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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.1001.58447" ID-GBIF-Dataset="698d424c-7e30-45c8-b96b-5dda6328b59e" ID-PMC="PMC7744391" ID-Pensoft-Pub="1313-2970-1001-1" ID-Pensoft-UUID="CED36F5A3D7453A2919A619A767B3367" ID-PubMed="33363428" ID-ZooBank="5914D476D746459ABCBFF86C8BD0A78B" ModsDocID="1313-2970-1001-1" checkinTime="1607628142719" checkinUser="pensoft" docAuthor="Caterino, Michael S. &amp; Tishechkin, Alexey K." docDate="2020" docId="B76E0F6B0A745ED59A6F18EB70B0ABBF" docLanguage="en" docName="ZooKeys 1001: 1-154" docOrigin="ZooKeys 1001" docPubDate="2020-12-09" docSource="http://dx.doi.org/10.3897/zookeys.1001.58447" docTitle="Phelister matatlantica Caterino &amp; Tishechkin 2020, sp. nov." docType="treatment" docUuid="D22D3203-C4EE-4544-A8C1-6979D38A0DA6" docUuidSource="ZooBank" docVersion="4" id="CED36F5A3D7453A2919A619A767B3367" lastPageNumber="1" masterDocId="CED36F5A3D7453A2919A619A767B3367" masterDocTitle="Recognition and revision of the Phelister blairi group (Histeridae, Histerinae, Exosternini)" masterLastPageNumber="154" masterPageNumber="1" pageNumber="1" updateTime="1668146370867" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Recognition and revision of the Phelister blairi group (Histeridae, Histerinae, Exosternini)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Caterino, Michael S.</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-2597-5707</mods:nameIdentifier>
<mods:affiliation>Department of Plant &amp; Environmental Sciences, Clemson University, Clemson, SC 29634, USA</mods:affiliation>
<mods:nameIdentifier type="email">mcateri@clemson.edu</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Tishechkin, Alexey K.</mods:namePart>
<mods:affiliation>California Dept. of Food and Agriculture Plant Pest Diagnostics Center, Sacramento, CA 95832, USA</mods:affiliation>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
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<mods:part>
<mods:date>2020</mods:date>
<mods:detail type="pubDate">
<mods:number>2020-12-09</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>1001</mods:number>
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<mods:extent unit="page">
<mods:start>1</mods:start>
<mods:end>154</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.1001.58447</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.1001.58447</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-1001-1</mods:identifier>
<mods:identifier type="ZooBank">5914D476D746459ABCBFF86C8BD0A78B</mods:identifier>
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<treatment ID-GBIF-Taxon="170725739" LSID="urn:lsid:zoobank.org:act:D22D3203-C4EE-4544-A8C1-6979D38A0DA6" httpUri="http://treatment.plazi.org/id/B76E0F6B0A745ED59A6F18EB70B0ABBF" lastPageNumber="1" pageId="0" pageNumber="1">
<subSubSection pageId="0" pageNumber="1" type="nomenclature">
<paragraph pageId="0" pageNumber="1">
47.
<taxonomicName LSID="http://zoobank.org/D22D3203-C4EE-4544-A8C1-6979D38A0DA6" authority="Caterino &amp; Tishechkin, 2020" authorityName="Caterino &amp; Tishechkin" authorityYear="2020" class="Insecta" family="Histeridae" genus="Phelister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phelister matatlantica" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="matatlantica" status="sp. nov.">Phelister matatlantica</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="1">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 29" captionStartId="F45" captionText="Figure 29. A, B Phelister microdens: A dorsal habitus B ventral habitus C, D P. matatlantica: C dorsal habitus D ventral habitus." figureDoi="10.3897/zookeys.1001.58447.figure29" httpUri="https://binary.pensoft.net/fig/484974" pageId="0" pageNumber="1">Figs 29C, D</figureCitation>
<figureCitation captionStart="Figure 30" captionStartId="F46" captionText="Figure 30. Male genitalia A, B Phelister microdens: A aedeagus, dorsal B aedeagus, lateral C, D P. matatlantica: C aedeagus, dorsal D aedeagus, lateral." figureDoi="10.3897/zookeys.1001.58447.figure30" httpUri="https://binary.pensoft.net/fig/484975" pageId="0" pageNumber="1">, 30C, D</figureCitation>
<figureCitation captionStart="Map 16" captionStartId="F44" captionText="Map 16. Collecting records for Phelister chicomendesi (stars), P. microdens (white circles), and P. matatlantica (squares)." figureDoi="10.3897/zookeys.1001.58447.map16" httpUri="https://binary.pensoft.net/fig/484973" pageId="0" pageNumber="1">, Map 16</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="type material">
<paragraph pageId="0" pageNumber="1">Type material.</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">
<emphasis italics="true" pageId="0" pageNumber="1">Holotype</emphasis>
male
</emphasis>
: &quot;
<emphasis bold="true" pageId="0" pageNumber="1">Brazil</emphasis>
:
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 50 km SE Mogi das Cruzes, Serra do Mar, Est. Biol.
<normalizedToken originalValue="Boracéia">Boraceia</normalizedToken>
, 800-900m [-23.7, -46], 28-30.IV.1997, F.
<normalizedToken originalValue="Génier">Genier</normalizedToken>
&amp; S. Ide, ex. feces trap, cloud forest&quot; / &quot;Caterino/Tishechkin
<taxonomicName authorityName="Bickhardt" authorityYear="1914" lsidName="" pageId="0" pageNumber="1" rank="tribe" tribe="Exosternini">Exosternini</taxonomicName>
Voucher EXO-00384&quot; (CMNC).
<emphasis bold="true" pageId="0" pageNumber="1">
<emphasis italics="true" pageId="0" pageNumber="1">Paratypes</emphasis>
(58): Brazil
</emphasis>
:
<normalizedToken originalValue="Espírito">Espirito</normalizedToken>
Santo, Pico Pedra Azul (
<geoCoordinate degrees="20.4333" direction="south" orientation="latitude" precision="5" value="-20.4333">-20.4333</geoCoordinate>
,
<geoCoordinate degrees="40.9833" direction="west" orientation="longitude" precision="5" value="-40.9833">-40.9833</geoCoordinate>
), 1500 m, January 1969, F. Vaz-de-Mello, EXO-03326 (CHND, 1ex.);
<normalizedToken originalValue="Espírito">Espirito</normalizedToken>
Santo, Venda Nova do Imigrante (
<geoCoordinate degrees="20.2667" direction="south" orientation="latitude" precision="5" value="-20.2667">-20.2667</geoCoordinate>
,
<geoCoordinate degrees="41.4167" direction="west" orientation="longitude" precision="5" value="-41.4167">-41.4167</geoCoordinate>
), December 2000, FIT, F.Z. Vaz-de-Mello, EXO-03327 (AKTC, 1ex.); Minas Gerais,
<normalizedToken originalValue="Águas">Aguas</normalizedToken>
Vermelhas (-15.75, -41.4667), December 1997, FIT, A.M. Bello &amp; F.Z. Vaz-de-Mello (CEMT, 2ex.); Minas Gerais,
<normalizedToken originalValue="Águas">Aguas</normalizedToken>
Vermelhas (-15.75, -41.4667), December 1983, Cad. de serpent (CHND, 2ex.); Minas Gerais, Aguas Vermelhas (-15.75, -41.4667), December 1997, F. Vaz-de-Mello (CHND, 20ex.); Minas Gerais, Lavras (-21.2333, -45), September 2001, FIT, Dry forest, F.Z. Vaz-de-Mello, EXO-03324 (AKTC, 1ex.); Minas Gerais,
<normalizedToken originalValue="Viçosa">Vicosa</normalizedToken>
(-20.75, -42.8833), December 1998, FIT, F.Z. Vaz-de-Mello, EXO-03325 (AKTC, 1ex.); Minas Gerais,
<normalizedToken originalValue="Viçosa">Vicosa</normalizedToken>
, Mata do Paraiso (
<geoCoordinate degrees="20.805" direction="south" orientation="latitude" precision="55" value="-20.805">-20.805</geoCoordinate>
,
<geoCoordinate degrees="42.8556" direction="west" orientation="longitude" precision="5" value="-42.8556">-42.8556</geoCoordinate>
), 750 m, 2/3/00, dg. tps., prim. mesophilic semideciduous Atlantic for., F.
<normalizedToken originalValue="Génier">Genier</normalizedToken>
(CMNC, 4ex.); Minas Gerais, Parque Estadual do Itacolomi, Trilha do Forno at
<geoCoordinate degrees="20.4290" direction="south" orientation="latitude" precision="5" value="-20.429">20.4290°S</geoCoordinate>
,
<geoCoordinate degrees="43.5075" direction="west" orientation="longitude" precision="5" value="-43.5075">43.5075°W</geoCoordinate>
, 1350 m, flight intercept trap, 8-10.ii.2014, A. K. Tishechkin (AKTC, 4ex.);
<normalizedToken originalValue="Paraná">Parana</normalizedToken>
, Piraquara Mananciais da Serra (
<geoCoordinate degrees="25.4934" direction="south" orientation="latitude" precision="5" value="-25.4934">-25.4934</geoCoordinate>
,
<geoCoordinate degrees="48.9786" direction="west" orientation="longitude" precision="5" value="-48.9786">-48.9786</geoCoordinate>
), 12/9/11-12/11/11, FIT, F.W.T. Leivas, M.S. Caterino &amp; A.K. Tishechkin, EXO-00929 (MSCC, 1ex.); Rio de Janeiro, 17 km E Nova Friburgo (
<geoCoordinate degrees="22.3844" direction="south" orientation="latitude" precision="5" value="-22.3844">-22.3844</geoCoordinate>
,
<geoCoordinate degrees="42.5583" direction="west" orientation="longitude" precision="5" value="-42.5583">-42.5583</geoCoordinate>
), 750 m, 1/23/00, carrion trap, secondary mount. Atlantic for., F.
<normalizedToken originalValue="Génier">Genier</normalizedToken>
&amp; S. Ide, EXO-03317 (CMNC, 1ex.); Rio de Janeiro, 17 km E Nova Friburgo (
<geoCoordinate degrees="22.3844" direction="south" orientation="latitude" precision="5" value="-22.3844">-22.3844</geoCoordinate>
,
<geoCoordinate degrees="42.5583" direction="west" orientation="longitude" precision="5" value="-42.5583">-42.5583</geoCoordinate>
), 750 m, 1/29/00, carrion trap, secondary mount. Atlantic for., F.
<normalizedToken originalValue="Génier">Genier</normalizedToken>
&amp; S. Ide (CMNC, 2ex.); Rio de Janeiro, N. Friburgo, Sitio Bacco (-22.3, -42.5), 4760ft, 3/26/91-4/9/91, FIT, K.P. Bland (NHMUK, 2ex.); Rio de Janeiro, Nova Friburgo (
<geoCoordinate degrees="22.2667" direction="south" orientation="latitude" precision="5" value="-22.2667">-22.2667</geoCoordinate>
,
<geoCoordinate degrees="42.5333" direction="west" orientation="longitude" precision="5" value="-42.5333">-42.5333</geoCoordinate>
), 10/26/09-10/31/09, FIT (CHND, 2ex.); Rio de Janeiro, Nova Friburgo, 17 km S (
<geoCoordinate degrees="22.3845" direction="south" orientation="latitude" precision="5" value="-22.3845">-22.3845</geoCoordinate>
,
<geoCoordinate degrees="42.5583" direction="west" orientation="longitude" precision="5" value="-42.5583">-42.5583</geoCoordinate>
), 750 m, 1/23/00, carrion trap, secondary montane Atlantic forest, F.
<normalizedToken originalValue="Génier">Genier</normalizedToken>
&amp; S. Ide, SM0808780 (SEMC, 1ex.); Rio de Janeiro, Nova Friburgo,
<normalizedToken originalValue="Macaé">Macae</normalizedToken>
de Cima (
<geoCoordinate degrees="22.2667" direction="south" orientation="latitude" precision="5" value="-22.2667">-22.2667</geoCoordinate>
,
<geoCoordinate degrees="42.5333" direction="west" orientation="longitude" precision="5" value="-42.5333">-42.5333</geoCoordinate>
), 1500 m, February 2000, F. Vaz-de-Mello (CHND, 2ex.); Rio de Janeiro,
<normalizedToken originalValue="Macaé">Macae</normalizedToken>
de Cima, Nova Friburgo (
<geoCoordinate degrees="22.3816" direction="south" orientation="latitude" precision="5" value="-22.3816">-22.3816</geoCoordinate>
,
<geoCoordinate degrees="42.4819" direction="west" orientation="longitude" precision="5" value="-42.4819">-42.4819</geoCoordinate>
), 1030 m, 10/1/03-10/31/07, FIT (AKTC &amp; CHND, 3ex.); Rio de Janeiro, Nova Friburgo, Sans Souci (-22.3, -42.6), 11/9/09-11/15/09, FIT, E. Grossi (DZUP, CESP, 3ex.); Santa Catarina, Chapecozinho (-26.8, -52), January 1958, litter primeral forest, F. Plaumann, FMNH-INS 0000 069 245 (FMNH, 1ex.); Santa Catarina, Nova Teutonia (
<geoCoordinate degrees="27.1833" direction="south" orientation="latitude" precision="5" value="-27.1833">-27.1833</geoCoordinate>
,
<geoCoordinate degrees="52.3833" direction="west" orientation="longitude" precision="5" value="-52.3833">-52.3833</geoCoordinate>
), 10/7/52, bei
<taxonomicName class="Insecta" family="Formicidae" genus="Acromyrmex" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Acromyrmex" order="Hymenoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="1">Acromyrmex</emphasis>
</taxonomicName>
, F. Plaumann, FMNH-INS 0000 069 241 (FMNH, 1ex.); Santa Catarina, Nova Teutonia (
<geoCoordinate degrees="27.1833" direction="south" orientation="latitude" precision="5" value="-27.1833">-27.1833</geoCoordinate>
,
<geoCoordinate degrees="52.3833" direction="west" orientation="longitude" precision="5" value="-52.3833">-52.3833</geoCoordinate>
), 5/10/60, Berlese: floor litter, F. Plaumann, FMNH-INS 0000 069 236 (FMNH, 1ex.); Santa Catarina, Rio das Antas (-26.9, -51.1), January 1953, Camargo, EXO-03328 (FMNH, 1ex.);
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, Est. Biol.
<normalizedToken originalValue="Boracéia">Boraceia</normalizedToken>
, 50 km SE Mogi das Cruzes, Serra do Mar (-23.7, -46), 800-900 m, 4/28/97-5/30/97, feces trap, cloud forest, F.
<normalizedToken originalValue="Génier">Genier</normalizedToken>
&amp; S. Ide (CMNC, 2ex.);
<emphasis bold="true" pageId="0" pageNumber="1">Paraguay</emphasis>
: Concepcion, Estancia Cororo (-23.4, -56.5), 11/26/99-11/30/99, Tampa con intestino de vacuno, C. Aguilar, EXO-03330 (CHND, 1ex.); Itapua, Itapua Poty (
<geoCoordinate degrees="26.5833" direction="south" orientation="latitude" precision="5" value="-26.5833">-26.5833</geoCoordinate>
,
<geoCoordinate degrees="55.5667" direction="west" orientation="longitude" precision="5" value="-55.5667">-55.5667</geoCoordinate>
), 2/2/91-2/28/91, C. Aguilar, EXO-03329 (CHND, 1ex.).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="description">
<paragraph pageId="0" pageNumber="1">Diagnostic description.</paragraph>
<paragraph pageId="0" pageNumber="1">
Length: 2.05-2.25 mm (avg. 2.13 mm); width: 1.85-2.01 mm (avg. 1.95 mm). Body rather round, elongate oval, weakly depressed, rufescent, often faintly bicolored, with the central portion of the elytra darkened; ground punctation rather fine and sparse; frons and epistoma depressed in middle, supraorbital stria complete, meeting frontal at sides, frontal stria complete, well impressed, weakly sinuate through depression; epistoma with sides raised; labrum weakly emarginate; each mandible with very small basal tooth; prescutellar impression oval, barely larger than scutellum; pronotal disk with few secondary punctures near sides; marginal pronotal stria complete along sides and front, strongly crenulate in front; lateral submarginal stria complete; median pronotal gland openings annulate, slightly&gt; 1/2 pronotal length behind anterior margin; elytron with single, complete epipleural stria; outer subhumeral stria present in apical one-third, inner absent; dorsal stria one weakened to obsolete in apical 1/2, stria two and three complete, striae four and five both present in apical 1/2, sutural stria in apical 2/3; propygidium with secondary punctures dense almost throughout, sparse only along posterior margin; pygidial secondary punctures smaller and sparser; prosternal keel shallowly emarginate at base, striae well-impressed joined along basal margin, slightly sinuate, united anteriorly; prosternal keel-lobe junction slightly narrowed, lobe short, rounded, with complete marginal stria; mesoventrite wide and short, produced anteriorly at middle, with fine marginal stria often interrupted at middle, continued at side by short postmesocoxal stria which ends behind mesocoxa; meso-metaventral stria bluntly angulate at middle, reaching anterior third of mesoventrite; lateral metaventral stria directed posterolaterally, nearly reaching middle of metacoxa; metaventrite impunctate; 1st abdominal ventrite with complete inner and abbreviated outer lateral striae; protibia with outer margin sinuate,
<normalizedToken originalValue="‘scalloped’">'scalloped'</normalizedToken>
, more weakly so in basal 1/2; meso- and metatibiae with fine marginal spines those of mesotibia more numerous and robust. Male: S8 with apical setal fringe; aedeagus with basal piece nearly 1/2 length of tegmen; tegmen with sides slightly widened to middle, evenly narrowed to bluntly rounded apices, apex slightly curved ventrally in lateral view; medioventral process very strong, produced at basal fourth; median lobe ~ 3/4 tegmen length, basal apodemes abruptly narrowed in basal 1/3.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="etymology">
<paragraph pageId="0" pageNumber="1">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="1">The species name is a contraction of Mata Atlantica, the Brazilian Atlantic Forest globally recognized hotspot on which this species largely depends.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="distribution">
<paragraph pageId="0" pageNumber="1">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="1">This species is known mainly from coastal (Atlantic forest) areas of southeastern Brazil. However, it also extends inland through Santa Catarina and into Paraguay.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="remarks">
<paragraph pageId="0" pageNumber="1">Remarks.</paragraph>
<paragraph pageId="0" pageNumber="1">While similar in most characters to other species in this group, this species is considerably larger than the others. They are also frequently faintly bicolored (redder anterad), have the lateral submarginal stria complete, and have the prosternal keel striae well impressed, united anteriorly.</paragraph>
</subSubSection>
</treatment>
</document>

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<treatment id="B76E28ECAC110815A33009D1F8700872" ID-DOI="http://doi.org/10.5281/zenodo.6269407" ID-GBIF-Taxon="100126027" ID-Zenodo-Dep="6269407" LSID="urn:lsid:plazi:treatment:B76E28ECAC110815A33009D1F8700872" httpUri="http://treatment.plazi.org/id/B76E28ECAC110815A33009D1F8700872" lastPageNumber="3" pageNumber="2">
<subSubSection id="8C3227AADE129FC5D015E74E87BDF43C" type="nomenclature">
<paragraph id="8A3C043B9566E5E2AB1A3F751019050C" pageNumber="2">
<taxonomicName id="9C50C421199ABFAEB1410290A80597FA" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:184276" class="Insecta" family="Formicidae" genus="Discothyrea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-HNS="Discothyrea sringerensis Zacharias &amp; Rajan" order="Hymenoptera" pageNumber="2" phylum="Arthropoda" rank="species" species="sringerensis">Discothyrea sringerensis</taxonomicName>
<taxonomicNameLabel id="2F075FB59CFD67E069E91C472F2ECE48" rank="species">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="51D443FCEC39E426CA624AA4904240B5" type="description">
<paragraph id="56763A168FB42582F27EF928770A6419" pageNumber="2">Description of holotype. Worker (Fig. 1). Total body length 3.05mm, Head length 0.80mm, Head width 0.54mm, Cephalic Index (HW/HL x 100) 67, scape length 0.46mm, petiole height 0.25mm, dorsal petiole length 0.10mm, dorsal petiole width 0.28mm.</paragraph>
<paragraph id="73DE41393398307167114CF3EFAD9C65" lastPageNumber="3" pageNumber="2">Head from front broadly oval with almost straight posterior margin. Clypeus extending forward so that it overhangs the rear part of the mandibles when they are closed. Anterior margin of the clypeus crenulated. Palp formula 5, 4. Antennae 10 segmented; scape massively clavate. Second antennal segment 0.06mm long, sub globular. 3rd to 6th antennal segments almost equal in length (each 0.07mm). 7th-9th antennal segments distinctly wider than long. Apical antennal segment large, twice as long as wide (Fig. 2). Depressed scrobal area distinct. Frontal carinae extending back about two-thirds the length of the head, dilated and horizontally flattened in the middle. Posterior end of frontal lobe opened and rounded anteroposteriorly and continuous with the regular convex surface of the head. Eyes placed on side of head nearer to the base of the mandibles than to the posterior border of the head, composed of at least 12 ommatidia in each with hairs in between the ommatidia.</paragraph>
<paragraph id="318B284C6B93E9F12F66CF04A42B3AE7" pageNumber="3">Mesosoma stout, with blunt, rounded humeral angles. Propodeum with more distinct propodeal teeth. Legs stout, spur of posterior leg pectinate, spur on middle leg absent.</paragraph>
<paragraph id="A05BF408274C66547AA7618BAD0BE4E8" pageNumber="3">Petiole in the form of a thick disc, 2.80 times broader than long dorsally and with two low rims running along the anterior and posterior margins of the petiole. The petiole is attached over a large portion of its posterior surface to the gaster and is lower in height than the gaster. In profile, petiole with weakly concave dorsal margin and a sloping anterior surface. Subpetiolar process on the ventral margin, a prominent compressed translucent tooth.</paragraph>
<paragraph id="A5CA5EB86C8FD6E9B2406D4D2A3B1FA6" pageNumber="3">Head, mesosoma and abdomen coarsely punctate. The scrobal depressions sparsely punctate with horizontal striations that extend from the lateral walls of the frontal lobe. The entire body densely clothed with short suberect to erect whitish pubescence, which is slightly shorter on the head and antennae.</paragraph>
<paragraph id="DEAFFE862C2273751E28741F192CFE94" pageNumber="3">Body colour reddish brown with mandibles, antennae, legs and tip of gaster yellowish brown.</paragraph>
</subSubSection>
<subSubSection id="DE62DF62465CFC8D802C75751D1830F7" type="nomenclature">
<paragraph id="CF609D761466B3AF4B28D2012E80EBD7" pageNumber="3">
<materialsCitation id="BA5A4D360B33E6465B329B6A66E26D3D" ID-GBIF-Occurrence="923893326" collectingDate="2001-03-26" collectingMethod="pitfall trap" collectorName="P.A. Sinu" country="India" latitude="13.488611" location="Kumbarakodu Reserve Forest of Sringeri" longitude="75.19778" stateProvince="Karnataka" typeStatus="Holotype">
<typeStatus id="F8BFBDFC4D2E686F53BC7D165D9A15CF">Holotype</typeStatus>
. Worker,
<collectingCountry id="97B7E43E3F9B9C9E08ACAB09CB0155B6">India</collectingCountry>
,
<collectingRegion id="B091785578A7E50F976183D566BBE9E3">Karnataka</collectingRegion>
,
<location id="FBA916DA9B359C1314B2D2A13634269A" LSID="urn:lsid:plazi:treatment:B76E28ECAC110815A33009D1F8700872:FBA916DA9B359C1314B2D2A13634269A" country="India" latitude="13.488611" longitude="75.19778" name="Kumbarakodu Reserve Forest of Sringeri" stateProvince="Karnataka">Kumbarakodu Reserve Forest of Sringeri</location>
(
<geoCoordinate id="22BB6F94F558F02CF4E3DE41C05E4C60" direction="north" orientation="latitude" value="13.488611">13°29'19'' N</geoCoordinate>
and
<geoCoordinate id="97B54FFDF68640FA9283B267DB952430" direction="east" orientation="longitude" value="75.19778">75°11'52'' E</geoCoordinate>
).
<collectingDate id="F4ED9659D276DA04794B4FE20AC1600F" day="26" month="03" value="2001-03-26" year="2001">26. iii. 2001</collectingDate>
.
<collectorName id="2BDD5A63E5C0285159ECB0F39EA77CEB">P.A. Sinu</collectorName>
. Collected from evergreen forest leaf litter using
<colletingMethod id="0747D88CB124C7D1CBC9CEBC107022D3">pitfall trap</colletingMethod>
.
</materialsCitation>
</paragraph>
<paragraph id="C5CDECCF808D059F41B714B74A604083" pageNumber="3">
<materialsCitation id="D2082A1912C3B41BD69CEF4ED3CB5F40" ID-GBIF-Occurrence="923893327" collectingDate="2001-03-26" collectingMethod="pitfall trap" collectorName="P.A. Sinu" country="India" latitude="13.488611" location="Kumbarakodu Reserve Forest of Sringeri" longitude="75.19778" stateProvince="Karnataka" typeStatus="Paratypes">
<typeStatus id="DD02DB44D6D76DCBD713627D1530EC0F">Paratypes</typeStatus>
. 6 workers, data same as holotype.
</materialsCitation>
</paragraph>
<paragraph id="E0357AA8943BA41D2CFAD58B29957336" pageNumber="3">Type repository. Holotype will be deposited at the Museum of the Zoological Survey of India, Calcutta and the paratypes will be deposited at the British Museum of Natural History and Insect Museum of Ashoka Trust for Research in Ecology and the Environment (ATREE), Bangalore, Karnataka, India.</paragraph>
</subSubSection>
<subSubSection id="CBF46D3D929616F611FC80DDF20E0B1E" type="etymology">
<paragraph id="E7480E5596459217AE022AD1607D6B7C" pageNumber="3">Etymology. The species name sringerensis indicates its type locality.</paragraph>
</subSubSection>
<subSubSection id="264458E846950D5686A820BA2E826DD5" type="discussion">
<paragraph id="FBD0BB70D2B18CAAE49B98E475A4F457" pageNumber="3">Remarks</paragraph>
<paragraph id="6BE1E361EA405D0F93EFEAB9DDAB7403" pageNumber="3">
This species is similar to one Malaysian species,
<taxonomicName id="63278D305EB88B35A46849EC562D9F29" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:28609" class="Insecta" family="Formicidae" genus="Discothyrea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-HNS="Discothyrea bryanti (Wheeler)" order="Hymenoptera" pageNumber="3" phylum="Arthropoda" rank="species" species="bryanti">Discothyrea bryanti Wheeler</taxonomicName>
, and one Australian species,
<taxonomicName id="9934A5EC026BB10E0EB6F1C0904FED3F" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:28633" class="Insecta" family="Formicidae" genus="Discothyrea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-HNS="Discothyrea velutina (Wheeler)" order="Hymenoptera" pageNumber="3" phylum="Arthropoda" rank="species" species="velutina">Discothyrea velutina</taxonomicName>
, in the following characteristics: (1) Antennae with 10 segments, and (2) absence of spur in middle tibiae.
<taxonomicName id="82124B729A3F772A63ABED68ADC1FD05" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:184276" class="Insecta" family="Formicidae" genus="Discothyrea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-HNS="Discothyrea sringerensis Zacharias &amp; Rajan" order="Hymenoptera" pageNumber="3" phylum="Arthropoda" rank="species" species="sringerensis">Discothyrea sringerensis</taxonomicName>
is similar to
<taxonomicName id="144C782EF71F66E4EA59EA6BD9E10E02" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:28609" class="Insecta" family="Formicidae" genus="Discothyrea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-HNS="Discothyrea bryanti (Wheeler)" order="Hymenoptera" pageNumber="3" phylum="Arthropoda" rank="species" species="bryanti">D. bryanti</taxonomicName>
in the following characters: (1) petiole with a prominent compressed translucent tooth on its ventral surface, and (2) more distinct propodeal teeth. However, the Indian species differs from
<taxonomicName id="687267728C23EA6CD234C22EC21E0E4F" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:28609" class="Insecta" family="Formicidae" genus="Discothyrea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-HNS="Discothyrea bryanti (Wheeler)" order="Hymenoptera" pageNumber="3" phylum="Arthropoda" rank="species" species="bryanti">D. bryanti</taxonomicName>
in (1) the funiculus being distinctly longer than the antennal scape (Fig. 2), (2) the petiole being less than four times as broad as long, and (3) the cheeks without a prominent blunt tooth in front of the eyes.
