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<mods:title id="72D703D63FF0AB3C6BCB7B16CF3D0260">Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascars rainforest</mods:title>
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<mods:namePart id="6847023CB07DBCF1D5D51E09550C6E0D">Crottini, Angelica</mods:namePart>
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<emphasis id="35D6531A1307BF16FF28FBE4FEE5464F" bold="true" box="[151,374,1122,1149]" italics="true" pageId="45" pageNumber="46">Lygodactylus fritzi</emphasis>
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<emphasis id="35D6531A1307BF16FF28FB21FE8E468C" box="[151,285,1191,1215]" italics="true" pageId="45" pageNumber="46">Lygodactylus</emphasis>
sp.
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11:
<bibRefCitation id="6333F2F91307BF16FED2FB21FDD5468C" author="Gippner, S. &amp; Travers S. L. &amp; Scherz M. D. &amp; Colston T. J. &amp; Lyra M. L. &amp; Mohan A. V. &amp; Multzsch M. &amp; Nielsen S. V. &amp; Rancilhac L. &amp; Glaw F. &amp; Bauer A. M. &amp; Vences M." box="[365,582,1191,1215]" pageId="45" pageNumber="46" pagination="107311" refId="ref37826" refString="Gippner, S., Travers S. L., Scherz M. D., Colston T. J., Lyra M. L., Mohan A. V., Multzsch M., Nielsen 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="35D6531A1307BF16FE79FB2EFE6A468C" box="[454,505,1191,1215]" italics="true" pageId="45" pageNumber="46">et al.</emphasis>
(2021)
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.
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<emphasis id="35D6531A1307BF16FF28FB6BFEB24734" box="[151,289,1261,1287]" italics="true" pageId="45" pageNumber="46">Justification</emphasis>
. This new species from coastal areas in Madagascars Northern Central East has previously been called
<emphasis id="35D6531A1307BF16FF28FA94FF344718" box="[151,167,1298,1323]" italics="true" pageId="45" pageNumber="46">L</emphasis>
. sp.
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<bibRefCitation id="6333F2F91307BF16FE9BFA97FD894718" author="Gippner, S. &amp; Travers S. L. &amp; Scherz M. D. &amp; Colston T. J. &amp; Lyra M. L. &amp; Mohan A. V. &amp; Multzsch M. &amp; Nielsen S. V. &amp; Rancilhac L. &amp; Glaw F. &amp; Bauer A. M. &amp; Vences M." box="[292,538,1297,1323]" pageId="45" pageNumber="46" pagination="107311" refId="ref37826" refString="Gippner, S., Travers S. L., Scherz M. D., Colston T. J., Lyra M. L., Mohan A. V., Multzsch M., Nielsen 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="35D6531A1307BF16FE36FA94FE504718" box="[393,451,1297,1323]" italics="true" pageId="45" pageNumber="46">et al.</emphasis>
(2021)
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. It corresponds to a lineage forming part of subclade A5 of
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<emphasis id="35D6531A1307BF16FB6EFA97FAF74718" box="[1233,1380,1297,1323]" italics="true" pageId="45" pageNumber="46">Domerguella</emphasis>
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, and occurs in a general area where
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<emphasis id="35D6531A1307BF16FE4AFAB0FDC5477C" box="[501,598,1334,1359]" italics="true" pageId="45" pageNumber="46">L. miops</emphasis>
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is also present. It is characterized by the presence of distinct lateral spinelike scales at the base of the tail in males, as are found in several representatives of subclade A5 but not in other
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<emphasis id="35D6531A1307BF16FF28FAFBFEB947A4" box="[151,298,1405,1431]" italics="true" pageId="45" pageNumber="46">Domerguella</emphasis>
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. The smallest genetic distance is 9.0% to specimens assigned to
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<emphasis id="35D6531A1307BF16FBABFAF8FBE447A4" box="[1044,1143,1406,1431]" italics="true" pageId="45" pageNumber="46">L. miops</emphasis>
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. However, it differs from this species by phylogenetic position and lack of haplotype sharing in nuclear-encoded genes. While species status of these two lineages is validated by the molecular evidence, the assignment of the
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of
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, and of the types of the other earlier names
<taxonomicName id="C0A2F48B1307BF16FDB6FA6FFD714430" authorityName="Angel" authorityYear="1942" box="[521,738,1513,1539]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="septemtuberculatus">
<emphasis id="35D6531A1307BF16FDB6FA6FFD714430" box="[521,738,1513,1539]" italics="true" pageId="45" pageNumber="46">septemtuberculatus</emphasis>
</taxonomicName>
and
<taxonomicName id="C0A2F48B1307BF16FCA3FA6FFC144430" authorityName="Boettger" authorityYear="1913" box="[796,903,1513,1539]" class="Reptilia" family="Gekkonidae" genus="Microscalabotes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="spinulifer">
<emphasis id="35D6531A1307BF16FCA3FA6FFC144430" box="[796,903,1513,1539]" italics="true" pageId="45" pageNumber="46">spinulifer</emphasis>
</taxonomicName>
, to either of them requires further justification (see also the account of
<taxonomicName id="C0A2F48B1307BF16FE1AF988FD954414" authorityName="Gunther" authorityYear="1891" box="[421,518,1550,1575]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="miops">
<emphasis id="35D6531A1307BF16FE1AF988FD954414" box="[421,518,1550,1575]" italics="true" pageId="45" pageNumber="46">L. miops</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="4FB8DC831307BF15FF78F9B7FEEA443A" lastPageId="46" lastPageNumber="47" pageId="45" pageNumber="46" type="description">
<paragraph id="071D8F081307BF16FF78F9B7FB4B445C" blockId="45.[151,1437,1261,1648]" pageId="45" pageNumber="46">
The following arguments support our decision to assign the
<typeStatus id="D81931AA1307BF16FCD3F9B7FC5D4478" box="[876,974,1585,1611]" pageId="45" pageNumber="46" type="holotype">holotype</typeStatus>
of
<emphasis id="35D6531A1307BF16FC4AF9B4FA8C4478" box="[1013,1311,1585,1611]" italics="true" pageId="45" pageNumber="46">
<taxonomicName id="C0A2F48B1307BF16FC4AF9B4FBA84478" authorityName="Gunther" authorityYear="1891" box="[1013,1083,1586,1611]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="miops">miops</taxonomicName>
,
<taxonomicName id="C0A2F48B1307BF16FBF9F9B7FA8C4478" authorityName="Angel" authorityYear="1942" box="[1094,1311,1585,1611]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="septemtuberculatus">septemtuberculatus</taxonomicName>
</emphasis>
and
<taxonomicName id="C0A2F48B1307BF16FAE9F9B4FF55445C" authorityName="Boettger" authorityYear="1913" class="Reptilia" family="Gekkonidae" genus="Microscalabotes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="spinulifer">
<emphasis id="35D6531A1307BF16FAE9F9B4FF55445C" italics="true" pageId="45" pageNumber="46">spinulifer</emphasis>
</taxonomicName>
to what we will in this section call the “widespread lowland lineage” rather than to
<emphasis id="35D6531A1307BF16FBCEF9D0FB1B445C" box="[1137,1160,1622,1647]" italics="true" pageId="45" pageNumber="46">L.</emphasis>
sp. 11:
</paragraph>
<paragraph id="071D8F081307BF16FF28F91BFAD74574" blockId="45.[151,1437,1693,2008]" pageId="45" pageNumber="46">
(1) The specimens of the widespread lowland lineage (including the three historical
<typeStatus id="D81931AA1307BF16FBFFF91BFBEE4484" box="[1088,1149,1693,1719]" pageId="45" pageNumber="46">types</typeStatus>
) differ from
<emphasis id="35D6531A1307BF16FAB4F918FAB14484" box="[1291,1314,1694,1719]" italics="true" pageId="45" pageNumber="46">L.</emphasis>
sp. 11 by a 36 vs. 2 dorsolateral tubercles between limbs; dorsal scale count 207242 vs. 247; ventral scale count 102113 vs. 98; lateral tubercles at tail base recognizable (also in females) vs. barely recognizable. While each of these meristic differences by itself is rather weak, taken together they characterize
<emphasis id="35D6531A1307BF16FB95F88CFBD24510" box="[1066,1089,1802,1827]" italics="true" pageId="45" pageNumber="46">L.</emphasis>
sp. 11 as morphologically distinct, and the
<specimenCount id="11A444811307BF16FEE1F8ABFE5E457B" box="[350,461,1837,1864]" pageId="45" pageNumber="46" type="generic" typeStatus="type">three type</specimenCount>
specimens of the earlier names as better fitting the widespread eastern lineage.
