treatments-xml/data/D9/AE/2F/D9AE2F5E87C0FB6DB7A89CDD5D6CD51A.xml

<document ID-DOI="http://dx.doi.org/10.3897/zookeys.759.24742" ID-GBIF-Dataset="bffe2edf-5eb7-430f-b1f0-c73c0d066d33" ID-PMC="PMC5974005" ID-Pensoft-Pub="1313-2970-759-99" ID-PubMed="29861648" ID-ZBK="F536AF25E7924C99B4D6FA4B76FDA45D" ModsDocAuthor="" ModsDocDate="2018" ModsDocID="1313-2970-759-99" ModsDocOrigin="ZooKeys 759" ModsDocTitle="Anomaloglossusmeansi sp. n., a new Pantepui species of the Anomaloglossusbeebei group (Anura, Aromobatidae)" checkinTime="1527037475429" checkinUser="pensoft" docAuthor="Kok, Philippe J. R., Nicolai, Michael P. J., Lathrop, Amy &amp; MacCulloch, Ross D." docDate="2018" docId="D9AE2F5E87C0FB6DB7A89CDD5D6CD51A" docLanguage="en" docName="ZooKeys 759: 99-116" docOrigin="ZooKeys 759" docSource="http://dx.doi.org/10.3897/zookeys.759.24742" docTitle="Anomaloglossus meansi Kok, Nicolai, Lathrop &amp; MacCulloch, 2018, sp. n." docType="treatment" docUuid="84C73332-67F9-4412-8140-CF70F1FB419C" docUuidSource="ZooBank" docVersion="5" lastPageNumber="99" masterDocId="9376EE3EFFD5600FFFB6EE00FF9F417E" masterDocTitle="Anomaloglossusmeansi sp. n., a new Pantepui species of the Anomaloglossusbeebei group (Anura, Aromobatidae)" masterLastPageNumber="116" masterPageNumber="99" pageNumber="99" updateTime="1668165812176" updateUser="ExternalLinkService">
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<mods:title>Anomaloglossusmeansi sp. n., a new Pantepui species of the Anomaloglossusbeebei group (Anura, Aromobatidae)</mods:title>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Kok, Philippe J. R.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Nicolai, Michael P. J.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Lathrop, Amy</mods:namePart>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>MacCulloch, Ross D.</mods:namePart>
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<mods:title>ZooKeys</mods:title>
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<mods:date>2018</mods:date>
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<mods:number>759</mods:number>
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<mods:url>http://dx.doi.org/10.3897/zookeys.759.24742</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.759.24742</mods:identifier>
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<treatment ID-GBIF-Taxon="144067148" LSID="urn:lsid:zoobank.org:act:84C73332-67F9-4412-8140-CF70F1FB419C" httpUri="http://treatment.plazi.org/id/D9AE2F5E87C0FB6DB7A89CDD5D6CD51A" lastPageNumber="99" pageId="0" pageNumber="99">
<subSubSection pageId="0" pageNumber="99" type="nomenclature">
<paragraph pageId="0" pageNumber="99">
<taxonomicName LSID="http://zoobank.org/84C73332-67F9-4412-8140-CF70F1FB419C" class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="99">sp. n.</taxonomicNameLabel>
Figures 3, 4; Table 1
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="reference_group">
<paragraph pageId="0" pageNumber="99">
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="genus">Anomaloglossus</taxonomicName>
sp. Ayanganna 
<bibRefCitation author="Grant, T" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" url="https://doi.org/10.1206/0003-0090(2006)299[1:PSODFA]2.0.CO;2" year="2006">Grant et al. 2006</bibRefCitation>
: 120-121, 
<bibRefCitation author="Grant, T" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" url="https://doi.org/10.2994/SAJH-D-17-00017.1" year="2017">2017</bibRefCitation>
: S66.
</paragraph>
<paragraph pageId="0" pageNumber="99">
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus praderioi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="praderioi">Anomaloglossus cf. praderioi</taxonomicName>
<bibRefCitation author="Kok, PJR" journalOrPublisher="Papeis Avulsos de Zoologia" pageId="0" pageNumber="99" pagination="51 - 68" title="A redescription of Anomaloglossuspraderioi (La Marca, 1998) (Anura: Aromobatidae: Anomaloglossinae), with description of its tadpole and call." volume="50" year="2010">Kok 2010</bibRefCitation>
: 66.
