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4.3. Lygodactylus verticillatus Lygodactylus verticillatus from Madagascar and from Europa Island do not differ in scalation (Boettger 1913; Pasteur 1965). The slight difference in the number of precloacal pores mentioned by Pasteur (1965) was not supported by this study. Both populations show a very similar variation in coloration and in patterning. Thus, L. verticillatus from Madagascar and Europa cannot be distinguished by their external appearance or by the number of precloacal pores. The most basal node within L. verticillatus separates specimens from the southern Malagasy localities Sakaraha and Isalo from all other samples. Sakaraha lies approximately 135 km from Toliara on the main road to the Isalo National Park. The remaining sister subgroup consists of specimens exclusively from the coastal regions of Madagascar, approximately from Morondava southwards to Toliara, and the insular population. We cannot exclude an inland origin of the L. verticillatus from Europa Island. However, based on our morphological and genetic dataset, it seems more likely that the nearest relatives of the insular L. verticillatus are conspecifics from populations in the coastal regions of south-western Madagascar. We found no relevant morphological differences, and only weak genetic differences between the Malagasy and the insular population, and only weak genetic variation within the monophyletic lineage of Europa Island. This agrees with the hypothesis of a relatively recent colonization as suggested by Pasteur (1965). Europa Island is 355 km west-northwest from Toliara or 300 km southwest from Cap Saint-Vincent close to Morombe, 529 km east-northeast from Inhambane (Mozambique) and approximately 600 km south of Juan de Nova (Caceres 2003; Fricke et al. 2013). It is nearly circular (6 × 7 km) with a total area of 30 km 2 (Caceres 2003) and represents the largest island of the Îles Éparses. Europa Island belongs to the group of ‘modern’ isolated seamounts in the Channel, possibly developed during Oligocene to Miocene times (Courgeon et al. 2016). Lygodactylus verticillatus could have reached Europa Island recently by natural transoceanic dispersal or by human-mediated dispersal. While a natural dispersal would be supported by flows westward from Madagascar to mainland Africa (Ali & Huber 2010), a much simpler and thus more parsimonious alternative explanation is ship-borne dispersal. Lygodactylus verticillatus could have arrived at Europa Island e.g., with ships of Malagasy fishermen seasonally searching for nesting chelonians, or with ships of European (French) settlers who started their settlement from Toliara and lived on the island from approximately 1860 to the 1920s (Fricke et al. 2013). The amount of genetic variation found within Europa (three haplotypes differing by up to two mutations in 16S) would be in agreement with either (i) ship-borne dispersal or (ii) natural dispersal of multiple individuals, or (iii) natural dispersal of a single individual at a somewhat deeper point in time, with subsequent in-situ genetic diversification. Support for either of these hypotheses could come from a denser sampling of Malagasy populations: the presence of haplotypes identical to those on Europa in Malagasy populations would allow to reject the third hypothesis, and given that simultaneous or repeated natural dispersal to such a small islet is rather unlikely, ship-borne introduction would in this case remain as the most probable hypothesis. Only if the presence of the Europa haplotypes in Madagascar could be reasonably excluded, the third hypothesis would be supported. Lygodactylus verticillatus is adapted to dry climatic conditions. In south-western Madagascar, the species inhabits shrubs and trees in semi-arid areas as well as fences and walls in human settlements (Puente et al. 2009). On Europa Island, it is widely distributed and occurs in most natural habitats on rocks, shrubs, trees and in a dry euphorbia forest as well as in human settlements (Brygoo 1966; Sanchez et al. 2015, 2019). The conditions for L. verticillatus in the coastal regions of south-western Madagascar and on Europa are very similar, facilitating the survival on the island., Published as part of Röll, Beate, Sanchez, Mickaël, Gippner, Sven, Bauer, Aaron M., Travers, Scott L., Glaw, Frank, Hawlitschek, Oliver & Vences, Miguel, 2023, Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean, pp. 