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Measurements in mm, and two scale counts (SUPL and LAM3T) taken from all Uroplatus specimens included in this study. HT, holotype; PT, paratype; NA, not available, NM, not measured. See materials and methods for other abbreviations. All measurements taken by the first author except those marked by an asterisk. 

Measurements in mm, and two scale counts (SUPL and LAM3T) taken from all Uroplatus specimens included in this study. HT, holotype; PT, paratype; NA, not available, NM, not measured. See materials and methods for other abbreviations. All measurements taken by the first author except those marked by an asterisk. 

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We describe a new leaf-tailed gecko species of the Uroplatus ebenaui group from the eastern central rainforests of Madagascar, which had previously been considered as a confirmed candidate species. Our description of Uroplatus fiera sp. nov. relies on integrating evidence from molecular and morphological characters and is based on newly collected m...

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Context 1
... visual comparison of the tails of U. fiera with those of the four nominal species in the U. ebenaui group, plus U. ebenaui [Ca1], suggests clear and constant differences in general tail size of the new species from all species except U. ebenaui (Fig. 3), regardless of sex. The ratio TAL/ SVL ( Fig. 4; original data in Table 1) yields similar values for U. fiera, U. ebenaui [Ca1] and U. ebenaui, but tails are relatively much longer in U. phantasticus, U. finiavana and U. malama. Where sample size is large enough for a statistical testing (U. fiera versus U. phantasticus) this difference is significant (Mann-Whitney U test; P=0.002). ...
Context 2
... important character of the U. ebenaui group is the pigmentation of the oral mucosa: it is black in U. phantasticus, U. ebenaui, and U. malama, but unpigmented in U. finiavana. We confirmed that all studied specimens of U. fiera and U. ebenaui [Ca1] (those listed in Table 1 plus additional specimens seen in collections and in the field) have an unpigmented oral mucosa, thus clearly differing from especially U. ebenaui, which by tail size and shape cannot be reliably distinguished from these lineages. A further difference between the new species U. fiera and U. ebenaui is observed in the number of lamellae under the third toe (Figs. ...
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... morphological data were based only on the original description (Boulenger 1888) and a photo of the preserved type specimen figured by Böhme & Henkel (1995). We recently re-examined the holotype of U. phantasticus (BMNH 1946.8.26.64) and provide our own measurements of this specimen (Table 1). Based on our re-examination, we confirm that the holotype (Fig. 6) conforms more closely to the long-and wide-tailed form from eastern Madagascar, to which it is usually assigned. ...
Context 4
... 0.30) than the long-tailed form considered to represent U. phantasticus 0.27-0.30, mean 0.29, U. phantasticus holotype not included), but this difference is not statistically significant and the values largely overlap (U-test, P=0.24; original data in Table 1). Still it should be mentioned that the value for the phantasticus holotype according to our own new measurements (0.296) is closer to the mean of the long-tailed form. ...
Context 5
... male in good condition with intact tail and everted hemipenes. SVL 66.2 mm, tail length 19.9 mm, maximum tail width 3.8 mm, for further measurements see Table 1. Head triangular in dorsal view, postorbital region 6.8 mm; snout length 7.3 mm; canthus rostralis indistinct; snout sloping strongly and continuously downward anteriorly; snout depressed, short (1.2 times longer than eye diameter); eyes large (eye diameter 6.1 mm), bulging slightly above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small (horizontal diameter 1.0-1.1 mm), its opening facing posterolaterally, but also posteroventrally (ear opening clearly visible in ventral view but not in dorsal view); nostrils laterally oriented; body somewhat laterally compressed, without lateral fringes; limbs well developed, without fringes, forelimb reaches beyond tip of snout when adpressed forward and almost to the groin when adpressed backwards along body (forelimb length/axilla-groin distance 29.1/25.7 mm =113%), hind limb reaches beyond axilla when adpressed forward along body (hind limb length/axilla-groin distance 36.3/34.9 ...
Context 6
... general, the paratypes agree well with the holotype in morphology. For measurements, see Table 1. The number of head spines varies from 4 (FRC 651) to 25 (ZSM 211/2002). ...
