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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

Uroplatus fivehy sp. nov. LSID: urn:lsid:zoobank.org:act: AC24C7D3-1AB5-451F-9DE2-935DE7174FDD Figs 3, 4, 6���8 Remarks. This new species corresponds to the lineage previously named Uroplatus sp. A by Raxworthy et al. (2008), Uroplatus sp. 4 by Ratsoavina et al. (2011), and Uroplatus ebenaui [Ca4] by Ratsoavina et al. (2012, 2013, 2015, 2017, 2019a,b). Holotype. ZSM 1721 /2012 (field number FGZC 3590), adult male, collected on 26 November 2012 by F. Glaw, O. Hawlitschek, T. Rajoafiarison, A. Rakotoarison, F. M. Ratsoavina and A. Razafimanantsoa on the Sorata Massif, Sava Region, northern Madagascar, in the vicinity of the campsite, near 13.6805��S, 49.4451��E, 1417 m a.s.l. Paratypes. A total of 12 specimens: ZSM 1722 /2012 (field number FGZC 3591), adult female, with same collection data as the holotype; UADBA-R 70851 (FGZC 3694), adult male, collected by same collectors as holotype on 29 November 2012 on the Sorata Massif at 13.6868��S, 49.4412��E, 1272 m a.s.l.; UADBA-R 70850 (FGZC 3715), adult male, collected by same collectors as holotype on 30 November 2012 on the Sorata Massif near a creek at 13.6829��S, 49.4403��E, 1325 a.s.l.; ZSM 634 /2009 (ZCMV 11308), adult male, and UADBA-R 70966 (ZCMV 11309), adult female, collected between 22���25 June 2009 by F. Ratsoavina, R. D. Randrianiaina, E. Rajeriarison, T. Rajoafiarison, D. R. Vieites and M. Vences near Angozongahy campsite on the western side of the Makira plateau in the Northern Central East of Madagascar, near 15.437��S, 49.1167��E, 1009 m a.s.l.; UADBA 29428 (APR 05955) and UADBA 29427 (APR 05921), two adult males collected on 22 November 2004 and UADBA 29426 (APR 05859), subadult, collected on 19 November 2004 by A.P. Raselimanana at Anjiabolo site (ca. 16.2800��S, 48.8017��E; respectively at 1100 m and 1000 m a.s.l) in Marotandrano Special Reserve, Sofia Region, Northern Central East Madagascar; UADBA 29430 (APR 05984), UADBA 29431 (APR 05985), two males and UADBA 29429 (APR 05983), adult female, collected on 25 November 2004; and UADBA 29432 (APR 06118), adult male, collected on 29 November 2004 by A.P. Raselimanana at Riamalandy site (16.2850��S, 48.8150��E, between 825���875 m a.s.l.) in Marotandrano Special Reserve. Etymology. The species epithet fivehy is a noun in apposition to the genus name, meaning ���paddle��� in local Malagasy dialect, and referring to the paddle-like shape of its tail. Diagnosis. Uroplatus fivehy sp. nov. is assigned to the Uroplatus ebenaui species group based on its relatively small size, leaf-like, laterally-compressed body shape, short tail, and triangular head with supraocular spines. The following unique combination of characters characterizes the new species: (1) short (TAL/SVL 0.21���0.41) and slender (TAW/SVL 0.06���0.13) tail, (2) oral mucosa without black pigment, and (3) relatively smooth (not spiny) appearance in males. Additionally, it is distinguished by high genetic pairwise-distances> 10% in DNA sequences of the mitochondrial 16S gene from all other nominal species of Uroplatus. By its overall leaf-like appearance, the species can easily be distinguished from all members of the U. fimbriatus species group, from U. lineatus, and from the phenetic U. alluaudi species group. Within the U. ebenaui species group, U. fivehy sp. nov. can be distinguished from U. finaritra and U. malama by smaller body size (SVL 60.8���72.4 mm versus 77.5���95.3 mm); from U. malama, U. phantasticus, U. finiavana and U. finaritra by shorter tail (TAL/SVL 0.21���0.41 versus 0.48���0.76); from U. fangorn, U. ebenaui, U. fetsy, and U. kelirambo by its unpigmented oral mucosa (pale pinkish all over versus black or deep red pigment present); from U. kelirambo by more expanded tail edges (versus strongly spear-shaped; TAW/SVL 0.06���0.13 versus 0.04���0.05 in U. kelirambo). U. fivehy sp. nov. is morphologically most similar to U. fiera and to U. fotsivava, which however