232-250 in Zootaxa 5311 (2) on pages 245-246, DOI: 10.11646/zootaxa.5311.2.4, http://zenodo.org/record/8094276, {"references":["Boettger, O. (1913) Reptilien und Amphibien von Madagascar, den Inseln und dem Festland Ostfarikas. In: Voeltzkow, A. (Ed.), Reise in Ostafrika in den Jahren 1903 - 1905. Wissenschaftliche Ergebnisse. Vol. 3. Systematische Arbeiten. Schweizerbartsche Verlagsbuchhandlung, Nagele und Sprosser, Stuttgart, pp. 269 - 375.","Pasteur, G. (1965) Recherches sur l'evolution des lygodactyles, lezards afromalgaches actuels. Travaux de l'Institut Scientifique Cherifien, Serie Zoologie, 29, 1 - 132.","Caceres, S. (2003) Etude prealable pour le classement en Reserve Naturelle des Iles Eparses. Memoire de DESS Sciences et Gestion de l'Environnement Tropical. Univ. Reunion (Laboratoire ECOMAR) & DIREN Reunion, 136 pp. Available from: http: // poupin. joseph. free. fr / pdf / caceres- 2003 - iles-eparses-classement. pdf (accessed 13 June 2023)","Fricke, R., Durville, P., Bernardi, G., Borsa, P., Mou-Tham, G. & Chabanet, P. (2013) Checklist of the shore fishes of Europa Island, Mozambique Channel, southwestern Indian Ocean, including 302 new records. Stuttgarter Beitrage zur Naturkunde, A, Neue Serie, 6, 247 - 276.","Courgeon, S., Jorry, S. J., Camoin, G. F., BouDagher-Fadel, M. K., Jouet, G., Revillon, S., Bachelery, P., Pelleter, E., Borgomano, J., Poli, E. & Droxler, A. W., (2016) Growth and demise of Cenozoic isolated carbonate platforms: New insights from the Mozambique Channel seamounts (SW Indian Ocean). Marine Geology, 380, 90 - 105. https: // doi. org / 10.1016 / j. margeo. 2016.07.006","Ali, J. R. & Huber, M. (2010) Mammalian biodiversity on Madagascar controlled by ocean currents. Nature, 463, 653 - 656. https: // doi. org / 10.1038 / nature 08706","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","Brygoo, E. R. (1966) Note sur les reptiles terrestres recoltes a Europa en avril 1964. Memoires du Museum national d'histoire naturelle, Serie A, 41, 29 - 32.","Sanchez, M. & Probst, J. - M. (2015) L'herpetofaune terrestre de l'ile d'Europa (Ocean Indien, Canal du Mozambique): synthese des connaissances et nouvelles donnees sur la repartition de l'ecologie des especes en vue de leur conservation. Bulletin de la Societe Herpetologique de France, 145, 63 - 76."]}

4.1. Lygodactylus pakenhami At present, the taxon ‘ pakenhami’ is considered a subspecies of L. grotei in most publications. In our study, L. grotei and L. g. pakenhami are genetically distinct lineages, confirming previous results (Gippner et al. 2021). Lygodactylus g. pakenhami from Pemba Island differs greatly in 16S and ND2 pairwise distances from mainland individuals considered as L. grotei. While L. grotei and L. g. pakenhami as far as known cannot be distinguished in scalation, they clearly differ in coloration. Furthermore, hatchlings of L. grotei and L. g. pakenhami are easily identifiable and distinguishable by the coloration of their trunks and especially of their tails. The reciprocal monophyly suggested by mitochondrial DNA, conspicuously different coloration—both in adults and hatchlings—and the high genetic distance suggest that these taxa most likely represent two distinct species. This is also in agreement with differences in the RAG1 sequences (although these data are not fully conclusive due to low sample sizes: one sequence available each for grotei and pakenhami). Therefore, we elevate L. g. pakenhami to full species status, as Lygodactylus pakenhami Loveridge, 1941, being aware that this taxonomic hypothesis requires further testing, especially from nuclear-encoded DNA data sets, ideally at the phylogenomic level. Lygodactylus pakenhami is endemic on Pemba Island, the northernmost and second largest island of the Zanzibar Archipelago. Pemba lies off the continental shelf and is surrounded by water 500 to 850 m deep (Moreau & Pakenham 1941). Geological evidence indicates that Pemba Island was separated from the African mainland by faulting that produced the Pemba Channel, possibly during the late Miocene or early Pliocene, 6 million years ago (Stockley 1942; Clarke & Burgess 2000). Corresponding to its long period of isolation, Pemba is characterized by a remarkable number of endemic species, including plants, mammals and reptiles (R̂dder et al. 2010). The simplest explanation for the existence of L. pakenhami on Pemba is vicariance. Its nearest extant