</paragraph>
<paragraph id="071D8F081307BF15FF28F8D7FB9142E6" blockId="45.[151,1437,1693,2008]" lastBlockId="46.[151,1437,151,1545]" lastPageId="46" lastPageNumber="47" pageId="45" pageNumber="46">
(2)
<emphasis id="35D6531A1307BF16FF79F8D4FF4E4558" box="[198,221,1874,1899]" italics="true" pageId="45" pageNumber="46">L.</emphasis>
sp. 11 is only known from two low-elevation localities near sea level (
<quantity id="C05A22ED1307BF16FBBEF8D7FBCB455F" box="[1025,1112,1873,1900]" metricMagnitude="1" metricUnit="m" metricValue="1.0" metricValueMax="2.0" metricValueMin="0.0" pageId="45" pageNumber="46" unit="m" value="10.0" valueMax="20.0" valueMin="0.0">020 m</quantity>
a.s.l.), and seems to be relatively localized; for instance, it has not been found at other nearby localities at slightly higher elevation such as Sahafina or Betampona, despite a substantial number of specimens sequenced from Betampona, which all belonged to the widespread lowland lineage. It is therefore less likely that historical specimens of
<emphasis id="35D6531A1307BF16FABDF838FA8A45E4" box="[1282,1305,1982,2007]" italics="true" pageId="45" pageNumber="46">L.</emphasis>
sp. 11 were collected historically, and that its range extends into the
<typeStatus id="D81931AA1304BF15FC89FF11FCF54282" box="[822,870,151,177]" pageId="46" pageNumber="47">type</typeStatus>
localities of the three historical nomina, especially up to Moramanga (the
<typeStatus id="D81931AA1304BF15FE78FF3DFE6442E6" box="[455,503,187,213]" pageId="46" pageNumber="47">type</typeStatus>
locality of
<taxonomicName id="C0A2F48B1304BF15FDC8FF3DFCC342E6" authorityName="Angel" authorityYear="1942" box="[631,848,187,213]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="septemtuberculatus">
<emphasis id="35D6531A1304BF15FDC8FF3DFCC342E6" box="[631,848,187,213]" italics="true" pageId="46" pageNumber="47">septemtuberculatus</emphasis>
</taxonomicName>
and
<taxonomicName id="C0A2F48B1304BF15FC39FF3DFC6242E6" authorityName="Boettger" authorityYear="1913" box="[902,1009,187,213]" class="Reptilia" family="Gekkonidae" genus="Microscalabotes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="spinulifer">
<emphasis id="35D6531A1304BF15FC39FF3DFC6242E6" box="[902,1009,187,213]" italics="true" pageId="46" pageNumber="47">spinulifer</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="071D8F081304BF15FF28FF59FB144196" blockId="46.[151,1437,151,1545]" pageId="46" pageNumber="47">
(3) The
<typeStatus id="D81931AA1304BF15FF46FF59FEBA42CA" box="[249,297,223,249]" pageId="46" pageNumber="47">type</typeStatus>
locality of
<taxonomicName id="C0A2F48B1304BF15FE12FF59FD9C42CB" authorityName="Gunther" authorityYear="1891" box="[429,527,223,248]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="miops">
<emphasis id="35D6531A1304BF15FE12FF59FD9C42CB" box="[429,527,223,248]" italics="true" pageId="46" pageNumber="47">L. miops</emphasis>
</taxonomicName>
, “Senbendrana” according to the original description (
<bibRefCitation id="6333F2F91304BF15FBC6FF59FA8942CA" author="Gunther, A." box="[1145,1306,223,249]" pageId="46" pageNumber="47" pagination="287 - 288" refId="ref38543" refString="Gunther, A. (1891) Eleventh Contribution to the knowledge of the Fauna of Madagascar. Annals and Magazine of Natural History, 8, 287 - 288. https: // doi. org / 10.1080 / 00222939109460436" type="journal article" year="1891">Günther 1891</bibRefCitation>
), cannot be located reliably at present. Senbendrana has been reported with the addition “near Tamatave” (=
<collectingRegion id="C56641EA1304BF15FB43FE85FAE4432E" box="[1276,1399,259,285]" country="Madagascar" name="Toamasina" pageId="46" pageNumber="47">Toamasina</collectingRegion>
) as a collecting locality of spiders (
<bibRefCitation id="6333F2F91304BF15FD99FEA1FD2E4372" author="Pocock, R. L." box="[550,701,294,321]" pageId="46" pageNumber="47" pagination="187 - 197" refId="ref40075" refString="Pocock, R. L. (1895) XXVIII. - Descriptions of new genera and species of Trap-door Spiders belonging to the Group Trionychi. Annals and Magazine of Natural History, Series, 6, 16, 187 - 197. https: // doi. org / 10.1080 / 00222939508680252" type="journal article" year="1895">Pocock 1895</bibRefCitation>
); or as corresponding to Sahembendrana or Sahambendrana. This latter synonymy is supported by the fact that the
<typeStatus id="D81931AA1304BF15FD5BFECDFC874356" box="[740,788,331,357]" pageId="46" pageNumber="47">type</typeStatus>
of
<taxonomicName id="C0A2F48B1304BF15FC87FECDFC0B4357" authorityName="Gunther" authorityYear="1891" box="[824,920,331,356]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="miops">
<emphasis id="35D6531A1304BF15FC87FECDFC0B4357" box="[824,920,331,356]" italics="true" pageId="46" pageNumber="47">L. miops</emphasis>
</taxonomicName>
was provided by “Majastre” (see
<bibRefCitation id="6333F2F91304BF15FAADFECDFE9143BA" author="Puente, M. &amp; Glaw, F. &amp; Vieites, D. R. &amp; Vences, M." pageId="46" pageNumber="47" pagination="1 - 76" refId="ref40406" refString="Puente, M., Glaw, F., Vieites, D. R. &amp; Vences, M. (2009) Review of the systematics, morphology and distribution of Malagasy dwarf geckos, genera Lygodactylus and Microscalabotes (Squamata: Gekkonidae). Zootaxa, 2103, 1 - 76. https: // doi. org / 10.11646 / zootaxa. 2103.1.1" type="journal article" year="2009">
Puente
<emphasis id="35D6531A1304BF15FADAFECDFA0E4356" box="[1381,1437,331,357]" italics="true" pageId="46" pageNumber="47">et al.</emphasis>
2009
</bibRefCitation>
), probably corresponding to A. Majastre, a collector who provided specimens of many animals and plants from this area. Some of Majastres collections are labelled “Sahambendrana”, e.g., the
<typeStatus id="D81931AA1304BF15FB2EFE15FB52439E" box="[1169,1217,403,429]" pageId="46" pageNumber="47">type</typeStatus>
of the orchid
<taxonomicName id="C0A2F48B1304BF15FAE1FE15FE2743E2" class="Liliopsida" family="Orchidaceae" genus="Eulophia" kingdom="Plantae" order="Asparagales" pageId="46" pageNumber="47" phylum="Tracheophyta" rank="species" species="grandibracteata">
<emphasis id="35D6531A1304BF15FAE1FE15FE2743E2" italics="true" pageId="46" pageNumber="47">Eulophia grandibracteata</emphasis>
</taxonomicName>
(see
<bibRefCitation id="6333F2F91304BF15FE53FE31FD1043E2" author="Schultz, M." box="[492,643,439,465]" pageId="46" pageNumber="47" pagination="5 - 251" refId="ref42039" refString="Schultz, M. (2013) The orchid types of the Herbarium Hamburgense (HBG). Verhandlungen des Naturwissenschaftlichen Vereins Hamburg NF, 47, 5 - 251." type="journal article" year="2013">Schultz 2013</bibRefCitation>
). The locality apparently was often misspelled; for example, we assume that “Sen Bendrana” (
<bibRefCitation id="6333F2F91304BF15FE7CFE5DFD1843C6" author="Michaelsen, W." box="[451,651,475,501]" pageId="46" pageNumber="47" pagination="1 - 72" refId="ref39397" refString="Michaelsen, W. (1891) Beschreibung der von Herrn Dr. Fr. Stuhlmann auf Sansibar und dem gegenuberliegenden Festlande gesammelten Terricolen. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 9, 1 - 72." type="journal article" year="1891">Michaelsen 1891</bibRefCitation>
), “Senbendra” (Sharp &amp; Ogilvie-Grant 1898), or “Schambendrama” (Bott 1963) refer to the same site as well.
