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<mods:title id="38FEF0B41AEA68F7EB75A3B1DCFF9AAD">Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean</mods:title>
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<mods:affiliation id="99C58C66AC124F0C00E8A1D08B7A36C9">Zoological Institute, Technical University of Braunschweig, Mendelssohnstr. 4, 38106 Braunschweig, Germany; s. gippner @ tu-braunschweig. de; https: // orcid. org / 0000 - 0003 - 2621 - 1001</mods:affiliation>
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<mods:affiliation id="7740CBD8D38061B9EC4C04560CEDEEE9">Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University, 800 Lancaster Avenue, Villanova, PA 19085, USA;</mods:affiliation>
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<mods:affiliation id="C603ACC770FEE9F8EC0ED811F77C6651">Department of Biological Sciences, Rutgers University-Newark, 195 University Avenue, Newark, NJ 07102, USA; scott. travers @ rutgers. edu; https: // orcid. org / 0000 - 0003 - 4656 - 4613</mods:affiliation>
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<mods:affiliation id="311BD1AC231C1A470E7F14BE501C3F90">Zoological Institute, Technical University of Braunschweig, Mendelssohnstr. 4, 38106 Braunschweig, Germany; s. gippner @ tu-braunschweig. de; https: // orcid. org / 0000 - 0003 - 2621 - 1001</mods:affiliation>
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4.2.
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<emphasis id="B96FEA9FF544E56327B29A740AA8F8A5" bold="true" box="[203,474,1807,1834]" italics="true" pageId="12" pageNumber="244">Lygodactylus insularis</emphasis>
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<taxonomicName id="4C1B4D0EF544E56327EE9A2C0AE1F8FE" authorityName="Boettger" authorityYear="1913" box="[151,403,1879,1905]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="244" phylum="Chordata" rank="species" species="insularis">
<emphasis id="B96FEA9FF544E56327EE9A2C0AE1F8FE" box="[151,403,1879,1905]" italics="true" pageId="12" pageNumber="244">Lygodactylus insularis</emphasis>
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is an endemic species on Juan de Nova, a very small island with a total area of approximately
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, 280 km
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from mainland Africa (East
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) and
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to the nearest point on the Malagasy coast (
<bibRefCitation id="EF8A4B7CF544E56327E69AE40A4BF836" author="Caceres, S." box="[159,313,1951,1977]" pageId="12" pageNumber="244" refId="ref12274" refString="Caceres, S. (2003) Etude prealable pour le classement en Reserve Naturelle des Iles Eparses. Memoire de DESS Sciences et Gestion de l'Environnement Tropical. Univ. Reunion (Laboratoire ECOMAR) &amp; DIREN Reunion, 136 pp. Available from: http: // poupin. joseph. free. fr / pdf / caceres- 2003 - iles-eparses-classement. pdf (accessed 13 June 2023)" type="book" year="2003">Caceres 2003</bibRefCitation>
;
<bibRefCitation id="EF8A4B7CF544E563263A9AE4097FF836" author="Lambs, L. &amp; Mangion, P. &amp; Mougin, E. &amp; Fromard, F." box="[323,525,1951,1977]" pageId="12" pageNumber="244" pagination="311 - 320" refId="ref13445" refString="Lambs, L., Mangion, P., Mougin, E. &amp; Fromard, F. (2016) Water cycle and salinity dynamics in the mangrove forests of Europa and Juan de Nova Islands, southwest Indian Ocean. Rapid Communications in Mass Spectrometry, 30, 311 - 320. https: // doi. org / 10.1002 / rcm. 7435" type="journal article" year="2016">
Lambs
<emphasis id="B96FEA9FF544E56326ED9ADB0AB9F836" box="[404,459,1951,1977]" italics="true" pageId="12" pageNumber="244">et al.</emphasis>
2016
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).
<bibRefCitation id="EF8A4B7CF544E56325649ADB09CDF835" author="Pasteur, G." box="[541,703,1951,1978]" pageId="12" pageNumber="244" pagination="1 - 132" refId="ref14163" refString="Pasteur, G. (1965) Recherches sur l'evolution des lygodactyles, lezards afromalgaches actuels. Travaux de l'Institut Scientifique Cherifien, Serie Zoologie, 29, 1 - 132." type="journal article" year="1965">Pasteur (1965)</bibRefCitation>
already included
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<emphasis id="B96FEA9FF544E56324F89ADB088DF836" box="[897,1023,1951,1977]" italics="true" pageId="12" pageNumber="244">L. insularis</emphasis>
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in the
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<emphasis id="B96FEA9FF544E563233E9ADB0FB0F836" box="[1095,1218,1952,1977]" italics="true" pageId="12" pageNumber="244">L. capensis</emphasis>
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group and assumed that
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<emphasis id="B96FEA9FF544E56327B09ABF0BABF852" box="[201,217,1988,2013]" italics="true" pageId="12" pageNumber="244">L</emphasis>
.