Context 7
... number of recognizable transverse lines across the head (in addition to the one between supraciliaries) varies from zero (ZSM 211/2002) to three (in various specimens). FRC 650 is the largest specimens (SVL 78.0 mm; Table 1). The colour pattern is highly variable. ...

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... This is also reflected by a lack of taxonomic information, indicated by numerous deep genetic lineages still not revised and scientifically named (Nagy et al. 2012). These candidate species are relatively evenly distributed across all main clades of Malagasy reptiles, and consequently, many recent taxonomic papers revised and named single or a limited number of new species, for instance in geckos (e.g., Glaw et al. 2010Glaw et al. , 2014Glaw et al. , 2018Jono et al. 2015;Crottini et al. 2011Crottini et al. , 2015Köhler et al. 2019;Ratsoavina et al. 2011Ratsoavina et al. , 2015Ratsoavina et al. , 2017Ratsoavina et al. , 2019aRatsoavina et al. ,b, 2020Miralles et al. 2021). In some groups of geckos and other squamates, the amount of cryptic genetic variation detected was so enormous that long-term efforts integrating multiple lines of evidence-including genetics, external morphology, osteology, and genital morphology, as well as new targeted fieldwork to collect fresh tissue samples and voucher specimens-were necessary to circumscribe species, assign historical types, and eventually describe multiple new species in a series of papers: e.g., chameleons of the Calumma nasutum group or leaf-tailed geckos of the Uroplatus ebenaui group, where initial diversity was detected by DNA barcoding (Gehring et al. 2012;Ratsoavina et al. 2013) and species subsequently named and described over several years (Ratsoavina et al. 2015(Ratsoavina et al. , 2017(Ratsoavina et al. , 2019a(Ratsoavina et al. ,b, 2020Prötzel et al. 2017Prötzel et al. , 2018Prötzel et al. , 2020. ...
... These candidate species are relatively evenly distributed across all main clades of Malagasy reptiles, and consequently, many recent taxonomic papers revised and named single or a limited number of new species, for instance in geckos (e.g., Glaw et al. 2010Glaw et al. , 2014Glaw et al. , 2018Jono et al. 2015;Crottini et al. 2011Crottini et al. , 2015Köhler et al. 2019;Ratsoavina et al. 2011Ratsoavina et al. , 2015Ratsoavina et al. , 2017Ratsoavina et al. , 2019aRatsoavina et al. ,b, 2020Miralles et al. 2021). In some groups of geckos and other squamates, the amount of cryptic genetic variation detected was so enormous that long-term efforts integrating multiple lines of evidence-including genetics, external morphology, osteology, and genital morphology, as well as new targeted fieldwork to collect fresh tissue samples and voucher specimens-were necessary to circumscribe species, assign historical types, and eventually describe multiple new species in a series of papers: e.g., chameleons of the Calumma nasutum group or leaf-tailed geckos of the Uroplatus ebenaui group, where initial diversity was detected by DNA barcoding (Gehring et al. 2012;Ratsoavina et al. 2013) and species subsequently named and described over several years (Ratsoavina et al. 2015(Ratsoavina et al. , 2017(Ratsoavina et al. , 2019a(Ratsoavina et al. ,b, 2020Prötzel et al. 2017Prötzel et al. , 2018Prötzel et al. , 2020. ...
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... This applies also to the geckos of the genus Uroplatus, which have been the subject of several morphological and molecular studies (see e.g., [13][14][15][16][17][18][19][20][21][22][23]), but only U. phantasticus (Boulenger, 1888) has a known karyotype, leaving the chromosome diversity of the genus completely unexplored. Overall, the karyotypes of geckos exhibit a wide variability in terms of the total number of chromosomes, number of uni-armed and bi-armed chromosomes, localization of different chromosome markers and presence or absence of differentiated sex chromosomes [6,8,9,24]. ...
... A fragment of about 450 bp of the mitochondrial 12S rRNA gene was amplified using the primer pair 12Sa 5 -AAACTGGGATTAGATACCCCACTAT−3 and 12Sb 5 -GAGGGTGAGGGCGGTG-TGT−3 [28]. This marker was chosen considering its wide use on Uroplatus geckos and the number of available sequences in public repositories [13,[15][16][17][18][19][20][21][22][23]. ...