concordantly differ in mitochondrial and nuclear DNA sequences and are not its immediate relatives based on molecular phylogenetic analysis; it differs from U. fotsivava by a generally longer and wider tail, although the values do overlap (TAL/SVL 0.21���0.41 versus 0.15���0.32; TAW/SVL 0.06���0.13 versus 0.05���0.10), and from U. fiera possibly in a slightly larger ratio TAL/TAW (0.22���0.37 versus 0.19���0.21). Description of the holotype. Adult male in good condition with an intact tail and everted hemipenes. SVL 63.1 mm, tail length 25.9 mm, maximum tail width 5.7 mm, for further measurements see Table 1. Head triangular in dorsal view; canthus rostralis recognizable and almost straight; snout sloping strongly and continuously downward anteriorly; snout weakly depressed, short; eyes large (eye diameter 4.6 mm), not bulging above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small but distinct (horizontal diameter 0.6 mm), its opening facing posterolaterally, but also posteroventrally (ear opening clearly visible in lateral view but not in dorsal view); nostrils laterally oriented; body somewhat laterally compressed, without lateral dermal fringes; limbs without fringes and practically without spines on forelimbs and hindlimbs, except for a small flap on the knee; forelimb reaches beyond tip of snout when adpressed forward and does not reach the groin when adpressed backwards along body (forelimb length/axilla���groin distance 27.7/ 29.7 mm = 93%), hindlimb reaches beyond axilla when adpressed forward along body (hindlimb length/axilla���groin distance 37.4/ 29.7 mm = 126%); original tail length 41% of snout���vent length, membranous borders of the denticulated tail rather asymmetrical, broadening on either side and then narrowing again toward the tip), completely absent from the slightly spatulate tail-tip. Nares separated from each other by eight small granular scales, from the first supralabial by one scale, and from the rostral scale by two scales; the first supralabial taller than the others; rostral entire, much wider than tall; mental scale small, not differentiated from infralabial scales; 21/20 (right/left) supralabials and 19/21 infralabials (as defined in methods above); scales at the periphery of the chin (toward the infralabial scales) slightly larger than central chin scales; the dorsal scales of head, neck, body, limbs and tail small, granular, juxtaposed and largely of homogeneous size interspersed with very few larger and partly raised tubercles; the numerous curved transverse lines on the head and dorsal side of the body are not marked by significantly enlarged scales. Very few spines on head and hindlimbs, a dermal flap on each knee, bearing no spine; no flaps or spines on elbow and forelimb; a prominent pointed flap on the posterior portion of each upper eyelid (supraocular spine); posterior border of eye fringed. Axillary pits not recognizable. Scales of the ventral abdomen distinctly larger than dorsal body scales and arranged more homogeneously. Hemipenes everted, bearing two lobes. Calyx with protuberance bearing honeycomb-like structures, especially on the asulcal side. Area of sulcus spermaticus is smooth. Each lobe with a dense field of pointed papillae at its apex. Coloration of the holotype. After eight years of preservation in 70% ethanol the color pattern remains similar to that of the living animal (Figs 6H, 8) but its vividness and intensity have slightly faded. All dorsal surfaces are mottled beige and blackish in base color. The tail has a beige pattern on its dorsal pygal section, bordered by blackish color. The dorsum has rather distinctly defined posteroventrally oriented lighter and darker beige chevrons. The dorsal surfaces of the head are dark brown with a series of six distinct transverse narrow beige lines, running either straight or converging anteriorly or posteriorly. No dark band between the eyes, but a dark V-shaped area on the nape is well distinguishable. Additional dark flecks are present on the