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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

Cornufer (Batrachylodes) exedrus sp. nov. Figs. 2a–c, 4, 5 Holotype. South Australia Museum (SAMA) R64760 (SJR Field No. 10784), adult male, collected by S. J. Richards at Tompoi Camp, 1590 m above sea level (05°20.623'S, 151°18.873'E; WGS-84), Nakanai Mountains, East New Britain Province, Papua New Guinea, 19 April 2009. Paratypes. SAMA R64762 and R64764 (SJR Field Nos. 10815 and 10820 respectively), two adult males, and SAMA R64763 (SJR 10819), adult female, all collected at the type locality, 21 April 2009; SAMA R64761 and R71008 (SJR 10798 and 10821 respectively), two adult males, same locality, 20 and 21 April 2009, respectively; SAMA R64765 (SJR 10843), juvenile, and SAMA R71007 (SJR 10846), adult male, same locality, 23 April 2009. Etymology. The specific epithet is derived from the masculine formation of the Greek adjective exedros, meaning away from home, strange, or extraordinary. The surprising discovery of Cornufer (Batrachylodes) exedrus from a high elevation site in New Britian is a taxonomic and biogeographic enigma, given that all other species in the subgenus are restricted to the Solomon Archipelago. Diagnosis. The new species is a member of the genus Cornufer based on molecular phylogenetic evidence. There are currently no morphological synapomorphies known for the genus Cornufer, however, based on the updated taxonomic arrangement of this clade (Brown et al. 2015) into several morpholgically diagnosible subgenera the new species can be assigned to the subgenus Batrachylodes, as defined by Brown et al. (2015), on the basis of its small, triangular shaped body, pointed snout, absence of interdigital webbing, absence of vomerine teeth, presence of a darkened loreal stripe extending diagonally across the flanks, and molecular phylogenetic placement (Brown et al. 2015:fig. 2). Cornufer exedrus is distinguished from congeners by a combination of (1) adult body size small (SVL 17.6–18.8 mm for six adult males, mean = 18.3 mm; SVL 18.3 mm for one adult female), (2) body shape triangular with a small, pointed head, (3) terminal disks of the fingers and toes narrowly expanded (among adults: Fin3DW 0.3–0.6 mm, mean = 0.5 mm; Toe4DW 0.4–0.7 mm, mean = 0.6 mm) (Figs. 2, 4, 5), (4) vomerine teeth absent, (5) digits lacking all vestiges of lateral flange and interdigital webbing, (6) dorsal skin smooth to finely granular, lacking prominent dermal tubercles or ridges, (7) subarticular tubercles of fingers, toes, hands and feet uniformly low and flattened, (8) dark band extending along the canthus rostralis and onto the flank with a smooth dorsal margin demarcating lighter dorsal and darker lateral colors, and (9) unique advertisement call consisting of a series of brief, high-frequency atonal notes. Comparisons with other species. Among members of the subgenus Batrachylodes, Cornufer exedrus can be easily distinguished from C. elegans, C. gigas, C. montanus, C. vertebralis, and C. wolfi by the narrowly expanded (versus widely expanded) terminal finger disks except for the first finger (Fin3DW/SVL = 0.02–0.03, Fin3DW/ Fin3L = 0.1–0.2, and Fin3DW/TD = 0.2–0.3 in C. exedrus versus Fin3DW/SVL> 0.03, Fin3DW/Fin3L> 0.2, and Fin3DW/TD> 0.3 in the aforementioned species) and smaller adult body size (SVL 17.6–18.8 mm in C. exedrus versus SVL> 20.0 mm in the aforementioned species) (Table 2). Of the Batrachylodes that lack prominently dilated finger disks, Cornufer exedrus differs from C. mediodiscus in having a smaller adult body size (SVL 17.6– 18.8 mm in C. exedrus versus 20.2–26.0 mm in C. mediodiscus), slightly smaller third finger disk width (reflected in the Fin3DW/SVL, Fin3DW/Fin3L, and Fin3DW/TD ratios, Table 2), and lacking raised dorsal skin ridges (Brown & Parker 1970). Cornufer exedrus differs from C. minutus by its slightly larger body size (SVL 17.6–18.8 mm in C. exedrus versus 15.2–17.3 mm in C. minutus) and by differences in the proportions related to hind limb length (reflected in the HW/TBL, TBL/SVL, and PL/SVL ratios, Table 2). Cornufer minutus also differs in color pattern, with the dark band along the canthus rostralis more poorly defined, often broken or with an irregular dorsal margin, than in Cornufer exedrus. The new species differs from C. trossulus by its slightly smaller body size (SVL 17.6–18.8 mm in C. exedrus versus 18.9–24.3 mm in C. trossulus) and by differences in the proportions related to hind limb length (reflected in