relative, L. grotei, is widely distributed in south-eastern Tanzania and northern Mozambique, including the coastal regions. Presumably, a population of an ancestor of L. grotei and L. pakenhami already existed on Pemba Island before its separation from the mainland. Thereafter, there was no further genetic exchange between the continental and the insular population, so that the latter evolved isolated from its continental congener. While a natural dispersal from Tanzania after the separation of Pemba Island cannot be excluded, it is considered as unlikely because the geographic position of Pemba suggests that the East African Coastal Current or a similar paleocurrent conveyed any drifting material into the open ocean very rapidly (Moreau & Pakenham 1940; Hawlitschek et al. 2016a). A recent, human-mediated transportation is unlikely, as after such a short period of isolation we would expect the sharing of mitochondrial haplotypes between mainland and island populations, which we did not detect. Although only two samples yielding DNA sequences of L. grotei have precise georeferenced information, one of these was collected on the mainland directly opposite Pemba Island. However, as is always the case with inferences of allopatric occurrence, we cannot fully exclude that undiscovered populations of L. pakenhami may occur on the African mainland, which would invalidate our biogeographic hypotheses but not our taxonomic conclusions. In contrast to Pemba Island, Zanzibar and Mafia islands lie on the continental shelf and are separated from the mainland only by shallow waters with an average depth of 30 to 35 m, rarely 50 m (Moreau & Pakenham 1941). Geological data of coastal eastern Africa point to a land connection of Zanzibar and Mafia islands with the African mainland up to the end of the Pleistocene, probably only 10,000 –18,000 years ago (Stockley 1942; Clarke & Burgess 2000). Zanzibar and Mafia are inhabited by L. grotei, L. picturatus and L. viscatus, all conspecific with continental populations. Presumably, the distribution of these three species on the islands is probably best explained by recent vicariant events. In contrast to L. pakenhami on Pemba Island, the species on Zanzibar and Mafia have been isolated on the islands for a relatively short time, about 10,000 years. Additionally, both natural dispersal over water after the separation of Zanzibar and Mafia from the mainland and/or a human-mediated transportation due to long-standing trading in the recent past cannot be excluded., Published as part of Röll, Beate, Sanchez, Mickaël, Gippner, Sven, Bauer, Aaron M., Travers, Scott L., Glaw, Frank, Hawlitschek, Oliver & Vences, Miguel, 2023, Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean, pp. 232-250 in Zootaxa 5311 (2) on page 244, DOI: 10.11646/zootaxa.5311.2.4, http://zenodo.org/record/8094276, {"references":["Gippner, S., Travers, S. L., Scherz, M. D., Colston, T. J., Lyra, M. L., Ashwini, V. M., Multzsch, M., Nielson, S. V., Rancilhac, L., Glaw, F., Bauer, A. M. & 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","Loveridge, A. (1941) New geckos (Phelsuma and Lygodactylus), snake (Lepotyphlops) and frog (Phrynobarachits) from Pemba Island, East Africa. Proceedings of the Biological Society, Washington, 54 (8), 175 - 178.","Stockley, G. M. (1942) The geology of the Zanzibar Protectorate and its relation to the East African mainland. Geological Magazine, 79 (4), 233 - 240. https: // doi. org / 10.1017 / S 0016756800073921","Clarke, G. P. & Burgess, N. D. (2000) Geology and geomorphology. In: Burgess, N. D. & Clarke, G. P. (Eds.), Coastal Forests of Eastern Africa. IUCN, Gland, pp. 29 - 40.","Moreau, R. E. & Pakenham, R. H. W. (1940) The land vertebrates of Pemba, Zanzibar, and Mafia: a zoogeographical study. Proceedings of the Zoological Society, London, 110 (A), 97 - 128. https: // doi. org / 10.1111 / j. 1469 - 7998.1941. tb 08463. x","Hawlitschek, O., Garrido, S. R. & Glaw, G. (2016 a). How marine currents influenced the widespread natural overseas dispersal of reptiles in the Western Indian Ocean. Journal of Biogeography, 44, 1426 - 1440. https: // doi. org / 10.1111 / jbi. 12940"]}

