" pageNumber="317" type="diagnosis">
<paragraph id="6D61EF4FFFDDFFB2F8DB6843F83135EB" blockId="16.[151,1437,1261,2006]" pageId="16" pageNumber="317">
<emphasis id="5FAA335DFFDDFFB2F8DB6843FB733640" bold="true" box="[201,316,1369,1394]" pageId="16" pageNumber="317">Diagnosis</emphasis>
. The advertisement call (
<figureCitation id="F5E5F3CAFFDDFFB2FA496843F8D13643" box="[603,670,1369,1393]" captionStart="FIGURE 8" captionStartId="18.[151,250,918,941]" captionTargetBox="[174,1408,210,897]" captionTargetId="figure@18.[174,1414,195,897]" captionTargetPageId="18" captionText="FIGURE 8. Advertisement calls of H. benguellensis Humpata (left) and H. friedemanni Karonga (right)." httpUri="https://zenodo.org/record/215991/files/figure.png" pageId="16" pageNumber="317">Fig. 8</figureCitation>
) is a brief note consisting of five pulses, followed by 14 pulses at a slower rate, with a duration of
<geoCoordinate id="08EA8988FFDDFFB2F9FE6867F87E36A7" box="[492,561,1405,1429]" direction="south" orientation="latitude" pageId="16" pageNumber="317" precision="555" value="-0.41">0.41 s</geoCoordinate>
. It can be distinguished from the brief calls consisting only of a single note,
<taxonomicName id="AADE94CCFFDDFFB2FD936864FB48368B" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="adspersus">
<emphasis id="5FAA335DFFDDFFB2FD936864FB48368B" italics="true" pageId="16" pageNumber="317">H. adspersus</emphasis>
</taxonomicName>
and
<taxonomicName id="AADE94CCFFDDFFB2F92F68B8FB94368B" box="[317,475,1441,1465]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="lupiroensis" status="sp. nov.">
<emphasis id="5FAA335DFFDDFFB2F92F68B8FB94368B" box="[317,475,1441,1465]" italics="true" pageId="16" pageNumber="317">H. lupiroensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="44998E26FFDDFFB2F9F368B8F875368B" box="[481,570,1442,1466]" pageId="16" pageNumber="317" rank="species">
<emphasis id="5FAA335DFFDDFFB2F9F368B8F87C3688" bold="true" box="[481,563,1442,1466]" pageId="16" pageNumber="317">sp. nov</emphasis>
.
</taxonomicNameLabel>
and those consisting only of a series of pulses,
<taxonomicName id="AADE94CCFFDDFFB2FC5F68B8FE96368B" box="[1101,1241,1442,1465]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="acuticeps">
<emphasis id="5FAA335DFFDDFFB2FC5F68B8FE96368B" box="[1101,1241,1442,1465]" italics="true" pageId="16" pageNumber="317">H. acuticeps</emphasis>
</taxonomicName>
,
<taxonomicName id="AADE94CCFFDDFFB2FCF468B8FF3C368B" box="[1254,1395,1441,1465]" class="Amphibia" family="Hyperoliidae" genus="Hyperolious" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="jacobseni" status="sp. nov.">
<emphasis id="5FAA335DFFDDFFB2FCF468B8FF3C368B" box="[1254,1395,1441,1465]" italics="true" pageId="16" pageNumber="317">H. jacobseni</emphasis>
</taxonomicName>
<taxonomicNameLabel id="44998E26FFDDFFB2FD6868B8FA8936EF" pageId="16" pageNumber="317" rank="species">
<emphasis id="5FAA335DFFDDFFB2FD6868B8FA8E36EF" bold="true" pageId="16" pageNumber="317">sp. nov</emphasis>
.