... For taxonomic attribution, the newly determined sequences were compared with available homologous traits deposited in GenBank which were used in previous phylogenetic and taxonomic studies on the genus Uroplatus (see e.g., [13][14][15][16][17][18][19][20][21][22][23]. ...
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We provide here the first karyotype description of eight Uroplatus species and a characterization of their chromosomal diversity. We performed a molecular taxonomic assessment of several Uroplatus samples using the mitochondrial 12S marker and a comparative cytogenetic analysis with standard karyotyping, silver staining (Ag-NOR) and sequential C-banding + Giemsa, +Chromomycin A3 (CMA3), +4′,6-diamidino-2-phenylindole (DAPI). We found chromosomal variability in terms of chromosome number (2n = 34–38), heterochromatin composition and number and localization of loci or Nucleolar Organizer Regions (NORs) (alternatively on the 2nd, 6th, 10th or 16th pair). Chromosome morphology is almost constant, with karyotypes composed of acrocentric chromosomes, gradually decreasing in length. C-banding evidenced a general low content of heterochromatin, mostly localized on pericentromeric and telomeric regions. Centromeric bands varied among the species studied, resulting in CMA3 positive and DAPI negative or positive to both fluorochromes. We also provide evidence of a first putative heteromorphic sex chromosome system in the genus. In fact, in U. alluaudi the 10th pair was highly heteromorphic, with a metacentric, largely heterochromatic W chromosome, which was much bigger than the Z. We propose an evolutionary scenario of chromosome reduction from 2n = 38 to 2n = 34, by means of translocations of microchromosomes on larger chromosomes (often involving the NOR-bearing microchromosomes). Adding our data to those available from the literature, we show that similar processes characterized the evolutionary radiation of a larger gecko clade. Finally, we hypothesize that sex chromosome diversification occurred independently in different genera.
... Especially the small-sized, leaf-mimicking forms in the U. ebenaui group have been subject to intensive research in recent years. After initial assessments of substantial cryptic diversity and definition of candidate species (Greenbaum et al. 2007;Raxworthy et al. 2008;Ratsoavina et al. 2013), six new species were formally described and named (Ratsoavina et al. 2011(Ratsoavina et al. , 2015(Ratsoavina et al. , 2017(Ratsoavina et al. , 2019a. This stepwise procedure-instead of a comprehensive revisionary work of the entire species group-was necessary because each of the new species presented idiosyncratic challenges for taxonomic revision, including limited availability of specimens for morphological study and extreme morphological similarity to other species, as well as problems in combining information from various research teams due to the use of DNA sequences from different genes (see Ratsoavina et al. 2013). ...
... For molecular analysis, total genomic DNA was extracted following a standard salt extraction protocol using proteinase K digestion in a concentration of 10 mg/ml (Bruford et al. 1992). DNA sequences from previous studies (Ratsoavina et al. 2011(Ratsoavina et al. , 2012(Ratsoavina et al. , 2013(Ratsoavina et al. , 2015(Ratsoavina et al. , 2017(Ratsoavina et al. , 2019a were complemented with new sequences for the new species named herein and U. finiavana, plus sequences retrieved from GenBank from the study of Raxworthy et al. (2008). ...
... The phylogenetic analysis is based on concatenated partial sequences of the four mitochondrial genes for NADH dehydrogenase subunit 4 (ND4), cytochrome oxidase I (COXI), 16S ribosomal RNA (16S rRNA or 16S), and 12S ribosomal RNA (12S rRNA or 12S), as well as the nuclear locus for oocyte maturation factor Mos (CMOS), which were amplified and sequenced with primers and PCR protocols as in previous studies (Ratsoavina et al. 2011(Ratsoavina et al. , 2012(Ratsoavina et al. , 2013(Ratsoavina et al. , 2015. In addition, we sequenced segments of the two nuclear genes sacsin (SACS), with primers and a nested PCR approach of Shen et al. (2012), and recombination-activating gene 1 (RAG1) with primers and a nested approach as described in Rakotoarison et al. (2015) from samples of U. finiavana and of the candidate species Ca3 and Ca4. ...
