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<mods:title id="2330C05FC340B76A541CDD36EF285C26">A revision of the Dwarf Geckos, genus Lygodactylus (Squamata: Gekkonidae) from Angola, with the description of three new species</mods:title>
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<mods:namePart id="2F28CAD1DF2D27544C5C3842E7E63E30">Marques, Mariana P.</mods:namePart>
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<mods:affiliation id="2DF486F49038A5891913763D29DDBDB2">Research Center in Biodiversity and Genetic Resources (CIBIO), InBIO, University of Porto, Rua Padre Armando Quintas 7, Vairão, 4485 - 661 Porto, Portugal &amp; mpltmarques @ gmail. com; https: // orcid. org / 0000 - 0002 - 1712 - 2632 &amp; Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Rua da Escola Politécnica, 58, 1269 - 102 Lisboa, Portugal &amp; mpltmarques @ gmail. com; https: // orcid. org / 0000 - 0002 - 1712 - 2632 &amp; lmceriaco @ mhnc. up. pt; https: // orcid. org / 0000 - 0002 - 0591 - 9978 &amp; Museu de História Natural e da Ciência da Universidade do Porto, Praça Gomes Teixeira 4099 - 002 Porto, Portugal &amp; lmceriaco @ mhnc. up. pt; https: // orcid. org / 0000 - 0002 - 0591 - 9978 &amp; mpltmarques @ gmail. com; https: // orcid. org / 0000 - 0002 - 1712 - 2632</mods:affiliation>
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<mods:namePart id="04883659AFDFE6F0C7B7FE68FCCD4E48">Ceríaco, Luis M. P.</mods:namePart>
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<mods:affiliation id="574805C8045D71E9EFCC76B75025DD36">Museu de História Natural e da Ciência da Universidade do Porto, Praça Gomes Teixeira 4099 - 002 Porto, Portugal &amp; lmceriaco @ mhnc. up. pt; https: // orcid. org / 0000 - 0002 - 0591 - 9978 &amp; Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Rua da Escola Politécnica, 58, 1269 - 102 Lisboa, Portugal &amp; mpltmarques @ gmail. com; https: // orcid. org / 0000 - 0002 - 1712 - 2632 &amp; lmceriaco @ mhnc. up. pt; https: // orcid. org / 0000 - 0002 - 0591 - 9978</mods:affiliation>
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<mods:namePart id="798E9C08D6CE02E59D6DC84069FE76AC">Buehler, Matthew D.</mods:namePart>
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<mods:affiliation id="DB0284164EA9CE55695D1D2967434A09">Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085 - 1699, USA &amp; mbuehle 3 @ gmail. com; https: // orcid. org / 0000 - 0001 - 5460 - 7770 &amp; adez. bandeira @ gmail. com; https: // orcid. org / 0000 - 0002 - 7496 - 9119 &amp; Department of Biological Sciences &amp; Museum of Natural History, Auburn University, Auburn, Alabama, 36849, USA &amp; mbuehle 3 @ gmail. com; https: // orcid. org / 0000 - 0001 - 5460 - 7770</mods:affiliation>
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<mods:affiliation id="03E14ED91D6BC0A30FB32CA31263A1D2">Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085 - 1699, USA &amp; mbuehle 3 @ gmail. com; https: // orcid. org / 0000 - 0001 - 5460 - 7770 &amp; adez. bandeira @ gmail. com; https: // orcid. org / 0000 - 0002 - 7496 - 9119 &amp; Instituto Nacional da Biodiversidade e Áreas de Conservação, Ministério do Ambiente de Angola, Centralidade do Kilamba, Rua 26 de Fevereiro, Quarteirão Nimi Ya Lukemi, edifício Q 11, 3 º andar, Luanda, Angola. &amp; adez. bandeira @ gmail. com; https: // orcid. org / 0000 - 0002 - 7496 - 9119</mods:affiliation>
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<mods:namePart id="CBDBDE0DBA108E3479882E8876DB5C5A">Janota, Joyce M.</mods:namePart>
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<mods:affiliation id="FFBD655D82D183FB57A0ED5035E41012">Instituto Nacional da Biodiversidade e Áreas de Conservação, Ministério do Ambiente de Angola, Centralidade do Kilamba, Rua 26 de Fevereiro, Quarteirão Nimi Ya Lukemi, edifício Q 11, 3 º andar, Luanda, Angola. &amp; adez. bandeira @ gmail. com; https: // orcid. org / 0000 - 0002 - 7496 - 9119</mods:affiliation>
