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The Malagasy genus Gephyromantis contains 51 species of primarily terrestrial or scansorial frogs. Although many species are morphologically weakly divergent from each other, the combination of molecular and bioacoustic evidence has led to a continuous flow of species discoveries in the last years. Previous works have notably shown the existence of numerous additional deep mitochondrial lineages of uncertain status in the nominal subgenus Gephyromantis, some of these considered as confirmed or unconfirmed candidate species, some as deep conspecific lineages. Here we use DNA sequences of one mitochondrial and one nuclear marker, as well as morphological and bioacoustic data, to conduct an integrative revision of the subgenus Gephyromantis. The analyses reveal at least 12 distinct and independent evolutionary lineages belonging to the G. blanci and G. boulengeri species complexes. Evidence for the species status of these lineages included multiple cases of syntopic occurrence without genetic admixture, as well as differences in advertisement calls or morphological differentiation without intermediate forms, suggesting reproductive isolation. We discuss the relevance of these different lines of evidence and describe six new species of Gephyromantis., The work of AM was supported by the Deutsche Forschungsgemeinschaft (grant MI 2748/1-1) and the work of AC was supported by the Portuguese National Funds through FCT (Fundação para a Ciência e a Tecnologia, contract 2020.00823. CEECIND/CP1601/CT0003).

Based on sequences of three mitochondrial and two nuclear-encoded genes, we examine genetic variation in the leaf tailed geckos Uroplatus sameiti and U. sikorae, and morphological and chromatic characteristics of the genetic clusters identified. The mitochondrial phylogeny reveals a puzzling diversity of 16 deep lineages (4 in U. sameiti and 12 in U. sikorae) differing by 2.9–9.9% uncorrected pairwise distance in a fragment of the 16S rRNA gene. Populations from Analalava in the North and Zahamena in the Northern Central East form two mitochondrial lineages clustering with U. sameiti but being deeply divergent (>8% 16S distance); however, no information on their morphology is available. In U. sikorae, the mitochondrial lineages identified form several major geographic clades, two of which (from the northernmost and southernmost populations, respectively) received substantial support in the phylogenetic analysis. No instance of sympatry of two or more mitochondrial lineages was observed, precluding an unambiguous assessment of species status under the biological species criterion without experimental approaches or detailed hybrid zone analyses. In the fragment of the nuclear encoded gene SACS we observed haplotype sharing between species and mitochondrial lineages, while in the fragment of KIAA1239 no haplotype sharing was detected although neither species nor mitochondrial lineages formed coherent phylogroups in the respective network. A screening of colour patterns from live photos, partly of the genotyped individuals, confirmed a large variation within species and populations, with a possible sexual dichromatism where a longitudinally striped phenotype is restricted to males. All individuals from populations of the U. sikorae clade from the Southern Central East and South East had an unpigmented oral mucosa just like U. sameiti, while all other U. sikorae populations are characterized by a black oral mucosa pigmentation. The extremely strong phylogeographic structure in the U. sikorae complex without obvious species-level divergences is unprecedented for large-sized squamates in Madagascar and calls for further taxonomic scrutiny using phylogenomic approaches; and it exemplifies how the loss of any major block of the remaining rainforests in Madagascar will inevitably lead to a substantial loss of genetic diversity – even if often intraspecific – in rainforest-specialized species.

Lygodactylus guibei Pasteur, 1965 Lygodactylus (Domerguella) guibei Pasteur, 1965 Partial chresonymy Lygodactylus guibei: Kluge (1991); Glaw & Vences (1992, 1994, 2007); Puente et al. (2005, 2009); Röll et al. (2010); Gippner et al. (2021) Lygodactylus (Domerguella) guibei: Rösler (2000b). Name-bearing type: holotype MNHN 1993.60 from “Périnet (Est)” (=Andasibe), according to the original description.—Other types: According to the original description, there were two paratypes; we only were able to locate MNHN 1933.156.—Etymology: Eponym for Jean Guibé. Identity and Diagnosis. The holotype agrees well morphologically with most other individuals assigned to this species by relatively low longitudinal counts of dorsal scales (L. miops have higher counts (>200 />100). Despite some overlap in these variables, the differences between the two lineages seem to allow a distinction of most individuals. Furthermore, L. guibei does not reach the high INFL and NNS counts of some L. miops individuals, reaches larger body sizes, and males are characterized by more distinct lateral tubercles at the base of the tail, judging from the specimens morphologically examined herein. Specimens appear to have a rather indistinct dorsal pattern (Fig. 14). Two photographed individuals have a conspicuous stripe-like row of dark spots on the chest (Fig. 14C, E) but this pattern is absent in most other individuals examined. Distribution. L. guibei is known from several localities in the Northern Central East of Madagascar: (1) the type locality Andasibe, (2) Vohidrazana, (3) Moramanga, (4) Anjozorobe, (5) Mahasoa Forest (based on ND4 sequences of Gippner et al. 2021), and (6) Angozongahy on the west slope of the Makira Reserve., Published as part of Vences, Miguel, Multzsch, Malte, Gippner, Sven, Miralles, Aurélien, Crottini, Angelica, Gehring, Philip-Sebastian, Rakotoarison, Andolalao, Ratsoavina, Fanomezana M., Glaw, Frank & Scherz, Mark D., 2022, Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar's rainforest, pp. 1-61 in Zootaxa 5179 (1) on pages 23-25, DOI: 10.11646/zootaxa.5179.1.1, http://zenodo.org/record/7040745, {"references":["Kluge, A. G. (1991) Checklist of Gekkonoid Lizards. Smithsonian Herpetological Information Service 85, 36 pp. https: // doi. org / 10.5479 / si. 23317515.85.1","Glaw, F. & Vences, M. (1992) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 331 pp. [First Edition.]","Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 480 pp. [Second Edition.]","Puente, M., Thomas, M. & Vences, M. (2005) Phylogeny and biogeography of Malagasy dwarf geckos, Lygodactylus Gray, 1864: Preliminary data from mitochondrial DNA sequences (Squamata: Gekkonidae). In: Huber, B. A. & Lampe, K. H. (Eds.), African Biodiversity: Molecules, Organisms, Ecosystems. Proc. 5 th Intern. Symp. Trop. Biol., Museum Koenig, Bonn. Springer, pp. 229 - 235. https: // doi. org / 10.1007 / 0 - 387 - 24320 - 8 _ 21","Puente, M., Glaw, F., Vieites, D. R. & 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","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. & 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","Rosler, H. (2000 b) Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28 - 153."]}

Lygodactylus miops Günther, 1891 Lygodactylus miops Günther, 1891 Synonyms Microscalabotes spinulifer Boettger, 1913 Lygodactylus septemtuberculatus Angel, 1942 Chresonyms: Lygodactylus septemtuberculatus: Kluge (1991) Lygodactylus (Domerguella) miops: Pasteur (1965a) Lygodactylus (Lygodactylus) septemtuberculatus: Rösler (2000b) Lygodactylus miops: Kluge (1991); Glaw & Vences (1992, 1994, 2007); Puente et al. (2005, 2009); Röll et al. (2010); Gippner et al. (2021) Name-bearing type: holotype, BMNH 1946.8.22.55, female.—Type locality: “Senbendrana”, Madagascar according to the original description (probably referring to Sahembendrana; see Blommers-Schlösser & Blanc 1991).— Other types: none according to original description.— Etymology: From Latin (originally Greek) miops = short sighted and probably referring to the large eyes of the species highlighted in the original description. Identity and Diagnosis. Our data show the presence of five genetic lineages in the general area of the Northern Central East of Madagascar whence L. miops has been described. These include the lineages commonly named L. guibei and L. miops, as well as the candidate species L. sp. 11, L. sp. 19, and L. sp. 20, all belonging to subclade A5. Of these lineages, no material for morphological examination was available for L. sp. 19 and L. sp. 20. At the same time, there are four historical nomina described from this general region, all without genetic data for the name-bearing types: Lygodactylus miops Günther, 1891; Microscalabotes spinulifer Boettger, 1913; Lygodactylus septemtuberculatus Angel, 1942; Lygodactylus guibei Pasteur, 1965a. We here continue to define the relatively small-sized lineage that is widespread mostly in low elevations along much of Madagascar’s east coast as L. miops (as in Puente et al. 2009), based on the following rationale: (i) several of the specimens genetically assigned to this lineage share with the L. miops holotype a high count of infralabial scales (INFL = 8), which is not observed in specimens assigned to L. guibei (INFL = 6 or 7); (ii) the count of internasal scales (IN = 3) of the L. miops holotype is higher than in any specimen assigned to L. guibei (IN = 1 or 2) but is found in two other individuals of this genetic lineage; (iii) with an SVL of 29.9 mm (according to our own, new measurements) the holotype fits well the size range of other individuals usually assigned to L. miops (27.2–31.2 mm), while several specimens of L. guibei reach SVLs between 34.0– 39.5 mm; (iv) most importantly, the longitudinal counts of dorsal and ventral scales are larger than in all individuals assigned to L. guibei (LCDS 233 vs. 170–220; LCVS 113 vs. 87–109), and agree with those of other specimens usually assigned to L. miops (LCDS 205–242, LCVS 98–113); (v) the tail base tubercles are distinct and medium-sized as in other males usually assigned to L. miops; (vi) finally, the L. miops holotype has a distinct pattern with light dorsolateral bands (already mentioned in the original description), which is rarely found in subclade A5 but observed in genotyped individuals from Betampona (e.g. Fig. 13C). The L. miops holotype also differs from the sole voucher specimen of L. sp. 11 available for morphological examination by a higher number of dorsal tubercles, lower dorsal scale count, higher ventral scale count, and more distinct tubercles at tail base. We here consider L. sp. 11 as a distinct species, L. fritzi sp. nov., and provide additional comparisons and justifications (including a detailed discussion of the L. miops type locality) in the diagnosis of that species below. However, based on the available data we cannot fully exclude that the L. miops holotype is conspecific with L. sp. 19 or L. sp. 20 for which no morphological data are available. Synonyms. We consider two nomina to be synonyms of L. miops, in agreement with current taxonomy: Microscalabotes spinulifer Boettger, 1913 with the lectotype (designated by Mertens 1967) SMF 8931, collected by F. Sikora at Moramanga; and Lygodactylus septemtuberculatus Angel, 1942 with the holotype MNHN 1893.63 as well from Moramanga. Both these nomina agree with the lineage here considered to represent L. miops by their relatively high longitudinal counts of dorsal and ventral scales (LCDS 240 (spinulifer) and 225 (septemtuberculatus); LCVS 107 and 102, vs. LCDS 205–242 and LCVS 98–113 for specimens assigned to L. miops; Table 1), and relatively small body size (28.5 and 29.0 mm, vs. 27.2–31.2 mm for specimens assigned to L. miops; Table 1). The same morphological characters are also found in L. sp. 11, but this lineage is known from coastal localities and has not been found in or nearby Moramanga so far. Natural history. In Betampona this species is very common and can be found both in disturbed areas and in densely forested habitat. Here the species is often found in the leaflitter, on twigs or along the partially aerial roots of larger trees. This species generally roosts on the leaves of small bushes (including the invasive strawberry guava). Distribution. L. miops as understood here is one of the most widespread species of the L. madagascariensis group, occurring in multiple localities along the eastern coast of Madagascar, which encompass the regions South East, Southern Central East, Northern Central East, and North East. It is known from (1) the type locality Senbendrana (=Sahembendrana or Sahambendrana? For a detailed discussion of this locality, see the account of L. fritzi sp. nov. below), and the type locality of its two synonyms, (2) Moramanga. Furthermore, genetically verified records (in a south–north direction) originate from (3) Manantantely, (4) Andohahela, (5) a site north of Andohahela, (6) Sainte Luce, (7) Sampanandrano, (8) Tsitongambarika, (9) Ranomafana, (10) Ambohitsara, (11) Mahakajy, (12) Anosibe Anala, (13) Vohimana, (14) Sahafina, (15) Betampona, (16) Makira (Ambodivoahangy).., Published as part of Vences, Miguel, Multzsch, Malte, Gippner, Sven, Miralles, Aurélien, Crottini, Angelica, Gehring, Philip-Sebastian, Rakotoarison, Andolalao, Ratsoavina, Fanomezana M., Glaw, Frank & Scherz, Mark D., 2022, Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar's rainforest, pp. 1-61 in Zootaxa 5179 (1) on pages 21-23, DOI: 10.11646/zootaxa.5179.1.1, http://zenodo.org/record/7040745, {"references":["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","Boettger, O. (1913) Reptilien und Amphibien von Madagascar, den Inseln und dem Festland Ostafrikas. Voeltzkow, A., Reise in Ostafrika, 3 (4), 1 - 269.","Angel, F. (1942) Les lezards de Madagascar. Memoires de l'Academie Malgache, 36, 1 - 139.","Kluge, A. G. (1991) Checklist of Gekkonoid Lizards. Smithsonian Herpetological Information Service 85, 36 pp. https: // doi. org / 10.5479 / si. 23317515.85.1","Pasteur, G. (1965 a) Notes preliminaries sur les lygodactyles (Gekkonides). IV Diagnoses de quelques formes africaines et malgaches. Bulletin du Museum national d'Histoire naturelle, 36, 311 - 314.","Rosler, H. (2000 b) Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28 - 153.","Glaw, F. & Vences, M. (1992) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 331 pp. [First Edition.]","Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 480 pp. [Second Edition.]","Puente, M., Thomas, M. & Vences, M. (2005) Phylogeny and biogeography of Malagasy dwarf geckos, Lygodactylus Gray, 1864: Preliminary data from mitochondrial DNA sequences (Squamata: Gekkonidae). In: Huber, B. A. & Lampe, K. H. (Eds.), African Biodiversity: Molecules, Organisms, Ecosystems. Proc. 5 th Intern. Symp. Trop. Biol., Museum Koenig, Bonn. Springer, pp. 229 - 235. https: // doi. org / 10.1007 / 0 - 387 - 24320 - 8 _ 21","Puente, M., Glaw, F., Vieites, D. R. & 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","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. & 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","Blommers-Schlosser, R. M. A. & Blanc, C. P. (1991) Amphibiens (premiere partie). Faune de Madagascar, 75 (1), 1 - 379.","Mertens, R. (1967) Die herpetologische Sektion des Naturmuseums und Forschungsinstitutes Senckenberg in Frankfurt a. M. nebst einem Verzeichnis ihrer Typen. Senckenbergiana Biologica, 48, Sonderheft A, 1 - 106."]}

Lygodactylus roellae sp. nov. Lygodactylus sp. 17: Gippner et al. (2021). Holotype. ZSM 49 /2016 (MSZC 0072), adult female, collected by M.D. Scherz, J. Borrell, L. Ball, T. Starnes, E. Razafimandimby, D.H. Nomenjanahary, and J. Rabearivony at Ampotsidy mountains, 15.7 km NNW of Bealanana (8.7 km NNW of Beandrarezona), northern Madagascar, at geographical coordinates S14.41974, E48.71935, 1344 m a.s.l., on 22 December 2015 (Fig. 15). Paratypes. ZSM 556 /2014 (DRV 6289), adult male, collected by F.M. Ratsoavina, D. Vieites, M. Vences, R. D. Randrianiaina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, and T. Rajoafiarison at Andrevorevo, a site south of the Tsaratanana Massif, northern Madagascar, at geographical coordinates S14.3464, E49.1028, 1717 m a.s.l., on 21 June 2010; UADBA-R 70855 (MSZC 0010), adult female, with the same collection data as the holotype but collected at S14.41878 E48.71896, 1354 m a.s.l. on 18 December 2015 at 20h20. Diagnosis. Lygodactylus roellae sp. nov. corresponds to a genetically highly distinct lineage from northern Madagascar that is the sister species of L. salvi described above, but differs by high genetic divergence and several scale counts. It belongs to subclade A3 within Domerguella as defined herein. It can also be assigned to the subgenus Domerguella by an undivided mental scale with two postmentals, absence of a claw on the first finger, and 5 preanal pores in males. Within Domerguella, the new species is only known from two localities in northern Madagascar and differs from the other nominal species of Domerguella occurring in this part of the island as follows: from L. expectatus by a different color pattern, without scapular semi-ocellus and with a striped pattern apparently in most individuals (vs. scapular semi-ocellus usually present, and striped pattern unknown); from L. rarus by lack of regular crossbands on tail (vs. presence) and different body shape without elongated limbs (relative hindlimb length 0.41–0.45 vs.>0.55); from L. madagascariensis, L. petteri, L. salvi, and L. tantsaha by smaller longitudinal dorsal scale count (159–169 vs.>188) and smaller longitudinal ventral scale count (83–92 vs.>96). Genetically, the new species is highly distinct from all species in subclade A5, and differs at least from L. guibei by a less distinctly expressed lateral spine at the tail base of males (vs. presence of a distinct, large spine). Furthermore, the longitudinal dorsal scale count is smaller than in all known individuals of subclade A5. The new species differs from its sister lineage, L. salvi (described above), by a lower longitudinal dorsal scale count (159–169 vs. 211–217) and a lower ventral scale count (83–92 vs. 107–112). The two sister species also differ by a high genetic divergence of 11.2–12.6% in the 16S gene, and do not share haplotypes in RAG1 despite occurring in geographical proximity. For a distinction from additional species newly named and described herein, see the respective diagnoses below. Etymology. We are pleased to dedicate this beautiful gecko species to Beate Röll, in recognition for her substantial contributions to Lygodactylus biology and phylogeny. The name is a matronym (i.e., a noun in the genitive case). Description of the holotype. Adult female, in a good state of preservation, the right hind limb is partly removed as a source of tissue for molecular analysis (Fig. 15). SVL 35.9 mm, TAL 39.6 mm; for other measurements see Table 1. Head and neck thick, body broader than head. The distance from the tip of the snout to the anterior border of the eye (3.8 mm) is less than the interorbital distance anteriorly (4.2 mm), 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 five scales: rostral, first supralabial, and three supranasals. Mental scale undivided; no contact between posterior projection of mental scale and first infralabial; two asymmetrical postmental scale with five postpostmental scales; seven infralabial scales; seven supralabial scales; two internasal scales; granular dorsal scales; dorsum with small, homogeneous, granular, and unkeeled scales of similar size to those on trunk, smaller than on head and tail, the scales on limbs can be slightly larger; 169 dorsal scales longitudinal along the body; 92 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; no dorsolateral tubercles; tail without whorls; small lateral spines at the base of the tail. Based on available photographs (Fig. 18A–B), the life coloration of the holotype exhibited a distinct pattern of dark brownish to yellowish stripes on the dorsum reaching from the eye to base of the tail. The stripes continue in an irregular pattern of more elongated dark and brighter spots on the caudal spine. The dark wide area on the back is slightly emarginated at the level of the hindlimbs before ending at the base of the tail. The flanks and limbs are gray to brownish (Fig. 18A). The ventral side is whitish with few small brown spots predominately on the throat and the tail (Fig. 18B). The specimen has darkened after 6 years of preservation in ethanol; however, the striped pattern is still distinctly visible. Variation. The coloration of the dorsum is characteristic with a distinct pattern of dark brownish to yellowish stripes on the dorsum with a variable strength of expression on the tail (Fig. 18A and 18E), which appears to weaken greatly or be lost upon regeneration (Fig. 18C). ......continued on the next page TABLE 1. (Continued) ......continued on the next page TABLE 1. (Continued) ......continued on the next page TABLE 1. (Continued) ......continued on the next page TABLE 1. (Continued) ......continued on the next page TABLE 1. (Continued) The examined male paratype specimen (ZSM 556/2014) is almost the same size as the holotype with a SVL of 36.0 mm and has a longer tail (45.3 mm) and hindlimbs (HIL/SVL 0.45). On the fourth toe it has four instead of three subdigital lamellae, which differs from all other examined specimens. Unlike the holotype it has tubercles between the limbs (6) that consist of one scale each. It has fewer dorsal (159) and ventral scales (83). These differences could be due to different sex or just random variation. Natural history. Specimens of this species were collected sleeping at night on roosts up to 1 m above the ground in Ampotsidy. UADBA-R 70855 was found sleeping on the tip of a Pandanus frond. It occurs in close sympatry with Lygodactylus winki sp. nov., described below. Distribution. L. roellae is known from (1) the type locality Ampotsidy and (2) Andrevorevo.