<taxonomicName id="7706FF651662979C5E15A787228AF9BF" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:184276" class="Insecta" family="Formicidae" genus="Discothyrea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-HNS="Discothyrea sringerensis Zacharias &amp; Rajan" order="Hymenoptera" pageNumber="3" phylum="Arthropoda" rank="species" species="sringerensis">Discothyrea sringerensis</taxonomicName>
shows some affinity to
<taxonomicName id="79EACD822B8422A72D462564DB9F8752" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:28633" class="Insecta" family="Formicidae" genus="Discothyrea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-HNS="Discothyrea velutina (Wheeler)" order="Hymenoptera" pageNumber="3" phylum="Arthropoda" rank="species" species="velutina">D. velutina</taxonomicName>
in the following characters: (1) cheeks without a blunt tooth in front of eyes, and (2) funiculus longer than scape. However, it differs from
<taxonomicName id="B63562A9C400959907705B8D463961FF" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:28633" class="Insecta" family="Formicidae" genus="Discothyrea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-HNS="Discothyrea velutina (Wheeler)" order="Hymenoptera" pageNumber="3" phylum="Arthropoda" rank="species" species="velutina">D. velutina</taxonomicName>
in the following characters: (1) distinct propodeal teeth, (2) petiole with a prominent compressed translucent tooth on its ventral surface, and (3) palp formula 5,4 (
<taxonomicName id="B3891BC91610A9ABD0872675BAC569D9" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:28633" class="Insecta" family="Formicidae" genus="Discothyrea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-HNS="Discothyrea velutina (Wheeler)" order="Hymenoptera" pageNumber="3" phylum="Arthropoda" rank="species" species="velutina">D. velutina</taxonomicName>
has a palp formula of 4,4). The maxillary and labial palps are variously segmented, according to species (Brown 1958). The palp formula of
<taxonomicName id="25D1A41C56595F6E89A61F1E80A5BA7A" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:28609" class="Insecta" family="Formicidae" genus="Discothyrea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-HNS="Discothyrea bryanti (Wheeler)" order="Hymenoptera" pageNumber="3" phylum="Arthropoda" rank="species" species="bryanti">D. bryanti</taxonomicName>
is unavailable.
</paragraph>
</subSubSection>
</treatment>
</document>

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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.199.3245" ID-GBIF-Dataset="e5a9e1c9-4a18-4224-9192-d41fa6200a99" ID-PMC="PMC3368282" ID-Pensoft-Pub="1313-2970-199-71" ID-PubMed="22711996" ModsDocAuthor="" ModsDocDate="2012" ModsDocID="1313-2970-199-71" ModsDocOrigin="ZooKeys 199" ModsDocTitle="A revision of the genus Kaszabister Mazur (Histeridae, Histerinae, Exosternini)" checkinTime="1451248970292" checkinUser="pensoft" docAuthor="Degallier, Nicolas, Mazur, Slawomir, Tishechkin, Alexey K. &amp; Caterino, Michael S." docDate="2012" docId="B76E34DE15C07FBBA117B4DCAEF0465E" docLanguage="en" docName="ZooKeys 199: 71-89" docOrigin="ZooKeys 199" docSource="http://dx.doi.org/10.3897/zookeys.199.3245" docTitle="Kaszabister Mazur 1972" docType="treatment" docVersion="3" lastPageNumber="73" masterDocId="F94CB3646F5CFFB58D31FFDEFFC64B13" masterDocTitle="A revision of the genus Kaszabister Mazur (Histeridae, Histerinae, Exosternini)" masterLastPageNumber="89" masterPageNumber="71" pageNumber="72" updateTime="1668153980577" updateUser="ExternalLinkService">
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<treatment ID-GBIF-Taxon="152035996" LSID="urn:lsid:plazi:treatment:B76E34DE15C07FBBA117B4DCAEF0465E" httpUri="http://treatment.plazi.org/id/B76E34DE15C07FBBA117B4DCAEF0465E" lastPageId="3" lastPageNumber="73" pageId="1" pageNumber="72">
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<paragraph pageId="1" pageNumber="72">
Genus
<taxonomicName LSID="http://species-id.net/wiki/Kaszabister" authority="Mazur, 1972" authorityName="Mazur" authorityYear="1972" class="Insecta" family="Histeridae" genus="Kaszabister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaszabister" order="Coleoptera" pageId="1" pageNumber="72" phylum="Arthropoda" rank="genus">Kaszabister Mazur, 1972</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="72" type="reference_group">
<paragraph pageId="1" pageNumber="72">
<taxonomicName class="Insecta" family="Histeridae" genus="Kaszabister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaszabister" order="Coleoptera" pageId="1" pageNumber="72" phylum="Arthropoda" rank="genus">Kaszabister</taxonomicName>
Mazur, 1972: 189.
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="72" type="type species">
<paragraph pageId="1" pageNumber="72">Type species.</paragraph>
<paragraph pageId="1" pageNumber="72">
<taxonomicName class="Insecta" family="Histeridae" genus="Kaszabister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaszabister mahunkai" order="Coleoptera" pageId="1" pageNumber="72" phylum="Arthropoda" rank="species" species="mahunkai">Kaszabister mahunkai</taxonomicName>
Mazur, 1972 (now regarded as a junior synonym of
<taxonomicName class="Insecta" family="Histeridae" genus="Kaszabister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaszabister rubellus" order="Coleoptera" pageId="1" pageNumber="72" phylum="Arthropoda" rank="species" species="rubellus">Kaszabister rubellus</taxonomicName>
(Erichson, 1834)), original designation.
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="72" type="diagnosis">
<paragraph pageId="1" pageNumber="72">Diagnosis.</paragraph>
<paragraph pageId="1" pageNumber="72">
<taxonomicName class="Insecta" family="Histeridae" genus="Kaszabister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaszabister" order="Coleoptera" pageId="1" pageNumber="72" phylum="Arthropoda" rank="genus">Kaszabister</taxonomicName>
can be easily separated from other Neotropical
<taxonomicName lsidName="" pageId="1" pageNumber="72" rank="tribe" tribe="Exosternini">Exosternini</taxonomicName>
by its strongly carinate frontal stria (Fig. 3); epipleural, subhumeral, and dorsal elytral striae 1 apically carinate and convergent to posterolateral corner (Fig. 1B, 2A); and narrow, edentate meso- and metatibiae which bear only a few small spines (Figs 1B, 2A, 4). The narrowly depressed lateral pronotal margin is also rare in other genera (Figs 1A, 2B).
</paragraph>
</subSubSection>
<subSubSection lastPageId="2" lastPageNumber="73" pageId="1" pageNumber="72" type="description">
<paragraph pageId="1" pageNumber="72">Description.</paragraph>
<paragraph lastPageId="2" lastPageNumber="73" pageId="1" pageNumber="72">
Body length 1.7-2.3mm, width 1.3-1.7mm, oval or oblong, more or less convex, reddish brown, glabrous. Head: Frons bordered by a prominent, moderately to strongly carinate frontal stria; antennae inserted under the rim of the frons in front of eyes; antennal scape slightly setose; antennal club oval, tomentose, lacking sutures or annuli, with small oval subapical sensoria on upper and lower surfaces; epistoma flat to convex, bordered by striae or carinae; labrum short, broad, rounded at sides and emarginate at middle; mandibles with strong furrows along lower outer margins and very weak subapical teeth on incisor edge; gena setose and weakly depressed; gular sutures impressed; submentum with numerous fringed setae, projecting slightly between maxillar bases; mentum about one-fourth broader than long, sides rounded, tapering apically, margin faintly emarginate; palpi relatively short, with truncate apices. Pronotum: pronotum widest at base, sides rounded, anterior angles acute; prescutellar impression absent; gland openings annulate, situated about one-third from anterior margin, behind inner edge of eye on each side; with 3 pores along each side; marginal stria complete, continuous with anterior marginal stria; lateral stria absent. Elytra: Dorsal striae of elytra simple or carinate, variously abbreviated; dorsal stria 1, subhumeral striae, and epipleural stria carinate and convergent apically. Prosternum: Antennal cavities of the prosternum visible in ventral view, located in the anterior angles of pronotum; prosternal lobe short, broad, reaching hypomeron laterally, with marginal stria at least medially; base of prosternal keel weakly emarginate; with complete carinate striae diverging
<pageBreakToken pageId="2" pageNumber="73" start="start">anterad</pageBreakToken>
and posterad, not joined. Mesoventrite: Disk flat, weakly projecting at middle, with complete marginal stria; mesometaventral stria present, angulate forward onto mesoventral disk. Metaventrite: Metaventral disk with postcoxal and lateral striae, both extending laterad toward metepisternum. Abdomen: Propygidium short, moderately convex, with annulate gland openings in anterolateral corners; pygidium lacking apical stria; abdominal ventrite 1 with one or two lateral striae; ventrites 2-5 with or without posterior marginal striae. Legs: Protrochanter with seta; protibia lacking teeth, but with 8-10 stout marginal spines; protibial spurs short, strong; protarsus with fine ventral spines, pretarsal claws simple and equal; meso- and metatrochanters lacking setae; meso- and metatibiae nearly parallel-sided, the former with few weak marginal spines; meso- and metatarsomeres with single pair of apicoventral setae. Male: Eighth tergite with accessory sclerites, shallowly narrowly incised at subtruncate apex, with basal membrane attachment distad basal emargination; ventral apodemes of 8th tergite broadly rounded, not meeting at midline; 8th sternite approximately parallel-sided, halves not joined along midline, with apical guides gradually more strongly elevated toward apices; 9th tergite with median emargination deep, ventral apodemes situated just behind midpoint, strongly toothed; spiculum gastrale (9th sternite) narrowed in distal two-thirds, with thin apical arms and short median apical flanges; halves of 10th tergite well developed, separated along midline. Female: Eighth tergite forming a single, apically emarginate plate; 8th sternite divided into two lateral plates, with thin, separate basal baculi which are articulated with the disk of S8; 9th sternite present, elongate, connected to apex of S8; tenth tergite present, without basal apodemes; valvifers elongate, enlarged basally; coxites with two strong and one weak inner tooth; gonostyle present, free, setose; bursa copulatrix small; spermatheca short, sclerotized, forming a ventral concave disk over oviduct; spermathecal gland attached at base of spermatheca, elongate, gradually expanded to apex.
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="73" type="distribution">
<paragraph pageId="2" pageNumber="73">Distribution.</paragraph>
<paragraph pageId="2" pageNumber="73">The distribution of the species of this genus is interestingly discontinuous, with three species concentrated in subtropical South America, and a single species from Central America, with few records from the northern half of South America.</paragraph>
</subSubSection>
<subSubSection lastPageId="3" lastPageNumber="74" pageId="2" pageNumber="73" type="key to species">
<paragraph pageId="2" pageNumber="73">Key to species</paragraph>
<paragraph lastPageId="3" lastPageNumber="74" pageId="2" pageNumber="73">
<table lastPageId="3" lastPageNumber="74" pageId="2" pageNumber="73">
<tr pageId="2" pageNumber="73">
<td colspan="1" pageId="2" pageNumber="73" rowspan="1">Fig. 4AFig. 4AFig. 3A</td>
<td colspan="1" pageId="2" pageNumber="73" rowspan="1">
<taxonomicName class="Insecta" family="Histeridae" genus="Kaszabister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaszabister barrigai" order="Coleoptera" pageId="2" pageNumber="73" phylum="Arthropoda" rank="species" species="barrigai">Kaszabister barrigai</taxonomicName>
</td>
</tr>
<tr pageId="2" pageNumber="73">
<td colspan="1" pageId="2" pageNumber="73" rowspan="1">Fig. 4B, DFig. 4C</td>
</tr>
<tr pageId="2" pageNumber="73">
<td colspan="1" pageId="2" pageNumber="73" rowspan="1">Fig. 4D</td>
<td colspan="1" pageId="2" pageNumber="73" rowspan="1">
<taxonomicName class="Insecta" family="Histeridae" genus="Kaszabister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaszabister carinatus" order="Coleoptera" pageId="2" pageNumber="73" phylum="Arthropoda" rank="species" species="carinatus">Kaszabister carinatus</taxonomicName>
</td>
</tr>
<tr pageId="2" pageNumber="73">
<td colspan="1" pageId="2" pageNumber="73" rowspan="1">Figs 4A-C</td>
</tr>
<tr pageId="3" pageNumber="74">
<td colspan="1" pageId="3" pageNumber="74" rowspan="1">
<pageBreakToken pageId="3" pageNumber="74" start="start">Fig</pageBreakToken>
. 3BFig. 4B
</td>
<td colspan="1" pageId="3" pageNumber="74" rowspan="1">
<taxonomicName class="Insecta" family="Histeridae" genus="Kaszabister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaszabister ferrugineus" order="Coleoptera" pageId="3" pageNumber="74" phylum="Arthropoda" rank="species" species="ferrugineus">Kaszabister ferrugineus</taxonomicName>
</td>
</tr>
<tr pageId="3" pageNumber="74">
<td colspan="1" pageId="3" pageNumber="74" rowspan="1">Fig. 3CFig. 4C</td>
<td colspan="1" pageId="3" pageNumber="74" rowspan="1">
<taxonomicName class="Insecta" family="Histeridae" genus="Kaszabister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaszabister rubellus" order="Coleoptera" pageId="3" pageNumber="74" phylum="Arthropoda" rank="species" species="rubellus">Kaszabister rubellus</taxonomicName>
</td>
</tr>
</table>
</paragraph>
</subSubSection>
</treatment>
</document>

358
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<document ID-DOI="10.1371/journal.pone.0041744" ID-GBIF-Dataset="eb1f92d6-b231-49e8-942a-23a304304304" ID-PMC="PMC3440430" ID-PubMed="22984399" ID-Zenodo-Dep="4244560" approvedBy="poa" checkinTime="1604502670596" checkinUser="tatiana" docAuthor="Peter J. Taylor, Samantha Stoffberg, Ara Monadjem, Martinus Corrie Schoeman, Julian Baylis &amp; Fenton P. D. Cotterill" docDate="2012" docId="B76E487AFFE0FFD4FC81034FFDA913A8" docLanguage="en" docName="PLoSOne.7.9.e41744.pdf.imf" docOrigin="PLoS ONE 7 (9)" docStyle="DocumentStyle{}" docTitle="Rhinolophus cohenae Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill 2012, new species" docType="treatment" docVersion="6" lastPageNumber="17" masterDocId="4B573002FFEFFFC4FFC9012BFF941755" masterDocTitle="Four New Bat Species (Rhinolophus hildebrandtii Complex) Reflect Plio-Pleistocene Divergence of Dwarfs and Giants across an Afromontane Archipelago" masterLastPageNumber="23" masterPageNumber="1" pageNumber="16" updateTime="1668143260826" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title>Four New Bat Species (Rhinolophus hildebrandtii Complex) Reflect Plio-Pleistocene Divergence of Dwarfs and Giants across an Afromontane Archipelago</mods:title>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Peter J. Taylor</mods:namePart>
<mods:affiliation>Department of Ecology and Resource Management, University of Venda, Thohoyandou, South Africa, Durban Natural Science Museum, Durban, South Africa, School of Life Sciences, University of KwaZulu-Natal, Durban, South Africa</mods:affiliation>
<mods:nameIdentifier type="email">peter.taylor@univen.ac.za</mods:nameIdentifier>
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<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Samantha Stoffberg</mods:namePart>
<mods:affiliation>Evolutionary Genomics Group, Department of Botany and Zoology, University of Stellenbosch, Stellenbosch, South Africa</mods:affiliation>
</mods:name>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ara Monadjem</mods:namePart>
<mods:affiliation>All Out Africa Research Unit, Department of Biological Sciences, University of Swaziland, Kwaluseni, Swaziland</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Martinus Corrie Schoeman</mods:namePart>
<mods:affiliation>School of Life Sciences, University of KwaZulu-Natal, Durban, South Africa</mods:affiliation>
</mods:name>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Julian Baylis</mods:namePart>
<mods:affiliation>Mulanje Mountain Conservation Trust, Mulanje, Malawi, Conservation Science Group, Department of Zoology, University of Cambridge, Cambridge, United Kingdom</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Fenton P. D. Cotterill</mods:namePart>
<mods:affiliation>Africa Earth Observatory Network, Geoecodynamics Research Hub, Department of Botany and Zoology, University of Stellenbosch, Stellenbosch, South Africa</mods:affiliation>
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<mods:number>7</mods:number>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.4328641" ID-GBIF-Taxon="168754889" ID-Zenodo-Dep="4328641" LSID="urn:lsid:zoobank.org:act:D0E1C9DD-7D36-4C01-AE0D-60AD474960A0" httpUri="http://treatment.plazi.org/id/B76E487AFFE0FFD4FC81034FFDA913A8" lastPageId="16" lastPageNumber="17" pageId="15" pageNumber="16">
<subSubSection pageId="15" pageNumber="16" type="nomenclature">
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<heading box="[840,1221,612,636]" fontSize="10" level="2" pageId="15" pageNumber="16" reason="1">
<taxonomicName authority="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill, 2012" authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[840,1074,612,636]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="cohenae" status="new species">
<emphasis box="[840,1074,612,636]" italics="true" pageId="15" pageNumber="16">Rhinolophus cohenae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1083,1221,612,636]" pageId="15" pageNumber="16" rank="species">new species</taxonomicNameLabel>
</heading>
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urn:lsid:zoobank.org:act:
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</paragraph>
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<subSubSection pageId="15" pageNumber="16" type="description">
<paragraph blockId="15.[840,1479,612,1473]" box="[864,1093,699,721]" pageId="15" pageNumber="16">Cohen’s Horseshoe Bat</paragraph>
<paragraph blockId="15.[840,1479,612,1473]" box="[864,1390,727,749]" pageId="15" pageNumber="16">
<figureCitation box="[864,948,727,749]" captionStart="Figure 4" captionStartId="8.[840,905,1377,1396]" captionTargetBox="[844,1474,166,1343]" captionTargetId="figure-696@8.[840,1478,162,1343]" captionTargetPageId="8" captionText="Figure 4. Morphometric variation in a series representing the R. hildebrandtii complex from Lutope-Ngolangola, Zimbabwe: a) biplot of forearm length versus noseleaf width and b) PCA of five craniometric variables (M3M3, CM3, IOC, NW, NH) in 26 individuals of known (37 or 46 kHz) and unknown (?) frequency. Females indicated by open circlesı males by closed circles or crosses or asterisk. Voucher specimens for molecular sequencing study indicated by asterisk (Clade 1e: = smithersii sp. nov.; see Taxonomic Conclusions) and crosses (Clade 2: = mossambicus sp. nov.; see Taxonomic Conclusions). Hereafterı all individuals with a frequency of 37 kHz were assumed to belong to Clade 2 (mossambicus sp. nov.) and the 46 kHz individual was assumed to belong to Clade 1e (smithersi sp. nov.). doi:10.1371/journal.pone.0041744.g004" figureDoi="http://doi.org/10.5281/zenodo.4244568" httpUri="https://zenodo.org/record/4244568/files/figure.png" pageId="15" pageNumber="16">Fig. 4, 4</figureCitation>
,
<figureCitation box="[960,1031,727,749]" captionStart="Figure 1" captionStartId="1.[155,220,1840,1859]" captionTargetBox="[158,790,800,1803]" captionTargetId="figure-761@1.[155,793,797,1806]" captionTargetPageId="1" captionText="Figure 1. Portraits of (a) Rhinolophus smithersi species novo, and (b) Rhinolophus mossambicus species novo, two of four new cryptic species described herein within the R. hildebrandtii complex. doi:10.1371/journal.pone.0041744.g001" figureDoi="http://doi.org/10.5281/zenodo.4244562" httpUri="https://zenodo.org/record/4244562/files/figure.png" pageId="15" pageNumber="16">Fig. S1</figureCitation>
,
<tableCitation box="[1043,1121,727,749]" captionStart="Table 3" captionStartId="7.[171,235,1322,1343]" captionText="Table 3. Summary of data for peak echolocation frequency of R. hildebrandtii s.l. populations." pageId="15" pageNumber="16">Table 3</tableCitation>
,
<tableCitation box="[1134,1148,727,749]" captionStart="Table 4" captionStartId="15.[856,920,1521,1542]" captionText="Table 4. Eigenvectors for PCA of recently collected voucher specimens of the Rhinolophus hildebrandtii complex associated with molecular and acoustic analysesı which includes type specimens of R. hildebrandtii and R. eloquens (see Fig. 5a). doi:10.1371/journal.pone.0041744.t004" pageId="15" pageNumber="16">4</tableCitation>
,
<tableCitation box="[1160,1250,727,749]" captionStart="Table 1" captionStartId="6.[171,235,1481,1502]" captionText="Table 1. ‘Uncorrected p’ pairwise sequence distances for the 12S gene below the diagonal and Chd1 gene above the diagonal for the clades (1ı 2ı 3) and some lineages (1aı 1bı 1c) identified in the molecular analyses of the Rhinolophus hildebrandtii complex." pageId="15" pageNumber="16">Table S1</tableCitation>
, Appendix S1
</paragraph>
</subSubSection>
<subSubSection pageId="15" pageNumber="16" type="materials_examined">
<paragraph blockId="15.[840,1479,612,1473]" pageId="15" pageNumber="16">
<materialsCitation ID-GBIF-Occurrence="2908237306" collectingDate="2004-09-28" collectionCode="DM" collectorName="L. Cohen" country="South Africa" elevation="690" latitude="-25.7188889" location="Mountainland Nature Reserve" longitude="031.2661111" municipality="Barberton" pageId="15" pageNumber="16" specimenCode="DM 8626" specimenCount="2" specimenCount-adult="1" specimenCount-male="1" stateProvince="Mpumalanga Province" typeStatus="holotype">
<emphasis bold="true" box="[863,967,756,775]" pageId="15" pageNumber="16">
<typeStatus box="[863,963,756,775]" pageId="15" pageNumber="16">Holotype</typeStatus>
.
</emphasis>
<specimenCode box="[992,1096,754,776]" pageId="15" pageNumber="16">DM 8626</specimenCode>
;
<specimenCount box="[1114,1163,754,776]" pageId="15" pageNumber="16" type="adult">adult</specimenCount>
<specimenCount box="[1177,1226,754,776]" pageId="15" pageNumber="16" type="male">male</specimenCount>
, alcohol skin, skull and prepared baculum (
<figureCitation box="[1032,1095,782,804]" captionStart="Figure 9" captionStartId="13.[155,220,1417,1436]" captionTargetBox="[178,1109,164,1382]" captionTargetId="figure-203@13.[155,1113,162,1383]" captionTargetPageId="13" captionText="Figure 9. Dorsal (D), ventral (V) and lateral (L) view of bacula (tips on right) from four individuals (a–d) from Mpumalanga (Clade 1a = cohenae sp. nov.), two (e–f) from lowland sites in Mozambique (Clade 2 = mossambicus sp. nov.) and one (g) from Mt Mabu in Mozambique (Clade 1b = mabuensis sp. nov.). a = DM 11558 (Sudwala); b = DM 11620 (Barberton Tunnel; Topotype of cohenae); c = DM 11560 (Mayo); d = DM 11618 (Barberton Tunnel); e = DM 8580 (Gorongosa); f = DM 8578 (Niassa GR; Holotype of mossambicus); g = DM 10842 (Mt Mabu; Holotype of mabuensis). Bacula of Clade 1a (cohenae sp. nov.) have spatulate tip (rounded in Clades 2 (mossambicus sp. nov.) and 1b (mabuensis sp. nov.))ı typically emarginated basal portion (less so in Clades 2 and 1b) and shaft laterally compressed (cylindrical in Clades 2 and 1b) and sloping downwards in lateral view (horizontal in Clades 2 and 1b). doi:10.1371/journal.pone.0041744.g009" figureDoi="http://doi.org/10.5281/zenodo.4244578" httpUri="https://zenodo.org/record/4244578/files/figure.png" pageId="15" pageNumber="16">Fig. 9f</figureCitation>
), collected by
<collectorName box="[1236,1331,782,804]" pageId="15" pageNumber="16">L. Cohen</collectorName>
,
<date pageId="15" pageNumber="16" value="2004-09-28">
<collectingDate pageId="15" pageNumber="16" value="2004-09-28">28 September 2004</collectingDate>
</date>
. Skull and mandible in good condition.
</materialsCitation>
</paragraph>
<paragraph blockId="15.[840,1479,612,1473]" pageId="15" pageNumber="16">
<emphasis bold="true" box="[863,1017,839,858]" pageId="15" pageNumber="16">Type locality.</emphasis>
<collectingMunicipality box="[1041,1141,837,859]" pageId="15" pageNumber="16">Barberton</collectingMunicipality>
,
<collectingCounty box="[1160,1474,837,859]" pageId="15" pageNumber="16">Mountainland Nature Reserve</collectingCounty>
,
<locationDeviation box="[840,1027,865,887]" pageId="15" pageNumber="16">
<quantity box="[840,902,865,887]" metricMagnitude="4" metricUnit="m" metricValue="6.8" pageId="15" pageNumber="16" unit="km" value="68.0">68 km</quantity>
SE Sudwala
</locationDeviation>
,
<collectingRegion box="[1038,1261,865,887]" country="South Africa" name="Mpumalanga" pageId="15" pageNumber="16">Mpumalanga Province</collectingRegion>
,
<collectingCountry box="[1271,1396,865,887]" name="South Africa" pageId="15" pageNumber="16" value="South Africa">South Africa</collectingCountry>
,
<geoCoordinate pageId="15" pageNumber="16">
25
<emphasis bold="true" box="[1430,1439,865,886]" pageId="15" pageNumber="16">°</emphasis>
<date pageId="15" pageNumber="16" value="1943-08-09">
43
<emphasis box="[1470,1478,865,887]" italics="true" pageId="15" pageNumber="16">'</emphasis>
8
</date>
<emphasis box="[852,860,892,914]" italics="true" pageId="15" pageNumber="16">&quot;</emphasis>
S
</geoCoordinate>
;
<geoCoordinate box="[887,1025,892,915]" pageId="15" pageNumber="16">
31
<emphasis bold="true" box="[911,920,893,914]" pageId="15" pageNumber="16">°</emphasis>
<date box="[928,992,892,915]" pageId="15" pageNumber="16" value="1958-09-15">
15
<emphasis box="[952,960,892,914]" italics="true" pageId="15" pageNumber="16">'</emphasis>
58
</date>
<emphasis box="[992,1000,892,914]" italics="true" pageId="15" pageNumber="16">&quot;</emphasis>
E
</geoCoordinate>
; elevation
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<elevation box="[1131,1194,893,915]" metricMagnitude="2" metricUnit="m" metricValue="6.9" pageId="15" pageNumber="16" unit="m" value="690.0">690 m</elevation>
</quantity>
asl.