<emphasis id="B96FEA9FF544E56327919AB80A38F852" box="[232,330,1987,2013]" italics="true" pageId="12" pageNumber="244">insularis</emphasis>
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originated from either
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L. c.
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</emphasis>
or
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L. c.
<taxonomicName id="4C1B4D0EF544E56324289ABF08E1F852" authorityName="Sternfeld" authorityYear="1911" box="[849,915,1988,2013]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="244" phylum="Chordata" rank="species" species="grotei">grotei</taxonomicName>
</emphasis>
.
<taxonomicName id="4C1B4D0EF544E56324DB9AB80FEDF852" baseAuthorityName="Smith" baseAuthorityYear="1849" box="[930,1183,1987,2013]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="244" phylum="Chordata" rank="species" species="capensis">
<emphasis id="B96FEA9FF544E56324DB9AB80FEDF852" box="[930,1183,1987,2013]" italics="true" pageId="12" pageNumber="244">Lygodactylus capensis</emphasis>
</taxonomicName>
and
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<emphasis id="B96FEA9FF544E56323A19ABF0E2BF852" box="[1240,1369,1987,2013]" italics="true" pageId="12" pageNumber="244">L. insularis</emphasis>
</taxonomicName>
share the same number of precloacal pores and a superficially similar coloration of the adults. However, they strongly differ in the status of the subcaudals and in the appearance of their hatchlings. It is thus impossible to discern the relationships of
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<emphasis id="B96FEA9FF545E56226359DA40AB9FF76" box="[332,459,223,249]" italics="true" pageId="13" pageNumber="245">L. insularis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C1B4D0EF545E56226A09DA40925FF77" baseAuthorityName="Smith" baseAuthorityYear="1849" box="[473,599,223,248]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="capensis">
<emphasis id="B96FEA9FF545E56226A09DA40925FF77" box="[473,599,223,248]" italics="true" pageId="13" pageNumber="245">L. capensis</emphasis>
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and
<taxonomicName id="4C1B4D0EF545E56225F79DA4099FFF77" authorityName="Sternfeld" authorityYear="1911" box="[654,749,223,248]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="grotei">
<emphasis id="B96FEA9FF545E56225F79DA4099FFF77" box="[654,749,223,248]" italics="true" pageId="13" pageNumber="245">L. grotei</emphasis>
</taxonomicName>
by means of scalation or coloration alone.
</paragraph>
<paragraph id="8BA4368DF545E56227BE9C780810FD42" blockId="13.[151,1437,151,1581]" pageId="13" pageNumber="245">
Our molecular phylogenetic analyses reveal that
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<emphasis id="B96FEA9FF545E56225939C780818FE92" box="[746,874,259,285]" italics="true" pageId="13" pageNumber="245">L. insularis</emphasis>
</taxonomicName>
is embedded within the
<taxonomicName id="4C1B4D0EF545E56223059C780F8BFE93" baseAuthorityName="Smith" baseAuthorityYear="1849" box="[1148,1273,259,284]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="capensis">
<emphasis id="B96FEA9FF545E56223059C780F8BFE93" box="[1148,1273,259,284]" italics="true" pageId="13" pageNumber="245">L. capensis</emphasis>
</taxonomicName>
group, clearly indicating its African origin and, in so far, supporting
<bibRefCitation id="EF8A4B7CF545E56225929C5C08E0FECE" author="Pasteur, G." box="[747,914,294,321]" pageId="13" pageNumber="245" pagination="1 - 132" refId="ref14163" refString="Pasteur, G. (1965) Recherches sur l'evolution des lygodactyles, lezards afromalgaches actuels. Travaux de l'Institut Scientifique Cherifien, Serie Zoologie, 29, 1 - 132." type="journal article" year="1965">Pasteur (1965)</bibRefCitation>
. While the single gene as well as the multigene analyses fully support closer relationships between
<taxonomicName id="4C1B4D0EF545E562259C9C300814FEEA" authorityName="Boettger" authorityYear="1913" box="[741,870,331,357]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="insularis">
<emphasis id="B96FEA9FF545E562259C9C300814FEEA" box="[741,870,331,357]" italics="true" pageId="13" pageNumber="245">L. insularis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1B4D0EF545E56224D89C300F52FEEB" baseAuthorityName="Smith" baseAuthorityYear="1849" box="[929,1056,331,356]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="capensis">