</paragraph>
<paragraph pageId="0" pageNumber="99">
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="genus">Anomaloglossus</taxonomicName>
sp. B 
<bibRefCitation author="Kok, PJR" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" url="https://doi.org/10.1016/j.cub.2012.06.034" year="2012">Kok et al. 2012</bibRefCitation>
: supplementary information.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="holotype">
<paragraph pageId="0" pageNumber="99">Holotype.</paragraph>
<paragraph pageId="0" pageNumber="99">
ROM 43896, adult male from the vicinity of Camp 2 on the Wokomung Massif, Potaro-Siparuni District, Guyana (
<geoCoordinate direction="north" orientation="latitude" precision="1" value="5.1097217">05°06.5833'N</geoCoordinate>
; 
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-59.821114">059°49.2667'W</geoCoordinate>
), 1234 m elevation, collected by A. Lathrop and R. James on 30 October 2004.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="paratypes">
<paragraph pageId="0" pageNumber="99">Paratypes</paragraph>
<paragraph pageId="0" pageNumber="99">
(n = 10). An adult male (ROM 39639) from the northeast plateau of Mount Ayanganna, Cuyuni-Mazaruni District, Guyana (
<geoCoordinate direction="north" orientation="latitude" precision="92" value="5.4016666">05°24.1'N</geoCoordinate>
; 
<geoCoordinate direction="west" orientation="longitude" precision="92" value="-59.956665">059°57.4'W</geoCoordinate>
), 1490 m elevation, collected by R. D. MacCulloch, A. Lathrop and C. Cox on 26 October 2000; four adult females (ROM 43320, ROM 43329, ROM 43331, ROM 43332) from the vicinity of Camp 2 on the Wokomung Massif, Potaro-Siparuni District, Guyana (
<geoCoordinate direction="north" orientation="latitude" precision="1" value="5.1097217">05°06.5833'N</geoCoordinate>
; 
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-59.821114">059°49.2667'W</geoCoordinate>
), 1234 m elevation, collected by A. Lathrop, R. D. MacCulloch and S. Khan between 26-31 October 2004; one adult female (ROM 43323) from the vicinity of Camp 3 on the Wokomung Massif, Potaro-Siparuni District, Guyana (
<geoCoordinate direction="north" orientation="latitude" precision="9" value="5.0941668">05°05.65'N</geoCoordinate>
; 
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-59.843056">059°50.5833'W</geoCoordinate>
), 1411 m elevation, collected by A. Lathrop, R. D. MacCulloch and S. Khan on 3 November 2004; one juvenile (ROM 43322) from the vicinity of Camp 2 on the Wokomung Massif, Potaro-Siparuni District, Guyana (
<geoCoordinate direction="north" orientation="latitude" precision="1" value="5.1097217">05°06.5833'N</geoCoordinate>
; 
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-59.821114">059°49.2667'W</geoCoordinate>
), 1234 m elevation, collected by C. Alban on 26 October 2004; two juveniles (ROM 43324, ROM 43325) from the vicinity of Camp 2 on the Wokomung Massif, Potaro-Siparuni District, Guyana (
<geoCoordinate direction="north" orientation="latitude" precision="1" value="5.1097217">05°06.5833'N</geoCoordinate>
; 
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-59.821114">059°49.2667'W</geoCoordinate>
), 1234 m elevation, collected by A. Lathrop, R. D. MacCulloch and S. Khan between 28-31 October 2004; and one adult male (CPI11000) from Falls Camp on the Wokomung Massif, Potaro-Siparuni District, Guyana (
<geoCoordinate direction="north" orientation="latitude" precision="1" value="5.090555">05°05.4333'N</geoCoordinate>
; 
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-59.838055">059°50.2833'W</geoCoordinate>
), ca. 1371 m elevation, collected by D. Bruce Means on 24 July 2003.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="diagnosis">
<paragraph pageId="0" pageNumber="99">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="99">
The following characteristics pertain to preserved specimens unless otherwise noted. A medium-sized 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="genus">Anomaloglossus</taxonomicName>
differing from other species in the genus by the following combination of characters: (1) mean SVL in males 18.53 mm (18.15-18.86 mm, n = 3), mean SVL in females 19.15 mm (17.66-21.26, n = 5); (2) skin on dorsum shagreened, venter smooth; (3) tympanic annulus visible anteroventrally; (4) Fingers I and II subequal in length, FI = FII when fingers adpressed; (5) tip of Finger IV not surpassing the base of the distal subarticular tubercle on Finger III when fingers adpressed; (6) distal subarticular tubercle on Finger III and IV present; (7) Finger III swollen in males (conspicuous pre- and postaxial swelling in breeding males); (8) fringes on fingers absent; (9) toes basally webbed, fringes on toes absent; (10) tarsal keel well defined, slightly tubercle-like and weakly curved at proximal end; (11) black arm gland absent, glandular supracarpal pad present in both sexes (larger and more glandular in males); (12) cloacal tubercles absent; (13) pale paracloacal mark present; (14) in life, thin dorsolateral stripe present, from tip of snout to tip of urostyle (not visible, or only barely distinguishable in preservative); (15) ventrolateral stripe absent, but presence of irregular white blotches on the lower flank; (16) oblique lateral stripe absent; (17) sexual dichromatism in throat colour pattern: throat heavily pigmented with melanophores in males (dark brown to black in life), immaculate cream in females (yellowish-orange in life); (18) sexual dichromatism in ventral colour pattern: belly pigmented with melanophores in males, immaculate cream in females; (19) in life, iris metallic reddish bronze with fine dark brown reticulation; (20) large intestine extensively pigmented; (21) testes cream, unpigmented; (22) mature oocytes partly pigmented; (23) median lingual process small, longer than wide, tapered; (24) maxillary teeth present, small.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="comparisons">
<paragraph pageId="0" pageNumber="99">Comparisons.</paragraph>
<paragraph pageId="0" pageNumber="99">
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. can mainly be distinguished from the four described species belonging to the degranvillei group [sensu 
<bibRefCitation author="Vacher, J-P" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="99" pagination="158 - 173" title="Cryptic diversity in Amazonian frogs: integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield." url="https://doi.org/10.1016/j.ympev.2017.04.017" volume="112" year="2017">Vacher et al. 2017</bibRefCitation>
and 
<bibRefCitation author="Fouquet, A" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" pagination="001 - 023" title="On the brink of extinction: two new species of Anomaloglossus from French Guiana and amended definition of Anomaloglossusdegranvillei and A. surinamensis (Anura: Aromobatidae)." url="https://doi.org/10.11646/zootaxa.4379.1.1" volume="4379" year="2018">Fouquet et al. 2018</bibRefCitation>
, i.e. 