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<treatment id="1970878C5454FFEC9EB0FF691AA5FC23" ID-DOI="http://doi.org/10.5281/zenodo.4506725" ID-GBIF-Taxon="177213303" ID-Zenodo-Dep="4506725" LSID="urn:lsid:plazi:treatment:1970878C5454FFEC9EB0FF691AA5FC23" httpUri="http://treatment.plazi.org/id/1970878C5454FFEC9EB0FF691AA5FC23" lastPageNumber="340" pageId="39" pageNumber="340">
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<taxonomicName id="56D94D195454FFEC9EB0FF691A2DFF56" ID-CoL="3WQ49" authority="Hewitt, 1926" authorityName="Hewitt" authorityYear="1926" box="[151,600,151,178]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="39" pageNumber="340" phylum="Chordata" rank="species" species="lawrencei">
<emphasis id="A3ADEA885454FFEC9EB0FF691A2DFF56" bold="true" box="[151,600,151,178]" pageId="39" pageNumber="340">
<emphasis id="A3ADEA885454FFEC9EB0FF6919C7FF55" bold="true" box="[151,434,151,178]" italics="true" pageId="39" pageNumber="340">Lygodactylus lawrencei</emphasis>
Hewitt, 1926
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(
<tableCitation id="DC5B03215454FFEC9EB8FF451974FF32" box="[159,257,187,213]" captionStart="TABLE 4" captionStartId="18.[151,239,404,428]" captionText="TABLE 4. Comparison between the different species of Lygodactylus occurring in Angola. Measurements are presented in millimeters (mm). Abbreviations are those described in Materials and Methods section." pageId="39" pageNumber="340">Tables 4</tableCitation>
;
<figureCitation id="09E22A1F5454FFEC9F2AFF451919FF32" box="[269,364,187,213]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURE 3" captionStart-3="FIGURE 4" captionStartId-0="19.[151,250,1799,1823]" captionStartId-1="20.[151,250,1754,1778]" captionStartId-2="21.[151,248,1463,1487]" captionStartId-3="22.[151,250,1008,1032]" captionTargetBox-0="[203,1396,960,1759]" captionTargetBox-1="[151,1436,200,1730]" captionTargetBox-2="[161,1426,190,1426]" captionTargetBox-3="[151,1437,181,985]" captionTargetId-0="figure-386@19.[180,1408,915,1775]" captionTargetId-1="figure-22@20.[151,1436,200,1730]" captionTargetId-2="figure-19@21.[151,1436,181,1440]" captionTargetId-3="figure-22@22.[151,1437,181,985]" captionTargetPageId-0="19" captionTargetPageId-1="20" captionTargetPageId-2="21" captionTargetPageId-3="22" captionText-0="FIGURE 1. Maximum likelihood phylogeny of Lygodactylus using ND2. Nodes supported by ML and BI are indicated by black dots. nodes only supported by the ML tree are indicated with half-filled black dots. nodes only supported by BI are indicated by half-filled red dots. and nodes with low support in both the ML and BI trees have red dots. Species of Lygodactylus occurring in Angola have a star next to them in the tree. Species within the Madgascan Lygodactylus clade include L. tolampyae, L. heterurus, L. arnoulti. Species in the Greater Drakensberg group include; L. nigropunctatus, L. graniticolus, L. ocellatus, L. soutpansbergensis. L. incognitus, L. montiscaeruli, L. waterbergensis, and L. methueni. Note that this tree should be used only as a guide to shallow relationships in the genus and to indicate relative divergence." captionText-1="FIGURE 2. Comparison of the number of precloacal pores in Angolan individuals of Lygodactylus angolensis, L. baptistai sp. nov., L. nyaneka sp. nov., L. lawrencei and L. tchokwe sp. nov." captionText-2="FIGURE 3. Comparison of the nostrils position inAngolan individuals of:A. L. angolensis (AMB 9216); B. Lygodactylus baptistai sp. nov. (CAS 263557); C. L. nyaneka sp. nov. (UF 187229); D. L. lawrencei (CAS 264259); E. L. tchokwe sp. nov. (MD 5312b)." captionText-3="FIGURE 4. Comparison of the tail scales arrangement in Angolan individuals of A. Lygodactylus angolensis (MHNC-UP/REP 533); B. L. nyaneka sp. nov. (CAS 263531); C. L. baptistai sp. nov. (CAS 263557); D. L. lawrencei (CAS 264259); E. L. tchokwe sp. nov. (MD 5312d)." figureDoi-0="http://doi.org/10.5281/zenodo.4410729" figureDoi-1="http://doi.org/10.5281/zenodo.4410731" figureDoi-2="http://doi.org/10.5281/zenodo.4410733" figureDoi-3="http://doi.org/10.5281/zenodo.4410735" httpUri-0="https://zenodo.org/record/4410729/files/figure.png" httpUri-1="https://zenodo.org/record/4410731/files/figure.png" httpUri-2="https://zenodo.org/record/4410733/files/figure.png" httpUri-3="https://zenodo.org/record/4410735/files/figure.png" pageId="39" pageNumber="340">Fig. 1–4</figureCitation>