</taxonomicNameLabel>
,
<taxonomicName id="AADE94CCFFDDFFB2F8C968DCFB1536EF" box="[219,346,1478,1501]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="nasutus">
<emphasis id="5FAA335DFFDDFFB2F8C968DCFB1536EF" box="[219,346,1478,1501]" italics="true" pageId="16" pageNumber="317">H. nasutus</emphasis>
</taxonomicName>
and
<taxonomicName id="AADE94CCFFDDFFB2F98968DCF87436EF" box="[411,571,1477,1501]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="dartevellei">
<emphasis id="5FAA335DFFDDFFB2F98968DCF87436EF" box="[411,571,1477,1501]" italics="true" pageId="16" pageNumber="317">H. dartevellei</emphasis>
</taxonomicName>
. The other species with advertisement calls consisting of an initial note followed by some discrete pulses can be distinguished either by their short duration, less than
<geoCoordinate id="08EA8988FFDDFFB2FCE068F3FF633533" box="[1266,1324,1513,1537]" direction="south" orientation="latitude" pageId="16" pageNumber="317" precision="5555" value="-0.2">0.2 s</geoCoordinate>
, as in
<taxonomicName id="AADE94CCFFDDFFB2FD9368F0FB503517" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="friedemanni" status="sp. nov.">
<emphasis id="5FAA335DFFDDFFB2FD9368F0FB503517" italics="true" pageId="16" pageNumber="317">H. friedemanni</emphasis>
</taxonomicName>
<taxonomicNameLabel id="44998E26FFDDFFB2F93B6B17FBCB3517" box="[297,388,1549,1573]" pageId="16" pageNumber="317" rank="species">
<emphasis id="5FAA335DFFDDFFB2F93B6B17FB313517" bold="true" box="[297,382,1549,1573]" pageId="16" pageNumber="317">sp. nov</emphasis>
.
</taxonomicNameLabel>
,
<taxonomicName id="AADE94CCFFDDFFB2F9856B14F8413517" box="[407,526,1549,1573]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="howelli" status="sp. nov.">
<emphasis id="5FAA335DFFDDFFB2F9856B14F8413517" box="[407,526,1549,1573]" italics="true" pageId="16" pageNumber="317">H. howelli</emphasis>
</taxonomicName>
<taxonomicNameLabel id="44998E26FFDDFFB2FA0B6B17F83A3517" box="[537,629,1549,1573]" pageId="16" pageNumber="317" rank="species">
<emphasis id="5FAA335DFFDDFFB2FA0B6B17F8203517" bold="true" box="[537,623,1549,1573]" pageId="16" pageNumber="317">sp. nov</emphasis>
.
</taxonomicNameLabel>
,
<taxonomicName id="AADE94CCFFDDFFB2FA956B14F96A3517" box="[647,805,1549,1573]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="igbettensis">
<emphasis id="5FAA335DFFDDFFB2FA956B14F96A3517" box="[647,805,1549,1573]" italics="true" pageId="16" pageNumber="317">H. igbettensis</emphasis>
</taxonomicName>
,
<taxonomicName id="AADE94CCFFDDFFB2FB246B14F9E53517" box="[822,938,1550,1573]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="poweri">
<emphasis id="5FAA335DFFDDFFB2FB246B14F9E53517" box="[822,938,1550,1573]" italics="true" pageId="16" pageNumber="317">H. poweri</emphasis>
</taxonomicName>
and
<taxonomicName id="AADE94CCFFDDFFB2FBF56B14FE203517" box="[999,1135,1549,1573]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="rwandae" status="sp. nov.">
<emphasis id="5FAA335DFFDDFFB2FBF56B14FE203517" box="[999,1135,1549,1573]" italics="true" pageId="16" pageNumber="317">H. rwandae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="44998E26FFDDFFB2FC6B6B17FE9A3517" box="[1145,1237,1549,1573]" pageId="16" pageNumber="317" rank="species">
<emphasis id="5FAA335DFFDDFFB2FC6B6B17FE803517" bold="true" box="[1145,1231,1549,1573]" pageId="16" pageNumber="317">sp. nov</emphasis>
.