</paragraph>
</subSubSection>
<subSubSection lastPageId="16" lastPageNumber="17" pageId="15" pageNumber="16" type="diagnosis">
<paragraph blockId="15.[840,1479,612,1473]" lastBlockId="16.[155,793,164,1277]" lastPageId="16" lastPageNumber="17" pageId="15" pageNumber="16">
<emphasis bold="true" box="[863,976,922,941]" pageId="15" pageNumber="16">Diagnosis.</emphasis>
Lowest recorded peak frequency (33.0 kHz in the
<typeStatus box="[840,926,949,971]" pageId="15" pageNumber="16">holotype</typeStatus>
; mean 32.8±0.24 kHz, n =
<quantity box="[1226,1273,949,971]" metricMagnitude="-1" metricUnit="m" metricValue="1.778" pageId="15" pageNumber="16" unit="in" value="7.0">7 in</quantity>
<typeStatus box="[1289,1330,949,971]" pageId="15" pageNumber="16">type</typeStatus>
series) which immediately distinguishes this species from all others. Noseleaf extremely wide,
<quantity box="[1002,1143,1004,1026]" metricMagnitude="-2" metricUnit="m" metricValue="1.49" metricValueMax="1.63" metricValueMin="1.35" pageId="15" pageNumber="16" unit="mm" value="14.9" valueMax="16.3" valueMin="13.5">13.5–16.3 mm</quantity>
(
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in
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), exceeding the previously described range (12–15) for the species [
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]. Body size and cranial size very large (
<tableCitation box="[1151,1228,1060,1082]" captionStart="Table 5" captionStartId="17.[153,217,487,508]" captionText="Table 5. Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the Rhinolophus hildebrandtii complex." pageId="15" pageNumber="16">Table 5</tableCitation>
; FL
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;
<typeStatus box="[1395,1478,1060,1082]" pageId="15" pageNumber="16">holotype</typeStatus>
<quantity box="[840,931,1088,1110]" metricMagnitude="-2" metricUnit="m" metricValue="6.59" pageId="15" pageNumber="16" unit="mm" value="65.9">65.9 mm</quantity>
; GSL
<quantity box="[998,1107,1088,1110]" metricMagnitude="-2" metricUnit="m" metricValue="2.95" metricValueMax="3.0" metricValueMin="2.9" pageId="15" pageNumber="16" unit="mm" value="29.5" valueMax="30.0" valueMin="29.0">29–30 mm</quantity>
; CCL
<quantity box="[1177,1285,1088,1110]" metricMagnitude="-2" metricUnit="m" metricValue="2.5" metricValueMax="2.6" metricValueMin="2.4" pageId="15" pageNumber="16" unit="mm" value="25.0" valueMax="26.0" valueMin="24.0">24–26 mm</quantity>
;
<typeStatus box="[1298,1381,1088,1110]" pageId="15" pageNumber="16">holotype</typeStatus>
<quantity box="[1390,1478,1088,1110]" metricMagnitude="-2" metricUnit="m" metricValue="2.9899999999999998" pageId="15" pageNumber="16" unit="mm" value="29.9">29.9 mm</quantity>
and
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respectively). In body and cranial size this species overlaps with
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[981,1064,1143,1165]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="mabuensis">
<emphasis box="[981,1064,1143,1165]" italics="true" pageId="15" pageNumber="16">mabuensis</emphasis>
</taxonomicName>
; however the latter species has a distinct echolocation frequency (
<emphasis box="[1076,1094,1171,1193]" italics="true" pageId="15" pageNumber="16">ca</emphasis>
38 kHz) and a distinct baculum (‘‘
<typeStatus box="[1425,1477,1172,1194]" pageId="15" pageNumber="16">Type</typeStatus>
2’’). Anterior upper premolar conspicuous and located within tooth row or entirely absent. In the
<typeStatus box="[1245,1328,1227,1249]" pageId="15" pageNumber="16">holotype</typeStatus>
and in three additional specimens, the anterior premolar is prominent and located within the toothrow. In another three specimens, the anterior premolar is absent (and the canine and posterior premolar in contact). In another specimen the prominent anterior premolar is situated in the toothrow on the right side but absent on the left side, leaving a small gap between canine and posterior premolar. In no cases were specimens found to have small anterior premolars situated external to the toothrow, thus distinguishing this species from
<taxonomicName authorityName="Peters" authorityYear="1878" box="[211,314,164,186]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" sensu="stricto" species="hildebrandtii">
<emphasis box="[211,314,164,186]" italics="true" pageId="16" pageNumber="17">hildebrandtii</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[324,352,164,186]" pageId="16" pageNumber="17" sensu="stricto">s.s.</taxonomicNameLabel>
[
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]. The baculum (total length
<quantity box="[688,764,164,186]" metricMagnitude="-3" metricUnit="m" metricValue="3.5" pageId="16" pageNumber="17" unit="mm" value="3.5">3.5 mm</quantity>
in
<typeStatus box="[155,238,192,214]" pageId="16" pageNumber="17">holotype</typeStatus>
) has a unique shape (‘‘
<typeStatus box="[465,517,192,214]" pageId="16" pageNumber="17">Type</typeStatus>
1’’;
<figureCitation box="[563,619,192,214]" captionStart="Figure 9" captionStartId="13.[155,220,1417,1436]" captionTargetBox="[178,1109,164,1382]" captionTargetId="figure-203@13.[155,1113,162,1383]" captionTargetPageId="13" captionText="Figure 9. Dorsal (D), ventral (V) and lateral (L) view of bacula (tips on right) from four individuals (a–d) from Mpumalanga (Clade 1a = cohenae sp. nov.), two (e–f) from lowland sites in Mozambique (Clade 2 = mossambicus sp. nov.) and one (g) from Mt Mabu in Mozambique (Clade 1b = mabuensis sp. nov.). a = DM 11558 (Sudwala); b = DM 11620 (Barberton Tunnel; Topotype of cohenae); c = DM 11560 (Mayo); d = DM 11618 (Barberton Tunnel); e = DM 8580 (Gorongosa); f = DM 8578 (Niassa GR; Holotype of mossambicus); g = DM 10842 (Mt Mabu; Holotype of mabuensis). Bacula of Clade 1a (cohenae sp. nov.) have spatulate tip (rounded in Clades 2 (mossambicus sp. nov.) and 1b (mabuensis sp. nov.))ı typically emarginated basal portion (less so in Clades 2 and 1b) and shaft laterally compressed (cylindrical in Clades 2 and 1b) and sloping downwards in lateral view (horizontal in Clades 2 and 1b). doi:10.1371/journal.pone.0041744.g009" figureDoi="http://doi.org/10.5281/zenodo.4244578" httpUri="https://zenodo.org/record/4244578/files/figure.png" pageId="16" pageNumber="17">Fig. 9</figureCitation>
) characterised by narrow, laterally compressed and ventrally deflected shaft, emarginated distal portion and spatulate tip, which is not found in other species of the
<taxonomicName authorityName="Peters" authorityYear="1878" box="[378,481,275,297]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[378,481,275,297]" italics="true" pageId="16" pageNumber="17">hildebrandtii</emphasis>
</taxonomicName>
complex in which the baculum has been described (
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[361,466,303,325]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="mossambicus">
<emphasis box="[361,466,303,325]" italics="true" pageId="16" pageNumber="17">mossambicus</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[524,607,303,325]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="mabuensis">
<emphasis box="[524,607,303,325]" italics="true" pageId="16" pageNumber="17">mabuensis</emphasis>
</taxonomicName>
). Genotypes of
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="cohenae">
<emphasis italics="true" pageId="16" pageNumber="17">R.cohenae</emphasis>
</taxonomicName>
are members of Clade 1a (
<figureCitation box="[488,544,332,354]" captionStart="Figure 2" captionStartId="2.[155,220,1881,1900]" captionTargetBox="[160,1475,167,1843]" captionTargetId="figure-0@2.[155,1480,162,1848]" captionTargetPageId="2" captionText="Figure 2. Map of southern, central and eastern Africa indicating localities of individuals of R. hildebrandtii species-complex included in this study. Grey-shaded area represents elevations in excess of 600 m a.s.l. Closed squares indicate museum specimens from which craniometric data were obtained. Open symbols indicate specimens genotyped in this study. The distribution of the three major clades is based on cytochrome b (see Figure 2): open circles = Clade 1; open squares = Clade 2; open diamonds = Clade 3. Closed squares enclosed in open symbols indicate localities where both molecular and morphological data were available for selected specimens. Numbers refer to respective localities listed in Table S1. ‘‘T’’ indicates the type localities of R. eloquens in Uganda and R. hildebrandtii in Kenyaı respectively. doi:10.1371/journal.pone.0041744.g002" figureDoi="http://doi.org/10.5281/zenodo.4244564" httpUri="https://zenodo.org/record/4244564/files/figure.png" pageId="16" pageNumber="17">Fig. 2</figureCitation>
).
</paragraph>
<paragraph blockId="16.[155,793,164,1277]" pageId="16" pageNumber="17">
<materialsCitation ID-GBIF-Occurrence="2908237301" accessionNumber="JQ929270, JQ929206, JQ929230, JQ929287" collectingDate="2004-09-27" collectingDateMax="2009-09-27" collectingDateMin="2004-09-27" collectionCode="DM" collectorName="L. Cohen &amp; S. Stoffberg" country="South Africa" location="Mountainland Nature Reserve" municipality="Barberton" pageId="16" pageNumber="17" specimenCode="DM 7886" specimenCount="1" specimenCount-adult="1" specimenCount-male="1" typeStatus="paratype">
<emphasis bold="true" box="[177,292,361,380]" pageId="16" pageNumber="17">
<typeStatus box="[177,287,361,380]" pageId="16" pageNumber="17">Paratypes</typeStatus>
.
</emphasis>
<specimenCode box="[316,417,360,382]" pageId="16" pageNumber="17">DM 7886</specimenCode>
(
<specimenCount box="[436,484,360,382]" pageId="16" pageNumber="17" type="adult">adult</specimenCount>
<specimenCount box="[497,546,360,382]" pageId="16" pageNumber="17" type="male">male</specimenCount>
, alcohol skin and skull, collected by
<collectorName box="[278,371,388,410]" pageId="16" pageNumber="17">L. Cohen</collectorName>
on
<date box="[411,602,388,410]" pageId="16" pageNumber="17" value="2004-09-27">
<collectingDate box="[411,602,388,410]" pageId="16" pageNumber="17" value="2004-09-27">27 September 2004</collectingDate>
</date>
from type locality),
</materialsCitation>
<materialsCitation ID-GBIF-Occurrence="2908237308" accessionNumber="JQ929265, JQ929215, JQ929235, JQ929293" box="[155,261,416,438]" collectingDate="2004-09-27" collectingDateMax="2009-09-27" collectingDateMin="2004-09-27" collectionCode="DM" collectorName="L. Cohen &amp; S. Stoffberg" country="South Africa" municipality="Barberton" pageId="16" pageNumber="17" specimenCode="DM11620" specimenCount="1" specimenCount-adult="1" specimenCount-male="1" typeStatus="paratype">
<specimenCode box="[155,258,416,438]" pageId="16" pageNumber="17">DM11620</specimenCode>
,
</materialsCitation>
<materialsCitation ID-GBIF-Occurrence="2908237310" collectingDate="2004-09-27" collectingDateMax="2009-09-27" collectingDateMin="2004-09-27" collectionCode="DM" collectorName="L. Cohen &amp; S. Stoffberg" country="South Africa" pageId="16" pageNumber="17" specimenCode="DM 11919, DM 11918" specimenCount="2" specimenCount-adult="2" specimenCount-male="2" typeStatus="paratype">
<specimenCode box="[268,378,416,438]" pageId="16" pageNumber="17">DM 11919</specimenCode>
,
<specimenCode box="[389,497,416,438]" pageId="16" pageNumber="17">DM 11918</specimenCode>
(
<specimenCount box="[511,559,416,438]" pageId="16" pageNumber="17" type="adult">adult</specimenCount>
<specimenCount box="[566,623,416,438]" pageId="16" pageNumber="17" type="male">males</specimenCount>
, alcohol skin and skull, collected by
<collectorName box="[341,458,444,466]" pageId="16" pageNumber="17">S. Stoffberg</collectorName>
and
<collectorName box="[514,610,444,466]" pageId="16" pageNumber="17">L. Cohen</collectorName>
on
<date pageId="16" pageNumber="17" value="2009-09-27">
<collectingDate pageId="16" pageNumber="17" value="2009-09-27">27 September 2009</collectingDate>
</date>
)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="description">
<paragraph blockId="16.[155,793,164,1277]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[177,311,501,520]" pageId="16" pageNumber="17">Description.</emphasis>
Dorsal and ventral colour similar to
<taxonomicName authorityName="Peters" authorityYear="1878" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" sensu="stricto" species="hildebrandtii">
<emphasis italics="true" pageId="16" pageNumber="17">R. hildebrandtii</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[277,306,528,550]" pageId="16" pageNumber="17" sensu="stricto">s.s.</taxonomicNameLabel>
, but larger in noseleaf, skull and external dimensions as described above. The noseleaf shows the typical low, rounded shape of the connecting process in lateral view, in the
<typeStatus box="[155,238,612,634]" pageId="16" pageNumber="17">holotype</typeStatus>
(
<figureCitation box="[252,319,612,634]" captionStart="Figure 8" captionStartId="12.[155,220,993,1012]" captionTargetBox="[163,1000,167,956]" captionTargetId="figure-464@12.[155,1007,162,960]" captionTargetPageId="12" captionText="Figure 8. Photographs showing lateral views of noseleafs of selected individuals (including holotypes of new species) of the Rhinolophus hildebrandtii complex representing molecular Lineage 1a (= cohenae sp. nov.; a–b), Lineage 1b (= mabuensis sp. nov.; c) and Clade 2 (= mossambicus sp. nov.; d–f). a = DM 7886 (cohenae sp. nov.;Barberton Tunnel, Mpumalanga Province, South Africa); b = DM 8626 (cohenae sp. nov.; Barberton Tunnel, Mpumalanga Province, South Africa; Holotype); c = DM 10842 (mabuensis sp. nov.; Mt Mabu, Mozambique; Holotype); d = DM 8578 (mossambicus sp. nov.; Niassa Game Reserve, Mozambique; Holotype); e = DM 8579 (mossambicus sp. nov.; Chinizuia, Mozambique); f = DM 8577 (mossambicus sp. nov.; Namapa, Mozambique). doi:10.1371/journal.pone.0041744.g008" figureDoi="http://doi.org/10.5281/zenodo.4244576" httpUri="https://zenodo.org/record/4244576/files/figure.png" pageId="16" pageNumber="17">Fig. 8b</figureCitation>
) and in other specimens examined (e.g.
<figureCitation box="[714,781,612,634]" captionStart="Figure 8" captionStartId="12.[155,220,993,1012]" captionTargetBox="[163,1000,167,956]" captionTargetId="figure-464@12.[155,1007,162,960]" captionTargetPageId="12" captionText="Figure 8. Photographs showing lateral views of noseleafs of selected individuals (including holotypes of new species) of the Rhinolophus hildebrandtii complex representing molecular Lineage 1a (= cohenae sp. nov.; a–b), Lineage 1b (= mabuensis sp. nov.; c) and Clade 2 (= mossambicus sp. nov.; d–f). a = DM 7886 (cohenae sp. nov.;Barberton Tunnel, Mpumalanga Province, South Africa); b = DM 8626 (cohenae sp. nov.; Barberton Tunnel, Mpumalanga Province, South Africa; Holotype); c = DM 10842 (mabuensis sp. nov.; Mt Mabu, Mozambique; Holotype); d = DM 8578 (mossambicus sp. nov.; Niassa Game Reserve, Mozambique; Holotype); e = DM 8579 (mossambicus sp. nov.; Chinizuia, Mozambique); f = DM 8577 (mossambicus sp. nov.; Namapa, Mozambique). doi:10.1371/journal.pone.0041744.g008" figureDoi="http://doi.org/10.5281/zenodo.4244576" httpUri="https://zenodo.org/record/4244576/files/figure.png" pageId="16" pageNumber="17">Fig. 8a</figureCitation>
). Lower lip with single mental groove. The robust skull is represented in the mean cranial dimensions (
<tableCitation box="[602,680,668,690]" captionStart="Table 5" captionStartId="17.[153,217,487,508]" captionText="Table 5. Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the Rhinolophus hildebrandtii complex." pageId="16" pageNumber="17">Table 5</tableCitation>
;
<figureCitation box="[693,751,668,690]" captionStart="Figure 5" captionStartId="9.[155,220,1403,1422]" captionTargetBox="[159,791,166,1365]" captionTargetId="figure-699@9.[155,795,162,1369]" captionTargetPageId="9" captionText="Figure 5. Canonical variates analysis (CVA) (a) of 10 cranial variables in five groups of the Rhinolophus hildebrandtii complex defined by molecular analysis; and PCA (b) of five cranial variables for sample in (a) with type series of hildebrandtii (‘‘H¹’’) and eloquens (‘‘E¹’’) added. Open circles = Clade 1a (= cohenae sp. nov.); closed circles = Clade 1b (= mabuensis sp. nov.); shaded circles = Clade 1d (= smithersi sp. nov.; Pafuri); asterisk enclosed in circle = Clade 1e (= smithersi sp. nov.; Zimbabwe); open squares = Clade 2 (mossambicus sp. nov.; Mozambique); shaded squares = Clade 2 (mossambicus sp. nov.; Lutopeı Zimbabwe); open diamonds = R. eloquens type series (Clade 3); crosses in circles = R. hildebrandtii type and co-type (Clade 1c). doi:10.1371/journal.pone.0041744.g005" figureDoi="http://doi.org/10.5281/zenodo.4244570" httpUri="https://zenodo.org/record/4244570/files/figure.png" pageId="16" pageNumber="17">Fig. 5</figureCitation>
). In lateral view the skull has a prominent rostral chamber that extends to the same height as the sagittal crest; albeit the cranium is distinctly elongated and flattened in lateral profile (
<figureCitation box="[687,748,751,773]" captionStart="Figure 6" captionStartId="10.[155,220,1294,1313]" captionTargetBox="[159,1100,166,1256]" captionTargetId="figure-302@10.[155,1113,162,1260]" captionTargetPageId="10" captionText="Figure 6. Relative warps analysis (RWA) of 13 dorsal cranial landmarks from 22 individuals of R. hildebrandtii s.l. belonging to two molecular clades and two lineages of Clade 1 (see Fig. 3). Revised taxon names are provided in parentheses (see Taxonomic Conclusions). Skulls which were included in this analysis are indicated in Table S1. Symbols as in Fig. 5. Thin plate splines (grids) show landmark distortions represented by extremes of variation on RW1 (left = negative; right = positive) and RW2 (bottom = negative; top = positive) axes. The two skull photographs at the bottom are of actual specimens representing the negative (left: TM 41997ı smithersi from Pafuri) and positive (right: DM 11560ı cohenae from Mayoı Mpumalanga Province) extremes of variation on RW1. Landmark positions (filled circles) are shown in the photograph in the centre. doi:10.1371/journal.pone.0041744.g006" figureDoi="http://doi.org/10.5281/zenodo.4244572" httpUri="https://zenodo.org/record/4244572/files/figure.png" pageId="16" pageNumber="17">Fig. 6</figureCitation>
). In dorsal view the V-shaped inter-orbital basin formed by the supraorbital ridges is very prominent, with the anterior root of the sagittal crest displaced posteriorly (
<figureCitation box="[496,552,835,857]" captionStart="Figure 6" captionStartId="10.[155,220,1294,1313]" captionTargetBox="[159,1100,166,1256]" captionTargetId="figure-302@10.[155,1113,162,1260]" captionTargetPageId="10" captionText="Figure 6. Relative warps analysis (RWA) of 13 dorsal cranial landmarks from 22 individuals of R. hildebrandtii s.l. belonging to two molecular clades and two lineages of Clade 1 (see Fig. 3). Revised taxon names are provided in parentheses (see Taxonomic Conclusions). Skulls which were included in this analysis are indicated in Table S1. Symbols as in Fig. 5. Thin plate splines (grids) show landmark distortions represented by extremes of variation on RW1 (left = negative; right = positive) and RW2 (bottom = negative; top = positive) axes. The two skull photographs at the bottom are of actual specimens representing the negative (left: TM 41997ı smithersi from Pafuri) and positive (right: DM 11560ı cohenae from Mayoı Mpumalanga Province) extremes of variation on RW1. Landmark positions (filled circles) are shown in the photograph in the centre. doi:10.1371/journal.pone.0041744.g006" figureDoi="http://doi.org/10.5281/zenodo.4244572" httpUri="https://zenodo.org/record/4244572/files/figure.png" pageId="16" pageNumber="17">Fig. 6</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="distribution">
<paragraph blockId="16.[155,793,164,1277]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[177,317,864,883]" pageId="16" pageNumber="17">Distribution.</emphasis>
Known only from three closely-spaced localities in
<collectingRegion box="[225,449,891,913]" country="South Africa" name="Mpumalanga" pageId="16" pageNumber="17">Mpumalanga Province</collectingRegion>
of
<collectingCountry box="[488,615,891,913]" name="South Africa" pageId="16" pageNumber="17" value="South Africa">South Africa</collectingCountry>
in the vicinity of Nelspruit.
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="biology_ecology">
<paragraph blockId="16.[155,793,164,1277]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[177,267,949,968]" pageId="16" pageNumber="17">Ecology.</emphasis>
The
<typeStatus box="[344,385,947,969]" pageId="16" pageNumber="17">type</typeStatus>
locality of Barberton is located in the Savanna Biome [
<bibRefCitation author="Mucina L &amp; Rutherford MC" box="[335,375,975,997]" journalOrPublisher="Strelitzia 19, South African National Biodiversity Institute, Pretoria" pageId="16" pageNumber="17" refId="ref25379" refString="114. Mucina L, Rutherford MC (2006) The Vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19, South African National Biodiversity Institute, Pretoria." title="The Vegetation of South Africa, Lesotho and Swaziland" type="book" year="2006">114</bibRefCitation>
] at
<quantity box="[428,493,975,997]" metricMagnitude="2" metricUnit="m" metricValue="6.9" pageId="16" pageNumber="17" unit="m" value="690.0">690 m</quantity>
, close to the border of the Grassland Biome. Two nearby additional localities (Sudwala and Mayo Mines) are located in the Grassland Biome (Mesic Highveld Grassland) at altitudes between 900 and
<quantity box="[555,633,1059,1081]" metricMagnitude="3" metricUnit="m" metricValue="1.1" pageId="16" pageNumber="17" unit="m" value="1100.0">1100 m</quantity>
.
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="etymology">
<paragraph blockId="16.[155,793,164,1277]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[177,298,1089,1108]" pageId="16" pageNumber="17">Etymology.</emphasis>
We selected the specific epithet to recognize Lientjie Cohen who collected the
<typeStatus box="[536,577,1115,1137]" pageId="16" pageNumber="17">type</typeStatus>
specimen. She has contributed significantly to the conservation of bats in South Africa, particularly in Mpumalanga Province. The species name combines the surname Cohen and genitive singular case-ending ‘‘ae’’ indicative of feminine gender.
</paragraph>
<paragraph blockId="16.[155,793,164,1277]" box="[177,573,1255,1277]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[177,412,1256,1275]" pageId="16" pageNumber="17">Specimens examined.</emphasis>
<collectingRegion box="[436,470,1255,1277]" country="South Sudan" name="Lakes" pageId="16" pageNumber="17">See</collectingRegion>
<tableCitation box="[478,569,1255,1277]" captionStart="Table 1" captionStartId="6.[171,235,1481,1502]" captionText="Table 1. ‘Uncorrected p’ pairwise sequence distances for the 12S gene below the diagonal and Chd1 gene above the diagonal for the clades (1ı 2ı 3) and some lineages (1aı 1bı 1c) identified in the molecular analyses of the Rhinolophus hildebrandtii complex." pageId="16" pageNumber="17">Table S1</tableCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>

321
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<mods:title>Four New Bat Species (Rhinolophus hildebrandtii Complex) Reflect Plio-Pleistocene Divergence of Dwarfs and Giants across an Afromontane Archipelago</mods:title>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Peter J. Taylor</mods:namePart>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Samantha Stoffberg</mods:namePart>
<mods:affiliation>Evolutionary Genomics Group, Department of Botany and Zoology, University of Stellenbosch, Stellenbosch, South Africa</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ara Monadjem</mods:namePart>
<mods:affiliation>All Out Africa Research Unit, Department of Biological Sciences, University of Swaziland, Kwaluseni, Swaziland</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Martinus Corrie Schoeman</mods:namePart>
<mods:affiliation>School of Life Sciences, University of KwaZulu-Natal, Durban, South Africa</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Julian Baylis</mods:namePart>
<mods:affiliation>Mulanje Mountain Conservation Trust, Mulanje, Malawi, Conservation Science Group, Department of Zoology, University of Cambridge, Cambridge, United Kingdom</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Fenton P. D. Cotterill</mods:namePart>
<mods:affiliation>Africa Earth Observatory Network, Geoecodynamics Research Hub, Department of Botany and Zoology, University of Stellenbosch, Stellenbosch, South Africa</mods:affiliation>
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<mods:number>7</mods:number>
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<mods:identifier type="PubMed">22984399</mods:identifier>
<mods:identifier type="Zenodo-Dep">4244560</mods:identifier>
</mods:mods>
<treatment ID-DOI="http://doi.org/10.5281/zenodo.4328639" ID-GBIF-Taxon="168754888" ID-Zenodo-Dep="4328639" LSID="urn:lsid:plazi:treatment:B76E487AFFFDFFD6FF5205F6FB7F1113" httpUri="http://treatment.plazi.org/id/B76E487AFFFDFFD6FF5205F6FB7F1113" lastPageNumber="19" pageId="18" pageNumber="19">
<paragraph blockId="18.[155,793,1245,1952]" pageId="18" pageNumber="19">
<subSubSection box="[155,541,1245,1269]" pageId="18" pageNumber="19" type="nomenclature">
<heading box="[155,541,1245,1269]" fontSize="10" level="2" pageId="18" pageNumber="19" reason="1">
<taxonomicName authority="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill, 2012" authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[155,394,1245,1269]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="smithersi" status="new species">
<emphasis box="[155,394,1245,1269]" italics="true" pageId="18" pageNumber="19">Rhinolophus smithersi</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[404,541,1245,1269]" pageId="18" pageNumber="19" rank="species">new species</taxonomicNameLabel>
</heading>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="description">
<uri pageId="18" pageNumber="19">
urn:lsid:zoobank.org:act:
<uuid pageId="18" pageNumber="19">E78B0F44-7534-4991-975AF176CB668DDE</uuid>
</uri>
</subSubSection>
</paragraph>
<subSubSection box="[179,428,1333,1355]" pageId="18" pageNumber="19" type="vernacular_names">
<paragraph blockId="18.[155,793,1245,1952]" box="[179,428,1333,1355]" pageId="18" pageNumber="19">Smithers’s Horseshoe Bat</paragraph>
</subSubSection>
<subSubSection box="[179,679,1362,1384]" pageId="18" pageNumber="19" type="description">
<paragraph blockId="18.[155,793,1245,1952]" box="[179,679,1362,1384]" pageId="18" pageNumber="19">
<figureCitation box="[179,237,1362,1384]" captionStart="Figure 4" captionStartId="8.[840,905,1377,1396]" captionTargetBox="[844,1474,166,1343]" captionTargetId="figure-696@8.[840,1478,162,1343]" captionTargetPageId="8" captionText="Figure 4. Morphometric variation in a series representing the R. hildebrandtii complex from Lutope-Ngolangola, Zimbabwe: a) biplot of forearm length versus noseleaf width and b) PCA of five craniometric variables (M3M3, CM3, IOC, NW, NH) in 26 individuals of known (37 or 46 kHz) and unknown (?) frequency. Females indicated by open circlesı males by closed circles or crosses or asterisk. Voucher specimens for molecular sequencing study indicated by asterisk (Clade 1e: = smithersii sp. nov.; see Taxonomic Conclusions) and crosses (Clade 2: = mossambicus sp. nov.; see Taxonomic Conclusions). Hereafterı all individuals with a frequency of 37 kHz were assumed to belong to Clade 2 (mossambicus sp. nov.) and the 46 kHz individual was assumed to belong to Clade 1e (smithersi sp. nov.). doi:10.1371/journal.pone.0041744.g004" figureDoi="http://doi.org/10.5281/zenodo.4244568" httpUri="https://zenodo.org/record/4244568/files/figure.png" pageId="18" pageNumber="19">Fig. 4</figureCitation>
,
<figureCitation box="[249,319,1362,1384]" captionStart="Figure 1" captionStartId="1.[155,220,1840,1859]" captionTargetBox="[158,790,800,1803]" captionTargetId="figure-761@1.[155,793,797,1806]" captionTargetPageId="1" captionText="Figure 1. Portraits of (a) Rhinolophus smithersi species novo, and (b) Rhinolophus mossambicus species novo, two of four new cryptic species described herein within the R. hildebrandtii complex. doi:10.1371/journal.pone.0041744.g001" figureDoi="http://doi.org/10.5281/zenodo.4244562" httpUri="https://zenodo.org/record/4244562/files/figure.png" pageId="18" pageNumber="19">Fig. S1</figureCitation>
,
<tableCitation box="[332,410,1362,1384]" captionStart="Table 3" captionStartId="7.[171,235,1322,1343]" captionText="Table 3. Summary of data for peak echolocation frequency of R. hildebrandtii s.l. populations." pageId="18" pageNumber="19">Table 3</tableCitation>
,
<tableCitation box="[423,437,1362,1384]" captionStart="Table 4" captionStartId="15.[856,920,1521,1542]" captionText="Table 4. Eigenvectors for PCA of recently collected voucher specimens of the Rhinolophus hildebrandtii complex associated with molecular and acoustic analysesı which includes type specimens of R. hildebrandtii and R. eloquens (see Fig. 5a). doi:10.1371/journal.pone.0041744.t004" pageId="18" pageNumber="19">4</tableCitation>
,
<tableCitation box="[448,539,1362,1384]" captionStart="Table 1" captionStartId="6.[171,235,1481,1502]" captionText="Table 1. ‘Uncorrected p’ pairwise sequence distances for the 12S gene below the diagonal and Chd1 gene above the diagonal for the clades (1ı 2ı 3) and some lineages (1aı 1bı 1c) identified in the molecular analyses of the Rhinolophus hildebrandtii complex." pageId="18" pageNumber="19">Table S1</tableCitation>
, Appendix S1
</paragraph>
</subSubSection>
<materialsCitation ID-GBIF-Occurrence="2974116301" collectingDate="2000-10-26" collectionCode="NMZB" collectorName="F. P. D. Cotterill &amp; P. J. Taylor. Skull" country="Zimbabwe" elevation="1000" latitude="-18.2847222" location="Ngolangola Gorge at confluence with Lutope River" longitude="028.0833333" municipality="Sebungwe District" pageId="18" pageNumber="19" specimenCode="NMZB 33647" specimenCount="1" specimenCount-female="1" stateProvince="Gokwe Communal Land" typeStatus="holotype">
<subSubSection pageId="18" pageNumber="19" type="materials_examined">
<paragraph blockId="18.[155,793,1245,1952]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[177,282,1392,1411]" pageId="18" pageNumber="19">
<typeStatus box="[177,277,1392,1411]" pageId="18" pageNumber="19">Holotype</typeStatus>
.