<bibRefCitation id="6333F2F91304BF15FD1BFE79FBD4402A" author="Blommers-Schlosser, R. M. A. &amp; Blanc, C. P." box="[676,1095,511,537]" pageId="46" pageNumber="47" pagination="1 - 379" refId="ref36135" refString="Blommers-Schlosser, R. M. A. &amp; Blanc, C. P. (1991) Amphibiens (premiere partie). Faune de Madagascar, 75 (1), 1 - 379." type="journal article" year="1991">Blommers-Schlösser &amp; Blanc (1991)</bibRefCitation>
located Sahembendrana close to Akkoraka (at higher elevations in eastern
<collectingCountry id="7FB5CF981304BF15FD0FFDA5FCAF400E" box="[688,828,547,573]" name="Madagascar" pageId="46" pageNumber="47">Madagascar</collectingCountry>
), but it is likely that Majastres collecting site was situated closer to
<collectingRegion id="C56641EA1304BF15FEDEFDC1FE4D4052" box="[353,478,583,609]" country="Madagascar" name="Toamasina" pageId="46" pageNumber="47">Toamasina</collectingRegion>
. We could not locate current or historical maps mentioning a site with this or a similar name, but
<bibRefCitation id="6333F2F91304BF15FE84FDEDFDB240B6" author="Ramananjara, H. C." box="[315,545,619,645]" pageId="46" pageNumber="47" refId="ref40696" refString="Ramananjara, H. C. (2009) Etude des sources archivistiques et documentaires sur les terres dans la Basse Vallee de l'Ivoloina (Toamasina) de 1897 a 1939: Essai documentaire / Memoire de maitrise. Antananarivo: Universite d'Antananarivo / F. L. S. H / Departement d'Histoire / Formation en Archivistique- Documentation-Museologie, 255 pp." type="book" year="2009">Ramananjara (2009)</bibRefCitation>
documents the sale of a property in 1931, in the “Canton dAntetezambaro; sur la rivière Sahambendrana”, and more specifically “au sud dAmbodisatrana”, which likely refers to a coastal village about
<quantity id="C05A22ED1304BF15FEE3FD35FE3140FE" box="[348,418,691,717]" metricMagnitude="4" metricUnit="m" metricValue="3.0" pageId="46" pageNumber="47" unit="km" value="30.0">30 km</quantity>
north of
<collectingRegion id="C56641EA1304BF15FDBBFD35FDEE40FE" box="[516,637,691,717]" country="Madagascar" name="Toamasina" pageId="46" pageNumber="47">Toamasina</collectingRegion>
that can be located in historical maps from 1934 (Service Géographique de
<collectingCountry id="7FB5CF981304BF15FF54FD51FEE740C2" box="[235,372,727,753]" name="Madagascar" pageId="46" pageNumber="47">Madagascar</collectingCountry>
, map “Fénérive”, 1/500,000). On the other hand,
<bibRefCitation id="6333F2F91304BF15FC07FD51FB1940C2" author="Rosa, G. M. &amp; Andreone, F. &amp; Crottini, A. &amp; Hauswald, J. S. &amp; Noel, J. &amp; Rabibisoa, N. H. &amp; Randriambahiniarime, M. O. &amp; Rebelo, R. &amp; Raxworthy, C. J." box="[952,1162,727,753]" pageId="46" pageNumber="47" pagination="1531 - 1559" refId="ref41679" refString="Rosa, G. M., Andreone, F., Crottini, A., Hauswald, J. S., Noel, J., Rabibisoa, N. H., Randriambahiniarime, M. O., Rebelo, R. &amp; Raxworthy, C. J. (2012) The amphibians of the relict Betampona low-elevation rainforest, eastern Madagascar: an application of the integrative taxonomy approach to biodiversity assessments. Biodiversity and Conservation, 21, 1531 - 1559. https: // doi. org / 10.1007 / s 10531 - 012 - 0262 - x" type="journal article" year="2012">
Rosa
<emphasis id="35D6531A1304BF15FC45FD51FBA540C2" box="[1018,1078,727,753]" italics="true" pageId="46" pageNumber="47">et al.</emphasis>
(2012)
</bibRefCitation>
report a campsite from Betampona Reserve (about
<quantity id="C05A22ED1304BF15FDBFFD7CFDDA4126" box="[512,585,762,789]" metricMagnitude="4" metricUnit="m" metricValue="3.5" pageId="46" pageNumber="47" unit="km" value="35.0">35 km</quantity>
north-east of
<collectingRegion id="C56641EA1304BF15FD56FD7DFCF74126" box="[745,868,763,789]" country="Madagascar" name="Toamasina" pageId="46" pageNumber="47">Toamasina</collectingRegion>
) locally known as Sahambendrana, at coordinates
<geoCoordinate id="6296E9CF1304BF15FF79FC99FEA7410A" box="[198,308,799,825]" degrees="17.8984" direction="south" orientation="latitude" pageId="46" pageNumber="47" precision="5" value="-17.8984">S17.8984</geoCoordinate>
,
<geoCoordinate id="6296E9CF1304BF15FE81FC98FE3E410A" box="[318,429,798,825]" degrees="49.2154" direction="east" orientation="longitude" pageId="46" pageNumber="47" precision="5" value="49.2154">E49.2154</geoCoordinate>
,
<quantity id="C05A22ED1304BF15FE08FC98FE6E410A" box="[439,509,798,825]" metricMagnitude="2" metricUnit="m" metricValue="4.58" pageId="46" pageNumber="47" unit="m" value="458.0">458 m</quantity>
a.s.l. Other sources refer to a Sahambendrana river on the northern versant of Betampona (
<bibRefCitation id="6333F2F91304BF15FEA0FCC5FE78416E" author="Randriatavy, L. D." box="[287,491,835,861]" pageId="46" pageNumber="47" refId="ref40819" refString="Randriatavy, L. D. (2003) Etude de regeneration forestiere et de quelques especies vegetales utilisees a la fois par Varecia variegata et par l'homme dans la Reserve Naturelle Integrale No 1 de Betampona. DEA dissertation, Universite d'Antananarivo, Departement de Biologie et Ecologie Vegetales. Antananarivo, 114 pp." type="book" year="2003">Randriatavy 2003</bibRefCitation>
;
<bibRefCitation id="6333F2F91304BF15FE49FCC5FC87416E" author="Randrianarimanana, H. L. L." box="[502,788,835,861]" pageId="46" pageNumber="47" refId="ref40759" refString="Randrianarimanana, H. L. L. (2009) Etude comparative de l'alimentation et du comportement des deux especes sympatriques d'indriides: Propithecus diadema et Indri indri dans la Reserve Naturelle Integrale N 01 de Betampona (Toamasina). DEA dissertation, Universite d'Antananarivo, Departement de Paleontologie et d'Anthropologie Biologique. Antananarivo, 88 pp." type="book" year="2009">Randrianarimanana 2009</bibRefCitation>
). Whether any of these sites corresponds to Majastres collecting locality cannot be decided without further evidence, but these references demonstrate that the toponym has been and is in use for sites to the north and northeast in the vicinity of
<collectingRegion id="C56641EA1304BF15FBB9FC0DFB104196" box="[1030,1155,907,933]" country="Madagascar" name="Toamasina" pageId="46" pageNumber="47">Toamasina</collectingRegion>
.
</paragraph>
<paragraph id="071D8F081304BF15FF79FC29FAFD47AE" blockId="46.[151,1437,151,1545]" pageId="46" pageNumber="47">
The available evidence, however, points to the original collecting site being not directly at sea level. According to
<bibRefCitation id="6333F2F91304BF15FEAEFC55FE5141DE" author="Gunther, A." box="[273,450,979,1005]" pageId="46" pageNumber="47" pagination="287 - 288" refId="ref38543" refString="Gunther, A. (1891) Eleventh Contribution to the knowledge of the Fauna of Madagascar. Annals and Magazine of Natural History, 8, 287 - 288. https: // doi. org / 10.1080 / 00222939109460436" type="journal article" year="1891">Günther (1891)</bibRefCitation>
, the same collection that included the
<taxonomicName id="C0A2F48B1304BF15FCC5FC55FC4F41DF" authorityName="Gunther" authorityYear="1891" box="[890,988,979,1004]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="miops">
<emphasis id="35D6531A1304BF15FCC5FC55FC4F41DF" box="[890,988,979,1004]" italics="true" pageId="46" pageNumber="47">L. miops</emphasis>
</taxonomicName>
<typeStatus id="D81931AA1304BF15FC5AFC55FBD441DE" box="[997,1095,979,1005]" pageId="46" pageNumber="47" type="holotype">holotype</typeStatus>
also contained “
<taxonomicName id="C0A2F48B1304BF15FAB8FC55FE9A4622" authorityName="Boulenger" authorityYear="1882" class="Amphibia" family="Rhacophoridae" genus="Rhacophorus" kingdom="Animalia" order="Anura" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="luteus">
<emphasis id="35D6531A1304BF15FAB8FC55FE9A4622" italics="true" pageId="46" pageNumber="47">Rhacophorus luteus</emphasis>
</taxonomicName>
”, which almost certainly corresponds to a treefrog species of the
<taxonomicName id="C0A2F48B1304BF15FBB7FC71FB224622" baseAuthorityName="Boulenger" baseAuthorityYear="1882" box="[1032,1201,1015,1041]" class="Amphibia" family="Mantellidae" genus="Boophis" kingdom="Animalia" order="Anura" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="luteus">
<emphasis id="35D6531A1304BF15FBB7FC71FB224622" box="[1032,1201,1015,1041]" italics="true" pageId="46" pageNumber="47">Boophis luteus</emphasis>
</taxonomicName>
group, which is not known from coastal sites in
<collectingCountry id="7FB5CF981304BF15FE41FB9DFD164606" box="[510,645,1051,1077]" name="Madagascar" pageId="46" pageNumber="47">Madagascar</collectingCountry>
(but known to be present in Betampona), and for instance has not been collected at Vohibola or Ankaninny Nofy (
<bibRefCitation id="6333F2F91304BF15FD16FBB9FC14466A" author="Gehring, P. S. &amp; Ratsoavina, F. M. &amp; Vences, M." box="[681,903,1087,1113]" pageId="46" pageNumber="47" pagination="214 - 234" refId="ref37622" refString="Gehring, P. S., Ratsoavina, F. M. &amp; Vences, M. (2010) Filling the gaps - amphibian and reptile records from lowland rainforests in eastern Madagascar. Salamandra, 46, 214 - 234." type="journal article" year="2010">