Lygodactylus ulli sp. nov. Lygodactylus sp. 21: Gippner et al. (2021) Holotype. ZSM 154 /2005 (FGZC 2811), adult male, collected by F. Glaw, M. Vences, and R. D. Randrianiaina at Marojejy National Park, at Camp 1 “Mantella”, North-East of Madagascar, geographical coordinates S14.4377, E49.7756, 481 m a.s.l., on 14 February 2005 (Fig. 15). Referred material. UADBA-R uncatalogued (MSZC 0272), female, collected by M.D. Scherz, J. Razafindraibe, and A. Razafimanantsoa at the same locality as the holotype, at night on 23 November 2016. Diagnosis. Lygodactylus ulli sp. nov. corresponds to a lineage forming part of subclade A5 of Domerguella, and is the second representative of this subclade reaching northern Madagascar. It is characterized by the presence of distinct lateral spine-like scales at the base of the tail in males, as is found in several representatives of A5 from eastern Madagascar but not in other Domerguella. The smallest genetic distances are 10.7% uncorrected 16S distance to specimens of L. guibei. It can also be assigned to the subgenus Domerguella by an undivided mental scale with two postmentals, absence of a claw on the first finger, and 7 preanal pores in males. Within Domerguella, the new species (together with L. winki) is one of only two species of subclade A5 known from northern Madagascar, and the only species of Domerguella so far known from the rainforests of the Marojejy Massif. It differs from the nominal species of Domerguella occurring in northern Madagascar and belonging to subclades A1–A4 as follows: from L. expectatus by non-enlarged dorsolateral scales (longitudinal count of dorsal scales>250 vs. L. rarus by lack of regular crossbands on tail (vs. presence) and different body shape with less elongated limbs (relative hindlimb length 0.47 vs.>0.55); from L. petteri, L. tantsaha, L. salvi, L. roellae, and L. hapei by a higher longitudinal count of dorsal scales (253 vs. L. madagascariensis as well as most of the previously mentioned species by a small but distinct spine-like tubercle at the base of the tail in males. From the other nominal species in subclade A5 (L. miops, and L. guibei and L. winki) the new species differs by higher longitudinal counts of dorsal scales (253 vs. 170–242) and ventral scales (110 vs. 87–109). We did not detect haplotype sharing in RAG1 or CMOS between L. ulli and the other nominal species in subclade A5 (L. miops, and L. guibei and L. winki). Haplotpe sharing was detected only at the CMOS marker with L. sp. 20. For a distinction from additional species newly named and described herein, see the respective diagnoses below. Etymology. We are pleased to dedicate this species to Ulrich “Ulli” Joger, director emeritus of the Braunschweig Natural History Museum, in recognition of his contribution to the taxonomy of reptiles, especially geckos. The species epithet name is defined as a noun in apposition (not a noun in the genitive case) to avoid ending with a non-euphonious double-i. Description of the holotype. Adult male, hemipenes everted, in moderate state of preservation, tail is broken and missing, right forelimb is removed as source of tissue for molecular analysis (Fig. 15). SVL 28.8 mm, TAL 2.6 mm; for other measurements see Table 1. Head broader than body. The distance from the tip of the snout to the anterior border of the eye (3.5 mm) is lesser than the interorbital distance anteriorly (3.9 mm), 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 five scales: rostral, first supralabial, and three supranasals. Mental scale undivided; no contact between posterior projection of mental scale and first infralabial; three asymmetrical postmental scales with five postpostmental scales; six infralabial scales; seven 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; 253 dorsal scales longitudinally along the body; 110 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; seven preanal pores; tail without whorls; small lateral spines at the base of the tail. Based on available photographs (Fig. 21), the live holotype displays a brownish to grayish pattern on the dorsum and the limbs. From the snout, a narrow black stripe runs irregularly to two elongated spots on the shoulder. At the forelimb level two symmetrical black spots are present on the spine (Fig. 21A). After 16 years in ethanol, the preserved specimen is more grayish. The ventral side is whitish. While there are only a few small brown spots on the venter, multiple larger brown spots are irregularly scattered on the throat. Variation. Comparing the two photographed specimens, dorsal coloration and patterns are less pronounced on the male (Fig. 21A), maybe because it appears to be close to skin shedding. The female has a blackish and grayish alternating dorsal pattern, surrounded by a yellowish base layer (Fig. 21B +C). The female is laterally darker than the male (Fig. 21A +B), has a white throat and yellow venter and ventral tail. Natural history. Specimens were collected in primary rainforest. Distribution. L. ulli is only known from its type locality, the Marojejy Massif in the North East region of Madagascar., Published as part of Vences, Miguel, Multzsch, Malte, Gippner, Sven, Miralles, Aurélien, Crottini, Angelica, Gehring, Philip-Sebastian, Rakotoarison, Andolalao, Ratsoavina, Fanomezana M., Glaw, Frank & Scherz, Mark D., 2022, Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar's rainforest, pp. 1-61 in Zootaxa 5179 (1) on pages 44-46, DOI: 10.11646/zootaxa.5179.1.1, http://zenodo.org/record/7040745, {"references":["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. & 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"]}

Lygodactylus expectatus Pasteur & Blanc, 1967 Lygodactylus (Domerguella) expectatus Pasteur & Blanc, 1967 Chresonyms: Lygodactylus expectatus: Kluge (1991); Glaw & Vences (1992, 1994, 2007); Puente et al. (2009); Röll et al. (2010); Gippner et al. (2021) Lygodactylus (Domerguella) expectatus: Rösler (2000b) Name-bearing type: male holotype MNHN 1990.1 (original number BP 640).—Type locality: “Karst d’Ambilobe (Ankarana), à une douzaine de kilomètres au NNW de cette localité”, according to the original description.—Other types: according to the original description, five specimens were examined but explicitly only two of these were designated as paratypes, namely MNHN 1990.2 – 3 (BP 641, female, and BP 642, young female, according to original description).—Etymology: From Latin expectatus = expected. As explained in the original description, G. Pasteur and C.P. Blanc were expecting to find a new species in the karstic regions of the Ambilobe region. Identity and Diagnosis. According to the diagnosis given by Puente et al. (2009), the species differed from all species in the L. madagascariensis group known at the time by its dorsolateral scales, which are enlarged relative to the dorsal and lateral scales (not distinctly enlarged in the other species), and by the presence of two dark spots in the region of the neck (not distinct in the other species). The enlarged scales in the dorsolateral region, contrasting with the very small scales in the vertebral region, indeed represent a diagnostic character of this species that we could not observe in any other species of Domerguella (Fig. 6). This typical character state of L. expectatus is visible in all of the genetically characterized specimens collected, as well as the holotype (examined in June 2021, in relatively poor state of preservation). It is also reflected by a low longitudinal count of dorsal scales, of 130 scales or less if counting the enlarged scales (slightly more, with a maximum of 164, if counting the small vertebral scales, but also this value is still smaller than in all other nominal Domerguella, overlapping with only one candidate species, L. sp. 17). The dark spot in the region of the neck is located anterodorsal to the forelimb region, roughly in the scapular region, and we here name it the scapular semi-ocellus, considering that it is bordered by a whitish row of tubercles dorsally, giving the impression of an ocellus but lacking a ventral light lining. This semi-ocellus is typical for L. expectatus, but sometimes weakly expressed, and in such cases easy to confuse with dark lateral markings that can also be seen in other species of Domerguella, but often in slightly different positions (Figs. 7–8). Given these two diagnostic character states, which both have been verified in the holotype and in the genotyped specimens, along with the provenance of all these specimens from the Ankarana Massif, there is no doubt about the correct attribution of our specimens to L. expectatus. The species is rather small sized, with adult SVL 24.3–29.7 mm vs. a maximum size larger than 30 mm in several other species. There are no dorsolateral tubercles and no spiny tubercles at the tail base as they are characteristic for several other Domerguella, and no distinct, regular broad crossbands on the tail as in L. rarus (see below). According to the available counts, the species has 87–98 ventral scales longitudinally. Distribution. L. expectatus is only known from its type locality, the Ankarana Massif. According to the original description (Pasteur & Blanc 1967), additional specimens also came from “Ambilobé” and from “Region de DiégoSuarez”, but we have not verified the identity of the respective vouchers, and the localities are not precise enough for firmly concluding they are not in the Ankarana Massif (which is geographically located inbetween the towns of Ambilobe and Antsiranana (=Diego-Suarez).

Lygodactylus fritzi sp. nov. Lygodactylus sp. 11: Gippner et al. (2021). Justification. This new species from coastal areas in Madagascar’s Northern Central East has previously been called L. sp. 11 in Gippner et al. (2021). It corresponds to a lineage forming part of subclade A5 of Domerguella, and occurs in a general area where L. miops 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 Domerguella. The smallest genetic distance is 9.0% to specimens assigned to L. miops. 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 holotype of L. miops, and of the types of the other earlier names septemtuberculatus and spinulifer, to either of them requires further justification (see also the account of L. miops). The following arguments support our decision to assign the holotype of miops, septemtuberculatus and spinulifer to what we will in this section call the “widespread lowland lineage” rather than to L. sp. 11: (1) The specimens of the widespread lowland lineage (including the three historical types) differ from L. sp. 11 by a 3–6 vs. 2 dorsolateral tubercles between limbs; dorsal scale count 207–242 vs. 247; ventral scale count 102–113 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 L. sp. 11 as morphologically distinct, and the three type specimens of the earlier names as better fitting the widespread eastern lineage. (2) L. sp. 11 is only known from two low-elevation localities near sea level (0–20 m 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 L. sp. 11 were collected historically, and that its range extends into the type localities of the three historical nomina, especially up to Moramanga (the type locality of septemtuberculatus and spinulifer). (3) The type locality of L. miops, “Senbendrana” according to the original description (Günther 1891), cannot be located reliably at present. Senbendrana has been reported with the addition “near Tamatave” (= Toamasina) as a collecting locality of spiders (Pocock 1895); or as corresponding to Sahembendrana or Sahambendrana. This latter synonymy is supported by the fact that the type of L. miops was provided by “Majastre” (see Puente et al. 2009), probably corresponding to A. Majastre, a collector who provided specimens of many animals and plants from this area. Some of Majastre’s collections are labelled “Sahambendrana”, e.g., the type of the orchid Eulophia grandibracteata (see Schultz 2013). The locality apparently was often misspelled; for example, we assume that “Sen Bendrana” (Michaelsen 1891), “Senbendra” (Sharp & Ogilvie-Grant 1898), or “Schambendrama” (Bott 1963) refer to the same site as well. Blommers-Schlösser & Blanc (1991) located Sahembendrana close to Akkoraka (at higher elevations in eastern Madagascar), but it is likely that Majastre’s collecting site was situated closer to Toamasina. We could not locate current or historical maps mentioning a site with this or a similar name, but Ramananjara (2009) documents the sale of a property in 1931, in the “Canton d’Antetezambaro; sur la rivière Sahambendrana”, and more specifically “au sud d’Ambodisatrana”, which likely refers to a coastal village about 30 km north of Toamasina that can be located in historical maps from 1934 (Service Géographique de Madagascar, map “Fénérive”, 1/500,000). On the other hand, Rosa et al. (2012) report a campsite from Betampona Reserve (about 35 km north-east of Toamasina) locally known as Sahambendrana, at coordinates S17.8984, E49.2154, 458 m a.s.l. Other sources refer to a Sahambendrana river on the northern versant of Betampona (Randriatavy 2003; Randrianarimanana 2009). Whether any of these sites corresponds to Majastre’s 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 Toamasina. The available evidence, however, points to the original collecting site being not directly at sea level. According to Günther (1891), the same collection that included the L. miops holotype also contained “ Rhacophorus luteus ”, which almost certainly corresponds to a treefrog species of the Boophis luteus group, which is not known from coastal sites in Madagascar (but known to be present in Betampona), and for instance has not been collected at Vohibola or Ankanin’ny Nofy (Gehring et al. 2010) where L. sp. 11 occurs. Furthermore, Pellegrin (1933) reported fish specimens of the genus Sicyopterus from a “rivière Sahembendrana (région de Tamatave)”. These specimens were identified by Sparks & Nelson (2004) as S. franouxi, 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 L. sp. 11. The fact that Sahambendrana / Sahembendrana has on various occasions been used to refer to a river (e.g., Pellegrin 1993; Randriatavy 2003; Randrianarimanana 2009; Ramananjara 2009) allows for the possibility of an upstream collecting site of the L. miops holotype, 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 Domerguella lineage. In summary, the available evidence thus suggests that none of the earlier available names miops, septemtuberculatus, or spinulifer is likely to apply to L. sp. 11, which we therefore formally name as species new to science, L. fritzi sp. nov. Holotype. ZSM 651 /2009 (ZCMV 8902), female, collected by P.-S. Gehring, F. Ratsoavina, and E. Rajeriarison at Ankanin’ny Nofy, east coast of Madagascar, geographical coordinates - S18.6058, E49.2138, roughly at sea level, on 8 April 2009. Diagnosis. Lygodactylus fritzi sp. nov. is a species of the Lygodactylus subgenus Domerguella based on molecular phylogenetic relationships, and it can also be assigned to the subgenus Domerguella by an undivided mental scale with two postmentals, absence of a claw on the first finger, and 7 preanal pores in males. Within Domerguella, the new species is one of several species of subclade A5 known from the Northern Central East of Madagascar. It differs from the nominal species of Domerguella occurring in northern Madagascar and belonging to subclades A1–A4 as follows: from L. expectatus by non-enlarged dorsolateral scales (longitudinal count of dorsal scales>250 vs. L. rarus by lack of regular crossbands on tail (vs. presence) and different body shape with less elongated limbs (relative hindlimb length 0.47 vs.>0.55); from L. petteri, L. tantsaha, L. salvi, L. roellae, and L. hapei by a higher longitudinal count of dorsal scales (253 vs. L. madagascariensis as well as most of the previously mentioned species of clades A1‒A4 by a rudimentary spine-like tubercle at the base of the tail in the only known female (vs. absence). From other species of subclade A5, the new species differs as follows: from L. miops by fewer dorsolateral tubercles between limbs (2 vs. 3–6), higher longitudinal dorsal scale count (247 vs. 207–242), lower longitudinal ventral scale count (98 vs. 102–113), and weakly expressed lateral tubercles at tail base vs. clearly recognizable in males and females; from L. guibei by higher longitudinal dorsal scale count (247 vs. 170–220), and weakly expressed lateral tubercles at tail base vs. clearly recognizable, usually large in males and females; from L. winki by fewer dorsolateral tubercles between limbs (2 vs. 5–8), higher longitudinal dorsal scale count (247 vs. 187–222), and weakly expressed lateral tubercles at tail base vs. clearly recognizable usually large in males and females; from L. ulli 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. For a distinction from one other species newly named and described in the following, see the respective diagnosis below. Etymology. 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). Description of the holotype. Female in a good state of preservation, tail partly detached. SVL 26.4 mm, TAL 26.5 mm; for other measurements see Table 1. Body broader than head. The distance from the tip of the snout to the anterior border of the eye (3.1 mm), is less than the interorbital distance anteriorly (3.5 mm), 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. Coloration of the holotype 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. Variation. Morphometric and meristic data are only known from a single voucher specimen. However, color patterns could be assessed from photographs of three additional individuals (Fig. 22), some of which were dorsally gray-beige with an irregular contrasted pattern of larger light and smaller dark spots (Fig. 22A), others more uniform with a more or less symmetrical pattern of dark spots (Fig. 22E), or dark brown with weakly contrasted light brown dorsolateral bands with a somewhat reddish tone (Fig. 22B–C). Ventrally irregularly dark spotted (Fig. 22D). Distribution. L. fritzi is only known from (1) the type locality Ankanin’ny Nofy and (2) Vohibola, two coastal lowland localities (0–20 m a.s.l.) in the Northern Central East of Madagascar. Littoral forest harbours a high species richness especially of plants, with several genera endemic to this habitat (de Gouvenain & Silander 2003; Bollen & Donati 2005). Fisher & Girman (2000) identified littoral forests as one of four major areas for ant endemism in Madagascar. Previous studies in south-eastern littoral forests found no vertebrate species strictly endemic to that forest type (Ganzhorn et al. 2000; Goodman & Ramanamanjato 2007) but recent taxonomic revisions have revealed numerous amphibians and reptiles restricted to small areas of forest directly adjacent to the Madagascar’s coast, such as for example species of the miniaturized frog genus Mini (Scherz et al. 2019) or Pandanus-dwelling frogs of Guibemantis (Lehtinen et al. 2011), or the chameleon Calumma vohibola (Gehring et al. 2011), Lygodactylus fritzi adds to this growing list of species specialized to these highly threatened coastal forests. Natural history. One adult specimen was photographed millimeters from the posterior end of a bug (Fig. 22), and was presumably consuming honeydew excreted by the insect, as is known from gecko species (Fölling et al. 2001).

Lygodactylus winki sp. nov. Lygodactylus sp. 18— Gippner et al. (2021). Holotype. ZSM 47 /2016 (MSZC 0075), adult male, collected by M.D. Scherz, J. Borrell, L. Ball, T. Starnes, E. Razafimandimby, D.H. Nomenjanahary, and J. Rabearivony, at Ampotsidy, 15.7 km NNW of Bealanana (8.7 km NNW of Beandrarezona), northern Madagascar, at geographical coordinates S14.41900, E48.71883, 1364 m a.s.l., on 22 December 2015 (Fig. 15). Paratypes. ZSM 48 /2016 (MSZC 0110), adult male, collected by same collectors and at same locality as holotype, at geographical coordinates S14.42843, E48.72285, 1315 m a.s.l., on 29 December 2015; UADBA-R 70856– 70859 (MSZC 0011, 0019, 0023, 0077), two males and two females, respectively, collected by same collectors and at same locality as holotype, between coordinates S14.41455 –14.42317, E48.71149 –48.71916, 1320–1404 m a.s.l., on 18–22 December 2015; ZSM 1763 /2010 (ZCMV 12502), by M. Vences, D. R. Vieites, R. D. Randrianiaina, F.M. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, and T. Rajoafiarison, at Bemanevika, Antsirakala campsite, geographical coordinates S14.43061, E48.60179, 1466 m a.s.l., on 27 June 2010; ZSM 555 /2014 (DRV 6288), collected by F.M. Ratsoavina, D. R. Vieites, M. Vences, R. D. Randrianiaina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, and T. Rajoafiarison at Andrevorevo, geographical coordinates S14.3464, E49.1028, 1717 m a.s.l., on 21 June 2010. Diagnosis. Lygodactylus winki sp. nov. corresponds to a lineage forming part of the subclade A5 of Domerguella, and is one of only two representatives of this subclade known to reach northern Madagascar. The lowest genetic divergences of the lineages are 8.7% uncorrected 16S distance to L. guibei and 10.3% to L. miops. It is characterized by the presence of very distinct lateral spine-like scales at the base of the tail in males, as are found in several representatives of subclade A5 but not in other Domerguella. It can also be assigned to the subgenus Domerguella by an undivided mental scale with two postmentals, absence of a claw on the first finger, and 7 preanal pores in males. Within Domerguella, the new species is one of only two species of subclade A5 known from northern Madagascar. It differs from the other nominal species of Domerguella occurring in the same general area as follows: from L. expectatus by non-enlarged dorsolateral scales (longitudinal count of dorsal scales>185 vs. L. rarus by lack of regular crossbands on tail (vs. presence) and different body shape with less elongated limbs (relative hindlimb length 0.49–0.54 vs.>0.55); from L. madagascariensis, L. petteri, and L. salvi by a lower longitudinal count of ventral scales (83–98 vs.>100); from L. roellae and L. hapei, by a higher longitudinal count of dorsal scales (187–222 vs. 159–179); and from L. tantsaha by a lower longitudinal count of dorsal scales (187–222 vs. 239–240). Furthermore, the new species differs from all of these species of the subclades A1–A4 by the presence of a distinct lateral spine at the base of the tail, especially large in males but also clearly recognizable in females (vs. more weakly expressed or absent in the other species). From the other two nominal species in subclade A5 (L. guibei and L. miops, according to current taxonomy; see above) the new species differs as follows: from L. miops by a lower longitudinal count of ventral scales despite minimal overlap (83–98 vs. 98–113); and from L. guibei by apparently relatively longer hindlimbs (HIL/SVL 0.49–0.54 vs. 0.42–0.49). Further comparative examination of specimens also revealed a different head shape in L. winki compared to L. guibei, with apparently more expressed supraocular bulges (also visible in specimens in life; Fig. 14 vs. Fig. 20). Additional measurements taken on selected specimens in good, fully comparable state of preservation (Table 3) revealed that L. winki individuals have proportionally longer and higher heads than L. guibei, with non-overlapping values (relative snout tip to tympanum distance in percent, 24.7–28.8 vs. 23.0–24.6; relative head height in percent, 12.5–14.2 vs. 11.1–12.4; see Table 3). L. winki sp. nov. does not share haplotypes in CMOS or RAG1 with L. miops, and only one instance of haplotype sharing in RAG1 is detected with L. guibei. For a distinction from additional species newly named and described herein, see the respective diagnoses below. Etymology. This species is dedicated to Michael Wink, pharmacologist, herpetologist, ornithologist and professor emeritus of the University of Heidelberg, in recognition for his support of research in squamate systematics. The name is a patronym (i.e., a noun in the genitive case). Description of the holotype. Adult male, hemipenes everted, in good state of preservation, tail is broken and missing, second toe on the left forelimb is removed as source of tissue for molecular analysis (Fig. 15). SVL 29.5 mm, TAL 15.6 mm; for other measurements see Table 1. Head and neck short, head broader than body. The distance from the tip of the snout to the anterior border of the eye (3.9 mm) is less than the interorbital distance anteriorly (4.0 mm), 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 five scales: rostral, first supralabial, and three supranasals. Mental scale undivided; no contact between posterior projection of mental scale and first infralabial; two symmetrical postmental scales with four postpostmental scales; six infralabial scales; seven supralabial scales; two internasal scales; granular dorsal scales; dorsum with small, homogeneous, granular, and unkeeled scales of similar size to those on trunk, smaller than on head and tail, the scales on limbs can be slightly larger; 222 dorsal scales longitudinally along the body; 93 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; eight not very distinct dorsolateral tubercles, consisting of one scale; seven preanal pores; tail without whorls; large lateral spines at the base of the tail. Based on available photographs (Fig. 20), the holotype displayed a light marbled yellow grayish dorsal coloration with more brownish flanks before preservation. Distinct yellow spots are present on the head, the flank, the limbs, and the tail. Two black markings are present on the shoulder on either side of the body, reminiscent of double scapular semi-ocelli. Along the spine a narrow brown line reaches from the neck to the base of the tail. Adjacent to this line two pairs of symmetrical dark spots are present (forming two disrupted chevrons), one pair on forelimb level and one pair 10 mm posterior (Fig. 20D). After six years of preservation in ethanol, the preserved specimen is more uniformly brownish and most of the marbled pattern is faded. The ventral side is fawn with few small brown spots, most of them in the gular region. Variation. Males display a light marbled yellow grayish dorsal coloration with more brownish flanks and yellow spots. The dorsal and ventral coloration on females is darker without a pattern except for a few dark dorsal spots (Fig. 20E and 20I in comparison to Fig. 20A–G). Three additional specimens (two males [ZSM 555 /2014, ZSM 48 /2016], one female [ZSM 1763 /2010]) were examined. The SVL ranges between 29.8 and 33.4 mm with the female being the largest. The two males have a TAL of 37.8 and 45.0 mm. The relative hindlimb length is 0.49 to 0.53 with the female having the smallest. The female also has the smallest eyes relative to the size of the body and with a medium size smaller tubercles at the tail base than the males, which have large tubercles. The number of dorsal scales ranges between 187 and 208. The ventral scales range between 83 and 98. Natural history. The holotype was collected in primary rainforest, on the trunk of a big tree, 0.2 m above the ground. The paratypes from Ampotsidy were mostly collected at night sleeping on leaves, twigs, or vines. UADBA-R 70856 was collected in the afternoon, on the ground during heavy rain. Distribution. L. winki is known from three localities in the North and Sambirano regions in northern Madagascar: (1) the type locality, Ampotsidy, (2) Andrevorevo, and (3) Bemanevika.