</taxonomicNameLabel>
, or by the lower number of slow pulses, less than 10, as in
<taxonomicName id="AADE94CCFFDDFFB2FA7B6B28F8BD357B" box="[617,754,1586,1609]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="inyangae">
<emphasis id="5FAA335DFFDDFFB2FA7B6B28F8BD357B" box="[617,754,1586,1609]" italics="true" pageId="16" pageNumber="317">H. inyangae</emphasis>
</taxonomicName>
and
<taxonomicName id="AADE94CCFFDDFFB2FB356B28F9DF357B" box="[807,912,1585,1609]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="viridis">
<emphasis id="5FAA335DFFDDFFB2FB356B28F9DF357B" box="[807,912,1585,1609]" italics="true" pageId="16" pageNumber="317">H. viridis</emphasis>
</taxonomicName>
. See Table 3 for a summary of call parameters. The snout is shark-like in profile, protruding forward of the mouth in a straight line, before forming a sharp tip. It can be distinguished from the truncated, sharply- or bluntly rounded snout profiles as in
<emphasis id="5FAA335DFFDDFFB2FCCA6B60FB8D3587" italics="true" pageId="16" pageNumber="317">
<taxonomicName id="AADE94CCFFDDFFB2FCCA6B60FF2135A3" box="[1240,1390,1658,1681]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="acuticeps">H. acuticeps</taxonomicName>
,
<taxonomicName id="AADE94CCFFDDFFB2FD936B60FB453587" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="adspersus">H. adspersus</taxonomicName>
,
<taxonomicName id="AADE94CCFFDDFFB2F9056B84FB8D3587" box="[279,450,1693,1717]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="friedemanni" status="sp. nov.">H. friedemanni</taxonomicName>
</emphasis>
<taxonomicNameLabel id="44998E26FFDDFFB2F9D96B87F8693587" box="[459,550,1693,1717]" pageId="16" pageNumber="317" rank="species">
<emphasis id="5FAA335DFFDDFFB2F9D96B87F8503587" bold="true" box="[459,543,1693,1717]" pageId="16" pageNumber="317">sp. nov</emphasis>
.
</taxonomicNameLabel>
,
<emphasis id="5FAA335DFFDDFFB2FA276B84F93F3587" box="[565,880,1693,1717]" italics="true" pageId="16" pageNumber="317">
<taxonomicName id="AADE94CCFFDDFFB2FA276B84F89B3587" box="[565,724,1693,1717]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="igbettensis">H. igbettensis</taxonomicName>
,
<taxonomicName id="AADE94CCFFDDFFB2FAF36B84F93F3587" box="[737,880,1693,1717]" class="Amphibia" family="Hyperoliidae" genus="Hyperolious" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="jacobseni" status="sp. nov.">H. jacobseni</taxonomicName>
</emphasis>
<taxonomicNameLabel id="44998E26FFDDFFB2FB6B6B87F99B3587" box="[889,980,1693,1717]" pageId="16" pageNumber="317" rank="species">
<emphasis id="5FAA335DFFDDFFB2FB6B6B87F9823587" bold="true" box="[889,973,1693,1717]" pageId="16" pageNumber="317">sp. nov</emphasis>
.
</taxonomicNameLabel>
,
<taxonomicName id="AADE94CCFFDDFFB2FBF66B84FECC3587" box="[996,1155,1693,1717]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="lupiroensis" status="sp. nov.">
<emphasis id="5FAA335DFFDDFFB2FBF66B84FECC3587" box="[996,1155,1693,1717]" italics="true" pageId="16" pageNumber="317">H. lupiroensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="44998E26FFDDFFB2FC9E6B87FEA83587" box="[1164,1255,1693,1717]" pageId="16" pageNumber="317" rank="species">
<emphasis id="5FAA335DFFDDFFB2FC9E6B87FEAE3587" bold="true" box="[1164,1249,1693,1717]" pageId="16" pageNumber="317">sp. nov</emphasis>
.