Gehring
<emphasis id="35D6531A1304BF15FCB1FBB9FCD5466A" box="[782,838,1087,1113]" italics="true" pageId="46" pageNumber="47">et al.</emphasis>
2010
</bibRefCitation>
) where
<emphasis id="35D6531A1304BF15FC5FFBB9FC64466B" box="[992,1015,1087,1112]" italics="true" pageId="46" pageNumber="47">L.</emphasis>
sp. 11 occurs. Furthermore,
<bibRefCitation id="6333F2F91304BF15FA88FBB9FE83464E" author="Pellegrin, J." pageId="46" pageNumber="47" pagination="1 - 222" refId="ref40036" refString="Pellegrin, J. (1933) Les poissons des eaux douces de Madagascar et des iles voisines (Comores, Seychelles, Mascareignes). Memoires de L'Academie Malgache, Tananarive, 14, 1 - 222." type="journal article" year="1933">Pellegrin (1933)</bibRefCitation>
reported fish specimens of the genus
<taxonomicName id="C0A2F48B1304BF15FD0EFBE5FCA7464E" authorityName="T.N.Gill" authorityYear="1860" box="[689,820,1123,1149]" class="Actinopterygii" family="Gobiidae" genus="Sicyopterus" kingdom="Animalia" order="Perciformes" pageId="46" pageNumber="47" phylum="Chordata" rank="genus">
<emphasis id="35D6531A1304BF15FD0EFBE5FCA7464E" box="[689,820,1123,1149]" italics="true" pageId="46" pageNumber="47">Sicyopterus</emphasis>
</taxonomicName>
from a “rivière Sahembendrana (région de Tamatave)”. These specimens were identified by
<bibRefCitation id="6333F2F91304BF15FDDAFB01FCE94692" author="Sparks, J. S. &amp; Nelson, D. W." box="[613,890,1159,1185]" pageId="46" pageNumber="47" pagination="1 - 20" refId="ref42107" refString="Sparks, J. S. &amp; Nelson, D. W. (2004) Review of the Malagasy sicydiine gobies (Teleostei: Gobiidae), with description of a new species and comments on the taxonomic status of Gobius lagocephalus Pallas, 1770. American Museum Novitates, 3440, 1 - 20. https: // doi. org / 10.1206 / 0003 - 0082 (2004) 440 % 3 C 0001: ROTMSG % 3 E 2.0. CO; 2" type="journal article" year="2004">Sparks &amp; Nelson (2004)</bibRefCitation>
as
<taxonomicName id="C0A2F48B1304BF15FC1DFB01FB8D4692" baseAuthorityName="Pellegrin" baseAuthorityYear="1935" box="[930,1054,1159,1185]" class="Actinopterygii" family="Gobiidae" genus="Sicyopterus" kingdom="Animalia" order="Perciformes" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="franouxi">
<emphasis id="35D6531A1304BF15FC1DFB01FB8D4692" box="[930,1054,1159,1185]" italics="true" pageId="46" pageNumber="47">S. franouxi</emphasis>
</taxonomicName>
, a species that according to these authors inhabits clear, swift-flowing waters and is frequently captured quite far inland, again in agreement that this site is within the range of the widespread eastern lineage but not a coastal locality within the range of
<emphasis id="35D6531A1304BF15FAE0FB49FAE546DB" box="[1375,1398,1231,1256]" italics="true" pageId="46" pageNumber="47">L.</emphasis>
sp. 11. The fact that Sahambendrana / Sahembendrana has on various occasions been used to refer to a river (e.g., Pellegrin 1993;
<bibRefCitation id="6333F2F91304BF15FEC4FA91FDD44702" author="Randriatavy, L. D." box="[379,583,1303,1329]" pageId="46" pageNumber="47" refId="ref40819" refString="Randriatavy, L. D. (2003) Etude de regeneration forestiere et de quelques especies vegetales utilisees a la fois par Varecia variegata et par l'homme dans la Reserve Naturelle Integrale No 1 de Betampona. DEA dissertation, Universite d'Antananarivo, Departement de Biologie et Ecologie Vegetales. Antananarivo, 114 pp." type="book" year="2003">Randriatavy 2003</bibRefCitation>
;
<bibRefCitation id="6333F2F91304BF15FDEBFA91FCE14702" author="Randrianarimanana, H. L. L." box="[596,882,1303,1329]" pageId="46" pageNumber="47" refId="ref40759" refString="Randrianarimanana, H. L. L. (2009) Etude comparative de l'alimentation et du comportement des deux especes sympatriques d'indriides: Propithecus diadema et Indri indri dans la Reserve Naturelle Integrale N 01 de Betampona (Toamasina). DEA dissertation, Universite d'Antananarivo, Departement de Paleontologie et d'Anthropologie Biologique. Antananarivo, 88 pp." type="book" year="2009">Randrianarimanana 2009</bibRefCitation>
;
<bibRefCitation id="6333F2F91304BF15FCC0FA91FBC64702" author="Ramananjara, H. C." box="[895,1109,1303,1329]" pageId="46" pageNumber="47" refId="ref40696" refString="Ramananjara, H. C. (2009) Etude des sources archivistiques et documentaires sur les terres dans la Basse Vallee de l'Ivoloina (Toamasina) de 1897 a 1939: Essai documentaire / Memoire de maitrise. Antananarivo: Universite d'Antananarivo / F. L. S. H / Departement d'Histoire / Formation en Archivistique- Documentation-Museologie, 255 pp." type="book" year="2009">Ramananjara 2009</bibRefCitation>
) allows for the possibility of an upstream collecting site of the
<taxonomicName id="C0A2F48B1304BF15FDF7FABDFD394767" authorityName="Gunther" authorityYear="1891" box="[584,682,1339,1364]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="miops">
<emphasis id="35D6531A1304BF15FDF7FABDFD394767" box="[584,682,1339,1364]" italics="true" pageId="46" pageNumber="47">L. miops</emphasis>
</taxonomicName>
<typeStatus id="D81931AA1304BF15FD0CFABDFC8B4766" box="[691,792,1339,1365]" pageId="46" pageNumber="47" type="holotype">holotype</typeStatus>
, at some distance from the coast, and thus at a moderate elevation as indicated by the accompanying fish and amphibian fauna; and probably in the area close to Betampona where our collections have only yielded individuals of the widespread lowland
<taxonomicName id="C0A2F48B1304BF15FBC3FA05FA9C47AE" authorityName="Pasteur" authorityYear="1964" box="[1148,1295,1411,1437]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="55" phylum="Chordata" rank="subGenus" subGenus="Domerguella">
<emphasis id="35D6531A1304BF15FBC3FA05FA9C47AE" box="[1148,1295,1411,1437]" italics="true" pageId="46" pageNumber="47">Domerguella</emphasis>
</taxonomicName>
lineage.
</paragraph>
<paragraph id="071D8F081304BF15FF79FA21FEEA443A" blockId="46.[151,1437,151,1545]" pageId="46" pageNumber="47">
In summary, the available evidence thus suggests that none of the earlier available names
<emphasis id="35D6531A1304BF15FB06FA21FE8E47D6" italics="true" pageId="46" pageNumber="47">
<taxonomicName id="C0A2F48B1304BF15FB06FA21FB6C47F3" authorityName="Gunther" authorityYear="1891" box="[1209,1279,1447,1472]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="miops">miops</taxonomicName>
,
<taxonomicName id="C0A2F48B1304BF15FAB4FA21FE8947D6" authorityName="Angel" authorityYear="1942" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="septemtuberculatus">septemtuberculatus</taxonomicName>
,
</emphasis>
or
<taxonomicName id="C0A2F48B1304BF15FEFEFA4DFE3F47D6" authorityName="Boettger" authorityYear="1913" box="[321,428,1483,1509]" class="Reptilia" family="Gekkonidae" genus="Microscalabotes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="spinulifer">
<emphasis id="35D6531A1304BF15FEFEFA4DFE3F47D6" box="[321,428,1483,1509]" italics="true" pageId="46" pageNumber="47">spinulifer</emphasis>
</taxonomicName>
is likely to apply to
<emphasis id="35D6531A1304BF15FD32FA4DFD3747D7" box="[653,676,1483,1508]" italics="true" pageId="46" pageNumber="47">L.</emphasis>
sp. 11, which we therefore formally name as species new to science,
<taxonomicName id="C0A2F48B1304BF15FF79FA69FE8B443A" authority="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz, 2022" authorityName="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz" authorityYear="2022" box="[198,280,1519,1545]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="fritzi" status="sp. nov.">
<emphasis id="35D6531A1304BF15FF79FA69FE8B443A" box="[198,280,1519,1545]" italics="true" pageId="46" pageNumber="47">L. fritzi</emphasis>
</taxonomicName>
<emphasis id="35D6531A1304BF15FEA0FA69FEEA443A" bold="true" box="[287,377,1519,1545]" pageId="46" pageNumber="47">
<taxonomicNameLabel id="2EE5EE611304BF15FEA0FA69FEEA443A" box="[287,377,1519,1545]" pageId="46" pageNumber="47" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="4FB8DC831304BF15FF28F9B1FEDC44AA" pageId="46" pageNumber="47" type="materials_examined">
<paragraph id="071D8F081304BF15FF28F9B1FEDC44AA" blockId="46.[151,1437,1590,2013]" pageId="46" pageNumber="47">
<materialsCitation id="B7CA85551304BF15FF28F9B1FEDC44AA" ID-GBIF-Occurrence="3897988320" collectingDate="2009-04-08" collectionCode="ZSM, ZCMV" collectorName="P. - S. Gehring &amp; Gehring, F &amp; E. Rajeriarison" country="Madagascar" latitude="-18.6058" location="Madagascar" longLatPrecision="7" longitude="49.2138" pageId="46" pageNumber="47" specimenCode="ZSM 651, ZCMV 8902" specimenCount="1" typeStatus="holotype">
<emphasis id="35D6531A1304BF15FF28F9B1FE984462" bold="true" box="[151,267,1591,1617]" pageId="46" pageNumber="47">
<typeStatus id="D81931AA1304BF15FF28F9B1FE954462" box="[151,262,1591,1617]" pageId="46" pageNumber="47" type="holotype">Holotype</typeStatus>
.