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The northern part of Madagascar is well known for its high species diversity and endemism. Exceptional species richness is related to the existence of large forest blocks and mountain complexes. These areas shelter a diverse variety of habitats occupied by a wide diversity of species, including leaf-tailed geckos of the genus Uroplatus. Based on morphological and molecular evidence, we here formally name two evolutionary lineages as new species that previously had been considered as candidate species (Uroplatus spp. Ca3 and Ca4), both small-sized species of the Uroplatus ebenaui group. Genetically, both new species are related to U. finiavana with a genetic divergence (uncorrected pairwise distance) of 10.3-12.8% in a fragment of the mitochondrial 16S rRNA gene, and separated from each other by 10.
... The leaf-mimicking forms belong to a single species-rich clade, the U. ebenaui species group. Recent taxonomic work on this clade has resulted in the description of several new species and in the identification of additional candidate species (Ratsoavina et al. 2011(Ratsoavina et al. , 2012(Ratsoavina et al. , 2013(Ratsoavina et al. , 2015(Ratsoavina et al. , 2017(Ratsoavina et al. , 2019. ...
... genetically it was highly divergent (25.7% distance from U. ebenaui in a segment of the 12S rRNA gene). Given this situation we excluded this species from subsequent accounts (Ratsoavina et al. 2015(Ratsoavina et al. , 2017(Ratsoavina et al. , 2019, awaiting confirmation of the genetic results. For the present study, we disposed of a further specimen and an additional tissue sample, and we included several new samples of its putative closest relative, U. ebenaui. ...
... Total genomic DNA was extracted following a standard salt extraction protocol using proteinase K digestion in a concentration of 10 mg/ml (Bruford et al. 1992). For molecular analysis, we complemented DNA sequences from previous studies (Ratsoavina et al. 2011(Ratsoavina et al. , 2012(Ratsoavina et al. , 2013(Ratsoavina et al. , 2015(Ratsoavina et al. , 2017(Ratsoavina et al. , 2019 with new sequences for the new species described herein, and for U. ebenaui and U. malama. ...
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Previous studies on leaf-tailed geckos of the genus Uroplatus identified a lineage from the Ankarana karst massif in northern Madagascar as genetically highly divergent, but only fragmentary information was available on these geckos. Here, we provide an integrative analysis based on molecular and morphological data, including a newly collected specimen from this locality. Phylogenetic analysis placed the Ankarana lineage sister to U. ebenaui, but with a surprisingly high genetic divergence of over 19% uncorrected pairwise distance in the mitochondrial 16S rRNA gene. We formally describe the Ankarana lineage as Uroplatus fetsy sp. nov. and provide new information on the distribution and genetic differentiation of its sister species, Uroplatus ebenaui. Based on DNA sequences of newly examined samples this latter species is confirmed as widespread, ranging from low-elevation dry forests from Beanka in western Madagascar to Forêt d'Ambre and Analalava/Fanambana in northern Madagascar, and thus far has not been found in the Ankarana Massif, where U. fetsy occurs.
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... Yet much work still remains in the integrative taxonomy of the leaf-mimicking, smaller-bodied species of the Uroplatus ebenaui group (Ratsoavina et al. 2013). Several candidate species are recorded in the northern part of Madagascar (Ratsoavina et al. 2011(Ratsoavina et al. , 2013(Ratsoavina et al. , 2015(Ratsoavina et al. , 2017 and in need of integrative studies. In order to contribute to a better picture of Madagascar's biodiversity, we here report the description of one of these candidate species of leaf-tailed gecko from Marojejy National Park. ...