<emphasis id="B96FEA9FF545E56224D89C300F52FEEB" box="[929,1056,331,356]" italics="true" pageId="13" pageNumber="245">L. capensis</emphasis>
</taxonomicName>
than between
<taxonomicName id="4C1B4D0EF545E56223B09C300E39FEEA" authorityName="Boettger" authorityYear="1913" box="[1225,1355,331,357]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="insularis">
<emphasis id="B96FEA9FF545E56223B09C300E39FEEA" box="[1225,1355,331,357]" italics="true" pageId="13" pageNumber="245">L. insularis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1B4D0EF545E56222FF9C300BABFE07" authorityName="Sternfeld" authorityYear="1911" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="grotei">
<emphasis id="B96FEA9FF545E56222FF9C300BABFE07" italics="true" pageId="13" pageNumber="245">L. grotei</emphasis>
</taxonomicName>
, the position of
<taxonomicName id="4C1B4D0EF545E56226E09C140969FE06" authorityName="Boettger" authorityYear="1913" box="[409,539,367,393]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="insularis">
<emphasis id="B96FEA9FF545E56226E09C140969FE06" box="[409,539,367,393]" italics="true" pageId="13" pageNumber="245">L. insularis</emphasis>
</taxonomicName>
within the clade formed by
<taxonomicName id="4C1B4D0EF545E562241D9C140894FE06" authorityName="Boettger" authorityYear="1913" box="[868,998,367,393]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="insularis">
<emphasis id="B96FEA9FF545E562241D9C140894FE06" box="[868,998,367,393]" italics="true" pageId="13" pageNumber="245">L. insularis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C1B4D0EF545E562248F9C140F01FE06" box="[1014,1139,367,393]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="nyaneka">
<emphasis id="B96FEA9FF545E562248F9C140F01FE06" box="[1014,1139,367,393]" italics="true" pageId="13" pageNumber="245">L. nyaneka</emphasis>
</taxonomicName>
,
<emphasis id="B96FEA9FF545E56223FD9C140EEFFE07" box="[1156,1437,367,392]" italics="true" pageId="13" pageNumber="245">
<taxonomicName id="4C1B4D0EF545E56223FD9C140E71FE07" baseAuthorityName="Smith" baseAuthorityYear="1849" box="[1156,1283,367,392]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="capensis">L. capensis</taxonomicName>
<taxonomicNameLabel id="A25C57E4F545E56222749C140EEFFE07" box="[1293,1437,367,392]" pageId="13" pageNumber="245" sensu="stricto">sensu stricto</taxonomicNameLabel>
</emphasis>
and
<specimenCount id="9D1DFD04F545E56227B19CE80A75FE23" box="[200,263,403,429]" pageId="13" pageNumber="245" type="generic">
six
<taxonomicName id="4C1B4D0EF545E56227899CE80A75FE23" baseAuthorityName="Smith" baseAuthorityYear="1849" box="[240,263,403,428]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" isUncertain="true" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="capensis">
<emphasis id="B96FEA9FF545E56227899CE80A75FE23" box="[240,263,403,428]" italics="true" pageId="13" pageNumber="245">L.</emphasis>
</taxonomicName>
</specimenCount>
aff.
<emphasis id="B96FEA9FF545E56226459CE80AEEFE23" box="[316,412,403,428]" italics="true" pageId="13" pageNumber="245">capensis</emphasis>
lineages (provisionally named as “groups”; here, the last two of these are referred to as
<emphasis id="B96FEA9FF545E56222FF9CE80A1FFE5E" italics="true" pageId="13" pageNumber="245">
<taxonomicName id="4C1B4D0EF545E56222FF9CE80B85FE5F" baseAuthorityName="Smith" baseAuthorityYear="1849" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="capensis">L. capensis</taxonomicName>
<taxonomicNameLabel id="A25C57E4F545E56227849CCC0A1FFE5E" box="[253,365,439,465]" pageId="13" pageNumber="245" sensu="lato">sensu lato</taxonomicNameLabel>
</emphasis>
) is not clearly resolved. For a taxonomic revision of the
<taxonomicName id="4C1B4D0EF545E56224A39CCC0F24FE5F" baseAuthorityName="Smith" baseAuthorityYear="1849" box="[986,1110,439,464]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="capensis">
<emphasis id="B96FEA9FF545E56224A39CCC0F24FE5F" box="[986,1110,439,464]" italics="true" pageId="13" pageNumber="245">L. capensis</emphasis>
</taxonomicName>
complex—one is stated to be in progress by Travers