4.2. Lygodactylus insularis Lygodactylus insularis is an endemic species on Juan de Nova, a very small island with a total area of approximately 5 km 2, 280 km from mainland Africa (East Mozambique) and 120 km to the nearest point on the Malagasy coast (Caceres 2003; Lambs et al. 2016). Pasteur (1965) already included L. insularis in the L. capensis group and assumed that L. insularis originated from either L. c. capensis or L. c. grotei. Lygodactylus capensis and L. insularis share the same number of precloacal pores and a superficially similar coloration of the adults. However, they strongly differ in the status of the subcaudals and in the appearance of their hatchlings. It is thus impossible to discern the relationships of L. insularis, L. capensis and L. grotei by means of scalation or coloration alone. Our molecular phylogenetic analyses reveal that L. insularis is embedded within the L. capensis group, clearly indicating its African origin and, in so far, supporting Pasteur (1965). While the single gene as well as the multigene analyses fully support closer relationships between L. insularis and L. capensis than between L. insularis and L. grotei, the position of L. insularis within the clade formed by L. insularis, L. nyaneka, L. capensis sensu stricto and six L. aff. capensis lineages (provisionally named as “groups”; here, the last two of these are referred to as L. capensis sensu lato) is not clearly resolved. For a taxonomic revision of the L. capensis complex—one is stated to be in progress by Travers et al. unpublished (in Marques et al. 2020)—we strongly recommend a broader incorporation of genetic markers to resolve the conflicting topologies since our multigene dataset consists of only few markers especially for L. nyaneka and for the numerous groups forming L. capensis sensu lato. The clade including L. grotei and L. pakenhami is defined as the sister clade of L. insularis, L. nyaneka and L. capensis sensu lato. Lygodactylus insularis is recovered as a monophyletic group based on mtDNA, probably resulting from a single dispersal event. Furthermore, the species based on the sampling herein exhibits little intraspecific genetic variability, possibly due to the tiny size of the island resulting in a panmictic population. Members of the genus Lygodactylus successfully crossed the Mozambique Channel from mainland Africa back to Madagascar one or perhaps two times (R̂ll et al. 2010; Mezzasalma et al. 2017; Gippner et al. 2021). The ancestor of L. insularis probably began at least one further traverse in eastward direction ending on Juan de Nova, an island in the middle of the channel. Juan de Nova developed as the uplifted top of a seamount that probably was formed during the Cretaceous along the Davie Fracture (Riaux-Gobin & Witkowski 2012). Ali & Hedges (2022) suggest that it may have remained emerged as a low elevation atoll since the Early Palaeocene. In such a scenario, the dispersal of the ancestors of L. insularis may have taken place as early as 22.3 (confidence interval 15.6–30.2) mya, the stem age of the L. capensis group estimated in the Lygodactylus timetree of Gippner et al. (2021). Results of palaeogeographic reconstructions and palaeo-oceanographic modelling suggest strong surface currents in the Mozambique Channel flowing from northeast Mozambique and Tanzania eastward towards Madagascar during the Palaeogene (66 to 23 mya), which would support this scenario (Ali & Huber 2010). While the main contemporary direction of oceanic surface currents is westward, the situation in the Mozambique Channel is very complex with numerous eddies and possible countercurrents (Hawlitschek et al. 2016a). This also suggests that dispersal could have occurred during a more recent period of mainly westward marine currents. An alternative hypothesis, dispersal via short-lived Cenozoic land-bridges between mainland Africa and Madagascar (e.g., Mazza et al. 2019; Masters et al. 2020), is disputed and has been ruled out based on current geological reconstructions (e.g., Ali & Huber 2010; Ali & Hedges 2022). It also must be emphasized that despite a reasonable sampling of Eastern Africa, especially Mozambique (with L. capensis group sequences from 14 localities available so far), it is possible that a still undiscovered species representing the sister lineage of L. insularis may exist in this region. A somewhat similar situation was uncovered in the frogs of the Ptychadena mascariensis complex in which the sister lineage of the Malagasy species has only been recorded from a small area of Malawi and could have easily been overlooked without intensive sampling efforts (Zimkus et al. 2017). If a closer relative of L. insularis was found on the African