<taxonomicName lsidName="A. blanci" pageId="0" pageNumber="99" rank="species" species="blanci">A. blanci</taxonomicName>
Fouquet, Vacher, Courtois, Villette, Reizine, Gaucher, Jairam, Ouboter &amp; Kok, 2018, 
<taxonomicName lsidName="A. degranvillei" pageId="0" pageNumber="99" rank="species" species="degranvillei">A. degranvillei</taxonomicName>
(Lescure, 1975), 
<taxonomicName lsidName="A. dewynteri" pageId="0" pageNumber="99" rank="species" species="dewynteri">A. dewynteri</taxonomicName>
Fouquet, Vacher, Courtois, Villette, Reizine, Gaucher, Jairam, Ouboter &amp; Kok, 2018 and 
<taxonomicName lsidName="A. surinamensis" pageId="0" pageNumber="99" rank="species" species="surinamensis">A. surinamensis</taxonomicName>
Ouboter &amp; Jairam, 2012; characters in parentheses] by having FI = FII when fingers adpressed (FI&gt; FII), the tympanic annulus anteroventrally conspicuous (inconspicuous), and a conspicuous thin dorsolateral stripe from tip of snout to tip of urostyle (absent or inconspicuous).
</paragraph>
<paragraph pageId="0" pageNumber="99">
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. can mainly be distinguished from the four described species belonging to the 
<taxonomicName lsidName="stepheni" pageId="0" pageNumber="99" rank="species" species="stepheni">stepheni</taxonomicName>
group [sensu 
<bibRefCitation author="Vacher, J-P" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="99" pagination="158 - 173" title="Cryptic diversity in Amazonian frogs: integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield." url="https://doi.org/10.1016/j.ympev.2017.04.017" volume="112" year="2017">Vacher et al. 2017</bibRefCitation>
, i.e. 
<taxonomicName lsidName="A. apiau" pageId="0" pageNumber="99" rank="species" species="apiau">A. apiau</taxonomicName>
Fouquet, Souza, Nunes, Kok, Curcio, Carvalho, Grant &amp; Rodrigues, 2015, 
<taxonomicName lsidName="A. baeobatrachus" pageId="0" pageNumber="99" rank="species" species="baeobatrachus">A. baeobatrachus</taxonomicName>
(Boistel &amp; de Massary, 1999), 
<taxonomicName lsidName="A. leopardus" pageId="0" pageNumber="99" rank="species" species="leopardus">A. leopardus</taxonomicName>
Ouboter &amp; Jairam, 2012 and 
<taxonomicName lsidName="A. stepheni" pageId="0" pageNumber="99" rank="species" species="stepheni">A. stepheni</taxonomicName>
(Martins, 1989); characters in parentheses] in lacking an oblique lateral stripe (present, even if short and discontinuous in 
<taxonomicName lsidName="A. apiau" pageId="0" pageNumber="99" rank="species" species="apiau">A. apiau</taxonomicName>
), and in having a conspicuous thin dorsolateral stripe from tip of snout to tip of urostyle (absent).
</paragraph>
<paragraph pageId="0" pageNumber="99">
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. can mainly be distinguished from the three described species belonging to the 
<taxonomicName lsidName="megacephalus" pageId="0" pageNumber="99" rank="species" species="megacephalus">megacephalus</taxonomicName>
group [sensu 
<bibRefCitation author="Grant, T" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" url="https://doi.org/10.2994/SAJH-D-17-00017.1" year="2017">Grant et al. 2017</bibRefCitation>
, i.e. 
<taxonomicName lsidName="A. megacephalus" pageId="0" pageNumber="99" rank="species" species="megacephalus">A. megacephalus</taxonomicName>
Kok, MacCulloch, Lathrop, Willaert &amp; Bossuyt, 2010, 
<taxonomicName lsidName="A. verbeeksnyderorum" pageId="0" pageNumber="99" rank="species" species="verbeeksnyderorum">A. verbeeksnyderorum</taxonomicName>
<normalizedToken originalValue="Barrio-Amorós">Barrio-Amoros</normalizedToken>
, Santos &amp; Jovanovic, 2010, 
<taxonomicName lsidName="A. wothuja" pageId="0" pageNumber="99" rank="species" species="wothuja">A. wothuja</taxonomicName>
(
<normalizedToken originalValue="Barrio-Amorós">Barrio-Amoros</normalizedToken>
, Fuentes-Ramos &amp; Rivas-Fuenmayor, 2004); characters in parentheses] in having only basal toe webbing (moderate to extensive), in lacking an oblique lateral stripe (present, even if short and/or discontinuous), and in having a conspicuous thin dorsolateral stripe from tip of snout to tip of urostyle (absent).
</paragraph>
<paragraph pageId="0" pageNumber="99">
Compared to the other five species belonging to the 
<taxonomicName lsidName="beebei" pageId="0" pageNumber="99" rank="species" species="beebei">beebei</taxonomicName>
group [sensu 
<bibRefCitation author="Grant, T" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" url="https://doi.org/10.2994/SAJH-D-17-00017.1" year="2017">Grant et al. 2017</bibRefCitation>
, i.e. 
