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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"]}

Uroplatus fivehy sp. nov. LSID: urn:lsid:zoobank.org:act: AC24C7D3-1AB5-451F-9DE2-935DE7174FDD Figs 3, 4, 6���8 Remarks. This new species corresponds to the lineage previously named Uroplatus sp. A by Raxworthy et al. (2008), Uroplatus sp. 4 by Ratsoavina et al. (2011), and Uroplatus ebenaui [Ca4] by Ratsoavina et al. (2012, 2013, 2015, 2017, 2019a,b). Holotype. ZSM 1721 /2012 (field number FGZC 3590), adult male, collected on 26 November 2012 by F. Glaw, O. Hawlitschek, T. Rajoafiarison, A. Rakotoarison, F. M. Ratsoavina and A. Razafimanantsoa on the Sorata Massif, Sava Region, northern Madagascar, in the vicinity of the campsite, near 13.6805��S, 49.4451��E, 1417 m a.s.l. Paratypes. A total of 12 specimens: ZSM 1722 /2012 (field number FGZC 3591), adult female, with same collection data as the holotype; UADBA-R 70851 (FGZC 3694), adult male, collected by same collectors as holotype on 29 November 2012 on the Sorata Massif at 13.6868��S, 49.4412��E, 1272 m a.s.l.; UADBA-R 70850 (FGZC 3715), adult male, collected by same collectors as holotype on 30 November 2012 on the Sorata Massif near a creek at 13.6829��S, 49.4403��E, 1325 a.s.l.; ZSM 634 /2009 (ZCMV 11308), adult male, and UADBA-R 70966 (ZCMV 11309), adult female, collected between 22���25 June 2009 by F. Ratsoavina, R. D. Randrianiaina, E. Rajeriarison, T. Rajoafiarison, D. R. Vieites and M. Vences near Angozongahy campsite on the western side of the Makira plateau in the Northern Central East of Madagascar, near 15.437��S, 49.1167��E, 1009 m a.s.l.; UADBA 29428 (APR 05955) and UADBA 29427 (APR 05921), two adult males collected on 22 November 2004 and UADBA 29426 (APR 05859), subadult, collected on 19 November 2004 by A.P. Raselimanana at Anjiabolo site (ca. 16.2800��S, 48.8017��E; respectively at 1100 m and 1000 m a.s.l) in Marotandrano Special Reserve, Sofia Region, Northern Central East Madagascar; UADBA 29430 (APR 05984), UADBA 29431 (APR 05985), two males and UADBA 29429 (APR 05983), adult female, collected on 25 November 2004; and UADBA 29432 (APR 06118), adult male, collected on 29 November 2004 by A.P. Raselimanana at Riamalandy site (16.2850��S, 48.8150��E, between 825���875 m a.s.l.) in Marotandrano Special Reserve. Etymology. The species epithet fivehy is a noun in apposition to the genus name, meaning ���paddle��� in local Malagasy dialect, and referring to the paddle-like shape of its tail. Diagnosis. Uroplatus fivehy sp. nov. is assigned to the Uroplatus ebenaui species group based on its relatively small size, leaf-like, laterally-compressed body shape, short tail, and triangular head with supraocular spines. The following unique combination of characters characterizes the new species: (1) short (TAL/SVL 0.21���0.41) and slender (TAW/SVL 0.06���0.13) tail, (2) oral mucosa without black pigment, and (3) relatively smooth (not spiny) appearance in males. Additionally, it is distinguished by high genetic pairwise-distances> 10% in DNA sequences of the mitochondrial 16S gene from all other nominal species of Uroplatus. By its overall leaf-like appearance, the species can easily be distinguished from all members of the U. fimbriatus species group, from U. lineatus, and from the phenetic U. alluaudi species group. Within the U. ebenaui species group, U. fivehy sp. nov. can be distinguished from U. finaritra and U. malama by smaller body size (SVL 60.8���72.4 mm versus 77.5���95.3 mm); from U. malama, U. phantasticus, U. finiavana and U. finaritra by shorter tail (TAL/SVL 0.21���0.41 versus 0.48���0.76); from U. fangorn, U. ebenaui, U. fetsy, and U. kelirambo by its unpigmented oral mucosa (pale pinkish all over versus black or deep red pigment present); from U. kelirambo by more expanded tail edges (versus strongly spear-shaped; TAW/SVL 0.06���0.13 versus 0.04���0.05 in U. kelirambo). U. fivehy sp. nov. is morphologically most similar to U. fiera and to U. fotsivava, which however concordantly differ in mitochondrial and nuclear DNA sequences and are not its immediate relatives based on molecular phylogenetic analysis; it differs from U. fotsivava by a generally longer and wider tail, although the values do overlap (TAL/SVL 0.21���0.41 versus 0.15���0.32; TAW/SVL 0.06���0.13 versus 0.05���0.10), and from U. fiera possibly in a slightly larger ratio TAL/TAW (0.22���0.37 versus 0.19���0.21). Description of the holotype. Adult male in good condition with an intact tail and everted hemipenes. SVL 63.1 mm, tail length 25.9 mm, maximum tail width 5.7 mm, for further measurements see Table 1. Head triangular in dorsal view; canthus rostralis recognizable and almost straight; snout sloping strongly and continuously downward anteriorly; snout weakly depressed, short; eyes large (eye diameter 4.6 mm), not bulging above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small but distinct (horizontal diameter 0.6 mm), its opening facing posterolaterally, but also posteroventrally (ear opening clearly visible in lateral view but not in dorsal view); nostrils laterally oriented; body somewhat laterally compressed, without lateral dermal fringes; limbs without fringes and practically without spines on forelimbs and hindlimbs, except for a small flap on the knee; forelimb reaches beyond tip of snout when adpressed forward and does not reach the groin when adpressed backwards along body (forelimb length/axilla���groin distance 27.7/ 29.7 mm = 93%), hindlimb reaches beyond axilla when adpressed forward along body (hindlimb length/axilla���groin distance 37.4/ 29.7 mm = 126%); original tail length 41% of snout���vent length, membranous borders of the denticulated tail rather asymmetrical, broadening on either side and then narrowing again toward the tip), completely absent from the slightly spatulate tail-tip. Nares separated from each other by eight small granular scales, from the first supralabial by one scale, and from the rostral scale by two scales; the first supralabial taller than the others; rostral entire, much wider than tall; mental scale small, not differentiated from infralabial scales; 21/20 (right/left) supralabials and 19/21 infralabials (as defined in methods above); scales at the periphery of the chin (toward the infralabial scales) slightly larger than central chin scales; the dorsal scales of head, neck, body, limbs and tail small, granular, juxtaposed and largely of homogeneous size interspersed with very few larger and partly raised tubercles; the numerous curved transverse lines on the head and dorsal side of the body are not marked by significantly enlarged scales. Very few spines on head and hindlimbs, a dermal flap on each knee, bearing no 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. Axillary pits not recognizable. Scales of the ventral abdomen distinctly larger than dorsal body scales and arranged more homogeneously. 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 of the holotype. After eight years of preservation in 70% ethanol the color pattern remains similar to that of the living animal (Figs 6H, 8) but its vividness and intensity have slightly faded. All dorsal surfaces are mottled beige and blackish in base color. The tail has a beige pattern on its dorsal pygal section, bordered by blackish color. The dorsum has rather distinctly defined posteroventrally oriented lighter and darker beige chevrons. The dorsal surfaces of the head are dark brown with a series of six distinct transverse narrow beige lines, running either straight or converging anteriorly or posteriorly. No dark band between the eyes, but a dark V-shaped area on the nape is well distinguishable. Additional dark flecks are present on the mid-dorsum, and several smaller asymmetrical spots on other parts of the body, including legs and nape. One light cream marking is present posteriorly below each eye, descending to the supralabials. The supralabials and the infralabials are generally dark gray in color. The venter, including the ventral limbs except for the lower hindlimbs, is slightly lighter beige than the dorsum, and less mottled. The pectoral region bears two elongated, poorly delimited whitish spots. The postpygal section of the tail is of similar color dorsally and ventrally. The oral mucosa is unpigmented and whitish in preservative. The iris of the eye is gray, and the vertical pupil is blackish. Variation. As described for U. fangorn above, there is a substantial color variation (Figs 6���7), and sexual dimorphism in the edges of the tail. The color of the eye varies from beige/gray to dark brown and reddish-brown. Dorsal color can be uniform beige to reddish-brown with or without white lichen-like markings. Two sharply delimited vertebral black patches on the anterior and posterior part of the dorsum are present in several individuals. Relative tail length and tail width are characterized by substantial variability among individuals (Fig. 4). Here, we have included in the paratype series and in the morphological comparisons a series of individuals from Marotandrano Special Reserve that we attribute to this species. However, it must be stressed that the species identity has been confirmed by DNA sequences only for one of these (UADBA 29428 = APR5955). In the unlikely case that some of these individuals will turn out to belong to another species of Uroplatus, the morphometric variation of U. fivehy may be narrower than reported here. ......continued on the next page Distribution, natural history and conservation status. According to the material collected and examined by us, U. fivehy is known from (1) the type locality on the Sorata Massif, (2) the western slope of the Makira Reserve, and (3) Marotandrano Special Reserve. In addition, mitochondrial DNA sequences published by Ratsoavina et al. (2012) suggest its occurrence in (4) the Anjanaharibe-Sud Massif, and mitochondrial DNA sequences from Raxworthy et al. (2008) furthermore suggest the occurrence of the species in (5) Marojejy, (6) Ankitsika and (7) Betaolana. The confirmed elevational range from our own samples is between 1009 and 1417 m a.s.l., and the record from Ankitsika would extend this range downwards to 830 m a.s.l., characterizing the species as rather wide-ranging from the Northern Central East to the North East of Madagascar from mid-elevations to highlands (see also Ratsoavina et al. 2013). All specimens were found in intact rainforest, at night, perched on branches of trees. At Marotandrano, an adult male was found hidden in a Pandanus leaf axil during the day in closed canopy humid forest, and two adult individuals (male and female) were found at night close to each other in the same tree on 25 November 2004, suggesting it was probably the reproduction period for this species. Individuals were often observed active on small branches, leaves, and lianas at 0.5���3 m above the forest floor. All individuals were collected on slope or ridge forest. In Marotandrano, the species appeared to be quite common in forests associated with vine-like bamboo and shrub vegetation. A minimum convex polygon of the known distribution of this species covers an area of ca 15,100 km 2, qualifying the species for a status of Vulnerable under the IUCN Red List criterion B (IUCN 2012). This species is known from several protected areas (Marotandrano Special Reserve, Makira Reserve, COMATSA Nord, Marojejy National Park, Anjanaharibe-Sud National Park) but cumulatively just seven locations (B subcriterion a). These locations, and especially the unprotected areas between them, are experiencing substantial ongoing declines in the extent and quality of habitat (B subcriterion b(iii)). Therefore, we propose to list this species as Vulnerable under IUCN criterion B1ab(iii)., Published as part of Ratsoavina, Fanomezana M., Glaw, Frank, Raselimanana, Achille P., Rakotoarison, Andolalao, Vieites, David R., Hawlitschek, Oliver, Vences, Miguel & Scherz, Mark D., 2020, Towards completion of the species inventory of small-sized leaf-tailed geckos: two new species of Uroplatus from northern Madagascar, pp. 251-271 in Zootaxa 4895 (2) on pages 261-268, DOI: 10.11646/zootaxa.4895.2.5, http://zenodo.org/record/4322511, {"references":["Raxworthy, C. J., Pearson, R. G., Zimkus, B. M., Reddy, S., Deo, A. J., Nussbaum, R. A. & Ingram, C. M. (2008) 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 (1), 39 - 57. https: // doi. org / 10.11646 / zootaxa. 3022.1.3","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 distribu- tion 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 (1), 143 - 160. https: // doi. org / 10.11646 / zootaxa. 4006.1.7","Ratsoavina, F. M., Gehring, P. S., Scherz, M. D., Vieites, D. R., Glaw, F. & Vences, M. (2017) Two new species of leaf-tailed geckos (Uroplatus) from the Tsaratanana mountain massif in northern Madagascar. Zootaxa, 4347 (3), 446 - 464. https: // doi. org / 10.11646 / zootaxa. 4347.3.2","Ratsoavina, F. M., Raselimanana, A. P., Scherz, M. D., Rakotoarison, A., Razafindraibe, J. H., Glaw, F. & Vences, M. (2019 a) Finaritra! A new leaf-tailed gecko (Uroplatus) species from Marojejy National Park in north-eastern Madagascar. Zootaxa, 4545 (4), 563 - 577. https: // doi. org / 10.11646 / zootaxa. 4545.4.7","IUCN. (2012) IUCN Red List Categories and Criteria: Version 3.1. IUCN, Gland and Cambridge."]}