,
<figureCitation id="09E22A1F5454FFEC9F5FFF4519B5FF32" box="[376,448,187,213]" captionStart-0="FIGURE 18" captionStart-1="FIGURE 19" captionStart-2="FIGURE 20" captionStartId-0="39.[151,250,1815,1839]" captionStartId-1="40.[151,250,918,942]" captionStartId-2="40.[151,250,2017,2041]" captionTargetBox-0="[151,1437,988,1792]" captionTargetBox-1="[258,1328,181,894]" captionTargetId-0="figure-359@39.[151,1437,988,1792]" captionTargetId-1="figure-22@40.[258,1328,181,894]" captionTargetId-2="figure-52@40.[300,1287,1003,1963]" captionTargetPageId-0="39" captionTargetPageId-1="40" captionTargetPageId-2="40" captionText-0="FIGURE 18. Dorsal and ventral whole-body views (left) and dorsal. lateral and ventral views of the head (right) of Lygodactylus lawrencei (CAS 264259)." captionText-1="FIGURE 19. Lygodactylus lawrencei in life (CAS 264259). Maungo [-14.538018º, 12.744793º, 365 m a.s.l.]. Namibe Province. southwestern Angola Photos by Luis M. P. Ceríaco." captionText-2="FIGURE 20. Distribution of Lygodactylus lawrencei in the country." figureDoi-0="http://doi.org/10.5281/zenodo.4410765" figureDoi-1="http://doi.org/10.5281/zenodo.4410767" figureDoi-2="http://doi.org/10.5281/zenodo.4506715" httpUri-0="https://zenodo.org/record/4410765/files/figure.png" httpUri-1="https://zenodo.org/record/4410767/files/figure.png" httpUri-2="https://zenodo.org/record/4506715/files/figure.png" pageId="39" pageNumber="340">18–20</figureCitation>
)
</paragraph>
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<taxonomicName id="56D94D195454FFEC9EB0FEFE19F2FEFF" ID-CoL="3WQ49" authorityName="Hewitt" authorityYear="1926" box="[151,391,256,280]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="39" pageNumber="340" phylum="Chordata" rank="species" species="lawrencei">
<emphasis id="A3ADEA885454FFEC9EB0FEFE19F2FEFF" box="[151,391,256,280]" italics="true" pageId="39" pageNumber="340">Lygodactylus lawrencei</emphasis>
</taxonomicName>
(
<treatmentCitation id="1078108B5454FFEC9FB3FEFE1A3BFEFF" author="Branch, W. R." box="[404,590,256,280]" page="247" pageId="39" pageNumber="340" year="1998">
<bibRefCitation id="F5484B6B5454FFEC9FB3FEFE1A3BFEFF" author="Branch, W. R." box="[404,590,256,280]" pageId="39" pageNumber="340" refId="ref25855" refString="Branch, W. R. (1998) Field Guide to Snakes and Other Reptiles of Southern Africa. 3 rd Edition. Ralph Curtis Books, Sanibel Island, Florida, 399 pp. https: // doi. org / 10.1111 / evo. 13062" type="book" year="1998">Branch 1998: 247</bibRefCitation>
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;
<bibRefCitation id="F5484B6B5454FFEC9C7FFEFE1B52FEFF" author="Marques, M. P. &amp; Ceriaco, L. M. P. &amp; Blackburn, D. C. &amp; Bauer, A. M." box="[600,807,256,280]" pageId="39" pageNumber="340" pagination="1 - 501" refId="ref28241" refString="Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. &amp; Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola-Atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement 2), 1 - 501." type="journal article" year="2018">
Marques
<emphasis id="A3ADEA885454FFEC9C90FEFE1A9EFEFF" box="[695,747,256,280]" italics="true" pageId="39" pageNumber="340">et al.</emphasis>
2018
</bibRefCitation>
: 191,192)
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<paragraph id="9166369A5454FFEC9EB0FEB81C0BFDBB" blockId="39.[151,1437,326,964]" pageId="39" pageNumber="340">