<taxonomicName lsidName="A. beebei" pageId="0" pageNumber="99" rank="species" species="beebei">A. beebei</taxonomicName>
(Noble, 1923), 
<taxonomicName lsidName="A. kaiei" pageId="0" pageNumber="99" rank="species" species="kaiei">A. kaiei</taxonomicName>
(Kok, Sambhu, Roopsind, Lenglet &amp; Bourne, 2006), 
<taxonomicName lsidName="A. praderioi" pageId="0" pageNumber="99" rank="species" species="praderioi">A. praderioi</taxonomicName>
, 
<taxonomicName lsidName="A. roraima" pageId="0" pageNumber="99" rank="species" species="roraima">A. roraima</taxonomicName>
(La Marca, 1997) and 
<taxonomicName lsidName="A. rufulus" pageId="0" pageNumber="99" rank="species" species="rufulus">A. rufulus</taxonomicName>
(Gorzula, 1990)], 
<taxonomicName lsidName="A. meansi" pageId="0" pageNumber="99" rank="species" species="meansi">A. meansi</taxonomicName>
sp. n. can easily be distinguished from 
<taxonomicName lsidName="A. beebei" pageId="0" pageNumber="99" rank="species" species="beebei">A. beebei</taxonomicName>
by its larger size in males (maximum SVL 18.86 mm in 
<taxonomicName lsidName="A. meansi" pageId="0" pageNumber="99" rank="species" species="meansi">A. meansi</taxonomicName>
[n = 3,] versus maximum SVL 16.80 mm [n=18] in 
<taxonomicName lsidName="A. beebei" pageId="0" pageNumber="99" rank="species" species="beebei">A. beebei</taxonomicName>
), smooth ventral skin (granular in 
<taxonomicName lsidName="A. beebei" pageId="0" pageNumber="99" rank="species" species="beebei">A. beebei</taxonomicName>
), basal toe webbing (moderate in 
<taxonomicName lsidName="A. beebei" pageId="0" pageNumber="99" rank="species" species="beebei">A. beebei</taxonomicName>
), and in having a conspicuous thin dorsolateral stripe from tip of snout to tip of urostyle (absent or originating from the posterior corner of eye); from 
<taxonomicName lsidName="A. kaiei" pageId="0" pageNumber="99" rank="species" species="kaiei">A. kaiei</taxonomicName>
in having a conspicuous thin dorsolateral stripe from tip of snout to tip of urostyle (originating from the posterior corner of eye in 
<taxonomicName lsidName="A. kaiei" pageId="0" pageNumber="99" rank="species" species="kaiei">A. kaiei</taxonomicName>
) and a black throat in preserved males (greyish, never black in 
<taxonomicName lsidName="A. kaiei" pageId="0" pageNumber="99" rank="species" species="kaiei">A. kaiei</taxonomicName>
); from 
<taxonomicName lsidName="A. roraima" pageId="0" pageNumber="99" rank="species" species="roraima">A. roraima</taxonomicName>
by its larger size in females (maximum SVL 21.26 mm in 
<taxonomicName lsidName="A. meansi" pageId="0" pageNumber="99" rank="species" species="meansi">A. meansi</taxonomicName>
[n = 3,] versus maximum SVL 19.30 mm [n = 18] in 
<taxonomicName lsidName="A. roraima" pageId="0" pageNumber="99" rank="species" species="roraima">A. roraima</taxonomicName>
), smooth ventral skin (granular in 
<taxonomicName lsidName="A. roraima" pageId="0" pageNumber="99" rank="species" species="roraima">A. roraima</taxonomicName>
), and in having a conspicuous thin dorsolateral stripe from tip of snout to tip of urostyle (when present originating from the anterior or posterior corner of eye in 
<taxonomicName lsidName="A. roraima" pageId="0" pageNumber="99" rank="species" species="roraima">A. roraima</taxonomicName>
); from 
<taxonomicName lsidName="A. rufulus" pageId="0" pageNumber="99" rank="species" species="rufulus">A. rufulus</taxonomicName>
in having a conspicuous thin dorsolateral stripe from tip of snout to tip of urostyle (absent in 
<taxonomicName lsidName="A. rufulus" pageId="0" pageNumber="99" rank="species" species="rufulus">A. rufulus</taxonomicName>
) and the posterior part of belly unmarked (heavily marbled in 
<taxonomicName lsidName="A. rufulus" pageId="0" pageNumber="99" rank="species" species="rufulus">A. rufulus</taxonomicName>
). 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. is most similar to 
<taxonomicName lsidName="A. praderioi" pageId="0" pageNumber="99" rank="species" species="praderioi">A. praderioi</taxonomicName>
with which it shares a conspicuous thin dorsolateral stripe from tip of snout to tip of urostyle but is immediately distinguished by its smaller size in males (maximum SVL 18.86 mm in 
<taxonomicName lsidName="A. meansi" pageId="0" pageNumber="99" rank="species" species="meansi">A. meansi</taxonomicName>
[n = 3,] versus maximum SVL 22.40 mm [n = 11] in 
<taxonomicName lsidName="A. praderioi" pageId="0" pageNumber="99" rank="species" species="praderioi">A. praderioi</taxonomicName>
), the absence of fringes on toes (extensive in 
<taxonomicName lsidName="A. praderioi" pageId="0" pageNumber="99" rank="species" species="praderioi">A. praderioi</taxonomicName>
), Finger III with pre- and postaxial swelling in breeding males (preaxial swelling only in 
<taxonomicName lsidName="A. praderioi" pageId="0" pageNumber="99" rank="species" species="praderioi">A. praderioi</taxonomicName>
), less toe webbing (compare Figure 3 with figure 2 in 
<bibRefCitation author="Kok, PJR" journalOrPublisher="Papeis Avulsos de Zoologia" pageId="0" pageNumber="99" pagination="51 - 68" title="A redescription of Anomaloglossuspraderioi (La Marca, 1998) (Anura: Aromobatidae: Anomaloglossinae), with description of its tadpole and call." volume="50" year="2010">Kok 2010</bibRefCitation>
), and the lack of black spots on chest and lower flanks in males (present in 
<taxonomicName lsidName="A. praderioi" pageId="0" pageNumber="99" rank="species" species="praderioi">A. praderioi</taxonomicName>
).