Description of Calumma ratnasariae sp. nov. ZOOBANK urn:lsid:zoobank.org:act: D822D6EA-7CB8-425B-9AD1-473D1DCFB2C8 Remark: This new species refers to clade I of Fig. 2 and GEHRING et al. (2012). Holotype: ZSM 35 /2016 (MSZC 0066), adult male, collected in the Ampotsidy mountains (14.4146°S, 48.7115°E, 1400 m a.s.l.), Sofia Region, northern Madagascar, on 22 December 2015 by M.D. Scherz, J. Borrell, L. Ball, T. Starnes, T. S.E. Razafimandimby, D.H. Nomenjanahary, J. Rabearivony. Paratypes: ZSM 1724 /2010 (ZCMV 12483), adult male and ZSM 2884 /2010 (ZCMV 12273), adult female, both collected in Analabe Forest, near Antambato village (Ambodimanga mountain, 14.5048°S, 48.8760°E, 1361 m a.s.l.) on 24 June 2010, ZSM 517 / 2014 (DRV 6283), adult male collected in Andrevorevo (14.3464°S, 49.1028°E, 1717 m a.s.l.), Sofia Region, northern Madagascar on 21 June 2010, all three by M. Vences, D. R. Vieites, R. D. Randrianiaina, F.M. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, T. Rajoafiarison; ZSM 36 /2016 (MSZC 0140), adult female, collected on the Ampotsidy mountains (14.4099°S, 48.7155°E, 1647 m a.s.l.) on 3 January 2016, and ZSM 37 /2016 (MSZC 0169), adult female, collected on the Ampotsidy mountains (14.4193°S, 48.7193°E, 1337 m a.s.l.), on 8 January 2016, both by M.D. Scherz, J. Borrell, L. Ball, T. S.E. Razafimandimby, D.H. Nomenjanahary, J. Rabearivony. Diagnosis (based on the type series; osteology based on micro-CT scans of ZSM 35/2016 and ZSM 517/2014, both males): Calumma ratnasariae sp. nov. is characterised by (1) a large size (male SVL 43.9– 52.0 mm, female SVL 48.7 – 51.5 mm; male TL 97.1 – 110.7, female TL 95.3 – 101.0); (2) a short (1.8 – 2.3 mm in males, 2.1 – 2.2 mm in females) and distally rounded rostral appendage, (3) rostral scale not integrated into the rostral appendage, (4 – 7) rostral, lateral, temporal (one tubercle on either side), and cranial crests present, (8) parietal crest distinct and running the length of the parietal bone, (9) a distinctly raised casque in males with a height of 1.3– 1.5 mm, (10) a dorsal crest of 7 – 12 cones in males, generally present in females (6 – 8 cones), (11) 10 – 13 supralabial scales with a straight upper margin, (12) absence of axillary pits, (13) diameter of the largest scale in the temporal region of the head 1.2 – 1.6 mm, (14) a frontoparietal fenestra in the skull, (15) parietal and squamosal in contact (n = 2), (16) parietal bone width at midpoint 17.8 – 18.5% of skull length, (17) a generally yellowish body colouration in males, greyish body colouration in females, (18) rostral appendage not accentuated from the body colouration, (19) a blue and yellow cheek colouration, (20) yellow in males and beige in females, and (21) brown stripe crossing the eye. Calumma ratnasariae sp. nov. is unique among the C. nasutum complex in having an elevated bony knob on the anterodorsal edge of the maxillary facial process (this character is similar to that seen in C. uetzi). Calumma ratnasariae sp. nov. can be distinguished from all species of the C. boettgeri complex (see above) by the absence of occipital lobes; from C. gallus by different length, shape and colour of its rostral appendage (see above); from all other species of the C. nasutum group without occipital lobes except C. fallax by the presence of a frontoparietal fenestra. It is also quite unusual in having an overall yellowish body colouration in males. In addition, it can be distinguished from C. vatosoa easily by the presence of a rostral appendage (vs absence); from C. vohibola by longer rostral appendage (RRS 3.8 – 4.8% vs 0.2 – 3.1%), parietal crest present (vs absent), fewer supralabials (10 – 13 vs 13 – 16) with a straight upper margin (vs serrated), larger temporal scale (1.2 – 1.6 mm vs 1.0 mm), broader parietal bone with a continuous parietal crest (vs smooth parietal); from C. nasutum as here redefined by a larger maximum total length in males (110.7 mm vs 89.0 – 100.8 mm), a distinct parietal crest (vs absent or indistinct), dorsal crest generally present in both sexes (vs generally absent and absent in females); from C. radamanus by larger total length (95.3– 110.7 mm vs 77.0 – 93.5 mm), tail length in males longer than SVL (vs shorter), rostral scale not integrated into the rostral appendage (vs generally integrated), parietal crest present (vs absent), supralabials with a straight upper margin (vs serrated), larger temporal scale (1.2 – 1.6 mm vs 0.6 – 0.9 mm), and parietal and squamosal in contact (vs widely separated); from C. emelinae sp. nov. by parietal crest distinct (vs general absence), higher casque in males (1.3 – 1.5 mm vs 0.5 – 1.1 mm), dorsal crest consisting of cones (vs spines) in males; larger temporal scale (1.2 – 1.6 mm vs 0.6 – 1.0 mm), and broad postparietal process (vs strongly tapering); from C. tjiasmantoi sp. nov. by larger body length of females (SVL 48.7 – 51.5 mm vs 43.9 – 46.1 mm), dorsal crest generally present in females (vs absent), fewer supralabials (11 – 12 vs 15 – 17), and larger diameter of temporal scale (1.2 – 1.6 mm vs 0.6 – 0.8 mm); and from C. fallax by generally shorter relative rostral appendage length in females (RRS 4.1 – 4.5% vs 4.2 – 7.6%), cranial crest present (vs generally absent), parietal crest longer and more distinct, dorsal crest generally present in females (vs generally absent), and a wider mid-parietal width (17.8 – 18.5% of skull length vs 6.7 – 15.7%). Description of the holotype (Figs. 4F, 16 A,B): Adult male, with mouth closed, in good state of preservation, with everted hemipenes, one completely (on the left) and one incompletely (on the right); SVL 48.2 mm, tail length 54.2 mm, for other measurements, see suppl. Table 1; distinct rostral ridges that give the snout a right angle; laterally compressed dermal rostral appendage of oval tubercle scales that projects slightly downwards over a length of 2.3 mm with a diameter of 1.9 mm; 13 infralabial and 12 supralabial scales, both relatively large; supralabials with a smooth dorsal margin; distinct lateral crest running horizontally; temporal crest consisting of one tubercle per side; short cranial crest; distinct and long parietal crest ending in the tip of the casque with a height of 1.5 mm; no occipital lobes; dorsal crest present, consisting of 7 broad cones; no traces of gular or ventral crest. Body laterally compressed with fine homogeneous scalation and distinctly larger scales on extremities and head region, largest scale in temporal region with diameter of 1.3 mm and in cheek region of 1.1 mm; no axillary or inguinal pits. Skull osteology of the holotype (Fig. 14D): Skull length 11.9 mm; snout-casque length 14.4 mm; narrow paired nasals anterior slightly connected; anterior tip of frontal exceeding the middle of the prefrontal fontanelle, which is fused with the naris; prominent and broad prefrontal; frontal and parietal smooth without any tubercles; frontal with a width of 2.6 mm (21.8% of skull length) at border to prefrontal extending to 4.4 mm (37.0%) at border to postorbitofrontal; large frontoparietal fenestra with a width of 2.5 mm (21.0%); broad parietal with distinct parietal crest tapering slightly from a width of 4.0 mm (33.6%) at the border to postorbitofrontal to a width at midpoint of 2.2 mm (18.6%); the posterodorsally broadened end is in weak contact with the squamosals; squamosals thin with a few tubercles. For further measurements, see Table 2. Hemipenial morphology, based on ZSM 1724/2010 (Fig. 5E), ZSM 517/2014, and ZSM 35/2016: small calyces (hemipenial character A); two pairs of finely denticulated rotulae of different size, on sulcal side large with about 12 – 15 tips, on asulcal side small with about 5 – 7 tips (B); papillary field of small, unpaired papillae (C); pair of short cornucula gemina (D), only visible when hemipenis fully everted. Variation: For variation in measurements, see Table 1. Sexual dimorphism: Males and females do not seem to differ in body size. Tail length is longer in males than in females (RTaSV 112 – 121% vs 94 – 98%). Relative rostral appendage length does not differ. Dorsal crest does not differ between males and females. Colouration in life (Fig. 16): Strong sexual dichromatism with males of yellow body colouration and turquoise stripes and females generally uniformly beige. Males with turquoise annulated tail and extremities and two brown blotches on the body side that is crossed by a diffuse white lateral stripe; no pattern in females; throat and ventral region slightly brighter than the flank; indistinct rostral appendage not accentuated from the body/head, can be spotted with blue or yellow dots; in males cheek region and eyelids with turquoise dots, in females blue dots can occur on rostral ridges and eyelids when stressed; a diffuse brown stripe crosses the eye in both sexes. Etymology: The specific epithet is named after Yulia Ratnasari, in recognition of her support for taxonomic research and nature conservation projects in Madagascar through the BIOPAT initiative (http://biopat.de/en/). Distribution (Fig. 9): Calumma ratnasariae sp. nov. is only known from the Bealanana District of northern Madagascar. It is distributed from Analabe Forest in the south to Andrevorevo, about 20 km further north (for coordinates, see above), from an elevation of 1337– 1717 m a.s.l., Published as part of Prötzel, David, Scherz, Mark D., Ratsoavina, Fanomezana M., Vences, Miguel & Glaw, Frank, 2020, Untangling the trees: Revision of the Calumma nasutum complex (Squamata: Chamaeleonidae), pp. 23-59 in Vertebrate Zoology 70 (1) on pages 52-54, DOI: 10.26049/vz70-1-2020-3, http://zenodo.org/record/4394821, {"references":["GEHRING, P. - S., TOLLEY, K. A., ECKHARDT, F. S., TOWNSEND, T. M., ZIEGLER, T., RATSOAVINA, F., GLAW, F. & VENCES, M. (2012). Hiding deep in the trees: Discovery of divergent mitochondrial lineages in Malagasy chameleons of the Calumma nasutum group. Ecology and Evolution, 2, 1468 - 1479."]}

Key for male specimens of the Calumma nasutum group The following key is based on external morphology only and some characters of species lacking occipital lobes (highlighted by underlined numbers, except for Calumma roaloko) are illustrated in Fig. 17 with numbers referring to the sections of the key. For diagnostic characters of colouration, see Table 4, and for further morphological characters, see Table 1. Females of this complex are more homogeneous in morphology and thus generally show fewer diagnostic characters (e.g. shorter and indistinct rostral appendages, usually lacking a dorsal crest) and were not considered in the identification key, except for the identification of C. juliae where only females are known so far. The key for the Calumma boettgeri complex of PRÖTZEL et al. (2018b) was modified and supplemented with C. roaloko. Illustrations of characters for underlined numbers in the key are given in Fig. 17. 1a Occipital lobes absent (Calumma nasutum c omplex)...................................................................... 2 1b Occipital lobes present (Calumma boettgeri complex)..................................................................... 9 2a Rostral appendage long (5.0 – 11.0 mm) and spearlike, dorsal crest absent, homogeneous scalation on body and head.............. Calumma gallus complex 2b Rostral appendage shorter and rounded, dorsal crest can be present or absent, more or less heterogeneous scalation on body and head.................................. 3 3a Rostral scale integrated in rostral appendage, rostral appendage oriented downwards, parietal crest absent, absence of cornucula gemina on hemipenis.................................. Calumma radamanus complex 3b Rostral scale not integrated in rostral appendage, rostral appendage straight or oriented upwards, parietal crest present (can be absent in C. nasutum and C. vohibola), cornucula gemina present............... 4 4a Rostral appendage short (≤ 1.4 mm), small total length (≤ 89.0 mm), dorsal crest present and consisting of spines....................... Calumma vohibola 4b Rostral appendage completely lacking, large total length (> 120.0 mm), dorsal crest absent..................................................................... Calumma vatosoa 4c Rostral appendage longer (> 1.4 mm), total length medium sized (89.0 – 110.0 mm), dorsal crest present or absent.......................................................... 5 5a Low casque (≤ 0.6 mm) and dorsal crest consisting of spines..................... Calumma emelinae sp. nov. 5b Casque higher (> 0.6 mm) and dorsal crest consisting of small cones, if present (spines can also occur in C. radamanus and C. vohibola)......................... 6 6a Casque height 7 6b Casque height> 1.5 mm........................................ 8 7a Relatively homogeneous scalation on the head with diameter of largest temporal scale ≤ 0.8 mm, high number of supra- and infralabial scales (n ≥ 15), frontoparietal fenestra absent (you can feel the closed skull roof by gently pushing the top of the head of a preserved specimen)................................................................ Calumma tjiasmantoi sp. nov. 7b Heterogeneous scalation on the head with diameter of largest temporal scale ≥ 1.3 mm, lower number of supra- and infralabial scales (10 – 14), frontoparietal fenestra present (the top of the head of a preserved specimen feels soft) … Calumma ratnasariae sp. nov. 8a Parietal crest present, frontoparietal fenestra present, casque rather rounded (convex), axillary pits absent, upper margin of supralabial scales usually serrated........................................ Calumma fallax 8b Parietal crest usually absent, frontoparietal fenestra absent, casque rather pointed (concave), axillary pits can occur, upper margin of supralabial scales usually in line.......................... Calumma nasutum 9a Small tubercle scales (diameter ≤ 0.5 mm) on extremities, small body size (10 9b Large tubercle scales (diameter 0.5– 0.9 mm) on extremities, large body size (generally> 110 mm total length,> 45 mm SVL)....................................... 12 10a Tubercle scales on extremities isolated from each other, upper margin of supralabial scales serrated ..................................................... Calumma boettgeri 10b Tubercle scales on extremities bordering each other, upper margin of supralabial scales in line........... 11 11a Temporal crest present, dorsal crest consisting of 13 – 14 cones or spines, medium sized rostral appendage (≤ 3.9 mm) ...................... Calumma uetzi 11b Temporal crest absent, dorsal crest consisting of 1 – 2 cones, longer rostral appendage (≥ 3.9 mm)....................................................... Calumma roaloko 12a Occipital lobes not or only slightly separated (notch 0 – 0.8 mm), no frontoparietal fenestra (closed skull roof)..................................................................... 13 12b Occipital lobes clearly separated (notch> 0.5 mm), frontoparietal fenestra present (can be felt through the skin in alcohol-preserved specimens).......... 14 13a Temporal crest of one or two tubercles present, parietal crest present, dorsal crest in females absent or consisting of up to 6 dorsal cones.... Calumma linotum 13b Temporal crest absent, parietal crest absent, dorsal crest present in females and consisting of 9 – 14 cones............................................ Calumma juliae 14a Large frontoparietal fenestra, occipital lobes widely separated, dorsal and caudal crest absent, rostral appendage rounded......................... Calumma guibei 14b Small frontoparietal fenestra, occipital lobes slightly connected, dorsal crest of 7 – 15 distinct cones, no caudal crest, rostral appendage rounded................................................................... Calumma gehringi 14c Medium-sized frontoparietal fenestra, occipital lobes completely separated, dorsal crest of> 20 small conical scales, caudal crest present, rostral appendage pointed> 5.5 mm........................ Calumma lefona, Published as part of Prötzel, David, Scherz, Mark D., Ratsoavina, Fanomezana M., Vences, Miguel & Glaw, Frank, 2020, Untangling the trees: Revision of the Calumma nasutum complex (Squamata: Chamaeleonidae), pp. 23-59 in Vertebrate Zoology 70 (1) on pages 54-55, DOI: 10.26049/vz70-1-2020-3, http://zenodo.org/record/4394821, {"references":["PROTZEL, D., VENCES, M., HAWLITSCHEK, O., SCHERZ, M. D., RATSOA- VINA, F. M. & GLAW, F. (2018 b). Endangered beauties: Micro- CT cranial osteology, molecular genetics and external morphology reveal three new species of chameleons in the Calumma boettgeri complex (Squamata: Chamaeleonidae). Zoological Journal of the Linnean Society, 184, 471 - 498."]}