</emphasis>
<specimenCode box="[306,444,1391,1413]" pageId="18" pageNumber="19">NMZB 33647</specimenCode>
, alcohol and cleaned skull. Collector Number FWC 4764.
<specimenCount box="[367,437,1419,1441]" pageId="18" pageNumber="19" type="female">Female</specimenCount>
collected
<date box="[538,706,1419,1441]" pageId="18" pageNumber="19" value="2000-10-26">
<collectingDate box="[538,706,1419,1441]" pageId="18" pageNumber="19" value="2000-10-26">26 October 2000</collectingDate>
</date>
by
<collectorName pageId="18" pageNumber="19">F. P. D. Cotterill</collectorName>
and
<collectorName box="[341,539,1447,1469]" pageId="18" pageNumber="19">P. J. Taylor. Skull</collectorName>
and mandible in good condition.
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="distribution">
<paragraph blockId="18.[155,793,1245,1952]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[177,324,1505,1524]" pageId="18" pageNumber="19">Type locality.</emphasis>
<location LSID="urn:lsid:plazi:treatment:B76E487AFFFDFFD6FF5205F6FB7F1113:3A18AFB7FFFDFFD6FE9404CBFF471146" country="NW Zimbabwe" latitude="-9.083333" longitude="9.0" municipality="Sebungwe District" name="Ngolangola Gorge at confluence with Lutope River" pageId="18" pageNumber="19" stateProvince="Gokwe Communal Land">Ngolangola Gorge at confluence with Lutope River</location>
,
<collectingMunicipality box="[236,429,1533,1555]" pageId="18" pageNumber="19">Sebungwe District</collectingMunicipality>
,
<collectingRegion box="[454,727,1533,1555]" pageId="18" pageNumber="19">Gokwe Communal Land</collectingRegion>
,
<collectingCountry pageId="18" pageNumber="19">NW Zimbabwe</collectingCountry>
;
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18
<emphasis bold="true" box="[296,305,1560,1581]" pageId="18" pageNumber="19">°</emphasis>
<date box="[313,369,1560,1583]" pageId="18" pageNumber="19" value="2005-09-17">
17
<emphasis box="[337,345,1560,1582]" italics="true" pageId="18" pageNumber="19">'</emphasis>
05
</date>
<emphasis box="[369,377,1560,1582]" italics="true" pageId="18" pageNumber="19">&quot;</emphasis>
S
</geoCoordinate>
;
<geoCoordinate box="[404,533,1560,1583]" pageId="18" pageNumber="19">
28
<emphasis bold="true" box="[428,437,1560,1581]" pageId="18" pageNumber="19">°</emphasis>
<date box="[445,509,1560,1583]" pageId="18" pageNumber="19" value="2000-09-05">
05
<emphasis box="[469,477,1560,1582]" italics="true" pageId="18" pageNumber="19">'</emphasis>
00
</date>
<emphasis box="[508,516,1560,1582]" italics="true" pageId="18" pageNumber="19">&quot;</emphasis>
E
</geoCoordinate>
; elevation
<quantity box="[640,714,1561,1583]" metricMagnitude="3" metricUnit="m" metricValue="1.0" pageId="18" pageNumber="19" unit="m" value="1000.0">
<elevation box="[640,714,1561,1583]" metricMagnitude="3" metricUnit="m" metricValue="1.0" pageId="18" pageNumber="19" unit="m" value="1000.0">1000 m</elevation>
</quantity>
asl.
</paragraph>
</subSubSection>
</materialsCitation>
<subSubSection pageId="18" pageNumber="19" type="diagnosis">
<paragraph blockId="18.[155,793,1245,1952]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[177,291,1590,1609]" pageId="18" pageNumber="19">Diagnosis.</emphasis>
Peak echolocation frequency at 44–46 kHz which is considerably higher than all other members of this speciescomplex (33–42 kHz). Very small-cranium (mean CCL
<quantity box="[698,789,1646,1668]" metricMagnitude="-2" metricUnit="m" metricValue="2.34" pageId="18" pageNumber="19" unit="mm" value="23.4">23.4 mm</quantity>
; 23.0–
<quantity box="[209,300,1674,1696]" metricMagnitude="-2" metricUnit="m" metricValue="2.43" pageId="18" pageNumber="19" unit="mm" value="24.3">24.3 mm</quantity>
; n = 6;
<quantity box="[373,461,1674,1696]" metricMagnitude="-2" metricUnit="m" metricValue="2.32" pageId="18" pageNumber="19" unit="mm" value="23.2">23.2 mm</quantity>
in
<typeStatus box="[496,581,1674,1696]" pageId="18" pageNumber="19">holotype</typeStatus>
), the
<typeStatus box="[637,720,1674,1696]" pageId="18" pageNumber="19">holotype</typeStatus>
having a disproportionately wide noseleaf (
<quantity box="[508,580,1703,1725]" metricMagnitude="-2" metricUnit="m" metricValue="1.4" pageId="18" pageNumber="19" unit="mm" value="14.0">14 mm</quantity>
; 53% of skull length) (although this is not as pronounced in the series from Pafuri, NE South Africa; mean noseleaf width
<quantity box="[494,585,1760,1782]" metricMagnitude="-2" metricUnit="m" metricValue="1.0699999999999998" pageId="18" pageNumber="19" unit="mm" value="10.7">10.7 mm</quantity>
; 39% of skull length), it is immediately distinguishable from other members of the
<taxonomicName authorityName="Petersi" authorityYear="1878" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis italics="true" pageId="18" pageNumber="19">R. hildebrandtii</emphasis>
</taxonomicName>
complex on size and peak frequency (
<figureCitation box="[684,755,1816,1838]" captionStart="Figure 3" captionStartId="6.[155,220,1057,1076]" captionTargetBox="[160,1460,171,1017]" captionTargetId="figure-89@6.[155,1464,162,1023]" captionTargetPageId="6" captionText="Figure 3. Consensus tree for the cytochrome b dataset for representative genotyped specimens of the Rhinolophus hildebrandtii complex. The topology represents the consensus topology from a 20 million MCMC run implemented in BEAST. Estimates of divergence times (million years ago; Mya) are indicated adjacent to nodes or above branches and grey bars indicate 95% HPD values. The split between the Hipposideridae and Rhinolophidae was used as the calibration point. Taxa names include museum/field numbers which correspond to Appendix S1 or GenBank accession numbers and abbreviations are: RcfH - R. cf. hildebrandtiiı RD - R. darlingiı RE - R. eloquensı RF - R. fumigatusı RH - R. hildebrandtii s.l.ı RL - R. landeri and RR - R. ruwenzorii. Localitiesı where availableı are providedı abbreviations include SA - South Africaı MZ - Mozambiqueı and ZW - Zimbabweı and the numbers in parentheses correspond with place names in Table S1 and Fig. 2 for Clade 1 and 2 individuals. doi:10.1371/journal.pone.0041744.g003" figureDoi="http://doi.org/10.5281/zenodo.4244566" httpUri="https://zenodo.org/record/4244566/files/figure.png" pageId="18" pageNumber="19">Figs. 3</figureCitation>
;
<figureCitation box="[774,788,1816,1838]" captionStart="Figure 4" captionStartId="8.[840,905,1377,1396]" captionTargetBox="[844,1474,166,1343]" captionTargetId="figure-696@8.[840,1478,162,1343]" captionTargetPageId="8" captionText="Figure 4. Morphometric variation in a series representing the R. hildebrandtii complex from Lutope-Ngolangola, Zimbabwe: a) biplot of forearm length versus noseleaf width and b) PCA of five craniometric variables (M3M3, CM3, IOC, NW, NH) in 26 individuals of known (37 or 46 kHz) and unknown (?) frequency. Females indicated by open circlesı males by closed circles or crosses or asterisk. Voucher specimens for molecular sequencing study indicated by asterisk (Clade 1e: = smithersii sp. nov.; see Taxonomic Conclusions) and crosses (Clade 2: = mossambicus sp. nov.; see Taxonomic Conclusions). Hereafterı all individuals with a frequency of 37 kHz were assumed to belong to Clade 2 (mossambicus sp. nov.) and the 46 kHz individual was assumed to belong to Clade 1e (smithersi sp. nov.). doi:10.1371/journal.pone.0041744.g004" figureDoi="http://doi.org/10.5281/zenodo.4244568" httpUri="https://zenodo.org/record/4244568/files/figure.png" pageId="18" pageNumber="19">4</figureCitation>
;
<tableCitation box="[155,231,1845,1867]" captionStart="Table 5" captionStartId="17.[153,217,487,508]" captionText="Table 5. Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the Rhinolophus hildebrandtii complex." pageId="18" pageNumber="19">Table 5</tableCitation>
). Genotypes of
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[386,485,1844,1866]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="smithersi">
<emphasis box="[386,485,1844,1866]" italics="true" pageId="18" pageNumber="19">R. smithersi</emphasis>
</taxonomicName>
are members of Clades 1d and 1e (
<figureCitation box="[192,249,1873,1895]" captionStart="Figure 2" captionStartId="2.[155,220,1881,1900]" captionTargetBox="[160,1475,167,1843]" captionTargetId="figure-0@2.[155,1480,162,1848]" captionTargetPageId="2" captionText="Figure 2. Map of southern, central and eastern Africa indicating localities of individuals of R. hildebrandtii species-complex included in this study. Grey-shaded area represents elevations in excess of 600 m a.s.l. Closed squares indicate museum specimens from which craniometric data were obtained. Open symbols indicate specimens genotyped in this study. The distribution of the three major clades is based on cytochrome b (see Figure 2): open circles = Clade 1; open squares = Clade 2; open diamonds = Clade 3. Closed squares enclosed in open symbols indicate localities where both molecular and morphological data were available for selected specimens. Numbers refer to respective localities listed in Table S1. ‘‘T’’ indicates the type localities of R. eloquens in Uganda and R. hildebrandtii in Kenyaı respectively. doi:10.1371/journal.pone.0041744.g002" figureDoi="http://doi.org/10.5281/zenodo.4244564" httpUri="https://zenodo.org/record/4244564/files/figure.png" pageId="18" pageNumber="19">Fig. 2</figureCitation>
,
<figureCitation box="[261,330,1873,1895]" captionStart="Figure 1" captionStartId="1.[155,220,1840,1859]" captionTargetBox="[158,790,800,1803]" captionTargetId="figure-761@1.[155,793,797,1806]" captionTargetPageId="1" captionText="Figure 1. Portraits of (a) Rhinolophus smithersi species novo, and (b) Rhinolophus mossambicus species novo, two of four new cryptic species described herein within the R. hildebrandtii complex. doi:10.1371/journal.pone.0041744.g001" figureDoi="http://doi.org/10.5281/zenodo.4244562" httpUri="https://zenodo.org/record/4244562/files/figure.png" pageId="18" pageNumber="19">Fig. S1</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="materials_examined">
<paragraph blockId="18.[155,793,1245,1952]" lastBlockId="18.[840,1479,164,1606]" pageId="18" pageNumber="19">
<materialsCitation ID-GBIF-Occurrence="2908237316" accessionNumber="JQ929228" collectingDate="2000-10-26" collectionCode="NMZB" collectorName="F. P. D. Cotterill" country="Zimbabwe" location="Zimbabwe" pageId="18" pageNumber="19" specimenCode="NMZB 33652" specimenCount="1" specimenCount-male="1" typeStatus="paratype">
<emphasis bold="true" box="[177,281,1903,1922]" pageId="18" pageNumber="19">
<typeStatus box="[177,276,1903,1922]" pageId="18" pageNumber="19">Paratype</typeStatus>
.
</emphasis>
<specimenCode box="[305,450,1901,1923]" pageId="18" pageNumber="19">NMZB 33652</specimenCode>
,
<specimenCount box="[467,514,1901,1923]" pageId="18" pageNumber="19" type="male">male</specimenCount>
alcohol and cleaned skull, Collector Number FWC 4769. Collected
<date box="[569,760,1930,1952]" pageId="18" pageNumber="19" value="2000-10-26">
<collectingDate box="[569,760,1930,1952]" pageId="18" pageNumber="19" value="2000-10-26">26th October 2000</collectingDate>
</date>
by
<collectorName box="[840,992,164,186]" pageId="18" pageNumber="19">F.P.D. Cotterill</collectorName>
and P.J. Taylor from Lutope-Ngolangola Gorge,
<collectingCountry box="[840,946,192,214]" name="Zimbabwe" pageId="18" pageNumber="19" value="Zimbabwe">Zimbabwe</collectingCountry>
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="description">
<paragraph blockId="18.[840,1479,164,1606]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[863,996,222,241]" pageId="18" pageNumber="19">Description.</emphasis>
Apart from its distrinctly smaller size, conforming generally to the keys and published description for
<taxonomicName authorityName="Petersi" authorityYear="1878" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" sensu="lato" species="hildebrandtii">
<emphasis italics="true" pageId="18" pageNumber="19">R. hildebrandtii</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[955,981,277,299]" pageId="18" pageNumber="19" sensu="lato">s.l.</taxonomicNameLabel>
[
<bibRefCitation author="Csorba G &amp; Ujhelyip P &amp; Thomas N" box="[999,1027,277,299]" journalOrPublisher="Shropshire: Alana books" pageId="18" pageNumber="19" refId="ref21181" refString="12. Csorba G, Ujhelyip P, Thomas N (2003) Horseshoe Bats of the World (Chiroptera: Rhinolophidae). Shropshire: Alana books." title="Horseshoe Bats of the World (Chiroptera: Rhinolophidae)" type="book" year="2003">12</bibRefCitation>
], i.e. a relatively large-sized (forearm length
<quantity box="[840,928,306,328]" metricMagnitude="-2" metricUnit="m" metricValue="6.07" pageId="18" pageNumber="19" unit="mm" value="60.7">60.7 mm</quantity>
in
<typeStatus box="[976,1059,306,328]" pageId="18" pageNumber="19">holotype</typeStatus>
) bat with medium or large ears, wide horseshoe (
<quantity box="[949,1053,334,356]" metricMagnitude="-2" metricUnit="m" metricValue="1.2" metricValueMax="1.4" metricValueMin="1.0" pageId="18" pageNumber="19" unit="mm" value="12.0" valueMax="14.0" valueMin="10.0">10–14 mm</quantity>
) covering the muzzle, sella constricted at its proximal third, almost parallel-sided above and having long hairs, lancet relatively long and straight-sized, lower lip with single median groove, upper parts greyish-brown, with individual hairs long and unicoloured, underparts same colour or very slightly paler, skull large and heavily built. Lower lip with single mental groove. The sagittal crest is very well developed in relation to the nasal inflation and the rostral depression is weakly developed. The anterior premolar is small-sized and external to the toothrow (
<tableCitation box="[847,923,590,612]" captionStart="Table 5" captionStartId="17.[153,217,487,508]" captionText="Table 5. Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the Rhinolophus hildebrandtii complex." pageId="18" pageNumber="19">Table 5</tableCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="distribution">
<paragraph blockId="18.[840,1479,164,1606]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[863,1003,619,638]" pageId="18" pageNumber="19">Distribution.</emphasis>
Known from Lutope-Ngolangola Gorge south of the Zambezi Escarpment in NW Zimbabwe and also from Pafuri in the Limpopo Valley in the foothills of the Soutpansberg Mountains of northern Limpopo Province, South Africa but likely more widespread across savanna woodlands of the Limpopo and Zambezi valleys, and their escarpments (the Gwembe horst, and the Soutpansberg and Waterberg Mountains, respectively). Accurate delimitation of this species’ range is subject to further collecting and reappraisal of existing museum material. This applies to papers by M.B. Fenton and co-workers based on intensive studies of the bat fauna, which included populations of this species complex in the Sengwa Wildlife Research Area, only,
<quantity box="[860,910,959,981]" metricMagnitude="3" metricUnit="m" metricValue="5.0" pageId="18" pageNumber="19" unit="km" value="5.0">5 km</quantity>
north of the
<typeStatus box="[1135,1176,959,981]" pageId="18" pageNumber="19">type</typeStatus>
locality of
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[1355,1477,959,981]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="smithersi">
<emphasis box="[1355,1477,959,981]" italics="true" pageId="18" pageNumber="19">R. smithersi</emphasis>
</taxonomicName>
[120,121,122,123,124,125,126]. These all reported the largest rhinolophid studied as
<taxonomicName authorityName="Petersi" authorityYear="1878" box="[1060,1190,1016,1038]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[1060,1190,1016,1038]" italics="true" pageId="18" pageNumber="19">R. hildebrandtii</emphasis>
</taxonomicName>
, but these data would refer to both
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[892,1026,1044,1066]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="mossambicus">
<emphasis box="[892,1026,1044,1066]" italics="true" pageId="18" pageNumber="19">R. mossambicus</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[1078,1178,1044,1066]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="smithersi">
<emphasis box="[1078,1178,1044,1066]" italics="true" pageId="18" pageNumber="19">R. smithersi</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="biology_ecology">
<paragraph blockId="18.[840,1479,164,1606]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[863,953,1074,1093]" pageId="18" pageNumber="19">Ecology.</emphasis>
Poorly known; this species occurs sympatrically with
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[840,980,1101,1123]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="mossambicus">
<emphasis box="[840,980,1101,1123]" italics="true" pageId="18" pageNumber="19">R. mossambicus</emphasis>
</taxonomicName>
at one locality in miombo savanna on Karoo Sandstone, dominated by trees of
<taxonomicName box="[1213,1420,1129,1151]" class="Magnoliopsida" family="Fabaceae" genus="Brachystegia" kingdom="Plantae" order="Fabales" pageId="18" pageNumber="19" phylum="Tracheophyta" rank="species" species="glaucescens">
<emphasis box="[1213,1420,1129,1151]" italics="true" pageId="18" pageNumber="19">Brachystegia glaucescens</emphasis>
</taxonomicName>
, and including large specimens of baobabs,
<taxonomicName box="[1245,1405,1158,1180]" class="Magnoliopsida" family="Malvaceae" genus="Adansonia" kingdom="Plantae" order="Malvales" pageId="18" pageNumber="19" phylum="Tracheophyta" rank="species" species="digitata">
<emphasis box="[1245,1405,1158,1180]" italics="true" pageId="18" pageNumber="19">Adansonia digitata</emphasis>
</taxonomicName>
; albeit much of the landscape has been converted to cotton fields. More diverse riparian woodland fringes the Lutope and Ngolanola rivers as well as along the
<collectingRegion box="[1031,1119,1244,1266]" country="South Africa" name="Limpopo" pageId="18" pageNumber="19">Limpopo</collectingRegion>
River at Pafuri. Daylight roosts were not located but these bats could use caves in the sandstone cliffs and/or hollows in baobabs.
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="etymology">
<paragraph blockId="18.[840,1479,164,1606]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[863,984,1330,1349]" pageId="18" pageNumber="19">Etymology.</emphasis>
We selected the specific epithet in recognition of the late Reay Henry Noble Smithers (
<date box="[1201,1310,1357,1379]" pageId="18" pageNumber="19" value="1907" valueMax="1987">1907–1987</date>
), former Director of the National Museums of Zimbabwe, prodigious collector and researcher of mammals including bats, and author of important regional texts on the mammalogy of Botswana, Zimbabwe and Mozambique, notably his definitive monograph –
<emphasis italics="true" pageId="18" pageNumber="19">The Mammals of the Southern African Subregion</emphasis>
subsequently updated and revised [
<bibRefCitation author="Smithers RHN" box="[847,889,1527,1549]" pageId="18" pageNumber="19" refId="ref25854" refString="127. Smithers RHN (1983) The Mammals of the Southern African Subregion. Pretoria: University of Pretoria Press. 736 p." type="book" year="1983">127</bibRefCitation>
]. The species name combines the surname Smithers and genitive singular case-ending ‘‘i’’ indicative of masculine gender.
</paragraph>
<paragraph blockId="18.[840,1479,164,1606]" box="[863,1259,1584,1606]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[863,1097,1585,1604]" pageId="18" pageNumber="19">Specimens examined.</emphasis>
<collectingRegion box="[1122,1156,1584,1606]" country="South Sudan" name="Lakes" pageId="18" pageNumber="19">See</collectingRegion>
<tableCitation box="[1163,1254,1584,1606]" captionStart="Table 1" captionStartId="6.[171,235,1481,1502]" captionText="Table 1. ‘Uncorrected p’ pairwise sequence distances for the 12S gene below the diagonal and Chd1 gene above the diagonal for the clades (1ı 2ı 3) and some lineages (1aı 1bı 1c) identified in the molecular analyses of the Rhinolophus hildebrandtii complex." pageId="18" pageNumber="19">Table S1</tableCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>

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<mods:title>Four New Bat Species (Rhinolophus hildebrandtii Complex) Reflect Plio-Pleistocene Divergence of Dwarfs and Giants across an Afromontane Archipelago</mods:title>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Peter J. Taylor</mods:namePart>
<mods:affiliation>Department of Ecology and Resource Management, University of Venda, Thohoyandou, South Africa, Durban Natural Science Museum, Durban, South Africa, School of Life Sciences, University of KwaZulu-Natal, Durban, South Africa</mods:affiliation>
<mods:nameIdentifier type="email">peter.taylor@univen.ac.za</mods:nameIdentifier>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Samantha Stoffberg</mods:namePart>
<mods:affiliation>Evolutionary Genomics Group, Department of Botany and Zoology, University of Stellenbosch, Stellenbosch, South Africa</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ara Monadjem</mods:namePart>
<mods:affiliation>All Out Africa Research Unit, Department of Biological Sciences, University of Swaziland, Kwaluseni, Swaziland</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Martinus Corrie Schoeman</mods:namePart>
<mods:affiliation>School of Life Sciences, University of KwaZulu-Natal, Durban, South Africa</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Julian Baylis</mods:namePart>
<mods:affiliation>Mulanje Mountain Conservation Trust, Mulanje, Malawi, Conservation Science Group, Department of Zoology, University of Cambridge, Cambridge, United Kingdom</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Fenton P. D. Cotterill</mods:namePart>
<mods:affiliation>Africa Earth Observatory Network, Geoecodynamics Research Hub, Department of Botany and Zoology, University of Stellenbosch, Stellenbosch, South Africa</mods:affiliation>
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<mods:date>2012</mods:date>
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<mods:number>2012-09-12</mods:number>
</mods:detail>
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<mods:number>7</mods:number>
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<mods:number>9</mods:number>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.4328645" ID-GBIF-Taxon="168754887" ID-Zenodo-Dep="4328645" LSID="urn:lsid:zoobank.org:act:0D97DC39-F9CD-415E-A7AE-4DCF70C807B4" httpUri="http://treatment.plazi.org/id/B76E487AFFFDFFD7FC810740FDA910F1" lastPageId="19" lastPageNumber="20" pageId="18" pageNumber="19">
<paragraph blockId="18.[840,1478,1643,1952]" pageId="18" pageNumber="19">
<subSubSection box="[840,1276,1643,1667]" pageId="18" pageNumber="19" type="nomenclature">
<heading box="[840,1276,1643,1667]" pageId="18" pageNumber="19">
<taxonomicName authority="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill, 2012" authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[840,1129,1643,1667]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="mossambicus" status="new species">
<emphasis box="[840,1129,1643,1667]" italics="true" pageId="18" pageNumber="19">Rhinolophus mossambicus</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1138,1276,1643,1667]" pageId="18" pageNumber="19" rank="species">new species</taxonomicNameLabel>
</heading>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="description">
<uri pageId="18" pageNumber="19">
urn:lsid:zoobank.org:act:
<uuid pageId="18" pageNumber="19">0D97DC39-F9CD-415E-A7AE4DCF70C807B4</uuid>
</uri>
</subSubSection>
</paragraph>
<subSubSection box="[864,1142,1731,1753]" pageId="18" pageNumber="19" type="vernacular_names">
<paragraph blockId="18.[840,1478,1643,1952]" box="[864,1142,1731,1753]" pageId="18" pageNumber="19">Mozambican Horseshoe Bat</paragraph>
</subSubSection>
<subSubSection box="[864,1390,1760,1782]" pageId="18" pageNumber="19" type="description">
<paragraph blockId="18.[840,1478,1643,1952]" box="[864,1390,1760,1782]" pageId="18" pageNumber="19">
<figureCitation box="[864,948,1760,1782]" captionStart="Figure 4" captionStartId="8.[840,905,1377,1396]" captionTargetBox="[844,1474,166,1343]" captionTargetId="figure-696@8.[840,1478,162,1343]" captionTargetPageId="8" captionText="Figure 4. Morphometric variation in a series representing the R. hildebrandtii complex from Lutope-Ngolangola, Zimbabwe: a) biplot of forearm length versus noseleaf width and b) PCA of five craniometric variables (M3M3, CM3, IOC, NW, NH) in 26 individuals of known (37 or 46 kHz) and unknown (?) frequency. Females indicated by open circlesı males by closed circles or crosses or asterisk. Voucher specimens for molecular sequencing study indicated by asterisk (Clade 1e: = smithersii sp. nov.; see Taxonomic Conclusions) and crosses (Clade 2: = mossambicus sp. nov.; see Taxonomic Conclusions). Hereafterı all individuals with a frequency of 37 kHz were assumed to belong to Clade 2 (mossambicus sp. nov.) and the 46 kHz individual was assumed to belong to Clade 1e (smithersi sp. nov.). doi:10.1371/journal.pone.0041744.g004" figureDoi="http://doi.org/10.5281/zenodo.4244568" httpUri="https://zenodo.org/record/4244568/files/figure.png" pageId="18" pageNumber="19">Fig. 4, 4</figureCitation>
,
<figureCitation box="[960,1031,1760,1782]" captionStart="Figure 1" captionStartId="1.[155,220,1840,1859]" captionTargetBox="[158,790,800,1803]" captionTargetId="figure-761@1.[155,793,797,1806]" captionTargetPageId="1" captionText="Figure 1. Portraits of (a) Rhinolophus smithersi species novo, and (b) Rhinolophus mossambicus species novo, two of four new cryptic species described herein within the R. hildebrandtii complex. doi:10.1371/journal.pone.0041744.g001" figureDoi="http://doi.org/10.5281/zenodo.4244562" httpUri="https://zenodo.org/record/4244562/files/figure.png" pageId="18" pageNumber="19">Fig. S1</figureCitation>
,
<tableCitation box="[1043,1121,1760,1782]" captionStart="Table 3" captionStartId="7.[171,235,1322,1343]" captionText="Table 3. Summary of data for peak echolocation frequency of R. hildebrandtii s.l. populations." pageId="18" pageNumber="19">Table 3</tableCitation>
,
<tableCitation box="[1134,1148,1760,1782]" captionStart="Table 4" captionStartId="15.[856,920,1521,1542]" captionText="Table 4. Eigenvectors for PCA of recently collected voucher specimens of the Rhinolophus hildebrandtii complex associated with molecular and acoustic analysesı which includes type specimens of R. hildebrandtii and R. eloquens (see Fig. 5a). doi:10.1371/journal.pone.0041744.t004" pageId="18" pageNumber="19">4</tableCitation>
,
<tableCitation box="[1160,1250,1760,1782]" captionStart="Table 1" captionStartId="6.[171,235,1481,1502]" captionText="Table 1. ‘Uncorrected p’ pairwise sequence distances for the 12S gene below the diagonal and Chd1 gene above the diagonal for the clades (1ı 2ı 3) and some lineages (1aı 1bı 1c) identified in the molecular analyses of the Rhinolophus hildebrandtii complex." pageId="18" pageNumber="19">Table S1</tableCitation>
, Appendix S1
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="materials_examined">
<paragraph blockId="18.[840,1478,1643,1952]" pageId="18" pageNumber="19">
<materialsCitation ID-GBIF-Occurrence="2908237314" accessionNumber="JQ929255, JQ929203, JQ929234, JQ929289" collectingDate="2006-07-01" collectionCode="DM" collectorName="A. Monadjem" country="Mozambique" elevation="489" latitude="-12.1822222" location="Niassa Game Reserve (Maputo Camp)" longitude="037.5500000" pageId="18" pageNumber="19" specimenCode="DM8578" specimenCount="1" specimenCount-male="1" stateProvince="northern Mozambique" typeStatus="holotype">
<emphasis bold="true" box="[863,967,1790,1809]" pageId="18" pageNumber="19">
<typeStatus box="[863,963,1790,1809]" pageId="18" pageNumber="19">Holotype</typeStatus>
.
</emphasis>
<specimenCode box="[992,1082,1788,1810]" pageId="18" pageNumber="19">DM8578</specimenCode>
,
<specimenCount box="[1095,1144,1788,1810]" pageId="18" pageNumber="19" type="male">male</specimenCount>
, alcohol skin, skull and baculum, collected by
<collectorName box="[963,1098,1817,1839]" pageId="18" pageNumber="19">A. Monadjem</collectorName>
on
<date box="[1137,1253,1817,1839]" pageId="18" pageNumber="19" value="2006-07-01">
<collectingDate box="[1137,1253,1817,1839]" pageId="18" pageNumber="19" value="2006-07-01">1 July 2006</collectingDate>
</date>
. Skull and mandible in good condition.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="distribution">
<paragraph blockId="18.[840,1478,1643,1952]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[863,1021,1874,1894]" pageId="18" pageNumber="19">
<typeStatus box="[863,917,1875,1894]" pageId="18" pageNumber="19">Type</typeStatus>
locality.