mid-dorsum, and several smaller asymmetrical spots on other parts of the body, including legs and nape. One light cream marking is present posteriorly below each eye, descending to the supralabials. The supralabials and the infralabials are generally dark gray in color. The venter, including the ventral limbs except for the lower hindlimbs, is slightly lighter beige than the dorsum, and less mottled. The pectoral region bears two elongated, poorly delimited whitish spots. The postpygal section of the tail is of similar color dorsally and ventrally. The oral mucosa is unpigmented and whitish in preservative. The iris of the eye is gray, and the vertical pupil is blackish. Variation. As described for U. fangorn above, there is a substantial color variation (Figs 6���7), and sexual dimorphism in the edges of the tail. The color of the eye varies from beige/gray to dark brown and reddish-brown. Dorsal color can be uniform beige to reddish-brown with or without white lichen-like markings. Two sharply delimited vertebral black patches on the anterior and posterior part of the dorsum are present in several individuals. Relative tail length and tail width are characterized by substantial variability among individuals (Fig. 4). Here, we have included in the paratype series and in the morphological comparisons a series of individuals from Marotandrano Special Reserve that we attribute to this species. However, it must be stressed that the species identity has been confirmed by DNA sequences only for one of these (UADBA 29428 = APR5955). In the unlikely case that some of these individuals will turn out to belong to another species of Uroplatus, the morphometric variation of U. fivehy may be narrower than reported here. ......continued on the next page Distribution, natural history and conservation status. According to the material collected and examined by us, U. fivehy is known from (1) the type locality on the Sorata Massif, (2) the western slope of the Makira Reserve, and (3) Marotandrano Special Reserve. In addition, mitochondrial DNA sequences published by Ratsoavina et al. (2012) suggest its occurrence in (4) the Anjanaharibe-Sud Massif, and mitochondrial DNA sequences from Raxworthy et al. (2008) furthermore suggest the occurrence of the species in (5) Marojejy, (6) Ankitsika and (7) Betaolana. The confirmed elevational range from our own samples is between 1009 and 1417 m a.s.l., and the record from Ankitsika would extend this range downwards to 830 m a.s.l., characterizing the species as rather wide-ranging from the Northern Central East to the North East of Madagascar from mid-elevations to highlands (see also Ratsoavina et al. 2013). All specimens were found in intact rainforest, at night, perched on branches of trees. At Marotandrano, an adult male was found hidden in a Pandanus leaf axil during the day in closed canopy humid forest, and two adult individuals (male and female) were found at night close to each other in the same tree on 25 November 2004, suggesting it was probably the reproduction period for this species. Individuals were often observed active on small branches, leaves, and lianas at 0.5���3 m above the forest floor. All individuals were collected on slope or ridge forest. In Marotandrano, the species appeared to be quite common in forests associated with vine-like bamboo and shrub vegetation. A minimum convex polygon of the known distribution of this species covers an area of ca 15,100 km 2, qualifying the species for a status of Vulnerable under the IUCN Red List criterion B (IUCN 2012). This species is known from several protected areas (Marotandrano Special Reserve, Makira Reserve, COMATSA Nord, Marojejy National Park, Anjanaharibe-Sud National Park) but cumulatively just seven locations (B subcriterion a). These locations, and especially the unprotected areas between them, are experiencing substantial ongoing declines in the extent and quality of habitat (B subcriterion b(iii)). Therefore, we propose to list this species as Vulnerable