</taxonomicNameLabel>
,
<emphasis id="5FAA335DFFDDFFB2FCE56B84FB3835EB" italics="true" pageId="16" pageNumber="317">
<taxonomicName id="AADE94CCFFDDFFB2FCE56B84FF3C3587" box="[1271,1395,1694,1717]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="nasutus">H. nasutus</taxonomicName>
,
<taxonomicName id="AADE94CCFFDDFFB2FD936B84FAA735EB" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="poweri">H. poweri</taxonomicName>
,
<taxonomicName id="AADE94CCFFDDFFB2F8E16BD8FB3835EB" box="[243,375,1729,1753]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="rwandae" status="sp. nov.">H. rwandae</taxonomicName>
</emphasis>
<taxonomicNameLabel id="44998E26FFDDFFB2F96C6BD8FB9835EB" box="[382,471,1730,1754]" pageId="16" pageNumber="317" rank="species">
<emphasis id="5FAA335DFFDDFFB2F96C6BD8FB9F35E8" bold="true" box="[382,464,1730,1754]" pageId="16" pageNumber="317">sp. nov</emphasis>
.
</taxonomicNameLabel>
and
<taxonomicName id="AADE94CCFFDDFFB2FA1F6BD8F83835EB" box="[525,631,1729,1753]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="16" pageNumber="317" phylum="Chordata" rank="species" species="viridis">
<emphasis id="5FAA335DFFDDFFB2FA1F6BD8F83835EB" box="[525,631,1729,1753]" italics="true" pageId="16" pageNumber="317">H. viridis</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="25C4BCC4FFDDFFB3F8DB6BFFF8493562" lastPageId="17" lastPageNumber="318" pageId="16" pageNumber="317" type="description">
<paragraph id="6D61EF4FFFDDFFB3F8DB6BFFFF08329E" blockId="16.[151,1437,1261,2006]" lastBlockId="17.[151,1437,151,1977]" lastPageId="17" lastPageNumber="318" pageId="16" pageNumber="317">
<emphasis id="5FAA335DFFDDFFB2F8DB6BFFF83635CC" bold="true" box="[201,633,1765,1790]" pageId="16" pageNumber="317">Description of a Humpata specimen</emphasis>
. Body long and slender, widest at mid-body, slightly tapering to groin; head comparatively small (HL/SUL 0.33, HW/SUL 0.30), not wider than trunk, longer than wide (HL/HW 1.11); snout long (SL/HL 0.42), bluntly rounded in dorsal view, acute, sharklike in profile (
<figureCitation id="F5E5F3CAFFDDFFB2FCB76A37FEBE3477" box="[1189,1265,1837,1861]" captionStart="FIGURE 6" captionStartId="14.[151,250,1961,1984]" captionTargetBox="[175,1394,193,1914]" captionTargetId="figure@14.[175,1394,193,1916]" captionTargetPageId="14" captionText="FIGURE 6. Snout profiles of representatives of the species in the Hyperolius nasutus group." httpUri="https://zenodo.org/record/215989/files/figure.png" pageId="16" pageNumber="317">Fig. 6</figureCitation>
), considerably projecting beyond lower jaw, wider than long (SL/EE 0.68); canthus rostralis distinct, moderately sharp, slightly concave from eye to just beyond nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril directed dorsolaterally; situated much closer to tip of snout than to eye (EN/NS 1.64), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.11); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.33); eye diameter shorter than snout (ED/SL 0.78); interorbital distance much wider than upper eyelid (IO/EW 1.71), and greater than internarial distance (IO/ NN 2.1); tympanum not visible externally; upper jaw with dentition; teeth on premaxilla larger than those on maxilla; choanae small, oval, located far anterolaterally at margins of roof of the mouth, vomer processes and teeth absent; tongue long 5.8, and narrow (2.7 at widest point), free for about three-fourths of length, bifurcated distally for about one-third of length; median lingual process absent; vocal sac single, median, subgular, mostly unpigmented and translucent when fully inflated; gular flap cream-coloured, granular; vocal sac aperture on each side of the mouth, situated lateral from and close to base of tongue, slit-like, long, directed posterolaterally.