Lygodactylus tantsaha sp. nov. Lygodactylus sp. 10: Gippner et al. (2021) Holotype. ZSM 196 /2018 (field number MSZC 0772), adult male, collected by M.D. Scherz, J.H. Razafindraibe, A. Razafimanantsoa, and S.M. Rasolonjavato at Montagne d’Ambre, west slope, northern Madagascar, at geographical coordinates S12.58503, E49.11596, 817 m. a.s.l., on 8 December 2017 at 22h20 (Fig. 15). Paratype. ZSM 197 /2018 (MSZC 0771), collected by M.D. Scherz, J.H. Razafindraibe, A. Razafimanantsoa, and S.M. Rasolonjavato at Montagne d’Ambre, west slope, Madagascar, at geographical coordinates S12.58548, E49.11697, 820 m a.s.l., on 8 December 2017 at 21h36. Diagnosis. Lygodactylus tantsaha sp. nov. corresponds to a genetically highly distinct lineage from northern Madagascar that is not closely related to any nominal species of Lygodactylus as defined in the previous sections. It belongs to subclade A2 within Domerguella as defined herein. It can also be assigned to the subgenus Domerguella by an undivided mental scale with two postmentals, absence of a claw on the first finger, and 7 preanal pores in males. Within Domerguella, the new species is only known from Montagne d’Ambre in northern Madagascar and differs from the other Domerguella occurring in this region as follows: from L. expectatus by non-enlarged dorsolateral scales (longitudinal count of dorsal scales>230 vs. L. rarus by lack of regular crossbands on tail (vs. presence) and different body shape without elongated limbs (relative hindlimb length 0.45–0.50 vs.>0.55); from L. madagascariensis by asymmetrical postmental scales (vs. symmetrical); from L. petteri by a larger longitudinal count of dorsal scales (239–240 vs. 189–222). Genetically, the new species is highly distinct from all species in subclade A5, and differs from almost all of them (potentially not L. fritzi sp. nov. described below) by an absent or only weakly expressed lateral spine at the tail base of males (vs. presence of a distinct spine). Tentatively, L. tantsaha sp. nov. differs in coloration from other Domerguella by the distinctly white upper lip (vs. brown in all other species) and white spots along the flank (vs. absent or at most light gray in L. madagascariensis and L. miops). The new species, on Montagne d’Ambre, occurs sympatrically with L. madagascariensis and L. petteri and is morphologically quite similar to these species, differing only in faint meristic characters as specified above. However, the fully concordant differentiation in mitochondrial genes (deep divergence in 16S:>13% to both L. madagascariensis and L. petteri) and in the unlinked loci CMOS and RAG-1, despite close syntopy, confirms this lineage represents a distinct species with restricted or absent gene flow to other co-occurring Domerguella. For a distinction from other species newly named and described herein, see the respective diagnoses below. Etymology. We are pleased to dedicate this species to Aaron M. Bauer in recognition of his extraordinary work fostering our knowledge about gecko diversity, biology, and evolution. The species name is derived from the Malagasy word tantsaha = farmer, in allusion to the original root of Aaron’s surname Bauer (German) = farmer. Coincidentally, individuals assignable to this species were found at the edge of an area of illegal farming within the park on the west slope, giving the name a second local meaning. Description of the holotype. Adult male, hemipenes everted, in moderately good state of preservation (Fig. 15), although the tail is detached, and the right forelimb is largely removed as a tissue sample for molecular analysis. SVL 31.9 mm, original tail (TAL 36.9 mm); for other measurements see Table 1. Head slightly broader than body. The distance from the tip of the snout to the anterior border of the eye (4.0 mm) is greater than the interorbital distance anteriorly (3.7 mm), and slightly greater than the distance between the eye and ear opening. Snout covered with enlarged granular scales, larger anteriorly on snout, becoming smaller laterally and anteriorly above the eye. Nostril surrounded by three scales: rostral, first supralabial and one supranasal. Mental scale undivided; only slight contact between posterior projection of mental scale and first infralabial; two asymmetrical postmental scales; four postpostmental scales; seven infralabial scales; seven supralabial scales; three internasal scales; granular dorsal scales; dorsum with small, homogeneous, granular and unkeeled scales of similar size to those on trunk, the scales on limbs are distinctly larger; 239 dorsal scales longitudinally along the body; 111 ventral scales between mental and cloaca; venter with large homogeneous smooth scales; no obvious lateral spines at the base of the tail; first finger present but very small, without bearing a claw; three pairs of subdigital lamellae on the fourth toe; one weakly expressed dorsolateral tubercle on either side, each composed of 1–2 scales; 7 preanal pores; tail without whorls. The holotype’s coloration in life based on available photographs was dorsally brown with a diffuse pattern consisting of dark and light spots, venter whitish. Flanks brighten towards venter with a diffuse ocelli-like pattern. Brown color on head with distinct border on supralabials to whitish venter. Six black stripes radially arranged around the eye. Tail slightly brighter than dorsum with pairs of black and white spots running posteriorly along the caudal spine (Fig. 16A). After four years of preservation in ethanol, the specimen darkened and patterns faded. Preserved specimen displays dark irregular spots on whitish gular region expanding to the anterior ventral torso. Variation. The coloration of this species appears tentatively to be characteristic, with a series of white spots always present along the flank in life (Fig. 16). The upper lip is also white. Natural history. All individuals were encountered and collected at night sleeping at the ends of very thin twigs, narrower than their bodies (Fig. 16D). Distribution. L. tantsaha is only known from the type locality, western Montagne d’Ambre., Published as part of Vences, Miguel, Multzsch, Malte, Gippner, Sven, Miralles, Aurélien, Crottini, Angelica, Gehring, Philip-Sebastian, Rakotoarison, Andolalao, Ratsoavina, Fanomezana M., Glaw, Frank & Scherz, Mark D., 2022, Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar's rainforest, pp. 1-61 in Zootaxa 5179 (1) on pages 25-28, DOI: 10.11646/zootaxa.5179.1.1, http://zenodo.org/record/7040745, {"references":["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. & 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"]}

Lygodactylus hapei sp. nov. Lygodactylus sp. 26: Gippner et al. (2021). Holotype. ZSM 298 /2018, female, collected at Djangoa (Djohahely) in the Sambirano Region in north-western Madagascar, approximately at geographical coordinates S13.7993, E48.3361, 20 m a.s.l. (Fig. 15), by unspecified local collectors. Diagnosis. Lygodactylus hapei sp. nov. corresponds to a genetically highly distinct lineage from a poorly known site in north-western Madagascar, and forms a clade with L. tantsaha (described above) and L. sp. 24, both from Montagne d’Ambre in the North. Considering this lineage as a new species is justified by its very deep genetic divergence of over 14% to all other Domerguella (16.3–16.4% to L. tantsaha), differences in scale counts, and a distinct longitudinally striped pattern on the throat not known from any other Domerguella. The new species belongs to subclade A2 within Domerguella as defined herein. It can also be assigned to the subgenus Domerguella by an undivided mental scale with two postmentals, and absence of a claw on the first finger. Within Domerguella, the new species is only known from one locality in the Sambirano region in northern Madagascar, and differs from the other nominal species of Domerguella by the presence of a longitudinally striped pattern on the throat, and additionally from the species occurring in northern Madagascar as follows: from L. expectatus by non-enlarged dorsolateral scales (longitudinal count of dorsal scales>185 vs. L. rarus by lack of regular crossbands on tail (vs. presence) and different body shape without elongated limbs (relative hindlimb length 0.43 vs.>0.55); from L. madagascariensis, L. petteri, L. salvi, and L. tantsaha by smaller longitudinal dorsal scale count (179 vs.>188) and smaller longitudinal ventral scale count (87 vs.>96). The new species appears to be very similar to L. roellae, a species from subclade A3, in scale counts and color pattern, but it may differ by smaller body size (SVL 26.3 vs. 35.9–36.0). The new species is genetically highly distinct from all species in subclade A5, based on concordant differentiation in mitochondrial genes (with deep divergence in 16S to all other species:>14%) and the unlinked loci CMOS and RAG-1. In addition it appears to differ by the absence of a spine at the tail base, which is weakly recognizable also in the females of all subclade A5 species except L. fritzi. Furthermore, the longitudinal dorsal scale count is smaller than in all known individuals of this subclade. For a distinction from additional species newly named and described herein, see the respective diagnoses below. Etymology. We dedicate this species to Hans-Peter “HaPe” Berghof, in recognition of his contributions to the knowledge of Madagascar geckos, especially Phelsuma. The name is a patronym (i.e., a noun in the genitive case). Description of the holotype. Adult female, in good state of preservation, tail regenerated, fourth toe on the left hind limb is removed as source of tissue for molecular analysis (Fig. 15). SVL 26.3 mm, TAL 27.4 mm; for other measurements see Table 1. Head slender with long neck, body broader than head. The distance from the tip of the snout to the anterior border of the eye (3.5 mm) is greater than the interorbital distance anteriorly (3.2 mm), and greater than the distance between the eye and ear opening. Snout covered with granular scales equally sized compared to the rest of the dorsum. Nostril surrounded by three scales: rostral, first supralabial, and two supranasal. Mental scale undivided; no contact between posterior projection of mental scale and first infralabial; two symmetrical postmental scales with five postpostmental scales; seven infralabial scales; eight supralabial scales; three internasal scales; granular dorsal scales; dorsum with small, homogeneous, granular, and unkeeled scales of similar size to those on trunk, the scales on limbs are not distinctly larger; 179 dorsal scales longitudinally along the body; 87 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; no dorsolateral tubercle; tail without whorls; no obvious lateral spines at the base of the tail. ......continued on the next page TABLE 2. (Continued) Based on available photograph (Fig. 19), the holotype in life displayed a broad brown stripe on the back with a brighter center running along the spine reaching from the snout to the base of the tail. Along the brighter center, irregularly scattered black spots are present. Flanks are yellowish brown with irregular small dark spots. A distinct black stripe is running from the snout through the eye to the shoulder ending in a black marking somewhat reminiscent of a scapular semi-ocellus, but positioned more posteriorly. Above this, a second whitish and broader stripe is present, reaching from the eye to the shoulder. Dorsally brown with a diffuse pattern consisting of dark and light spots, venter whitish. Flanks brighten towards venter with a diffuse ocelli-like pattern. Brown color on head with distinct border on supralabials to whitish venter. Six black stripes radially arranged around the eye. Tail slightly brighter than dorsum with pairs of black and white spots running posteriorly along the caudal spine (Fig. 19). During preservation in ethanol, the specimen darkened and patterns faded. Preserved specimen displays dark irregular spots on whitish gular region expanding to the anterior ventral torso. Variation. Only a single individual of this species (the holotype) is known. Natural history. The only known specimen was photographed millimeters from the posterior end of a planthopper larva (Fig. 19), and was presumably consuming honeydew excreted by the insect, as is known from other gecko species (Fölling et al. 2001). Distribution. L. hapei is only known from its type locality, Djohahely., Published as part of Vences, Miguel, Multzsch, Malte, Gippner, Sven, Miralles, Aurélien, Crottini, Angelica, Gehring, Philip-Sebastian, Rakotoarison, Andolalao, Ratsoavina, Fanomezana M., Glaw, Frank & Scherz, Mark D., 2022, Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar's rainforest, pp. 1-61 in Zootaxa 5179 (1) on pages 38-41, DOI: 10.11646/zootaxa.5179.1.1, http://zenodo.org/record/7040745, {"references":["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. & 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","Folling, M., Knogge, C. & Bohme, W. (2001) Geckos are milking honeydew-producing planthoppers in Madagascar. Journal of Natural History, 35, 279 - 284. https: // doi. org / 10.1080 / 00222930150215378"]}

Lygodactylus rarus Pasteur & Blanc, 1973 Lygodactylus (Domerguella) rarus Pasteur & Blanc, 1973 Chresonyms: Lygodactylus rarus: Kluge (1991); Glaw & Vences (1992, 1994, 2007); Puente et al. (2005, 2009); Röll et al. (2010); Gippner et al. (2021) Lygodactylus (Domerguella) rarus: Rösler (2000b) Name-bearing type: female holotype, MNHN 1990.6 —Type locality: “haute de la falaise orientale du karst d’Ambilobe (extrémité nord-est du Massif de l’Ankarana)”, according to the original description.—Other types: none according to original description.—Etymology: derived from Latin rarus (rare, unusual). Identity and Diagnosis. According to the diagnosis of Puente et al. (2009) this is a rather large-sized endemic of limestone karst areas of northern Madagascar, characterized by a long-legged, long-tailed and slender appearance. It differs from all species in the L. madagascariensis group by the presence of broad crossbands in the tail, of alternate light gray/brown color (Figs. 9–10). Although other Domerguella also can have tail crossbands, these are usually irregular, typically with alternating sections which start light brown or beige, gradually become darker to end in a somewhat posteriorly concave narrow dark line that then posteriorly borders sharply on the next light portion. In contrast, the crossbands of L. rarus typically consist of alternating brownish vs. gray portions which rather sharply border at each other, the brown portions typically being broader than the gray portions (Fig. 9). This typical pattern is also visible in the holotype which, upon examination in 2021, was in a quite poor state of preservation. The species also differs from all other Domerguella by the highest number of longitudinal ventral scales along the body (119–139, with 125 longitudinal ventral scales in the holotype; all other Domerguella have at most 110 ventral scales). In addition, this species is also characterized by a particularly slender body and long limbs (Fig. 10): relative hindlimb length (HIL/SVL) is 0.56–0.60 in L. rarus, vs. a maximum of 0.50 in all but one other Domerguella. The only other Domerguella species with long hindlimbs>0.5 is L. sp. 18, but also this species only reaches a ratio value of 0.54, thus shorter than in L. rarus. The three diagnostic character states (tail crossbands, large number of ventral scales, long hindlimbs) are all recognizable in the holotype, and in the genetically characterized specimens collected by us. All these specimens were collected in the Ankarana Massif. Therefore, there is no doubt about the identity of L. rarus, and the molecular data herein can confidently be assigned to this species. Furthermore, L. rarus is distinguished from L. miops and especially L. guibei by the absence (vs. presence) of dorsolateral tubercles and spiny tubercles at the tail base. It is further distinguished from the sympatric L. expectatus by its non-enlarged dorsolateral scales (vs. enlarged), absence of dark spots on the neck (vs. presence), and larger size (adult SVL 31.6–36.5 mm vs. 27.0– 29.7 mm). Distribution. L. rarus is reliably only known from its type locality, the Ankarana Massif. Pasteur & Blanc (1973) also report the species from Mangindrano (located at 1300 m a.s.l. on the Tsaratanana Massif), based on two juveniles that hatched from eggs collected in an abandoned bird nest. We here consider this record as in need of confirmation, given the uncertain attribution of these two hatchlings., Published as part of Vences, Miguel, Multzsch, Malte, Gippner, Sven, Miralles, Aurélien, Crottini, Angelica, Gehring, Philip-Sebastian, Rakotoarison, Andolalao, Ratsoavina, Fanomezana M., Glaw, Frank & Scherz, Mark D., 2022, Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar's rainforest, pp. 1-61 in Zootaxa 5179 (1) on pages 16-17, DOI: 10.11646/zootaxa.5179.1.1, http://zenodo.org/record/7040745, {"references":["Pasteur, G. & Blanc, C. P. (1973) Nouvelles etudes sur les lygodactyles (Sauriens Gekkonides). I. Donnees recentes sur Domerguella et sur ses rapports avec la Phytogeographie malgache. Bulletin de la Societe Zoologique de France, 98, 165 - 174.","Kluge, A. G. (1991) Checklist of Gekkonoid Lizards. Smithsonian Herpetological Information Service 85, 36 pp. https: // doi. org / 10.5479 / si. 23317515.85.1","Glaw, F. & Vences, M. (1992) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 331 pp. [First Edition.]","Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 480 pp. [Second Edition.]","Puente, M., Thomas, M. & Vences, M. (2005) Phylogeny and biogeography of Malagasy dwarf geckos, Lygodactylus Gray, 1864: Preliminary data from mitochondrial DNA sequences (Squamata: Gekkonidae). In: Huber, B. A. & Lampe, K. H. (Eds.), African Biodiversity: Molecules, Organisms, Ecosystems. Proc. 5 th Intern. Symp. Trop. Biol., Museum Koenig, Bonn. Springer, pp. 229 - 235. https: // doi. org / 10.1007 / 0 - 387 - 24320 - 8 _ 21","Puente, M., Glaw, F., Vieites, D. R. & 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","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. & 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","Rosler, H. (2000 b) Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28 - 153."]}

Lygodactylus madagascariensis (Boettger, 1881) Scalabotes madagascariensis Boettger, 1881 Chresonyms: Lygodactylus madagascariensis: Boulenger (1885), Puente et al. (2005) Lygodactylus madagascariensis (partim; including petteri as subspecies): Kluge (1991); Glaw & Vences (1992, 1994, 2007); Puente et al. (2009); Röll et al. (2010); Gippner et al. (2021); Lygodactylus (Domerguella) madagascariensis: Pasteur (1965a) Lygodactylus (Domerguella) madagascariensis (partim; including petteri as subspecies): Pasteur & Blanc (1967) Lygodactylus (Domerguella) madagascariensis madagascariensis: Rösler (2000b) Name-bearing type: male lectotype SMF 8937 (designated by Mertens 1967), collected by A. Stumpff. Krüger (2001) considered SMF 8937 as holotype.—Type locality: Nosy Be; “hab. in insula Nossi-Bé rarus”, according to the original description.—Other types: One paralectotype (the description was based on two specimens “(2 spec.)” according to the original description).—Etymology: name derived from its general provenance, Madagascar. Identity and Diagnosis. The lectotype of Lygodactylus madagascariensis was collected on the offshore island of Nosy Be. It is an adult male characterized by the absence of character states diagnostic for other species: it has no enlarged dorsolateral tubercles (as L. expectatus), no regular crossbands on the tail and not particularly long hindlimbs (as L. rarus), and no enlarged tubercles at the base of the tail. According to our molecular data, only one genetic lineage of Domerguella has been found on Nosy Be (two sequences available). This same lineage also occurs in several forests of relatively low elevation in the Sambirano region (to which Nosy Be also belongs): Tsaratanana (Andampy), Manongarivo, Maromiandra. These localities also host many other species of amphibians and reptiles occurring on Nosy Be (e.g. Penny et al. 2017), supporting the biogeographic assignment of the name L. madagascariensis to this lineage. The available material of this lineage also agrees in all studied morphological characters with the holotype, for instance in the number of longitudinal ventral scales (106 in the holotype vs. 106–138 in the other specimens) and dorsal scales (246 vs. 205–258). The species is comparatively small (SVL 28.5 –34.0 mm) and in many specimens shows a rather typical color pattern of irregular beige patches arranged in longitudinal rows on the brown dorsum, along with irregular dark brown pattern (Fig. 11). A genetically slightly divergent variant of L. madagascariensis is also present on Montagne d’Ambre, an isolated mountain in extreme northern Madagascar. This is of relevance because L. madagascariensis petteri has been described from this mountain as a subspecies. In the subsequent species account we will show that the name petteri does not apply to the L. madagascariensis specimens from Montagne d’Ambre but to another, sympatric lineage, thus justifying the elevation of petteri to species status. Distribution. L. madagascariensis is reliably known from (1) its type locality Nosy Be, (2) Manarikoba Forest on the western slope of the Tsaratanana Massif (Andampy Campsite), (3) Manongarivo, (4) Maromiandra, (5) Andrafainkona, and (6) Montagne d’Ambre. These localities are in the Sambirano region and the North regions of Madagascar., Published as part of Vences, Miguel, Multzsch, Malte, Gippner, Sven, Miralles, Aurélien, Crottini, Angelica, Gehring, Philip-Sebastian, Rakotoarison, Andolalao, Ratsoavina, Fanomezana M., Glaw, Frank & Scherz, Mark D., 2022, Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar's rainforest, pp. 1-61 in Zootaxa 5179 (1) on pages 18-19, DOI: 10.11646/zootaxa.5179.1.1, http://zenodo.org/record/7040745, {"references":["Boettger, O. (1881) Diagnoses reptilium et batrachiorum novorum ab ill. Antonio Stumpff in insula Nossi-Be Madagascariensis lectorum. Zoologischer Anzeiger, 4, 358 - 362.","Boulenger, G. A. (1885) Catalogue of the lizards in the British Museum (Nat. Hist.) I. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. Printed by order of the Trustees London, 450 pp.","Puente, M., Thomas, M. & Vences, M. (2005) Phylogeny and biogeography of Malagasy dwarf geckos, Lygodactylus Gray, 1864: Preliminary data from mitochondrial DNA sequences (Squamata: Gekkonidae). In: Huber, B. A. & Lampe, K. H. (Eds.), African Biodiversity: Molecules, Organisms, Ecosystems. Proc. 5 th Intern. Symp. Trop. Biol., Museum Koenig, Bonn. Springer, pp. 229 - 235. https: // doi. org / 10.1007 / 0 - 387 - 24320 - 8 _ 21","Kluge, A. G. (1991) Checklist of Gekkonoid Lizards. Smithsonian Herpetological Information Service 85, 36 pp. https: // doi. org / 10.5479 / si. 23317515.85.1","Glaw, F. & Vences, M. (1992) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 331 pp. [First Edition.]","Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 480 pp. [Second Edition.]","Puente, M., Glaw, F., Vieites, D. R. & 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","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. & 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","Pasteur, G. (1965 a) Notes preliminaries sur les lygodactyles (Gekkonides). IV Diagnoses de quelques formes africaines et malgaches. Bulletin du Museum national d'Histoire naturelle, 36, 311 - 314.","Pasteur, G. & Blanc, C. P. (1967) Les lezards du sous-genre malgache de lygodactyles Domerguella (Gekkonides). Bulletin de la Societe Zoologique de France, 92, 583 - 597.","Rosler, H. (2000 b) Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28 - 153.","Mertens, R. (1967) Die herpetologische Sektion des Naturmuseums und Forschungsinstitutes Senckenberg in Frankfurt a. M. nebst einem Verzeichnis ihrer Typen. Senckenbergiana Biologica, 48, Sonderheft A, 1 - 106.","Kruger, J. (2001) Die madagassischen Gekkoniden. Teil II: Die Geckos der Gattung Lygodactylus GRAY 1864 (Reptilia: Sauria: Gekkonidae). Gekkota, 3, 3 - 28.","Penny, S. G., Crottini, A., Andreone, F., Bellati, A., Rakotozafy, L. M. S., Holderied, M. W., Schwitzer, C. & Rosa, G. M. (2017) Combining old and new evidence to increase the known biodiversity value of the Sahamalaza Peninsula, Northwest Madagascar. Contributions to Zoology, 86, 273 - 296. https: // doi. org / 10.1163 / 18759866 - 08604002"]}