</emphasis>
<specimenCode id="570427731304BF15FEADF9B1FEE84462" box="[274,379,1590,1617]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="46" pageNumber="47" type="Museum">ZSM 651</specimenCode>
/2009 (
<specimenCode id="570427731304BF15FE74F9B1FDF24462" box="[459,609,1591,1617]" collectionCode="ZCMV" pageId="46" pageNumber="47">ZCMV 8902</specimenCode>
), female, collected by
<collectorName id="AA57EADE1304BF15FCE1F9B1FC6B4462" box="[862,1016,1591,1617]" pageId="46" pageNumber="47">P.-S. Gehring</collectorName>
, F. Ratsoavina, and
<collectorName id="AA57EADE1304BF15FB66F9B1FA124462" box="[1241,1409,1591,1617]" pageId="46" pageNumber="47">E. Rajeriarison</collectorName>
at Ankaninny Nofy, east coast of
<collectingCountry id="7FB5CF981304BF15FDBEF9DDFD194446" box="[513,650,1627,1653]" name="Madagascar" pageId="46" pageNumber="47">Madagascar</collectingCountry>
, geographical coordinates -
<geoCoordinate id="6296E9CF1304BF15FC7FF9DCFBBC4446" box="[960,1071,1626,1653]" degrees="18.6058" direction="south" orientation="latitude" pageId="46" pageNumber="47" precision="5" value="-18.6058">S18.6058</geoCoordinate>
,
<geoCoordinate id="6296E9CF1304BF15FB83F9DDFB384446" box="[1084,1195,1627,1653]" degrees="49.2138" direction="east" orientation="longitude" pageId="46" pageNumber="47" precision="5" value="49.2138">E49.2138</geoCoordinate>
, roughly at sea level, on
<date id="731CA9C81304BF15FF05F9F9FEDF44AA" box="[186,332,1663,1689]" pageId="46" pageNumber="47" value="2009-04-08">
<collectingDate id="635850201304BF15FF05F9F9FEDF44AA" box="[186,332,1663,1689]" pageId="46" pageNumber="47" value="2009-04-08">8 April 2009</collectingDate>
</date>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="4FB8DC831304BF14FF78F924FF7240B6" lastPageId="47" lastPageNumber="48" pageId="46" pageNumber="47" type="diagnosis">
<paragraph id="071D8F081304BF15FF78F924FAE6451A" blockId="46.[151,1437,1590,2013]" pageId="46" pageNumber="47">
<emphasis id="35D6531A1304BF15FF78F924FED1448F" bold="true" box="[199,322,1698,1724]" pageId="46" pageNumber="47">Diagnosis.</emphasis>
<taxonomicName id="C0A2F48B1304BF15FEF5F925FD8F448E" authority="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz, 2022" authorityName="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz" authorityYear="2022" box="[330,540,1699,1725]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="fritzi" status="sp. nov.">
<emphasis id="35D6531A1304BF15FEF5F925FD8F448E" box="[330,540,1699,1725]" italics="true" pageId="46" pageNumber="47">Lygodactylus fritzi</emphasis>
</taxonomicName>
<emphasis id="35D6531A1304BF15FD9BF925FDEC448E" bold="true" box="[548,639,1699,1725]" pageId="46" pageNumber="47">
<taxonomicNameLabel id="2EE5EE611304BF15FD9BF925FDEC448E" box="[548,639,1699,1725]" pageId="46" pageNumber="47" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is a species of the
<taxonomicName id="C0A2F48B1304BF15FCE5F925FC7D448E" box="[858,1006,1699,1725]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="genus">
<emphasis id="35D6531A1304BF15FCE5F925FC7D448E" box="[858,1006,1699,1725]" italics="true" pageId="46" pageNumber="47">Lygodactylus</emphasis>
</taxonomicName>
subgenus
<taxonomicName id="C0A2F48B1304BF15FBD7F925FB68448E" authorityName="Pasteur" authorityYear="1964" box="[1128,1275,1699,1725]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="subGenus" subGenus="Domerguella">
<emphasis id="35D6531A1304BF15FBD7F925FB68448E" box="[1128,1275,1699,1725]" italics="true" pageId="46" pageNumber="47">Domerguella</emphasis>
</taxonomicName>
based on molecular phylogenetic relationships, and it can also be assigned to the subgenus
<taxonomicName id="C0A2F48B1304BF15FC42F941FB0344D2" authorityName="Pasteur" authorityYear="1964" box="[1021,1168,1735,1761]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="subGenus" subGenus="Domerguella">
<emphasis id="35D6531A1304BF15FC42F941FB0344D2" box="[1021,1168,1735,1761]" italics="true" pageId="46" pageNumber="47">Domerguella</emphasis>
</taxonomicName>
by an undivided mental scale with two postmentals, absence of a claw on the first finger, and 7 preanal pores in males. Within
<taxonomicName id="C0A2F48B1304BF15FABCF96DFA054536" authorityName="Pasteur" authorityYear="1964" box="[1283,1430,1771,1797]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="55" phylum="Chordata" rank="subGenus" subGenus="Domerguella">
<emphasis id="35D6531A1304BF15FABCF96DFA054536" box="[1283,1430,1771,1797]" italics="true" pageId="46" pageNumber="47">Domerguella</emphasis>
</taxonomicName>
, the new species is one of several species of subclade A5 known from the Northern Central East of
<collectingCountry id="7FB5CF981304BF15FB58F889FAE3451A" box="[1255,1392,1807,1833]" name="Madagascar" pageId="46" pageNumber="47">Madagascar</collectingCountry>
.
</paragraph>
<paragraph id="071D8F081304BF14FF78F8B5FDAA42E6" blockId="46.[151,1437,1590,2013]" lastBlockId="47.[151,1437,151,2013]" lastPageId="47" lastPageNumber="48" pageId="46" pageNumber="47">
It differs from the nominal species of
<taxonomicName id="C0A2F48B1304BF15FDC9F8B5FC9A457E" authorityName="Pasteur" authorityYear="1964" box="[630,777,1843,1869]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="55" phylum="Chordata" rank="subGenus" subGenus="Domerguella">
<emphasis id="35D6531A1304BF15FDC9F8B5FC9A457E" box="[630,777,1843,1869]" italics="true" pageId="46" pageNumber="47">Domerguella</emphasis>
</taxonomicName>
occurring in northern
<collectingCountry id="7FB5CF981304BF15FBB5F8B5FB02457E" box="[1034,1169,1843,1869]" name="Madagascar" pageId="46" pageNumber="47">Madagascar</collectingCountry>
and belonging to subclades A1A4 as follows: from
<taxonomicName id="C0A2F48B1304BF15FE54F8D1FDED4543" authorityName=": Rosler" authorityYear="2000" box="[491,638,1879,1904]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="expectatus">
<emphasis id="35D6531A1304BF15FE54F8D1FDED4543" box="[491,638,1879,1904]" italics="true" pageId="46" pageNumber="47">L. expectatus</emphasis>
</taxonomicName>
by non-enlarged dorsolateral scales (longitudinal count of dorsal scales&gt;250 vs. &lt;170); from
<taxonomicName id="C0A2F48B1304BF15FE34F8FDFE7745A7" authorityName=": Rosler" authorityYear="2000" box="[395,484,1915,1940]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="rarus">
<emphasis id="35D6531A1304BF15FE34F8FDFE7745A7" box="[395,484,1915,1940]" italics="true" pageId="46" pageNumber="47">L. rarus</emphasis>
</taxonomicName>
by lack of regular crossbands on tail (vs. presence) and different body shape with less elongated limbs (relative hindlimb length 0.47 vs.&gt;0.55); from
<emphasis id="35D6531A1304BF15FCD0F819FADF458A" box="[879,1356,1951,1977]" italics="true" pageId="46" pageNumber="47">
<taxonomicName id="C0A2F48B1304BF15FCD0F819FC49458B" authorityName="Pasteur &amp; Blanc" authorityYear="1967" box="[879,986,1951,1976]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="petteri">L. petteri</taxonomicName>
,
<taxonomicName id="C0A2F48B1304BF15FC59F819FBFB458A" authorityName="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz" authorityYear="2022" box="[998,1128,1951,1977]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="tantsaha">L. tantsaha</taxonomicName>
,
<taxonomicName id="C0A2F48B1304BF15FBCAF819FB5F458A" authorityName="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz" authorityYear="2022" box="[1141,1228,1951,1977]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="salvi">L. salvi</taxonomicName>
,
<taxonomicName id="C0A2F48B1304BF15FB67F819FADB458A" authorityName="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz" authorityYear="2022" box="[1240,1352,1951,1977]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="roellae">L. roellae</taxonomicName>
,
</emphasis>
and
<taxonomicName id="C0A2F48B1304BF15FA39F819FF4645EE" authorityName="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz" authorityYear="2022" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="hapei">
<emphasis id="35D6531A1304BF15FA39F819FF4645EE" italics="true" pageId="46" pageNumber="47">L. hapei</emphasis>
</taxonomicName>
by a higher longitudinal count of dorsal scales (253 vs. &lt;241); and from
<taxonomicName id="C0A2F48B1304BF15FBBAF845FB7F45EE" authorityName=": Pasteur" authorityYear="1965" box="[1029,1260,1987,2013]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="35D6531A1304BF15FBBAF845FB7F45EE" box="[1029,1260,1987,2013]" italics="true" pageId="46" pageNumber="47">L. madagascariensis</emphasis>
</taxonomicName>
as well as most of the previously mentioned species of clades A1A4 by a rudimentary spine-like tubercle at the base of the tail in the only known female (vs. absence).