Description of Calumma tjiasmantoi sp. nov. ZOOBANK urn:lsid:zoobank.org:act: 65471E38-67B4-4BC0-9CC9-402B8214A7A8 Remark: This new species refers to clade J of Fig. 2 and GEHRING et al. (2012). Due to uncertainties about the collection data of the only male specimen we designate a well-documented female as the holotype. Holotype: ZSM 735 /2003 (FG / MV 2002-497), adult female collected in Ranomafana National Park (21.2639°S, 47.4194°E, 983 m a.s.l.), Vatovavy-Fitovinany Region, eastern Madagascar, on 23 January 2003 by F. Glaw, M. Puente, L. Raharivololoniaina, M. Thomas, D. R. Vieites. Paratypes: ZSM 312 /2006 (ZCMV 2896), adult male, and UADBA uncatalogued (ZCMV 2895), female, both collected in Ranomafana, probably Ambatolahy (21.2439°S, 47.4262°E, 919 m a.s.l.) on 21 February 2006 by M. Vences; ZSM 723 /2003 (FG / MV 2002- 0396) and ZSM 736 /2003 (FG / MV 2002-0498), both adult females, same collection data as holotype; ZSM 380 /2016 (ZCMV 14835), adult female, collected in Sampanandrano (24.1399°S, 47.0742°E, 539 m a.s.l.), Atsimo-Antsinanana Region, southeastern Madagascar, on 16 December 2016 by A. Rakotoarison, E. Rajeriarison, J.W. Ranaivosolo. Diagnosis (based on the type series, osteology based on micro-CT scan of ZSM 735/2003, female): Calumma tjiasmantoi sp. nov. is characterised by (1) a small size (male SVL 43.9 – 46.8 mm, female TL 84.1 – 94.8 mm); (2) a medium sized (1.1 – 2.1 mm) and distally rounded rostral appendage, (3) rostral scale not integrated into the rostral appendage, (4 – 8) rostral, lateral, temporal, cranial, and parietal crests present, (9) casque medium sized in males (1.3 mm), (10) a dorsal crest of 7 – 9 spines can be present in males (based also on photographs), absent in females, (11) 15 – 17 supralabial scales with a mostly straight upper margin, (12) general absence of axillary pits, (13) diameter of the largest scale in the temporal region of the head 0.6 – 0.8 mm, (14) frontoparietal fenestra absent, (15) parietal and squamosal in contact, (16) parietal bone width at midpoint 16.1% of skull length (n = 1) with a characteristic broad tip to the postparietal process, (17) bright green or yellowish body colouration in males, females generally browner and less conspicuous, (18) rostral appendage colour generally inconspicuous, (19) cheek colouration greenish to turquoise, (20) five characteristic dorsoventral stripes of blue or brown colour, and (21) a diffuse brown stripe crossing the eye. Calumma tjiasmantoi sp. nov. can easily be distinguished from all species of the C. boettgeri complex (see above) by the absence of occipital lobes; from C. gallus by different length, shape and colour of its rostral appendage (see above); from C. vatosoa by presence of a rostral appendage (vs absence); from C. vohibola by longer rostral appendage in females (RRS 2.4 – 4.6% vs 0.2 – 0.7%), parietal crest present (vs absent), smaller temporal scale (0.6 – 0.8 mm vs 1.0 mm); from C. nasutum as redefined herein by the higher number of supralabials (15 – 17 vs 12 – 15), a shorter rostral appendage (4.3% of SVL vs 4.5 – 5.3% of SVL), a shorter parietal (36.3% of skull length vs 41.0 – 44.3%), broad postparietal process (vs narrow); from C. radamanus by a relatively longer tail in females (RTaSV 92 – 95% vs 79 – 89%), rostral scale not integrated in rostral appendage (vs generally integrated), parietal crest present (vs absent), more supralabials (15 – 17 vs 11 – 15) with a generally straight upper margin (vs serrated), parietal and squamosal in contact (vs widely separated); from C. emelinae sp. nov. by presence of parietal crest (vs usually absent), absence of lateral white stripe, and broader postparietal process; for diagnosis against C. fallax, see below. For diagnosis against the other species described herein, see its description below. Description of the holotype (Fig. 4D): Adult female, with mouth closed, in good state of preservation except for a ventrally sliced body, with four eggs still fixed in the oviduct; SVL 45.5 mm, tail length 42.5 mm, for other measurements, see suppl. Table 1; bulging rostral ridges; laterally flattened and distally rounded rostral appendage of tubercle scales that projects straight forward over a length of 1.7 mm with a diameter of 1.4 mm not including the rostral scale; 15 infralabial and 15 supralabial scales, all rather small; most of the supralabials with a straight upper margin, only anterior scales serrated; distinct lateral crest running horizontally; temporal crest consisting of two tubercles; short cranial crest; short parietal crest; no occipital lobes; medium sized casque of 1.1 mm height; no dorsal, gular or ventral crest. Body laterally compressed with fine homogeneous scalation and only slightly larger scales on extremities and head region, largest scale in temporal region with diameter of 0.6 mm and in cheek region of 0.6 mm; no axillary or inguinal pits. Skull osteology of the holotype (Fig. 11B): Skull length 12.4 mm; snout-casque length 14.4 mm; narrow paired nasals anteriorly still in contact; anterior tip of frontal not exceeding the middle of the prefrontal fontanelle, which is fused with the naris; prominent and broad prefrontal with laterally raised tubercles; frontal and parietal smooth with only a few tubercles; frontal with a width of 2.4 mm (19.4% of skull length) at border to prefrontal extending to 4.3 mm (34.7%) at border to postorbitofrontal; no frontoparietal fenestra; broad parietal with distinct parietal crest tapering slightly from a width of 4.3 mm (34.7%) at the border to postorbitofrontal to a width at midpoint of 2.0 mm (16.1%); the posterodorsally broadened end is in strong contact with the squamosals; squamosals broad with several tubercles. For further measurements, see Table 2. Hemipenial morphology, based on ZSM 312/2006 (no micro-CT scan available): medium sized calyces (hemipenial character A); two pairs of small rotulae on apex of about the same size (B), finely denticulated with about 7 – 9 tips each; papillary field of small, unpaired papillae (C); pair of medium sized cornucula gemina (D), only visible when hemipenis fully everted. Variation: For variation in measurements, see Table 1. For variation in colouration in life, see Fig. 13. The female ZSM 134/2005 (FGZC 2508) from Andohahela at high elevation (1548 m a.s.l.) belongs genetically to clade J but shows substantial morphological differences so that we do not designate it as a paratype. Aside from a larger total length (97.0 mm), a longer rostral appendage (2.6 mm), and a low number of supralabials (12) it also has a unique skull morphology with a FF. This correlates well with a previous study where only high elevation species tend to have a FF (PRÖTZEL et al., 2018b) but does not fit in the general pattern of C. tjiasmantoi sp. nov. Further studies are necessary to clarify this apparent contradiction in our dataset and potential taxonomic conclusions. Sexual dimorphism: Body size (SVL and TL) is slightly larger in males than females, tail length is longer in males than in females (RTaSV 103% vs 92 – 95%), and a dorsal crest is only present in males. Colouration in life: Strong sexual dichromatism with males of bright green or yellowish body colouration and turquoise extremities and females generally browner and less conspicuous. In both sexes five diffuse dorsoventral stripes on the body – in males of blue colour with light spots, in females of brown colour; no lateral stripe across the body; tail annulated, continuing the blotches from the body; throat and ventral region can be beige or of same colour as the body; indistinct rostral appendage not accentuated from the head; cheek region turquoise in males; a diffuse brown stripe may cross the eye. Etymology: The specific epithet is a patronym honouring Wewin Tjiasmanto in recognition of his support for taxonomic research and nature conservation projects in Madagascar through the BIOPAT initiative (http://biopat. de/en). Distribution (Fig. 9): Calumma tjiasmantoi sp. nov. is known from eastern Madagascar from Andohahela in the south to Ranomafana NP about 400 km further north (for coordinates, see above), from an elevation of 267 to 983 m a.s.l. (see Fig. 9)., Published as part of Prötzel, David, Scherz, Mark D., Ratsoavina, Fanomezana M., Vences, Miguel & Glaw, Frank, 2020, Untangling the trees: Revision of the Calumma nasutum complex (Squamata: Chamaeleonidae), pp. 23-59 in Vertebrate Zoology 70 (1) on pages 45-48, DOI: 10.26049/vz70-1-2020-3, http://zenodo.org/record/4394821, {"references":["GEHRING, P. - S., TOLLEY, K. A., ECKHARDT, F. S., TOWNSEND, T. M., ZIEGLER, T., RATSOAVINA, F., GLAW, F. & VENCES, M. (2012). Hiding deep in the trees: Discovery of divergent mitochondrial lineages in Malagasy chameleons of the Calumma nasutum group. Ecology and Evolution, 2, 1468 - 1479.","PROTZEL, D., VENCES, M., HAWLITSCHEK, O., SCHERZ, M. D., RATSOA- VINA, F. M. & GLAW, F. (2018 b). Endangered beauties: Micro- CT cranial osteology, molecular genetics and external morphology reveal three new species of chameleons in the Calumma boettgeri complex (Squamata: Chamaeleonidae). Zoological Journal of the Linnean Society, 184, 471 - 498."]}

Description of Calumma emelinae sp. nov. ZOOBANK urn:lsid:zoobank.org:act: EE10A0D2-832F-4CBE-81E0-10E6CADB4A76 Remark: This new species refers to clade B of Fig. 2 and GEHRING et al. (2012). Holotype: ZSM 618 /2009 (ZCMV 11292), adult male with completely everted hemipenes, right hemipenis cut off for micro-CT scanning, collected in the Makira plateau, Angozongahy or Ampofoko (about 15.44°S, 49.12°E, 1000 m a.s.l.), Analanjirofo Region, northeastern Madagascar, on 22– 25 June 2009 by M. Vences, D. R. Vieites, F.M. Ratsoavina, R. D. Randrianiaina, E. Rajeriarison, T. Rajofiarison, J. Patton. Paratypes: ZSM 660 /2014 (DRV 5899), ZSM 663 /2014 (DRV 5898), both adult males, collected in Angozongahy, western side of Makira plateau camp 1 (15.4370°S, 49.1186°E, 1009 m a.s.l.), Analanjirofo Region, northeastern Madagascar, on 26 June 2009 by M. Vences, D. R. Vieites, F.M. Ratsoavina, R. D. Randrianiaina, E. Rajeriarison, T. Rajofiarison, J. Patton; ZSM 553 /2001 (MV 2001- 239), adult female, collected in Andasibe (about 18.93°S, 48.42°E, 900 m a.s.l.), Alaotra-Mangoro Region, eastern Madagascar, on 16 – 18 February 2001 by M. Vences, D. R. Vieites; ZSM 135 /2005 (FGZC 2692) and ZSM 136 /2005 (FGZC 2693), both adult females and collected in Vohidrazana (about 18.95°S, 48.50°E, 700– 800 m a.s.l.), Alaotra-Mangoro Region, eastern Madagascar, on 09 February 2005 by F. Glaw, R.D. Randrianiaina, R. Dolch; ZSM 661 /2014 (DRV 5677) and ZSM 662 /2014 (DRV 5708), both adult females, collected in Mahasoa campsite near Ambodisakoa village (NE Vohimena, NE Lake Alaotra, 17.2977°S, 48.7019°E, 1032 m a.s.l.), Alaotra-Mangoro Region, eastern Madagascar, on 13– 15 February 2008 by D. R. Vieites, J. Patton, P. Bora, M. Vences; ZSM 148 /2016 (FGZC 5236), adult female, collected east of Moramanga in ‘ Julia Forest’ (18.9511°S, 48.2719°E, 941 m a.s.l.) on 6 January 2015; ZSM 147 /2016 (FGZC 5175), adult female collected south of Moramanga (19.0192°S, 48.2341°E, 903 m a.s.l.), Alaotra-Mangoro Region, eastern Madagascar, on 4 January 2016, both by F. Glaw, D. Prötzel, L. Randriamanana. Diagnosis (based on the type series; osteology based on micro-CT scan of ZSM 618/2009, male): Calumma emelinae sp. nov. is characterised by (1) a medium size (male SVL 46.6 – 48.7 mm, female SVL 40.1 – 49.1 mm; male TL 93.6 – 103.2 mm, female TL 82.7 – 95.8 mm), (2) a medium (2.3 – 2.9 mm in males 1.5 – 1.8 mm in females) and distally rounded rostral appendage, (3) rostral scale not integrated into the rostral appendage, (4) rostral crest present, (5) lateral crest present, (6) temporal crest present, (7) cranial crest variable, (8) parietal crest usually absent, (9) casque low in males with a height of 0.5 – 1.1 mm, (10) a dorsal crest of 7 – 10 spines in males, absent in females, (11) 12 – 16 supralabial scales with a mostly straight upper margin, serrated anteriorly, (12) absence of axillary pits, (13) diameter of the largest scale in the temporal region of the head 0.6 – 1.0 mm, (14) no frontoparietal fenestra, (15) parietal and squamosal in contact, (16) parietal bone width at midpoint 16.2% of skull length (n = 1), (17) a generally greyish to greenish body colouration, (18) rostral appendage colour generally unremarkable, (19) a green cheek colouration, (20) suggestions of two weak bluish lateral blotches, and (21) no strong eye colouration. Calumma emelinae sp. nov. can easily be distinguished from all species of the C. boettgeri complex (see above) by the absence of occipital lobes; from C. gallus by different length, shape and colour of its rostral appendage (see above); from C. vatosoa by presence of a rostral appendage (vs absence); from C. vohibola by generally longer relative rostral appendage length (RRS 3.1 – 6.1% vs 0.1 – 1.4%), dorsal crest always present in males (vs generally absent), and pointed tip of postparietal process (vs relatively broad), and crenate prefrontal (vs smooth); from C. nasutum as redefined herein by a lower casque in males (0.5 – 1.1 mm vs 1.5 – 2.0 mm), dorsal crest present in males consisting of spines (vs general absence or consisting of cones if present), and scales more homogeneous (largest temporal scale in males 0.7 mm vs 0.9 – 1.6 mm); from C. radamanus by relatively longer tail in males (longer than SVL vs shorter), longer rostral appendage in males (RRS 4.7 – 6.1% vs 2.9 – 3.6%), rostral scale not integrated into rostral appendage (vs integrated), supralabials with a largely straight upper margin (vs serrated), and parietal and squamosal in contact (vs widely separated); for diagnosis against C. fallax, see below. For diagnosis against the other species described herein, see their respective descriptions below. Description of the holotype (Fig. 4C): Adult male, with mouth closed, in good state of preservation, both hemipenes fully everted; SVL 47.9 mm, tail length 50.7 mm, for other measurements, see suppl. Table 1; rostral ridges that give the snout a right angle; laterally flattened and distally rounded rostral appendage of small tubercle scales that projects straight forward over a length of 2.9 mm with a diameter of 2.4 mm not including the rostral scale; 14 infralabial and 15 supralabial scales, both rather small; supralabials with a straight upper margin; distinct lateral crest running horizontally; no temporal or cranial crest; low parietal crest; no occipital lobes; very low casque of 0.5 mm height; dorsal crest consisting of 10 spines; no gular or ventral crest. Body laterally compressed with fine homogeneous scalation and slightly larger scales on extremities and head region, largest scale in temporal region with diameter of 0.7 mm and in cheek region of 0.9 mm; no axillary or inguinal pits. A B Skull osteology of the holotype (Fig. 11A): Skull length 11.7 mm; snout-casque length 13.8 mm; narrow paired nasals still slightly connected at anterior end; anterior tip of frontal extending to about the middle of the prefrontal fontanelle, which is fused with the naris; prominent and broad prefrontal with laterally raised tubercles; frontal and parietal covered with a few tubercles; frontal with a width at border to prefrontal of 3.0 mm (25.6% of skull length) extending to 4.4 mm (37.6%) at border to postorbitofrontal; no frontoparietal fenestra; narrow parietal with a width at the border to postorbitofrontal of 4.2 mm (35.9%) and a width at midpoint of 1.9 mm (16.2%) tapering strongly posterodorsally; parietal laterally in strong contact with the squamosals; squamosals relatively thick and covered with several tubercles. For further measurements, see Table 2. Hemipenial morphology, based on ZSM 618/2009 (Fig. 5C), ZSM 660/2014 and ZSM 663/2014: medium sized calyces (hemipenial character A); two pairs of small rotulae on apex of about the same size (B), roughly denticulated with about 9 – 12 tips each; papillary field of small, unpaired papillae (C); pair of short cornucula gemina (D), only visible when hemipenis fully everted. Variation: For variation in measurements, see Table 1. Sexual dimorphism: Males are usually larger than females (mean TL of 98.5 mm vs 88.3 mm). Tail length is generally longer in males than in females (RTaSV> 100% vs 2.0 mm vs Colouration in life (Fig. 12): Both sexes with an indistinct brown to beige body colouration, extremities and tail of same colour as the body; three diffuse/scattered brown dorsoventral blotches can occur on the body; males can show a beige lateral stripe; throat and ventral region beige; rostral appendage not accentuated; cheek region can be light green; eyelids crossed by a brown stripe and occasionally with radiating dark green stripes in both sexes. Etymology: The specific epithet is named after Emelina Widjojo, the mother of Wewin Tjiasmanto, in recognition of her support for taxonomic research and nature conservation projects in Madagascar through the BIOPAT initiative (http://biopat.de/en/). Distribution (Fig. 9): Calumma emelinae sp. nov. is known in eastern Madagascar from Anosibe An’Ala to Angozongahy (Makira) about 500 km further north (for coordinates, see above), from an elevation of 750 1030 m a.s.l., Published as part of Prötzel, David, Scherz, Mark D., Ratsoavina, Fanomezana M., Vences, Miguel & Glaw, Frank, 2020, Untangling the trees: Revision of the Calumma nasutum complex (Squamata: Chamaeleonidae), pp. 23-59 in Vertebrate Zoology 70 (1) on pages 44-45, DOI: 10.26049/vz70-1-2020-3, http://zenodo.org/record/4394821, {"references":["GEHRING, P. - S., TOLLEY, K. A., ECKHARDT, F. S., TOWNSEND, T. M., ZIEGLER, T., RATSOAVINA, F., GLAW, F. & VENCES, M. (2012). Hiding deep in the trees: Discovery of divergent mitochondrial lineages in Malagasy chameleons of the Calumma nasutum group. Ecology and Evolution, 2, 1468 - 1479."]}