</paragraph>
<paragraph id="6D61EF4FFFDCFFB3F8DB6CADFBDC32C6" blockId="17.[151,1437,151,1977]" pageId="17" pageNumber="318">Dorsal surfaces of head, trunk and limbs smooth; ventral surface of limbs and gular smooth, chin and abdomen slightly more areolate.</paragraph>
<paragraph id="6D61EF4FFFDCFFB3F8DB6F1AFAA931FE" blockId="17.[151,1437,151,1977]" pageId="17" pageNumber="318">
Fore limbs slender; hand moderately large (HND/SUL 0.30); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: I&lt;II&lt;IV&lt;III; subarticular tubercles rounded, well developed, with one on fingers I and II, two on fingers III and IV, with proximal tubercle on finger IV hardly discernible; webbing formula of the hand
<date id="1960C98FFFDCFFB3FA9E6F76F8EC31B6" box="[652,675,620,644]" pageId="17" pageNumber="318">
<emphasis id="5FAA335DFFDCFFB3FA9E6F76F8D831B6" bold="true" box="[652,663,620,644]" pageId="17" pageNumber="318">I</emphasis>
2
</date>
2.5
<emphasis id="5FAA335DFFDCFFB3FAC76F76F8A431B6" bold="true" box="[725,747,620,644]" pageId="17" pageNumber="318">II</emphasis>
2
<date id="1960C98FFFDCFFB3FB1A6F76F95B31B6" box="[776,788,620,644]" pageId="17" pageNumber="318">3</date>
<emphasis id="5FAA335DFFDCFFB3FB076F76F97931B6" bold="true" box="[789,822,620,644]" pageId="17" pageNumber="318">III</emphasis>
2.52.5
<emphasis id="5FAA335DFFDCFFB3FB986F76F9E631B6" bold="true" box="[906,937,620,644]" pageId="17" pageNumber="318">IV</emphasis>
(after Myers &amp; Duellman [1982]); thenar tubercle absent; palmar tubercles absent; metacarpals without supernumerary tubercles; nuptial pads or asperities absent.
</paragraph>
<paragraph id="6D61EF4FFFDCFFB3F8DB6FCDFBC030F5" blockId="17.[151,1437,151,1977]" pageId="17" pageNumber="318">
<collectingCountry id="15C9AFDFFFDCFFB3F8DB6FCDFB4E31DD" box="[201,257,727,751]" name="India" pageId="17" pageNumber="318">Hind</collectingCountry>
limbs slender, moderately long (LEG/SUL 1.55); tibio-tarsal articulation reaching to level of snout tip when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.53), shorter than thigh (TFL/THL 0.91); heels overlapping each other considerably when knees are flexed and thighs are held laterally at right angle to body; foot shorter than tibiofibula (FOT/TFL 0.83); relative length of toes: I&lt;II&lt;III&lt;V&lt;IV; discs of toes smaller than those of fingers; subarticular tubercles: one on toes I and II, two on toes III and V, and three on toe IV; pedal webbing formula (
<figureCitation id="F5E5F3CAFFDCFFB3F9756E96FBE43096" box="[359,427,908,932]" captionStart="FIGURE 7" captionStartId="15.[151,250,1970,1993]" captionTargetBox="[175,1414,193,1943]" captionTargetId="figure@15.[175,1414,193,1943]" captionTargetPageId="15" captionText="FIGURE 7. Webbing diagrams of representatives of the species in the Hyperolius nasutus group." httpUri="https://zenodo.org/record/215990/files/figure.png" pageId="17" pageNumber="318">Fig. 7</figureCitation>
)
<emphasis id="5FAA335DFFDCFFB3F9A96E96FB893096" bold="true" box="[443,454,908,932]" pageId="17" pageNumber="318">I</emphasis>
1