mainland, it would lead to younger age estimates of the colonization event of Juan de Nova by these geckos., Published as part of Röll, Beate, Sanchez, Mickaël, Gippner, Sven, Bauer, Aaron M., Travers, Scott L., Glaw, Frank, Hawlitschek, Oliver & Vences, Miguel, 2023, Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean, pp. 232-250 in Zootaxa 5311 (2) on pages 244-245, DOI: 10.11646/zootaxa.5311.2.4, http://zenodo.org/record/8094276, {"references":["Caceres, S. (2003) Etude prealable pour le classement en Reserve Naturelle des Iles Eparses. Memoire de DESS Sciences et Gestion de l'Environnement Tropical. Univ. Reunion (Laboratoire ECOMAR) & DIREN Reunion, 136 pp. Available from: http: // poupin. joseph. free. fr / pdf / caceres- 2003 - iles-eparses-classement. pdf (accessed 13 June 2023)","Lambs, L., Mangion, P., Mougin, E. & Fromard, F. (2016) Water cycle and salinity dynamics in the mangrove forests of Europa and Juan de Nova Islands, southwest Indian Ocean. Rapid Communications in Mass Spectrometry, 30, 311 - 320. https: // doi. org / 10.1002 / rcm. 7435","Pasteur, G. (1965) Recherches sur l'evolution des lygodactyles, lezards afromalgaches actuels. Travaux de l'Institut Scientifique Cherifien, Serie Zoologie, 29, 1 - 132.","Marques, M. P., Ceriaco, L. M. P., Buehler, M. D., Bandeira, S. A., Janota, J. M. & Bauer, A. M. (2020) A revision of the dwarf geckos, genus Lygodactylus (Squamata: Gekkonidae), from Angola, with description of three new species. Zootaxa, 4853 (3), 301 - 352. https: // doi. org / 10.11646 / zootaxa. 4853.3.1","Mezzasalma, M., Andreone, F., Aprea, G., Glaw, F., Odierna, G. & Guarino, F. M. (2017) Molecular phylogeny, biogeography and chromosome evolution of Malagasy geckos of the genus Lygodactylus (Squamata, Gekkonidae). Zoological Scripta, 46, 42 - 54. https: // doi. org / 10.1111 / zsc. 12188","Gippner, S., Travers, S. L., Scherz, M. D., Colston, T. J., Lyra, M. L., Ashwini, V. M., Multzsch, M., Nielson, S. V., Rancilhac, L., Glaw, F., Bauer, A. M. & 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","Riaux-Gobin, C. & Witkowski, A. (2012) Small-sized and discoid species of the genus Cocconeiopsis (Bacillariophyta) on Holothuria atra (Juan de Nova, Mozambique Channel). Phytotaxa, 54, 43 - 58. https: // doi. org / 10.11646 / phytotaxa. 54.1.5","Ali, J. R. & Hedges, S. B. (2022) A review of geological evidence bearing on proposed Cenozoic land connections between Madagascar and Africa and its relevance to biogeography. Earth-Science Reviews, 232, 104103. https: // doi. org / 10.1016 / j. earscirev. 2022.104103","Ali, J. R. & Huber, M. (2010) Mammalian biodiversity on Madagascar controlled by ocean currents. Nature, 463, 653 - 656. https: // doi. org / 10.1038 / nature 08706","Hawlitschek, O., Garrido, S. R. & Glaw, G. (2016 a). How marine currents influenced the widespread natural overseas dispersal of reptiles in the Western Indian Ocean. Journal of Biogeography, 44, 1426 - 1440. https: // doi. org / 10.1111 / jbi. 12940","Mazza, P. P. A., Buccianti, A. & Savorelli, A. (2019) Grasping at straws: a re-evaluation of sweepstakes colonisation of islands by mammals. Biological Reviews, 94, 1364 - 1380. https: // doi. org / 10.1111 / brv. 12506","Masters, J. C., Genin, F., Zhang, Y., Pellen, R., Huck, T., Mazza, P. P. A., Rabineau, M., Doucoure & M., Aslanian, D. (2020) Biogeographic mechanisms involved in the colonization of Madagascar by African vertebrates: Rifting, rafting and runways. Journal of Biogeography, 48, 492 - 510. https: // doi. org / 10.1111 / jbi. 14032","Zimkus, B. M., Lawson, L. P., Barej, M. F., Barratt, C. D., Channing, A., Dash, K. M., Dehling, J. M., Du Preez, L., Gehring, P. - S., Greenbaum, E., Gvozdik, V., Harvey, J., Kielgast, J., Kusamba, C., Nagy, Z. T., Pabijan, M., Penner, J., R ˆ del, M. - O., Vences, M. & L ˆ tters, S. (2017) Leapfrogging into new territory: How Mascarene ridged frogs diversified across Africa and Madagascar to maintain their ecological niche. Molecular Phylogenetics and Evolution, 106, 254 - 269. https: // doi. org / 10.1016 / j. ympev. 2016.09.018"]}

Röll, Beate, Sanchez, Mickaël, Gippner, Sven, Bauer, Aaron M., Travers, Scott L., Glaw, Frank, Hawlitschek, Oliver, Vences, Miguel (2023): Supplementary Materials for Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean. Zootaxa (supp.) 5311 (2): 1-18, DOI: http://doi.org/10.5281/zenodo.8102622

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"]}