Identity and re-description of Calumma fallax (Mocquard, 1900) Syntypes: MOCQUARD (1900a, 1900b) did not specify the specimen numbers of the types, but stated that they were one male and one female, collected in Forêt d’Ikongo by Grandidier (MOCQUARD, 1900a). By inference from collection data, we follow BRYGOO (1971), GEHRING et al. (2011) and KLAVER (2019) in accepting only two of potentially six specimens as syntypes, MNHN 1899.317, an adult male, and MNHN 1899.318, an adult female, from the locality Forêt d’Ikongo, collected in 1898 – 1899 by Guillaume Grandidier. Lectotype designation: We designate MNHN 1899.317, the adult male syntype, as the lectotype of the species, the remaining syntype, MNHN 1899.318, becoming the paralectotype. Assignment to genetic clade (based on a comparison of the lectotype with the male specimens assigned to clade H, ZSM 693/2003, ZSM 286/2010, ZSM 685/2003, and ZSM 694/2003): Calumma fallax belongs to clade H, according to the high casque (2.1 mm in the lectotype vs 1.3 – 2.5 mm), rounded, oval rostral appendage with a length of (2.7 mm vs 1.8 – 4.3 mm), heterogeneous scalation in head region with diameter of largest scale in temporal region of 1.6 mm vs 0.8 – 1.5 mm, distinct parietal crest ending in the casque, temporal crest present consisting of 1 or 2 tubercles (2 vs 1 – 2); osteology of the skull is almost identical, e.g. width of the frontoparietal fenestra with 16.4% of skull length vs 13.9 – 15.4%. Referred material: The specimens MNHN 1890.430, MNHN 1890.431, MNHN 1890.432, all three adult males, and MNHN 1888.24, adult female, are non-type specimens. In addition, in anticipation of our conclusions on the taxonomic identity of the species, we here refer the following specimens to C. fallax as it is here re-defined: ZSM 685 /2003 (FG / MV 2002-0291), ZSM 693 /2003 (FG / MV 2002-0317) and ZSM 694 /2003 (FG / MV 2002-0318), all three adult males, collected in Ranomafana NP, Vohiparara, near Kidonavo bridge (about 21.22°S, 47.37°E, about 1000 m a.s.l.), Vatovavy-Fitovinany Region, eastern Madagascar, on 16– 20 January 2003 by F. Glaw, M. Puente, L. Raharivololoniaina, M. Thomas, D. R. Vieites; ZSM 134 /2005 (FGZC 2508), female, collected in Andohahela NP (24.5440°S, 46.7141°E, 1548 m a.s.l.), Anosy Region, southeastern Madagascar, on 27 January 2005 by F. Glaw, M. Vences, P. Bora; ZSM 286 /2010 (FGZC 4588), adult male, collected east of Tsinjoarivo, between camps 2 and 1 (19.7103°S, 47.8182°E, 1465 m a.s.l.) on 23 April 2010 by F. Glaw, J. Köhler, P.-S. Gehring, J.L. Brown, E. Rajeriarison; ZSM 313 /2006 (ZCMV 2930), adult female, collected in Ranomafana NP, Vohiparara river and stream/swamp (about 21.25°S, 47.40°E, about 1100 m a.s.l.) on 20 February 2006 by M. Vences, E. Rajeriarison, Y. Chiari, E. Balian; ZSM 476 /2010 (FGZC 4352), adult female, collected in Anjozorobe region, Mananara Lodge (18.4629°S, 47.9381°E, 1298 m a.s.l.), Analamanga Region, eastern Madagascar, on 6 April 2010, ZSM 479 /2010 (FGZC 4575), adult female, collected east of Tsinjoarivo, camp 1 (19.6800°S, 47.7706°E, 1607 m a.s.l.) on 19 April 2010, both by F. Glaw, J. Köhler, P.-S. Gehring, M. Pabijan, K. Mebert, E. Rajeriarison, F. Randrianasolo, S. Rasamison; ZSM 149 /2016 (FGZC 5226) and ZSM 150 /2016 (FGZC 5225), both adult females, collected in Mandraka (18.9122°S, 47.9144°E, 1235 m a.s.l.), Analamanga Region, eastern Madagascar, on 5 January 2016 by F. Glaw, D. Prötzel, L. Randriamanana; ZSM 258 /2016 (FGZC 5291), adult female, collected in Mandraka (18.9133°S, 47.9145°E, 1260 m a.s.l.) on 3 August 2016 by F. Glaw, D. Prötzel, J. Forster, N. Raharinoro. Diagnosis (based on the type series and the referred material, see above; osteology based on micro-CT scans of MNHN 1899.317, MNHN 1890.430, ZSM 693/2003, and ZSM 286/2010, all four males): Calumma fallax is characterised by (1) a medium size (male SVL 42.9 – 50.6 mm, female SVL 40.8 – 50.7 mm; male TL 90.9 – 107.3 mm, female TL 77.3 – 99.8 mm), (2) a long (1.8 – 4.3 mm in males, 1.7 – 3.2 mm in females) and distally rounded rostral appendage, (3) rostral scale not integrated into the rostral appendage, (4) prominent rostral crest forming a concave cup on the snout, (5) lateral crests present, (6) temporal crest generally present, (7) cranial crest generally absent, (8) parietal crest generally present but short, (9) a distinctly raised casque in males with a height of 1.3 – 2.5 mm, (10) a dorsal crest of 6 – 11 cones in males, generally absent in females (one specimen with five cones), (11) 10– 16 supralabial scales with a straight upper margin, (12) absence of axillary pits, (13) diameter of the largest scale in the temporal region of the head 0.8 – 1.8 mm, (14) a frontoparietal fenestra in the skull, (15) parietal and squamosal generally in contact, (16) parietal bone width at midpoint 6.7 – 15.7% of skull length, (17) a generally greenish, greyish, or brownish body colouration, (18) a typically blue or grey nose in non-stressed colouration, (19) a green cheek colouration, (20) three blue dorsoventral stripes on the body and a white lateral stripe, and (21) a diffuse brown strip crossing the eye. C. fallax can be distinguished from all species of the C. boettgeri complex (see above) by the absence of occipital lobes; from C. gallus by different length, shape and colour of its rostral appendage (see above); from all other species of the C. nasutum group without occipital lobes (except for C. ratnasariae, see below) by the presence of a frontoparietal fenestra. In addition, it can be distinguished from C. vatosoa by the presence of a rostral appendage (vs absence); from C. vohibola by longer rostral appendage (RRS 4.2 – 8.5% vs 0.2 – 3.1%), supralabials with a straight upper margin (vs serrated), parietal crest generally present (vs absent); from C. nasutum as here redefined by general absence of cranial crest (vs present), a shorter frontal (39.4 – 50.4% of skull length vs 51.2 – 82.1%), blue rostral appendage (vs brown), and three blue lateral blotches (vs four brown blotches with light spots); from C. radamanus by rostral scale not integrated into the rostral appendage (vs generally integrated), parietal crest generally present (vs absent), supralabials with a straight upper margin (vs serrated), parietal and squamosal in contact or closely approaching (vs widely separated), and width of parietal at midpoint (6.7 – 15.7% vs 16.1 – 22.4%); from C. emelinae sp. nov. by general presence of parietal crest (vs general absence), higher casque in males (1.3 – 2.5 mm vs 0.5 – 1.1 mm), dorsal crest consisting of cones (vs spines) in males, and larger temporal scale in males (0.8 – 1.6 mm vs 0.7 mm); from C. tjiasmantoi sp. nov. by fewer supralabials (10 – 15 vs 15 – 17), larger diameter of temporal scale (1.0 – 1.8 mm vs 0.6 – 0.8 mm), and slightly narrower postparietal process. Re-description of the lectotype (Fig. 4E): Adult male, with mouth slightly opened, in good state of preservation, hemipenes not everted. SVL 42.9 mm, tail length 53.2 mm, for other measurements, see suppl. Table 1; distinct rostral ridges that render the dorsal surface of the snout a concave cup, laterally compressed dermal rostral appendage of oval tubercle scales that projects straight forward over a length of 2.7 mm with a diameter of 3.4 mm, oblong but taller than wide; 11 infralabial and 11 supralabial scales, all relatively large; supralabials with a smooth dorsal margin; distinct lateral crest running horizontally; distinct temporal crest consisting of two tubercles per side; no cranial crest; distinct parietal crest; no occipital lobes; highly elevated (2.1 mm) and rather acute casque; dorsal crest present, consisting of 11 cones that decrease in height posteriorly; no traces of gular or ventral crest. Body laterally compressed with fine homogeneous scalation and distinctly larger scales on extremities and head region, largest scale in temporal region with diameter of 1.6 mm and in cheek region of 1.4 mm; no axillary or inguinal pits. Skull osteology of the lectotype (Fig. 14A): Skull length 12.0 mm; snout-casque length 14.6 mm; narrow paired nasals completely separated from each other by the anterior tip of frontal that meets the premaxilla; prefrotal fontanelle and naris fused; prominent prefrontal with laterally raised tubercles exceeding more than the half of the prefrontal fontanelle; frontal and parietal smooth without any tubercles; frontal with a width of 2.7 mm (22.5% of skull length) at border to prefrontal extending to 4.4 mm (36.7%) at border to postorbitofrontal; large frontoparietal fenestra with a width of 2.4 mm (20.0%); curved parietal tapering strongly from a width of 3.9 mm (32.5%) at the border to postorbitofrontal to a width at midpoint of 0.8 mm (6.7%) and broadening slightly poterodorsally, where it is in weak contact with the squamosals; squamosals thin without any tubercles. For further measurements, see Table 2. Hemipenial morphology, based on the lectotype (no micro-CT scan available): small sized calyces (hemipenial character A); two pairs of finely denticulated rotulae of different size, on sulcal side large with about 16 tips, on asulcal side small with about 8 tips (B); papillary field of few, unpaired papillae (C); pair of medium sized cornucula gemina (D), only visible when hemipenis fully everted. Variation: The osteology of the male ZSM 693/2003 differs from the other specimens in having the parietal and squamosal bone not connected; probably this is due to its juvenile state. For variation in measurements, see Table 1. Sexual dimorphism: Body size (SVL and TL) is slightly larger in males than females. Tail length is longer in males than in females (RTaSV 102 – 124% vs 89 – 104%). Relative rostral appendage length does not differ. Dorsal crest is more pronounced in males than females. Colouration in life (Fig. 15): Weak sexual dichromatism, males slightly more colourful. In both sexes grey/ beige body colouration with three bright blue dorsoventral stripes on the body that can be crossed by a broad white lateral stripe; extremities and tail of same colour as the body, tail in males can be diffusely annulated; rostral appendage grey or blue; cheek region can be bright green; a diffuse brown stripe may cross the eye. Etymology: A Latin adjective meaning ‘deceptive’ or ‘fallacious’ in the neutral nominative, with unclear justification. Distribution (Fig. 9): Calumma fallax as redefined here, occurs in eastern Madagascar from Andohahela in the south to Mandraka about 650 km further north (coordinates, see above), from an elevation of 922– 1781 m a.s.l., Published as part of Prötzel, David, Scherz, Mark D., Ratsoavina, Fanomezana M., Vences, Miguel & Glaw, Frank, 2020, Untangling the trees: Revision of the Calumma nasutum complex (Squamata: Chamaeleonidae), pp. 23-59 in Vertebrate Zoology 70 (1) on pages 48-52, DOI: 10.26049/vz70-1-2020-3, http://zenodo.org/record/4394821, {"references":["MOCQUARD, M. F. (1900 a). Diagnoses d'especes nouvelles de reptiles de Madagascar. Bulletin du Museum national d'histoire naturelle, 6, 344 - 345.","MOCQUARD, M. F. (1900 b). Nouvelle contribution a la faune herpetologique de Madagascar. Bulletin de la Societe philomathique de Paris, 9, 93 - 111.","BRYGOO, E. R. (1971). Reptiles Sauriens Chamaeleonidae. Genre Chamaeleo. Faune de Madagascar, 33, 1 - 318.","GEHRING, P. - S., RATSOAVINA, F. M., VENCES, M. & GLAW, F. (2011). Calumma vohibola, a new chameleon species (Squamata: Chamaeleonidae) from the littoral forests of eastern Madagascar. African Journal of Herpetology, 60, 130 - 154.","KLAVER, C. (2019). Notes on the typification of several chameleon taxa (Sauria: Chamaeleonidae). Amphibia-Reptilia, DOI: 10.1163 / 15685381 - 20191230."]}