<date id="1960C98FFFDCFFB3F9F76E96F8493096" box="[485,518,908,932]" pageId="17" pageNumber="318">
1
<emphasis id="5FAA335DFFDCFFB3F9E26E96F8493096" bold="true" box="[496,518,908,932]" pageId="17" pageNumber="318">II</emphasis>
</date>
0.75
<date id="1960C98FFFDCFFB3FA586E96F81C3096" box="[586,595,908,932]" pageId="17" pageNumber="318">1</date>
<emphasis id="5FAA335DFFDCFFB3FA416E96F83B3096" bold="true" box="[595,628,908,932]" pageId="17" pageNumber="318">III</emphasis>
0
<date id="1960C98FFFDCFFB3FA866E96F8D13096" box="[660,670,908,932]" pageId="17" pageNumber="318">1</date>
<emphasis id="5FAA335DFFDCFFB3FA8F6E96F8F33096" bold="true" box="[669,700,908,932]" pageId="17" pageNumber="318">IV</emphasis>
1
<date id="1960C98FFFDCFFB3FAC96E96F8B63096" box="[731,761,908,932]" pageId="17" pageNumber="318">
1
<emphasis id="5FAA335DFFDCFFB3FAF56E96F8B63096" bold="true" box="[743,761,908,932]" pageId="17" pageNumber="318">V</emphasis>
</date>
; inner metatarsal tubercle small, oval, prominent; outer one low, almost indistinct.
</paragraph>
<paragraph id="6D61EF4FFFDCFFB3F8DB6EC9FF2537DA" blockId="17.[151,1437,151,1977]" pageId="17" pageNumber="318">
<emphasis id="5FAA335DFFDCFFB3F8DB6EC9FBE830DE" bold="true" box="[201,423,979,1004]" pageId="17" pageNumber="318">Colouration in life</emphasis>
. Skin more or less translucent. Dorsum and dorsal surface of head and limbs dark green (
<figureCitation id="F5E5F3CAFFDCFFB3F88C6EEDFA91373D" box="[158,222,1015,1039]" captionStart="FIGURE 4" captionStartId="11.[151,250,1898,1921]" captionTargetBox="[153,1434,193,1876]" captionTargetId="figure@11.[153,1434,193,1877]" captionTargetPageId="11" captionText="FIGURE 4. Hyperolius acuticeps Chongoni, Malawi, photo V. Mercurio (A); H. adspersus Bateka Nature Reserve, Gabon, photo G. Jongsma (B); H. benguellensis, Humpata, Angola, photo A. Channing (C); H. friedemanni sp. nov., Karonga, Malawi, photo V. Mercurio (D); H. howelli sp. nov. holotype, SAIAB 118979, Arusha, Tanzania, photo A. Channing (E); H. howelli sp. nov. female paratype, SAIAB 118980 - 1, Arusha, Tanzania, photo A. Channing (F)." httpUri="https://zenodo.org/record/215987/files/figure.png" pageId="17" pageNumber="318">Fig 4</figureCitation>
); lateral sides of head and scapular region dark green; light, yellowish-white, moderately broad dorsolateral stripe running along each side of the body from snout tip, over the eye to vent; Pale paravertebral stripes originating on posterior of snout, diverging to level of eyes, and then running parallel to vent; small dark brown to black dots and on dorsum, most densely bordering both sides of dorsolateral stripes; distal portions of fingers and toes, especially the tips, yellow; ventral side and parts of dorsal side of thigh and upper arm largely unpigmented but with irregular dark spots, appearing bluish-green; peritoneum white, shining through the translucent belly skin. Iris reddish-brown.
<emphasis id="5FAA335DFFDCFFB3F96669D5F8F237DA" bold="true" box="[372,701,1231,1256]" pageId="17" pageNumber="318">Colouration in preservative</emphasis>
. All colours have faded to yellow; gular flap whitish-yellow.