<emphasis id="B96FEA9FF545E56226EA9CA00AB8FE7A" box="[403,458,475,501]" italics="true" pageId="13" pageNumber="245">et al.</emphasis>
unpublished (in
<bibRefCitation id="EF8A4B7CF545E56225FD9CA00813FE7A" author="Marques, M. P. &amp; Ceriaco, L. M. P. &amp; Buehler, M. D. &amp; Bandeira, S. A. &amp; Janota, J. M. &amp; Bauer, A. M." box="[644,865,475,501]" pageId="13" pageNumber="245" pagination="301 - 352" refId="ref13670" refString="Marques, M. P., Ceriaco, L. M. P., Buehler, M. D., Bandeira, S. A., Janota, J. M. &amp; Bauer, A. M. (2020) A revision of the dwarf geckos, genus Lygodactylus (Squamata: Gekkonidae), from Angola, with description of three new species. Zootaxa, 4853 (3), 301 - 352. https: // doi. org / 10.11646 / zootaxa. 4853.3.1" type="journal article" year="2020">
Marques
<emphasis id="B96FEA9FF545E56225959CA00851FE7A" box="[748,803,475,501]" italics="true" pageId="13" pageNumber="245">et al.</emphasis>
2020
</bibRefCitation>
)—we strongly recommend a broader incorporation of genetic markers to resolve the conflicting topologies since our multigene dataset consists of only few markers especially for
<taxonomicName id="4C1B4D0EF545E562264D9F580ADCFDB2" box="[308,430,547,573]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="nyaneka">
<emphasis id="B96FEA9FF545E562264D9F580ADCFDB2" box="[308,430,547,573]" italics="true" pageId="13" pageNumber="245">L. nyaneka</emphasis>
</taxonomicName>
and for the numerous groups forming
<emphasis id="B96FEA9FF545E56224239F580F3CFDB2" box="[858,1102,547,573]" italics="true" pageId="13" pageNumber="245">
<taxonomicName id="4C1B4D0EF545E56224239F5808A5FDB3" baseAuthorityName="Smith" baseAuthorityYear="1849" box="[858,983,547,572]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="capensis">L. capensis</taxonomicName>
<taxonomicNameLabel id="A25C57E4F545E56224A79F580F3CFDB2" box="[990,1102,547,573]" pageId="13" pageNumber="245" sensu="lato">sensu lato</taxonomicNameLabel>
</emphasis>
. The clade including
<taxonomicName id="4C1B4D0EF545E56222479F580EEEFDB3" authorityName="Sternfeld" authorityYear="1911" box="[1342,1436,547,572]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="grotei">
<emphasis id="B96FEA9FF545E56222479F580EEEFDB3" box="[1342,1436,547,572]" italics="true" pageId="13" pageNumber="245">L. grotei</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1B4D0EF545E56227BE9F3C0A2DFDEE" authorityName="Loveridge" authorityYear="1941" box="[199,351,583,609]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="pakenhami">
<emphasis id="B96FEA9FF545E56227BE9F3C0A2DFDEE" box="[199,351,583,609]" italics="true" pageId="13" pageNumber="245">L. pakenhami</emphasis>
</taxonomicName>
is defined as the sister clade of
<emphasis id="B96FEA9FF545E56225BC9F3C08BFFDEE" box="[709,973,583,609]" italics="true" pageId="13" pageNumber="245">
<taxonomicName id="4C1B4D0EF545E56225BC9F3C0835FDEE" authorityName="Boettger" authorityYear="1913" box="[709,839,583,609]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="insularis">L. insularis</taxonomicName>
,
<taxonomicName id="4C1B4D0EF545E562242A9F3C08BFFDEE" box="[851,973,583,609]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="nyaneka">L. nyaneka</taxonomicName>
</emphasis>
and
<emphasis id="B96FEA9FF545E562237D9F3C0F88FDEE" box="[1028,1274,583,609]" italics="true" pageId="13" pageNumber="245">
<taxonomicName id="4C1B4D0EF545E562237D9F3C0FF3FDEF" baseAuthorityName="Smith" baseAuthorityYear="1849" box="[1028,1153,583,608]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="capensis">L. capensis</taxonomicName>
<taxonomicNameLabel id="A25C57E4F545E56223F09F3C0F88FDEE" box="[1161,1274,583,609]" pageId="13" pageNumber="245" sensu="lato">sensu lato</taxonomicNameLabel>
</emphasis>
.
<taxonomicName id="4C1B4D0EF545E56222719F3C0B8BFD0A" authorityName="Boettger" authorityYear="1913" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="insularis">
<emphasis id="B96FEA9FF545E56222719F3C0B8BFD0A" italics="true" pageId="13" pageNumber="245">Lygodactylus insularis</emphasis>
</taxonomicName>
is recovered as a monophyletic group based on mtDNA, probably resulting from a single dispersal event. Furthermore, the species based on the sampling herein exhibits little intraspecific genetic variability, possibly due to the tiny size of the island resulting in a panmictic population.