Uroplatus finaritra sp. nov. Figs. 1–5, Table 1 Uroplatus sp. G—Raxworthy et al. (2008) Uroplatus sp. Ca9— Ratsoavina et al. (2013) LSID: urn:lsid:zoobank.org:act: E6A1407B-7C9E-46F6-8FEC-D120A111A344 Holotype. UADBA-R 70489 (field number APR 12522), adult female, collected on the eastern slope of Marojejy by a tributary of the Ampanasatongotra River, Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.4559°S, 49.7749°E (780 m a.s.l.) on the night of 15 May 2016 by A. P. Raselimanana. Paratypes. UADBA-R 70493 (field number APR 12591), adult male, UADBA-R 70490 (APR 12590), adult male, and UADBA-R 70491 (APR 12589), adult female, three specimens collected in the same locality as the holotype (750–810 m a.s.l.) on 18 May 2016 by A. P. Raselimanana; UADBA-R 70492 (APR 12691), adult male, and UADBA-R 70494 (APR 12692), juvenile female, two specimens collected on the western slope of Marojejy by Tsiasisa River, 2.2 km east of Antanimbaribe, Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.5074°S, 49.6153°E (845 m a.s.l.) on the night of 28 May 2016 by A. P. Raselimanana; UADBA-R 70501 (MSZC 0253), subadult female, ZSM 458 /2016 (MSZC 0271), subadult female, two specimens collected near the path below Camp 1 (Camp Mantella) of Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.438°S, 49.776°E (450 m a.s.l.) on 21–23 November 2016 by M. D. Scherz, M. Bletz, A. Rakotoarison, J. Razafindraibe, A. Razafimanantsoa, and M. Vences. Etymology. Finaritra is a Malagasy word for special greetings but also means healthy and happy. We refer to our delight in describing this splendid and exceptionally large species from a clade of generally small-sized leaftailed geckos. The name is an invariable noun in apposition. Diagnosis. Uroplatus finaritra sp. nov. is assigned to the Uroplatus ebenaui group based on its relatively small size and its triangular head with supraocular spines, and overall leaf-mimicking aspect. It is characterised by (1) a dark red oral mucosa, (2) a comparatively large body size (82.5–95.3 mm SVL in adults), (3) 7–8 lamellae (exceptionally 6) under the third finger and toe, and (4) a long (length 53–65% of SVL), wide (width 16–18% of SVL), leaf-shaped 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 smaller size (SVL 82.5–95.3 mm vs. 85–200 mm) and the lack of lateral dermal fringes on any part of the body (vs. presence except in U. lineatus). Also, Uroplatus finaritra sp. nov. diverges from the species of the U. fimbriatus group by their lack of lateral body compression. The prominent triangular head, supraocular spines, and rather smooth skin distinguish Uroplatus finaritra sp. nov. from U. alluaudi, U. guentheri, U. malahelo and U. pietschmanni. Amongst the U. ebenaui group, the rather long tail (TAL/SVL 0.53–0.68) differentiates Uroplatus finaritra sp. nov. from U. ebenaui, U. fiera, U. fotsivava, and U. kelirambo (TAL/SVL 0.22–0.44), and furthermore it differs from U. fiera and U. fotsivava by a pigmented oral mucosa (vs. unpigmented). The new species differs from U. finiavana by a wider tail (TAW/SVL 0.16–0.18 vs. 0.05–0.14), and by its pigmented oral mucosa (vs. unpigmented). By its large tail and pigmented oral mucosa, the new species is most similar to U. malama and U. phantasticus. It differs from U. phantasticus by larger size (SVL 83–95 mm vs. 52–76 mm), a generally shorter tail (TAL/SVL 0.53–0.65 vs. 0.62–0.76), a typically higher number of lamellae under the third finger and toe (6–8 vs. 5–7), and the absence of black pigment of the oral mucosa (oral mucosa dark red vs. black). It differs from U. malama by a spinier integument in some adult males, by the absence of black pigment of the oral mucosa (oral mucosa dark red vs. black), and probably by a narrower tail (TAW/SVL 0.16–0.18 vs. 0.24). In the mitochondrial phylogeny, the new species occupies an isolated position within the U. ebenaui group, without affinities to the morphologically similar U. phantasticus and U. malama (which are not each other's closest relatives either). The nuclear gene c-mos reveals haplotype sharing of the new species with U. kelirambo and two undescribed candidate species (U. sp. Ca3 and Ca4) occurring in the same region of Madagascar, but morphologically very different by their short tails. TABLE 1. Morphological measurements and scale counts of Uroplatus finaritra sp. nov. For abbreviations used, refer to Methods; additional abbreviations: M, male; F, female; sa, subadult; j, juvenile; HT, holotype; PT, paratype; na, not applicable (due to missing original tail); nm, not measured. All measurements in mm. Description of the holotype. Adult female in fair condition, with a fully intact tail, both of its hind limbs fractured (Fig. 2). SVL 95.3 mm, tail length 50.9 mm, maximum tail width 17.0 mm; for further measurements see Table 1. Head triangular in dorsal view; canthus rostralis indistinct and concave; snout sloping weakly downwards anteriorly; snout weakly depressed, fairly long (1.7 times longer than eye diameter); eyes large (eye diameter 6.5 mm), bulging slightly above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small (horizontal diameter 1.1 mm), its opening facing posterolaterally, but also posteroventrally (ear opening clearly visible in ventral view but not in dorsal view); nostrils laterally oriented; body somewhat laterally compressed, without lateral dermal fringes; limbs without fringes and very few, miniscule spines on the hind limbs (one on the knee, as well as a small flap of skin), and two miniscule spines on the forelimb (on the elbow and near the wrist); forelimb reaches the canthus rostralis when adpressed forward and midbody when adpressed backwards along body (forelimb length/axilla–groin distance 33.4/ 50.2 mm = 67%), hindlimb reaches ¾ up the body when adpressed forward along body (hindlimb length/axilla–groin distance 41.4/ 50.2 mm = 83%); original tail length 53% of snout–vent length, membranous borders of the tail symmetrical, without any emarginations, long and leafshaped, with a thin, borderless spatulate tip. Toes bear small claws, with the distal phalange not much wider than the rest of the digit; the third finger bears 7 lamellae, the third toe 8 lamellae. Nares separated from each other by at least 10 small granular scales, from the first supralabial by 3 scales, and from the rostral scale by 5 scales; first supralabial not taller than the others; rostral entire, much wider than tall; mental scale very small, not differentiated from infralabial scales; 25/26 (right/left) supralabials and 25/25 infralabials; 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, except on the posterior ventral abdomen, where they are slightly larger than the rest of the body, and arranged almost uniformly. 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, fairly straight line (also formed by a row of slightly enlarged scales) is present between the eyes and connects the supraocular spines. Another, less distinct line is present connecting the anterior angle of the eyes over the snout in a bowed line. The body possesses very few dermal spines—a prominent pointed supraocular spine, a single spine at the posterior angle of the head, and a small number of spines on each limb (described above). No axillary pits present. Colouration. After two years of preservation in 70% ethanol the colour is slightly faded (Fig. 2). All dorsal surfaces of the specimen are a ruddy brown, fading to a salmon brown in the neck region and at the insertion of the hind limbs. A fine network of blackish lines is present across the body, with the following notable features: a somewhat straight line connecting the suprocular spines (described above); a v-shaped marking on the neck, which extends into a vertebral stripe that reaches all the way to the tail tip; a number of posteriorly oriented chevrons from the mid-dorsum to the belly, becoming thinner posteroventrally, three of which are most distinct. The exterior of the shank and outermost two toes are grey-black, whereas the inner foot is brown. The dorsal snout is darker than the rest of the head. The tail is not different in colour or pattern from the dorsal body. A distinct lateral line runs along the ventrolateral edges of the abdomen from the insertion of the leg to the axilla. The ventral colouration is uniformly orange-brown, with a faint, symmetrical pattern of dark lines on the chin. The soles of the hands and feet are grey. The ventral tail is the colour of the trunk mottled with grey splotches. A faint cream tear-like marking is present at the posterior corner of the eye above the supralabial. The oral mucosa is dark red, with a pink tongue and cream under the eyes and along the lower jaw. In life, the iris colour was silver externally and rusty around the pupil (Fig. 3d). Variation. Molecular data are also available from a previous paper (Raxworthy et al. 2008) for one specimen (RAN 42274) from Marojejy as Uroplatus sp. G, included for 12S and cytochrome b as U. sp. Ca 9 in Ratsoavina et al. (2013). The available sequence for 12S agrees with those of our specimens. The paratypes were sexed as adult males by the presence of everted hemipenes (UADBA-R 70493) or presence of distinctly enlarged tail base (UADBA-R 70490 and 70492), and distinctly notched tail edges; or as females based on the absence of hemipenes or hemipenial bulges, presence of visible oviduct or ovary structures after dissection, and generally smooth tail edges. In the smallest specimen (UADBA-R 70494) female inner reproductive organs were observed but were only weakly developed, and we therefore consider this specimen as juvenile. The sexual maturity of two further specimens (UADBA-R 70501 and ZSM 458/2016) is unclear, as they are considerably smaller than the large adult females (UADBA-R 70491 and 70489) but have similar gonad development. We tentatively consider these to be subadults. In general, the paratypes agree well with the holotype in morphology. For measurements, see Table 1. The number of head spines varies from 2–9, and is more in males than in females, as males are more spiny than females in general, and have more heterogeneous scalation. Forelimbs can bear a minimum of two spines, or as many as eight, with a small spine present on the elbow. Hind limbs can bear 2–15 spines, and the dermal knee flap almost always bears at least a small spine, which is smaller in females than males. In males the tail is moderately emarginated, with semicircular emarginations that tend to be asymmetrical. In females the tail is without emarginations and has smooth edges. Internarial scales range from nine to ten, supralabials from 25–29, and infralabials from 21–25. The third finger bears 7–8 lamellae (6 in one juvenile specimen, UADBA-R 70494), the third toe bears 7–8 lamellae. The colouration is varied, as is commonly the case in Uroplatus species (Figs. 3–4). The line on the head connecting the supraocular spines is generally bowed, but can be slightly pointed posteriorly, and in UADBA-R 70492, it is irregular at its midpoint. The dark vertebral stripe is only present in the holotype, UADBA-R 70491 and 70494, and in other specimens is either absent, or is cream and not dark (in UADBA-R 70492 and 70490). Overall dorsal colouration differs dramatically, from almost completely reddish brown (UADBA-R 70501) to highly mottled greys and blacks (UADBA-R 70493). Irregular flecks can be present or absent on any part of the body. The different colouration of the outer shank and foot is always present, and a cream tear-like marking below the eye is almost always present (absent in UADBA-R 70494 and 70501). The ventrolateral stripe is always present but not always distinct (indistinct in UADBA-R 70493). Ventral colouration is as variable as dorsal colouration. UADBA-R 70491 is extremely similar to the holotype, and to a lesser degree 70492, 70494, and 70501 resemble the holotype ventrally. UADBA-R 70490 and 70493 differ dramatically in being darker in colour, having irregular flecks of cream and reddish brown, and lacking the symmetrical markings on the chin. In life, the colouration of the iris is typically silvery but can be red (Figs. 3c, f), and the oral mucosa was consistently dark red (Fig. 5). Distribution. At present this species has only been collected on the Marojejy Massif, at altitudes from 450–845 m a.s.l. Natural History. The holotype was collected active at night on a stem 0.5 m above the forest floor in nearly closed canopy humid forest at the bottom of a slope, in the vicinity of a Uroplatus lineatus specimen. UADBA-R 70490, 70491, and 70493 were collected on small branches and leaves between 1.5 and 2.5 m above the ground in similar forest. UADBA-R 70492 and 70494 were also active on branches 3–4.5 m from the forest floor in nearly closed-canopy humid forest with a thick understory and ferns. UADBA-R 70501 and ZSM 458/2016 were collected active at night on thin branches (up to 4 m above the ground) in closed-canopy, low-altitude rainforest. We consider the various collection localities to represent a single Threat-Defined Location in the sense of the International Union for the Conservation of Nature (IUCN 2012), as we have recently done for several frog species from low- to mid-elevation forests in Marojejy (Scherz et al. 2016, Rakotoarison et al. 2017). In line with these assessments, we recognise that there is on-going habitat degradation due to illegal logging activity, especially at low altitudes in and around Marojejy National Park. The Extent of Occurrence and Area of Occupancy of the new species is certainly greater than 10 km 2, but is likely less than 500 km 2, and as such we recommend a status of Endangered for it under criterion B1ab(iii).

Hutter, Carl Richard, Andriampenomanana, Zo F., Razafindraibe, Jary H., Rakotoarison, Dr. Andolalao, Scherz, Mark 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 (39-40): 2497-2510, DOI: 10.1080/00222933.2018.1543732, URL: http://dx.doi.org/10.1080/00222933.2018.1543732