</paragraph>
<paragraph id="6D61EF4FFFDCFFB3F8DB69E9F868364A" blockId="17.[151,1437,151,1977]" pageId="17" pageNumber="318">
<emphasis id="5FAA335DFFDCFFB3F8DB69E9FB78363E" bold="true" box="[201,311,1267,1292]" pageId="17" pageNumber="318">Variation</emphasis>
. The female ZMB 77271 is similar to the male in measurements (Appendix 2). The female is larger than the male (SUL 23.5). Colouration is variable; ZMB 77271 has a pale green dorsum with irregular darker marks, and yellow eyelids, with fingers and toes also yellow, and a white underside. AC 3073 has pale dorsolateral stripes on a dark green background.
</paragraph>
<paragraph id="6D61EF4FFFDCFFB3F8DB689EF86A36AE" blockId="17.[151,1437,151,1977]" box="[201,549,1412,1436]" pageId="17" pageNumber="318">
<emphasis id="5FAA335DFFDCFFB3F8DB689EFBE736AE" bold="true" box="[201,424,1412,1436]" pageId="17" pageNumber="318">Eggs and tadpoles.</emphasis>
Unknown.
</paragraph>
<paragraph id="6D61EF4FFFDCFFB3F8DB68BDF8493562" blockId="17.[151,1437,151,1977]" pageId="17" pageNumber="318">
<emphasis id="5FAA335DFFDCFFB3F8DB68BDFB6A36F2" bold="true" box="[201,293,1447,1472]" pageId="17" pageNumber="318">Habitat</emphasis>
. We found the species only in open grassy habitats, along stream banks and man-made impoundments where sedges and other tall emergent vegetation were present (Humpata, Bicuar NP). Specimens were observed on leaves and stems of vegetation between
<quantity id="AA2642AAFFDCFFB3FA7268EAF8D6353A" box="[608,665,1520,1544]" metricMagnitude="-2" metricUnit="m" metricValue="5.0" pageId="17" pageNumber="318" unit="cm" value="5.0">5 cm</quantity>
and 1.0 m above water level. Males called from elevated positions. The following species were found sympatrically or even syntopically with the new species:
<taxonomicName id="AADE94CCFFDCFFB3FD326B0EFB6F3562" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="17" pageNumber="318" phylum="Chordata" rank="species" species="marmoratus">
<emphasis id="5FAA335DFFDCFFB3FD326B0EFB6F3562" italics="true" pageId="17" pageNumber="318">Hyperolius marmoratus</emphasis>
</taxonomicName>
and
<taxonomicName id="AADE94CCFFDCFFB3F9446B23FBB13562" box="[342,510,1592,1616]" class="Insecta" family="Curculionidae" genus="Xenopus" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="318" phylum="Arthropoda" rank="species" species="laevis">
<emphasis id="5FAA335DFFDCFFB3F9446B23FBB13562" box="[342,510,1592,1616]" italics="true" pageId="17" pageNumber="318">Xenopus laevis</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="25C4BCC4FFDCFFB3F8DB6B46FEFA348A" pageId="17" pageNumber="318" type="discussion">
<paragraph id="6D61EF4FFFDCFFB3F8DB6B46F82535AA" blockId="17.[151,1437,151,1977]" pageId="17" pageNumber="318">
<emphasis id="5FAA335DFFDCFFB3F8DB6B46FB793546" bold="true" box="[201,310,1628,1652]" pageId="17" pageNumber="318">Remarks</emphasis>
. The species is confirmed using molecular data from Rundu in the Caprivi Strip of
<collectingCountry id="15C9AFDFFFDCFFB3FD266B46FFD73546" box="[1332,1432,1628,1652]" name="Namibia" pageId="17" pageNumber="318">Namibia</collectingCountry>
, northern
<collectingCountry id="15C9AFDFFFDCFFB3F8EE6B9BFB2235AA" box="[252,365,1665,1688]" name="Botswana" pageId="17" pageNumber="318">Botswana</collectingCountry>
, and southern
<collectingCountry id="15C9AFDFFFDCFFB3FA026B9AF82935AA" box="[528,614,1664,1688]" name="Angola" pageId="17" pageNumber="318">Angola</collectingCountry>
.