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Article
The Marojejy Massif in northern Madagascar is a constant source of herpetological surprises. Herein we describe a new species of leaf-mimicking leaf-tailed gecko, Uroplatus finaritra sp. nov., based on morphological and molecular phylogenetic evidence. This new species inhabits the rainforests of Marojejy National Park at low elevations and is morphologically similar to Uroplatus phantasticus, but differs by having a larger body size, relatively shorter tail, and dark-red pigmentation of the oral mucosa. Molecular data reveals a distance of 14.0–14.7% in the 16S rRNA gene to U. phantasticus, and haplotype sharing in the c-mos nuclear gene only with species of much shorter tails (i.e., with U. kelirambo and two undescribed candidate species). Similar to other members of the leaf-mimicking Uroplatus species in northern Madagascar, Uroplatus finaritra sp. nov. likely has a small geographic distribution, and is currently known only from the lowland rainforests of Marojejy. We discuss the value of the buccal mucosa colouration and tail dimensions of Uroplatus as taxonomic characters, and the potential drivers of their evolution. In order to reduce the risk of international trade under incorrect species names we suggest that exported CITES species should be obligatorily accompanied by information about their precide geographic origin within Madagascar.
... For molecular genetic analysis, we follow the rationale of previous studies (e.g., Ratsoavina et al. 2011Ratsoavina et al. , 2015 in which we specifically searched for concordance between differentiation in mitochondrial and nuclear genes as support for the existence of independent evolutionary lineages with limited gene flow among them. While mtDNA due to its fourfold lower effective population size often shows a strong phylogeographic structure, this can be misleading because (1) it might reflect only a low vagility of females while males might ensure gene flow of nuclear genes among populations, and (2) mtDNA structure can be strongly affected by introgressive hybridization. ...
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The taxonomy of the Malagasy leaf-tailed geckos Uroplatus fimbriatus and U. giganteus is in need of revision since a molecular study casted doubt on the species status of U. giganteus from northern Madagascar. In this study we separately analyse DNA sequences of a mitochondrial gene (12S rRNA) and of four nuclear genes (CMOS, KIAA1239, RAG1, SACS), to test for concordant differentiation in these independent markers. In addition to the molecular data we provide a comprehensive review of colour variation of U. fimbriatus and U. giganteus populations from the entire distribution area based on photographs. The molecular evidence clearly supports a two-species taxonomy, with U. fimbriatus corresponding to a southern clade and U. giganteus to a northern clade. This conclusion relies on the high mitochondrial divergence among these units, and especially on the full concordance of the mitochondrial signal with differentiation in two of the four nuclear genes, which show no haplotype sharing among the northern and southern clade. This suggests limited or absent gene flow among these units, even in areas where they occur in rather close proximity to one another. The only consistent difference in colour variation between the two species is the whitish iris with rather indistinct brown vertical lines in U. giganteus, versus a yellowish iris with more distinct, continuous lines of typically a more reddish brown tone in U. fimbriatus. In the U. giganteus population of Montagne d’Ambre, a particular colour pattern on the head prevails which is not present in all the specimens studied from Marojejy. The Marojejy population of U. giganteus is also genetically distinct, as is one sample from Ankavanana included in a previous study, requiring in-depth future study. © 2018 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT), Mannheim, Germany.
... The leaf-tailed geckos of the genus Uroplatus are one such clade; their relationships and taxonomy have been assessed in various recent studies (Glaw et al. 2006;Greenbaum et al. 2007;Raxworthy et al. 2008b;Ratsoavina et al. 2012Ratsoavina et al. , 2013, and several candidate species await taxonomic revision and formal description. A subclade of small-bodied species, the Uroplatus ebenaui group, has the highest species diversity in the genus, with numerous new candidate species in northern Madagascar (Ratsoavina et al. 2011(Ratsoavina et al. , 2012(Ratsoavina et al. , 2013(Ratsoavina et al. , 2015. Species in the U. ebenaui group are characterized by the absence of dermal fringes, a triangular head and dermal spines on the head and especially the eyebrow. ...
... This group is currently composed of five nominal species of which U. ebenaui occurs in low-elevation forests of northern Madagascar, U. finiavana is restricted to mid-elevation rainforest of the Montagne d'Ambre massif in the North, U. fiera occurs in the Northern Central East, U. malama is restricted to the Southeast, and U. phantasticus is widespread in the Northern Central East and Southern Central East (Ratsoavina et al. 2011). Ratsoavina et al. (2013) identified five confirmed candidate species (Ca1, Ca2, Ca3, Ca4 and Ca7), as well as five unconfirmed candidate species (JX205421, EU596671, Ca6, Ca8 and Ca10) within the U. ebenaui group, and only one of these lineages (Ca7) has since been revised, and validated as new species U. fiera (Ratsoavina et al. 2015). Almost all candidate species enumerated in the U. ebenaui group show morphological affinities to the nominal taxon U. ebenaui, and most of them are restricted to the northern region of Madagascar. ...