</paragraph>
<caption pageId="0" pageNumber="99">
<paragraph pageId="0" pageNumber="99">
Figure 3. 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. in preservative. A male holotype ROM 43896, dorsal view B male holotype ROM 43896, ventral view C female paratype ROM 43323, dorsal view D female paratype ROM 43323, ventral view E male holotype ROM 43896, ventral view of right hand F male holotype ROM 43896, ventral view of right foot G female paratype ROM 43323, ventral view of left hand H female paratype ROM 43323, ventral view of left foot.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="99">
Compared to the remainder 12 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="genus">Anomaloglossus</taxonomicName>
species not yet assigned to any group [
<taxonomicName lsidName="A. ayarzaguenai" pageId="0" pageNumber="99" rank="species" species="ayarzaguenai">A. ayarzaguenai</taxonomicName>
(La Marca, 1997), 
<taxonomicName lsidName="A. breweri" pageId="0" pageNumber="99" rank="species" species="breweri">A. breweri</taxonomicName>
(
<normalizedToken originalValue="Barrio-Amorós">Barrio-Amoros</normalizedToken>
, 2006), 
<taxonomicName lsidName="A. guanayensis" pageId="0" pageNumber="99" rank="species" species="guanayensis">A. guanayensis</taxonomicName>
(La Marca, 1997), 
<taxonomicName lsidName="A. moffetti" pageId="0" pageNumber="99" rank="species" species="moffetti">A. moffetti</taxonomicName>
<normalizedToken originalValue="Barrio-Amorós">Barrio-Amoros</normalizedToken>
&amp; 
<normalizedToken originalValue="Brewer-Carías">Brewer-Carias</normalizedToken>
, 2008, 
<taxonomicName lsidName="A. murisipanensis" pageId="0" pageNumber="99" rank="species" species="murisipanensis">A. murisipanensis</taxonomicName>
(La Marca, 1997), 
<taxonomicName lsidName="A. parimae" pageId="0" pageNumber="99" rank="species" species="parimae">A. parimae</taxonomicName>
(La Marca, 1997), 
<taxonomicName lsidName="A. parkerae" pageId="0" pageNumber="99" rank="species" species="parkerae">A. parkerae</taxonomicName>
(Meinhardt &amp; Parmelee, 1996), 
<taxonomicName lsidName="A. shrevei" pageId="0" pageNumber="99" rank="species" species="shrevei">A. shrevei</taxonomicName>
(Rivero, 1961), 
<taxonomicName lsidName="A. tamacuarensis" pageId="0" pageNumber="99" rank="species" species="tamacuarensis">A. tamacuarensis</taxonomicName>
(Myers &amp; Donnelly, 1997), 
<taxonomicName lsidName="A. tepequem" pageId="0" pageNumber="99" rank="species" species="tepequem">A. tepequem</taxonomicName>
Fouquet, Souza, Nunes, Kok, Curcio, Carvalho, Grant &amp; Rodrigues, 2015, 
<taxonomicName lsidName="A. tepuyensis" pageId="0" pageNumber="99" rank="species" species="tepuyensis">A. tepuyensis</taxonomicName>
(La Marca, 1997) and 
<taxonomicName lsidName="A. triunfo" pageId="0" pageNumber="99" rank="species" species="triunfo">A. triunfo</taxonomicName>
(
<normalizedToken originalValue="Barrio-Amorós">Barrio-Amoros</normalizedToken>
, Fuentes-Ramos &amp; Rivas-Fuenmayor, 2004); characters in parentheses], 
<taxonomicName lsidName="A. meansi" pageId="0" pageNumber="99" rank="species" species="meansi">A. meansi</taxonomicName>
sp. n. mainly differs in having only basal toe webbing (moderate to extensive), and in having a conspicuous thin dorsolateral stripe from tip of snout to tip of urostyle (absent).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="description">
<paragraph pageId="0" pageNumber="99">Description of the holotype.</paragraph>
<paragraph pageId="0" pageNumber="99">Adult male (ROM 43896; Figure 3), 18.58 mm SVL, in suboptimal state of preservation (extensive ventral incisions, dorsal skin locally damaged); body robust; head as wide as long, HL 99.7% of HW, HW 32% of SVL; dorsal skin shagreened; ventral skin smooth; snout moderately long, SL 47% of HL, 128% of EL, round in dorsal view, protruding in lateral view, extending past lower jaw; nares located close to tip of snout, directed posterolaterally, visible from front, barely visible in dorsal and ventral views, EN 28% of HL, 77% of ED, EN 60% of SL, TSL 49% of SL; posterior rim of naris bordered posteriorly by an inconspicuous crescent-shaped ridge; IND 40% of HW; canthus rostralis rounded; loreal region concave; IOD 104% of EL, longer than upper eyelid; postrictal tubercles low and inconspicuous; tympanic membrane inconspicuous, round, concealed posterodorsally by a diffuse supratympanic swelling; tympanic annulus visible anteroventrally, TYM 52 % of EL; choanae small, circular, located anterolaterally. Maxillary teeth present, small. Tongue longer than wide, free posteriorly, with rounded margin, small median lingual process longer than wide, tapered. Vocal slits bilateral, large, extending from edge of tongue to angle of jaw.</paragraph>
<paragraph pageId="0" pageNumber="99">Forelimb swollen, robust, 94% of FAL. Ulnar fold absent, metacarpal ridge absent; swollen, glandular supracarpal pad present, heavily pigmented with melanophores; hand moderate in size, 24% of SVL, 75% of HW; relative length of fingers III&gt;II=I=IV; pre- and postaxial swelling on third finger (i.e. Finger III swollen); fingers without fringes; tip of Finger IV not reaching distal subarticular tubercle on Finger III when fingers adpressed; finger discs expanded, wider than long, about 1.4 times width of digit; width of disc on Finger III 0.52 mm; palmar tubercle large, egg shaped, 0.72 mm (larger than Finger III disc), thenar tubercle smaller, elliptical; one or two round to ovoid subarticular tubercles (one each on Fingers I and II, two each on Fingers III and IV, with distal tubercle on Finger IV less conspicuous).</paragraph>