</emphasis>
<location LSID="urn:lsid:plazi:treatment:B76E487AFFFDFFD7FC810740FDA910F1:3A18AFB7FFFDFFD6FBDF067AFA561032" box="[1046,1474,1873,1895]" country="Mozambique" latitude="-9.933333" longitude="9.0" name="Niassa Game Reserve (Maputo Camp)" pageId="18" pageNumber="19" stateProvince="northern Mozambique">Niassa Game Reserve (Maputo Camp)</location>
,
<collectingRegion box="[840,1070,1902,1924]" pageId="18" pageNumber="19">northern Mozambique</collectingRegion>
,
<geoCoordinate box="[1088,1213,1901,1924]" pageId="18" pageNumber="19">
12
<emphasis bold="true" box="[1112,1121,1901,1922]" pageId="18" pageNumber="19">°</emphasis>
<date box="[1135,1191,1901,1924]" pageId="18" pageNumber="19" value="1956-09-10">
10
<emphasis box="[1159,1167,1901,1923]" italics="true" pageId="18" pageNumber="19">'</emphasis>
56
</date>
<emphasis box="[1191,1199,1901,1923]" italics="true" pageId="18" pageNumber="19">&quot;</emphasis>
S
</geoCoordinate>
;
<geoCoordinate box="[1232,1374,1901,1924]" pageId="18" pageNumber="19">
37
<emphasis bold="true" box="[1256,1265,1901,1922]" pageId="18" pageNumber="19">°</emphasis>
33
<emphasis box="[1303,1311,1901,1923]" italics="true" pageId="18" pageNumber="19">'</emphasis>
00
<emphasis box="[1349,1357,1901,1923]" italics="true" pageId="18" pageNumber="19">&quot;</emphasis>
E
</geoCoordinate>
; elevation
<quantity box="[840,903,1930,1952]" metricMagnitude="2" metricUnit="m" metricValue="4.89" pageId="18" pageNumber="19" unit="m" value="489.0">
<elevation box="[840,903,1930,1952]" metricMagnitude="2" metricUnit="m" metricValue="4.89" pageId="18" pageNumber="19" unit="m" value="489.0">489 m</elevation>
</quantity>
asl.
</paragraph>
</subSubSection>
<subSubSection pageId="19" pageNumber="20" type="diagnosis">
<paragraph blockId="19.[155,793,163,1956]" pageId="19" pageNumber="20">
<emphasis bold="true" box="[177,291,164,183]" pageId="19" pageNumber="20">Diagnosis.</emphasis>
Peak frequency between 35–38 kHz (not recorded in
<typeStatus box="[217,300,191,213]" pageId="19" pageNumber="20">holotype</typeStatus>
but inferred from members of the same genetic clade). Members of this cryptic species are genetically distinct from
<taxonomicName authorityName="Petersi" authorityYear="1878" box="[155,292,247,269]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" sensu="stricto" species="hildebrandtii">
<emphasis box="[155,292,247,269]" italics="true" pageId="19" pageNumber="20">R. hildebrandtii</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[304,332,248,270]" pageId="19" pageNumber="20" sensu="stricto">s.s.</taxonomicNameLabel>
(7.7–9.0% cytochrome b uncorrected divergence; Clade
<quantity box="[291,331,276,298]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="19" pageNumber="20" unit="in" value="2.0">2 in</quantity>
<figureCitation box="[340,398,276,298]" captionStart="Figure 2" captionStartId="2.[155,220,1881,1900]" captionTargetBox="[160,1475,167,1843]" captionTargetId="figure-0@2.[155,1480,162,1848]" captionTargetPageId="2" captionText="Figure 2. Map of southern, central and eastern Africa indicating localities of individuals of R. hildebrandtii species-complex included in this study. Grey-shaded area represents elevations in excess of 600 m a.s.l. Closed squares indicate museum specimens from which craniometric data were obtained. Open symbols indicate specimens genotyped in this study. The distribution of the three major clades is based on cytochrome b (see Figure 2): open circles = Clade 1; open squares = Clade 2; open diamonds = Clade 3. Closed squares enclosed in open symbols indicate localities where both molecular and morphological data were available for selected specimens. Numbers refer to respective localities listed in Table S1. ‘‘T’’ indicates the type localities of R. eloquens in Uganda and R. hildebrandtii in Kenyaı respectively. doi:10.1371/journal.pone.0041744.g002" figureDoi="http://doi.org/10.5281/zenodo.4244564" httpUri="https://zenodo.org/record/4244564/files/figure.png" pageId="19" pageNumber="20">Fig. 2</figureCitation>
and
<figureCitation box="[452,522,276,298]" captionStart="Figure 1" captionStartId="1.[155,220,1840,1859]" captionTargetBox="[158,790,800,1803]" captionTargetId="figure-761@1.[155,793,797,1806]" captionTargetPageId="1" captionText="Figure 1. Portraits of (a) Rhinolophus smithersi species novo, and (b) Rhinolophus mossambicus species novo, two of four new cryptic species described herein within the R. hildebrandtii complex. doi:10.1371/journal.pone.0041744.g001" figureDoi="http://doi.org/10.5281/zenodo.4244562" httpUri="https://zenodo.org/record/4244562/files/figure.png" pageId="19" pageNumber="20">Fig. S1</figureCitation>
). Noseleaf with typical low rounded profile of connecting process in the
<typeStatus box="[615,698,304,326]" pageId="19" pageNumber="20">holotype</typeStatus>
(
<figureCitation box="[716,786,304,326]" captionStart="Figure 8" captionStartId="12.[155,220,993,1012]" captionTargetBox="[163,1000,167,956]" captionTargetId="figure-464@12.[155,1007,162,960]" captionTargetPageId="12" captionText="Figure 8. Photographs showing lateral views of noseleafs of selected individuals (including holotypes of new species) of the Rhinolophus hildebrandtii complex representing molecular Lineage 1a (= cohenae sp. nov.; a–b), Lineage 1b (= mabuensis sp. nov.; c) and Clade 2 (= mossambicus sp. nov.; d–f). a = DM 7886 (cohenae sp. nov.;Barberton Tunnel, Mpumalanga Province, South Africa); b = DM 8626 (cohenae sp. nov.; Barberton Tunnel, Mpumalanga Province, South Africa; Holotype); c = DM 10842 (mabuensis sp. nov.; Mt Mabu, Mozambique; Holotype); d = DM 8578 (mossambicus sp. nov.; Niassa Game Reserve, Mozambique; Holotype); e = DM 8579 (mossambicus sp. nov.; Chinizuia, Mozambique); f = DM 8577 (mossambicus sp. nov.; Namapa, Mozambique). doi:10.1371/journal.pone.0041744.g008" figureDoi="http://doi.org/10.5281/zenodo.4244576" httpUri="https://zenodo.org/record/4244576/files/figure.png" pageId="19" pageNumber="20">Fig. 8d</figureCitation>
) although slight variation was observed among members of this species with one specimen having a higher and rounded shape (
<figureCitation box="[161,226,388,410]" captionStart="Figure 8" captionStartId="12.[155,220,993,1012]" captionTargetBox="[163,1000,167,956]" captionTargetId="figure-464@12.[155,1007,162,960]" captionTargetPageId="12" captionText="Figure 8. Photographs showing lateral views of noseleafs of selected individuals (including holotypes of new species) of the Rhinolophus hildebrandtii complex representing molecular Lineage 1a (= cohenae sp. nov.; a–b), Lineage 1b (= mabuensis sp. nov.; c) and Clade 2 (= mossambicus sp. nov.; d–f). a = DM 7886 (cohenae sp. nov.;Barberton Tunnel, Mpumalanga Province, South Africa); b = DM 8626 (cohenae sp. nov.; Barberton Tunnel, Mpumalanga Province, South Africa; Holotype); c = DM 10842 (mabuensis sp. nov.; Mt Mabu, Mozambique; Holotype); d = DM 8578 (mossambicus sp. nov.; Niassa Game Reserve, Mozambique; Holotype); e = DM 8579 (mossambicus sp. nov.; Chinizuia, Mozambique); f = DM 8577 (mossambicus sp. nov.; Namapa, Mozambique). doi:10.1371/journal.pone.0041744.g008" figureDoi="http://doi.org/10.5281/zenodo.4244576" httpUri="https://zenodo.org/record/4244576/files/figure.png" pageId="19" pageNumber="20">Fig. 8f</figureCitation>
). Skull intermediate-sized (GSL
<quantity box="[545,635,388,410]" metricMagnitude="-2" metricUnit="m" metricValue="2.85" pageId="19" pageNumber="20" unit="mm" value="28.5">28.5 mm</quantity>
; CCL 25.0 mm in the
<typeStatus box="[231,317,416,438]" pageId="19" pageNumber="20">holotype</typeStatus>
; range of GSL
<quantity box="[493,601,416,438]" metricMagnitude="-2" metricUnit="m" metricValue="2.8" metricValueMax="2.9" metricValueMin="2.7" pageId="19" pageNumber="20" unit="mm" value="28.0" valueMax="29.0" valueMin="27.0">27–29 mm</quantity>
; CCL
<quantity box="[680,789,416,438]" metricMagnitude="-2" metricUnit="m" metricValue="2.5" metricValueMax="2.6" metricValueMin="2.4" pageId="19" pageNumber="20" unit="mm" value="25.0" valueMax="26.0" valueMin="24.0">24–26 mm</quantity>
;
<tableCitation box="[155,231,444,466]" captionStart="Table 5" captionStartId="17.[153,217,487,508]" captionText="Table 5. Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the Rhinolophus hildebrandtii complex." pageId="19" pageNumber="20">Table 5</tableCitation>
) and can be distinguished from larger (
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[627,688,444,466]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="cohenae">
<emphasis box="[627,688,444,466]" italics="true" pageId="19" pageNumber="20">cohenae</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[703,786,444,466]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="mabuensis">
<emphasis box="[703,786,444,466]" italics="true" pageId="19" pageNumber="20">mabuensis</emphasis>
</taxonomicName>
) and smaller (
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[300,371,472,494]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="smithersi">
<emphasis box="[300,371,472,494]" italics="true" pageId="19" pageNumber="20">smithersi</emphasis>
</taxonomicName>
) species, although it is overlapped by measurements of the
<typeStatus box="[384,425,500,522]" pageId="19" pageNumber="20">type</typeStatus>
(GSL
<quantity box="[505,593,500,522]" metricMagnitude="-2" metricUnit="m" metricValue="2.7399999999999998" pageId="19" pageNumber="20" unit="mm" value="27.4">27.4 mm</quantity>
) and cotype (GSL
<quantity box="[155,243,529,551]" metricMagnitude="-2" metricUnit="m" metricValue="2.76" pageId="19" pageNumber="20" unit="mm" value="27.6">27.6 mm</quantity>
) of
<taxonomicName authorityName="Petersi" authorityYear="1878" box="[284,416,528,550]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[284,416,528,550]" italics="true" pageId="19" pageNumber="20">R. hildebrandtii</emphasis>
</taxonomicName>
(
<tableCitation box="[431,507,529,551]" captionStart="Table 5" captionStartId="17.[153,217,487,508]" captionText="Table 5. Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the Rhinolophus hildebrandtii complex." pageId="19" pageNumber="20">Table 5</tableCitation>
). Frontal depression reduced in
<typeStatus box="[186,269,557,579]" pageId="19" pageNumber="20">holotype</typeStatus>
and five other members of this species examined; sagittal crest very pronounced, much higher than rostral chamber in lateral profile and commencing just posterior to the reduced frontal depression; in this respect it can be distinguished from
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[155,216,669,691]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="cohenae">
<emphasis box="[155,216,669,691]" italics="true" pageId="19" pageNumber="20">cohenae</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[273,356,669,691]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="mabuensis">
<emphasis box="[273,356,669,691]" italics="true" pageId="19" pageNumber="20">mabuensis</emphasis>
</taxonomicName>
in which the sagittal crest is reduced, and starts further back mid-dorsally in the inter-orbital region behind a well-developed frontal depression. The baculum conforms to the ‘‘
<typeStatus box="[164,217,753,775]" pageId="19" pageNumber="20">Type</typeStatus>
2’’ (shaft wide in dorsal view and horizontal in lateral view with rounded tip) and measures
<quantity box="[484,572,782,804]" metricMagnitude="-3" metricUnit="m" metricValue="3.14" pageId="19" pageNumber="20" unit="mm" value="3.14">3.14 mm</quantity>
in total length in the
<typeStatus box="[155,238,810,832]" pageId="19" pageNumber="20">holotype</typeStatus>
(
<figureCitation box="[252,308,810,832]" captionStart="Figure 9" captionStartId="13.[155,220,1417,1436]" captionTargetBox="[178,1109,164,1382]" captionTargetId="figure-203@13.[155,1113,162,1383]" captionTargetPageId="13" captionText="Figure 9. Dorsal (D), ventral (V) and lateral (L) view of bacula (tips on right) from four individuals (a–d) from Mpumalanga (Clade 1a = cohenae sp. nov.), two (e–f) from lowland sites in Mozambique (Clade 2 = mossambicus sp. nov.) and one (g) from Mt Mabu in Mozambique (Clade 1b = mabuensis sp. nov.). a = DM 11558 (Sudwala); b = DM 11620 (Barberton Tunnel; Topotype of cohenae); c = DM 11560 (Mayo); d = DM 11618 (Barberton Tunnel); e = DM 8580 (Gorongosa); f = DM 8578 (Niassa GR; Holotype of mossambicus); g = DM 10842 (Mt Mabu; Holotype of mabuensis). Bacula of Clade 1a (cohenae sp. nov.) have spatulate tip (rounded in Clades 2 (mossambicus sp. nov.) and 1b (mabuensis sp. nov.))ı typically emarginated basal portion (less so in Clades 2 and 1b) and shaft laterally compressed (cylindrical in Clades 2 and 1b) and sloping downwards in lateral view (horizontal in Clades 2 and 1b). doi:10.1371/journal.pone.0041744.g009" figureDoi="http://doi.org/10.5281/zenodo.4244578" httpUri="https://zenodo.org/record/4244578/files/figure.png" pageId="19" pageNumber="20">Fig. 9</figureCitation>
;
<tableCitation box="[319,395,810,832]" captionStart="Table 5" captionStartId="17.[153,217,487,508]" captionText="Table 5. Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the Rhinolophus hildebrandtii complex." pageId="19" pageNumber="20">Table 5</tableCitation>
). The anterior upper premolar is minute and external to the toothrow (in the
<typeStatus box="[507,590,838,860]" pageId="19" pageNumber="20">holotype</typeStatus>
and three additional specimens examined; in a fourth (DM 11276) the anterior premolar was absent), as found also in
<taxonomicName authorityName="Anderson" authorityYear="1905" box="[544,641,894,916]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="eloquens">
<emphasis box="[544,641,894,916]" italics="true" pageId="19" pageNumber="20">R. eloquens</emphasis>
</taxonomicName>
but not in any other members of the
<taxonomicName authorityName="Petersi" authorityYear="1878" box="[397,500,922,944]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[397,500,922,944]" italics="true" pageId="19" pageNumber="20">hildebrandtii</emphasis>
</taxonomicName>
complex. There is a single mental groove in the
<typeStatus box="[364,447,950,972]" pageId="19" pageNumber="20">holotype</typeStatus>
and in three other members of this species examined. Genotypes classify
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[521,655,978,1000]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="mossambicus">
<emphasis box="[521,655,978,1000]" italics="true" pageId="19" pageNumber="20">R. mossambicus</emphasis>
</taxonomicName>
in Clade 2.
</paragraph>
</subSubSection>
<subSubSection pageId="19" pageNumber="20" type="materials_examined">
<paragraph blockId="19.[155,793,163,1956]" pageId="19" pageNumber="20">
<materialsCitation ID-GBIF-Occurrence="2908237313" accessionNumber="JQ929254, JQ929210, JQ929288" collectionCode="DM" country="Mozambique" location="Namapa" pageId="19" pageNumber="20" specimenCode="DM 8577" specimenCount="1" specimenCount-female="1" stateProvince="northern Mozambique" typeStatus="paratype">
<emphasis bold="true" box="[177,292,1008,1027]" pageId="19" pageNumber="20">
<typeStatus box="[177,287,1008,1027]" pageId="19" pageNumber="20">Paratypes</typeStatus>
.
</emphasis>
<specimenCode box="[316,416,1006,1028]" pageId="19" pageNumber="20">DM 8577</specimenCode>
<specimenCount box="[427,551,1006,1028]" pageId="19" pageNumber="20" type="adult">adult female</specimenCount>
alcohol and skull from
<location LSID="urn:lsid:plazi:treatment:B76E487AFFFDFFD7FC810740FDA910F1:3A18AFB7FFFCFFD7FF520520FF641374" box="[155,240,1035,1057]" country="Mozambique" municipality="Chinizuia" name="Namapa" pageId="19" pageNumber="20" stateProvince="northern Mozambique">Namapa</location>
,
<collectingRegion box="[254,480,1035,1057]" pageId="19" pageNumber="20">northern Mozambique</collectingRegion>
;
</materialsCitation>
<materialsCitation ID-GBIF-Occurrence="2908237305" accessionNumber="JQ929257, JQ929213, JQ929291" collectionCode="DM" country="Mozambique" municipality="Chinizuia" pageId="19" pageNumber="20" specimenCode="DM 8579" specimenCount="1" specimenCount-adult="1" specimenCount-male="1" stateProvince="northern Mozambique" typeStatus="paratype">
<specimenCode box="[493,592,1035,1057]" pageId="19" pageNumber="20">DM 8579</specimenCode>
<specimenCount box="[602,707,1035,1057]" pageId="19" pageNumber="20" type="adult">adult male</specimenCount>
alcohol, skull and baculum from
<collectingMunicipality box="[414,511,1063,1085]" pageId="19" pageNumber="20">Chinizuia</collectingMunicipality>
,
<collectingRegion box="[527,736,1063,1085]" pageId="19" pageNumber="20">central Mozambique</collectingRegion>
;
</materialsCitation>
<materialsCitation ID-GBIF-Occurrence="2908237311" accessionNumber="JQ929256, JQ929212, JQ929290" collectionCode="DM" country="Mozambique" location="Gorongosa Caves" pageId="19" pageNumber="20" specimenCode="DM 8580" specimenCount="1" specimenCount-adult="1" specimenCount-male="1" stateProvince="northern Mozambique" typeStatus="paratype">
<specimenCode pageId="19" pageNumber="20">DM 8580</specimenCode>
<specimenCount box="[216,324,1091,1113]" pageId="19" pageNumber="20" type="male">adult male</specimenCount>
alcohol, skull and baculum from
<location LSID="urn:lsid:plazi:treatment:B76E487AFFFDFFD7FC810740FDA910F1:3A18AFB7FFFCFFD7FD670568FF431320" country="Mozambique" municipality="Chinizuia" name="Gorongosa Caves" pageId="19" pageNumber="20" stateProvince="northern Mozambique">Gorongosa Caves</location>
, central
<collectingCountry box="[297,428,1119,1141]" name="Mozambique" pageId="19" pageNumber="20">Mozambique</collectingCountry>
;
</materialsCitation>
<materialsCitation ID-GBIF-Occurrence="2908237315" accessionNumber="JQ929253, JQ929209, JQ929233, JQ929301" collectionCode="DM" country="Mozambique" location="Gerhard's Cave" pageId="19" pageNumber="20" specimenCode="DM 11276" specimenCount="1" specimenCount-female="1" stateProvince="southern Mozambique" typeStatus="paratype">
<specimenCode box="[438,546,1119,1141]" pageId="19" pageNumber="20">DM 11276</specimenCode>
,
<specimenCount box="[556,674,1119,1141]" pageId="19" pageNumber="20" type="female">adult female</specimenCount>
alcohol and skull from
<location LSID="urn:lsid:plazi:treatment:B76E487AFFFDFFD7FC810740FDA910F1:3A18AFB7FFFCFFD7FECD0550FE3413C4" box="[260,416,1147,1169]" country="Mozambique" municipality="Chinizuia" name="Gerhard's Cave" pageId="19" pageNumber="20" stateProvince="southern Mozambique">Gerhard’s Cave</location>
,
<collectingRegion box="[428,652,1147,1169]" pageId="19" pageNumber="20">southern Mozambique</collectingRegion>
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="19" pageNumber="20" type="description">
<paragraph blockId="19.[155,793,163,1956]" pageId="19" pageNumber="20">
<emphasis bold="true" box="[177,311,1176,1195]" pageId="19" pageNumber="20">Description.</emphasis>
Generally conforms to the previously published keys and description [
<bibRefCitation author="Csorba G &amp; Ujhelyip P &amp; Thomas N" box="[384,412,1203,1225]" journalOrPublisher="Shropshire: Alana books" pageId="19" pageNumber="20" refId="ref21181" refString="12. Csorba G, Ujhelyip P, Thomas N (2003) Horseshoe Bats of the World (Chiroptera: Rhinolophidae). Shropshire: Alana books." title="Horseshoe Bats of the World (Chiroptera: Rhinolophidae)" type="book" year="2003">12</bibRefCitation>
], i.e. a large-sized bat (forearm
<quantity metricMagnitude="-2" metricUnit="m" metricValue="6.25" metricValueMax="6.5" metricValueMin="6.0" pageId="19" pageNumber="20" unit="mm" value="62.5" valueMax="65.0" valueMin="60.0">60– 65 mm</quantity>
;
<quantity box="[242,330,1231,1253]" metricMagnitude="-2" metricUnit="m" metricValue="6.39" pageId="19" pageNumber="20" unit="mm" value="63.9">63.9 mm</quantity>
in
<typeStatus box="[370,453,1231,1253]" pageId="19" pageNumber="20">holotype</typeStatus>
) with medium or large ears (
<quantity metricMagnitude="-2" metricUnit="m" metricValue="3.25" metricValueMax="3.7" metricValueMin="2.8" pageId="19" pageNumber="20" unit="mm" value="32.5" valueMax="37.0" valueMin="28.0">28– 37 mm</quantity>
;
<quantity box="[245,333,1259,1281]" metricMagnitude="-2" metricUnit="m" metricValue="3.26" pageId="19" pageNumber="20" unit="mm" value="32.6">32.6 mm</quantity>
in
<typeStatus box="[380,465,1259,1281]" pageId="19" pageNumber="20">holotype</typeStatus>
), wide horseshoe (
<quantity box="[663,789,1259,1281]" metricMagnitude="-2" metricUnit="m" metricValue="1.3599999999999999" metricValueMax="1.42" metricValueMin="1.3" pageId="19" pageNumber="20" unit="mm" value="13.6" valueMax="14.2" valueMin="13.0">13–14.2 mm</quantity>
;
<quantity box="[155,243,1288,1310]" metricMagnitude="-2" metricUnit="m" metricValue="1.42" pageId="19" pageNumber="20" unit="mm" value="14.2">14.2 mm</quantity>
in
<typeStatus box="[277,360,1288,1310]" pageId="19" pageNumber="20">holotype</typeStatus>
) covering the muzzle, sella constricted at its proximal third, almost parallel-sided above and having long hairs, lancet relatively long and straight-sized, lower lip with single median groove, upper parts greyish-brown, with individual hairs long and unicoloured, underparts same colour or very slightly paler, skull large and heavily built (
<figureCitation box="[502,558,1428,1450]" captionStart="Figure 9" captionStartId="13.[155,220,1417,1436]" captionTargetBox="[178,1109,164,1382]" captionTargetId="figure-203@13.[155,1113,162,1383]" captionTargetPageId="13" captionText="Figure 9. Dorsal (D), ventral (V) and lateral (L) view of bacula (tips on right) from four individuals (a–d) from Mpumalanga (Clade 1a = cohenae sp. nov.), two (e–f) from lowland sites in Mozambique (Clade 2 = mossambicus sp. nov.) and one (g) from Mt Mabu in Mozambique (Clade 1b = mabuensis sp. nov.). a = DM 11558 (Sudwala); b = DM 11620 (Barberton Tunnel; Topotype of cohenae); c = DM 11560 (Mayo); d = DM 11618 (Barberton Tunnel); e = DM 8580 (Gorongosa); f = DM 8578 (Niassa GR; Holotype of mossambicus); g = DM 10842 (Mt Mabu; Holotype of mabuensis). Bacula of Clade 1a (cohenae sp. nov.) have spatulate tip (rounded in Clades 2 (mossambicus sp. nov.) and 1b (mabuensis sp. nov.))ı typically emarginated basal portion (less so in Clades 2 and 1b) and shaft laterally compressed (cylindrical in Clades 2 and 1b) and sloping downwards in lateral view (horizontal in Clades 2 and 1b). doi:10.1371/journal.pone.0041744.g009" figureDoi="http://doi.org/10.5281/zenodo.4244578" httpUri="https://zenodo.org/record/4244578/files/figure.png" pageId="19" pageNumber="20">Fig. 9</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="19" pageNumber="20" type="distribution">
<paragraph blockId="19.[155,793,163,1956]" pageId="19" pageNumber="20">
<emphasis bold="true" box="[177,317,1457,1476]" pageId="19" pageNumber="20">Distribution.</emphasis>
Based on genetic data, the species is known from five localities in Mozambique from altitudes of,
<quantity box="[729,792,1484,1506]" metricMagnitude="2" metricUnit="m" metricValue="5.0" pageId="19" pageNumber="20" unit="m" value="500.0">500 m</quantity>
(
<figureCitation box="[161,224,1512,1534]" captionStart="Figure 1" captionStartId="1.[155,220,1840,1859]" captionTargetBox="[158,790,800,1803]" captionTargetId="figure-761@1.[155,793,797,1806]" captionTargetPageId="1" captionText="Figure 1. Portraits of (a) Rhinolophus smithersi species novo, and (b) Rhinolophus mossambicus species novo, two of four new cryptic species described herein within the R. hildebrandtii complex. doi:10.1371/journal.pone.0041744.g001" figureDoi="http://doi.org/10.5281/zenodo.4244562" httpUri="https://zenodo.org/record/4244562/files/figure.png" pageId="19" pageNumber="20">Fig. 1</figureCitation>
: Chinizuia Forest, Gerhard’s Cave, Gorongosa Caves, Namapa and Niassa Game Reserve) and one locality, Lutope-Ngolangola Confluence in NW Zimbabwe. These scattered records across the subregion suggest that
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[618,761,1596,1618]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="mossambicus">
<emphasis box="[618,761,1596,1618]" italics="true" pageId="19" pageNumber="20">R. mossambicus</emphasis>
</taxonomicName>
is widespread across the savanna biome of Zimbabwe and Mozambique. Its overall range awaits elucidation (
<figureCitation box="[577,633,1653,1675]" captionStart="Figure 2" captionStartId="2.[155,220,1881,1900]" captionTargetBox="[160,1475,167,1843]" captionTargetId="figure-0@2.[155,1480,162,1848]" captionTargetPageId="2" captionText="Figure 2. Map of southern, central and eastern Africa indicating localities of individuals of R. hildebrandtii species-complex included in this study. Grey-shaded area represents elevations in excess of 600 m a.s.l. Closed squares indicate museum specimens from which craniometric data were obtained. Open symbols indicate specimens genotyped in this study. The distribution of the three major clades is based on cytochrome b (see Figure 2): open circles = Clade 1; open squares = Clade 2; open diamonds = Clade 3. Closed squares enclosed in open symbols indicate localities where both molecular and morphological data were available for selected specimens. Numbers refer to respective localities listed in Table S1. ‘‘T’’ indicates the type localities of R. eloquens in Uganda and R. hildebrandtii in Kenyaı respectively. doi:10.1371/journal.pone.0041744.g002" figureDoi="http://doi.org/10.5281/zenodo.4244564" httpUri="https://zenodo.org/record/4244564/files/figure.png" pageId="19" pageNumber="20">Fig. 2</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="19" pageNumber="20" type="biology_ecology">
<paragraph blockId="19.[155,793,163,1956]" pageId="19" pageNumber="20">
<emphasis bold="true" box="[177,267,1683,1702]" pageId="19" pageNumber="20">Ecology.</emphasis>
Associated with the southern savanna biome. Based on five localities in Mozambique, its range extends from
<quantity metricMagnitude="2" metricUnit="m" metricValue="2.8" metricValueMax="5.0" metricValueMin="0.6" pageId="19" pageNumber="20" unit="m" value="280.0" valueMax="500.0" valueMin="60.0">60– 500 m</quantity>
, and to
<quantity box="[303,378,1737,1759]" metricMagnitude="3" metricUnit="m" metricValue="1.0" pageId="19" pageNumber="20" unit="m" value="1000.0">1000 m</quantity>
in NW Zimbabwe.