Lygodactylus salvi sp. nov. Lygodactylus sp. 8: Gippner et al. (2021). Holotype. ZSM 783 /2001 (FGMV 2001.74), an adult male collected by F. Andreone, F. Mattioli, J. Randrianirina, and M. Vences on the western slope of the Tsaratanana massif (Manarikoba forest, Antsahamanara campsite), northern Madagascar, at geographical coordinates S14.0450, E48.7844, ca. 1000 m a.s.l., between 4–9 February 2001 (Fig. 15). Paratype. ZSM 557 /2014 (DRV 6327), a female collected by F.M. Ratsoavina, D. R. Vieites, M. Vences, R. D. Randrianiaina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, and T. Rajoafiarison, from Ambodikakazo, a site south of the Tsaratanana Massif, northern Madagascar, at geographical coordinates S14.2098, E48.8981, 1411 m a.s.l., on 16 June 2010. Diagnosis. Lygodactylus salvi sp. nov. corresponds to a genetically highly distinct lineage from northern Madagascar that is not closely related to any nominal species of Lygodactylus as defined in the previous sections. It belongs to subclade A3 within Domerguella as defined herein. It can also be assigned to the subgenus Domerguella by an undivided mental scale with two postmentals, absence of a claw on the first finger, and 6 preanal pores in males. Within Domerguella, the new species is only known from two localities in northern Madagascar and differs from the other nominal species of Domerguella occurring in this part of the island as follows: from L. expectatus by non-enlarged dorsolateral scales (longitudinal count of dorsal scales>210 vs. L. rarus by lack of regular crossbands on tail (vs. presence) and different body shape without elongated limbs (relative hindlimb length 0.48–0.50 vs.>0.55); from L. madagascariensis by smaller longitudinal dorsal scale count (211–217 vs. 219–258); and from L. petteri by a larger longitudinal count of ventral scales (107–112 vs. 101–105 in all but one individual of L. petteri); and from L. tantsaha by a smaller longitudinal count of dorsal scales (211–217 vs. 239–240). Given the somewhat uncertain allocation of the nomina to the eastern species in subclade A5, a morphological diagnosis from these species is convoluted. However, genetically, the new species is highly distinct from all species in this subclade, and differs from L. guibei as well as from many specimens of L. miops by a less distinctly expressed lateral spine at the tail base of males (vs. presence of a distinct, large spine in L. guibei, and an at least slightly more distinct spine in L. miops). Furthermore, the new species can be distinguished from L. guibei and L. miops by smaller relative eye diameter (ED is 4.8–5.0% of SVL, vs. 5.3–7.7% in L. guibei and L. miops). Finally, this species shows concordant differentiation in mitochondrial genes (deep divergence in 16S to all other species:>11%) and the unlinked loci CMOS and RAG-1 (haplotype sharing with lineage L. sp. 17 only in some CMOS haplotypes). For a distinction from the sister lineage, described below as. L. roellae, see Diagnosis of that species. For a distinction from additional species newly named and described herein, see the respective diagnoses below. Etymology. The name is dedicated to Salvador “Salvi” Carranza, Institut de Biologia Evolutiva (CSIC-UPF), Barcelona, in recognition for his substantial contributions to gecko taxonomy, and conservation of herpetofauna. The species epithet name is defined as a noun in apposition (not a noun in the genitive case) to avoid ending with a non-euphonious double-i. Description of the holotype. Adult male, hemipenes everted, in moderate state of preservation, tail regenerated, right forelimb is removed as source of tissue for molecular analysis (Fig. 15). SVL 29.9 mm, TAL 24.9 mm; for other measurements see Table 1. Long head with distinct neck, body broader than head. The distance from the tip of the snout to the anterior border of the eye (4.2 mm) is greater than the interorbital distance anteriorly (3.6 mm), and greater than the distance between the eye and ear opening. Snout covered with enlarged granular scales compared to 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 symmetrical postmental scales with four postpostmental scales; seven infralabial scales; seven supralabial scales; three internasal scales; granular dorsal scales; dorsum with small, homogeneous, granular and unkeeled scales of similar size to those on trunk, the scales on limbs are slightly larger; 217 dorsal scales longitudinally along the body; 107 ventral scales between mental and cloaca; venter with large homogeneous smooth scales that are a bit smaller in the gular region; first finger present but very small, not bearing a claw; three pairs of subdigital lamellae on the fourth toe; six dorsolateral tubercles, each consisting of one scale; six preanal pores; tail without whorls; small lateral spines at the base of the tail. Life coloration of the holotype based on available photographs (Fig. 17) was grayish with a slightly distinct red-brownish lateral stripe running from the eye to the base of the tail. Both colors are also distinctly present on the limbs. Overall, the appearance is cryptic with no distinct black markings (Fig. 17). Color after 20 years in preservative ethanol is almost uniformly gray brownish, with a weak dorsal pattern. In the preserved specimen, the ventral side is uniformly whitish with a slight yellowness. Small brown spots are present on the throat and the rest of the venter. Variation. The female paratype (ZSM 557 /2014) is bigger than the holotype, with an SVL of 36.2 mm, but has relatively shorter hindlimbs (HIL / SVL 0.48). The relative size of tubercles at the tail base is a bit smaller, while the lateral tubercles between the legs contain a few more scales (1–3). The animal has bigger eyes in relation to its size. While the dorsal scale count is a bit lower (211), the ventral scale count is a bit higher (112). Coloration in preservative is uniformly grayish-brown, without lateral stripe, and with very small symmetrical dark dorsal markings anterior to hindlimb insertion; ventrally with dense dark mottling on throat, chest and belly. The differences between holotype and paratype could be due to different sex or just random variation. Distribution. L. salvi is known from (1) the type locality, Manarikoba forest on the west slope of the Tsaratanana Massif, and (2) Ambodikakazo south of Tsaratanana. Natural history. Practically nothing is known of the natural history of this enigmatic species., Published as part of Vences, Miguel, Multzsch, Malte, Gippner, Sven, Miralles, Aurélien, Crottini, Angelica, Gehring, Philip-Sebastian, Rakotoarison, Andolalao, Ratsoavina, Fanomezana M., Glaw, Frank & Scherz, Mark D., 2022, Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar's rainforest, pp. 1-61 in Zootaxa 5179 (1) on pages 28-29, DOI: 10.11646/zootaxa.5179.1.1, http://zenodo.org/record/7040745, {"references":["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. & 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"]}

Lygodactylus petteri Pasteur & Blanc, 1967 Lygodactylus madagascariensis petteri Pasteur & Blanc, 1967 Chresonyms: Lygodactylus madagascariensis petteri: Kluge (1991); Glaw & Vences (1992, 1994, 2007); Puente et al. (2009); Gippner et al. (2021) Lygodactylus (Domerguella) madagascariensis petteri: Rösler (2000b) Name-bearing type: holotype MNHN 1990.4, female.—Type locality: “Montagne d’Ambre, forêt ancienne-Roussettes” according to the original description.—Other types: two paratypes; MNHN 1990.5, male; and MNHN 1893.194.—Etymology: eponym for Jean-Jacques Petter. Identity and Diagnosis. This nomen was coined for specimens from Montagne d’Ambre that were considered to be a subspecies of L. madagascariensis. According to the original description, this subspecies was purported to differ from typical L. madagascariensis by fewer scales in general (i.e., lower values in various scale counts), suggesting overall larger scales; a larger body size; a different coloration (beige vs. brown); and some other possible differences. Indeed, our measurements and scale counts of the name-bearing type (holotype) and one paratype confirmed these are relatively large-sized (SVL 33.2–35.0, thus at and slightly beyond the upper size limit of L. madagascariensis) and have lower longitudinal counts of dorsal scales (189 in the holotype; vs. 205–258 in L. madagascariensis) and ventral scales (102 in the holotype and 103 in one paratype; vs. 106–138 in L. madagascariensis). This suggests the name petteri should be applied to one of the Domerguella lineages occurring at Montagne d’Ambre. One of these (called L. sp. 10 by Gippner et al. 2021) appears to reach rather large-sizes (36.9 mm SVL in one specimen) but has relatively high longitudinal counts of dorsal scales (239–240 in two available specimens), thus differing from the types of L. petteri. This lineage (known only from the west slope of Montagne d’Ambre) represents a new species that will be formally named and described below. Another lineage from Montagne d’Ambre (L. sp. 24) is represented by only one genetic sample, the voucher of which was not available for examination. Unfortunately, no information at all on the coloration or morphology of this lineage is available. We can only hypothesize from its rarity (no further specimens found despite intensive surveys in Montagne d’Ambre) that it is unlikely to correspond to the types of petteri. The third lineage is the one that we have genetically assigned to L. madagascariensis above, and the one individual from Montagne d’Ambre examined (Table 1) agrees well with topotypical specimens of this species, but not with the petteri types. However, a fourth lineage from Montagne d’Ambre agrees in all morphological characters very well with the petteri types: it consists of relatively large specimens (SVL in our material 30.3–38.5 mm) with few ventral scales (101–113 vs. 102–103 in the types) and dorsal scales (209–222 vs. 189 in the holotype). We therefore are confident that the specimens belonging to this lineage are conspecific with the types of L. petteri. Since this lineage co-occurs on Montagne d’Ambre with L. madagascariensis with deep genetic differentiation in both mitochondrial and nuclear genes, we conclude that the nomen petteri applies to a full species, Lygodactylus petteri, and we therefore herewith formally elevate it to species level. It needs to be emphasized that due to a lack of comparative morphological data of the only specimen of L. sp. 24 we cannot fully exclude that this lineage also matches morphologically the holotype of L. petteri and may be conspecific with it. Collection of additional material of L. sp. 24, or alternatively, molecular “archival DNA” data from the holotype of L. petteri, is needed to fully ascertain the identity of these geckos from Montagne d’Ambre. However, independent from these remaining questions, it appears we can conclude with sufficient reliability that L. petteri is not conspecific with L. madagascariensis from which it differs morphologically, and we confirm it is distinct from L. sp. 10, which is described as a new species below. No clear and consistent differences in color or pattern were found between L. madagascariensis and L. petteri; both showed a considerable variation in dorsal pattern (near-uniform to heavily patterned in L. madagascariensis vs. asymmetrical series of rather small dorsolateral markings or striped phenotype in L. petteri). However, the two specimens of L. petteri for which life coloration is known (Fig. 12) do not show the longitudinal rows of large beige patches typical for many L. madagascariensis, and furthermore, the male specimen ZSM 195/2018 has yellow elements dorsally, which we have not seen in any L. madagascariensis. Ventrally the throat is yellowish and ranged from near unspotted to weakly and irregularly spotted in both species. According to the original description of L. petteri by Pasteur & Blanc (1967), it differs from L. madagascariensis by several characters, which we review here. First of all, L. petteri purportedly has fewer scales (and thus larger ones) in general (characters 9, 12, 13, 17, 31, 32, 33 of Pasteur & Blanc 1967). This agrees with our findings for longitudinal counts of dorsal and ventral scales, while for instance the number of supralabials (character 9 of Pasteur & Blanc 1967) does not clearly differ between the two species according to our data. The authors also reported a larger body size for L. petteri, which is in agreement with our data, as well as differences in coloration and in sexual dimorphism, and a possibly larger size of preanal pores in L. petteri. Once more extensive series of both species become available, it will be worth examining whether these characters may indeed constitute diagnostic differences. Natural history. A half-digested specimen of L. petteri was regurgitated by a young Compsophis sp. aff. laphystius (Hutter et al. 2018). Two specimens of L. petteri (ACZC 1407 and ACZC 1427) were found under the bark of Eucalyptus sp. trees at the Gîte d’Étape site on Montagne d’Ambre. Distribution. L. petteri is only known from its type locality, Montagne d’Ambre., Published as part of Vences, Miguel, Multzsch, Malte, Gippner, Sven, Miralles, Aurélien, Crottini, Angelica, Gehring, Philip-Sebastian, Rakotoarison, Andolalao, Ratsoavina, Fanomezana M., Glaw, Frank & Scherz, Mark D., 2022, Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar's rainforest, pp. 1-61 in Zootaxa 5179 (1) on pages 19-20, DOI: 10.11646/zootaxa.5179.1.1, http://zenodo.org/record/7040745, {"references":["Pasteur, G. & Blanc, C. P. (1967) Les lezards du sous-genre malgache de lygodactyles Domerguella (Gekkonides). Bulletin de la Societe Zoologique de France, 92, 583 - 597.","Kluge, A. G. (1991) Checklist of Gekkonoid Lizards. Smithsonian Herpetological Information Service 85, 36 pp. https: // doi. org / 10.5479 / si. 23317515.85.1","Glaw, F. & Vences, M. (1992) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 331 pp. [First Edition.]","Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 480 pp. [Second Edition.]","Puente, M., Glaw, F., Vieites, D. R. & 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","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. & 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","Rosler, H. (2000 b) Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28 - 153.","Hutter, C. R., Andriampenomanana, Z. F., Razafindraibe, J., Rakotoarison, A. & Scherz, M. D. (2018) New dietary data from Compsophis and Alluaudina species (Squamata: Lamprophiidae: Pseudoxyrhophiinae), and implications for their dietary complexity and evolution. Journal of Natural History, 52, 2497 - 2510. https: // doi. org / 10.1080 / 00222933.2018.1543732"]}

New genetic, bioacoustic and morphological data on green-coloured Boophis treefrogs from eastern Madagascar reveal an additional level of cryptic diversity in these frogs. Two candidate species, Boophis sp. Ca36 and Ca37, are closely related to each other and to B. sandrae, with uncorrected pairwise distances in the mitochondrial 16S rRNA gene as low as 2.2% between some individuals. However, the three lineages show full concordance between differentiation in the 16S and the nuclear-encoded SACS gene, despite confirmed syntopy of B. sandrae and B. sp. Ca37 in the Ranomafana region, and probable syntopy of B. sp. Ca36 and B. sp. Ca37 in the Andasibe region. Most likely, these lineages are also divergent in advertisement calls, but the available recordings cannot be reliably assigned to either of them. Based on new material collected from various new sites, we here formally name B. sp. Ca36 as new species B. asquithi sp. n., and suggest targeted fieldwork on calls and larval stages to allow for a complete and fully conclusive taxonomic revision of this species complex. The example of these frogs illustrates how continued underestimation of cryptic diversity in anurans can lead to incorrect assignment of specimens, and leads us to emphasize the importance of designating as name-bearing types (holotypes) of anurans only individuals whose identity is unambiguous by genetic data or, at least, call recordings reliably assignable to the type specimen.

Uroplatus kelirambo sp. nov. (Figures 5���6) Remark. This new species was previously referred to as Uroplatus sp. E by Raxworthy et al. (2008b), as U. sp. 2 by Ratsoavina et al. (2011), and as U. ebenaui [Ca2] by Ratsoavina et al. (2012, 2013, 2015). Holotype. ZSM 641/2014 (field number DRV 6192), adult male with everted hemipenes, collected at the Matsaborimaiky camp site in the Tsaratanana mountain massif, Antsiranana Province, northern Madagascar, 14.1526��S, 48.9573��E, 2021 m above sea level, on the night of 13 June 2010, by D. Vieites, M. Vences, F. M. Ratsoavina, F. Randrianasolo, R. D. Randrianiaina, S. Rasamison, A. Rakotoarison, E. Rajeriarison and T. Rajoafiarison. Paratypes. ZSM 642/2014 (DRV 6250) and ZSM 643/2014 (DRV 6251), two adult males, and ZSM 1832/ 2010 (ZCMV 12388), an adult female, all with same collection data as the holotype. Etymology. The specific epithet kelirambo is composed of the Malagasy words for small (kely) and tail (rambo). The name refers to the small and very thin tail of this species, but also points to the movie character Rambo and his toughness. This species shares with other members of the U. ebenaui group a grim, Rambo-esque expression, and is tough, surviving at exceptionally high altitudes, unique among Madagascar���s nocturnal geckos. The name is an invariable noun in apposition. Diagnosis. Uroplatus kelirambo sp. nov. is included in the Uroplatus ebenaui group of small-sized leaf-tailed geckos due to its triangular head with supraocular spines, laterally compressed body, and short tail. It differs from all members of the U. fimbriatus group (U. fimbriatus, U. giganteus, U. henkeli, U. sikorae and U. sameiti) and U. lineatus by its much smaller size (adult SVL 53.1���63.4 mm versus at least 85 mm), lack of lateral dermal fringes on any part of the body, and lateral compression of the body (versus depressed body shape with lateral integumentary fringes), and a relatively much shorter tail. The evident triangular head, rather smooth skin and short tail distinguish U. kelirambo from U. alluaudi, U. guentheri, U. malahelo and U. pietschmanni. Within the U. ebenaui group, the blackish oral mucosa distinguishes U. kelirambo from U. fiera, U. finiavana and U. fotsivava (oral mucosa unpigmented). Furthermore, the new species differs from all species by the unique elongated shape and small size of its tail. In particular, it differs from U. finiavana, U. malama and U. phantasticus by a shorter tail (adult TAL/SVL 0.35���0.44 vs. 0.48 in U. finiavana, 0.72 in U. malama, and 0.62���0.76 in U. phantasticus), and from U. fiera and U. fotsivava by a longer tail (adult TAL/SVL 0.35���0.44 vs. 0.28���0.34 in U. fiera and 0.15���0.32 in U. fotsivava). It differs from all other nominal species of the U. ebenaui group by a narrower tail; this is obvious from the lower values of the ratio TAW/SVL (0.04���0.05 vs. 0.24 in U. malama, 0.16���0.20 in U. phantasticus, 0.11 in U. finiavana, 0.06���0.07 in U. fiera, 0.05���0.09 in U. ebenaui and 0.07���0.1 in U. fotsivava), but especially in values of the ratio TAW/TAL (0.10���0.13 vs. 0.33 in U. malama, 0.25���0.29 in U. phantasticus, 0.23 in U. finiavana, 0.19���0.21 in U. fiera, 0.13���0.27 in U. ebenaui and 0.17���0.32 in U. fotsivava). In direct comparison to U. finiavana, the closest relative of U. kelirambo among the nominal species of Uroplatus, the new species is thus distinguished by its tail shape and size, and the pigmented (vs. unpigmented) oral mucosa. Description of the holotype. Adult male in good condition with an intact tail and everted hemipenes. SVL 57.3 mm, tail length 25.4 mm, maximum tail width 2.5 mm, for further measurements see Table 1. Head triangular in dorsal view; snout length 5.1 mm; canthus rostralis indistinct; snout sloping weakly and continuously downward anteriorly; snout undepressed in lateral view (1.1 times longer than eye diameter); eyes large (eye diameter 4.8 mm), bulging slightly above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small (horizontal diameter 0.7 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 dermal fringes; limbs without fringes, forelimb reaches beyond tip of snout when adpressed forward, but is quite far from the groin when adpressed backwards along body (forelimb length/axilla��� groin distance 24.6/29.8 mm = 83%), hindlimb reaches the axilla when adpressed forward along body (hindlimb length/axilla���groin distance 30.8/29.8 mm = 103%); tail length 44% of snout���vent length, almost without membranous borders of the tail, with just one small dermal spine on each side, with a slightly spatulate tail-tip. Nares separated from each other by seven small granular scales, from the first supralabial by one scale, and from the rostral scale by one scale; first supralabial taller than the others; rostral entire, much wider than tall; mental scale very small, not differentiated from infralabial scales; 22/22 (right/left) true supralabials and 21/20 infralabials (as defined in methods above); no enlarged postmental scales or chin shields; dorsal and ventral scales of head, neck, body, limbs, and tail small, strongly granular, juxtaposed and largely of uniform size, except for some larger, interspersed and partly raised tubercles and except for the irregular lines on the head and body which consist of series of slightly enlarged scales. Two curved lines (rows of slightly enlarged scales) extending from the posterolateral parts of the head (nuchal region) converge on the neck forming a V-shaped pattern (neck triangular line). A weakly curved, moderately distinct, and weakly posteriorly extending line (also formed by a row of slightly enlarged scales) is present between the eyes and connects the supraocular spines. Three additional weaklyvisible transverse lines, consisting of slightly enlarged scales, are recognizable on the frontal region of the head. A few well developed spines on the posterior part of the head (ca. 6), on hindlimbs (ca. 12 per limb), a dermal flap on each knee not bearing a spine; spines present on the elbow, three spines per arm; a long, pointed supraocular spine above each upper eyelid; posterior border of eye fringed. Axillary pits absent. Scales of the posterior ventral abdomen enlarged and arranged more uniformly than other scales, almost forming proper rows. Hemipenes everted, bearing two lobes. Calyx with protuberance bearing honeycomb-like structures, especially on the asulcal side. Area of sulcus spermaticus is smooth. Each lobe with dense field of pointed papillae at its apex. Coloration. After six and a half years of preservation in 70% ethanol, the color pattern remains the same as in the living animal (Fig. 5) but its vivacity and intensity have faded slightly. All dorsal surfaces are brown in base color, mottled darker and lighter. The tail is creamy chocolate above, darkening to a brown tip. The head is also creamy chocolate above, with the V-shaped marking of the nape darkening toward its posterior tip, bordered by lighter brown, making the marking highly contrasting. The dorsum has delicate lines of cream and dark brown, generally parallel to the ribs beneath them. A few small dark brown flecks are present on the dorsum, but none of note. Two light cream markings defined by brown borders are present below each eye, one at the posterior corner and one at the mid-eye, both descending to the supralabials. The supralabials are generally beige in color, but a few are burnt umber. The infralabials weakly alternate between cream and burnt umber. The exterior shank from the knee to the outer foot is beige in color, strongly contrasting the rather dark brown of the rest of the leg and foot. A cream dorsomedial line runs from the pelvic region to the mid-dorsum. The venter, including the ventral limbs except the lower hindlimbs, is a lighter brown than the dorsum, and much less mottled. The ventral trunk bears several round cream spots defined by brown borders, with two pairs of such markings in the anterior pectoral region, and one small pair at the glenoid socket and other such spots on the mid and posterior abdomen. The tail is slightly darker below than above, and has an oblong cream fleck at its base. The oral mucosa is dark in preservative. In life the eye was beige with reddish and yellowish brown mottling. Variation. In general, the paratypes agree well with the holotype in morphology. For measurements, see Table 1. The canthus rostralis of ZSM 643/2014 is quite distinct, but it is indistinct in all other type specimens. The number of head spines varies from four (ZSM 1832/2010) to eight (ZSM 642/2014 ��� 643/2014). Spines are always present on the elbow, but can be miniscule, and there can be almost no other spines on the arms (ZSM 1832/2010) or up to four spines per arm (ZSM 642/2014). The legs can have as few as five spines per leg (ZSM 1832/2010) to as many as seventeen (ZSM 642/2014). The oral mucosa is pigmented in all specimens. The supraocular spines of all paratypes are shorter than the holotype, and the line connecting them can be curved to a point. The hemipenes of ZSM 643/2014 strongly resemble those of the holotype; those of ZSM 642/2014 are not fully everted. The number of true supralabials ranges from 18 to 22, of true infralabials 17 to 21. Coloration is highly variable, as is typical of Uroplatus. ZSM 643/2014 is quite similar to the holotype, but the posterior trunk is darker than the chin, it lacks cream spots on the venter except at the base of the tail, it has a less distinct nape V and chevrons on the body, and it has large burnt umber spots, especially on the midline. ZSM 642/2014 is a light caramel brown in base color, with dark markings as in ZSM 643/2014, but a distinct nape marking as in the holotype; ventrally this specimen has dark chevrons on the chin and is mottled cream and brown, with paired cream spots on the posterior abdomen and one weak one on the throat. ZSM 1832/2010 is reddish brown in base color with large off-white blotches on the flanks and head, its ventral coloration similar to ZSM 642/2014, and possessing a triangle of three white spots at the base of the tail. It has only one marking below the eye. Tails of males can have two or three lateral spines, whereas females have a very narrow membranous border; again this indicates some degree of sexual dimorphism in tail shape (see Ratsoavina et al. 2015). Distribution. The species is known from only a single site, the environments of Matsaborimaiky (named Matsabory Maiky by Rakotoarison et al. 2012), the second campsite along the trail to the peak of Maromokotro in the Tsaratanana Massif (the highest mountain in Madagascar). Here, specimens were collected in the environments of the campsite, and on the trail towards the third campsite, up to an estimated elevation of 2200 m a.s.l. Natural history. All specimens of U. kelirambo were collected at sites close to Matsaborimaiky, on branches of small trees. The site is characterized by a wetland surrounded by a mixture of montane and bamboo trees. Classified as a humid montane forest, the canopy does not reach 20 meters height and trees are rather short, sinuous in shape and covered in lichen. The holotype of U. kelirambo was collected at night perching in a small tree at ca. 1.75 m height from the ground., Published as part of Ratsoavina, Fanomezana M., Gehring, Philip-Sebastian, Scherz, Mark D., Vieites, David R., Glaw, Frank & Vences, Miguel, 2017, Two new species of leaf-tailed geckos (Uroplatus) from the Tsaratanana mountain massif in northern Madagascar in Zootaxa 4347 (3), DOI: 10.11646/zootaxa.4347.3.2, http://zenodo.org/record/1048750, {"references":["Raxworthy, C. J., Pearson, R. G., Zimkus, B. M., Reddy, S., Deo, A. J., Nussbaum, R. A. & Ingram, C. M. (2008 b) Continental speciation in the tropics: contrasting biogeographic patterns of divergence in the Uroplatus leaf-tailed gecko radiation of Madagascar. Journal of Zoology, 275, 423 - 440. https: // doi. org / 10.1111 / j. 1469 - 7998.2008.00460. x","Ratsoavina, F. M., Louis, E. E. Jr., Crottini, A., Randrianiaina, R. D., Glaw, F. & Vences, M. (2011) A new leaf tailed gecko species from northern Madagascar with a preliminary assessment of molecular and morphological variability in the Uroplatus ebenaui group. Zootaxa, 3022, 39 - 57.","Ratsoavina, F. M., Vences, M. & Louis, E. E. Jr. (2012) Phylogeny and phylogeography of the Malagasy leaf-tailed geckos in the Uroplatus ebenaui group. African Journal of Herpetology, 61, 143 - 158. https: // doi. org / 10.1080 / 21564574.2012.729761","Ratsoavina, F. M., Raminosoa, N. R., Louis, E. E. Jr., Raselimanana, A. P., Glaw, F. & Vences, M. (2013) An overview of Madagascar's leaf tailed geckos (genus Uroplatus): species boundaries, candidate species and review of geographical distribution based on molecular data. Salamandra, 49, 115 - 148.","Ratsoavina, F. M., Ranjanaharisoa, F. A., Glaw, F., Raselimanana, A. P., Miralles, A. & Vences, M. (2015) A new leaf-tailed gecko of the Uroplatus ebenaui group (Squamata: Gekkonidae) from Madagascar's central eastern rainforests. Zootaxa, 4006, 143 - 160. https: // doi. org / 10.11646 / zootaxa. 4006.1.7","Rakotoarison, A., Glaw, F., Vieites, D. R., Raminosoa, N. R. & Vences, M. (2012) Taxonomy and natural history of arboreal microhylid frogs (Platypelis) from the Tsaratanana Massif in northern Madagascar, with description of a new species."]}