</paragraph>
<paragraph id="071D8F081305BF14FF78FF59FCAA400E" blockId="47.[151,1437,151,2013]" pageId="47" pageNumber="48">
From other species of subclade A5, the new species differs as follows: from
<taxonomicName id="C0A2F48B1305BF14FB82FF59FB3342CB" authorityName="Gunther" authorityYear="1891" box="[1085,1184,223,248]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="miops">
<emphasis id="35D6531A1305BF14FB82FF59FB3342CB" box="[1085,1184,223,248]" italics="true" pageId="47" pageNumber="48">L. miops</emphasis>
</taxonomicName>
by fewer dorsolateral tubercles between limbs (2 vs. 36), higher longitudinal dorsal scale count (247 vs. 207242), lower longitudinal ventral scale count (98 vs. 102113), and weakly expressed lateral tubercles at tail base vs. clearly recognizable in males and females; from
<taxonomicName id="C0A2F48B1305BF14FE6CFECDFDA44356" box="[467,567,331,357]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="guibei">
<emphasis id="35D6531A1305BF14FE6CFECDFDA44356" box="[467,567,331,357]" italics="true" pageId="47" pageNumber="48">L. guibei</emphasis>
</taxonomicName>
by higher longitudinal dorsal scale count (247 vs. 170220), and weakly expressed lateral tubercles at tail base vs. clearly recognizable, usually large in males and females; from
<taxonomicName id="C0A2F48B1305BF14FAA2FEE9FAEA43BA" authorityName="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz" authorityYear="2022" box="[1309,1401,367,393]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="winki">
<emphasis id="35D6531A1305BF14FAA2FEE9FAEA43BA" box="[1309,1401,367,393]" italics="true" pageId="47" pageNumber="48">L. winki</emphasis>
</taxonomicName>
by fewer dorsolateral tubercles between limbs (2 vs. 58), higher longitudinal dorsal scale count (247 vs. 187222), and weakly expressed lateral tubercles at tail base vs. clearly recognizable usually large in males and females; from
<taxonomicName id="C0A2F48B1305BF14FF28FE5DFF4F43C6" authorityName="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz" authorityYear="2022" box="[151,220,475,501]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="ulli">
<emphasis id="35D6531A1305BF14FF28FE5DFF4F43C6" box="[151,220,475,501]" italics="true" pageId="47" pageNumber="48">L. ulli</emphasis>
</taxonomicName>
possibly by more weakly expressed lateral tubercles at tail base and a lower longitudinal count of ventral scales (98 vs. 110). From all these species, it differs by phylogenetic position, at least 9% 16S distance, and absence of haplotype sharing in both nuclear-encoded genes studied.
</paragraph>
<paragraph id="071D8F081305BF14FF78FDC1FF7240B6" blockId="47.[151,1437,151,2013]" pageId="47" pageNumber="48">For a distinction from one other species newly named and described in the following, see the respective diagnosis below.</paragraph>
</subSubSection>
<subSubSection id="4FB8DC831305BF14FF78FD09FB9A4126" pageId="47" pageNumber="48" type="etymology">
<paragraph id="071D8F081305BF14FF78FD09FB9A4126" blockId="47.[151,1437,151,2013]" pageId="47" pageNumber="48">
<emphasis id="35D6531A1305BF14FF78FD09FEDD409A" bold="true" box="[199,334,655,681]" pageId="47" pageNumber="48">Etymology.</emphasis>
We are pleased to dedicate this species to Uwe Fritz, director of the Museum of Zoology, Dresden (part of the Senckenberg Natural History Collections), in recognition of his substantial contributions to the taxonomy of chelonians and squamates, and his tireless efforts to spearhead the fight for continued funding of basic taxonomic research. The name is a patronym (i.e., a noun in the genitive case).
</paragraph>
</subSubSection>
<subSubSection id="4FB8DC831305BF14FF78FC98FACF443A" pageId="47" pageNumber="48" type="description">
<paragraph id="071D8F081305BF14FF78FC98FC2E46F6" blockId="47.[151,1437,151,2013]" pageId="47" pageNumber="48">
<emphasis id="35D6531A1305BF14FF78FC98FD81410A" bold="true" box="[199,530,798,825]" pageId="47" pageNumber="48">
Description of the
<typeStatus id="D81931AA1305BF14FE1AFC99FD9D410A" box="[421,526,799,825]" pageId="47" pageNumber="48" type="holotype">holotype</typeStatus>
.
</emphasis>
Female in a good state of preservation, tail partly detached. SVL
<quantity id="C05A22ED1305BF14FB48FC99FACD410A" box="[1271,1374,799,825]" metricMagnitude="-2" metricUnit="m" metricValue="2.6399999999999997" pageId="47" pageNumber="48" unit="mm" value="26.4">26.4 mm</quantity>
, TAL
<quantity id="C05A22ED1305BF14FF28FCC4FF6D416E" box="[151,254,834,861]" metricMagnitude="-2" metricUnit="m" metricValue="2.65" pageId="47" pageNumber="48" unit="mm" value="26.5">26.5 mm</quantity>
; for other measurements see
<tableCitation id="4A20BAB31305BF14FDFFFCC5FD00416E" box="[576,659,835,861]" captionStart="TABLE 1" captionStartId="31.[151,239,459,485]" captionTargetPageId="31" captionText="TABLE 1. Selected morphometric and meristic data for voucher specimens of the Lygodactylus madagascariensis group examined for this study. All morphometric measurements (SVL, TAL, HIL, HW, ED, SED) in mm. All specimens in the table were genotyped by 16S sequences, except for historical types from BMNH, MNHN, and SMF, and two further individuals of L. expectatus. Abbreviations are those described in Materials and Methods section; additional abbreviations and symbols as follows: NA, not assessed or not applicable; * counts and measurements partly or fully taken by M. Puente (Puente et al. 2009); § specimen not genotyped; # tail broken or incomplete; ## tail regenerated. HT (sep) and HT (spi) mark the holotypes of Lygodactylus septemtuberculatus Angel, 1942 and Microscalabotes spinulifer Boettger, 1913, here considered to be junior synonyms of L. miops. First LCDS value for L. expectatus refers to the enlarged dorsolateral scales, the second value to the non-enlarged vertebral scales. See species account of L. fritzi sp. nov for the identity of the historical locality “Senbendrana”." httpUri="http://table.plazi.org/id/53DDDF801335BF24FF28FE4DFD0E408E" pageId="47" pageNumber="48" tableUuid="53DDDF801335BF24FF28FE4DFD0E408E">Table 1</tableCitation>
. Body broader than head. The distance from the tip of the snout to the anterior border of the eye (
<quantity id="C05A22ED1305BF14FE77FCE1FDB741B2" box="[456,548,871,897]" metricMagnitude="-3" metricUnit="m" metricValue="3.1" pageId="47" pageNumber="48" unit="mm" value="3.1">3.1 mm</quantity>
), is less than the interorbital distance anteriorly (
<quantity id="C05A22ED1305BF14FBF3FCE0FB3B41B2" box="[1100,1192,870,897]" metricMagnitude="-3" metricUnit="m" metricValue="3.5" pageId="47" pageNumber="48" unit="mm" value="3.5">3.5 mm</quantity>
), and greater than the distance between the eye and ear opening. Snout covered with granular scales larger than those on the rest of the dorsum. Nostril surrounded by four scales: rostral, first supralabial, and two supranasals. Mental scale undivided; no contact between posterior projection of mental scale and first infralabial; two asymmetrical postmental scales with four postpostmental scales; seven infralabial scales; eight supralabial scales; two internasal scales; granular dorsal scales; dorsum with small, homogeneous, granular, and unkeeled scales of similar size to those on trunk, no distinct size difference to scales on limbs; 247 dorsal scales longitudinally along the body; 98 ventral scales between mental and cloaca; venter with large homogeneous smooth scales; first finger present but very small, not bearing a claw; three pairs of subdigital lamellae on the fourth toe; two not very distinct dorsolateral tubercles, each consisting of one scale; tail without whorls; small lateral spines at the base of the tail.