<paragraph pageId="0" pageNumber="99">Hind limb robust, moderately long, with heel of adpressed leg reaching posterior corner of eye; skin granular with no cloacal tubercles discernible (but this could be an artefact of preservation); TL 46% of SVL, heels not in contact when hind limbs are flexed at right angle to sagittal plane of body; FL 38% of SVL; relative length of adpressed toes IV&gt;III&gt;V&gt;II&gt;I; Toe I very short, its tip barely reaching the base of subarticular tubercle of Toe II when adpressed; toe discs larger than width of toes; disc on Toe I only slightly larger than width of digit; width of disc on Toe IV 0.67 mm; toes basally webbed, lateral fringes absent; one to three round to ovoid subarticular tubercles (one each on Toes I and II, two each on Toes III and V, and three on Toe IV, with distal tubercle on Toe IV the smallest and least conspicuous). Inner metatarsal tubercle protuberant elliptical, 0.47 mm in length, outer metatarsal tubercle round, protuberant, pigmented, 0.35 mm in diameter. No medial metatarsal tubercle discernible. Tarsal keel slightly tubercle-like and weakly curved at proximal end, extending distally to preaxial edge of Toe I. Metatarsal fold not visible.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="colour of holotype in life">
<paragraph pageId="0" pageNumber="99">Colour of holotype in life.</paragraph>
<paragraph pageId="0" pageNumber="99">Dorsal ground colour chestnut brown with a short black middorsal line between shoulders. A black line from snout tip through eye, extending dorsolaterally to groin. A narrow pale brown dorsolateral stripe above this line, blending into the chestnut dorsal ground colour. Upper surface of limbs light brown proximally, becoming dark brown distally. Flanks reddish brown with yellow spots on lower flanks. Venter pale brown with dark brown mottling, throat very dark brown to black. Underside of limbs orange-red, changing to dark reddish brown on distal forearms.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="colour of holotype in preservative">
<paragraph pageId="0" pageNumber="99">Colour of holotype in preservative.</paragraph>
<paragraph pageId="0" pageNumber="99">After more than 13 years in preservative, dorsal ground colour became dark chestnut brown with a short middorsal black longitudinal line in the scapular region. No other dorsal marking present. Dorsal surface of arms varies from light brown proximally to dark brown, purplish-black towards the granular supracarpal pads. Dorsal surface of legs light brown with darker brown markings. Flanks dark brown to purplish-black with pale spots on lower flanks. Narrow pale brown dorsolateral stripes indistinguishable from dorsal ground colour, although the black dorsolateral stripe remains visible. Throat black, heavily pigmented with melanophores; belly cream, pigmented with melanophores (less densely distributed than on throat). Pale paracloacal marks are visible. Palms dark brown, soles medium brown (Figure 3).</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="description">
<paragraph pageId="0" pageNumber="99">Measurements of holotype</paragraph>
<paragraph pageId="0" pageNumber="99">(in mm).SVL = 18.58; HL = 5.91; HW = 5.93; IOD = 2.26; EN = 1.68; SL = 2.77; TSL = 1.35; EL = 2.16; TYM = 1.12; IND = 2.39; HAND I = 3.06; HAND II = 3.2; HAND III = 4.44; HAND IV = 3.09; WFD = 0.52; FAL = 3.81; THL = 7.80; TIL = 8.61; TAL = 4.48; FL = 7.08; WTD = 0.67.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="sexual dimorphism and variation within the type series">
<paragraph pageId="0" pageNumber="99">Sexual dimorphism and variation within the type series.</paragraph>
<paragraph pageId="0" pageNumber="99">Males are usually smaller than females, 18.15-18.86 mm SVL (n = 3) versus 17.66-21.26 mm SVL (n = 5) in females, with Finger III distinctly swollen in breeding males (Figure 3). Supracarpal pads are less extended and less glandular in females and juveniles than in males. Colouration is sexually dichromatic; throat heavily pigmented black in males (immaculate yellowish-orange in females), and belly yellowish-orange pigmented with melanophores in males (immaculate yellowish-orange in females) (Figure 3). Venter immaculate in juveniles, although small pigmented areas on throat may occur (presumably in juvenile males).</paragraph>
<paragraph pageId="0" pageNumber="99">Morphometric variation is summarized in Table 1, illustrations of a male and a female paratype in life are in Figure 4. Snout in dorsal and ventral views varies from round to truncate (the latter more particularly in females, see Figure 3).</paragraph>
<caption pageId="0" pageNumber="99">
<paragraph pageId="0" pageNumber="99">
Figure 4. 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. in life. A female paratype ROM 43332, dorsal view B female paratype ROM 43329, dorsolateral view C male paratype CPI 11000, dorsolateral view. Photographs (A, B) by AL; photograph (C) courtesy D. Bruce Means.