Uroplatus fotsivava sp. nov. (Figures 4���5) Remark. This new species was previously referred as Uroplatus sp. F (1) by Raxworthy et al. (2008b), as U. sp. 1 by Ratsoavina et al. (2011), and as U. ebenaui [Ca1] by Ratsoavina et al. (2012, 2013, 2015). Holotype. ZSM 1830/2010 (field number ZCMV 12279), adult male, collected in the Analabe forest on the Ambodimanga mountain near Antambato village, Antsiranana Province, northern Madagascar, 14.5048��S, 48.8760��E, 1361 m above sea level, on the night of 6 June 2010, by M. Vences, D. R.Vieites, R. D. Randrianiaina, F. M. Ratsoavina, F. Randrianasolo, S. Rasamison, A. Rakotoarison, E. Rajeriarison and T. Rajoafiarison. Paratypes. ZSM 1831/2010 (ZCMV 12280) and ZSM 639/2014 (DRV 6067), two adult males, and ZSM 1829/2010 (ZCMV 1227), an adult female, all with the same collecting data as the holotype; ZSM 646/2014 (DRV 6326), adult male, collected at Ambodikakazo, 14.2098��S, 48.8981��E, 1411 m above sea level, on the night of 16 June 2010 by F. M. Ratsoavina and F. Randrianasolo; ZSM 648/2014 (DRV 6263), adult female, collected at Vinanitelo forest, Ambinanitelo, 14.2098��S, 48.9702��E, 1280 m above sea level, on the night of 22 June 2010 by D.R. Vieites, M. Vences, R. D. Randrianiaina, S. Rasamison, A. Rakotoarison, E. Rajeriarison and T. Rajoafiarison; ZSM 647/2014 (DRV 6409), adult female, collected close to the village of Bemanevika, 14.3600��S, 48.5903��E, 1538 m above sea level, on the night of 28 June 2010 by M. Vences, D.R. Vieites, R. D. Randrianiaina, F. M. Ratsoavina, F. Randrianasolo, S. Rasamison, A. Rakotoarison, E. Rajeriarison and T. Rajoafiarison; ZSM 645/2014 (DRV 6324), adult female, collected at the Ambodikakazo campsite, 14.2098��S, 48.8981��E, 1473 m above sea level, on the night of 17 June 2010 by F. M. Ratsoavina and F. Randrianasolo; ZSM 51/2016 ��� 55/2016 (MSZC 57, 81, 104, 113, 141) and UADBA uncatalogued (MSZC 67 and 118), three males and four females from Ampotsidy, 15 km north of Bealanana, (14.412���14.422��S, 48.712���48.722��E), 1293���1489 m above sea level, collected at night between the 18th December 2015 and 4th January 2016 by M. D. Scherz, J. Borrell, L. Ball, T. Starnes, E. Razafimandimby, D. H. Nomenjanahary and J. Rabearivony. Etymology. The specific epithet fotsivava is a composite of the Malagasy words for white (fotsy) and mouth (vava). The name refers to the unpigmented oral mucosa of the species, as it was the first character we used to distinguish this new species to the other forms occurring in the Tsaratanana massif. The name is an invariable noun in apposition. Diagnosis. Uroplatus fotsivava sp. nov. is included in the Uroplatus ebenaui group of small-sized leaf-tailed geckos due to its triangular head with supraocular spines, laterally compressed body, and short tail. It differs from all members of the U. fimbriatus group (U. fimbriatus, U. giganteus, U. henkeli, U. sikorae and U. sameiti) and U. lineatus by its much smaller size (adult SVL 49.5���70.7 mm versus at least 85 mm), lack of lateral dermal fringing on any part of the body, and lateral compression of the body (versus depressed body shape with lateral dermal fringes), and a relatively much shorter tail. The evident triangular head, rather smooth skin and short tail distinguish U. fotsivava from U. alluaudi, U. guentheri, U. malahelo and U. pietschmanni. Within the U. ebenaui group, the new species differs from U. finiavana, U. malama and U. phantasticus by a shorter tail (adult TAL/SVL 0.15���0.32 vs. 0.42 in U. finiavana, 0.72 in U. malama, 0.62���0.76 in U. phantasticus) and by a narrower tail (TAW/SVL 0.07���0.10 vs. 0.14���0.16 in U. finiavana, 0.24 in U. malama, 0.16���0.20 in U. phantasticus). It is further distinguished from U. ebenaui, U. malama and U. phantasticus by its unpigmented oral mucosa (versus at least partly blackish pigmented oral mucosa). Uroplatus fotsivava shares an unpigmented oral mucosa with its sister species U. fiera, and has overlaps in values of relative tail length and tail width with this species. However, the ratio TAW/TAL distinguishes almost all specimens of the two lineages (0.17���0.31 in U. fotsivava vs. 0.19���0.21 in U. fiera; except for MSZC 0 118, all specimens of U. fotsivava have values of 0.23 or larger). Furthermore, U. fotsivava in comparison to U. fiera has a higher number of supralabials and associated scales (28���31 versus 27���28; U-test, P = 0.005) and a tendency of fewer lamellae under the third toe (5���8 versus 7���8; U-test, P = 0.065). Description of the holotype. Adult male in good condition with an intact tail and everted hemipenes. SVL 56.0 mm, tail length 16.2 mm, maximum tail width 4.2 mm, for further measurements see Table 1. Head triangular in dorsal view; canthus rostralis recognizable and concave; snout sloping strongly and continuously downward anteriorly; snout weakly depressed, short (1.2 times longer than eye diameter); eyes large (eye diameter 4.6 mm), bulging slightly above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small (horizontal diameter 1.0 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 dermal fringes; limbs without fringes but with miniscule spines on the hindlimbs, and practically no spines on the forelimbs; forelimb reaches beyond tip of snout when adpressed forward and approaches the groin when adpressed backwards along body (forelimb length/axilla���groin distance 25.2/26.6 mm = 95%), hindlimb reaches beyond axilla when adpressed forward along body (hindlimb length/axilla���groin distance 30.7/26.6 mm = 115%); original tail length 29% of snout���vent length, membranous borders of the denticulated tail more or less symmetrical, somewhat trapezoidal (i.e. broadening on either side and then narrowing again posteriorly), completely absent from the slightly spatulate tail-tip. Nares separated from each other by at least nine small granular scales, from the first supralabial by one scale, and from the rostral scale by one scale; first supralabial taller than the others; rostral entire, much wider than tall; mental scale very small, not differentiated from infralabial scales; 22/20 (right/left) true supralabials and 21/20 infralabials (as defined in methods above); no enlarged postmental scales or chin shields; dorsal and ventral scales of head, neck, body, limbs, and tail small, granular, juxtaposed and largely of uniform size interspersed with a few larger and partly raised tubercles, except for the irregular lines on the head and body which consist of series of slightly enlarged scales. Two curved lines (rows of slightly enlarged scales) extending from the posterolateral parts of the head (nuchal region) converge on the neck forming a V-shaped pattern (neck triangular line). A similar, moderately distinct, angular and medially straight line (also formed by a row of slightly enlarged scales) is present between the eyes and connects the supraocular spines. Two additional transverse lines, consisting of slightly enlarged scales, are recognizable on the frontal region of the head. Numerous spines on the posterior and postorbital part of the head, on hindlimbs (ca. 30 per limb), a dermal flap on each knee, bearing a spine; no flaps or spines on elbow and forelimb; a prominent pointed flap on the posterior portion of each upper eyelid (supraocular spine); posterior border of eye fringed. Weakly hollowed axillary pits present. Scales of the posterior ventral abdomen enlarged and arranged more uniformly than other scales, almost forming proper rows. Hemipenes everted, bearing two lobes. Calyx with protuberance bearing honeycomb-like structures, especially on the asulcal side. Area of sulcus spermaticus is smooth. Each lobe with a dense field of pointed papillae at its apex. Coloration. After six and a half years of preservation in 70% ethanol the color pattern remains the same as in the living animal (Fig. 5) but its vivacity and intensity have faded. All dorsal surfaces are mottled beige in base color, with individual flecks of burnt umber. The tail is as the dorsum at its base, but beyond the hemipenes is dorsally lighter beige, with few small brownish spots. The dorsum has weakly defined posteroventrally oriented lighter and darker beige chevrons, running parallel to the ribs beneath them. Several burnt umber flecks are noteworthy, especially an hourglass-shaped marking on the snout, and a strong dark fleck on the mid-dorsum, as well as several asymmetrical flecks on other parts of the body, including the arms, legs, and nape. Several whitish flecks that were visible on the animal in life (Fig. 4) are no longer visible in preservative, particularly those on the flank. The V-shaped area on the nape is almost indistinguishable. Two light cream markings defined by brown borders are present below each eye, one at the posterior corner and one at the mid-eye, both descending to the supralabials. The supralabials are generally beige in color, but a few are burnt umber. The infralabials weakly alternate between cream and burnt umber. The venter, including the ventral limbs except the lower hindlimbs, is a lighter beige than the dorsum, and much less mottled. The venter bears several round cream spots defined by brown borders, with three pairs of such markings in the pectoral region, and other such spots on the posterior abdomen. The tail is slightly darker below than above, and has a few small cream flecks bordered with brown, and a few burnt umber spots. The oral mucosa is unpigmented and whitish in preservative. In life the eye was golden brown with dark striations. Variation. In general, the paratypes agree well with the holotype in morphology. For measurements, see Table 1. The number of head spines varies from 2���26, and tends to be associated with the overall spiny-ness of the specimens; those with more head spines tend to have more arm and leg spines as well, and generally have more heterogeneous scalation. Forelimbs can bear no spines, or as many as six, with at most a slightly raised scale on the elbow but never a clear spine. Hindlimbs can bear 2���30 spines, and the dermal knee flap almost always bears a spine. In males the tail is highly denticulated and tends to be asymmetrical, but it can be symmetrical as well; in females it is without denticulations and trapezoidal; this might reflect a general pattern of sexual dimorphism in tail shape in the U. ebenaui group that warrants further investigation (see also Ratsoavina et al. 2015). Internarial scales range from eight to nine, supralabials from 18���22, and infralabials from 17���21. The line on the head connecting the supraocular spines can be curved to a point, or almost straight. The coloration of the type series is immensely variable, and includes browns to reds, with some specimens being strongly contrastingly colored, while others have almost homogeneous brown coloration, sometimes with dark and light markings. Ventral coloration is equally variable, with the venter sometimes being cream, sometimes being brown, and the chin sometimes bearing weak posteriorly-oriented chevrons. The V-shape on the nape can be extremely dark and thus highly contrasting, or hardly noticeable. Cream markings below the eye can be present or absent. The identity of all paratypes from Ampotsidy (MSZC series) was ascertained by sequencing a fragment of the mitochondrial 12S rRNA gene (see Results); they showed no or negligible differentiation compared to the samples included in our main molecular phylogenetic analysis. ZSM 52/2016 (MSZC 0141) clustered in a separate haplotype lineage with ZSM 648/2014 from Ambinanitelo, whereas all other Ampotsidy samples clustered in the Analabe/Bemanevika haplotype lineage. ������continued on the next page TABLE ��. (Continueđ) Distribution. Summarizing all sites vouchered by molecular data, the species is known from six sites, all in mid- to high-elevations of the Tsaratanana area: (1) the type locality, Analabe forest; (2) Ambodikakazo; (3) Ambinanitelo; (4) Bemanevika; (5) Ampotsidy; and (6) Manarikoba forest (Antsahamanara campsite; located at 1100 m above sea level; Andreone et al. 2009). Taken together, these localities span an elevational range of 1100��� 1538 m a.s.l. Natural history. The holotype was found active and perching at night on a tree branch of the Analabe forest, as were the remaining specimens from this locality. The forest is isolated at the top of the Ambodimanga mountain, with some degradation especially towards the edge. It is a humid evergreen forest characterized by a few big trees and dense understory, and a series of small streams running down the mountain. In Ampotsidy individuals of the species were typically found in bushes and small trees, up to a maximum of four meters above the ground. Several females encountered in December 2015 were gravid, suggesting a mating period coinciding with December to January, or the start of the local rainy season., Published as part of Ratsoavina, Fanomezana M., Gehring, Philip-Sebastian, Scherz, Mark D., Vieites, David R., Glaw, Frank & Vences, Miguel, 2017, Two new species of leaf-tailed geckos (Uroplatus) from the Tsaratanana mountain massif in northern Madagascar in Zootaxa 4347 (3), DOI: 10.11646/zootaxa.4347.3.2, http://zenodo.org/record/1048750, {"references":["Raxworthy, C. J., Pearson, R. G., Zimkus, B. M., Reddy, S., Deo, A. J., Nussbaum, R. A. & Ingram, C. M. (2008 b) Continental speciation in the tropics: contrasting biogeographic patterns of divergence in the Uroplatus leaf-tailed gecko radiation of Madagascar. Journal of Zoology, 275, 423 - 440. https: // doi. org / 10.1111 / j. 1469 - 7998.2008.00460. x","Ratsoavina, F. M., Louis, E. E. Jr., Crottini, A., Randrianiaina, R. D., Glaw, F. & Vences, M. (2011) A new leaf tailed gecko species from northern Madagascar with a preliminary assessment of molecular and morphological variability in the Uroplatus ebenaui group. Zootaxa, 3022, 39 - 57.","Ratsoavina, F. M., Vences, M. & Louis, E. E. Jr. (2012) Phylogeny and phylogeography of the Malagasy leaf-tailed geckos in the Uroplatus ebenaui group. African Journal of Herpetology, 61, 143 - 158. https: // doi. org / 10.1080 / 21564574.2012.729761","Ratsoavina, F. M., Raminosoa, N. R., Louis, E. E. Jr., Raselimanana, A. P., Glaw, F. & Vences, M. (2013) An overview of Madagascar's leaf tailed geckos (genus Uroplatus): species boundaries, candidate species and review of geographical distribution based on molecular data. Salamandra, 49, 115 - 148.","Ratsoavina, F. M., Ranjanaharisoa, F. A., Glaw, F., Raselimanana, A. P., Miralles, A. & Vences, M. (2015) A new leaf-tailed gecko of the Uroplatus ebenaui group (Squamata: Gekkonidae) from Madagascar's central eastern rainforests. Zootaxa, 4006, 143 - 160. https: // doi. org / 10.11646 / zootaxa. 4006.1.7","Andreone, F., Glaw, F., Mattioli, F., Jesu, R., Schimmenti, G., Randrianirina, J. & Vences, M. (2009) The peculiar herpetofauna of some Tsaratanana rainforests and its affinities with Manongarivo and other massifs and forests of northern Madagascar. Italian Journal of Zoology, 76, 92 - 110. https: // doi. org / 10.1080 / 11250000802088603"]}