... In this study we focus on revising two confirmed candidate species from the Tsaratanana massif previously named as Uroplatus ebenaui [Ca1] and [Ca2] (e.g., Ratsoavina et al. 2012Ratsoavina et al. , 2013Ratsoavina et al. , 2015. We complement the sampling of these lineages with newly collected material, provide expanded evidence from mitochondrial and nuclear genes as well as morphology for their status as independent evolutionary lineages, and provide their formal description as new species. ...
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The Tsaratanana Massif is the highest mountain massif of Madagascar and is characterized by a high species-level endemism of its biota. Here we describe two new small-sized species of leaf-tailed geckos of the Uroplatus ebenaui group from this region. Named in a preliminary way as confirmed candidate species Uroplatus ebenaui [Ca1] and [Ca2] in previous studies, we here provide detailed data on their molecular and morphological differentiation and describe them as Uroplatus fotsivava sp. nov. and Uroplatus kelirambo sp. nov. Their closest relatives are U. fiera and U. finiavana, respectively, as revealed by a phylogeny based on DNA sequences of four mitochondrial genes. However, integration of various lines of evidence confirms that the new species are independent evolutionary lineages, differing from other Uroplatus by high mitochondrial divergences, lack of haplotype sharing in the nuclear CMOS gene, and shape and relative size of the tail. While U. fotsivava is known from multiple sites at elevations between 1100–1538 m above sea level, U. kelirambo was found only at high elevations between 2000–2200 m a.s.l.; it therefore is the only known forest-dwelling nocturnal gecko reaching such high elevations in Madagascar, and might be narrowly endemic to montane forests of the Tsaratanana Massif.
... As a result, even many comparatively species-rich groups are now well studied in terms of their taxonomy, phylogeny, and biogeography. This is especially true for many gecko genera such as Phelsuma (Rocha et al. 2007Rocha et al. , 2009), Uroplatus (Greenbaum et al. 2007; Raxworthy et al. 2008; Ratsoavina et al. 2012 Ratsoavina et al. , 2015), and Paroedura (Jackman et al. 2008; ). In contrast , the genus Geckolepis, despite its apparent low diversity, is widely considered a taxonomically difficult group. ...
Article
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Fish scale geckos (Geckolepis) are taxonomically poorly resolved, mainly because of the difficulty of applying standard morphological characters to diagnose taxa. Three species, Geckolepis maculata, G. polylepis, and G. typica, are currently recognized from Madagascar and the Comoro Islands. Molecular studies suggested a number of operational taxonomical units within the G. maculata complex, but none of these has been formally described. The Comoran population was described as Geckolepis humbloti Vaillant 1887 but later synonymized. Prior to our study, no genetic data and little other information were available for this taxon. We revised the status of G. humbloti using molecular genetics, external morphology, and osteological characters retrieved from 3D skeletal models created using micro-computed tomography (micro-CT). Our results demonstrate that G. humbloti represents a genetic lineage strongly distinct from all other Geckolepis species. It is furthermore distinguished by a combination of external morphological characters and probably by osteology. We therefore resurrect G. humbloti Vaillant, 1887 from synonymy with G. maculata. Remarkably, this lineage is not restricted to the Comoros: A specimen from Tsingy de Bemaraha in western Madagascar falls as a closely related sister lineage to all Comoran Geckolepis in our molecular phylogenetic analysis and is osteologically almost identical with a specimen from the type locality Grand Comoro. We therefore include it in G. humbloti. The phylogenetic topology and the intraspecific genetic divergences suggest that the Comoros were colonized naturally from western Madagascar by overseas dispersal. G. humbloti is not considered as threatened, but its presence is indicative of natural or near-natural habitats.