</paragraph>
<paragraph id="8BA4368DF545E56227BE9FAC0F64FB4A" blockId="13.[151,1437,151,1581]" pageId="13" pageNumber="245">
Members of the genus
<taxonomicName id="4C1B4D0EF545E56226BC9FAC092BFD7E" box="[453,601,727,753]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="genus">
<emphasis id="B96FEA9FF545E56226BC9FAC092BFD7E" box="[453,601,727,753]" italics="true" pageId="13" pageNumber="245">Lygodactylus</emphasis>
</taxonomicName>
successfully crossed the
<collectingCountry id="F30C761DF545E562240B9FAC0F74FD7E" box="[882,1030,727,753]" name="Mozambique" pageId="13" pageNumber="245">Mozambique</collectingCountry>
Channel from mainland Africa back to
<collectingCountry id="F30C761DF545E56227CE9F800A4CFC9A" box="[183,318,763,789]" name="Madagascar" pageId="13" pageNumber="245">Madagascar</collectingCountry>
one or perhaps two times (R̂ll
<emphasis id="B96FEA9FF545E56225C79F80098BFC9A" box="[702,761,763,789]" italics="true" pageId="13" pageNumber="245">et al.</emphasis>
2010;
<bibRefCitation id="EF8A4B7CF545E56224349F800F16FC9A" author="Mezzasalma, M. &amp; Andreone, F. &amp; Aprea, G. &amp; Glaw, F. &amp; Odierna, G. &amp; Guarino, F. M." box="[845,1124,763,789]" pageId="13" pageNumber="245" pagination="42 - 54" refId="ref13913" refString="Mezzasalma, M., Andreone, F., Aprea, G., Glaw, F., Odierna, G. &amp; Guarino, F. M. (2017) Molecular phylogeny, biogeography and chromosome evolution of Malagasy geckos of the genus Lygodactylus (Squamata, Gekkonidae). Zoological Scripta, 46, 42 - 54. https: // doi. org / 10.1111 / zsc. 12188" type="journal article" year="2017">
Mezzasalma
<emphasis id="B96FEA9FF545E562249A9F800F6DFC9A" box="[995,1055,763,789]" italics="true" pageId="13" pageNumber="245">et al.</emphasis>
2017
</bibRefCitation>
;
<bibRefCitation id="EF8A4B7CF545E562230A9F800E29FC9A" author="Gippner, S. &amp; Travers, S. L. &amp; Scherz, M. D. &amp; Colston, T. J. &amp; Lyra, M. L. &amp; Ashwini, V. M. &amp; Multzsch, M. &amp; Nielson, S. V. &amp; Rancilhac, L. &amp; Glaw, F. &amp; Bauer, A. M. &amp; Vences, M." box="[1139,1371,763,789]" pageId="13" pageNumber="245" pagination="107311" refId="ref12804" refString="Gippner, S., Travers, S. L., Scherz, M. D., Colston, T. J., Lyra, M. L., Ashwini, V. M., Multzsch, M., Nielson, S. V., Rancilhac, L., Glaw, F., Bauer, A. M. &amp; Vences, M. (2021) A comprehensive phylogeny of dwarf geckos of the genus Lygodactylus, with insights into their systematics and morphological variation. Molecular Phylogenetics and Evolution, 165, 107311. https: // doi. org / 10.1016 / j. ympev. 2021.107311" type="journal article" year="2021">
Gippner
<emphasis id="B96FEA9FF545E56223A09F800E67FC9A" box="[1241,1301,763,789]" italics="true" pageId="13" pageNumber="245">et al.</emphasis>
2021
</bibRefCitation>
). The ancestor of
<taxonomicName id="4C1B4D0EF545E56226629E640AE9FCB6" authorityName="Boettger" authorityYear="1913" box="[283,411,799,825]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="insularis">
<emphasis id="B96FEA9FF545E56226629E640AE9FCB6" box="[283,411,799,825]" italics="true" pageId="13" pageNumber="245">L. insularis</emphasis>
</taxonomicName>
probably began at least one further traverse in eastward direction ending on Juan de Nova, an island in the middle of the channel. Juan de Nova developed as the uplifted top of a seamount that probably was formed during the Cretaceous along the Davie Fracture (
<bibRefCitation id="EF8A4B7CF545E56224649E1C0FE5FC0E" author="Riaux-Gobin, C. &amp; Witkowski, A." box="[797,1175,871,897]" pageId="13" pageNumber="245" pagination="43 - 58" refId="ref14515" refString="Riaux-Gobin, C. &amp; Witkowski, A. (2012) Small-sized and discoid species of the genus Cocconeiopsis (Bacillariophyta) on Holothuria atra (Juan de Nova, Mozambique Channel). Phytotaxa, 54, 43 - 58. https: // doi. org / 10.11646 / phytotaxa. 54.1.5" type="journal article" year="2012">Riaux-Gobin &amp; Witkowski 2012</bibRefCitation>
).