Phelsuma gouldi sp. nov. (Figs. 2���3) Holotype. ZSM 804 / 2010 (ZCMV 13056), adult female, collected in Anja Reserve (Fig. 1), 13 km south of Ambalavao 21 �� 51 ' 2.64 '' S, 46 �� 50 ' 33.80 '' E, 949 m a.s.l., Haute Matsiatra Region, Fianarantsoa province, southern central Madagascar, on 9 th December 2009 by Angelica Crottini, D. James Harris, Iker A. Irisarri, Alexandra Lima, Solohery Rasamison and Emile Rajeriarison. Paratypes. None. Diagnosis. A presumably medium-sized, greyish-brown Phelsuma (SVL 45.1 mm, TL 81.5 mm) with an irregular pattern of brown to black lines and spots (Fig. 2). It differs from all other Phelsuma species, except for P. b o r a i, P. mutabilis, P. breviceps, P. standingi, and P. masohoala Raxworthy & Nussbaum by its cryptic life colouration without traces of green. Phelsuma gouldi differs from P. standingi by its smaller total length (TL 81 mm vs. up to 279 mm), number of supralabials (7 / 6 vs. 9���12), number of infralabials (6 vs. 7���8) and shows a distinct life colouration (brown-greyish dorsal colouration with irregular blackish stripes and dots vs. grey or bluish-green with dark reticulation and with greenish head and bluish tail). It differs from P. masohoala by the number of supralabials (7 / 6 vs. 8���9), number of infralabials (6 vs. 7), smooth dorsal scales on body and tail vs. keeled dorsal scales, a distinct life colouration (brown-greyish dorsal colouration with irregular blackish stripes and dots vs. a white and black pigmentation), and by a single V-shaped chevron along the lower suture of infralabials vs. up to three V-shaped chevrons on the throat. Phelsuma gouldi differs from P. breviceps by a slender snout vs. a very stout snout, by the number of transversely enlarged subdigital lamellae under the fourth toe of left/right foot (7 / 9 vs. 11 / 10), by the presence of a single Vshaped chevron along the lower suture of infralabials vs. absence, and by a set of throat scalation characteristics (for comparison see Fig. 3 of this study and Fig. 3 in Glaw et al. 2009): a triangular-shaped mental in P. gouldi (vs. a bell-shaped mental in P. breviceps), two postmental scales (vs. one postmental scale), four rows of enlarged postmentals (vs. one), the minimum number of scales needed to connect the suture between the second and third infralabial from the left to the right is 10 in P. gouldi vs. 14 in P. breviceps. The most similar species to P. gouldi are P. borai and P. mutabilis. Phelsuma gouldi differs from P. borai by a lower number of transversely enlarged subdigital lamellae under the fourth toe (7 / 9 vs. -/ 11), a lower number of supralabials (7 / 6 vs. 10 / 9), one internasal scale (vs. three), the absence of a distinct concave groove between the nasals (vs. presence), by the presence of a V-shaped chevron along the lower suture of infralabials (vs. absence), and by a different configuration of the throat scalation (see Fig. 3, comparison A with C): two postmental scales bordering the mental scale for about one-third in P. gouldi vs. two postmental scales border about one half of the mental scale in P. borai, the minimum number of scales needed to connect the suture between the second and third infralabial from the left to the right is 10 vs. 5, presence of 4 rows of enlarged postmentals vs. 5, and by a horizontally divided third infralabial vs. undivided. Phelsuma gouldi differs from the P. mutabilis specimens examined in Glaw et al. (2009) comprising thirteen specimens from Toliara (ZSM 945 / 2003, ZSM 948 / 2003), Toliara-Arboretum (ZSM 587 / 2000, ZSM 588 / 2000), Ampanihy-Tranoroa (ZSM 186 / 2004), unknown localities (MNHN 1895.152, MNHN 1895.154), Androy Nord (MNHN 1901.150, MNHN 1901.151) and Menabe (SMF 9470-9473) by a lower number of transversely enlarged subdigital lamellae under the fourth toe (7 / 9 vs. min. 9 / 10 to max. 11 / 11) and by a different configuration of the throat scalation (see Fig. 3, comparison A with B): triangular-shaped mental scale vs. bell-shaped, the minimum number of scales needed to connect the suture between the second and third infralabial from the left to the right is 10 in P. gouldi vs. 5 in P. mutabilis, 4 rows of enlarged postmentals vs. 3 rows, and by a horizontally divided third infralabial vs. undivided. Furthermore, P. gouldi differs from P. mutabilis, P. b o r a i, and P. breviceps, and all other Phelsuma species for which molecular data are available, by a substantial genetic differentiation (see Fig. 4). Description of the holotype. Well preserved, with regenerated tail autotomized after the first half. Autotomized tail part preserved separately. The tail tip (ca. 3 mm) was removed as a tissue sample. Body and head flattened dorsoventrally. Head slightly wider than neck, about as wide as body. Ear opening roundish. Tail shorter than snout-vent length, dorsoventrally flattened in cross section. No distinct tail whorls recognisable. Digits strongly expanded at tips, first finger and first toe vestigial, comparative finger and toe length 1 Measurements: SVL 45.1 mm; tail length 36.4 mm (plus ca. 3 mm that were taken as tissue sample); head width (at widest point) 8.6 mm; snout length (anterior edge of eye to tip of snout) 5.4 mm; horizontal eye diameter 2.1 mm; ear opening diameter 0.7 mm; eye-ear distance 3.1 mm; internarial distance 1.5 mm; nostril-eye distance 5.3 mm, axilla-groin distance 20.5 mm; forelimb length (from axilla to tip of longest finger) 14.6 mm; hindlimb length (from groin to tip of longest toe) 18.9 mm. Colouration: Colour after nearly one year in alcohol similar to that shortly after being euthanised (Fig. 2). Unfortunately no pictures of the holotype in life are available, but a photograph provided by H.-P. Berghof resembling the holotype and possibly showing a P. gouldi from near Betroka (Fig. 5) suggests that colouration in life and in preservative are probably similar as is also the case in related species. Ground colour of head, body, tail and dorsal parts of limbs dorsally and laterally brown-grey with irregular blackish stripes and dots. Several dark framed light brown spots dorsally on neck, body and limbs. A thin, dark medio-dorsal line extending from mid-body towards neck, where it divides into two thin dark lines extending in direction of snout tip converging between eye socket and nostril. A black band from nostril to anterior eye, continuing from posterior eye above ear opening, fading out as single thin dark line shortly behind neck. Another dark band between mouth corner and ear opening. A distinct dark lateral band starting from neck extending posteriorly until the tailbase, shortly behind the groin which marks the colour border between dorsal and ventral part of animal. Between axilla and groin and underneath the lateral band an irregular line of single spots, which continues on the tail where it fades out in posterior half. Supralabials and infralabials greyish-white with single dark pigmentations. Thin dark line along first two infralabials extending in posterior direction on gular scales, curving in direction to ear opening, fading shortly before that. Throat, chest, venter, and ventral parts of forelimbs and hindlimbs greyish-white. Toe tips on hand and feet dark grey-coloured. Distribution, conservation and IUCN Red List status. The new species is currently known only from the type locality within the Anja Reserve (Fig. 1), but further investigations are required to understand its actual distribution. It is possible that records of P. mutabilis from the central areas of Madagascar actually refer to P. gouldi. For instance, Hallmann et al. (2008) cite an observation by H.-P. Berghof who found P. mutabilis between the Horombe Plateau and the Andringitra massif at an altitude above 1000 m a. s. l. On request H.-P. Berghof (pers. comm.) was able to define this locality more precisely as near Betroka which is some 160 km south-west of the type locality, suggesting that P. gouldi might not be a local endemic of the Ambalavao region. In the area of the type locality there was no evidence of mineral or precious stone extraction or collecting for the pet-trade, but deforestation for agriculture, logging and cattle grazing was observed in all surrounding areas. It is likely that P. gouldi will qualify for inclusion in one of the threatened categories, but due to the currently restricted knowledge on this species we suggest to consider its conservation status as ���Data Deficient��� according to IUCN criteria (IUCN 2001). Habitat and habits. The holotype of P. gouldi was found around 4 p. m. on a tree trunk at a roosting height of about 2.5 meters. The tree was in an open and sunny spot inside a forest fragment. However, unpublished records report the sighting of P. mutabilis -like specimens on granitic rocks in Anja (B��hle, pers. comm.), and this record most probably refers to P. gouldi. No other Phelsuma species were found around the type locality. Etymology. We name this new species in honour of Stephen Jay Gould, a paleontologist, evolutionary biologist, historian of science and supreme writer of popular science who also provided invaluable contributions to the public appreciation of natural history and of science in general. Available names. Two junior synonyms are currently recognised for Phelsuma mutabilis and need to be considered as possible earlier available names for P. g o u l d i (see Hallmann et al. 2008; Glaw et al. 2009): Phelsuma androyensis Mocquard (originally described as Phelsuma androyense) and Phelsuma micropholis Boettger. The synonymy with P. mutabilis of both taxa was discussed in detail in Glaw et al. (2009). Based on comparisons of throat scalation and the number of transversely enlarged subdigital lamellae under fourth toe (11 / 10 and 11 / 11 in P. androyensis and 10 /11, 11/10, 10/10, 11/ 11 in P. micropholis; vs. 7 / 9 in P. gouldi) it is evident that these two nomina are not conspecific with Phelsuma gouldi (see Table 1 in Glaw et al. 2009 for additional details). Mitochondrial DNA variation, differentiation and phylogenetic relationships. The phylogenetic analyses resulted in a tree with largely unresolved basal relationships (Fig. 4) but with good support for the monophyly of P. mutabilis samples from across its range and clearly indicating the distinctness of P. gouldi from all the other three currently recognised species of the P. mutabilis group. The uncorrected genetic distance of P. gouldi to the three other Phelsuma species is: 8.1 % to P. mutabilis, 12.8 % to P. breviceps, and 8.5 % to P. borai. The phylogenetic analyses identified three major mitochondrial clades within P. mutabilis. Clade 1, although not resolved, encompasses samples of P. mutabilis from the south east (Tranomaro), south west (Ifaty and Ejeda), and west (Makay and Antsalova); clade 2 includes the samples of P. mutabilis from the north west (Ankarafantsika) and from a locality between Tranoroa and Ampaniny (south west); finally, only samples coming from Toliara in the south west belong to the clade 3 (see Fig. 4). Genetic distances within P. mutabilis amount to 4.7 % between the clades and range between 0.2 % to 4.5 % within clades., Published as part of Crottini, Angelica, Gehring, Philip-Sebastian, Glaw, Frank, Harris, James, Lima, Alexandra & Vences, Miguel, 2011, Deciphering the cryptic species diversity of dull-coloured day geckos Phelsuma (Squamata: Gekkonidae) from Madagascar, with description of a new species, pp. 40-48 in Zootaxa 2982 on pages 42-46, DOI: 10.5281/zenodo.208124, {"references":["Hallmann, G., Kruger, J. & Trautmann, G. (2008) Faszinierende Taggeckos. Die Gattung Phelsuma. Second Edition. Natur und Tier-Verlag, Munster, Germany, 253 pp.","IUCN (2001) IUCN red list categories and criteria. Version 3.1. IUCN Species Survival Commission, IUCN, Gland, Switzerland and Cambridge, United Kingdom, 30 pp."]}

Blaesodactylus ambonihazo sp. nov. ( Figures 2���5) Holotype. ZSM 469 / 2001 (field number MV 2001 - 340), adult male (Fig. 2 A); Ankarafantsika National Park (Ampijoroa), Mahajanga Province, northwestern Madagascar (16 �� 18 ���S, 46 �� 49 ���E, 173 m above sea level), collected by M. Vences, D. Vieites, G. Garc��a, V. Raherisoa, and A. Rasoamamonjinirina, on 26 February 2001. Paratypes. ZSM 470 / 2001 (field number MV 2001 - 341), subadult/adult male, ZSM 471 / 2001 (field number MV 2001 - 345), adult female; both with same data as holotype; ZFMK 51141, collected by F. W. Henkel and J. Sameit in 1988-1989; ZFMK 52290, collected by F. W. Henkel and W. Schmidt without precise collecting date; ZFMK 55257, collected by F. W. Henkel, without precise collecting date, three adult specimens from Ankarafantsika, Mahajanga Province, northwestern Madagascar, probably all females. Etymology. The species name is composed of the two Malagasy words hazo (tree) and ambony (on) and refers to the tree-dwelling habits of the new species (the terminal "y" of ambony is replaced by "i" as typical in Malagasy composite words). The species name is used as noun in apposition. Diagnosis. A mid-sized (max. SVL at least 108 mm), robust-bodied Blaesodactylus with tail much longer than SVL (Fig. 2). Snout elongate, gular granules small and regular, internarial region convex. 136���155 scale rows around midbody, 17���20 longitudinal rows of enlarged, keeled tubercles on dorsum. Fourth digit of pes with 15 undivided subdigital lamellae. 8���10 tubercles per row at base of tail, caudal tubercles limited to basal portion of tail. Body with 5���6 broad, more-or-less well-developed brown cross bands between nape and sacrum, original tail boldly banded, venter pale. Blaesodactylus ambonihazo sp. nov. may be differentiated from B. boivini by its smaller size (maximum SVL 108 mm versus 144 mm), homogeneous (versus heterogeneous) gular granules (Fig. 4) and uniform pale venter (versus mottled with areas of dark pigmentation). It differs from B. sakalava in its uniform (versus usually heterogeneous) gular granules, longer snout (Fig. 5 A) with internarial depression (versus convex internarial region), more restricted caudal tubercles (on proximal 40 % only versus entire length of original tail, and some dorsal tubercles bearing fine radial keels as well as a central longitudinal keel (Fig. 5 B; versus longitudinal keel only). It is distinguished from B. antongilensis, with which it has previously been confused by its uniform gular granules (versus enlarged anterior granules), lower number of dorsal tubercle rows (17���20 versus up to 24), and less strongly keeled and more widely spaced tubercles (2���3 intertubercular granules versus 1���2). Description of the holotype. ZSM 469 / 2001, adult male. SVL 107.9 mm. Skin torn on top of head, above left shoulder, and on right flank; muscle removed from ventral surface of right hind leg for tissue sample. Head relatively long (HeadL/SVL = 0.26) and very wide (HeadW/HeadL = 0.80), not depressed (HeadD/HeadL = 0.46); distinct from neck. Loreal region strongly inflated, frontonasal region strongly concave, snout elongate (SnEye/HeadL 0.46), blunt, longer than eye diameter (OrbD/SnEye = 0.54); scales on snout and forehead small, granular, heterogeneous, with minute rugosities or pustulations; scales on snout larger than those on occipital region except for scattered conical tubercles (~ 4���5 times size of adjacent scales); 24 scales across narrowest point of frontals, 54 between superciliary scale rows. Eye large (OrbD/HeadL = 0.25); pupil vertical with crenelated margins; superciliaries forming a short brillar fold with small spines at anterior and posterior margins. Ear opening obliquely oval, small (EarL/HeadL = 0.13); eye to ear distance longer than diameter of eye (EyeEar/OrbD = 1.23). Rostral quadrangular, much wider (4.6 mm) than high (2.3 mm), with no median groove. Enlarged supranasals separated by two transverse series of internasal scales, 3 anteriorly, bordering rostral, and 2 posteriorly; rostral in contact with first supralabials, supranasals, and 3 internasals; nostrils round, each surrounded by supranasal, rostral, first supralabial and two postnasals, and a crescentic nasal, itself bordered posteriorly by 3 postnasal scales; 3���4 rows of small scales separate rim of orbit from supralabials. Mental rhomboidal, wider (3.7 mm) than deep (2.1 mm); median pair of postmentals elongated (3.1 mm long), each bordered anteromedially by mental, medially in broad contact with other postmental along entire length, bordered anterolaterally by first infralabial, laterally by second postmental, posteriorly by 3 enlarged chin scales (5 in total, median chinshield bordered by both median postmentals); 11 (right) to 12 (left) supralabials, 11 (right) to 12 (left) infralabials on both side. Right side of head posterior to torn skin with healed wound exhibiting early stages of scale differentiation. Body stout, relatively short (TrunkL/SVL 0.43) with moderately developed ventrolateral folds. Dorsal scales smooth, granular to conical and recumbent, juxtaposed; 136 scale rows around midbody, intermixed with enlarged, smooth to longitudinally keeled to tubercles (~ 6 times size of adjacent scales, largest on flanks, and smallest in occipital region), some with fine radial striations, extending from occipital and postorbital regions to tail base, more conical and strongly keeled over sacrum and on tail base; tubercles in 17���20 rows at midbody. Ventral scales much larger than dorsals, smooth, imbricate, free margin rounded on chest, becoming more pointed posteriorly, largest on abdomen except for rows immediately anterior to vent; 34 scale rows across venter between ventrolateral folds; gular region with relatively homogeneous, smooth, rounded to oval scales, juxtaposed anteriorly to subimbricate posteriorly. No precloacal or femoral pores, no enlarged femoral scales. Preaxial scalation of limbs subimbricate, postaxial granular, dorsal surface of limbs without tubercles. Scales of palms and soles smooth, flattened, round, subimbricate. Limbs short and robust (ForeaL/SVL 0.13; CrusL/SVL 0.19). Digits broadly dilated, distal portion of digits II��� V free of pad, bearing a prominent recurved claw partly sheathed between a pair of scales, distal portion of digit I not free of pad, claw minute and lying in a groove in the adhesive pad; number of broad lamellae beneath each digit (9��� 13 ��� 14 ��� 14 ��� 12 manus; 12���13 ��� 14���15 ��� 11 pes); all lamellae, except distalmost slightly bowed, undivided except for distalmost lamellae of digit I; interdigital webbing weakly developed. Relative length of digits (manus): IV> III> III> II>V> I; (pes): IV>III> V> II> I. Mostly regenerated tail slender, tapering to tip; much longer than snout-vent length (TailL/SVL = 1.35), dorsoventrally depressed. Tail base with 3 smooth cloacal spurs on each side, largest anteriormost. Scales of tail dorsum heterogeneous ��� rectangular to pentagonal or hexagonal, subimbricate; midventral subcaudal scales transversely enlarged but less than half of tail width; tail segmented, each segment 8 dorsal scale rows and 3 transversely enlarged subcaudal scales in length; posterior edge of each segment with a row of enlarged keeled tubercles, 10 across basalmost segment, decreasing in number distally; lateral tubercles decreasing in size and becoming keelless on distsal portion of original tail. Regenerated portion of tail with some segmentation evident but scalation irregular and tubercles lacking. Hemipenes partly everted, pedicel and truncus not well demarcated from one another, apex bilobed, surface of apical lobes granular, lacking distinctive calyces. Coloration (in preservative). Body dorsum grayish brown with a series of weakly developed darker brown crossbars, one across occiput and nape, one across shoulders, two between axilla and groin, and one on anterior part of sacral region, another bar on tail base. A faint pale vertebral line bisecting anterior 3 crossbars. Flanks with some faint mottling. Limbs mottled. Tail with faint alternating brown and white bands, becoming bolder at approximately one third of tail length from cloaca, faint again on distal portion of tail. Head medium brown with beige labial scales, rictus, and posteroventral orbital margin. Venter cream, palms and soles grayish. Coloration in life (Fig. 3) generally similar to that in preservative, but darker and more contrasting. The iris was amber in colour. Variation. Variation in mensural and selected meristic features is summarized in Table 1. The ZSM paratypes shared the same number of ventral scale rows and internasal scales as the holotype and the male paratype ZSM 470 / 2001 had the same number of cloacal spurs. Caudal tubercles on the original tail of ZSM 470 / 2001 are in a maximum of 8 per row and are present only on whorls of the basal approximately 40 % of the tail. Dorsal coloration of male paratype ZSM 470 / 2001 is much bolder than in the holotype, with chocolate brown cross markings on a beige background, 3 crossbars between axilla and groin, and 12 brown bands on the postpygal portion of the tail, very bold from the fourth band distally (Fig. 2 B). Crown of head with diffuse, irregularly distributed cross markings; internasal and anterior frontal region dark brown; a vague dark stripe on each loreal region to anterior midpoint of orbit, continuing behind eye and above ear to join with brown nape marking. Female paratype ZSM 471 / 2001 with faded pattern similar to holotype, nape marking faint, partly fused with shoulder crossbar (Fig. 2 C). No enlarged cloacal spurs, endolymphatic sacs greatly enlarged and bulging along neck. Both paratypes exhibit some skin damage; on the left chest of the male and on the nape of the female, which also has a healed wound on the right flank. The hemipenes are everted in ZSM 470 / 2001. No detailed mensural data were taken from the ZFMK paratypes, but all diagnostic characters were verified in these specimens. The three specimens are in good state of preservation. SVL is 94.4 mm for ZFMK 55257, 99.4 mm for ZFMK 52290, and 97.1 mm for ZFMK 51141. The three specimens have 17���18 dorsal tubercle rows and uniform gular granules. Their tails are either regenerated or damaged, but in one of the specimens it is recognizable that the caudal tubercles become indistinct after the proximal one-third of the tail. Fine radial keels on some dorsal tubercles are present and especially clear in ZFMK 52290, less distinct in the other two specimens possibly because of fixation differences. Photographs of Blaesodactylus taken by A. Hartig at Ankarafantsika (not shown) confirm the characters described as diagnostic for the new species. Four of the five photographed individuals have original tails with tubercles visible on the proximal part of the tail only. All original tails are boldly banded either with black and white or with black and brown. The colouration of the back varies from indistinctly marbled with different tones of brown to a distinct pattern of largely symmetrical whitish blotches or even an almost rectangular network pattern. Mori et al. (2006) provided a colour photograph of one specimen and morphological data for nine Blaesodactylus individuals from Ampijoroa, most likely belonging to B. ambonihazo. According to their data snout-vent length ranged from 66.6-108.3 mm (mean 84.7 mm), tail length (original and regenerated) from 40-150 mm (mean 99.5 mm), and body mass was 6.7-30.8 g (mean 13.9 g). Distribution. So far, the species is reliably only known from Ankarafantsika National Park in north-western Madgascar, but we assume that it is more widespread and might occur in other remnants of dry forest in this largely deforested region (see Discussion below). Natural history. Specimens were collected on large tree trunks around Lac Ravelobe, next to Ampijoroa forest station. No further details of the collection circumstances were noted. Mori et al. (2006) observed individuals of Blaesodactylus most likely belonging to B. ambonihazo in the Ampijoroa forest within the Ankarafantsika reserve, perching on trees in the "Jardin Botanique A" and on walls of the buildings in the Ampijoroa station. These authors found seven individuals on trees in the morning which were partially hidden in shelters or crevices in shade and seemed to be inactive. These observations suggest some diurnal activity in this generally nocturnal species., Published as part of Bauer, Aaron M., Glaw, Frank, Gehring, Philip-Sebastian & Vences, Miguel, 2011, New species of Blaesodactylus (Squamata: Gekkonidae) from Ankarafantsika National Park in north-western Madagascar, pp. 57-68 in Zootaxa 2942 on pages 60-65, DOI: 10.5281/zenodo.205273, {"references":["Mori, A., Ikeuchi, I. & Hasegawa, M. (2006) Herpetofauna of Ampijoroa, Ankarafantsika Strict Nature Reserve, a dry forest in northwestern Madagascar. Herpetological Natural History, 10 (1), 31 - 60."]}