</paragraph>
<paragraph id="6D61EF4FFFDCFFB3F8DB6BBEFF4535D2" blockId="17.[151,1437,151,1977]" pageId="17" pageNumber="318">
The
<typeStatus id="B26551EDFFDCFFB3F9136BBFFB7E358F" box="[257,305,1701,1725]" pageId="17" pageNumber="318">type</typeStatus>
locality of
<taxonomicName id="AADE94CCFFDCFFB3F9D16BBFF8CC358E" box="[451,643,1700,1724]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="17" pageNumber="318" phylum="Chordata" rank="species" species="benguellensis">
<emphasis id="5FAA335DFFDCFFB3F9D16BBFF8CC358E" box="[451,643,1700,1724]" italics="true" pageId="17" pageNumber="318">H. benguellensis</emphasis>
</taxonomicName>
is Caota,
<collectingCountry id="15C9AFDFFFDCFFB3FB166BBEF915358E" box="[772,858,1700,1724]" name="Angola" pageId="17" pageNumber="318">Angola</collectingCountry>
. The
<typeStatus id="B26551EDFFDCFFB3FBB06BBFF99D358F" box="[930,978,1701,1725]" pageId="17" pageNumber="318">type</typeStatus>
series (MBL 17.220-223; now Museu Bocage, National Museum of Natural History, University of Lisbon) has been destroyed (Frost 2011).
</paragraph>
<paragraph id="6D61EF4FFFDCFFB3F8DB6BF6FEFA348A" blockId="17.[151,1437,151,1977]" pageId="17" pageNumber="318">
Our specimens agree with the original description of a sharp snout and small dorsal speckles. The specimens from Humpata show a range of colour patterns, from a uniform finely spotted dorsum, to pale dorsolateral stripes, to dorsolateral and paravertebral stripes. This variation was absent from the
<typeStatus id="B26551EDFFDCFFB3FBE06A2FFE6D347F" box="[1010,1058,1845,1869]" pageId="17" pageNumber="318">type</typeStatus>
description. The genetic material is from the same drainage basin as the
<typeStatus id="B26551EDFFDCFFB3FA676A43F8E63443" box="[629,681,1881,1905]" pageId="17" pageNumber="318">type</typeStatus>
.
<taxonomicName id="AADE94CCFFDCFFB3FAAB6A42F9813442" box="[697,974,1880,1904]" class="Amphibia" family="Hyperoliidae" genus="Hyperolius" kingdom="Animalia" order="Anura" pageId="17" pageNumber="318" phylum="Chordata" rank="species" species="oxyrhynchus">
<emphasis id="5FAA335DFFDCFFB3FAAB6A42F9813442" box="[697,974,1880,1904]" italics="true" pageId="17" pageNumber="318">Hyperolius oxyrhynchus</emphasis>
</taxonomicName>
is regarded as a synonym as the
<typeStatus id="B26551EDFFDCFFB3FD796A43FFD43443" box="[1387,1435,1881,1905]" pageId="17" pageNumber="318">type</typeStatus>
description matches this species. The species is presently only confirmed from southern
<collectingCountry id="15C9AFDFFFDCFFB3FC7D6A66FE8B34A6" box="[1135,1220,1916,1940]" name="Angola" pageId="17" pageNumber="318">Angola</collectingCountry>
, northern
<collectingCountry id="15C9AFDFFFDCFFB3FD266A66FFD834A6" box="[1332,1431,1916,1940]" name="Namibia" pageId="17" pageNumber="318">Namibia</collectingCountry>
, and northern
<collectingCountry id="15C9AFDFFFDCFFB3F93E6ABBFBD1348A" box="[300,414,1953,1976]" name="Botswana" pageId="17" pageNumber="318">Botswana</collectingCountry>
. We suggest that the conservation status Least Concern be maintained.
</paragraph>
</subSubSection>
</treatment>
</document>