<bibRefCitation id="EF8A4B7CF545E56223D39E1C0EEEFC0E" author="Ali, J. R. &amp; Hedges, S. B." box="[1194,1436,871,897]" pageId="13" pageNumber="245" pagination="104103" refId="ref11532" refString="Ali, J. R. &amp; Hedges, S. B. (2022) A review of geological evidence bearing on proposed Cenozoic land connections between Madagascar and Africa and its relevance to biogeography. Earth-Science Reviews, 232, 104103. https: // doi. org / 10.1016 / j. earscirev. 2022.104103" type="journal article" year="2022">Ali &amp; Hedges (2022)</bibRefCitation>
suggest that it may have remained emerged as a low elevation atoll since the Early Palaeocene. In such a scenario, the dispersal of the ancestors of
<taxonomicName id="4C1B4D0EF545E562257A9ED409F1FC46" authorityName="Boettger" authorityYear="1913" box="[515,643,943,969]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="insularis">
<emphasis id="B96FEA9FF545E562257A9ED409F1FC46" box="[515,643,943,969]" italics="true" pageId="13" pageNumber="245">L. insularis</emphasis>
</taxonomicName>
may have taken place as early as 22.3 (confidence interval 15.630.2) mya, the stem age of the
<taxonomicName id="4C1B4D0EF545E56226D69EA8095EFC63" baseAuthorityName="Smith" baseAuthorityYear="1849" box="[431,556,979,1004]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="capensis">
<emphasis id="B96FEA9FF545E56226D69EA8095EFC63" box="[431,556,979,1004]" italics="true" pageId="13" pageNumber="245">L. capensis</emphasis>
</taxonomicName>
group estimated in the
<taxonomicName id="4C1B4D0EF545E562244D9EA808BAFC62" box="[820,968,979,1005]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="genus">
<emphasis id="B96FEA9FF545E562244D9EA808BAFC62" box="[820,968,979,1005]" italics="true" pageId="13" pageNumber="245">Lygodactylus</emphasis>
</taxonomicName>
timetree of
<bibRefCitation id="EF8A4B7CF545E56223299EA80E4DFC62" author="Gippner, S. &amp; Travers, S. L. &amp; Scherz, M. D. &amp; Colston, T. J. &amp; Lyra, M. L. &amp; Ashwini, V. M. &amp; Multzsch, M. &amp; Nielson, S. V. &amp; Rancilhac, L. &amp; Glaw, F. &amp; Bauer, A. M. &amp; Vences, M." box="[1104,1343,979,1005]" pageId="13" pageNumber="245" pagination="107311" refId="ref12804" refString="Gippner, S., Travers, S. L., Scherz, M. D., Colston, T. J., Lyra, M. L., Ashwini, V. M., Multzsch, M., Nielson, S. V., Rancilhac, L., Glaw, F., Bauer, A. M. &amp; Vences, M. (2021) A comprehensive phylogeny of dwarf geckos of the genus Lygodactylus, with insights into their systematics and morphological variation. Molecular Phylogenetics and Evolution, 165, 107311. https: // doi. org / 10.1016 / j. ympev. 2021.107311" type="journal article" year="2021">
Gippner
<emphasis id="B96FEA9FF545E56223CA9EA80F99FC62" box="[1203,1259,979,1005]" italics="true" pageId="13" pageNumber="245">et al.</emphasis>
(2021)
</bibRefCitation>
. Results of palaeogeographic reconstructions and palaeo-oceanographic modelling suggest strong surface currents in the
<collectingCountry id="F30C761DF545E56227EE99600A59FBBA" box="[151,299,1051,1077]" name="Mozambique" pageId="13" pageNumber="245">Mozambique</collectingCountry>
Channel flowing from northeast
<collectingCountry id="F30C761DF545E56225E799600840FBBA" box="[670,818,1051,1077]" name="Mozambique" pageId="13" pageNumber="245">Mozambique</collectingCountry>
and
<collectingCountry id="F30C761DF545E562241E996008BEFBBA" box="[871,972,1051,1077]" name="Tanzania" pageId="13" pageNumber="245">Tanzania</collectingCountry>
eastward towards
<collectingCountry id="F30C761DF545E56223E599600E51FBBA" box="[1180,1315,1051,1077]" name="Madagascar" pageId="13" pageNumber="245">Madagascar</collectingCountry>
during the Palaeogene (66 to 23 mya), which would support this scenario (
<bibRefCitation id="EF8A4B7CF545E562241C99440F4BFBD6" author="Ali, J. R. &amp; Huber, M." box="[869,1081,1087,1113]" pageId="13" pageNumber="245" pagination="653 - 656" refId="ref11490" refString="Ali, J. R. &amp; Huber, M. (2010) Mammalian biodiversity on Madagascar controlled by ocean currents. Nature, 463, 653 - 656. https: // doi. org / 10.1038 / nature 08706" type="journal article" year="2010">Ali &amp; Huber 2010</bibRefCitation>
). While the main contemporary direction of oceanic surface currents is westward, the situation in the
<collectingCountry id="F30C761DF545E56224D599180F32FBF2" box="[940,1088,1123,1149]" name="Mozambique" pageId="13" pageNumber="245">Mozambique</collectingCountry>
Channel is very complex with numerous eddies and possible countercurrents (
<bibRefCitation id="EF8A4B7CF545E56225CD99FC08A6FB2E" author="Hawlitschek, O. &amp; Garrido, S. R. &amp; Glaw, G." box="[692,980,1159,1185]" pageId="13" pageNumber="245" pagination="1426 - 1440" refId="ref12974" refString="Hawlitschek, O., Garrido, S. R. &amp; Glaw, G. (2016 a). How marine currents influenced the widespread natural overseas dispersal of reptiles in the Western Indian Ocean. Journal of Biogeography, 44, 1426 - 1440. https: // doi. org / 10.1111 / jbi. 12940" type="journal article" year="2016">
Hawlitschek
<emphasis id="B96FEA9FF545E562243099FC08F0FB2E" box="[841,898,1159,1185]" italics="true" pageId="13" pageNumber="245">et al.</emphasis>
2016a
</bibRefCitation>
). This also suggests that dispersal could have occurred during a more recent period of mainly westward marine currents.