Several authors had suggested the presence of
<taxonomicName id="56D94D195454FFEC9CBBFEB91B53FE87" authorityName="Hewitt" authorityYear="1926" box="[668,806,326,352]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="39" pageNumber="340" phylum="Chordata" rank="species" species="lawrencei">
<emphasis id="A3ADEA885454FFEC9CBBFEB91B53FE87" box="[668,806,326,352]" italics="true" pageId="39" pageNumber="340">L. lawrencei</emphasis>
</taxonomicName>
in southern
<collectingCountry id="E9CE760A5454FFEC9D89FEB81C75FE87" box="[942,1024,326,352]" name="Angola" pageId="39" pageNumber="340">Angola</collectingCountry>
(
<bibRefCitation id="F5484B6B5454FFEC9A28FEB81CD6FE87" author="Branch, W. R." box="[1039,1187,326,352]" pageId="39" pageNumber="340" refId="ref25855" refString="Branch, W. R. (1998) Field Guide to Snakes and Other Reptiles of Southern Africa. 3 rd Edition. Ralph Curtis Books, Sanibel Island, Florida, 399 pp. https: // doi. org / 10.1111 / evo. 13062" type="book" year="1998">Branch 1998</bibRefCitation>
;
<bibRefCitation id="F5484B6B5454FFEC9A89FEB81DE5FE87" author="Marques, M. P. &amp; Ceriaco, L. M. P. &amp; Blackburn, D. C. &amp; Bauer, A. M." box="[1198,1424,326,352]" pageId="39" pageNumber="340" pagination="1 - 501" refId="ref28241" refString="Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. &amp; Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola-Atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement 2), 1 - 501." type="journal article" year="2018">
Marques
<emphasis id="A3ADEA885454FFEC9B31FEB91D3BFE87" box="[1302,1358,326,352]" italics="true" pageId="39" pageNumber="340">et al.</emphasis>
2018
</bibRefCitation>
), but so far no vouchered records confirmed its presence in the country. The species was originally described from the
<collectingRegion id="531DF8785454FFEC9EE4FE701901FE4F" box="[195,372,398,424]" country="Angola" name="Cunene" pageId="39" pageNumber="340">Kunene Region</collectingRegion>
in
<collectingCountry id="E9CE760A5454FFEC9FBAFE701988FE4F" box="[413,509,398,424]" name="Namibia" pageId="39" pageNumber="340">Namibia</collectingCountry>
by
<bibRefCitation id="F5484B6B5454FFEC9C0BFE701AADFE4E" author="Hewitt, J." box="[556,728,398,425]" pageId="39" pageNumber="340" pagination="473 - 490" refId="ref27386" refString="Hewitt, J. (1926 a) Some new or little-known reptiles and batrachians from South Africa. Annals of the South African Museum, 20, 473 - 490." type="journal article" year="1926">Hewitt (1926a)</bibRefCitation>
and has to date been confirmed only in far northwestern
<collectingCountry id="E9CE760A5454FFEC9B55FE7018AFFE2B" name="Namibia" pageId="39" pageNumber="340">Namibia</collectingCountry>
. This rocky, dry savanna specialist is potentially the easiest species of
<taxonomicName id="56D94D195454FFEC9DCFFE4C1C09FE2B" box="[1000,1148,434,460]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="39" pageNumber="340" phylum="Chordata" rank="genus">
<emphasis id="A3ADEA885454FFEC9DCFFE4C1C09FE2B" box="[1000,1148,434,460]" italics="true" pageId="39" pageNumber="340">Lygodactylus</emphasis>
</taxonomicName>
to diagnose in the region, given its stereotypically low number of precloacal pores (two), when compared to its congeners. Our records from Maungo and
<quantity id="56219B7F5454FFEC9F0BFE041907FDF3" box="[300,370,506,532]" metricMagnitude="4" metricUnit="m" metricValue="1.1" pageId="39" pageNumber="340" unit="km" value="11.0">11 km</quantity>
N of Espinheira camp, Iona National Park camp are the first vouchered records of this species for
<collectingCountry id="E9CE760A5454FFEC9E9BFDE0197BFDDF" box="[188,270,542,568]" name="Angola" pageId="39" pageNumber="340">Angola</collectingCountry>
and mark the northernmost known distribution of the species. Comparison of the mitochondrial gene 16S (add GenBank no.) confirms that the Angolan and Namibian populations are conspecific.