</paragraph>
</subSubSection>
<subSubSection pageId="19" pageNumber="20" type="etymology">
<paragraph blockId="19.[155,793,163,1956]" pageId="19" pageNumber="20">
<emphasis bold="true" box="[177,298,1767,1786]" pageId="19" pageNumber="20">Etymology.</emphasis>
The name denotes the country of origin (Mozambique) of the
<typeStatus box="[354,395,1794,1816]" pageId="19" pageNumber="20">type</typeStatus>
specimen of this species. Additional sequenced specimens from Mozambique suggest that the species is widely distributed throughout this country. However
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="mossambicus">
<emphasis italics="true" pageId="19" pageNumber="20">R. mossambicus</emphasis>
</taxonomicName>
was not recorded on montane islands (Mts Mabu and Inago) to which populations of
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[507,619,1905,1927]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="mabuensis">
<emphasis box="[507,619,1905,1927]" italics="true" pageId="19" pageNumber="20">R. mabuensis</emphasis>
</taxonomicName>
are restricted.
</paragraph>
</subSubSection>
<subSubSection box="[177,573,1934,1956]" pageId="19" pageNumber="20" type="materials_examined">
<paragraph blockId="19.[155,793,163,1956]" box="[177,573,1934,1956]" pageId="19" pageNumber="20">
<emphasis bold="true" box="[177,412,1935,1954]" pageId="19" pageNumber="20">Specimens examined.</emphasis>
<collectingRegion box="[436,470,1934,1956]" country="South Sudan" name="Lakes" pageId="19" pageNumber="20">See</collectingRegion>
<tableCitation box="[478,569,1934,1956]" captionStart="Table 1" captionStartId="6.[171,235,1481,1502]" captionText="Table 1. ‘Uncorrected p’ pairwise sequence distances for the 12S gene below the diagonal and Chd1 gene above the diagonal for the clades (1ı 2ı 3) and some lineages (1aı 1bı 1c) identified in the molecular analyses of the Rhinolophus hildebrandtii complex." pageId="19" pageNumber="20">Table S1</tableCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>

341
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@ -0,0 +1,341 @@
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<mods:title>Four New Bat Species (Rhinolophus hildebrandtii Complex) Reflect Plio-Pleistocene Divergence of Dwarfs and Giants across an Afromontane Archipelago</mods:title>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Peter J. Taylor</mods:namePart>
<mods:affiliation>Department of Ecology and Resource Management, University of Venda, Thohoyandou, South Africa, Durban Natural Science Museum, Durban, South Africa, School of Life Sciences, University of KwaZulu-Natal, Durban, South Africa</mods:affiliation>
<mods:nameIdentifier type="email">peter.taylor@univen.ac.za</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Samantha Stoffberg</mods:namePart>
<mods:affiliation>Evolutionary Genomics Group, Department of Botany and Zoology, University of Stellenbosch, Stellenbosch, South Africa</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ara Monadjem</mods:namePart>
<mods:affiliation>All Out Africa Research Unit, Department of Biological Sciences, University of Swaziland, Kwaluseni, Swaziland</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Martinus Corrie Schoeman</mods:namePart>
<mods:affiliation>School of Life Sciences, University of KwaZulu-Natal, Durban, South Africa</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Julian Baylis</mods:namePart>
<mods:affiliation>Mulanje Mountain Conservation Trust, Mulanje, Malawi, Conservation Science Group, Department of Zoology, University of Cambridge, Cambridge, United Kingdom</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Fenton P. D. Cotterill</mods:namePart>
<mods:affiliation>Africa Earth Observatory Network, Geoecodynamics Research Hub, Department of Botany and Zoology, University of Stellenbosch, Stellenbosch, South Africa</mods:affiliation>
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<mods:date>2012</mods:date>
<mods:detail type="pubDate">
<mods:number>2012-09-12</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>7</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>9</mods:number>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.4328637" ID-GBIF-Taxon="168754886" ID-Zenodo-Dep="4328637" LSID="urn:lsid:plazi:treatment:B76E487AFFFFFFD4FF520409FB7F13B7" httpUri="http://treatment.plazi.org/id/B76E487AFFFFFFD4FF520409FB7F13B7" lastPageNumber="17" pageId="16" pageNumber="17">
<paragraph blockId="16.[155,793,1314,1956]" pageId="16" pageNumber="17">
<subSubSection box="[155,560,1314,1338]" pageId="16" pageNumber="17" type="nomenclature">
<heading box="[155,560,1314,1338]" fontSize="10" level="2" pageId="16" pageNumber="17" reason="1">
<taxonomicName authority="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill, 2012" authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[155,413,1314,1338]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="mabuensis" status="new species">
<emphasis box="[155,413,1314,1338]" italics="true" pageId="16" pageNumber="17">Rhinolophus mabuensis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[422,560,1314,1338]" pageId="16" pageNumber="17" rank="species">new species</taxonomicNameLabel>
</heading>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="description">
<uri pageId="16" pageNumber="17">
urn:lsid:zoobank.org:act:
<uuid pageId="16" pageNumber="17">C205928E-D72C-40D5-9647FDC9391CB1A6</uuid>
</uri>
</subSubSection>
</paragraph>
<subSubSection box="[179,462,1402,1424]" pageId="16" pageNumber="17" type="vernacular_names">
<paragraph blockId="16.[155,793,1314,1956]" box="[179,462,1402,1424]" pageId="16" pageNumber="17">Mount Mabu Horseshoe Bat</paragraph>
</subSubSection>
<subSubSection box="[179,705,1431,1453]" pageId="16" pageNumber="17" type="description">
<paragraph blockId="16.[155,793,1314,1956]" box="[179,705,1431,1453]" pageId="16" pageNumber="17">
<figureCitation box="[179,263,1431,1453]" captionStart="Figure 4" captionStartId="8.[840,905,1377,1396]" captionTargetBox="[844,1474,166,1343]" captionTargetId="figure-696@8.[840,1478,162,1343]" captionTargetPageId="8" captionText="Figure 4. Morphometric variation in a series representing the R. hildebrandtii complex from Lutope-Ngolangola, Zimbabwe: a) biplot of forearm length versus noseleaf width and b) PCA of five craniometric variables (M3M3, CM3, IOC, NW, NH) in 26 individuals of known (37 or 46 kHz) and unknown (?) frequency. Females indicated by open circlesı males by closed circles or crosses or asterisk. Voucher specimens for molecular sequencing study indicated by asterisk (Clade 1e: = smithersii sp. nov.; see Taxonomic Conclusions) and crosses (Clade 2: = mossambicus sp. nov.; see Taxonomic Conclusions). Hereafterı all individuals with a frequency of 37 kHz were assumed to belong to Clade 2 (mossambicus sp. nov.) and the 46 kHz individual was assumed to belong to Clade 1e (smithersi sp. nov.). doi:10.1371/journal.pone.0041744.g004" figureDoi="http://doi.org/10.5281/zenodo.4244568" httpUri="https://zenodo.org/record/4244568/files/figure.png" pageId="16" pageNumber="17">Fig. 4, 4</figureCitation>
,
<figureCitation box="[275,345,1431,1453]" captionStart="Figure 1" captionStartId="1.[155,220,1840,1859]" captionTargetBox="[158,790,800,1803]" captionTargetId="figure-761@1.[155,793,797,1806]" captionTargetPageId="1" captionText="Figure 1. Portraits of (a) Rhinolophus smithersi species novo, and (b) Rhinolophus mossambicus species novo, two of four new cryptic species described herein within the R. hildebrandtii complex. doi:10.1371/journal.pone.0041744.g001" figureDoi="http://doi.org/10.5281/zenodo.4244562" httpUri="https://zenodo.org/record/4244562/files/figure.png" pageId="16" pageNumber="17">Fig. S1</figureCitation>
,
<tableCitation box="[358,436,1431,1453]" captionStart="Table 3" captionStartId="7.[171,235,1322,1343]" captionText="Table 3. Summary of data for peak echolocation frequency of R. hildebrandtii s.l. populations." pageId="16" pageNumber="17">Table 3</tableCitation>
,
<tableCitation box="[448,462,1431,1453]" captionStart="Table 4" captionStartId="15.[856,920,1521,1542]" captionText="Table 4. Eigenvectors for PCA of recently collected voucher specimens of the Rhinolophus hildebrandtii complex associated with molecular and acoustic analysesı which includes type specimens of R. hildebrandtii and R. eloquens (see Fig. 5a). doi:10.1371/journal.pone.0041744.t004" pageId="16" pageNumber="17">4</tableCitation>
,
<tableCitation box="[474,565,1431,1453]" captionStart="Table 1" captionStartId="6.[171,235,1481,1502]" captionText="Table 1. ‘Uncorrected p’ pairwise sequence distances for the 12S gene below the diagonal and Chd1 gene above the diagonal for the clades (1ı 2ı 3) and some lineages (1aı 1bı 1c) identified in the molecular analyses of the Rhinolophus hildebrandtii complex." pageId="16" pageNumber="17">Table S1</tableCitation>
, Appendix S1
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="materials_examined">
<materialsCitation ID-GBIF-Occurrence="2908237312" accessionNumber="JQ929269, JQ929205, JQ292229" collectingDate="2008-10-13" collectionCode="DM" collectorName="M. Curran &amp; M. Kopp" country="South Africa" elevation="1043m" location="Mt Mabu" pageId="16" pageNumber="17" specimenCode="DM 10842" specimenCount="1" specimenCount-adult="1" specimenCount-male="1" stateProvince="northern Mozambique" typeStatus="holotype">
<paragraph blockId="16.[155,793,1314,1956]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[177,282,1460,1479]" pageId="16" pageNumber="17">
<typeStatus box="[177,277,1460,1479]" pageId="16" pageNumber="17">Holotype</typeStatus>
.
</emphasis>
<specimenCode box="[306,421,1459,1481]" pageId="16" pageNumber="17">DM 10842</specimenCode>
;
<specimenCount box="[437,486,1459,1481]" pageId="16" pageNumber="17" type="adult">adult</specimenCount>
<specimenCount box="[498,547,1459,1481]" pageId="16" pageNumber="17" type="male">male</specimenCount>
, alcohol skin, skull and prepared baculum, collected by
<collectorName box="[485,596,1487,1509]" pageId="16" pageNumber="17">M. Curran</collectorName>
and
<collectorName box="[656,753,1487,1509]" pageId="16" pageNumber="17">M. Kopp</collectorName>
,
<date pageId="16" pageNumber="17" value="2008-10-13">
<collectingDate pageId="16" pageNumber="17" value="2008-10-13">13 October 2008</collectingDate>
</date>
.
</paragraph>
<paragraph blockId="16.[155,793,1314,1956]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[177,329,1544,1563]" pageId="16" pageNumber="17">Type locality.</emphasis>
<location LSID="urn:lsid:plazi:treatment:B76E487AFFFFFFD4FF520409FB7F13B7:3A18AFB7FFFFFFD4FEA8072CFE531148" box="[353,455,1543,1565]" name="Mt Mabu" pageId="16" pageNumber="17" stateProvince="northern Mozambique">Mt Mabu</location>
,
<collectingRegion box="[471,699,1543,1565]" pageId="16" pageNumber="17">northern Mozambique</collectingRegion>
,
<elevation pageId="16" pageNumber="17">
<geoCoordinate pageId="16" pageNumber="17">
16
<emphasis bold="true" box="[739,748,1542,1563]" pageId="16" pageNumber="17">°</emphasis>
<date pageId="16" pageNumber="17" value="2017-02-09">
17
<emphasis box="[784,792,1542,1564]" italics="true" pageId="16" pageNumber="17">'</emphasis>
2
</date>
<emphasis box="[167,175,1570,1592]" italics="true" pageId="16" pageNumber="17">&quot;</emphasis>
S
</geoCoordinate>
;
<geoCoordinate box="[212,361,1570,1593]" pageId="16" pageNumber="17">
36
<emphasis bold="true" box="[236,245,1570,1591]" pageId="16" pageNumber="17">°</emphasis>
<date box="[263,336,1570,1593]" pageId="16" pageNumber="17" value="1953-09-23">
23
<emphasis box="[287,295,1570,1592]" italics="true" pageId="16" pageNumber="17">'</emphasis>
53
</date>
<emphasis box="[336,344,1570,1592]" italics="true" pageId="16" pageNumber="17">&quot;</emphasis>
E
</geoCoordinate>
</elevation>
; elevation
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<elevation box="[488,562,1571,1593]" metricMagnitude="3" metricUnit="m" metricValue="1.043" pageId="16" pageNumber="17" unit="m" value="1043.0">1043 m</elevation>
</quantity>
asl.
</paragraph>
</materialsCitation>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="diagnosis">
<paragraph blockId="16.[155,793,1314,1956]" lastBlockId="16.[840,1479,164,1250]" pageId="16" pageNumber="17">
Diagnosis. Peak frequency of 37.5 and 37.9 kHz for two animals captured and released in sub-montane and montane forests at
<quantity box="[678,741,1627,1649]" metricMagnitude="2" metricUnit="m" metricValue="5.5" pageId="16" pageNumber="17" unit="m" value="550.0">550 m</quantity>
and
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respectively on
<collectingRegion box="[397,495,1655,1677]" pageId="16" pageNumber="17">Mt Mabu</collectingRegion>
(Curran and Kopp, personal communication). The echolocation call frequencies of the
<typeStatus box="[709,792,1683,1705]" pageId="16" pageNumber="17">holotype</typeStatus>
from
<collectingRegion box="[216,317,1711,1733]" pageId="16" pageNumber="17">Mt Mabu</collectingRegion>
and the
<typeStatus box="[427,511,1711,1733]" pageId="16" pageNumber="17">paratype</typeStatus>
from Mt Inago were not measured. Body, cranium and noseleaf very large (FL
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in
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, 69.0 in
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; GSL
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in
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;
<quantity box="[704,792,1766,1788]" metricMagnitude="-2" metricUnit="m" metricValue="3.03" pageId="16" pageNumber="17" unit="mm" value="30.3">30.3 mm</quantity>
in
<typeStatus box="[184,271,1794,1816]" pageId="16" pageNumber="17">paratype</typeStatus>
; CCL
<quantity box="[345,433,1794,1816]" metricMagnitude="-2" metricUnit="m" metricValue="2.52" pageId="16" pageNumber="17" unit="mm" value="25.2">25.2 mm</quantity>
in
<typeStatus box="[474,560,1794,1816]" pageId="16" pageNumber="17">holotype</typeStatus>
,
<quantity box="[574,661,1794,1816]" metricMagnitude="-2" metricUnit="m" metricValue="2.59" pageId="16" pageNumber="17" unit="mm" value="25.9">25.9 mm</quantity>
in
<typeStatus box="[702,789,1794,1816]" pageId="16" pageNumber="17">paratype</typeStatus>
; NLW 15.0 mm in
<typeStatus box="[342,428,1822,1844]" pageId="16" pageNumber="17">holotype</typeStatus>
,
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in
<typeStatus box="[562,649,1822,1844]" pageId="16" pageNumber="17">paratype</typeStatus>
;
<tableCitation box="[661,738,1822,1844]" captionStart="Table 5" captionStartId="17.[153,217,487,508]" captionText="Table 5. Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the Rhinolophus hildebrandtii complex." pageId="16" pageNumber="17">Table 5</tableCitation>
); size and position of anterior upper premolar variable; either relatively largeand situated partially within the toothrow, with gap between canine and posterior premolar (
<typeStatus box="[481,564,1906,1928]" pageId="16" pageNumber="17">paratype</typeStatus>
) or small and located external to the toothrow (
<typeStatus box="[416,501,1934,1956]" pageId="16" pageNumber="17">holotype</typeStatus>
). Much larger than all other members of the
<taxonomicName authorityName="Peters" authorityYear="1878" box="[999,1130,164,186]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[999,1130,164,186]" italics="true" pageId="16" pageNumber="17">R. hildebrandtii</emphasis>
</taxonomicName>
complex, except for
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[1335,1424,164,186]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="cohenae">
<emphasis box="[1335,1424,164,186]" italics="true" pageId="16" pageNumber="17">R. cohenae</emphasis>
</taxonomicName>
from which
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[909,992,192,214]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="mabuensis">
<emphasis box="[909,992,192,214]" italics="true" pageId="16" pageNumber="17">mabuensis</emphasis>
</taxonomicName>
can be distinguished by echolocation frequency (33 kHz in
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[949,1037,219,241]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="cohenae">
<emphasis box="[949,1037,219,241]" italics="true" pageId="16" pageNumber="17">R. cohenae</emphasis>
</taxonomicName>
;
<emphasis box="[1049,1067,219,241]" italics="true" pageId="16" pageNumber="17">ca</emphasis>
38 kHz in
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[1169,1279,219,241]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="mabuensis">
<emphasis box="[1169,1279,219,241]" italics="true" pageId="16" pageNumber="17">R. mabuensis</emphasis>
</taxonomicName>
) and baculum shape (
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<emphasis box="[955,1038,247,269]" italics="true" pageId="16" pageNumber="17">mabuensis</emphasis>
</taxonomicName>
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<emphasis box="[1163,1224,247,269]" italics="true" pageId="16" pageNumber="17">cohenae</emphasis>
</taxonomicName>
;
<figureCitation box="[1239,1298,248,270]" captionStart="Figure 9" captionStartId="13.[155,220,1417,1436]" captionTargetBox="[178,1109,164,1382]" captionTargetId="figure-203@13.[155,1113,162,1383]" captionTargetPageId="13" captionText="Figure 9. Dorsal (D), ventral (V) and lateral (L) view of bacula (tips on right) from four individuals (a–d) from Mpumalanga (Clade 1a = cohenae sp. nov.), two (e–f) from lowland sites in Mozambique (Clade 2 = mossambicus sp. nov.) and one (g) from Mt Mabu in Mozambique (Clade 1b = mabuensis sp. nov.). a = DM 11558 (Sudwala); b = DM 11620 (Barberton Tunnel; Topotype of cohenae); c = DM 11560 (Mayo); d = DM 11618 (Barberton Tunnel); e = DM 8580 (Gorongosa); f = DM 8578 (Niassa GR; Holotype of mossambicus); g = DM 10842 (Mt Mabu; Holotype of mabuensis). Bacula of Clade 1a (cohenae sp. nov.) have spatulate tip (rounded in Clades 2 (mossambicus sp. nov.) and 1b (mabuensis sp. nov.))ı typically emarginated basal portion (less so in Clades 2 and 1b) and shaft laterally compressed (cylindrical in Clades 2 and 1b) and sloping downwards in lateral view (horizontal in Clades 2 and 1b). doi:10.1371/journal.pone.0041744.g009" figureDoi="http://doi.org/10.5281/zenodo.4244578" httpUri="https://zenodo.org/record/4244578/files/figure.png" pageId="16" pageNumber="17">Fig. 9</figureCitation>
). Genotypes of
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="mabuensis">
<emphasis italics="true" pageId="16" pageNumber="17">R. mabuensis</emphasis>
</taxonomicName>
are members of Clade 1b (
<figureCitation box="[1197,1253,276,298]" captionStart="Figure 2" captionStartId="2.[155,220,1881,1900]" captionTargetBox="[160,1475,167,1843]" captionTargetId="figure-0@2.[155,1480,162,1848]" captionTargetPageId="2" captionText="Figure 2. Map of southern, central and eastern Africa indicating localities of individuals of R. hildebrandtii species-complex included in this study. Grey-shaded area represents elevations in excess of 600 m a.s.l. Closed squares indicate museum specimens from which craniometric data were obtained. Open symbols indicate specimens genotyped in this study. The distribution of the three major clades is based on cytochrome b (see Figure 2): open circles = Clade 1; open squares = Clade 2; open diamonds = Clade 3. Closed squares enclosed in open symbols indicate localities where both molecular and morphological data were available for selected specimens. Numbers refer to respective localities listed in Table S1. ‘‘T’’ indicates the type localities of R. eloquens in Uganda and R. hildebrandtii in Kenyaı respectively. doi:10.1371/journal.pone.0041744.g002" figureDoi="http://doi.org/10.5281/zenodo.4244564" httpUri="https://zenodo.org/record/4244564/files/figure.png" pageId="16" pageNumber="17">Fig. 2</figureCitation>
).
</paragraph>
<paragraph blockId="16.[840,1479,164,1250]" pageId="16" pageNumber="17">
<materialsCitation ID-GBIF-Occurrence="2908237302" accessionNumber="JQ929260, JQ929214, JQ929292" collectingDate="2009-09-05" collectionCode="DM" collectorName="J. Bayliss" country="Mozambique" location="Mt Inago" pageId="16" pageNumber="17" specimenCode="DM 11485" specimenCount="2" specimenCount-adult="1" specimenCount-female="1" typeStatus="paratype">
<emphasis bold="true" box="[863,966,305,324]" pageId="16" pageNumber="17">
<typeStatus box="[863,962,305,324]" pageId="16" pageNumber="17">Paratype</typeStatus>
.
</emphasis>
<specimenCode box="[991,1101,304,326]" pageId="16" pageNumber="17">DM 11485</specimenCode>
(
<specimenCount box="[1117,1165,304,326]" pageId="16" pageNumber="17" type="adult">adult</specimenCount>
<specimenCount box="[1176,1242,304,326]" pageId="16" pageNumber="17" type="female">female</specimenCount>
, alcohol skin and skull, collected
<collectorName box="[932,1019,332,354]" pageId="16" pageNumber="17">J. Bayliss</collectorName>
on
<date box="[1060,1240,332,354]" pageId="16" pageNumber="17" value="2009-09-05">
<collectingDate box="[1060,1240,332,354]" pageId="16" pageNumber="17" value="2009-09-05">5 September 2009</collectingDate>
</date>
from
<location LSID="urn:lsid:plazi:treatment:B76E487AFFFFFFD4FF520409FB7F13B7:3A18AFB7FFFFFFD4FADF0067FAE01637" box="[1302,1396,332,354]" country="Mozambique" name="Mt Inago" pageId="16" pageNumber="17">Mt Inago</location>
)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="description">
<paragraph blockId="16.[840,1479,164,1250]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[863,996,361,380]" pageId="16" pageNumber="17">Description.</emphasis>
External and noseleaf description and colour similar to
<taxonomicName authorityName="Peters" authorityYear="1878" box="[951,1087,388,410]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" sensu="stricto" species="hildebrandtii">
<emphasis box="[951,1087,388,410]" italics="true" pageId="16" pageNumber="17">R. hildebrandtii</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1101,1130,388,410]" pageId="16" pageNumber="17" sensu="stricto">s.s.</taxonomicNameLabel>
, but larger in skull and external dimensions (see above). Lower lip with single mental groove. Noseleaf shape variable with profile of connecting process forming a continuous arch as typical for
<taxonomicName authorityName="Peters" authorityYear="1878" box="[1203,1342,472,494]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[1203,1342,472,494]" italics="true" pageId="16" pageNumber="17">R. hildebrandtii</emphasis>
</taxonomicName>
[
<bibRefCitation author="Csorba G &amp; Ujhelyip P &amp; Thomas N" box="[1364,1391,472,494]" journalOrPublisher="Shropshire: Alana books" pageId="16" pageNumber="17" refId="ref21181" refString="12. Csorba G, Ujhelyip P, Thomas N (2003) Horseshoe Bats of the World (Chiroptera: Rhinolophidae). Shropshire: Alana books." title="Horseshoe Bats of the World (Chiroptera: Rhinolophidae)" type="book" year="2003">12</bibRefCitation>
] in the individual from Mt Inago, but highly distinctive in the Mt Mabu specimen, representing a small rounded bump (
<figureCitation box="[1299,1353,528,550]" captionStart="Figure 8" captionStartId="12.[155,220,993,1012]" captionTargetBox="[163,1000,167,956]" captionTargetId="figure-464@12.[155,1007,162,960]" captionTargetPageId="12" captionText="Figure 8. Photographs showing lateral views of noseleafs of selected individuals (including holotypes of new species) of the Rhinolophus hildebrandtii complex representing molecular Lineage 1a (= cohenae sp. nov.; a–b), Lineage 1b (= mabuensis sp. nov.; c) and Clade 2 (= mossambicus sp. nov.; d–f). a = DM 7886 (cohenae sp. nov.;Barberton Tunnel, Mpumalanga Province, South Africa); b = DM 8626 (cohenae sp. nov.; Barberton Tunnel, Mpumalanga Province, South Africa; Holotype); c = DM 10842 (mabuensis sp. nov.; Mt Mabu, Mozambique; Holotype); d = DM 8578 (mossambicus sp. nov.; Niassa Game Reserve, Mozambique; Holotype); e = DM 8579 (mossambicus sp. nov.; Chinizuia, Mozambique); f = DM 8577 (mossambicus sp. nov.; Namapa, Mozambique). doi:10.1371/journal.pone.0041744.g008" figureDoi="http://doi.org/10.5281/zenodo.4244576" httpUri="https://zenodo.org/record/4244576/files/figure.png" pageId="16" pageNumber="17">Fig. 8</figureCitation>
). The robust skull is reflected in the large means recorded for most cranial dimensions (
<tableCitation box="[964,1042,584,606]" captionStart="Table 5" captionStartId="17.[153,217,487,508]" captionText="Table 5. Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the Rhinolophus hildebrandtii complex." pageId="16" pageNumber="17">Table 5</tableCitation>
). In lateral view the skull has a prominent rostral chamber which extends to the same height as the relatively weak sagittal crest. In dorsal view the V-shaped inter-orbital basin (frontal depression [
<bibRefCitation author="Csorba G &amp; Ujhelyip P &amp; Thomas N" box="[1033,1059,668,690]" journalOrPublisher="Shropshire: Alana books" pageId="16" pageNumber="17" refId="ref21181" refString="12. Csorba G, Ujhelyip P, Thomas N (2003) Horseshoe Bats of the World (Chiroptera: Rhinolophidae). Shropshire: Alana books." title="Horseshoe Bats of the World (Chiroptera: Rhinolophidae)" type="book" year="2003">12</bibRefCitation>
]) formed by the supraorbital ridges is very prominent and deep, and the anterior root of the sagittal crest is displaced posteriorly.
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="distribution">
<paragraph blockId="16.[840,1479,164,1250]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[863,1003,753,772]" pageId="16" pageNumber="17">Distribution.</emphasis>
Known only from two mountains in northern Mozambique but quite possibly extending to nearby Mts. Namuli, Chiperone, Mulanje and the Malawi Rift.
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="biology_ecology">
<paragraph blockId="16.[840,1479,164,1250]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[863,953,837,856]" pageId="16" pageNumber="17">Ecology.</emphasis>
All known specimens were associated with montane or sub-montane forest on the two mountains where they were collected.
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="etymology">
<paragraph blockId="16.[840,1479,164,1250]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[863,984,921,940]" pageId="16" pageNumber="17">Etymology.</emphasis>
We selected the specific epithet to draw attention to the serious threats to the unique biodiversity isolated on the montane forest islands in northern Mozambique – notably Mts Mabu and Inago. None of these landforms lie within formally protected areas, and all are undergoing major habitat degradation and destruction from ever-increasing human activities - hunting, fires, timber harvesting and expanding agriculture [
<bibRefCitation author="Bayliss J &amp; Monteiro J &amp; Fishpool L &amp; Congdon C &amp; Bampton I &amp; Bruessow C &amp; Matimele H &amp; Banze A &amp; Timberlake JR" bookContentInfo="32 pp." box="[1384,1414,1088,1110]" journalOrPublisher="Mulanje Mountain Conservation Trust, Malawi" pageId="16" pageNumber="17" refId="ref21541" refString="22. Bayliss J, Monteiro J, Fishpool L, Congdon C, Bampton I, Bruessow C, Matimele H, Banze A, Timberlake JR (2010) Biodiversity and Conservation of Mount Inago, Mozambique. Report produced under Darwin Initiative Award 15 / 036. Mulanje Mountain Conservation Trust, Malawi. 32 pp." title="Biodiversity and Conservation of Mount Inago, Mozambique. Report produced under Darwin Initiative Award 15 / 036" type="book" year="2010">22</bibRefCitation>
]. The conservation status of this threatened biodiversity on Mts Mabu and Inago is highlighted by
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[1155,1275,1143,1165]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="mabuensis">
<emphasis box="[1155,1275,1143,1165]" italics="true" pageId="16" pageNumber="17">R. mabuensis</emphasis>
</taxonomicName>
, alongside recently discovered species of butterflies, crabs, snakes and chameleons [22;115,116,117,118].