</paragraph>
<paragraph id="071D8F081305BF14FF78FB49FA8D4766" blockId="47.[151,1437,151,2013]" pageId="47" pageNumber="48">
Coloration of the
<typeStatus id="D81931AA1305BF14FE32FB49FE7C46DA" box="[397,495,1231,1257]" pageId="47" pageNumber="48" type="holotype">holotype</typeStatus>
is only described from the specimen that was preserved in ethanol for 12 years. The dorsum is fawn to brownish with scattered darker brown spots, the flanks are darker than the dorsum. A distinctive dark stripe is displayed on the shoulder. The parietal exhibits two darker areas. The tail is fawn with equispaced darker brown stripes on it. Venter and the snout are uniformly whitish with small irregular brown spots.
</paragraph>
<paragraph id="071D8F081305BF14FF78FAD8FACF443A" blockId="47.[151,1437,151,2013]" pageId="47" pageNumber="48">
<emphasis id="35D6531A1305BF14FF78FAD8FEAC474B" bold="true" box="[199,319,1374,1400]" pageId="47" pageNumber="48">Variation.</emphasis>
Morphometric and meristic data are only known from a single voucher specimen. However, color patterns could be assessed from photographs of three additional individuals (
<figureCitation id="9F99938D1305BF14FC4EFA05FBD647AE" box="[1009,1093,1411,1437]" captionStart="FIGURE 22" captionStartId="48.[151,250,958,982]" captionTargetBox="[151,1436,181,934]" captionTargetId="figure-18@48.[151,1436,181,934]" captionTargetPageId="48" captionText="FIGURE 22. Photos of Lygodactylus fritzi sp. nov. specimens from Vohibola in life. All specimens correspond to genotyped individuals included in Fig. 1 that were not collected." figureDoi="http://doi.org/10.5281/zenodo.7040811" httpUri="https://zenodo.org/record/7040811/files/figure.png" pageId="47" pageNumber="48">Fig. 22</figureCitation>
), some of which were dorsally gray-beige with an irregular contrasted pattern of larger light and smaller dark spots (
<figureCitation id="9F99938D1305BF14FBFFFA21FB3447F2" box="[1088,1191,1447,1473]" captionStart="FIGURE 22" captionStartId="48.[151,250,958,982]" captionTargetBox="[151,1436,181,934]" captionTargetId="figure-18@48.[151,1436,181,934]" captionTargetPageId="48" captionText="FIGURE 22. Photos of Lygodactylus fritzi sp. nov. specimens from Vohibola in life. All specimens correspond to genotyped individuals included in Fig. 1 that were not collected." figureDoi="http://doi.org/10.5281/zenodo.7040811" httpUri="https://zenodo.org/record/7040811/files/figure.png" pageId="47" pageNumber="48">Fig. 22A</figureCitation>
), others more uniform with a more or less symmetrical pattern of dark spots (
<figureCitation id="9F99938D1305BF14FD4AFA4DFCC947D6" box="[757,858,1483,1509]" captionStart="FIGURE 22" captionStartId="48.[151,250,958,982]" captionTargetBox="[151,1436,181,934]" captionTargetId="figure-18@48.[151,1436,181,934]" captionTargetPageId="48" captionText="FIGURE 22. Photos of Lygodactylus fritzi sp. nov. specimens from Vohibola in life. All specimens correspond to genotyped individuals included in Fig. 1 that were not collected." figureDoi="http://doi.org/10.5281/zenodo.7040811" httpUri="https://zenodo.org/record/7040811/files/figure.png" pageId="47" pageNumber="48">Fig. 22E</figureCitation>
), or dark brown with weakly contrasted light brown dorsolateral bands with a somewhat reddish tone (
<figureCitation id="9F99938D1305BF14FD77FA69FCC3443A" box="[712,848,1519,1545]" captionStart="FIGURE 22" captionStartId="48.[151,250,958,982]" captionTargetBox="[151,1436,181,934]" captionTargetId="figure-18@48.[151,1436,181,934]" captionTargetPageId="48" captionText="FIGURE 22. Photos of Lygodactylus fritzi sp. nov. specimens from Vohibola in life. All specimens correspond to genotyped individuals included in Fig. 1 that were not collected." figureDoi="http://doi.org/10.5281/zenodo.7040811" httpUri="https://zenodo.org/record/7040811/files/figure.png" pageId="47" pageNumber="48">Fig. 22BC</figureCitation>
). Ventrally irregularly dark spotted (
<figureCitation id="9F99938D1305BF14FB58FA69FAC3443A" box="[1255,1360,1519,1545]" captionStart="FIGURE 22" captionStartId="48.[151,250,958,982]" captionTargetBox="[151,1436,181,934]" captionTargetId="figure-18@48.[151,1436,181,934]" captionTargetPageId="48" captionText="FIGURE 22. Photos of Lygodactylus fritzi sp. nov. specimens from Vohibola in life. All specimens correspond to genotyped individuals included in Fig. 1 that were not collected." figureDoi="http://doi.org/10.5281/zenodo.7040811" httpUri="https://zenodo.org/record/7040811/files/figure.png" pageId="47" pageNumber="48">Fig. 22D</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="4FB8DC831305BF14FF78F994FBE74542" pageId="47" pageNumber="48" type="distribution">
<paragraph id="071D8F081305BF14FF78F994FBE74542" blockId="47.[151,1437,151,2013]" pageId="47" pageNumber="48">
<emphasis id="35D6531A1305BF14FF78F994FEF2441F" bold="true" box="[199,353,1554,1580]" pageId="47" pageNumber="48">Distribution.</emphasis>
<taxonomicName id="C0A2F48B1305BF14FED7F995FE29441E" authorityName="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz" authorityYear="2022" box="[360,442,1555,1581]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="fritzi">
<emphasis id="35D6531A1305BF14FED7F995FE29441E" box="[360,442,1555,1581]" italics="true" pageId="47" pageNumber="48">L. fritzi</emphasis>
</taxonomicName>
is only known from (1) the
<typeStatus id="D81931AA1305BF14FD49F995FCB5441E" box="[758,806,1555,1581]" pageId="47" pageNumber="48">type</typeStatus>
locality Ankaninny Nofy and (2) Vohibola, two coastal lowland localities (
<quantity id="C05A22ED1305BF14FED4F9B1FE534462" box="[363,448,1591,1617]" metricMagnitude="1" metricUnit="m" metricValue="1.0" metricValueMax="2.0" metricValueMin="0.0" pageId="47" pageNumber="48" unit="m" value="10.0" valueMax="20.0" valueMin="0.0">020 m</quantity>
a.s.l.) in the Northern Central East of
<collectingCountry id="7FB5CF981305BF14FCD9F9B1FC7C4462" box="[870,1007,1591,1617]" name="Madagascar" pageId="47" pageNumber="48">Madagascar</collectingCountry>
. Littoral forest harbours a high species richness especially of plants, with several genera endemic to this habitat (
<bibRefCitation id="6333F2F91305BF14FC61F9DDFAD74446" author="de Gouvenain, R. C. &amp; Silander, J. A." box="[990,1348,1627,1653]" pageId="47" pageNumber="48" pagination="103 - 111" refId="ref36974" refString="de Gouvenain, R. C. &amp; Silander, J. A. (2003) Littoral forest. In: Goodman, S. M. &amp; Benstead, J. P. (Eds.), The Natural History of Madagascar. The University of Chicago Press, London, pp. 103 - 111" type="book chapter" year="2003">de Gouvenain &amp; Silander 2003</bibRefCitation>
;
<bibRefCitation id="6333F2F91305BF14FAEEF9DDFED544AA" author="Bollen, A. &amp; Donati, G." pageId="47" pageNumber="48" pagination="32 - 43" refId="ref36242" refString="Bollen, A. &amp; Donati, G. (2005) Phenology of the littoral forest of Sainte Luce, southeastern Madagascar. Biotropica, 37, 32 - 43 https: // doi. org / 10.1111 / j. 1744 - 7429.2005.04094. x" type="journal article" year="2005">Bollen &amp; Donati 2005</bibRefCitation>
).