Blommersia variabilis sp. nov. Holotype. ZSM 237 / 2010 (field number FGZC 4305), adult male (Fig. 5), from Ambodivoahangy, northwest of Maroantsetra, northeastern Madagascar (coordinates: 15 �� 17 ��� 23.8 '' S, 43 �� 37 ���13.0'' E, below 50 m a.s.l.), collected on 3 April 2010 by P.-S. Gehring, F. Glaw, J. K��hler and M. Pabijan. Paratypes. Eleven adult males: ZSM 236 / 2010 (field number FGZC 4289), UADBA uncatalogued (FGZC 4290), UADBA uncatalogued (FGZC 4306), with same data as holotype; ZFMK 52608���52610, collected by F. Glaw and M. Vences on 18 March 1991 in the coastal town of Maroantsetra (15 �� 26 ' S, 49 �� 44 ' E, ca. 20 m a.s.l.); ZFMK 52612���52615, collected by F. Glaw and M. Vences on 19 March 1991 at a site slightly inland from the coastal village of Voloina (15 �� 34 ' S, 49 �� 37 ' E) which is located south of Maroantsetra; ZMA 19509 (field number FGMV 2002.2214) collected by M. Vences and A. Sarovy on 12 February 2003 in Maroantsetra. Diagnosis. Assigned to the genus Blommersia in the Mantellidae by a combination of (1) presence of intercalary elements between ultimate and penultimate phalanges of fingers and toes (verified by external examination), (2) presence of femoral glands and absence of nuptial pads in males, (3) presence of a moderately distensible, inconspicuously coloured single subgular vocal sac in males, (4) small size (adult SVL Blommersia, B. variabilis is distinguished from B. blommersae by vomerine teeth present (vs. absent); from B. domerguei by different colouration (absence of light brown dorsum with dark longitudinal markings) and vomerine teeth present (vs. absent); from B. grandisonae by different colouration (absence of distinct black lateral line ventrally bordered by white, vs. presence) and vomerine teeth present (vs. absent); from B. kely by larger size (male SVL 19���25 mm vs. 14���16 mm) and vomerine teeth present (vs. absent); from B. sarotra by larger size (male SVL 19���25 mm vs. 15���17 mm) and vomerine teeth present (vs. absent); from B. angolafa by colouration (absence of uniformly brownish dorsal colour with white dots) and vomerine teeth present (vs. absent). The new species is morphologically most similar to B. wittei from which it mainly differs by the wider distance between inner margins of femoral glands (FGD 2.8���4.5 mm vs. 1.6 ���2.0 in the type series of B. wittei; see Vences et al. 2010; FGD 12���20 % of SVL vs. 7���9 %), connected metatarsalia in most specimens (vs. always separated by webbing), and advertisement calls consisting of a short series of 3-6 notes of a mean duration of 107 ms and a mean inter-note interval of 27 ms (vs. longer series of up to 25 notes with a mean duration of 23-46 ms and a mean interval duration of 30���72 ms). The new species forms a monophyletic group with the recently described B. dejongi and B. galani. It differs from B. galani by the presence of vomerine teeth (vs. absence) and smaller femoral glands (see Fig. 4; femoral gland length 3.5��� 5.2 vs. 4.8���9.2 mm). It differs from B. dejongi by a shorter distance between the inner margins of femoral glands (Fig. 4; FGD 2.8���4.5 mm vs. 7.8���9.8 mm). Furthermore, the new species differs from all nominal species of Blommersia by a substantial genetic divergence and by its advertisement call (as described below). Description of the holotype. Specimen in a good state of preservation, tongue removed as tissue sample for molecular analysis. SVL 23.7 mm, for further measurements see Table 1. Body slender; head longer than wide, wider than body; snout rounded in dorsal and lateral views, nostrils directed laterally, slightly protuberant, nearer to tip of snout than to eye; canthus rostralis distinct, straight; loreal region slightly concave; tympanum distinct, rounded, its diameter 54 % of eye diameter; supratympanic fold distinct posterior to tympanum, indistinct between eye and tympanum, curved; tongue removed, its shape therefore not verifiable; vomerine teeth present as two distinct protuberances immediately posterior-medially of choanae, maxillary teeth present; choanae small, rounded. Arms slender, subarticular tubercles single; fingers without webbing; relative length of fingers 1 et al. (2000), consisting of 17 distinct granules as verified from internal view (after reflexing of the ventral skin on thigh), distance between femoral glands 2.8 mm. Table 1. Measurements of the type series of Blommersia variabilis sp. nov. and comparative specimens of B. wittei (all males). ZSM 237 / 2010 is the holotype, all other B. variabilis specimens are paratypes. For abbreviations, see Material and Methods. Relative hindlimb length (RHL) is coded as follows: when hindlimb is adpressed along body, tibiotarsal articulation reaches (1) anterior eye corner, (2) eye center, (3) between eye and nostril, (4) nostril, (5) snout tip. After nine months in preservative, the dorsum appears greyish-brown because of many densely spaced, single dark melanophores. A translucent pink colour covers the entire dorsum, but is more prominent between and around the eye sockets and the tympanum. A thin, single medio-dorsal whitish line stretches from the neck in a posterior direction until the cloaca. There is a moderately distinct colour border between the flanks and the lighter dorsum. The hind limbs are light brown with four distinct dark brown crossbands on the thighs, three indistinct crossbands on tibia; tarsus light brown. The arms show indistinct, irregular dark patterns. Posterior to the tympanum a lateral dark streak runs along the supratympanic fold and ends at the forelimb insertion. Ventrally, the holotype was whitish with single, small and irregular dark melanophores. Throat almost uniformly whitish. Femoral glands largely white, contrasting with the yellowish colour of the inner thighs. In life the holotype (Fig. 5) had a brown dorsum with a reddish hue on the anterior half of the body, including the head and neck, whereas the posterior is grayish-brown. Indistinct, gray crossbands line the limbs and a distinct white vertebral stripe started in the neck region and continued down the length of the body. Iris golden dorsally, ventrally poorly recognizable due to the widely opened pupil. Ventrally, the throat is yellow, but the rest of the body is white with some yellow mottling along the sternum and along the sides. The undersides of the front limbs are pinkish-blue. The ventral side of the hindlimbs is a darker blue, especially on the digits and joints. The ventral side of the thighs is pinkish-blue with yellow, clearly discernible femoral glands. The skin on the venter was translucent such that larger blood vessels could be easily discerned. The ventral colouration extends slightly onto the flanks, merging rather indistinctly with the dorsal brown colour. Remark. This species was referred to as Blommersia sp. aff. blommersae ���Maroantsetra��� by Glaw & Vences (2007), Blommersia sp. aff. wittei (Maroantsetra) by Vences et al. (2006), and Blommersia sp. 2 Maroantsetra by Vieites et al. (2009) and Vences et al. (2010). Variation. Morphological variation of nine paratypes is provided in Table 1. In general the morphology of the paratypes was similar to the holotype. However, there is some variability in the separation of the lateral metatarsalia. For instance, in ZFMK 52608 they are separated, whereas in other specimens they are partly connected, and ZFMK 52612 shows connected metatarsalia on the left foot and nearly completely separated metatarsalia on the right foot. Dorsal colouration of the paratypes is rather variable. The reddish hue characterizing the holotype is not typically observed in other specimens which are usually light brown with dark brown markings. A narrow white median line can be present on the dorsum as well as a broader band that becomes triangular at the anterior part of the dorsum, covering the entire surface of the head. Brown or grey markings and spots can be present on the venter. In 2003 we also collected one specimen of very large body size in Maroantsetra, mixed with normal-sized specimens. Because of a labeling error we cannot ascertain with full reliability the current identity of this specimen and therefore did not include it in Table 1, almost certainly it is the male ZMA 19511 of 29.0 mm SVL with largely separated outer metatarsalia (intermediate between the connected and separated state). Etymology. The species name variabilis is an adjective referring to the remarkable variation observed in various characters of this new species, especially in the separation or connection of the lateral metatarsalia which otherwise is considered to be a very stable character within species of mantellids. Natural history. At the type locality in the vicinities of Ambodivoahangy village, B. variabilis was common outside of the forest in cultivated landscape. It was not seen or heard within primary or selectively logged forest. Calling males were observed in dense secondary vegetation on the borders of partially inundated ricefields, about 30 cm to 1 m off the ground. The male holotype was discovered while calling within a dense bush at the edge of a rice field. During the day specimens were difficult to find, hiding underneath the leaves of plants. Calling was most frequent in the evening and at night, but was also occasionally heard during the day, especially after rains. Calls were often intermingled with choruses of Aglyptodactylus sp., Guibemantis cf. kathrinae and Ptychadena mascareniensis. At Maroantsetra it is a common species living in ditches covered by dense vegetation within the town itself. Near Voloina we heard and collected the species in flooded areas in a mosaic of rice paddies, shrubs, and remains of primary rainforest. A single clutch of eggs deposited on the surface of a leaf was discovered within the village of Ambinanitelo on 1 April 2010. Mating in this species takes place on a leaf usually overhanging lentic water and has been observed by us in Voloina. As in other Blommersia, the male and female sit vertically on a leaf, the male on top of the female with his legs extending over the female���s dorsum (Glaw & Vences 1994). Vocalization. The advertisement call of B. variabilis (Fig. 6) was recorded at the type locality (Ambodivoahangy) on 3 April 2010 at an estimated air temperature of 26 ��C. It consists of a series of short inharmonious notes, repeated at regular intervals in fast succession with a repetition rate of approximately 7.4 notes/second. Each call is composed of 3���6 notes, whereas in most cases a long call is followed by a shorter call containing fewer notes, in combination resulting in a call series with two calls. Within these call series, the interval between first and second call ranges between 500 and 720 ms. Within calls, the initial note is longer and contains more pulses than subsequent secondary notes. Numerical call parameters are as follows: call duration 344���887 ms (532 �� 219; n = 9); note duration 54���223 ms (107 �� 48; n = 23); pulses/note 3���14 (6.4 �� 3.0; n = 23); inter-note interval 21���33 ms (26.7 �� 3.2 ms; n = 19). Overall frequency is distributed in a broad band from app. 2000���6500 Hz, with a dominant frequency at around 5100 Hz. Calls were repeated at irregular intervals. Similar calls were recorded in 1991 in Maroantsetra, but due to the poor quality of the recordings we refrain from a detailed analysis. Detailed call data for various populations of B. galani, B. dejongi, and B. wittei are presented in Vences et al. (2010). In comparison, the call of B. variabilis differs from that of B. wittei by longer note duration, shorter internote intervals and a lower note repetition rate. Calls of B. galani differ from those of B. variabilis by shorter notes and much longer inter-note intervals. In calls of B. dejongi, notes are barely spaced and partly fused and separate pulses within notes are unrecognizable. Distribution. B. variabilis is known from (1) Maroantsetra, (2) Voloina, (3) Ambodivoahangy and (4) Ambohinantely (15 �� 20 ' 50.71 '' S, 49 �� 35 '02.84'' E). The largest straight line distance between any of these sites is approximately 30 km (Ambodivoahangy-Voloina). The altitude was not precisely measured at the collecting localities, but all except Voloina are situated below 50 m a.s.l. We estimate an elevation of 100-300 m a.s.l. for the collecting locality near Voloina. The Voloina site lies close to a medium-sized river, but due to the lack of reliable coordinates for our precise collecting locality in 1991, we cannot ascertain if our samples originate from north or south of the river. The remaining three sites are in the immediate vicinity of the Antainambalana river., Published as part of Pabijan, Maciej, Gehring, Philip-Sebastian, K��hler, J��rn, Glaw, Frank & Vences, Miguel, 2011, A new microendemic frog species of the genus Blommersia (Anura: Mantellidae) from the east coast of Madagascar, pp. 34-50 in Zootaxa 2978 on pages 41-45, DOI: 10.5281/zenodo.207732, {"references":["Blommers-Schlosser, R. M. A. (1979) Biosystematics of the Malagasy frogs. I. Mantellinae (Ranidae). Beaufortia, 352, 1 - 77.","Glaw, F., Vences, M., & Gossmann, V. (2000) A new species of Mantidactylus from Madagascar, with a comparative survey of internal femoral gland structure in the genus (Amphibia: Ranidae: Mantellinae). Journal of Natural History, 34, 1135 - 1154.","Glaw, F., & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Third edition. Vences & Glaw Verlag, Cologne, 496 pp.","Glaw, F., & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Second edition. Vences & Glaw Verlag, Cologne, 480 pp."]}

Phelsuma klemmeri group (contains P. klemmeri) Affinities of this species to the P. lineata and P. laticauda groups are apparent and well-supported from our analysis (Fig. 7), but mitochondrial as well as nuclear markers place P. klemmeri at a basal position within the clade containing these two species groups. Because its colouration is very characteristic and strongly differs from all other Phelsuma, the isolated position of this species is best reflected by placing it in a separate species group. Phenotypically, several characters of P. klemmeri are in agreement with its affinities: non-gluing behaviour and absence of nostril-rostral contact are shared with the P. lineata and P. laticauda groups, and presence of smooth ventral and subcaudal scales, number of nasalia (3) as well as of size-reduced vertebral scales are similar to the P. laticauda group (Tables 1���2)., Published as part of Rocha, Sara, R��sler, Herbert, Gehring, Philip-Sebastian, Glaw, Frank, Posada, David, Harris, James & Vences, Miguel, 2010, Phylogenetic systematics of day geckos, genus Phelsuma, based on molecular and morphological data (Squamata: Gekkonidae), pp. 1-28 in Zootaxa 2429 on page 23, DOI: 10.5281/zenodo.194693

Phelsuma astriata group (contains P. astriata and P. sundbergi) These two species form a small endemic radiation on the Seychelles islands and various subspecies have been described for both. Variation within this Seychellois group was studied by Cheke (1982), Gardner (1984, 1986, 1987), and more recently, by Radtkey (1996). The available molecular data, nuclear and mitochondrial, is unambiguous in supporting monophyly of this group (Fig. 5: clade H), which furthermore was recovered in single-gene analyses of all genes except C- mos, where differentiation relative to other species in clade E (see Fig. 5) is very low. The mitochondrial and combined analyses indicate that, possibly, Phelsuma vanheygeni is the sister species of the Seychellois clade and may be in the future included in the P. astriata group (see section on this species below). Species in the Phelsuma astriata group are all green coloured with a weak dorsal pattern and no or only weak dorsal or lateral lines, keeled dorsalia and lateralia, size-reduced vertebral scales, and are non-gluers, but otherwise appear to be quite variable in morphological characters (Tables 1���2). Relationships within the P. madagascariensis group This group is in a well-supported clade (Fig. 5; clade E) with the Seychellean P. astriata group, and with P. vanheygeni. The basal topology of clade E is uncertain; the position of both P. vanheygeni and P. guttata is unresolved, mainly because of conflicts between the mitochondrial and nuclear datasets. In the mitochondrial, as well as in the combined data, P. vanheygeni is sister taxon to the P. astriata group and P. guttata is placed in the P. madagascariensis group. The nuclear data, however, place P. vanheygeni basal in clade E, and P. guttata completely outside of it, albeit both placements with less than 50 % PP/BS support. Because of its morphological affinities to the P. madagascariensis group we nevertheless include P. guttata in this group, but we emphasize that further study on its phylogenetic position is needed. Within the P. madagascariensis group, the P. guttata species group as previously defined (P. g u t t a t a, P. abbotti, P. masohoala, and P. seippi; Glaw & Vences 1994; Glaw et al. 1999; Van Heygen 2004) does not form a monophyletic group; instead, P. guttata is placed basal in the group, and P. s e i p p i and P. abbotti also do not appear to be sister species. Two members of this group originated by (long) distance colonization in a northwestwards direction: P. abbotti abbotti (Aldabra) and P. parkeri (Pemba). Both have their closest relatives in the northwestern/northern Malagasy coast, the most likely area in fact from which colonizations to occur from, considering surface sea-currents in the region (Ali & Huber 2010). One of the few nominal Phelsuma species for which no DNA sequences are available until now is P. masohoala. As discussed by Meier & B��hme (1996) it is possible that P. masohoala is related to P. abbotti and thus belongs to the P. madagascariensis group where we provisionally include it. Relationships among P. madagascariensis, P. grandis, and P. k o c h i In our molecular analysis, Phelsuma grandis, P. kochi, and P. madagascariensis which were long considered subspecies of a single species (P. madagascariensis) did not form a clade, although the mitochondrial and nuclear data suggested different relationships among these taxa. This confirms the results of Raxworthy et al. (2007) who recently proposed to elevate them to species level, based on data from a molecular analysis and from environmental niche modelling. Neither Raxworthy et al. (2007) nor ourselves had sequences of Phelsuma madagascariensis boehmei Meier, 1982, and the status of this taxon thus remains unclarified. We detected deep genetic divergences among P. grandis, P. kochi, and P. madagascariensis, reaching up to 14 % (uncorrected p-distances from cytochrome b). Within P. grandis and P. kochi distances of up to 1.1% and 9 % were, respectively, observed for cytochrome b. These surprisingly high distances prompted us to undertake a wider survey of variability in the P. madagascariensis group, based on mitochondrial 16 S rDNA sequences. The three taxa, P. grandis, P. kochi, and P. madagascariensis are large geckos that are thought to be allopatrically distributed mainly in coastal Madagascar: P. madagascariensis in the east, P. grandis in the north, and P. kochi in the west (Meier & B��hme 1991), with contiguous distributions and contact zones between them. The new data (Figs. 5 and 6) add various new perspectives to this picture: First, we detected a second deep lineage that we here assign to P. kochi in a preliminary way: a specimen (identified as P. k o c h i) from Manongarivo in the Sambirano region, far northwestern Madagascar, differed strongly from specimens from the west (Tsingy de Bemaraha and Ankarafantsika). For 16 S alone, it did not even form a monophyletic group with the remaining P. k o c h i (Fig. 6), and although it did in the combined analysis (Fig. 5), support was not significant (PPP. madagascariensis were deeply differentiated, with an eastern-southeastern clade (from Ambohitsara and Nosy Boraha) and a north-eastern clade (from Ankalampo), and with a differentiation between these of 7.5% uncorrected p-distance in the 16 S gene (Fig. 6). Differentiation within P. abbotti A last aspect obvious from the multi-gene phylogeny (Fig. 5) and the 16 S data (Fig. 6) is the low differentiation among the various subspecies of Phelsuma abbotti. In Madagascar, the subspecies P. a. chekei shows a slight differentiation between specimens from the north (Ankarana and Montagne des Francais) and west (Tsingy de Bemaraha). Phelsuma a. abbotti and P. a. s u m p t i o from Aldabra and Assumption islands show some differentiation but certainly have colonized these islands very recently, as expected. Phenotypic characterization of the P. madagascariensis group Species in the Phelsuma madagascariensis group are characterized by non-gluing egg laying behaviour, smooth ventral and subcaudal scales, keeled dorsal and lateral scales absence of nostril-rostral contact, and size-reduced vertebral scales, whereas other characters are variable among species in the group, possibly in part related to the size differences (Tables 1���2). For example, the number of preanofemoral pores is higher in the larger species (P. grandis, P. kochi, P. madagascariensis, P. parkeri). Several characters such as nongluing behaviour, and absence of rostral-nostril contact are shared by the P. madagascariensis group and the P. astriata group, in accordance with their placement in a monophyletic group by the molecular data, but P. vanheygeni which also belongs into this clade is an egg-gluer (see below). Definition and content of the group Molecular data strongly corroborate the previously suggested (Glaw & Vences 1994) evolutionary relationships between P. lineata, P. quadriocellata and P. pusilla. Other species belonging to this well supported group are P. k e l y, the Comoran P. c o m o re n s i s, and P. antanosy (Fig. 7 a; clade J) Two subgroups can further be defined: (1) all P. quadriocellata subspecies (including P. quadriocellata bimaculata, sometimes regarded as a distinct species ��� Glaw & Vences, 1994) and P. antanosy and (2) P. lineata, P. pusilla, P. k e l y and P. c o m o re n s i s. Although nuclear genes reveal unresolved or paraphyletic relationships between these (Figure 7 c), for the faster evolving mtDNA markers these groups already reached reciprocal monophyly and are recovered with strong support (Fig. 7 b). Phelsuma antanosy was previously thought to be more closely related to either the species from the Mascarenes (Raxworthy & Nussbaum 1994), i.e., the P. cepediana group as defined here, or to the P. laticauda group (Glaw & Vences 1994) but in the molecular analysis it is clearly nested within the P. lineata group, sister to P. quadriocellata. Although no molecular data are yet available, we here assign the recently described (Berghof & Trautmann 2009) Phelsuma hoeschi to the P. lineata group based on its dark lateral stripe and its similarities to P. pusilla and P. kely in scale characters, as also emphasized in the original description (see also Tables 1��� 2). Loveridge (1942) hypothesized about the dark spots of P. quadriocellata being derived from the dark lateral band of P. lineata. It is worth mentioning that P. l. bombetokensis, a geographically isolated taxon from dry forest in north-western Madagascar (still unstudied from a molecular perspective), has an intermediate pattern in this respect. Variation within the P. lineata complex and recognition of P. dorsivittata at species level Phelsuma lineata, being paraphyletic in respect to P. k e l y, P. c o m o re n s i s and P. p u s i l l a (if lineata is considered as including dorsivittata as subspecies), exhibits consistent deep differentiation (12.8% uncorrected p-distances for cytochrome b) between P. lineata dorsivittata from the far north of Madagascar (including a specimen of uncertain status from Marojejy in north-eastern Madagascar) and the remaining individuals. Genetic distances within remaining P. lineata individuals for cytochrome b reach 7.7% uncorrected p-distances. Also, p-distances reach as much as 8.8% within P. lineata dorsivittata. The expanded analysis based on 16 S sequences only, but including many more specimens (those sequenced by Boumans et al. 2007, plus 16 others) largely confirms these conclusions (Fig. 8). Phelsuma kely and P. c o m o re n s i s are deeply nested within the P. lineata clade. A taxonomic consequence from this fact could be to accept a paraphyletic species P. lineata or, given the known colour differences between populations (Kr��ger 1996 c), its partitioning into various species. The following deeply differentiated clades can be distinguished: (1) P. lineata populations from the highlands (Antananarivo, Ambohitantely, Fierenana), which would correspond to P. lineata elanthana according to Kr��ger (1996 c), (2) populations from the south east (Ste. Luce), (3) populations from the northern central east and southern central east (Andasibe, Besariaka, Ranomafana, Ifanadiana, Ambohitsara, Farafangana), and (4) as the most divergent lineage, the populations from the north, currently corresponding to P. l. dorsivittata. However, P. lineata sensu lato is almost continuously distributed over most of Madagascar's rainforests, with probable contact and possibly hybrid zones, especially between population groups from the highlands and the northern central east. Furthermore, the nuclear genes provide no evidence for separating lineages 2 versus 3 (Fig. 7 c; highland specimens of lineage 1 not included). The available evidence is therefore insufficient for conclusive statements on the taxonomic status of the lineages 1���3. On the contrary, lineage 4 from the northern localities Montagne d'Ambre, Iharana (= Vohemar) and Marojejy is consistently recovered by mitochondrial and nuclear markers (Fig. 7 b���c) and is grouped basal to the clade containing P. l i n e a t a lineages 1���3, P. pusilla, P. kely and P. c o m o re n s i s (Fig. 7 a). We see this as sufficient support for a status of the northern populations as separate species, Phelsuma dorsivittata. Typical specimens of Phelsuma dorsivittata have several red spots on the back, which can often fuse to larger markings. Usually a red vertebral stripe is present on the anterior back. A rather indistinct dark lateral spot is present behind the forelimb, and can be also present in front of the hindlimbs, forming the anterior and posterior part of a dark lateral line (distinctly recognizable in the holotype). However, the anterior spot is not isolated and not bordered by blue colour in life. In Phelsuma dorsivittata 8���9 supralabialia (min-max) and 6���9 infralabialia (min-max) are present. Selected morphological characters of the male holotype (SMF 59373, Fig. 9) of Phelsuma dorsivittata are as follows: SVL 55.3 mm; TL 124.4; 9 / 9 supralabials on the left and right side of the head; 8 / 7 infralabials; 1 internasal; nostril in contact with rostral and first supralabial; 28 preanofemoral pores; mental triangle-shaped; dorsals, ventrals, and subcaudals keeled. For a more detailed description of the holotype see Mertens (1964). The holotype, upon examination in 2009 and based on the original description, shows the colour pattern as described above (distinct anterior dark marking on the flanks, red dorsal markings forming a midventral line on the anterior part of the back. However, a specimen from Marojejy (the same as included in our molecular trees) as well as another specimen from the type locality Montagne d'Ambre (see Glaw & Vences 2007: p. 403 and their fig 1 f) show a rather uniform dorsum with no or few red markings which do not form a longitudinal line. Whether this variation is related to the substantial molecular variation observed among these populations and among specimens from Montagne d'Ambre requires further study. Variation within the P. quadriocellata complex and recognition of P. parva at species level Within P. quadriocellata, uncorrected p-distances for cytochrome b reach 13.4% and three main clades seem to be distinguishable by the mtDNA markers (Figs. 7���8), although nuclear markers do not provide resolution (and show evidence of ancestral polymorphism) at this level. One of these likely corresponds to typical P. quadriocellata and appears to be distributed over much of the mid-altitude localities in the northern central east and southern central east, from Sahatelaka (near Lake Alaotra) to Ranomafana. A second clade was found in the lowlands of the southern central east and south east (Ifanadiana and Ste. Luce). The specimens from these two sites are distinctly smaller and have a different pattern of red dorsal markings than those from nearby Ranomafana, and thus corresponded to the current subspecies P. quadriocellata parva. Specimens from lower elevations in the northern central east (Nosy Boraha and Mahasoa) clustered together (uncorrected p-distance for the 16 S gene 2.2%) (Fig. 8) and correspond morphologically to two subspecies: specimens from Mahasoa possibly to P. q. lepida and specimens from Nosy Boraha possibly to P. q. bimaculata. Given that especially quadriocellata and parva occur in close geographic proximity (Ranomafana vs. Ifanadiana) but each of them is genetically and morphologically constant over wide areas of eastern Madagascar (parva: Ste Luce in the southern central east to Ifanadiana in the southern central east; quadriocellata: Ranomafana in the southern central east to Sahatelaka in the northern central east; Fig. 8), we hypothesize that these two taxa are distinct at the species level, and consider Phelsuma parva as full species. Average cytochrome b distances between P. q. quadriocellata and P. parva are high (11.9%), and average distances within P. p a r v a and P. q. quadriocellata are high as well, 7.3% and 5.6% respectively, for cytochrome b. Phelsuma parva is one of the smallest species within the genus Phelsuma; its total length does not reach more than 90 mm (Meier 1983; Glaw & Vences 2007). The body has a slender shape with a bright green colouration. It has distinct red spots on the back and a dark spot behind the forelimb that is variable in size, more or less round and often surrounded by a blue ring. Phelsuma parva is characterised by 7���10 supralabialia (min-max), 7���8 infralabialia (min-max). Selected morphological characters of the male holotype (ZFMK 19315, Fig. 9) of Phelsuma parva are as follows: SVL 36.2 mm; TL 80.6; 7 / 7 supralabials; 7 / 7 infralabials; 1 internasal; nostril in contact with rostral and first supralabial; 22 preanofemoral pores; mental triangle-shaped; dorsals, ventrals, and subcaudals keeled. For a more detailed description of the holotype see Meier (1983). All specimens collected by us and included in our molecular trees agree in size and colour closely with the holotype. Specimens assigned to the newly defined species Phelsuma parva do not form a monophyletic group in the analysis based on nuclear DNA sequences (Fig. 7 c). However, the same would also be true for P. quadriocellata sensu lato, if parva was seen as subspecies of such an inclusive species. Moreover, the fact that rather well-defined species such as P. antanosy, P. comorensis, P. pusilla, and P. kely cluster among individuals of P. parva and P. l i n e a t a in the nuclear gene tree indicates that these placements are probably due to incomplete lineage sorting in these complexes of species. The recognition of P. parva at species level also rises questions about the status of individuals here assigned to P. q. bimaculata and P. q. lepida which in the mitochondrial analysis form a clade with P. parva rather than P. q. quadriocellata (Fig. 7). These forms have distinct differences in colour patterns from each other, from P. q. quadriocellata, and from P. p a r v a. We have little doubts that they are to be assigned to one or possibly even two separate species. However, the nomenclatural situation is convoluted because (1) a clear definition of the taxon bimaculata based on a re-examination of type material is missing, and (2) our analysis does not include topotypical samples of lepida (which was described from near Andapa in the North-East), so that the identity of this taxon requires confirmation too. We therefore here refrain from taxonomic conclusions and postpone these to forthcoming studies. Phenotypic characterization of the P. lineata group The Phelsuma lineata group is characterized by a predominant green colour with usually a pattern of lateral/sublateral stripes, typically with keeled ventral and subcaudal scales (except P. lineata punctulata P. lineata elanthana and P. c o m o re n s i s), absence of rostral-nostril contact, a relatively low number of midbody scales (below 98, except for P. antanosy) and absence of size-reduced vertebral scales (Tables 1���2). The P. lineata group shares with the P. laticauda group (see below), which is its sister group (Fig. 7), the non-gluing behaviour and absence of nostril-rostral contact, but species in the P. laticauda group have smooth ventral and subcaudal scales, and several of them h, Published as part of Rocha, Sara, R��sler, Herbert, Gehring, Philip-Sebastian, Glaw, Frank, Posada, David, Harris, James & Vences, Miguel, 2010, Phylogenetic systematics of day geckos, genus Phelsuma, based on molecular and morphological data (Squamata: Gekkonidae), pp. 1-28 in Zootaxa 2429 on pages 14-23, DOI: 10.5281/zenodo.194693, {"references":["Cheke, A. S. (1982) Phelsuma Gray 1825 in the Seychelles and neighboring islands: A re-appraisal of their taxonomy and description of two new forms. Senckenbergiana biologica, 62, 181 - 198.","Gardner, A. S. (1984) The evolutionary ecology and population systematics of day geckos (Phelsuma) in the Seychelles. Unpublished Ph. D Thesis, University of Aberdeen.","Gardner, A. S. (1986) Morphological evolution in the day gecko Phelsuma sundbergi in the Seychelles: A multivariate study. Biological Journal of the Linnean Society, 29, 223 - 244.","Gardner, A. S. (1987) The systematics of the Phelsuma madagascariensis species group of day geckos (Reptilia: Gekkonidae) in the Seychelles. Zoological Journal of the Linnean Society, 91, 93 - 105.","Radtkey, R. (1996) Adaptive radiation of day-geckos (Phelsuma) in the Seychelles Archipelago: A phylogenetic analysis. Evolution, 50, 604 - 623.","Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Second Edition. Vences and Glaw Verlag, Koln, 480 pp.","Glaw, F., Vences, M. & Ziegler, T. (1999) Bemerkungen zu Phelsuma dubia (Boettger, 1881): Wiederentdeckung des Holotypus, Verwandtschaftsbeziehungen und Daten zur Fortpflanzung. Salamandra, 35, 267 - 278.","Van Heygen, E. (2004) Remarks on the Phelsuma barbouri- and Phelsuma klemmeri- phenetic groups. Phelsuma, 12, 153 - 154.","Ali, J. R. & Huber, M. (2010) Mammalian biodiversity on Madagascar controlled by ocean currents. Nature, 653 - 656.","Meier, H. & Bohme, W. (1996) Zum taxonomischen Status des Formenkreises von Phelsuma abbotti Stejneger, 1893, mit Bemerkungen uber P. masohoala Raxworthy & Nussbaum, 1994. Salamandra, 32, 85 - 98.","Raxworthy, C. J., Ingram, C. M., Rabibisoa, N. & Pearson, R. G. (2007) Applications of ecological niche modelling for species delimitation: A review and empirical evaluation using day geckos (Phelsuma) from Madagascar. Systematic Biology, 56, 907 - 923.","Meier, H. (1982) Ergebnisse zur Taxonomie und Okologie einiger Arten und Unterarten der Gattung Phelsuma auf Madagaskar, gesammelt in den Jahren 1972 bis 1981, mit Beschreibung einer neuen Form (Reptilia: Sauria: Gekkonidae). Salamandra, 18, 168 - 190.","Meier, H. & Bohme, W. (1991) Zur Arealkunde von Phelsuma madagascariensis (Gray, 1831) anhand der Museumssammlungen A. Koenig und Senckenberg, mit Bemerkungen zur Variabilitat von P. m. kochi Mertens, 1954. Salamandra, 27, 143 - 151.","Raxworthy, C. J. & Nussbaum, R. A. (1994) A partial systematic revision of the day geckos, Phelsuma Gray, of Madagascar (Reptilia: Squamata: Gekkonidae). Zoological Journal of the Linnean Society, 112, 321 - 335.","Berghof, H. P. & Trautmann, G. (2009) Eine neue Art der Gattung Phelsuma Gray, 1825 (Sauria: Gekkonidae) von der Ostkuste Madagaskars. Sauria, 31, 5 - 14.","Loveridge, A. (1942) Revision of the Afro-Oriental geckos of the genus Phelsuma. Bulletin of the Museum Comparative Zoology, Harvard, 89, 439 - 482.","Boumans, L., Vieites, D. R., Glaw, F. & Vences, M. (2007) Geographical patterns of deep mitochondrial differentiation in widespread Malagasy reptiles. Molecular Phylogenetics and Evolution, 45, 822 - 839.","Kruger, J. (1996 c) Zur Nomenklatur der lineata - Gruppe in der Gattung Phelsuma (Reptilia: Sauria: Gekkonidae). Sauria, 18 (3), 37 - 42.","Mertens, R. (1964) Studien uber die Reptilienfauna Madagaskars V. Funf neue Rassen der Geckonengattung Phelsuma. Senckenbergiana biologica, 45, 99 - 112.","Meier, H. (1983) Neue Ergebnisse uber Phelsuma lineata pusilla Mertens, 1964, Phelsuma bimaculata Kaudern, 1922 und Phelsuma quadriocellata (Peters, 1883), mit Beschreibung von zwei neuen Unterarten (Sauria: Gekkonidae). Salamandra, 19, 108 - 122."]}