CALUMMA LEFONA SP . NOV . urn:lsid:zoobank.org:act:99 AF 4 F 23- EBDF- 4276- AE 41- 3 C 04 E 7443 FD 8 Remark: This species was considered as clade FII by Gehring et al. (2012). Holotype: ZSM 2 8 4 9/ 2 0 1 0 (DRV 6 2 8 7), adult male with incompletely everted hemipenes, left hemipenis removed for micro-CT scanning, collected in Andrevorevo southeast of Tsaratanana Massif (14.3464�� S, 49.1028�� E, 1717 m a.s.l.), Mahajanga Province, northern Madagascar, on 21 June 2010 by F. M. Ratsoavina and F. Randrianasola. Paratypes: None. Diagnosis: Calumma lefona sp. nov. is a member of the phenetic C. nasutum species group (Pr��tzel, Ruthensteiner & Glaw, 2016), on the basis of the presence of a soft, dermal, unpaired rostral appendage, absence of gular or ventral crests, and heterogeneous scalation on the lower arm, consisting mostly of tubercles of ~ 0.5 mm diameter. Within the genus, it is a small sized chameleon (SVL 51.3 mm, TL 113.7 mm) that is characterized by a long and pointed rostral appendage, occipital lobes that are widely notched and completely separated, a distinctly elevated rostral crest, a dorsal and caudal crest, absence of axillary pits and unique skull morphology. Calumma lefona sp. nov. differs from C. fallax, C. gallus, C. nasutum, C. peyrierasi, C. vatosoa, and C. vohibola of the C. nasutum group by the presence of occipital lobes; from C. boettgeri and C. linotum in the clearly notched occipital lobes with a depth of 1.8 mm (vs. not or slightly notched with 0���0.7 mm), presence of a frontoparietal fenestra with a width of 1.9 mm (14.7% of skull length; vs. completely closed brain case), prefrontal, frontal and parietal with many tubercles (vs. smooth or only a few tubercles); additionally, from C. boettgeri by the higher number (21 in line) of large, juxtaposed tubercle scales on the extremities (vs. seven to 14 isolated tubercles). From the other three taxa with notched occipital lobes, C. gehringi, C. guibei, and C. uetzi sp. nov., C. lefona sp. nov. differs by the long (5.6 mm) and pointed rostral appendage (vs. 3.1���5.4 mm in males, rounded), with 60 large scales (diameter> 0.3 mm) on the right side of the rostral appendage (vs. 20���42 in males), dorsal crest of 23 small conical scales (vs. absence or 5���15 large conical scales), clearly and widely separated occipital lobes with a notch of 1.8 mm (vs. tightly separated or connected with a notch of 0.5���1.5 mm in males); furthermore, in skull morphology by a narrower frontal, e.g. at the border to the postorbitofrontal with 30.5% of SkL (vs. 35.5���39.7%) and strongly raised maxillae; from C. gehringi and C. guibei in a broader parietal at midpoint with 18.3% of SkL (vs. 9.9���15.3%); from C. uetzi sp. nov. by possession of a frontoparietal fenestra (vs. completely closed brain case) and a narrower head (e.g. RFWPo of 30.5 vs. 35.6���39.4%); from C. guibei by a smaller frontoparietal fenestra with 14.7% of SkL (vs. 21.0%), prefrontal fontanelle and naris separated by contact of prefrontal with maxilla (vs. fused), thick squamosal (vs. thin) in broad dorsal contact with the parietal (vs. not meeting parietal or only in weak contact); and from C. gehringi by possession of a larger frontoparietal fenestra (21.0% of SkL vs. 5.8���9.1%). Description of the holotype (Fig. 3 B): Adult male, in a good state of preservation, left forelimb removed for DNA analysis, left hemipenis removed for micro-CT scanning; mouth slightly opened with tongue between the jaws; originally both hemipenes incompletely everted (Fig. 5 B), but left hemipenis cut off for micro-CT scanning and stored in a separate Eppendorf tube alongside the specimen; SVL 51.3 mm, tail length 62.4 mm; see Table 1 for other measurements; distinct and elevated rostral ridges that form a concave cup on the snout and fuse on the anterior snout at the base of a tapering, laterally compressed dermal rostral appendage that projects straight forward over a length of 5.6 mm with a diameter of 2.0 mm, pointed distally; 12 infralabial and 11 supralabial scales; supralabials dorsally serrated; no supra-orbital crest; distinct lateral crest running horizontally; short temporal crest consisting of two tubercles per side; indistinct parietal crest; occipital lobes clearly developed and completely separated by a deep, ���U���-shaped notch (1.8 mm); casque raised; dorsal crest present, starting 0.8 mm from the base of the notch between the occipital lobes, consisting of a row of 23 separated, small conical scales spaced at irregular intervals from 0.1 to 0.9 mm, and several more on the tail decreasing in size towards the tip; no traces of gular or ventral crest. Body laterally compressed, with fine homogeneous scalation with the exception of the extremities and head region; limbs with rounded tubercle scales with a maximal diameter of 0.5 mm; heterogeneous scalation on the head, with largest scale on temporal region with diameter of 0.6 mm and 60 oval tubercle scales (diameter> 0.3 mm) on the right side of the rostral appendage; no axillary or inguinal pits. Skull osteology of the holotype (Fig. 4 B, Supporting information, Video S 3): Skull length 13.1 mm; snout���casque length 15.9 mm; broad paired nasals meeting anteriorly, anterior tip of frontal exceeding more than half of the naris; prefrontal fontanelle and naris separated by contact of prefrontal with maxilla; elevated maxillae building a rectangular edge at anterior margin; prominent prefrontals with laterally raised tubercles; frontal and parietal with several tubercles, five on the midline forming a parietal crest; frontal with a width of 2.9 mm (22.1% of SkL) at border to prefrontal extending to 4.0 mm (30.5% of SkL) at border to postorbitofrontal; frontoparietal fenestra with transverse diameter of 1.9 mm (14.7% of SkL); parietal tapering more or less constantly, with a width of 3.9 mm (29.8% of SkL) at the border to frontal and 2.4 mm (18.3% of SkL) at its midpoint, and then tapering rather strongly to the posterior tip; posterodorsally directed parietal meets the squamosal laterally; squamosal thick, with several tubercles. For further measurements, see Table 2. Coloration of the holotype in preservative (Fig. 3 B): The body of the holotype in preservative is of a grey���blue colour, with an indistinct beige lateral stripe; ventral and temporal regions and throat also beige; dark blue line from dorsal part of rostral appendage crossing the eyes and the lateral crest, ending in occipital lobes; extremities speckled with bluish tubercle scales. The coloration in life is unknown. Hemipenial morphology based on diceCT scans: The hemipenis of the holotype was incompletely everted, and a diceCT scan resulted in an inadequate illustration (Fig. 5 B). Apparently, it showed deep calyces and two pairs of rotulae; changing the threshold revealed two pairs of cornucula that were not everted. Available names: Apart from C. gehringi, C. guibei, and C. uetzi sp. nov., there is no other valid species or synonym in the C. nasutum group with deeply notched occipital lobes. Etymology: Calumma lefona sp. nov. is the only species in the C. nasutum group with a relatively long and pointed/constantly tapering rostral appendage (with the exception of C. gallus). This shape reminds of a spearhead; accordingly, we chose the Malagasy word ���lefona��� (meaning ���spear���) as its species epithet. It is used as an invariable noun in apposition to the genus name. Distribution: Calumma lefona sp. nov. is, so far, known from only a single location in northern Madagascar, southeast of the Tsaratanana Massif, called Andrevorevo, at 1717 m a.s.l. (Fig. 6; for geographical coordinates see the ���holotype��� details above). It lives within the lower elevational range of its sister taxon C. guibei (from 1590���2250 m a.s.l.) and at higher elevation than C. gehringi (730���1540 m a.s.l.). Natural history and ecology: Calumma lefona sp. nov. was found roosting at night on tree branches ~ 2 m above the ground. The habitat consisted of primary forest with closed canopy cover and small streams. During the nocturnal observations on 20 and 21 June 2010, two adults and two juveniles were found along the forest path, but only one adult (the holotype) was collected. Recommended IUCN status: Practically no data are available on the distribution and condition of C. lefona sp. nov., and the species is known from a single specimen and a few additional observations. To avoid inflation of perceived risk, we recommend that this species be considered Data Deficient by the IUCN until more data become available., Published as part of David Pr��tzel, Miguel Vences, Oliver Hawlitschek, Mark D. Scherz, Fanomezana M. Ratsoavina & Frank Glaw, 2018, Endangered beauties: micro-CT cranial osteology, molecular genetics and external morphology reveal three new species of chameleons in the Calumma boettgeri complex (Squamata: Chamaeleonidae), pp. 1-28 in Zoological Journal of the Linnean Society 2018 (20) on pages 12-15, DOI: 10.1093/zoolinnean/zlx112, http://zenodo.org/record/1226487, {"references":["Protzel D, Ruthensteiner B, Glaw F. 2016. No longer single! Description of female Calumma vatosoa (Squamata, Chamaeleonidae) including a review of the species and its systematic position. Zoosystematics and Evolution 92: 13 - 21."]}

CALUMMA JULIAE SP.NOV. urn:lsid:zoobank.org:act:6 A 92 C 95 D- 6383-45DC-9 FFC- 0943 B 08 F 064 F Remark: Despite intensive research at the type locality in January (by L. Randriamanana, F. G. and D. P.), August (by N. Raharinoro, K. Glaw, T. Glaw, J. Forster, F. G. and D. P.) and November 2016 (by A. Rakotoarison and M.D. S.) in the rainy and dry seasons, only female specimens of this new species were found. In total, eight adult females, two subadult females and two juveniles were encountered (but not all of them collected). Holotype: ZSM 143/ 2016 (FGZC 5235), adult female, collected in small forest fragment 5 km east of Moramanga, just south of the Route Nationale 2 (18.9520�� S, 48.2707�� E, at 950 m a.s.l.), Toamasina Province, eastern Madagascar on 6 January 2016 by F. Glaw, D. Pr��tzel and L. Randriamanana. Paratypes: ZSM 142 / 2016 (FGZC 5233), FGZC 5232 and FGZC 5234 (two uncatalogued specimens in UADBA), all three adult females, collected from the same location as the holotype (18.9519�� S, 48.2705�� E, within a radius of 50 m, at 950 m a.s.l.) on 6 January 2016 by F. Glaw, D. Pr��tzel, and L. Randriamanana. ZSM 254 / 2016 (FGZC 5274), adult female, and ZSM 255/ 2016 (FGZC 5275) and FGZC 5276 (uncatalogued specimen in UADBA), both juveniles, all three collected on 30 July 2016; FGZC 5277 (uncatalogued specimen in UADBA), subadult, collected on 31 July 2016 at the same location as above by F. Glaw, D. Pr��tzel, and N. Raharinoro. Diagnosis: Male specimens are unknown so far, hence the diagnosis refers only to females of this species. Calumma juliae sp. nov. is a member of the phenetic C. nasutum species group (Pr��tzel, Ruthensteiner & Glaw, 2016), on the basis of the presence of a soft, dermal unpaired rostral appendage, absence of gular or ventral crests, and heterogeneous scalation on the lower arm, consisting mostly of tubercles of a diameter of 0.7���0.8 mm. Within the group, it is a large (TL 105.3���111.6 mm), grey���beige chameleon that is characterised by a long and distally rounded rostral appendage, a dorsal crest of 11���14 tubercles, occipital lobes that are clearly notched but not completely separated, and absence of axillary pits. Calumma juliae sp. nov. differs from C. fallax, C. gallus, C. nasutum, C. peyrierasi, C. vatosoa, and C. vohibola of the C. nasutum group by the presence of occipital lobes; from C. gehringi, C. guibei, and C. lefona sp. nov. in the completely closed brain case (vs. frontoparietal fenestra); additionally, from female C. gehringi and C. guibei in body size of 53.3���59.4 mm SVL (vs. 47.5���52.3 mm); from female C. gehringi in the shorter rostral appendage of 2.3���2.7 mm (vs. 3.2���4.4 mm); from C. guibei (both sexes) in the notch between the occipital lobes of 0.2���0.8 mm (vs. completely separated with notch of 1.2���1.9 mm; see Brygoo, 1971); from C. lefona sp. nov. (one male) in the shorter (2.3���2.7 mm) and rounded rostral appendage (vs. 5.6 mm, pointed), the absence of a temporal and parietal crest (vs. presence) and the number of dorsal cones of 11���14 (vs. 23); from C. boettgeri (both sexes) by the higher number of large (0.7���0.8 mm diameter) juxtaposed tubercle scales on the extremities (17���19 in line vs. 7���14, diameter of 0.2���0.5 mm and isolated from each other); from female C. uetzi sp. nov. in the larger body size of 53.3���59.4 mm SVL (vs. 42.0 mm SVL in females), the absence of a temporal and parietal crest (vs. presence of both) and the higher number of infralabial scales of 14���15 (vs. 11���12); from its most similar taxon C. linotum by the clearly notched occipital lobes with a depth of 0.2���0.8 mm (vs. not or slightly notched with depth 0���0.2 mm), presence of a dorsal crest in females consisting of 11���14 conical scales in C. juliae sp. nov. (vs. zero in C. linotum and six in C. cf. linotum; one specimen from Andampy), higher number of infralabial scales in females of 14���15 (vs. 12���13), absence of temporal and parietal crest in females (vs. both present), and in generally larger body size in females of 53.3���59.4 mm SVL (vs. 42.7���54.5 mm). In skull morphology, the squamosal and parietal do not meet in female C. juliae sp. nov. (vs. broad in contact in male and female C. linotum (Pr��tzel et al., 2015); frontal of triangular shape and narrower, e.g. 16.8���18.2% of SkL at border to prefrontal (vs. 22.1���24.4%), 30.7��� 31.1% of SkL at border to postorbitofrontal (vs. 34.4��� 35.8%) and 25.0���26.3% of SkL bordering the parietal (vs. 32.5���33.6%); also, parietal is narrower at border to frontal with 28.5���29.5% of SkL (vs. 34.1���35.1%). Description of the holotype (Fig. 3 C): Adult female, in a good state of preservation; SVL 55.8 mm, tail length 55.4 mm; for other measurements, see Table 1; distinct rostral crest, laterally compressed dermal rostral appendage that projects forward of the snout tip over a length of 2.7 mm with a diameter of 2.2 mm, rounded distally; 14 infralabial and 13 supralabial scales; supralabials dorsally serrated; no supra-orbital crest; distinct lateral crest running horizontally; no temporal crest; occipital lobes clearly developed and separated but still slightly connected by a notch of 0.5 mm; casque raised; dorsal crest present, starting 4.3 mm from the base of the notch between the occipital lobes, consisting of a row of 12 separated, small conical scales spaced at regular intervals of ~ 2 mm, continuing on the tail with smaller and narrower spaced cones; no traces of gular or ventral crest. Body laterally compressed, with fine homogeneous scalation with the exception of the extremities and head region; limbs with rounded tubercle scales of maximal diameter 0.7 mm; heterogeneous scalation on the head, with largest scale on temporal region having a diameter of 1.3 mm; no axillary or inguinal pits. Skull osteology of the holotype (Fig. 4 C, Supporting information, Video S4): Skull length 13.2 mm; snout��� casque length 16.1 mm; broad paired nasals meeting each other; anterior tip of frontal exceeding less than half of the naris and separated from premaxilla; prefrontal fontanelle and naris fused; frontal smooth, and parietal with only a few tubercles; frontal with width of 2.4 mm (18.2% of SkL) at border to prefrontal extending to 4.1 mm (31.1% of SkL) at border to postorbitofrontal; broad parietal tapering strongly from a width of 3.9 mm (29.5% of SkL) at the border to frontal to 0.8 mm (6.1% of SkL) at midpoint, then extending mostly straight beyond the posterodorsal extension of the squamosals, and finally tapering again at its tip; posterodorsally directed parietal does not meet the squamosal; squamosal thin without tubercles. For further measurements, see Table 2. Coloration of the holotype in preservative (Fig. 3C): The body of the holotype in preservative is of dark grey colour without any distinct pattern; inner side of extremities and tail beige; extremities and temporal and postocular region of the head speckled with greyish���blue tubercle scales. Variation: All adult female C. juliae sp. nov. that have been found so far show a consistent morphology and consequently agree well with the holotype. ZSM 254/2016 and FGZC 5234 have a slightly shorter rostral appendage than the holotype, at 2.4 and 2.3 mm (vs. 2.7 mm); the notch between the occipital lobes is deeper in FGZC 5232 at 0.8 mm and less deep in ZSM 142/ 2016 at 0.2 mm (vs. 0.5 mm); the dorsal crest of ZSM 255/ 2016 consists of only nine tubercles (vs. 12); this might be referred to its juvenile developmental stage. In osteology, no significant variations were found. Coloration in life (Fig. 7): Females are indistinctly grey���beige coloured; a netlike dark brown pattern on the skin between the scales and two dorsoventrally compressed blue blotches may occur on the body; tubercle scales on extremities and body may be bright green; rostral appendage not highlighted and of same colour as the body; a dark lateral stripe may stretch from the rostral appendage across the eyes to the occipital lobes. If stressed, only the head coloration changes to dark brown or green pattern at the dorsal head region, eyelids with radially aligned blue/violet spots, and rostral crest and appendage of turquoise/ green colour. Available names: Apart from C. boettgeri and C. linotum, there is no other valid species or synonym in the C. nasutum group with slightly notched occipital lobes. Etymology: D. P. dedicates the first new species he discovered himself to Julia Forster, in recognition of her generous support and understanding of our research on Madagascan chameleons and her help in collecting specimens of this species. Distribution: Calumma juliae sp. nov. has so far been recorded only from a fragment of degraded primary rainforest, covering an area of just 15 ha, east of Moramanga (Toamasina Province, eastern Madagascar; Figs 6, 8 A); for geographical coordinates, see the descriptions in the subsections ���Holotype��� and ���Paratypes��� above. The forest fragment spreads over a hill that rises to a maximum elevation of 1010 m a.s.l.; specimens were found only along the trail at 950 m a.s.l., but we expect them to occur on the hill as well. A further specimen was observed just north of the Route Nationale 2 (18.9513�� S, 48.2721�� E, 950 m a.s.l.) in secondary bushes. Natural history and ecology: Calumma juliae sp. nov. is an arboreal, diurnal species found in bushes and trees in a small forest fragment of degraded primary rainforest. Roosting sites at night were thin branches or, rarely, leaves that were not exposed, but hidden inside tree/bush cover 0.3���2 m above the ground. In contrast to the syntopically occurring population of C. cf. nasutum, the present species preferred a horizontal sleeping position (vs. head pointing downwards). When disturbed, some specimens dropped immediately and stayed curled and motionless on the ground. In appropriate habitat, specimens occurred a few metres from one another. In January 2016, only adult females were collected; in August 2016, additionally subadult and juvenile specimens; in November 2016, only one adult female was found. None of the collected females showed signs of being gravid. The size of the juveniles suggests that they hatched during the rainy season from approximately January to March. Under the occipital lobes of two specimens (ZSM 143/ 2016 and 142/2016) we found three and two mites, respectively; the lobes might function as mite pockets comparable to axillary pits in other chameleon species to limit and locate the damage caused by these ectoparasites (Arnold, 1986). Recommended IUCN status: As no attempts have yet been made to estimate the population size or status of C. juliae sp. nov. directly, we suggest that it should be assessed under the IUCN Red List criterion B (IUCN, 2012). The distribution area of the single known forest in which the species occurs has an area of ~ 0.15 km 2, which we interpret as the area of occupancy (AOO) of the species as defined by the IUCN (2012), but which at present is also equivalent to the EOO of the species (criterion B 1). This constitutes a single threat-defined location (criterion B, subcriterion a). The forest in which it occurs is under heavy active anthropogenic pressure and is, in our opinion, in imminent danger of disappearance [criterion B, subcriterion b(iii)]. As the AOO of the species is considerably C. boettgeri complex. Osteological measurements proved to be a useful tool to delimit these rather cryptic species. As a further important taxonomic character, C. gehringi (Fig. 10 A ��� C), C. guibei, and C. lefona sp. nov. have a distinct frontoparietal fenestra (FF; also pineal or parietal foramen of other authors; see Eakin, 1973) that differs in size between the species. This fenestra also occurs in other species of the C. nasutum group, e.g. C. fallax (Rieppel & Crumly, 1997) and C. cf. nasutum (Table 4). Compared with the elevational distribution of the species, a highly significant correlation [P = 3.8207 �� 10���6, r (Pearson) = 0.67] was found with presence/width of the fenestra and elevation (Fig. 10D). None of the species that live 1500 m a.s.l. have one. Additionally, FF also occur in C. peyrierasi and C. tsaratananense (Pr��tzel et al., 2018; A. van���t Padje et al., unpublished observations); both are montane species that occur> 1900 m a.s.l. (Brygoo, 1971, 1978), the latter of which is certainly not closely related to the C. nasutum group (Tolley et al., 2013). All other investigated Calumma species have a closed skull roof (Pr��tzel et al., 2018; van���t Padje et al., unpubl.)., Published as part of David Pr��tzel, Miguel Vences, Oliver Hawlitschek, Mark D. Scherz, Fanomezana M. Ratsoavina & Frank Glaw, 2018, Endangered beauties: micro-CT cranial osteology, molecular genetics and external morphology reveal three new species of chameleons in the Calumma boettgeri complex (Squamata: Chamaeleonidae), pp. 1-28 in Zoological Journal of the Linnean Society 2018 (20) on pages 15-20, DOI: 10.1093/zoolinnean/zlx112, http://zenodo.org/record/1226487, {"references":["Protzel D, Ruthensteiner B, Glaw F. 2016. No longer single! Description of female Calumma vatosoa (Squamata, Chamaeleonidae) including a review of the species and its systematic position. Zoosystematics and Evolution 92: 13 - 21.","Protzel D, Ruthensteiner B, Scherz MD, Glaw F. 2015. Systematic revision of the Malagasy chameleons Calumma boettgeri and C. linotum (Squamata: Chamaeleonidae). Zootaxa 4048: 211 - 231.","IUCN. 2012. IUCN red list categories and criteria: version 3.1. Gland, Switzerland and Cambridge, UK: IUCN.","Rieppel O, Crumly C. 1997. Paedomorphosis and skull structure in Malagasy chamaeleons (Reptilia: Chamaeleoninae). Journal of Zoology 243: 351 - 380.","Tolley KA, Townsend TM, Vences M. 2013. Large-scale phylogeny of chameleons suggests African origins and Eocene diversification. Proceedings of the Royal Society of London B: Biological Sciences, 280: 20130184."]}