<bibRefCitation id="6333F2F91305BF14FEE6F9F9FDFF44AA" author="Fisher, B. L. &amp; Girman, D." box="[345,620,1663,1689]" pageId="47" pageNumber="48" pagination="331 - 344" refId="ref37363" refString="Fisher, B. L. &amp; Girman, D. (2000) Biogeography of ants in eastern Madagascar. In: Lourenco, W. R. &amp; Goodman, S. M. (Eds.), Diversity and Endemism in Madagascar. Memoires de la Societe de Biogeographie, Paris, pp. 331 - 344." type="book chapter" year="2000">Fisher &amp; Girman (2000)</bibRefCitation>
identified littoral forests as one of four major areas for ant endemism in
<collectingCountry id="7FB5CF981305BF14FF28F925FEB3448E" box="[151,288,1699,1725]" name="Madagascar" pageId="47" pageNumber="48">Madagascar</collectingCountry>
. Previous studies in south-eastern littoral forests found no vertebrate species strictly endemic to that forest
<typeStatus id="D81931AA1305BF14FF62F941FE9E44D2" box="[221,269,1735,1761]" pageId="47" pageNumber="48">type</typeStatus>
(
<bibRefCitation id="6333F2F91305BF14FEA4F941FD9944D2" author="Ganzhorn, J. U. &amp; Goodman, S. M. &amp; Ramanamanjato, J. B. &amp; Rakotondavony, D. &amp; Rakotosamimanana, B. &amp; Vallan, D." box="[283,522,1735,1761]" pageId="47" pageNumber="48" pagination="155 - 164" refId="ref37529" refString="Ganzhorn, J. U., Goodman, S. M., Ramanamanjato, J. B., Rakotondavony, D., Rakotosamimanana, B. &amp; Vallan, D. (2000) Vertebrate species in fragmented littoral forest of Madagascar. In: Lourenco, W. R. &amp; Goodman, S. M. (Eds.), Diversity and endemism in Madagascar. Memoires de la Societe de Biogeographie Edition. Museum Histoire Naturelle, Paris, pp. 155 - 164." type="book chapter" year="2000">
Ganzhorn
<emphasis id="35D6531A1305BF14FE2EF941FE5B44D2" box="[401,456,1735,1761]" italics="true" pageId="47" pageNumber="48">et al.</emphasis>
2000
</bibRefCitation>
;
<bibRefCitation id="6333F2F91305BF14FDABF941FC0944D2" author="Goodman, S. M. &amp; Ramanamanjato, J. P." box="[532,922,1735,1761]" pageId="47" pageNumber="48" pagination="25 - 48" refId="ref38325" refString="Goodman, S. M. &amp; Ramanamanjato, J. P. (2007) A perspective on the Paleo - ecology and biogeography of extreme southeastern Madagascar, with special reference to animals. In: Ganzhorn, J. U., Goodman, S. M. &amp; Vincelette, M. (Eds.), Biodiversity, Ecology and Conservation of the Littoral Ecosystems of South - eastern Madagascar, Tolagnaro (Fort Dauphin). SI / MAB Series # 11. Smithsonian Institution, Washington DC, USA, pp. 25 - 48." type="book chapter" year="2007">Goodman &amp; Ramanamanjato 2007</bibRefCitation>
) but recent taxonomic revisions have revealed numerous amphibians and reptiles restricted to small areas of forest directly adjacent to the Madagascars coast, such as for example species of the miniaturized frog genus
<taxonomicName id="C0A2F48B1305BF14FC93F889FBD0451A" authority="(Scherz et al. 2019)" baseAuthorityName="Scherz" baseAuthorityYear="2019" box="[812,1091,1807,1833]" class="Amphibia" family="Microhylidae" genus="Mini" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="genus">
<emphasis id="35D6531A1305BF14FC93F889FCF2451B" box="[812,865,1807,1832]" italics="true" pageId="47" pageNumber="48">Mini</emphasis>
(
<bibRefCitation id="6333F2F91305BF14FCD0F889FBAF451A" author="Scherz, M. D. &amp; Hutter, C. R. &amp; Rakotoarison, A. &amp; Riemann, J. C. &amp; Rodel, M. O. &amp; Ndriantsoa, S. H. &amp; Glos, J. &amp; Hyde Roberts, S. &amp; Crottini, A. &amp; Vences, M. &amp; Glaw, F." box="[879,1084,1807,1833]" pageId="47" pageNumber="48" pagination="0213314" refId="ref41924" refString="Scherz, M. D., Hutter, C. R., Rakotoarison, A., Riemann, J. C., Rodel, M. O., Ndriantsoa, S. H., Glos, J., Hyde Roberts, S., Crottini, A., Vences, M. &amp; Glaw, F. (2019) Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera. PLoS One, 14, e 0213314. https: // doi. org / 10.1371 / journal. pone. 0213314" type="journal article" year="2019">
Scherz
<emphasis id="35D6531A1305BF14FC7CF889FC67451A" box="[963,1012,1807,1833]" italics="true" pageId="47" pageNumber="48">et al</emphasis>
. 2019
</bibRefCitation>
)
</taxonomicName>
or Pandanus-dwelling frogs of
<taxonomicName id="C0A2F48B1305BF14FF28F8B5FDB0457E" authority="(Lehtinen et al. 2011)" baseAuthorityName="Lehtinen" baseAuthorityYear="2011" box="[151,547,1843,1869]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="genus">
<emphasis id="35D6531A1305BF14FF28F8B5FEB5457E" box="[151,294,1843,1869]" italics="true" pageId="47" pageNumber="48">Guibemantis</emphasis>
(
<bibRefCitation id="6333F2F91305BF14FE8AF8B5FD88457E" author="Lehtinen, R. M. &amp; Glaw, F. &amp; Vences, M." box="[309,539,1843,1869]" pageId="47" pageNumber="48" pagination="95 - 112" refId="ref39163" refString="Lehtinen, R. M., Glaw, F. &amp; Vences, M. (2011) Two new plant-breeding frog species (Anura: Mantellidae, Guibemantis) from southeastern Madagascar. Herpetological Journal, 21, 95 - 112." type="journal article" year="2011">
Lehtinen
<emphasis id="35D6531A1305BF14FE1EF8B5FE41457E" box="[417,466,1843,1869]" italics="true" pageId="47" pageNumber="48">et al</emphasis>
. 2011
</bibRefCitation>
)
</taxonomicName>
, or the chameleon
<taxonomicName id="C0A2F48B1305BF14FD49F8B5FB53457E" authority="(Gehring et al. 2011)" baseAuthorityName="Gehring" baseAuthorityYear="2011" box="[758,1216,1843,1869]" class="Reptilia" family="Chamaeleonidae" genus="Calumma" kingdom="Animalia" order="Squamata" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="vohibola">
<emphasis id="35D6531A1305BF14FD49F8B5FC5F457E" box="[758,972,1843,1869]" italics="true" pageId="47" pageNumber="48">Calumma vohibola</emphasis>
(
<bibRefCitation id="6333F2F91305BF14FC65F8B5FB2A457E" author="Gehring, P. - S. &amp; Ratsoavina, F. M. &amp; Vences, M. &amp; Glaw, F." box="[986,1209,1843,1869]" pageId="47" pageNumber="48" pagination="130 - 154" refId="ref37666" refString="Gehring, P. - S., Ratsoavina, F. M., Vences, M. &amp; Glaw, F. (2011) Calumma vohibola, a new chameleon species (Squamata: Chamaeleonidae) from the littoral forests of eastern Madagascar. African Journal of Herpetology, 60, 130 - 154. https: // doi. org / 10.1080 / 21564574.2011.628412" type="journal article" year="2011">
Gehring
<emphasis id="35D6531A1305BF14FB80F8B5FBE4457E" box="[1087,1143,1843,1869]" italics="true" pageId="47" pageNumber="48">et al.</emphasis>
2011
</bibRefCitation>
)
</taxonomicName>
,
<taxonomicName id="C0A2F48B1305BF14FB73F8B5FA0F457E" authorityName="Vences &amp; Multzsch &amp; Gippner &amp; Miralles &amp; Crottini &amp; Gehring &amp; Rakotoarison &amp; Ratsoavina &amp; Glaw &amp; Scherz" authorityYear="2022" box="[1228,1436,1843,1869]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="fritzi">
<emphasis id="35D6531A1305BF14FB73F8B5FA0F457E" box="[1228,1436,1843,1869]" italics="true" pageId="47" pageNumber="48">Lygodactylus fritzi</emphasis>
</taxonomicName>
adds to this growing list of species specialized to these highly threatened coastal forests.
</paragraph>
</subSubSection>
<subSubSection id="4FB8DC831305BF14FF78F8FDFF7345EE" pageId="47" pageNumber="48" type="biology_ecology">
<paragraph id="071D8F081305BF14FF78F8FDFF7345EE" blockId="47.[151,1437,151,2013]" pageId="47" pageNumber="48">
<emphasis id="35D6531A1305BF14FF78F8FDFE1645A7" bold="true" box="[199,389,1914,1941]" pageId="47" pageNumber="48">Natural history.</emphasis>
One adult specimen was photographed millimeters from the posterior end of a bug (
<figureCitation id="9F99938D1305BF14FA84F8FDFA1C45A6" box="[1339,1423,1915,1941]" captionStart="FIGURE 22" captionStartId="48.[151,250,958,982]" captionTargetBox="[151,1436,181,934]" captionTargetId="figure-18@48.[151,1436,181,934]" captionTargetPageId="48" captionText="FIGURE 22. Photos of Lygodactylus fritzi sp. nov. specimens from Vohibola in life. All specimens correspond to genotyped individuals included in Fig. 1 that were not collected." figureDoi="http://doi.org/10.5281/zenodo.7040811" httpUri="https://zenodo.org/record/7040811/files/figure.png" pageId="47" pageNumber="48">Fig. 22</figureCitation>
), and was presumably consuming honeydew excreted by the insect, as is known from gecko species (
<bibRefCitation id="6333F2F91305BF14FAB7F819FF4045EE" author="Folling, M. &amp; Knogge, C. &amp; Bohme, W." pageId="47" pageNumber="48" pagination="279 - 284" refId="ref37425" refString="Folling, M., Knogge, C. &amp; Bohme, W. (2001) Geckos are milking honeydew-producing planthoppers in Madagascar. Journal of Natural History, 35, 279 - 284. https: // doi. org / 10.1080 / 00222930150215378" type="journal article" year="2001">
Fölling
<emphasis id="35D6531A1305BF14FADCF819FA0E458A" box="[1379,1437,1951,1977]" italics="true" pageId="47" pageNumber="48">et al.</emphasis>
2001
</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>