</paragraph>
<paragraph blockId="16.[840,1479,164,1250]" box="[863,1259,1228,1250]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[863,1097,1229,1248]" pageId="16" pageNumber="17">Specimens examined.</emphasis>
<collectingRegion box="[1122,1156,1228,1250]" country="South Sudan" name="Lakes" pageId="16" pageNumber="17">See</collectingRegion>
<tableCitation box="[1163,1254,1228,1250]" captionStart="Table 1" captionStartId="6.[171,235,1481,1502]" captionText="Table 1. ‘Uncorrected p’ pairwise sequence distances for the 12S gene below the diagonal and Chd1 gene above the diagonal for the clades (1ı 2ı 3) and some lineages (1aı 1bı 1c) identified in the molecular analyses of the Rhinolophus hildebrandtii complex." pageId="16" pageNumber="17">Table S1</tableCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>

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<mods:title>Four New Bat Species (Rhinolophus hildebrandtii Complex) Reflect Plio-Pleistocene Divergence of Dwarfs and Giants across an Afromontane Archipelago</mods:title>
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<mods:namePart>Peter J. Taylor</mods:namePart>
<mods:affiliation>Department of Ecology and Resource Management, University of Venda, Thohoyandou, South Africa, Durban Natural Science Museum, Durban, South Africa, School of Life Sciences, University of KwaZulu-Natal, Durban, South Africa</mods:affiliation>
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<mods:namePart>Samantha Stoffberg</mods:namePart>
<mods:affiliation>Evolutionary Genomics Group, Department of Botany and Zoology, University of Stellenbosch, Stellenbosch, South Africa</mods:affiliation>
</mods:name>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Ara Monadjem</mods:namePart>
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</mods:name>
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<mods:role>
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<mods:namePart>Martinus Corrie Schoeman</mods:namePart>
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<mods:namePart>Julian Baylis</mods:namePart>
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</mods:name>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Fenton P. D. Cotterill</mods:namePart>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.4328643" ID-GBIF-Taxon="168754885" ID-Zenodo-Dep="4328643" LSID="urn:lsid:plazi:treatment:B76E487AFFFFFFD6FC81042DFD8913ED" httpUri="http://treatment.plazi.org/id/B76E487AFFFFFFD6FC81042DFD8913ED" lastPageId="18" lastPageNumber="19" pageId="16" pageNumber="17">
<subSubSection box="[840,1266,1286,1310]" pageId="16" pageNumber="17" type="nomenclature">
<paragraph blockId="16.[840,1479,1286,1956]" box="[840,1266,1286,1310]" pageId="16" pageNumber="17">
<heading box="[840,1266,1286,1310]" pageId="16" pageNumber="17">
<taxonomicName authority="Petersi 1878" authorityName="Petersi" authorityYear="1878" box="[840,1266,1286,1310]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[840,1120,1286,1310]" italics="true" pageId="16" pageNumber="17">Rhinolophus hildebrandtii</emphasis>
Petersı 1878
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection box="[864,1147,1317,1339]" pageId="16" pageNumber="17" type="vernacular_names">
<paragraph blockId="16.[840,1479,1286,1956]" box="[864,1147,1317,1339]" pageId="16" pageNumber="17">Hildebrandt’s Horseshoe Bat</paragraph>
</subSubSection>
<subSubSection box="[864,1390,1346,1368]" pageId="16" pageNumber="17" type="description">
<paragraph blockId="16.[840,1479,1286,1956]" box="[864,1390,1346,1368]" pageId="16" pageNumber="17">
<figureCitation box="[864,948,1346,1368]" captionStart="Figure 4" captionStartId="8.[840,905,1377,1396]" captionTargetBox="[844,1474,166,1343]" captionTargetId="figure-696@8.[840,1478,162,1343]" captionTargetPageId="8" captionText="Figure 4. Morphometric variation in a series representing the R. hildebrandtii complex from Lutope-Ngolangola, Zimbabwe: a) biplot of forearm length versus noseleaf width and b) PCA of five craniometric variables (M3M3, CM3, IOC, NW, NH) in 26 individuals of known (37 or 46 kHz) and unknown (?) frequency. Females indicated by open circlesı males by closed circles or crosses or asterisk. Voucher specimens for molecular sequencing study indicated by asterisk (Clade 1e: = smithersii sp. nov.; see Taxonomic Conclusions) and crosses (Clade 2: = mossambicus sp. nov.; see Taxonomic Conclusions). Hereafterı all individuals with a frequency of 37 kHz were assumed to belong to Clade 2 (mossambicus sp. nov.) and the 46 kHz individual was assumed to belong to Clade 1e (smithersi sp. nov.). doi:10.1371/journal.pone.0041744.g004" figureDoi="http://doi.org/10.5281/zenodo.4244568" httpUri="https://zenodo.org/record/4244568/files/figure.png" pageId="16" pageNumber="17">Fig. 4, 4</figureCitation>
,
<figureCitation box="[960,1031,1346,1368]" captionStart="Figure 1" captionStartId="1.[155,220,1840,1859]" captionTargetBox="[158,790,800,1803]" captionTargetId="figure-761@1.[155,793,797,1806]" captionTargetPageId="1" captionText="Figure 1. Portraits of (a) Rhinolophus smithersi species novo, and (b) Rhinolophus mossambicus species novo, two of four new cryptic species described herein within the R. hildebrandtii complex. doi:10.1371/journal.pone.0041744.g001" figureDoi="http://doi.org/10.5281/zenodo.4244562" httpUri="https://zenodo.org/record/4244562/files/figure.png" pageId="16" pageNumber="17">Fig. S1</figureCitation>
,
<tableCitation box="[1043,1121,1346,1368]" captionStart="Table 3" captionStartId="7.[171,235,1322,1343]" captionText="Table 3. Summary of data for peak echolocation frequency of R. hildebrandtii s.l. populations." pageId="16" pageNumber="17">Table 3</tableCitation>
,
<tableCitation box="[1134,1148,1346,1368]" captionStart="Table 4" captionStartId="15.[856,920,1521,1542]" captionText="Table 4. Eigenvectors for PCA of recently collected voucher specimens of the Rhinolophus hildebrandtii complex associated with molecular and acoustic analysesı which includes type specimens of R. hildebrandtii and R. eloquens (see Fig. 5a). doi:10.1371/journal.pone.0041744.t004" pageId="16" pageNumber="17">4</tableCitation>
,
<tableCitation box="[1160,1250,1346,1368]" captionStart="Table 1" captionStartId="6.[171,235,1481,1502]" captionText="Table 1. ‘Uncorrected p’ pairwise sequence distances for the 12S gene below the diagonal and Chd1 gene above the diagonal for the clades (1ı 2ı 3) and some lineages (1aı 1bı 1c) identified in the molecular analyses of the Rhinolophus hildebrandtii complex." pageId="16" pageNumber="17">Table S1</tableCitation>
, Appendix S1
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="materials_examined">
<paragraph blockId="16.[840,1479,1286,1956]" pageId="16" pageNumber="17">
<materialsCitation ID-GBIF-Occurrence="2908237307" collectingDate="1877-07" collectionCode="ZMB" collectorName="S. Hildebrandt" country="Kenya" elevation="390" latitude="-02.7786111" location="Ndi" longitude="038.7716667" pageId="16" pageNumber="17" specimenCode="ZMB 5378" specimenCount="1" specimenCount-male="1" stateProvince="Taita" typeStatus="holotype">
<emphasis bold="true" box="[863,967,1376,1395]" pageId="16" pageNumber="17">
<typeStatus box="[863,963,1376,1395]" pageId="16" pageNumber="17">Holotype</typeStatus>
.
</emphasis>
<specimenCode box="[992,1102,1375,1397]" pageId="16" pageNumber="17">ZMB 5378</specimenCode>
(
<specimenCount box="[1118,1164,1375,1397]" pageId="16" pageNumber="17" type="male">male</specimenCount>
collected by S. Hildebrandt in
<date box="[840,938,1403,1425]" pageId="16" pageNumber="17" value="1877-07">
<collectingDate box="[840,938,1403,1425]" pageId="16" pageNumber="17" value="1877-07">July 1877</collectingDate>
</date>
from
<location LSID="urn:lsid:plazi:treatment:B76E487AFFFFFFD6FC81042DFD8913ED:3A18AFB7FFFFFFD4FC390450FB8312C4" box="[1008,1047,1403,1425]" country="Kenya" name="Ndi" pageId="16" pageNumber="17" stateProvince="Taita">Ndi</location>
,
<collectingRegion box="[1062,1116,1403,1425]" country="Kenya" name="Taita/Taveta" pageId="16" pageNumber="17">Taita</collectingRegion>
,
<collectingCountry box="[1132,1198,1403,1425]" name="Kenya" pageId="16" pageNumber="17" value="Kenya">Kenya</collectingCountry>
).
</materialsCitation>
<materialsCitation ID-GBIF-Occurrence="2908237304" collectingDate="1877-07" collectionCode="ZMB" collectorName="S. Hildebrandt" country="Kenya" elevation="390" latitude="-02.7786111" location="Ndi" longitude="038.7716667" pageId="16" pageNumber="17" specimenCode="ZMB 5379" specimenCount="1" specimenCount-male="1" stateProvince="Taita" typeStatus="cotype">
<typeStatus box="[1220,1290,1403,1425]" pageId="16" pageNumber="17">Cotype</typeStatus>
<specimenCode box="[1302,1414,1403,1425]" pageId="16" pageNumber="17">ZMB 5379</specimenCode>
(
<specimenCount box="[1432,1478,1403,1425]" pageId="16" pageNumber="17" type="male">male</specimenCount>
collected by
<collectorName box="[961,1105,1431,1453]" pageId="16" pageNumber="17">S. Hildebrandt</collectorName>
in
<date box="[1137,1229,1431,1453]" pageId="16" pageNumber="17" value="1877-07">
<collectingDate box="[1137,1229,1431,1453]" pageId="16" pageNumber="17" value="1877-07">July 1877</collectingDate>
</date>
from
<location LSID="urn:lsid:plazi:treatment:B76E487AFFFFFFD6FC81042DFD8913ED:3A18AFB7FFFFFFD4FAC004BCFAA412F8" box="[1289,1328,1431,1453]" country="Kenya" name="Ndi" pageId="16" pageNumber="17" stateProvince="Taita">Ndi</location>
,
<collectingRegion box="[1337,1391,1431,1453]" country="Kenya" name="Taita/Taveta" pageId="16" pageNumber="17">Taita</collectingRegion>
,
<collectingCountry box="[1402,1468,1431,1453]" name="Kenya" pageId="16" pageNumber="17">Kenya</collectingCountry>
).
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="17" type="distribution">
<paragraph blockId="16.[840,1479,1286,1956]" pageId="16" pageNumber="17">
<emphasis bold="true" box="[863,1008,1460,1479]" pageId="16" pageNumber="17">Type locality.</emphasis>
<location LSID="urn:lsid:plazi:treatment:B76E487AFFFFFFD6FC81042DFD8913ED:3A18AFB7FFFFFFD4FBC00498FBA4129C" box="[1033,1072,1459,1481]" country="Kenya" latitude="-9.716666" longitude="9.3" name="Ndi" pageId="16" pageNumber="17" stateProvince="Taita">Ndi</location>
,
<collectingRegion box="[1082,1136,1459,1481]" country="Kenya" name="Taita/Taveta" pageId="16" pageNumber="17">Taita</collectingRegion>
,
<collectingCountry box="[1146,1212,1459,1481]" name="Kenya" pageId="16" pageNumber="17" value="Kenya">Kenya</collectingCountry>
,
<geoCoordinate box="[1223,1335,1458,1481]" pageId="16" pageNumber="17">
2
<emphasis bold="true" box="[1235,1244,1458,1479]" pageId="16" pageNumber="17">°</emphasis>
46
<emphasis box="[1275,1283,1458,1480]" italics="true" pageId="16" pageNumber="17">'</emphasis>
43
<emphasis box="[1313,1321,1458,1480]" italics="true" pageId="16" pageNumber="17">&quot;</emphasis>
S
</geoCoordinate>
;
<geoCoordinate box="[1347,1474,1458,1481]" pageId="16" pageNumber="17">
38
<emphasis bold="true" box="[1371,1380,1458,1479]" pageId="16" pageNumber="17">°</emphasis>
<date box="[1387,1449,1458,1481]" pageId="16" pageNumber="17" value="1946-09-18">
46
<emphasis box="[1411,1419,1458,1480]" italics="true" pageId="16" pageNumber="17">'</emphasis>
18
</date>
<emphasis box="[1449,1457,1458,1480]" italics="true" pageId="16" pageNumber="17">&quot;</emphasis>
E
</geoCoordinate>
; elevation
<quantity box="[935,997,1487,1509]" metricMagnitude="2" metricUnit="m" metricValue="3.9" pageId="16" pageNumber="17" unit="m" value="390.0">
<elevation box="[935,997,1487,1509]" metricMagnitude="2" metricUnit="m" metricValue="3.9" pageId="16" pageNumber="17" unit="m" value="390.0">390 m</elevation>
</quantity>
asl.
</paragraph>
</subSubSection>
<subSubSection lastPageId="18" lastPageNumber="19" pageId="16" pageNumber="17" type="diagnosis">
<paragraph blockId="16.[840,1479,1286,1956]" lastBlockId="18.[155,794,164,1208]" lastPageId="18" lastPageNumber="19" pageId="16" pageNumber="17">
<emphasis bold="true" box="[863,976,1516,1535]" pageId="16" pageNumber="17">Diagnosis.</emphasis>
Individuals from two localities in Kenya, including the
<typeStatus box="[921,962,1543,1565]" pageId="16" pageNumber="17">type</typeStatus>
locality of Taita District, had a peak frequency of 42 kHz [
<bibRefCitation author="Jones G &amp; Van Parijs SM" box="[927,955,1571,1593]" journalOrPublisher="Series B Biological Sciences" pageId="16" pageNumber="17" pagination="119 - 1258" part="251" refId="ref21229" refString="15. Jones G, Van Parijs SM (1993) Bimodal echolocation in pipistrelle bats: are cryptic species present? Proceedings of the Royal Society Series B Biological Sciences 251: 119 - 1258." title="Bimodal echolocation in pipistrelle bats: are cryptic species present? Proceedings of the Royal Society" type="journal article" year="1993">15</bibRefCitation>
]. Intermediate in body and cranial size, with the
<typeStatus box="[1437,1478,1571,1593]" pageId="16" pageNumber="17">type</typeStatus>
and co-type overlapping in cranial dimensions with members of Clade 2 from which they are however easily distinguished genetically based on sequences from three mitochondrial and one nuclear gene (cytochrome b uncorrected genetic divergence 7.7–9.0%). Although there is minor overlap in individual cranial and forearm variables (
<tableCitation box="[1093,1178,1738,1760]" captionStart="Table 5" captionStartId="17.[153,217,487,508]" captionText="Table 5. Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the Rhinolophus hildebrandtii complex." pageId="16" pageNumber="17">Table 5</tableCitation>
), they can be distinguished completely by PCA of cranial dimensions from other members of Clade1 which are distinctly smaller (Lineages 1d, 1e) or larger (Lineages 1a, 1b) (
<figureCitation box="[1026,1098,1822,1844]" captionStart="Figure 5" captionStartId="9.[155,220,1403,1422]" captionTargetBox="[159,791,166,1365]" captionTargetId="figure-699@9.[155,795,162,1369]" captionTargetPageId="9" captionText="Figure 5. Canonical variates analysis (CVA) (a) of 10 cranial variables in five groups of the Rhinolophus hildebrandtii complex defined by molecular analysis; and PCA (b) of five cranial variables for sample in (a) with type series of hildebrandtii (‘‘H¹’’) and eloquens (‘‘E¹’’) added. Open circles = Clade 1a (= cohenae sp. nov.); closed circles = Clade 1b (= mabuensis sp. nov.); shaded circles = Clade 1d (= smithersi sp. nov.; Pafuri); asterisk enclosed in circle = Clade 1e (= smithersi sp. nov.; Zimbabwe); open squares = Clade 2 (mossambicus sp. nov.; Mozambique); shaded squares = Clade 2 (mossambicus sp. nov.; Lutopeı Zimbabwe); open diamonds = R. eloquens type series (Clade 3); crosses in circles = R. hildebrandtii type and co-type (Clade 1c). doi:10.1371/journal.pone.0041744.g005" figureDoi="http://doi.org/10.5281/zenodo.4244570" httpUri="https://zenodo.org/record/4244570/files/figure.png" pageId="16" pageNumber="17">Fig. 5b</figureCitation>
). The anterior upper premolar, when present is moderate-sized but situated external to the toothrow (with the canine and P
<quantity box="[1064,1100,1873,1900]" metricMagnitude="-1" metricUnit="m" metricValue="1.016" pageId="16" pageNumber="17" unit="in" value="4.0">
<superScript attach="left" box="[1064,1073,1873,1887]" fontSize="6" pageId="16" pageNumber="17">4</superScript>
in
</quantity>
contact) [
<bibRefCitation author="Csorba G &amp; Ujhelyip P &amp; Thomas N" box="[1198,1227,1878,1900]" journalOrPublisher="Shropshire: Alana books" pageId="16" pageNumber="17" refId="ref21181" refString="12. Csorba G, Ujhelyip P, Thomas N (2003) Horseshoe Bats of the World (Chiroptera: Rhinolophidae). Shropshire: Alana books." title="Horseshoe Bats of the World (Chiroptera: Rhinolophidae)" type="book" year="2003">12</bibRefCitation>
]. In this respect, they are distinguished from
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[1045,1106,1906,1927]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="cohenae">
<emphasis box="[1045,1106,1906,1927]" italics="true" pageId="16" pageNumber="17">cohenae</emphasis>
</taxonomicName>
which have relatively conspicuous anterior premolars within the toothrow and
<taxonomicName authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[1276,1347,1934,1955]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="smithersi">
<emphasis box="[1276,1347,1934,1955]" italics="true" pageId="16" pageNumber="17">smithersi</emphasis>
</taxonomicName>
in which the anterior premolar is small to minute and external to the toothrow. Genotypes of
<taxonomicName authorityName="Petersi" authorityYear="1878" box="[292,423,192,214]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[292,423,192,214]" italics="true" pageId="18" pageNumber="19">R. hildebrandtii</emphasis>
</taxonomicName>
are members of Clade 1c (
<figureCitation box="[694,750,192,214]" captionStart="Figure 2" captionStartId="2.[155,220,1881,1900]" captionTargetBox="[160,1475,167,1843]" captionTargetId="figure-0@2.[155,1480,162,1848]" captionTargetPageId="2" captionText="Figure 2. Map of southern, central and eastern Africa indicating localities of individuals of R. hildebrandtii species-complex included in this study. Grey-shaded area represents elevations in excess of 600 m a.s.l. Closed squares indicate museum specimens from which craniometric data were obtained. Open symbols indicate specimens genotyped in this study. The distribution of the three major clades is based on cytochrome b (see Figure 2): open circles = Clade 1; open squares = Clade 2; open diamonds = Clade 3. Closed squares enclosed in open symbols indicate localities where both molecular and morphological data were available for selected specimens. Numbers refer to respective localities listed in Table S1. ‘‘T’’ indicates the type localities of R. eloquens in Uganda and R. hildebrandtii in Kenyaı respectively. doi:10.1371/journal.pone.0041744.g002" figureDoi="http://doi.org/10.5281/zenodo.4244564" httpUri="https://zenodo.org/record/4244564/files/figure.png" pageId="18" pageNumber="19">Fig. 2</figureCitation>
).
</paragraph>
</subSubSection>
<caption box="[153,1540,487,508]" pageId="17" pageNumber="18">
<paragraph box="[153,1540,487,508]" pageId="17" pageNumber="18">
<emphasis bold="true" box="[153,246,487,508]" pageId="17" pageNumber="18">Table 5.</emphasis>
Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the
<taxonomicName authorityName="Petersi" authorityYear="1878" box="[1188,1439,487,508]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[1188,1439,487,508]" italics="true" pageId="17" pageNumber="18">Rhinolophus hildebrandtii</emphasis>
</taxonomicName>
complex.
</paragraph>
</caption>
<paragraph pageId="17" pageNumber="18">
<table box="[153,1929,572,1445]" gridcols="13" gridrows="13" pageId="17" pageNumber="18">
<tr box="[153,1929,572,656]" gridrow="0" pageId="17" pageNumber="18">
<th box="[153,204,572,656]" gridcol="0" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" box="[153,204,639,656]" pageId="17" pageNumber="18">Clade</emphasis>
</th>
<th box="[260,383,572,656]" gridcol="1" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" pageId="17" pageNumber="18">Species name (&amp; holotype number)</emphasis>
</th>
<th box="[403,555,572,656]" gridcol="2" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" box="[403,511,639,656]" pageId="17" pageNumber="18">Distribution</emphasis>
</th>
<th box="[581,652,572,656]" gridcol="3" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" pageId="17" pageNumber="18">Freq. (kHz)</emphasis>
</th>
<th box="[670,777,572,656]" gridcol="4" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" box="[670,712,639,656]" pageId="17" pageNumber="18">NLW</emphasis>
</th>
<th box="[795,869,572,656]" gridcol="5" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" box="[795,816,639,656]" pageId="17" pageNumber="18">FL</emphasis>
</th>
<th box="[884,984,572,656]" gridcol="6" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" box="[884,919,639,656]" pageId="17" pageNumber="18">GSL</emphasis>
</th>
<th box="[1009,1102,572,656]" gridcol="7" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" box="[1009,1044,639,656]" pageId="17" pageNumber="18">CCL</emphasis>
</th>
<th box="[1133,1313,572,656]" gridcol="8" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" pageId="17" pageNumber="18">Relative height of rostral chamber and sagittal crest in lateral view (Fig. 6)</emphasis>
</th>
<th box="[1347,1475,572,656]" gridcol="9" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" pageId="17" pageNumber="18">Depth of cranial profile overall (Fig. 6)</emphasis>
</th>
<th box="[1490,1612,572,656]" gridcol="10" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" pageId="17" pageNumber="18">Frontal depression in dorsal view (Fig. 6)</emphasis>
</th>
<th box="[1668,1745,572,656]" gridcol="11" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" box="[1668,1745,617,656]" pageId="17" pageNumber="18">Baculum (Fig. 8)</emphasis>
</th>
<th box="[1775,1929,572,656]" gridcol="12" gridrow="0" pageId="17" pageNumber="18">
<emphasis bold="true" pageId="17" pageNumber="18">Relative size and position of anterior upper premolar</emphasis>
</th>
</tr>
<tr box="[153,1929,679,741]" gridrow="1" pageId="17" pageNumber="18">
<th box="[153,204,679,741]" gridcol="0" gridrow="1" pageId="17" pageNumber="18">1a</th>
<td box="[260,383,679,741]" gridcol="1" gridrow="1" pageId="17" pageNumber="18">
<taxonomicName authority="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill, 2012" authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[260,324,679,696]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="cohenae" status="sp. nov">
<emphasis box="[260,324,679,696]" italics="true" pageId="17" pageNumber="18">cohenae</emphasis>
</taxonomicName>
<taxonomicNameLabel pageId="17" pageNumber="18" rank="species">sp. nov</taxonomicNameLabel>
</td>
<td box="[403,555,679,741]" gridcol="2" gridrow="1" pageId="17" pageNumber="18">Mpumalanga Provinceı South Africa</td>
<td box="[581,652,679,741]" gridcol="3" gridrow="1" pageId="17" pageNumber="18">33</td>
<td box="[670,777,679,741]" gridcol="4" gridrow="1" pageId="17" pageNumber="18">15.3; 13.5–16.3 (7)</td>
<td box="[795,984,679,741]" colspan="2" colspanRight="1" gridcol="5" gridrow="1" pageId="17" pageNumber="18">66.4; 29.8; 66–68 (10) 29–30 (10)</td>
<td box="[1009,1102,679,741]" gridcol="7" gridrow="1" pageId="17" pageNumber="18">25.3; 24–26 (10)</td>
<td box="[1133,1313,679,741]" gridcol="8" gridrow="1" pageId="17" pageNumber="18">Rostral chamber. = sagittal crest</td>
<td box="[1347,1475,679,741]" gridcol="9" gridrow="1" pageId="17" pageNumber="18">Flatter</td>
<td box="[1490,1612,679,741]" gridcol="10" gridrow="1" pageId="17" pageNumber="18">Conspicuous&amp; deep</td>
<td box="[1668,1745,679,741]" gridcol="11" gridrow="1" pageId="17" pageNumber="18">Type 1</td>
<td box="[1775,1929,679,741]" gridcol="12" gridrow="1" pageId="17" pageNumber="18">Prominentı in toothrow (or absent)</td>
</tr>
<tr box="[153,1929,758,820]" gridrow="2" pageId="17" pageNumber="18" rowspan-0="1">
<td box="[260,383,758,820]" gridcol="1" gridrow="2" pageId="17" pageNumber="18">(DM 8626)</td>
<td box="[403,555,758,820]" gridcol="2" gridrow="2" pageId="17" pageNumber="18">Mountainland Nature Reserveı Barberton</td>
<td box="[581,652,758,820]" gridcol="3" gridrow="2" pageId="17" pageNumber="18">33</td>
<td box="[670,777,758,820]" gridcol="4" gridrow="2" pageId="17" pageNumber="18">15.5</td>
<td box="[795,869,758,820]" gridcol="5" gridrow="2" pageId="17" pageNumber="18">65.9</td>
<td box="[884,984,758,820]" gridcol="6" gridrow="2" pageId="17" pageNumber="18">29.9</td>
<td box="[1009,1102,758,820]" gridcol="7" gridrow="2" pageId="17" pageNumber="18">25.6</td>
<td box="[1133,1313,758,820]" gridcol="8" gridrow="2" pageId="17" pageNumber="18">As above</td>
<td box="[1347,1475,758,820]" gridcol="9" gridrow="2" pageId="17" pageNumber="18">As above</td>
<td box="[1490,1612,758,820]" gridcol="10" gridrow="2" pageId="17" pageNumber="18">As above</td>
<td box="[1668,1745,758,820]" gridcol="11" gridrow="2" pageId="17" pageNumber="18">Type 1</td>
<td box="[1775,1929,758,820]" gridcol="12" gridrow="2" pageId="17" pageNumber="18">Prominent in toothrow</td>
</tr>
<tr box="[153,1929,836,876]" gridrow="3" pageId="17" pageNumber="18">
<th box="[153,204,836,876]" gridcol="0" gridrow="3" pageId="17" pageNumber="18">1b</th>
<td box="[260,383,836,876]" gridcol="1" gridrow="3" pageId="17" pageNumber="18">
<taxonomicName authority="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill, 2012" authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[260,341,836,853]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="mabuensis" status="sp. nov.">
<emphasis box="[260,341,836,853]" italics="true" pageId="17" pageNumber="18">mabuensis</emphasis>
</taxonomicName>
<taxonomicNameLabel pageId="17" pageNumber="18" rank="species">sp. nov.</taxonomicNameLabel>
</td>
<td box="[403,555,836,876]" gridcol="2" gridrow="3" pageId="17" pageNumber="18">Mts Mabu &amp; Inagoı N Mozambique</td>
<td box="[581,652,836,876]" gridcol="3" gridrow="3" pageId="17" pageNumber="18">38</td>
<td box="[670,777,836,876]" gridcol="4" gridrow="3" pageId="17" pageNumber="18">15.0; 15.0–15.1(2)</td>
<td box="[795,869,836,876]" gridcol="5" gridrow="3" pageId="17" pageNumber="18">67.5; 66–69 (2)</td>
<td box="[884,984,836,876]" gridcol="6" gridrow="3" pageId="17" pageNumber="18">29.7–30.3 (2)</td>
<td box="[1009,1102,836,876]" gridcol="7" gridrow="3" pageId="17" pageNumber="18">25.6; 25–26 (2)</td>
<td box="[1133,1313,836,876]" gridcol="8" gridrow="3" pageId="17" pageNumber="18">Rostral chamber. = sagittal crest</td>
<td box="[1347,1475,836,876]" gridcol="9" gridrow="3" pageId="17" pageNumber="18">Flatter</td>
<td box="[1490,1612,836,876]" gridcol="10" gridrow="3" pageId="17" pageNumber="18">Conspicuous &amp; deep</td>
<td box="[1668,1745,836,876]" gridcol="11" gridrow="3" pageId="17" pageNumber="18">Type 2</td>
<td box="[1775,1929,836,876]" gridcol="12" gridrow="3" pageId="17" pageNumber="18">Prominent - smallı internal-external</td>