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"]}

Morphology of Calumma boettgeri sensu lato External morphology. Measurements of important morphological parameters were taken from 23 specimens (11 males, 12 females) from Nosy Be, six specimens (five males, one female) from Montagne d’Ambre, and another nine specimens without exact locality data (Table 1). Because there was only one female from Montagne d’Ambre available, only males from both species were considered for comparison of body size, extremities and appendages. The specimens without localities were not included in mean value calculations but could be assigned according to their morphology to the Nosy Be morphotype (four specimens) and the Montagne d’Ambre morphotype (five specimens), respectively. , ratio of tail to snout-vent length; LRA, length of rostral appendage from snout tip; RRS, ratio of length of rostral appendage and snout-vent length; RAPSC, number of peripheral scales on rostral appendage; NPSCM, number of peripheral scales per mm on rostral appendage; RC, rostral crest present (+) or absent (-); NSL, number of supralabials; NIL, number of infralabials; OLD, lateral diameter of the occipital lobe; OLN, depth of the dorsal notch in occipital lobe; PC, parietal crest absent (-) or number of parietal cones; DC, dorsal crest absent (-) or number of dorsal cones; AP, axillary pits present (+) or absent (-); DSC, diameter of broadest scale on upper arm; RSB, ratio of broadest scale to snout-vent length; NSC, number of big scales on upper arm from lateral view; UAD, upper arm diameter; RAS, ratio of arm diameter to snout-vent length; all measurements in mm. ……continued on the next page …….continued on the next page Continued. collection no. species locality sex NIL OLD OLN PC DC AP DSC RSB NSC UAD RAS ……continued on the next page collection no. species locality sex NIL OLD OLN PC DC AP DSC RSB NSC UAD RAS SMF 16471 C. boettgeri Nosy Be f 10 4.2 0.0 - 0 - 0.4 0.007 9 2.2 0.044 SMF 16472 C. boettgeri Nosy Be f 13 3.5 0.1 - 11 - 0.4 0.009 10 1.8 0.039 ZSM 865 / 1920 C. boettgeri - f 13 2.8 0.1 - 0 - 0.3 0.006 9 1.9 0.038 ZFMK 51515 C. boettgeri - f 13 3.3 0.7 - 0 - 0.3 0.006 12 2.1 0.041 MCZ 5988 Ch. macrorhinus ‘Madagascar’ f 6.1 3.2 0.1 - 0 - 0.3 0.007 8 2.3 0.047 Individuals from Montagne d’Ambre show clear morphological differences from Nosy Be specimens (Fig. 7). The adult males examined from Montagne d’Ambre are larger than those from Nosy Be (mean values of TL 107.8 mm in Montagne d’Ambre vs. 99.3 mm in Nosy Be, Table 2), their rostral appendage is longer related to the snoutvent length (RRS 0.078 vs. 0.065) and the ratio of arm diameter to snout-vent length is larger (UAD /SVL 0.053 vs. 0.042). species C. boettgeri C. boettgeri C. linotum locality Nosy Be Nosy Be M. d'Ambre sex m f m Both populations show heterogeneous scalation, especially at the extremities and the rostral appendage. The enlarged rounded tubercles on the limbs are distinctly larger in animals from Montagne d’Ambre; the mean diameter of the largest tubercle of the upper arm is 0.64 mm compared to 0.36 mm (Nosy Be). Additionally the number of the enlarged tubercles on the upper arm of males from Montagne d’Ambre is approximately twice that of males from Nosy Be, with a mean of 20.2 compared to 9.9, respectively, and the tubercles are not bordering each other on Nosy Be individuals. This character is also confirmed from a juvenile of SVL 26.9 mm (ZFMK 48227, Nosy Be) and a juvenile of SVL 36.2 mm (ZFMK 52308, Joffreville). Another morphological difference between both populations can be found in the pholidosis of the rostral appendage. Although the rostral appendage is significantly smaller in males from Nosy Be (see above), they have more peripheral scales on it, with a mean of 18.9 compared to 15.5 in Montagne d’Ambre males. In relation to the size of the appendage, this means 6.1 scales per mm compared with 4.1 scales per mm in Montagne d’Ambre. In summary, specimens from Montagne d’Ambre show a more heterogeneous scalation with broader tubercles on extremities and the rostral appendage. This is true of females as well (note that only one female with locality data was examined). Montagne d’Ambre individuals differ also in both sexes from Nosy Be specimens in the presence of a parietal crest which is best visible in the micro-CT scan (see below). The other morphological features either were highly variable or did not differ between the populations. For example, the number of dorsal cones was 0–28 in Nosy Be, and 9–13 in Montagne d’Ambre. Likewise the edges of the occipital lobes vary between the specimens. They range between un-notched and clearly notched (up to 0.7 mm) in Nosy Be, and are either not or only slightly notched (up to 0.3 mm) in Montagne d’Ambre. All specimens have a rostral crest and none have axillary pits. Colouration. Males also show great differences between the populations in colouration (see Fig. 2 A, B). Males from Montagne d’Ambre were more colourful, with a true blue rostral appendage and greenish turquoise extremities (Fig. 2 A). The colour of the legs is induced only by the coloured tubercle scales. The body is pale green or light brown with two dark brown spots and (occasionally) a beige lateral stripe on each side that stops at the base of the tail. The tail is the same colour as the body and (in stress colouration) possesses black annulations. The head is also greenish or brown with a dark stripe from the snout crossing the eyes to the occipital lobes. The skin around the mouth and the throat can be white. The colour description is based on a total of seven pictures of the Montagne d’Ambre form, referred to as C. boettgeri in Schmidt et al. (2010), C. boettgeri (picture 1 c) in Glaw & Vences (2007) Calumma sp. in Nečas (2004) and C. boettgeri in Garbutt et al. (2001). The body and head of females is brown; the rostral appendage can be coloured bright blue, see picture of C. boettgeri in Schmidt et al. (2010). The body colouration of males from Nosy Be in contrast is yellowish or greenish brown with little dark brown rosettes, when stressed (Fig. 2 B). The legs are brown with little blue or green spots resulting from the tubercles. The colouration of the head is similar to the body colouration. The rostral appendage differs clearly from Montagne d’Ambre with the absence of any striking colour and is the same brown colour as the casque. Females are uniformly light or greenish brown coloured. Compare also a total of seven pictures of Calumma boettgeri in Hyde Roberts & Daly (2014), in Glaw & Vences (2007: 191) picture 1 a and 1 b, in Nečas (2004) and in Henkel & Schmidt (1995). Osteology of the skull based on micro-CT scans. Micro-CT scans of heads of two males and two females from Nosy Be and from Montagne d’Ambre exposed additional differences between the two forms. Specimens from Montagne d’Ambre (Fig. 4, D and E) bear tubercles on the parietal in both sexes. These form a little parietal crest in the middle with three to four tubercles, laterally followed by two tubercles on each side. The frontal is also irregularly spotted with tubercles. The parietal and frontal of animals from Nosy Be in contrast are smooth (Fig. 4, A and B). As in all species of the genus Calumma, the nasal bones are paired (Rieppel & Crumly 1997). These are broader in our specimens from Montagne d’Ambre (mean NW 0.35 mm vs. 0.24 mm in Nosy Be; mean NW/NL 0.18 vs. 0.14, Table 3, Fig. 4) and the anterior tip of the frontal bone does not exceed more than a half of the naris. In skulls from Nosy Be it does exceed this point, and the frontal meets the premaxilla, as described for C. nasutum (Rieppel & Crumly 1997). The parietal also varies between the two localities. In Nosy Be samples, the parietal tapers more tightly. Its diameter is at the tightest area on average 0.61 mm (vs. 1.06 mm) and 11 % of the largest diameter of the parietal (vs. 22 %, Table 3, Fig. 4). The parietal in Montagne d’Ambre samples appears wider and more compact. However, the form of the parietal is variable within localities and cannot be used as a diagnostic character. Although chameleons are sexually dimorphic animals, differences between sexes in skull structure were not proven (Table 3). Notes: m, male; f, female; NL, nasal length; NW, nasal width; RNWL, ratio of nasal width to length; PL, largest diameter; PS, parietal smallest diameter; RPSL, ratio of parietal smallest to largest diameter; PC, parietal crest absent (-) or number of tubercles. Hemipenial morphology based on micro-CT scans. The scans of hemipenes of specimens from each population enable a detailed view of their structure. The hemipenes are illustrated in sulcal and asulcal view with the apex on top (Fig. 5). Both populations show large and deep calyces with smooth ridges on the asulcal side of the truncus. The apex is ornamented with two pairs of long pointed papillae and two pairs of rotulae. The papillae rise from the sulcal side of the apex and are curved to the asulcal side. They can be completely everted (Fig. 5, C) or retracted in the apex (Fig. 5, D). One pair of rotulae is placed on the asulcal side (the smaller one) and one pair on the sulcal side. Here some differences between the populations are recognizable; in Nosy Be (n = 2) the rotulae are slightly more denticulated, with 6–11 tips on asulcal side and 14–16 tips on sulcal side, compared to Montagne d’Ambre (n = 3) with 6–8 tips on the rotulae of the asulcal side and 11–14 tips on both rotulae on the sulcal side.