</paragraph>
</subSubSection>
<subSubSection id="D9C365115454FFEC9EE0FD9B1AA5FC23" pageId="39" pageNumber="340" type="diagnosis">
<paragraph id="9166369A5454FFEC9EE0FD9B1AA5FC23" blockId="39.[151,1437,326,964]" pageId="39" pageNumber="340">
<emphasis id="A3ADEA885454FFEC9EE0FD9B1937FD98" bold="true" box="[199,322,613,639]" pageId="39" pageNumber="340">Diagnosis.</emphasis>
<taxonomicName id="56D94D195454FFEC9F6CFD981A23FD67" authorityName="Hewitt" authorityYear="1926" box="[331,598,614,640]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="39" pageNumber="340" phylum="Chordata" rank="species" species="lawrencei">
<emphasis id="A3ADEA885454FFEC9F6CFD981A23FD67" box="[331,598,614,640]" italics="true" pageId="39" pageNumber="340">Lygodactylus lawrencei</emphasis>
</taxonomicName>
is small dwarf gecko with a maximum SVL of
<quantity id="56219B7F5454FFEC9AA3FD981CACFD67" box="[1156,1241,614,640]" metricMagnitude="-2" metricUnit="m" metricValue="3.0" pageId="39" pageNumber="340" unit="mm" value="30.0">30 mm</quantity>
. It can be distinguished from other members of the genus occurring in
<collectingCountry id="E9CE760A5454FFEC9D2DFD741B29FD43" box="[778,860,650,676]" name="Angola" pageId="39" pageNumber="340">Angola</collectingCountry>
and surrounding regions by having 1) three scales surrounding the nostril (1
<superScript id="66AC9BD25454FFEC9F93FD5019CAFD5B" attach="left" box="[436,447,686,700]" fontSize="6" pageId="39" pageNumber="340">st</superScript>
supralabial, two supranasals); 2) nostrils separated from the rostral and pierced between supranasal and first labial; 3) mental followed by two postmentals; 4) first infralabial with &lt;25% overlap with postpostmental; 5) 16 to 17 rows of ventral scales at midbody; 6) only two precloacal pores in males; 7) subcaudals arranged in rows of imbricate scales about ⅓ (but mostly subequal) of width of tail (
<figureCitation id="09E22A1F5454FFEC9A78FCE41CCEFCD3" box="[1119,1211,794,820]" captionStart="FIGURE 4" captionStartId="22.[151,250,1008,1032]" captionTargetBox="[151,1437,181,985]" captionTargetId="figure-22@22.[151,1437,181,985]" captionTargetPageId="22" captionText="FIGURE 4. Comparison of the tail scales arrangement in Angolan individuals of A. Lygodactylus angolensis (MHNC-UP/REP 533); B. L. nyaneka sp. nov. (CAS 263531); C. L. baptistai sp. nov. (CAS 263557); D. L. lawrencei (CAS 264259); E. L. tchokwe sp. nov. (MD 5312d)." figureDoi="http://doi.org/10.5281/zenodo.4410735" httpUri="https://zenodo.org/record/4410735/files/figure.png" pageId="39" pageNumber="340">Fig. 4D</figureCitation>
); 8) venter white to cream; 9) black chevron markings on the gular region. Background usually ashy-grey with thin dark bands, usually broken. A large streak is visible from nostril to the forelimb insertion, passing through the eye. Often a pale chestnut lateral stripe is visible, extending from the neck to the anterior half of the tail. The Angolan individual has a very faded chevron pattern in the gular region (
<figureCitation id="09E22A1F5454FFEC9C57FC541AB6FC23" box="[624,707,938,964]" captionStart="FIGURE 18" captionStartId="39.[151,250,1815,1839]" captionTargetBox="[151,1437,988,1792]" captionTargetId="figure-359@39.[151,1437,988,1792]" captionTargetPageId="39" captionText="FIGURE 18. Dorsal and ventral whole-body views (left) and dorsal. lateral and ventral views of the head (right) of Lygodactylus lawrencei (CAS 264259)." figureDoi="http://doi.org/10.5281/zenodo.4410765" httpUri="https://zenodo.org/record/4410765/files/figure.png" pageId="39" pageNumber="340">Fig. 18</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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