Phelsuma barbouri group (contains P. barbouri and P. pronki) These two species were recovered as sister taxa with strong support by both the mitochondrial and nuclear sequences, and the combined analysis (clade Q in Rocha et al. 2009; not shown here). They were previously thought to be closely related (Glaw et al. 1999) but later P. pronki was reclassified within the P. klemmeri group based on being a non-egg-gluer (Van Heygen 2004), contrary to P. barbouri. The molecular data suggests instead that P. klemmeri does not belong to any of the previously identified groups; although it may be more related to the P. l i n e a t a and P. l a t i c a u d a species groups (see below). Phelsuma barbouri and P. pronki individuals exhibit a genetic distance of only 5.1% (uncorrected p-distance, cytochrome b), which is low compared to other interspecific differentiation values within this genus and points to their relatively recent differentiation. Phelsuma barbouri and P. pronki share a rather depressed body and head shape, and a rather similar and distinct colour pattern without clear predominance of green, but differ in the number of midbody scales (higher in P. barbouri), number of ventral scales and egg-gluing behaviour (Tables 1���2). P. barbouri is the most ground-dwelling species within the genus, found on and around the exposed rocky outcrops and boulders of unforested habitat areas of Madagascar's central mountain chain. In contrast, P. pronki is an arboreal species in the mid-altitude rainforests bordering Madagascar's central high plateau. The divergence between these two species is likely a recent one, and may have been driven by ecological specialization along the sharp ecotone between these habitat types. The distribution of P. pronki is very poorly known (Glaw & Vences 2007), and the species is probably rare and/or localized. Additional surveys are needed to understand if these two taxa are geographically isolated or have a contact or hybrid zone. Evidence for a close relationship between P. dubia, P. flavigularis, P. malamakibo, P. b e rg h o f i and P. hielscheri and of these with P. m o d e s t a and P. nigristriata is apparent from the molecular phylogenies (Fig. 2; clade M), corroborating morphological data. This group is strongly supported by the combined phylogeny and mtDNA data and also recovered (though not strongly supported ��� 89 PP, including also P. serraticauda) by the nuclear phylogeny. Within this clade, P. hielscheri, P. berghofi and P. malamakibo (clade O) plus P. flavigularis, form a generally strongly supported clade (Fig. 2), but the position of this latter species respective to others differs between the mitochondrial and combined phylogenies (where P. flavigularis is basal to remaining three species) and the nuclear data (which instead recovers P. flavigularis as sister to P. b e rg h o f i and P. malamakibo, with P. hiescheri basal to these three). The placement of this group of species (P. malamakibo, P. berghofi, P. hielscheri and P. flavigularis) with the P. dubia / P. ravenala clade is strongly supported by mtDNA data and, although weakly, also by the nuclear data. The placement of P. hielscheri (and P. malamakibo) in the P. dubia group is congruent with the morphological data (Table 1���2; see also Van Heygen 2004) but contrasts with recent results from Raxworthy et al. (2007) in which their individual of P. hielscheri was more closely related to P. lineata and P. laticauda. Specimens of P. h i e l s c h e r i have previously been considered as P. dubia (see Glaw et al. 1999), and the differences between the two species only recently noted (R��sler et al. 2001). In our analysis, even when it is not placed in a monophyletic group with P. m a l a m a k i b o and P. berghofi, P. hielscheri is always recovered as included in the P. dubia group, and never close to the P. lineata or P. laticauda species groups as in the analysis of Raxworthy et al. (2007). A reanalysis of the data of these authors as available from Genbank revealed that they had only a very short partial sequence of cytochrome b available for this species (193 bp; EF 434870); which, when aligned with our data, clustered deep inside the Phelsuma lineata group. Since our samples come from a well-identified specimen of P. hielscheri and the various markers we sequenced are congruent in placing it into the P. dubia group, we conclude that the deviant placement of the species in the phylogeny of Raxworthy et al. (2007) is likely erroneous. The mitochondrial genes reveal additional deep differentiation within P. dubia (although not apparent in the nuclear phylogeny, possibly due to incomplete lineage sorting). P. dubia has a patchy distribution along the northwestern coast of Madagascar, possibly with disjunct populations (Glaw & Vences 1994). We find a deep divergence between the individual from the northern isolate (Ambanja) and the southern one (Antsalova) (12.2% uncorrected p-distance in the cytochrome b marker). The included specimen from Moheli, Comoros, is very closely related to specimens from northwestern Madagascar (Ambanja), as are specimens from Zanzibar, Tanzania: see Rocha et al. 2007). Recently, Raxworthy et al. (2007) described P. ravenala from the eastern coast of Madagascar as a new species from the P. dubia group. Although genetic differentiation relative to P. dubia was minimal (~0.3%, 12 S rRNA uncorrected p-distance; 0.4���0.7% cytochrome b distance according to our data), allopatric distribution, morphological differentiation and better fit of distribution models when considering it a different species were invoked to argue for its specific status. Our P. dubia individuals from the Antsalova region in western Madagascar exhibit a much higher degree of differentiation relative to remaining P. dubia (14 % uncorrected p-distance at cytochrome b) and might also be allopatrically distributed. This indicates that the taxonomy in the P. dubia / ravenala complex of species merits further investigation; P. dubia is at the moment paraphyletic with respect to P. r a v e n a l a, and possibly contains yet further undescribed species, with the western Antsalova population being one candidate. Also, the validity of P. r a v e n a l a is in need of confirmation, especially regarding its morphological differentiation (in number of midbody scales, colour and number of preanofemoral pores; see Raxworthy et al. 2007 and Tables 1���2). The second major clade in the P. dubia group is composed of P. m o d e s t a and the Comoran P. nigristriata. The sister group relationship of these two species is supported by nuclear as well as mitochondrial data. The placement of this clade with the P. dubia group is strongly supported by the combined and mitochondrial data sets but unresolved with nuclear data alone. Our results are partly in agreement with the opinion of Nussbaum et al. (2000) who considered the subspecies of Phelsuma modesta as invalid forms representing colour variations. The level of differentiation between two of the subspecies (P. modesta modesta from Tolagnaro and P. m. isakae from near the type locality Isaka) is very low (0.1% uncorrected p-distance in cytochrome b) and the blue head colouration of males, the only character to distinguish P. m. i s a k a e from P. m. modesta is also seen in specimens from the Tolagnaro region, usually assigned to P. m. modesta. On the other hand, P. modesta leiogaster presents a higher level of differentiation (6 %), but the topology recovered differs between mitochondrial and nuclear genes. Species in the Phelsuma dubia group share egg-gluing behaviour (except for P. nigristriata), absence of nostril-rostral contact, a relatively low number of preanofemoral pores in males (30 or less), and keeled dorsal and lateral scales (except for P. ravenala). Most species have a weak expression of green colour, and in some species of the group the ventrals and/or subcaudals are keeled (Tables 1���2)., Published as part of Rocha, Sara, R��sler, Herbert, Gehring, Philip-Sebastian, Glaw, Frank, Posada, David, Harris, James & Vences, Miguel, 2010, Phylogenetic systematics of day geckos, genus Phelsuma, based on molecular and morphological data (Squamata: Gekkonidae), pp. 1-28 in Zootaxa 2429 on pages 9-13, DOI: 10.5281/zenodo.194693, {"references":["Rocha S., Vences M., Glaw F., Posada D., Harris D. J., 2009. Multigene phylogeny of Malagasy day geckos of the genus Phelsuma. Molecular Phylogenetics and Evolution, 52, 530 - 537.","Glaw, F., Vences, M. & Ziegler, T. (1999) Bemerkungen zu Phelsuma dubia (Boettger, 1881): Wiederentdeckung des Holotypus, Verwandtschaftsbeziehungen und Daten zur Fortpflanzung. Salamandra, 35, 267 - 278.","Van Heygen, E. (2004) Remarks on the Phelsuma barbouri- and Phelsuma klemmeri- phenetic groups. Phelsuma, 12, 153 - 154.","Raxworthy, C. J., Ingram, C. M., Rabibisoa, N. & Pearson, R. G. (2007) Applications of ecological niche modelling for species delimitation: A review and empirical evaluation using day geckos (Phelsuma) from Madagascar. Systematic Biology, 56, 907 - 923.","Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Second Edition. Vences and Glaw Verlag, Koln, 480 pp.","Rocha S., Posada D., Carretero M. A., Harris D. J., 2007. Phylogenetic affinities of Comoroan and East African day geckos (genus Phelsuma): multiple natural colonisations, introductions and island radiations. Molecular Phylogenetics and Evolution, 43, 685 - 692.","Nussbaum, R. A., Raxworthy, C. J., Raselimanana, A. P. & Ramanamanjato, J. B. (2000) New species of Day Gecko, Phelsuma Gray (Reptilia: Squamata: Gekkonidae), from the Reserve Naturelle Integrale d'Andohahela, southern Madagascar. Copeia, 2000, 763 - 770."]}

Phelsuma laticauda group (contains P. laticauda, P. pasteuri, P. robertmertensi and P. v-nigra) None of the members of P. laticauda group as defined by Glaw et al. (1999) (P. serraticauda and P. antanosy) is actually closely related to P. laticauda, according to our analysis: P. antanosy is in fact more closely related to P. quadriocellata and P. serraticauda cannot be clearly allocated to any species group (see below). Instead, P. laticauda is recovered as the sister species to a clade containing the Comoran species P. robertmertensi, P. pasteuri and P. v-nigra (Fig. 7; 100 PP in the mtDNA and combined analysis). Nuclear markers per se do not provide enough resolution to recover these groups relationship, however, the monophyly of these three Comoroan species is strongly supported by both mtDNA and combined phylogenies (100 PP; clade L in Fig. 7 a). Phelsuma pasteuri and P. robertmertensi which are both endemic to the oldest island of the Comoros archipelago (Mayotte) form a monophyletic group, strongly corroborated by all gene phylogenies, confirming previous results (Rocha et al. 2007) and supporting the specific status of P. pasteuri. The P. laticauda group species (clade K in Fig. 7 a) are characterized by being non-gluers with smooth ventral and sub-caudal scales, three nasalia and without nostril-rostral contact. They are mostly bright green coloured and have no lateral or dorsal pattern of stripes (Tables 1���2)., Published as part of Rocha, Sara, R��sler, Herbert, Gehring, Philip-Sebastian, Glaw, Frank, Posada, David, Harris, James & Vences, Miguel, 2010, Phylogenetic systematics of day geckos, genus Phelsuma, based on molecular and morphological data (Squamata: Gekkonidae), pp. 1-28 in Zootaxa 2429 on page 23, DOI: 10.5281/zenodo.194693, {"references":["Glaw, F., Vences, M. & Ziegler, T. (1999) Bemerkungen zu Phelsuma dubia (Boettger, 1881): Wiederentdeckung des Holotypus, Verwandtschaftsbeziehungen und Daten zur Fortpflanzung. Salamandra, 35, 267 - 278.","Rocha S., Posada D., Carretero M. A., Harris D. J., 2007. Phylogenetic affinities of Comoroan and East African day geckos (genus Phelsuma): multiple natural colonisations, introductions and island radiations. Molecular Phylogenetics and Evolution, 43, 685 - 692."]}

Phelsuma mutabilis group (contains P. b o r a i [named P. sp. aff. mutabilis in Rocha et al. 2009], P. breviceps, and P. m u t a b i l i s) The three species in this group inhabit southwestern (P. breviceps, P. mutabilis) and western Madagascar (P. mutabilis, P. borai). Close relationships between P. b re v i c e p s and P. m u t a b i l i s had already been hypothesized based on morphometric and scale features (Loveridge 1942); in fact, the two species have been synonymized and resurrected several times (Loveridge 1942; Blanc 1972; Kluge 1991; Raxworthy & Nussbaum 1994). The mitochondrial and combined molecular data recover this group as monophyletic, although the nuclear data alone fail to support this grouping (Fig. 3). Furthermore, the combined analysis recovers a monophyletic P. mutabilis sister to the very divergent P. b o r a i. Genetic distances within this clade are high: average uncorrected p-distance between P. breviceps and P. mutabilis for cytochrome b is of 18.9%, with the distance of P. borai to P. breviceps (22.1%) and to P. mutabilis (23.5%) being even higher. Cytochrome b p-distances within P. m u t a b i l i s reach 8 % (see also Glaw et al. 2009). Species in the P. mutabilis group share, among other character states, the absence of bright green colour, non-gluing egg laying behaviour, a relatively low number of sublabials (5���6), smooth ventrals and subcaudals, and absence of nostril-rostral contact (Tables 1-2). The combined mitochondrial data (Fig. 3 b) indicate possible affinities of the P. mutabilis group with P. andamanense. However, this grouping received less than 50 % bootstrap and very low Bayesian support and warrants further analysis. The lineage containing all species from the Mascarene Islands was previously studied in detail by Austin et al. (2004) with a considerably better sampling including the two extinct species P. edwardnewtonii and P. gigas. Corroborating previous results, we also consistently recovered it as a strongly supported monophyletic clade (clade B in Fig. 4). Clearly recovered by both the mitochondrial and combined data, P. guentheri is basal to the remaining species, and appears to have differentiated from these at an early stage of the Mascarene radiation although the two extinct species, not included in our study, might have been even more basal (Austin et al. 2004). The extant species in the P. cepediana group are quite variable in many morphological and chromatic characters but share egg-gluing behaviour, smooth ventralia and subcaudalia, a contact between rostral and nostril (apparently all except P. rosagularis), and a high number of midbody scales (87 or more) (Tables 1���2). Phelsuma guentheri differs from all extant species in the P. cepediana group (including P. edwardnewtonii), and all other Phelsuma species, by the ability to change the pupil to a vertical slit. This and its associated "crepuscular" activity, as in the closely related (and extinct) P. gigas, are most probably of secondary origin (Austin 2004)., Published as part of Rocha, Sara, R��sler, Herbert, Gehring, Philip-Sebastian, Glaw, Frank, Posada, David, Harris, James & Vences, Miguel, 2010, Phylogenetic systematics of day geckos, genus Phelsuma, based on molecular and morphological data (Squamata: Gekkonidae), pp. 1-28 in Zootaxa 2429 on page 13, DOI: 10.5281/zenodo.194693, {"references":["Rocha S., Vences M., Glaw F., Posada D., Harris D. J., 2009. Multigene phylogeny of Malagasy day geckos of the genus Phelsuma. Molecular Phylogenetics and Evolution, 52, 530 - 537.","Loveridge, A. (1942) Revision of the Afro-Oriental geckos of the genus Phelsuma. Bulletin of the Museum Comparative Zoology, Harvard, 89, 439 - 482.","Blanc, C. P. (1972) Les Reptiles de Madagascar et des Iles Voisines. In: Battisttini, R. & Richard-Vindard, G. (Eds.), Biogeography and Ecology in Madagascar. W Junk, The Hague, pp. 501 - 614.","Kluge, A. R. (1991) Checklist of gekkonid lizards. Smithsonian Herpetological Information Service, 85, 1 - 35.","Raxworthy, C. J. & Nussbaum, R. A. (1994) A partial systematic revision of the day geckos, Phelsuma Gray, of Madagascar (Reptilia: Squamata: Gekkonidae). Zoological Journal of the Linnean Society, 112, 321 - 335.","Austin, J. J., Arnold, E. N. & Jones, C. G. (2004) Reconstructing an island radiation using ancient and recent DNA: the extinct and living day geckos (Phelsuma) of the Mascarene islands. Molecular Phylogenetics and Evolution, 31, 102 - 109."]}