under IUCN criterion B1ab(iii)., Published as part of Ratsoavina, Fanomezana M., Glaw, Frank, Raselimanana, Achille P., Rakotoarison, Andolalao, Vieites, David R., Hawlitschek, Oliver, Vences, Miguel & Scherz, Mark D., 2020, Towards completion of the species inventory of small-sized leaf-tailed geckos: two new species of Uroplatus from northern Madagascar, pp. 251-271 in Zootaxa 4895 (2) on pages 261-268, DOI: 10.11646/zootaxa.4895.2.5, http://zenodo.org/record/4322511, {"references":["Raxworthy, C. J., Pearson, R. G., Zimkus, B. M., Reddy, S., Deo, A. J., Nussbaum, R. A. & Ingram, C. M. (2008) Continental speciation in the tropics: contrasting biogeographic patterns of divergence in the Uroplatus leaf-tailed gecko radiation of Madagascar. Journal of Zoology, 275, 423 - 440. https: // doi. org / 10.1111 / j. 1469 - 7998.2008.00460. x","Ratsoavina, F. M., Louis, E. E. Jr., Crottini, A., Randrianiaina, R. D., Glaw, F. & Vences, M. (2011) A new leaf tailed gecko species from northern Madagascar with a preliminary assessment of molecular and morphological variability in the Uroplatus ebenaui group. Zootaxa, 3022 (1), 39 - 57. https: // doi. org / 10.11646 / zootaxa. 3022.1.3","Ratsoavina, F. M., Vences, M. & Louis, E. E. Jr. (2012) Phylogeny and phylogeography of the Malagasy leaf-tailed geckos in the Uroplatus ebenaui group. African Journal of Herpetology, 61, 143 - 158. https: // doi. org / 10.1080 / 21564574.2012.729761","Ratsoavina, F. M., Raminosoa, N. R., Louis, E. E. Jr., Raselimanana, A. P., Glaw, F. & Vences, M. (2013) An overview of Madagascar's leaf tailed geckos (genus Uroplatus): species boundaries, candidate species and review of geographical distribu- tion based on molecular data. Salamandra, 49, 115 - 148.","Ratsoavina, F. M., Ranjanaharisoa, F. A., Glaw, F., Raselimanana, A. P., Miralles, A. & Vences, M. (2015) A new leaf-tailed gecko of the Uroplatus ebenaui group (Squamata: Gekkonidae) from Madagascar's central eastern rainforests. Zootaxa, 4006 (1), 143 - 160. https: // doi. org / 10.11646 / zootaxa. 4006.1.7","Ratsoavina, F. M., Gehring, P. S., Scherz, M. D., Vieites, D. R., Glaw, F. & Vences, M. (2017) Two new species of leaf-tailed geckos (Uroplatus) from the Tsaratanana mountain massif in northern Madagascar. Zootaxa, 4347 (3), 446 - 464. https: // doi. org / 10.11646 / zootaxa. 4347.3.2","Ratsoavina, F. M., Raselimanana, A. P., Scherz, M. D., Rakotoarison, A., Razafindraibe, J. H., Glaw, F. & Vences, M. (2019 a) Finaritra! A new leaf-tailed gecko (Uroplatus) species from Marojejy National Park in north-eastern Madagascar. Zootaxa, 4545 (4), 563 - 577. https: // doi. org / 10.11646 / zootaxa. 4545.4.7","IUCN. (2012) IUCN Red List Categories and Criteria: Version 3.1. IUCN, Gland and Cambridge."]}

We describe two new frog species of the Madagascar-endemic genus Gephyromantis, belonging to the G. tandroka complex in the subgenus Duboimantis. Gephyromantis (Duboimantis) saturnini sp. nov. is known from specimens collected at Ampotsidy, in the District de Bealanana of the Région Sofia of northern Madagascar. It is a large and genetically distinct species with small interocular tubercles and indistinct femoral glands that emits calls in series composed of couplets and triplets. Morphologically and bioacoustically, it is most similar to G. tandroka, but genetically it was recovered as sister to G. schilfi, from which it is separated by an uncorrected p-distance of 5.1 % in the mitochondrial 16S rRNA gene. Gephyromantis (Duboimantis) grosjeani sp. nov. was collected on the Sorata massif, in the District de Vohemar of the Région Sava of northern Madagascar. It is a medium-sized species with large and distinct femoral glands, which are a distinctive characteristic from other members of this complex. Genetically, it is distinguished from all other Gephyromantis species by uncorrected p-distances of at least 5.1 % in the 16S rRNA gene.

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

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