Identity of Calumma boettgeri (Boulenger, 1888) Morphological measurements and pholidosis of the holotype require the assignment of Chamaeleo macrorhinus to the Nosy Be form of Calumma boettgeri. The measurements are similar to the mean values of the females from Nosy Be (Tables 1���2): diameter of the broadest tubercle on the upper arm 0.34 mm (vs. mean of Nosy Be females of 0.32 mm, SD 0.048), number of enlarged tubercles on the upper arm 8 (vs. 10.6, SD 1.8); ratio of the upper arm diameter to the body size, 0.046 (vs. 0.041, SD 0.0038), total length, 93.1 mm (vs. 96.3 mm, SD 8.1 mm) and length of the rostral appendage, 2.8 mm (vs. 2.7 mm, SD 0.5 mm). Diagnosis. A small-sized chameleon (SVL 41.1���55.5 mm, TL 83.8 ���108.0 mm) that is characterised by a soft dermal, distally rounded, typically brown rostral appendage, slightly notched occipital lobes, the absence of a parietal crest, a low casque, the absence of axillary pits, small rounded tubercles not bordering each other on the extremities, presence or absence of a dorsal crest in males, low casque, and absence of gular and ventral crests. It differs from C. guibei by unnotched or only slightly (max. 0.7 mm) notched versus completely notched occipital lobes. For a distinction from C. linotum, see above. Colouration in life. The body and head colouration of males in relaxed state ranges from light brown to yellow without any obvious colour patterns. When stressed, dark colour patterns become prominent and the tail becomes annulated. A dark line runs from the snout tip across the eyes to the occipital lobes. The skin around the mouth and the throat can be white. Remarkable is the inconspicuous brown colour of the rostral appendage. The extremities appear brown also, except for a few green or blue coloured tubercle scales (Fig. 2 B). The colouration of the females can vary from beige to a reddish or greenish brown ground colouration in a relaxed state. The rostral appendage and the extremities show the same colour as the body, except for a few green tubercle scales on the legs. In a stressed state, three parallel bright blue spots appear on the upper half of the eyelids. Distribution. All confirmed distribution records of C. boettgeri are confined to the biogeographic Sambirano region in northwest Madagascar. It was found in both primary rainforest (Lokobe) and secondary forests (near Andoany) of Nosy Be (Andreone et al. 2003), in Manongarivo (Rakotomalala 2002; Gehring et al. 2012), and on Nosy Komba (Hyde Roberts & Daly 2014). Additionally Nagy et al. (2012, suppl. Fig. 2) identified a population with similar gene sequences to C. boettgeri from a forest fragment locally known as Maromiandra (13 �� 99 ��� 65 ������S, 48 �� 21 ��� 77 ������E, 283 m). According to molecular phylogenetic data, C. boettgeri from Nosy Be and one individual from Manongarivo (FGMV 2002 -813, 13�� 58 ��� 62 ������S, 48 �� 25 ��� 32 ������E, 751 m a.s.l.) form their own clade (Gehring et al. 2012). The elevations of all these localities range from 0 to 751 m a.s.l. (Fig. 6). We consider all additional records of C. boettgeri, all located in northeastern Madagascar, as in need of confirmation: Ambolokopatrika, 810���860 m a.s.l. (Andreone et al. 2000), Antalaha (Brygoo 1971), Andrakaraka forest station ca. 10 km from Antalaha (Ramanantsoa 1974), and Marojejy, 1100���1200 m a.s.l. (Raselimanana et al. 2000)., Published as part of Pr��tzel, David, Ruthensteiner, Bernhard, Scherz, Mark D. & Glaw, Frank, 2015, Systematic revision of the Malagasy chameleons Calumma boettgeri and C. linotum (Squamata: Chamaeleonidae), pp. 211-231 in Zootaxa 4048 (2) on pages 226-227, DOI: 10.11646/zootaxa.4048.2.4, http://zenodo.org/record/233651, {"references":["Boulenger, G. A. (1888) Descriptions of two new chamaeleons from Nossi-Be, Madagascar. Annales and Magazine of Natural History, Series 6, 1 (1), 22 - 23. http: // dx. doi. org / 10.1080 / 00222938809460666","Andreone, F., Glaw, F., Nussbaum, R. A., Raxworthy, C. J., Vences, M., Randrianirina, J. E. (2003) The amphibians and reptiles of Nosy Be (NW Madagascar) and nearby islands: a case study of diversity and conservation of an insular fauna. Journal of Natural History, 37 (17), 2119 - 2149. http: // dx. doi. org / 10.1080 / 00222930210130357","Rakotomalala, D. (2002) Diversite des reptiles et amphibiens de la Reserve Speciale de Manongarivo, Madagascar. Boissiera, 59, 339 - 358.","Hyde Roberts, S. & Daly, C. (2014) A rapid herpetofaunal assessment of Nosy Komba Island, northwestern Madagascar, with new locality records for seventeen species. Salamandra, 50, 18 - 26.","Nagy, Z. T., Sonet, G., Glaw, F. & Vences, M. (2012) First large-scale DNA barcoding assessment of reptiles in the biodiversity hotspot of Madagascar, based on newly designed COI primers. PLoS ONE, 7 (3), e 34506. http: // dx. doi. org / 10.1371 / journal. pone. 0034506","Andreone, F., Randrianirina, J. E., Jenkins, P. D. & Aprea, G. (2000) Species diversity of Amphibia, Reptilia and Lipotyphla (Mammalia) at Ambolokopatrika, a rainforest between the Anjanaharibe-Sud and Marojejy massifs, NE Madagascar. Biodiversity and Conservation, 9, 1587 - 1622. http: // dx. doi. org / 10.1023 / A: 1026559728808","Brygoo, E. R. (1971) Reptiles Sauriens Chamaeleonidae - genre Chamaeleo. Faune de Madagascar, 33, 1 - 318.","Ramanantsoa, G. - A. (1974) Connaissance des cameleonides commun de la province de Diego Suarez par la population paysanne. Bulletin de l'Academie Malgache, 51, 147 - 149.","Raselimanana, A. P., Raxworthy, C. J. & Nussbaum, R. A. (2000) Herpetofaunal species diversity and elevational distribution within the Parc National de Marojejy, Madagascar. Fieldiana Zoology, 157 - 174."]}

Morphology of Calumma boettgeri sensu lato External morphology. Measurements of important morphological parameters were taken from 23 specimens (11 males, 12 females) from Nosy Be, six specimens (five males, one female) from Montagne d���Ambre, and another nine specimens without exact locality data (Table 1). Because there was only one female from Montagne d���Ambre available, only males from both species were considered for comparison of body size, extremities and appendages. The specimens without localities were not included in mean value calculations but could be assigned according to their morphology to the Nosy Be morphotype (four specimens) and the Montagne d���Ambre morphotype (five specimens), respectively. , ratio of tail to snout-vent length; LRA, length of rostral appendage from snout tip; RRS, ratio of length of rostral appendage and snout-vent length; RAPSC, number of peripheral scales on rostral appendage; NPSCM, number of peripheral scales per mm on rostral appendage; RC, rostral crest present (+) or absent (-); NSL, number of supralabials; NIL, number of infralabials; OLD, lateral diameter of the occipital lobe; OLN, depth of the dorsal notch in occipital lobe; PC, parietal crest absent (-) or number of parietal cones; DC, dorsal crest absent (-) or number of dorsal cones; AP, axillary pits present (+) or absent (-); DSC, diameter of broadest scale on upper arm; RSB, ratio of broadest scale to snout-vent length; NSC, number of big scales on upper arm from lateral view; UAD, upper arm diameter; RAS, ratio of arm diameter to snout-vent length; all measurements in mm. ������continued on the next page ������.continued on the next page Continued. collection no. species locality sex NIL OLD OLN PC DC AP DSC RSB NSC UAD RAS ������continued on the next page collection no. species locality sex NIL OLD OLN PC DC AP DSC RSB NSC UAD RAS SMF 16471 C. boettgeri Nosy Be f 10 4.2 0.0 - 0 - 0.4 0.007 9 2.2 0.044 SMF 16472 C. boettgeri Nosy Be f 13 3.5 0.1 - 11 - 0.4 0.009 10 1.8 0.039 ZSM 865 / 1920 C. boettgeri - f 13 2.8 0.1 - 0 - 0.3 0.006 9 1.9 0.038 ZFMK 51515 C. boettgeri - f 13 3.3 0.7 - 0 - 0.3 0.006 12 2.1 0.041 MCZ 5988 Ch. macrorhinus ���Madagascar��� f 6.1 3.2 0.1 - 0 - 0.3 0.007 8 2.3 0.047 Individuals from Montagne d���Ambre show clear morphological differences from Nosy Be specimens (Fig. 7). The adult males examined from Montagne d���Ambre are larger than those from Nosy Be (mean values of TL 107.8 mm in Montagne d���Ambre vs. 99.3 mm in Nosy Be, Table 2), their rostral appendage is longer related to the snoutvent length (RRS 0.078 vs. 0.065) and the ratio of arm diameter to snout-vent length is larger (UAD /SVL 0.053 vs. 0.042). species C. boettgeri C. boettgeri C. linotum locality Nosy Be Nosy Be M. d'Ambre sex m f m Both populations show heterogeneous scalation, especially at the extremities and the rostral appendage. The enlarged rounded tubercles on the limbs are distinctly larger in animals from Montagne d���Ambre; the mean diameter of the largest tubercle of the upper arm is 0.64 mm compared to 0.36 mm (Nosy Be). Additionally the number of the enlarged tubercles on the upper arm of males from Montagne d���Ambre is approximately twice that of males from Nosy Be, with a mean of 20.2 compared to 9.9, respectively, and the tubercles are not bordering each other on Nosy Be individuals. This character is also confirmed from a juvenile of SVL 26.9 mm (ZFMK 48227, Nosy Be) and a juvenile of SVL 36.2 mm (ZFMK 52308, Joffreville). Another morphological difference between both populations can be found in the pholidosis of the rostral appendage. Although the rostral appendage is significantly smaller in males from Nosy Be (see above), they have more peripheral scales on it, with a mean of 18.9 compared to 15.5 in Montagne d���Ambre males. In relation to the size of the appendage, this means 6.1 scales per mm compared with 4.1 scales per mm in Montagne d���Ambre. In summary, specimens from Montagne d���Ambre show a more heterogeneous scalation with broader tubercles on extremities and the rostral appendage. This is true of females as well (note that only one female with locality data was examined). Montagne d���Ambre individuals differ also in both sexes from Nosy Be specimens in the presence of a parietal crest which is best visible in the micro-CT scan (see below). The other morphological features either were highly variable or did not differ between the populations. For example, the number of dorsal cones was 0���28 in Nosy Be, and 9���13 in Montagne d���Ambre. Likewise the edges of the occipital lobes vary between the specimens. They range between un-notched and clearly notched (up to 0.7 mm) in Nosy Be, and are either not or only slightly notched (up to 0.3 mm) in Montagne d���Ambre. All specimens have a rostral crest and none have axillary pits. Colouration. Males also show great differences between the populations in colouration (see Fig. 2 A, B). Males from Montagne d���Ambre were more colourful, with a true blue rostral appendage and greenish turquoise extremities (Fig. 2 A). The colour of the legs is induced only by the coloured tubercle scales. The body is pale green or light brown with two dark brown spots and (occasionally) a beige lateral stripe on each side that stops at the base of the tail. The tail is the same colour as the body and (in stress colouration) possesses black annulations. The head is also greenish or brown with a dark stripe from the snout crossing the eyes to the occipital lobes. The skin around the mouth and the throat can be white. The colour description is based on a total of seven pictures of the Montagne d���Ambre form, referred to as C. boettgeri in Schmidt et al. (2010), C. boettgeri (picture 1 c) in Glaw & Vences (2007) Calumma sp. in Nečas (2004) and C. boettgeri in Garbutt et al. (2001). The body and head of females is brown; the rostral appendage can be coloured bright blue, see picture of C. boettgeri in Schmidt et al. (2010). The body colouration of males from Nosy Be in contrast is yellowish or greenish brown with little dark brown rosettes, when stressed (Fig. 2 B). The legs are brown with little blue or green spots resulting from the tubercles. The colouration of the head is similar to the body colouration. The rostral appendage differs clearly from Montagne d���Ambre with the absence of any striking colour and is the same brown colour as the casque. Females are uniformly light or greenish brown coloured. Compare also a total of seven pictures of Calumma boettgeri in Hyde Roberts & Daly (2014), in Glaw & Vences (2007: 191) picture 1 a and 1 b, in Nečas (2004) and in Henkel & Schmidt (1995). Osteology of the skull based on micro-CT scans. Micro-CT scans of heads of two males and two females from Nosy Be and from Montagne d���Ambre exposed additional differences between the two forms. Specimens from Montagne d���Ambre (Fig. 4, D and E) bear tubercles on the parietal in both sexes. These form a little parietal crest in the middle with three to four tubercles, laterally followed by two tubercles on each side. The frontal is also irregularly spotted with tubercles. The parietal and frontal of animals from Nosy Be in contrast are smooth (Fig. 4, A and B). As in all species of the genus Calumma, the nasal bones are paired (Rieppel & Crumly 1997). These are broader in our specimens from Montagne d���Ambre (mean NW 0.35 mm vs. 0.24 mm in Nosy Be; mean NW/NL 0.18 vs. 0.14, Table 3, Fig. 4) and the anterior tip of the frontal bone does not exceed more than a half of the naris. In skulls from Nosy Be it does exceed this point, and the frontal meets the premaxilla, as described for C. nasutum (Rieppel & Crumly 1997). The parietal also varies between the two localities. In Nosy Be samples, the parietal tapers more tightly. Its diameter is at the tightest area on average 0.61 mm (vs. 1.06 mm) and 11 % of the largest diameter of the parietal (vs. 22 %, Table 3, Fig. 4). The parietal in Montagne d���Ambre samples appears wider and more compact. However, the form of the parietal is variable within localities and cannot be used as a diagnostic character. Although chameleons are sexually dimorphic animals, differences between sexes in skull structure were not proven (Table 3). Notes: m, male; f, female; NL, nasal length; NW, nasal width; RNWL, ratio of nasal width to length; PL, largest diameter; PS, parietal smallest diameter; RPSL, ratio of parietal smallest to largest diameter; PC, parietal crest absent (-) or number of tubercles. Hemipenial morphology based on micro-CT scans. The scans of hemipenes of specimens from each population enable a detailed view of their structure. The hemipenes are illustrated in sulcal and asulcal view with the apex on top (Fig. 5). Both populations show large and deep calyces with smooth ridges on the asulcal side of the truncus. The apex is ornamented with two pairs of long pointed papillae and two pairs of rotulae. The papillae rise from the sulcal side of the apex and are curved to the asulcal side. They can be completely everted (Fig. 5, C) or retracted in the apex (Fig. 5, D). One pair of rotulae is placed on the asulcal side (the smaller one) and one pair on the sulcal side. Here some differences between the populations are recognizable; in Nosy Be (n = 2) the rotulae are slightly more denticulated, with 6���11 tips on asulcal side and 14���16 tips on sulcal side, compared to Montagne d���Ambre (n = 3) with 6���8 tips on the rotulae of the asulcal side and 11���14 tips on both rotulae on the sulcal side., Published as part of Pr��tzel, David, Ruthensteiner, Bernhard, Scherz, Mark D. & Glaw, Frank, 2015, Systematic revision of the Malagasy chameleons Calumma boettgeri and C. linotum (Squamata: Chamaeleonidae), pp. 211-231 in Zootaxa 4048 (2) on pages 214-221, DOI: 10.11646/zootaxa.4048.2.4, http://zenodo.org/record/233651, {"references":["Schmidt, W., Tamm, K. & Wallikewitz, E. (2010) Chamaleons - Drachen unserer Zeit. Natur- und Tier-Verlag, Munster, 333 pp.","Glaw, F. & Vences, M. (2007) A field guide to the amphibians and reptiles of Madagascar. Vences and Glaw Verlag, Cologne, 496 pp.","Necas, P. (2004) Chamaleons - bunte Juwelen der Natur. Edition Chimaira, Frankfurt am Main, 338 pp.","Garbutt, N., Bradt, H. & Schuurman, D. (2001) Madagascar wildlife - a visitor's guide. Bradt Travel Guides, Bucks, 138 pp.","Hyde Roberts, S. & Daly, C. (2014) A rapid herpetofaunal assessment of Nosy Komba Island, northwestern Madagascar, with new locality records for seventeen species. Salamandra, 50, 18 - 26.","Henkel, F. W. & Schmidt, W. (1995) Amphibien und Reptilien Madagaskars, der Maskarenen, Seychellen und Komoren. Eugen Ulmer Verlag, Stuttgart, 311 pp.","Rieppel, O. & Crumly, C. (1997) Paedomorphosis and skull structure in Malagasy chamaeleons (Reptilia: Chamaeleonidae). Journal of Zoology, 243 (2), 351 - 380. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1997. tb 02788. x"]}

Prötzel, David, Ruthensteiner, Bernhard, Scherz, Mark D., Glaw, Frank (2015): Systematic revision of the Malagasy chameleons Calumma boettgeri and C. linotum (Squamata: Chamaeleonidae). Zootaxa 4048 (2): 211-231, DOI: http://dx.doi.org/10.11646/zootaxa.4048.2.4