</paragraph>
<paragraph id="8BA4368DF545E56227BE99B408BAFABE" blockId="13.[151,1437,151,1581]" pageId="13" pageNumber="245">
An alternative hypothesis, dispersal via short-lived Cenozoic land-bridges between mainland Africa and
<collectingCountry id="F30C761DF545E56227EE99880A6CFA82" box="[151,286,1267,1293]" name="Madagascar" pageId="13" pageNumber="245">Madagascar</collectingCountry>
(e.g.,
<bibRefCitation id="EF8A4B7CF545E56226129988094DFA82" author="Mazza, P. P. A. &amp; Buccianti, A. &amp; Savorelli, A." box="[363,575,1267,1293]" pageId="13" pageNumber="245" pagination="1364 - 1380" refId="ref13857" refString="Mazza, P. P. A., Buccianti, A. &amp; Savorelli, A. (2019) Grasping at straws: a re-evaluation of sweepstakes colonisation of islands by mammals. Biological Reviews, 94, 1364 - 1380. https: // doi. org / 10.1111 / brv. 12506" type="journal article" year="2019">
Mazza
<emphasis id="B96FEA9FF545E56226C699880A88FA82" box="[447,506,1267,1293]" italics="true" pageId="13" pageNumber="245">et al.</emphasis>
2019
</bibRefCitation>
;
<bibRefCitation id="EF8A4B7CF545E562253799880843FA82" author="Masters, J. C. &amp; Genin, F. &amp; Zhang, Y. &amp; Pellen, R. &amp; Huck, T. &amp; Mazza, P. P. A. &amp; Rabineau, M. &amp; Aslanian, D." box="[590,817,1267,1293]" pageId="13" pageNumber="245" pagination="492 - 510" refId="ref13764" refString="Masters, J. C., Genin, F., Zhang, Y., Pellen, R., Huck, T., Mazza, P. P. A., Rabineau, M., Doucoure &amp; M., Aslanian, D. (2020) Biogeographic mechanisms involved in the colonization of Madagascar by African vertebrates: Rifting, rafting and runways. Journal of Biogeography, 48, 492 - 510. https: // doi. org / 10.1111 / jbi. 14032" type="journal article" year="2020">
Masters
<emphasis id="B96FEA9FF545E56225C999880996FA82" box="[688,740,1267,1293]" italics="true" pageId="13" pageNumber="245">et al</emphasis>
. 2020
</bibRefCitation>
), is disputed and has been ruled out based on current geological reconstructions (e.g.,
<bibRefCitation id="EF8A4B7CF545E562257A986C09A6FABE" author="Ali, J. R. &amp; Huber, M." box="[515,724,1303,1329]" pageId="13" pageNumber="245" pagination="653 - 656" refId="ref11490" refString="Ali, J. R. &amp; Huber, M. (2010) Mammalian biodiversity on Madagascar controlled by ocean currents. Nature, 463, 653 - 656. https: // doi. org / 10.1038 / nature 08706" type="journal article" year="2010">Ali &amp; Huber 2010</bibRefCitation>
;
<bibRefCitation id="EF8A4B7CF545E56225A4986C08CEFABE" author="Ali, J. R. &amp; Hedges, S. B." box="[733,956,1303,1329]" pageId="13" pageNumber="245" pagination="104103" refId="ref11532" refString="Ali, J. R. &amp; Hedges, S. B. (2022) A review of geological evidence bearing on proposed Cenozoic land connections between Madagascar and Africa and its relevance to biogeography. Earth-Science Reviews, 232, 104103. https: // doi. org / 10.1016 / j. earscirev. 2022.104103" type="journal article" year="2022">Ali &amp; Hedges 2022</bibRefCitation>
).