</tr>
<tr box="[153,1929,892,931]" gridrow="4" pageId="17" pageNumber="18" rowspan-0="1">
<td box="[260,383,892,931]" gridcol="1" gridrow="4" pageId="17" pageNumber="18">(DM 10842)</td>
<td box="[403,555,892,931]" gridcol="2" gridrow="4" pageId="17" pageNumber="18">Mt Mabu</td>
<td box="[581,652,892,931]" gridcol="3" gridrow="4" pageId="17" pageNumber="18">
<emphasis box="[581,599,892,909]" italics="true" pageId="17" pageNumber="18">ca</emphasis>
38*
</td>
<td box="[670,777,892,931]" gridcol="4" gridrow="4" pageId="17" pageNumber="18">15.0</td>
<td box="[795,869,892,931]" gridcol="5" gridrow="4" pageId="17" pageNumber="18">66.1</td>
<td box="[884,984,892,931]" gridcol="6" gridrow="4" pageId="17" pageNumber="18">29.7</td>
<td box="[1009,1102,892,931]" gridcol="7" gridrow="4" pageId="17" pageNumber="18">25.2</td>
<td box="[1133,1313,892,931]" gridcol="8" gridrow="4" pageId="17" pageNumber="18">As above</td>
<td box="[1347,1475,892,931]" gridcol="9" gridrow="4" pageId="17" pageNumber="18">As above</td>
<td box="[1490,1612,892,931]" gridcol="10" gridrow="4" pageId="17" pageNumber="18">As above</td>
<td box="[1668,1745,892,931]" gridcol="11" gridrow="4" pageId="17" pageNumber="18">Type 2</td>
<td box="[1775,1929,892,931]" gridcol="12" gridrow="4" pageId="17" pageNumber="18">Prominentı partially in toothrow</td>
</tr>
<tr box="[153,1929,947,987]" gridrow="5" pageId="17" pageNumber="18">
<th box="[153,204,947,987]" gridcol="0" gridrow="5" pageId="17" pageNumber="18">1c</th>
<td box="[260,383,947,987]" gridcol="1" gridrow="5" pageId="17" pageNumber="18">
<taxonomicName authorityName="Petersi" authorityYear="1878" box="[260,356,947,964]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[260,356,947,964]" italics="true" pageId="17" pageNumber="18">hildebrandtii</emphasis>
</taxonomicName>
</td>
<td box="[403,555,947,987]" gridcol="2" gridrow="5" pageId="17" pageNumber="18">East Africa (Kenyaı Tanzania)</td>
<td box="[581,652,947,987]" gridcol="3" gridrow="5" pageId="17" pageNumber="18">42</td>
<td box="[670,777,947,987]" gridcol="4" gridrow="5" pageId="17" pageNumber="18">11.0; 9.8–13.1 (9)</td>
<td box="[795,869,947,987]" gridcol="5" gridrow="5" pageId="17" pageNumber="18">62.5 60–68 (8)</td>
<td box="[884,984,947,987]" gridcol="6" gridrow="5" pageId="17" pageNumber="18">28.0 27–29 (4)</td>
<td box="[1009,1102,947,987]" gridcol="7" gridrow="5" pageId="17" pageNumber="18">24.0 23–25 (6)</td>
<td box="[1133,1313,947,987]" gridcol="8" gridrow="5" pageId="17" pageNumber="18">-</td>
<td box="[1347,1475,947,987]" gridcol="9" gridrow="5" pageId="17" pageNumber="18">-</td>
<td box="[1490,1612,947,987]" gridcol="10" gridrow="5" pageId="17" pageNumber="18">
Shallow to moderate [
<bibRefCitation author="Csorba G &amp; Ujhelyip P &amp; Thomas N" box="[1576,1597,970,987]" journalOrPublisher="Shropshire: Alana books" pageId="17" pageNumber="18" refId="ref21181" refString="12. Csorba G, Ujhelyip P, Thomas N (2003) Horseshoe Bats of the World (Chiroptera: Rhinolophidae). Shropshire: Alana books." title="Horseshoe Bats of the World (Chiroptera: Rhinolophidae)" type="book" year="2003">12</bibRefCitation>
]
</td>
<td box="[1668,1745,947,987]" gridcol="11" gridrow="5" pageId="17" pageNumber="18">-</td>
<td box="[1775,1929,947,987]" gridcol="12" gridrow="5" pageId="17" pageNumber="18">Moderate - smallı external</td>
</tr>
<tr box="[153,1929,1003,1020]" gridrow="6" pageId="17" pageNumber="18" rowspan-0="1">
<td box="[260,383,1003,1020]" gridcol="1" gridrow="6" pageId="17" pageNumber="18">(ZMB 5378)</td>
<td box="[403,555,1003,1020]" gridcol="2" gridrow="6" pageId="17" pageNumber="18">Ndiı Taitaı Kenya</td>
<td box="[581,652,1003,1020]" gridcol="3" gridrow="6" pageId="17" pageNumber="18">Ca 42 **</td>
<td box="[670,777,1003,1020]" gridcol="4" gridrow="6" pageId="17" pageNumber="18">9.8</td>
<td box="[795,869,1003,1020]" gridcol="5" gridrow="6" pageId="17" pageNumber="18">63.5</td>
<td box="[884,984,1003,1020]" gridcol="6" gridrow="6" pageId="17" pageNumber="18">27.6</td>
<td box="[1009,1102,1003,1020]" gridcol="7" gridrow="6" pageId="17" pageNumber="18">-</td>
<td box="[1133,1313,1003,1020]" gridcol="8" gridrow="6" pageId="17" pageNumber="18">-</td>
<td box="[1347,1475,1003,1020]" gridcol="9" gridrow="6" pageId="17" pageNumber="18">-</td>
<td box="[1490,1612,1003,1020]" gridcol="10" gridrow="6" pageId="17" pageNumber="18">-</td>
<td box="[1668,1745,1003,1020]" gridcol="11" gridrow="6" pageId="17" pageNumber="18">-</td>
<td box="[1775,1929,1003,1020]" gridcol="12" gridrow="6" pageId="17" pageNumber="18">-</td>
</tr>
<tr box="[153,1929,1036,1099]" gridrow="7" pageId="17" pageNumber="18">
<th box="[153,204,1036,1099]" gridcol="0" gridrow="7" pageId="17" pageNumber="18">1d/e</th>
<td box="[260,383,1036,1099]" gridcol="1" gridrow="7" pageId="17" pageNumber="18">
<taxonomicName authority="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill, 2012" authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[260,328,1036,1053]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="smithersi" status="sp. nov.">
<emphasis box="[260,328,1036,1053]" italics="true" pageId="17" pageNumber="18">smithersi</emphasis>
</taxonomicName>
<taxonomicNameLabel pageId="17" pageNumber="18" rank="species">sp. nov.</taxonomicNameLabel>
</td>
<td box="[403,555,1036,1099]" gridcol="2" gridrow="7" pageId="17" pageNumber="18">Pafuriı South Africa &amp; Lutope- Ngolangola Gorge</td>
<td box="[581,652,1036,1099]" gridcol="3" gridrow="7" pageId="17" pageNumber="18">44–46</td>
<td box="[670,777,1036,1099]" gridcol="4" gridrow="7" pageId="17" pageNumber="18">11.3; 10.1–14.4 (6)</td>
<td box="[795,869,1036,1099]" gridcol="5" gridrow="7" pageId="17" pageNumber="18">62.6; 61–64 (5)</td>
<td box="[884,984,1036,1099]" gridcol="6" gridrow="7" pageId="17" pageNumber="18">27.3; 27–28 (4)</td>
<td box="[1009,1102,1036,1099]" gridcol="7" gridrow="7" pageId="17" pageNumber="18">23.5 23–24 (6)</td>
<td box="[1133,1313,1036,1099]" gridcol="8" gridrow="7" pageId="17" pageNumber="18">Sagittal crest. rostral chamber</td>
<td box="[1347,1475,1036,1099]" gridcol="9" gridrow="7" pageId="17" pageNumber="18">Intermediate</td>
<td box="[1490,1612,1036,1099]" gridcol="10" gridrow="7" pageId="17" pageNumber="18">Weakly developed</td>
<td box="[1668,1745,1036,1099]" gridcol="11" gridrow="7" pageId="17" pageNumber="18">-</td>
<td box="[1775,1929,1036,1099]" gridcol="12" gridrow="7" pageId="17" pageNumber="18">Smallı external</td>
</tr>
<tr box="[153,1929,1115,1154]" gridrow="8" pageId="17" pageNumber="18" rowspan-0="1">
<td box="[260,383,1115,1154]" gridcol="1" gridrow="8" pageId="17" pageNumber="18">(NMBZ 33647)</td>
<td box="[403,555,1115,1154]" gridcol="2" gridrow="8" pageId="17" pageNumber="18">Lutope-Ngolangola Gorgeı Zimbabwe</td>
<td box="[581,652,1115,1154]" gridcol="3" gridrow="8" pageId="17" pageNumber="18">46</td>
<td box="[670,777,1115,1154]" gridcol="4" gridrow="8" pageId="17" pageNumber="18">14.4</td>
<td box="[795,869,1115,1154]" gridcol="5" gridrow="8" pageId="17" pageNumber="18">60.7</td>
<td box="[884,984,1115,1154]" gridcol="6" gridrow="8" pageId="17" pageNumber="18">26.5</td>
<td box="[1009,1102,1115,1154]" gridcol="7" gridrow="8" pageId="17" pageNumber="18">23.1</td>
<td box="[1133,1313,1115,1154]" gridcol="8" gridrow="8" pageId="17" pageNumber="18">-</td>
<td box="[1347,1475,1115,1154]" gridcol="9" gridrow="8" pageId="17" pageNumber="18">-</td>
<td box="[1490,1612,1115,1154]" gridcol="10" gridrow="8" pageId="17" pageNumber="18">-</td>
<td box="[1668,1745,1115,1154]" gridcol="11" gridrow="8" pageId="17" pageNumber="18">-</td>
<td box="[1775,1929,1115,1154]" gridcol="12" gridrow="8" pageId="17" pageNumber="18">-</td>
</tr>
<tr box="[153,1929,1170,1233]" gridrow="9" pageId="17" pageNumber="18">
<th box="[153,204,1170,1233]" gridcol="0" gridrow="9" pageId="17" pageNumber="18">2</th>
<td box="[260,383,1170,1233]" gridcol="1" gridrow="9" pageId="17" pageNumber="18">
<taxonomicName authority="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill, 2012" authorityName="Peter J. Taylor &amp; Samantha Stoffberg &amp; Ara Monadjem &amp; Martinus Corrie Schoeman &amp; Julian Baylis &amp; Fenton P. D. Cotterill" authorityYear="2012" box="[260,362,1171,1188]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="mossambicus" status="sp. nov.">
<emphasis box="[260,362,1171,1188]" italics="true" pageId="17" pageNumber="18">mossambicus</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[260,323,1193,1210]" pageId="17" pageNumber="18" rank="species">sp. nov.</taxonomicNameLabel>
</td>
<td box="[403,555,1170,1233]" gridcol="2" gridrow="9" pageId="17" pageNumber="18">Mozambiqueı extending to NW Zimbabwe</td>
<td box="[581,652,1170,1233]" gridcol="3" gridrow="9" pageId="17" pageNumber="18">35–38</td>
<td box="[670,777,1170,1233]" gridcol="4" gridrow="9" pageId="17" pageNumber="18">13.5; 12.9–14.2(3)</td>
<td box="[795,869,1170,1233]" gridcol="5" gridrow="9" pageId="17" pageNumber="18">62.8; 60–65 (5)</td>
<td box="[884,984,1170,1233]" gridcol="6" gridrow="9" pageId="17" pageNumber="18">28.1; 27–29 (5)</td>
<td box="[1009,1102,1170,1233]" gridcol="7" gridrow="9" pageId="17" pageNumber="18">24.4; 24–25 (5)</td>
<td box="[1133,1313,1170,1233]" gridcol="8" gridrow="9" pageId="17" pageNumber="18">Sagittal crest» rostral chamber</td>
<td box="[1347,1475,1170,1233]" gridcol="9" gridrow="9" pageId="17" pageNumber="18">Higher &amp; domed</td>
<td box="[1490,1612,1170,1233]" gridcol="10" gridrow="9" pageId="17" pageNumber="18">Reduced</td>
<td box="[1668,1745,1170,1233]" gridcol="11" gridrow="9" pageId="17" pageNumber="18">Type 2</td>
<td box="[1775,1929,1170,1233]" gridcol="12" gridrow="9" pageId="17" pageNumber="18">Minuteı external</td>
</tr>
<tr box="[153,1929,1249,1311]" gridrow="10" pageId="17" pageNumber="18" rowspan-0="1">
<td box="[260,383,1249,1311]" gridcol="1" gridrow="10" pageId="17" pageNumber="18">(DM 8578)</td>
<td box="[403,555,1249,1311]" gridcol="2" gridrow="10" pageId="17" pageNumber="18">Niassa Game Reserveı N Mozambique</td>
<td box="[581,652,1249,1311]" gridcol="3" gridrow="10" pageId="17" pageNumber="18">
<emphasis box="[581,599,1249,1266]" italics="true" pageId="17" pageNumber="18">ca</emphasis>
35–38 ***
</td>
<td box="[670,777,1249,1311]" gridcol="4" gridrow="10" pageId="17" pageNumber="18">14.2</td>
<td box="[795,869,1249,1311]" gridcol="5" gridrow="10" pageId="17" pageNumber="18">63.9</td>
<td box="[884,984,1249,1311]" gridcol="6" gridrow="10" pageId="17" pageNumber="18">28.5</td>
<td box="[1009,1102,1249,1311]" gridcol="7" gridrow="10" pageId="17" pageNumber="18">25.0</td>
<td box="[1133,1313,1249,1311]" gridcol="8" gridrow="10" pageId="17" pageNumber="18">As above</td>
<td box="[1347,1475,1249,1311]" gridcol="9" gridrow="10" pageId="17" pageNumber="18">As above</td>
<td box="[1490,1612,1249,1311]" gridcol="10" gridrow="10" pageId="17" pageNumber="18">As above</td>
<td box="[1668,1745,1249,1311]" gridcol="11" gridrow="10" pageId="17" pageNumber="18">As above</td>
<td box="[1775,1929,1249,1311]" gridcol="12" gridrow="10" pageId="17" pageNumber="18">As above</td>
</tr>
<tr box="[153,1929,1327,1367]" gridrow="11" pageId="17" pageNumber="18">
<th box="[153,204,1327,1367]" gridcol="0" gridrow="11" pageId="17" pageNumber="18">3</th>
<td box="[260,383,1327,1367]" gridcol="1" gridrow="11" pageId="17" pageNumber="18">
<taxonomicName authorityName="Anderson" authorityYear="1905" box="[260,327,1327,1344]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="eloquens">
<emphasis box="[260,327,1327,1344]" italics="true" pageId="17" pageNumber="18">eloquens</emphasis>
</taxonomicName>
</td>
<td box="[403,555,1327,1367]" gridcol="2" gridrow="11" pageId="17" pageNumber="18">East Africa</td>
<td box="[581,652,1327,1367]" gridcol="3" gridrow="11" pageId="17" pageNumber="18">-</td>
<td box="[670,777,1327,1367]" gridcol="4" gridrow="11" pageId="17" pageNumber="18">9.7; 9.4–10.6(4)</td>
<td box="[795,869,1327,1367]" gridcol="5" gridrow="11" pageId="17" pageNumber="18">55.5; (54–57(4)</td>
<td box="[884,984,1327,1367]" gridcol="6" gridrow="11" pageId="17" pageNumber="18">25.2; 25.1–25.3(3)</td>
<td box="[1009,1102,1327,1367]" gridcol="7" gridrow="11" pageId="17" pageNumber="18">22.0; 21.7–22.2(5)</td>
<td box="[1133,1313,1327,1367]" gridcol="8" gridrow="11" pageId="17" pageNumber="18">-</td>
<td box="[1347,1475,1327,1367]" gridcol="9" gridrow="11" pageId="17" pageNumber="18">-</td>
<td box="[1490,1612,1327,1367]" gridcol="10" gridrow="11" pageId="17" pageNumber="18">
Shallow[
<bibRefCitation author="Csorba G &amp; Ujhelyip P &amp; Thomas N" box="[1558,1576,1327,1344]" journalOrPublisher="Shropshire: Alana books" pageId="17" pageNumber="18" refId="ref21181" refString="12. Csorba G, Ujhelyip P, Thomas N (2003) Horseshoe Bats of the World (Chiroptera: Rhinolophidae). Shropshire: Alana books." title="Horseshoe Bats of the World (Chiroptera: Rhinolophidae)" type="book" year="2003">12</bibRefCitation>
]
</td>
<td box="[1668,1745,1327,1367]" gridcol="11" gridrow="11" pageId="17" pageNumber="18">-</td>
<td box="[1775,1929,1327,1367]" gridcol="12" gridrow="11" pageId="17" pageNumber="18">Minuteı external</td>
</tr>
<tr box="[153,1929,1383,1445]" gridrow="12" pageId="17" pageNumber="18" rowspan-0="1">
<td box="[260,383,1383,1445]" gridcol="1" gridrow="12" pageId="17" pageNumber="18">(BM 1899.8.4.4)</td>
<td box="[403,555,1383,1445]" gridcol="2" gridrow="12" pageId="17" pageNumber="18">Entebbeı Uganda</td>
<td box="[581,652,1383,1445]" gridcol="3" gridrow="12" pageId="17" pageNumber="18">-</td>
<td box="[670,777,1383,1445]" gridcol="4" gridrow="12" pageId="17" pageNumber="18">10.6</td>
<td box="[795,869,1383,1445]" gridcol="5" gridrow="12" pageId="17" pageNumber="18">56.1</td>
<td box="[884,984,1383,1445]" gridcol="6" gridrow="12" pageId="17" pageNumber="18">25.1</td>
<td box="[1009,1102,1383,1445]" gridcol="7" gridrow="12" pageId="17" pageNumber="18">21.7</td>
<td box="[1133,1313,1383,1445]" gridcol="8" gridrow="12" pageId="17" pageNumber="18">Sagittal crest» rostral chamber</td>
<td box="[1347,1475,1383,1445]" gridcol="9" gridrow="12" pageId="17" pageNumber="18">High and domed</td>
<td box="[1490,1612,1383,1445]" gridcol="10" gridrow="12" pageId="17" pageNumber="18">Moderately developed and deep</td>
<td box="[1668,1745,1383,1445]" gridcol="11" gridrow="12" pageId="17" pageNumber="18">-</td>
<td box="[1775,1929,1383,1445]" gridcol="12" gridrow="12" pageId="17" pageNumber="18">Minuteı external</td>
</tr>
</table>
</paragraph>
<paragraph pageId="17" pageNumber="18">
Freq.= peak (time expansion or full-spectrum detector) or maximum (ANABAT detector) frequency; NLW = nasal width; FL = forearm length; GSL = greatest skull length; CCL = condylo-canine skull length. *Echolocation frequency of holotype not measured but assumed on basis of two released individuals of the same species at Mt Mabu recorded during the same month (October 2008). **We assume the holotype and paratype specimens from the Taita District may have had a CF frequency of
<emphasis box="[1020,1038,1520,1537]" italics="true" pageId="17" pageNumber="18">ca</emphasis>
42 kHz based on recent recordings from the type locality. ***The holotype was not recorded but we assume a CF frequency of 35–38 kHz based on recordings of four bats taken from nearby localities in northern Mozambique. doi:10.1371/journal.pone.0041744.t005
</paragraph>
<subSubSection pageId="18" pageNumber="19" type="materials_examined">
<paragraph blockId="18.[155,794,164,1208]" pageId="18" pageNumber="19">
<materialsCitation ID-GBIF-Occurrence="2908237303" collectingDate="1877" collectionCode="BM" collectorName="S. Hildebrandt" country="Kenya" location="Taita" pageId="18" pageNumber="19" specimenCode="BM 79.1.21.1" specimenCount="1" stateProvince="Taita" typeStatus="paratype">
<emphasis bold="true" box="[177,281,222,241]" pageId="18" pageNumber="19">
<typeStatus box="[177,276,222,241]" pageId="18" pageNumber="19">Paratype</typeStatus>
.
</emphasis>
<specimenCode box="[305,440,221,243]" pageId="18" pageNumber="19">BM 79.1.21.1</specimenCode>
,
<collectingRegion box="[450,502,221,243]" country="Kenya" name="Taita/Taveta" pageId="18" pageNumber="19">Taita</collectingRegion>
(
<specimenCount box="[515,600,221,243]" pageId="18" pageNumber="19">un-sexed</specimenCount>
individual collected in
<collectingDate box="[182,230,249,271]" pageId="18" pageNumber="19">1877</collectingDate>
by
<collectorName box="[269,417,249,271]" pageId="18" pageNumber="19">S. Hildebrandt</collectorName>
)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="description">
<paragraph blockId="18.[155,794,164,1208]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[177,311,278,297]" pageId="18" pageNumber="19">Description.</emphasis>
Follows the general description for the species [
<bibRefCitation author="Csorba G &amp; Ujhelyip P &amp; Thomas N" box="[162,190,306,328]" journalOrPublisher="Shropshire: Alana books" pageId="18" pageNumber="19" refId="ref21181" refString="12. Csorba G, Ujhelyip P, Thomas N (2003) Horseshoe Bats of the World (Chiroptera: Rhinolophidae). Shropshire: Alana books." title="Horseshoe Bats of the World (Chiroptera: Rhinolophidae)" type="book" year="2003">12</bibRefCitation>
]; however,
<taxonomicName authorityName="Petersi" authorityYear="1878" box="[308,411,305,327]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" sensu="stricto" species="hildebrandtii">
<emphasis box="[308,411,305,327]" italics="true" pageId="18" pageNumber="19">hildebrandtii</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[420,448,306,328]" pageId="18" pageNumber="19" sensu="stricto">s.s.</taxonomicNameLabel>
as defined here (restricted to East Africa) is intermediate-sized in most external and cranial dimensions and varies from the range of values reported previously for
<taxonomicName authorityName="Petersi" authorityYear="1878" box="[189,319,390,412]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" sensu="lato" species="hildebrandtii">
<emphasis box="[189,319,390,412]" italics="true" pageId="18" pageNumber="19">R. hildebrandtii</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[326,352,391,413]" pageId="18" pageNumber="19" sensu="lato">s.l.</taxonomicNameLabel>
[
<bibRefCitation author="Csorba G &amp; Ujhelyip P &amp; Thomas N" box="[365,393,391,413]" journalOrPublisher="Shropshire: Alana books" pageId="18" pageNumber="19" refId="ref21181" refString="12. Csorba G, Ujhelyip P, Thomas N (2003) Horseshoe Bats of the World (Chiroptera: Rhinolophidae). Shropshire: Alana books." title="Horseshoe Bats of the World (Chiroptera: Rhinolophidae)" type="book" year="2003">12</bibRefCitation>
], for example noseleaf width in nine East African specimens examined varied from
<quantity box="[563,671,419,441]" metricMagnitude="-2" metricUnit="m" metricValue="1.2" metricValueMax="1.4" metricValueMin="1.0" pageId="18" pageNumber="19" unit="mm" value="12.0" valueMax="14.0" valueMin="10.0">10–14 mm</quantity>
, rather than
<quantity box="[155,261,448,470]" metricMagnitude="-2" metricUnit="m" metricValue="1.35" metricValueMax="1.5" metricValueMin="1.2" pageId="18" pageNumber="19" unit="mm" value="13.5" valueMax="15.0" valueMin="12.0">12–15 mm</quantity>
given by previous description [
<bibRefCitation author="Csorba G &amp; Ujhelyip P &amp; Thomas N" box="[569,598,448,470]" journalOrPublisher="Shropshire: Alana books" pageId="18" pageNumber="19" refId="ref21181" refString="12. Csorba G, Ujhelyip P, Thomas N (2003) Horseshoe Bats of the World (Chiroptera: Rhinolophidae). Shropshire: Alana books." title="Horseshoe Bats of the World (Chiroptera: Rhinolophidae)" type="book" year="2003">12</bibRefCitation>
].
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="distribution">
<paragraph blockId="18.[155,794,164,1208]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[177,317,477,496]" pageId="18" pageNumber="19">Distribution.</emphasis>
As here defined, and based only on limited material examined from Kenya and Tanzania in the Natural History Museum, London and Berlin Museum in this study (n = 11; mensural data reported in
<tableCitation box="[531,614,561,583]" captionStart="Table 5" captionStartId="17.[153,217,487,508]" captionText="Table 5. Matrix showing diagnostic traits and distribution of seven evolutionary lineages within the Rhinolophus hildebrandtii complex." pageId="18" pageNumber="19">Table 5</tableCitation>
), we restrict the distribution of
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<emphasis box="[295,398,589,611]" italics="true" pageId="18" pageNumber="19">hildebrandtii</emphasis>
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to East Africa, whilst recognising that northern populations in Democratic Republic of Congo, Ethiopia and southern Sudan (now South Sudan) are possibly referable to this species (as discussed by Koopman 1975, [
<bibRefCitation author="Monadjem A &amp; Taylor PJ &amp; Cotterill FPD &amp; Schoeman MC" box="[589,617,675,697]" editor="Wilson DE Reeder DM" journalOrPublisher="Baltimore: The John Hopkins University Press" pageId="18" pageNumber="19" pagination="313 - 529" refId="ref21181" refString="13. Monadjem A, Taylor PJ, Cotterill FPD, Schoeman MC (2010 a) Bats of Southern and Central Africa: A Biogeographic and Taxonomic Synthesis. Johannesburg: Wits University Press. 14. Simmons NB (2005) Order Chiroptera. In Wilson DE Reeder DM (eds.) Mammal Species of the World: A Taxonomic and Geographic Reference (3 rd edition). Baltimore: The John Hopkins University Press, 313 - 529." title="Bats of Southern and Central Africa: A Biogeographic and Taxonomic Synthesis. Johannesburg: Wits University Press. 14. Simmons NB (2005) Order Chiroptera" type="book chapter" volumeTitle="Mammal Species of the World: A Taxonomic and Geographic Reference (3 rd edition)" year="2010">13</bibRefCitation>
]. Moreover, given that two individuals with peak frequency of 37 kHz from Lutope-Ngolangola in NW Zimbabwe grouped genetically with individuals from coastal Mozambique assigned to Clade 2, it seems probable that
<taxonomicName authorityName="Petersi" authorityYear="1878" box="[304,407,788,810]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[304,407,788,810]" italics="true" pageId="18" pageNumber="19">hildebrandtii</emphasis>
</taxonomicName>
as here defined (coinciding with the divergent Lineage 1c) does not occur in southern Africa. Nevertheless, the taxonomic affinities of material from Zambia and Malawi remain unclear as craniometric analysis was unable to differentiate molecular clades and lineages at this scale (
<figureCitation box="[711,782,902,924]" captionStart="Figure 2" captionStartId="2.[155,220,1881,1900]" captionTargetBox="[160,1475,167,1843]" captionTargetId="figure-0@2.[155,1480,162,1848]" captionTargetPageId="2" captionText="Figure 2. Map of southern, central and eastern Africa indicating localities of individuals of R. hildebrandtii species-complex included in this study. Grey-shaded area represents elevations in excess of 600 m a.s.l. Closed squares indicate museum specimens from which craniometric data were obtained. Open symbols indicate specimens genotyped in this study. The distribution of the three major clades is based on cytochrome b (see Figure 2): open circles = Clade 1; open squares = Clade 2; open diamonds = Clade 3. Closed squares enclosed in open symbols indicate localities where both molecular and morphological data were available for selected specimens. Numbers refer to respective localities listed in Table S1. ‘‘T’’ indicates the type localities of R. eloquens in Uganda and R. hildebrandtii in Kenyaı respectively. doi:10.1371/journal.pone.0041744.g002" figureDoi="http://doi.org/10.5281/zenodo.4244564" httpUri="https://zenodo.org/record/4244564/files/figure.png" pageId="18" pageNumber="19">Fig. S2</figureCitation>
). Future studies need to incorporate molecular and bacular data from central, equatorial and northeastern Africa (particularly Zambia, Malawi, DRC and Ethiopia) to test the species limits of
<taxonomicName authorityName="Petersi" authorityYear="1878" box="[155,258,1015,1037]" class="Mammalia" family="Rhinolophidae" genus="Rhinolophus" kingdom="Animalia" order="Chiroptera" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="hildebrandtii">
<emphasis box="[155,258,1015,1037]" italics="true" pageId="18" pageNumber="19">hildebrandtii</emphasis>
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. It is possible that yet more cryptic species remain to be discovered.
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="19" type="biology_ecology">
<paragraph blockId="18.[155,794,164,1208]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[177,267,1074,1093]" pageId="18" pageNumber="19">Ecology.</emphasis>
This species was recorded from montane and submontane forests in the Usambara Mountains of the Eastern Arc Range in Tanzania; means and ranges given for external and cranial measurements for four individuals from this population conform very closely to our data [
<bibRefCitation author="Stanley WT &amp; Goodman SM" box="[488,530,1186,1208]" journalOrPublisher="Fieldiana Life and Earth Sciences" pageId="18" pageNumber="19" pagination="34 - 52" part="4" refId="ref25558" refString="119. Stanley WT, Goodman SM (2011) Small mammal inventories in the east and west Usambara and south Pare Mountains, Tanzania. 3. Chiroptera. Fieldiana Life and Earth Sciences 4: 34 - 52." title="Small mammal inventories in the east and west Usambara and south Pare Mountains, Tanzania. 3. Chiroptera" type="journal article" year="2011">119</bibRefCitation>
].
</paragraph>
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<bibRefCitation id="5430785C6E26CF9977B48ABACBA44245" author="Barreto-Reyes, P." journalOrPublisher="Anales de la Sociedad de Biologia de Bogota" pageId="0" pageNumber="4567" pagination="46 - 94" title="Lista de mosquitos de Colombia (Diptera: Culicidae)." volume="7" year="1955">Barreto-Reyes 1955</bibRefCitation>
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