</paragraph>
</caption>
<caption pageId="0" pageNumber="99">
<paragraph pageId="0" pageNumber="99">
Table 1. Morphometric measurements (in mm) of the type series of 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. Abbreviations are defined in the text. Means 
<normalizedToken originalValue="±">+/-</normalizedToken>
SD are followed by the range in parentheses.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="99">
<table pageId="0" pageNumber="99">
<tr pageId="0" pageNumber="99">
<th colspan="1" pageId="0" pageNumber="99" rowspan="1">Character</th>
<th colspan="1" pageId="0" pageNumber="99" rowspan="1">Male (n = 3)</th>
<th colspan="1" pageId="0" pageNumber="99" rowspan="1">Female (n = 5)</th>
<th colspan="1" pageId="0" pageNumber="99" rowspan="1">Juvenile (n = 3)</th>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">SVL</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">HW</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">IOD</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">HL</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">EN</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">SL</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">EL</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">TYM</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">IND</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">TSL</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">HAND</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">HAND</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">HAND</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">WFD</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">FAL</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">THL</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">TIL</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">TAL</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">FL</td>
</tr>
<tr pageId="0" pageNumber="99">
<td colspan="1" pageId="0" pageNumber="99" rowspan="1">WTD</td>
</tr>
</table>
</paragraph>
<paragraph pageId="0" pageNumber="99">There is substantial variation in colour among preserved individuals, obviously due to preservation artefact (CPI11000 for instance is much lighter than all other individuals). Lower lip pigmented in all male and juvenile individuals, but only in three out of five females. The interorbital region is usually darker than the dorsal ground colour. A short middorsal dark brown/black longitudinal line usually present in the scapular region. One female (ROM 43329) has a diffuse diamond shape marking on the anterior dorsum. Upper surface of arms and legs is cream to dark brown, with darker markings on legs. Palms and soles are light to dark brown. Flanks vary from cream to very dark purplish brown.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="distribution">
<paragraph pageId="0" pageNumber="99">Distribution and natural history.</paragraph>
<paragraph pageId="0" pageNumber="99">The only localities documented for the new species are depicted in Figure 2. Specimens were collected in cloud forest (Figure 5), on the ground or low vegetation. Most were collected after nightfall, although one adult and one juvenile were collected during daylight. Specimens were collected on mountain flanks, not summits; at 1490 m on Ayanganna, and at 1234 m, 1371 m and 1411 m on Wokomung. The majority of specimens (eight) were collected at 1234 m on Wokomung. Fewer were collected at higher elevations; only one each at 1490 m on Ayanganna, 1371 m and 1411 m on Wokomung. This may have been because of habitat differences; high-canopy open forest at lower elevation and dense, low-canopy vegetation at higher elevations (see Discussion).</paragraph>
<caption pageId="0" pageNumber="99">
<paragraph pageId="0" pageNumber="99">
Figure 5. Habitat of 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. on the Wokomung Massif A photograph (looking NE) of the highest part of the massif; the plateau in the centre of the photo is the tallest part of the entire Wokomung Massif B cloud forest at about 1385 m elevation, habitat of 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. Photographs courtesy D. Bruce Means.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="etymology">
<paragraph pageId="0" pageNumber="99">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="99">It is a great pleasure to name this new species after our friend and colleague D. Bruce Means, indefatigable explorer of the &quot;islands in the sky&quot;, and who collected one specimen of the new species and contributed with photographs and data. Thanks to his extensive fieldwork, Bruce Means greatly contributed to our understanding of the distribution, ecology, and taxonomy of Pantepui amphibians and reptiles. The specific epithet should be treated as a noun in the genitive case.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="phylogenetic relationships">
<paragraph pageId="0" pageNumber="99">Phylogenetic relationships.</paragraph>
<paragraph pageId="0" pageNumber="99">
The new species was recovered sister to 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus praderioi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="praderioi">Anomaloglossus praderioi</taxonomicName>
by 
<bibRefCitation author="Grant, T" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" url="https://doi.org/10.1206/0003-0090(2006)299[1:PSODFA]2.0.CO;2" year="2006">Grant et al. (2006</bibRefCitation>
, 
<bibRefCitation author="Grant, T" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" url="https://doi.org/10.2994/SAJH-D-17-00017.1" year="2017">2017</bibRefCitation>
) and 
<bibRefCitation author="Kok, PJR" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" url="https://doi.org/10.1016/j.cub.2012.06.034" year="2012">Kok et al. (2012)</bibRefCitation>
(see Figure 1). Uncorrected p distance in the 
<normalizedToken originalValue="“barcoding”">&quot;barcoding&quot;</normalizedToken>
fragment of the 16S rRNA gene [
<bibRefCitation author="Vences, M" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="99" url="https://doi.org/10.1186/1742-9994-2-5" year="2005">Vences et al. (2005)</bibRefCitation>
; based on the sequences used in 
<bibRefCitation author="Grant, T" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" url="https://doi.org/10.2994/SAJH-D-17-00017.1" year="2017">Grant et al. (2017)</bibRefCitation>
and calculated in PAUP 4.0a161 (
<bibRefCitation pageId="0" pageNumber="99">Swofford 2002</bibRefCitation>
)] is 4.3-4.8% between 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus praderioi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="praderioi">Anomaloglossus praderioi</taxonomicName>
and 
<taxonomicName lsidName="A. meansi" pageId="0" pageNumber="99" rank="species" species="meansi">A. meansi</taxonomicName>
sp. n. Genetic divergence between populations of 
<taxonomicName lsidName="A. praderioi" pageId="0" pageNumber="99" rank="species" species="praderioi">A. praderioi</taxonomicName>
from the slopes of Roraima-tepui and Maringma-tepui is 0.2%, whereas divergence between populations of 
<taxonomicName lsidName="A. meansi" pageId="0" pageNumber="99" rank="species" species="meansi">A. meansi</taxonomicName>
sp. n. from Mount Ayanganna and the Wokomung Massif is 0.9-1.0%.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="conservation status">
<paragraph pageId="0" pageNumber="99">Conservation status.</paragraph>
<paragraph pageId="0" pageNumber="99">
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. is only known from four localities and the 11 specimens used in the description. Virtually nothing is known about its ecology, breeding behaviour and population density. Given the uncertainty on its population status we suggest 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. to be listed as Data Deficient according to the IUCN Red List category guidelines (2014).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="discussion">
<paragraph pageId="0" pageNumber="99">Discussion.</paragraph>
<paragraph pageId="0" pageNumber="99">
Although Ayanganna and Wokomung are close neighbours, the habitats on their slopes are not exactly similar. The slopes of Ayanganna are a series of relatively flat poorly drained plateaus alternating with steeper slopes. Collecting activities were concentrated on the plateaus, where the vegetation consists of dense, low-canopy high-tepui forest, with a dense understory of woody shrubs and large terrestrial bromeliads (
<bibRefCitation author="MacCulloch, RD" journalOrPublisher="Royal Ontario Museum Science Contributions" pageId="0" pageNumber="99" pagination="1 - 36" title="Herpetofauna of Mount Ayanganna, Guyana. Results of the Royal Ontario Museum Ayanganna Expedition 2000." volume="4" year="2009">MacCulloch and Lathrop 2009</bibRefCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="99">
The slopes of Wokomung have no large flat plateaus. Habitat at the collecting sites consists of well-drained slopes covered in lower montane cloud forest with some epiphytes and medium density understory, including scattered terrestrial bromeliads. Streams were common on the slopes (
<bibRefCitation author="MacCulloch, RD" journalOrPublisher="Phyllomedusa" pageId="0" pageNumber="99" pagination="31 - 41" title="Exceptional diversity of Stefania (Anura: Cryptobatrachidae) II: six species from Mount Wokomung, Guyana." url="https://doi.org/10.11606/issn.2316-9079.v5i1p31-41" volume="5" year="2006">MacCulloch et al. 2006</bibRefCitation>
). The majority of specimens were found in this habitat, and this may indicate that 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus meansi" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="meansi">Anomaloglossus meansi</taxonomicName>
sp. n. prefers this to other habitat types; or is a reflection of collecting effort in these habitats.