</paragraph>
<paragraph id="8BA4368DF545E56227BE98400F58F9A2" blockId="13.[151,1437,151,1581]" pageId="13" pageNumber="245">
It also must be emphasized that despite a reasonable sampling of Eastern Africa, especially
<collectingCountry id="F30C761DF545E56223BF98400E28FADA" box="[1222,1370,1339,1365]" name="Mozambique" pageId="13" pageNumber="245">Mozambique</collectingCountry>
(with
<taxonomicName id="4C1B4D0EF545E56227EE98240A6AFAF7" baseAuthorityName="Smith" baseAuthorityYear="1849" box="[151,280,1375,1400]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="capensis">
<emphasis id="B96FEA9FF545E56227EE98240A6AFAF7" box="[151,280,1375,1400]" italics="true" pageId="13" pageNumber="245">L. capensis</emphasis>
</taxonomicName>
group sequences from 14 localities available so far), it is possible that a still undiscovered species representing the sister lineage of
<taxonomicName id="4C1B4D0EF545E562257298F809F9FA12" authorityName="Boettger" authorityYear="1913" box="[523,651,1411,1437]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="insularis">
<emphasis id="B96FEA9FF545E562257298F809F9FA12" box="[523,651,1411,1437]" italics="true" pageId="13" pageNumber="245">L. insularis</emphasis>
</taxonomicName>
may exist in this region. A somewhat similar situation was uncovered in the frogs of the
<taxonomicName id="4C1B4D0EF545E562261098DC09F5FA4F" baseAuthorityName="Duméril &amp; Bibron" baseAuthorityYear="1841" box="[361,647,1447,1473]" class="Amphibia" family="Ptychadenidae" genus="Ptychadena" kingdom="Animalia" order="Anura" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="mascariensis">
<emphasis id="B96FEA9FF545E562261098DC09F5FA4F" box="[361,647,1447,1473]" italics="true" pageId="13" pageNumber="245">Ptychadena mascariensis</emphasis>
</taxonomicName>
complex in which the sister lineage of the Malagasy species has only been recorded from a small area of
<collectingCountry id="F30C761DF545E562254398B009E2FA6A" box="[570,656,1483,1509]" name="Malawi" pageId="13" pageNumber="245">Malawi</collectingCountry>
and could have easily been overlooked without intensive sampling efforts (
<bibRefCitation id="EF8A4B7CF545E562278998940ABBF986" author="Zimkus, B. M. &amp; Lawson, L. P. &amp; Barej, M. F. &amp; Barratt, C. D. &amp; Channing, A. &amp; Dash, K. M. &amp; Dehling, J. M. &amp; Du Preez, L. &amp; Gehring, P. - S. &amp; Greenbaum, E. &amp; Gvozdik, V. &amp; Harvey, J. &amp; Kielgast, J. &amp; Kusamba, C. &amp; Nagy, Z. T. &amp; Pabijan, M. &amp; Penner, J. &amp; Vences, M." box="[240,457,1519,1545]" pageId="13" pageNumber="245" pagination="254 - 269" refId="ref15631" refString="Zimkus, B. M., Lawson, L. P., Barej, M. F., Barratt, C. D., Channing, A., Dash, K. M., Dehling, J. M., Du Preez, L., Gehring, P. - S., Greenbaum, E., Gvozdik, V., Harvey, J., Kielgast, J., Kusamba, C., Nagy, Z. T., Pabijan, M., Penner, J., R ˆ del, M. - O., Vences, M. &amp; L ˆ tters, S. (2017) Leapfrogging into new territory: How Mascarene ridged frogs diversified across Africa and Madagascar to maintain their ecological niche. Molecular Phylogenetics and Evolution, 106, 254 - 269. https: // doi. org / 10.1016 / j. ympev. 2016.09.018" type="journal article" year="2017">
Zimkus
<emphasis id="B96FEA9FF545E562263498940A0CF986" box="[333,382,1519,1545]" italics="true" pageId="13" pageNumber="245">et al</emphasis>
. 2017
</bibRefCitation>
). If a closer relative of
<taxonomicName id="4C1B4D0EF545E56225B49894083EF986" authorityName="Boettger" authorityYear="1913" box="[717,844,1519,1545]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="245" phylum="Chordata" rank="species" species="insularis">
<emphasis id="B96FEA9FF545E56225B49894083EF986" box="[717,844,1519,1545]" italics="true" pageId="13" pageNumber="245">L. insularis</emphasis>
</taxonomicName>
was found on the African mainland, it would lead to younger age estimates of the colonization event of Juan de Nova by these geckos.
</paragraph>
</subSubSection>
</treatment>
</document>