</paragraph>
<paragraph pageId="0" pageNumber="99">
Species in the 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus beebei" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="beebei">Anomaloglossus beebei</taxonomicName>
group are currently only known from east of the Rio 
<normalizedToken originalValue="Caroní">Caroni</normalizedToken>
, in the Eastern Pantepui District. 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus rufulus" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="species" species="rufulus">Anomaloglossus rufulus</taxonomicName>
is restricted to the highlands of the eastern part of the 
<normalizedToken originalValue="Chimantá">Chimanta</normalizedToken>
Massif in Venezuela, whereas 
<taxonomicName lsidName="A. kaiei" pageId="0" pageNumber="99" rank="species" species="kaiei">A. kaiei</taxonomicName>
has a rather large distribution in the uplands of the Pakaraima Mountains of Guyana (Figure 2). The sister species 
<taxonomicName lsidName="A. roraima" pageId="0" pageNumber="99" rank="species" species="roraima">A. roraima</taxonomicName>
and 
<taxonomicName lsidName="A. beebei" pageId="0" pageNumber="99" rank="species" species="beebei">A. beebei</taxonomicName>
are allopatric, 
<taxonomicName lsidName="A. roraima" pageId="0" pageNumber="99" rank="species" species="roraima">A. roraima</taxonomicName>
being restricted to the highlands of the eastern tip of the Eastern Tepui Chain, whereas 
<taxonomicName lsidName="A. beebei" pageId="0" pageNumber="99" rank="species" species="beebei">A. beebei</taxonomicName>
is reported further to the east in the uplands of Kaieteur National Park, the Wokomung Massif and Mount Ayanganna (Figure 2). A similar spatial distribution is detected in the sister species 
<taxonomicName lsidName="A. praderioi" pageId="0" pageNumber="99" rank="species" species="praderioi">A. praderioi</taxonomicName>
and 
<taxonomicName lsidName="A. meansi" pageId="0" pageNumber="99" rank="species" species="meansi">A. meansi</taxonomicName>
sp. n., which are also allopatric, with 
<taxonomicName lsidName="A. praderioi" pageId="0" pageNumber="99" rank="species" species="praderioi">A. praderioi</taxonomicName>
reported from the uplands of the Eastern Tepui Chain, whereas 
<taxonomicName lsidName="A. meansi" pageId="0" pageNumber="99" rank="species" species="meansi">A. meansi</taxonomicName>
sp. n. is only known further to the east in the uplands of Mount Wokomung and Mount Ayanganna.
</paragraph>
<paragraph pageId="0" pageNumber="99">
As mentioned above, virtually nothing is known about the ecology of 
<taxonomicName lsidName="A. meansi" pageId="0" pageNumber="99" rank="species" species="meansi">A. meansi</taxonomicName>
sp. n. Based on its phylogenetic position it is likely this species has an exotrophic, lentic tadpole (Figure 1). Comprehensive ecological data are crucial for the assessment of species conservation status, but these assessments are known for a few species only in the Pantepui region and there is a high risk that population declines remain unnoticed in such remote areas.
</paragraph>
<paragraph pageId="0" pageNumber="99">
Two additional phylogenetically distinct species of 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="genus">Anomaloglossus</taxonomicName>
remain to be described in the 
<taxonomicName lsidName="megacephalus" pageId="0" pageNumber="99" rank="species" species="megacephalus">megacephalus</taxonomicName>
group (see 
<bibRefCitation author="Grant, T" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" url="https://doi.org/10.2994/SAJH-D-17-00017.1" year="2017">Grant et al. 2017</bibRefCitation>
; the authors, in progress), but several locally restricted 
<taxonomicName class="Amphibia" family="Aromobatidae" genus="Anomaloglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Anomaloglossus" order="Anura" pageId="0" pageNumber="99" phylum="Chordata" rank="genus">Anomaloglossus</taxonomicName>
species probably await discovery (
<bibRefCitation author="Vacher, J-P" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="99" pagination="158 - 173" title="Cryptic diversity in Amazonian frogs: integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield." url="https://doi.org/10.1016/j.ympev.2017.04.017" volume="112" year="2017">Vacher et al. 2017</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>