the HW/TBL, TBL/SVL, and PL/SVL ratios, Table 2). On New Britain Island, C. exedrus can be differentiated from most congeners by its smaller adult body size (Cornufer species on New Britain (C. exedrus can be further distinguished from C. caesiops (and the morphologically similar C. browni, on nearby New Ireland Island; Allison & Kraus 2001; Kraus & Allison 2009) in having narrowly expanded finger disks (versus widely expanded in C. caesiops and C. browni); from C. akarithymus, C. bufonulus, and C. sulcatus in having a smooth dorsal surface (versus tuberculate or with dorsal ridges in C. akarithymus, C. bufonulus, and C. sulcatus); and from C. boulengeri (males and juveniles previously assigned to C. rhipiphalcus; Kraus 2008), by the lack of a fanlike array of dorsal ridges in the scapular region and a less prominent dark face mask, which does not extend onto the flanks in C. boulengeri. Description of holotype. Adult male, in good condition, preserved with mouth open, incision in left thigh where SAMA tag was threaded; habitus moderately robust; head not distinct, slightly narrower in dorsal aspect than body, length 41% of SVL; head length 120% of head width; snout short, tip acuminate in lateral aspect, rounded dorsal aspect, protruding prominently beyond lower jaw, unpigmented at tip; lower jaw notched at tip; eyes protrude only slightly beyond silhouette of head in both dorsal and lateral aspects; labial region rounded and smooth, lips not swollen, not extending past eyes when viewed in dorsal aspect; interorbital region slightly convex; eye diameter 90% of interorbital distance; pupil horizontally ovoid; canthus rostralis medially bowed; loreal region slightly concave; eye diameter 55% of snout length; nostrils slightly laterally protuberant; eye-narial distance 1.25 times the distance from nostril to snout tip; internarial region flat; tympanum distinct, its diameter 89% of eye diameter; slight supratympanic fold present, extending from posterior edge of eye, over dorsal edge of tympanum, and terminating at supra-axillary (post-rictal) region; tongue ovoid, with a slightly narrower anterior attachment; choanae round, minute, at anterolateral edge of palate, separated by a distance five to seven times greater than their diameter, obscured in ventral aspect by palatal shelf; vomerine teeth absent; openings to vocal sac minute slits, at level of the angle of jaw. Dorsal skin of body, head, and limbs generally smooth, lacking dermal folds, tubercles, or ridges; ventral surfaces of trunk, head, throat, and limbs smooth, with the exception of fanlike array of 12–14 small white tubercles on anterior margin of chin. Cornufer Batraychylodes......continued on the next page 9 5: 6;!#") C. montanus C. gigas & Manus length 50% of foot length; digits of manus narrow (Figs. 4–5), ovoid in cross-section; dermal flanges absent on lateral edges of digits; terminal finger disks barely expanded ( Hindlimbs short; tibia length 41% of snout-vent length, pes length 99% of tibia length; tarsus smooth, lacking folds, flaps, or tubercles; terminal disks of toes slightly expanded and ovoid ( Measurements of holotype. SVL 18.1; ED 1.8; TD 1.6; HL 7.4; SNL 3.3; IOD 2.0; HW 6.2; FL 8.1; TBL 7.5; TSL 4.5; PL 7.4; ML 3.7; FA 3.7; Toe4L 4.1; Fin1L 1.1; Fin3L 2.2; Fin1DW 0.3; Fin3DW 0.3; PpFin3 0.2; Toe4DW 0.4; PpToe4 0.3. Color of holotype in preservative. Dorsal surfaces of body and head exhibit nearly homogeneous light brown coloration; under magnification, background color of dorsal surfaces is pale cream and unpigmented with dense and nearly uniform speckling of brown chromatophores; less brown pigmentation present in interorbital region, extending posteriorly in a U-shaped pattern to a midpoint between the scapulae, giving slightly paler appearance to region along top of head than rest of body; two small, dark brown interorbital spots between anterior margin of eyes; tip of snout unpigmented; dorsal surface of limbs similar in background coloration to rest of body, but with darker brown transverse bars (three on forearm, three on femur, three on tibia, one on tarsus, two on metatarsus) with irregular margins and poorly defined; dorsal surfaces of hands, feet, and digits similarly colored as rest of the body and limbs with irregular dark spots and bars; the lateral surfaces of the head and body have a distinct dark brown band (darker than dorsal coloration) covering entire loreal and supralabial region, extending posteriorly along flanks diagonally (anteriodorsally to posteroventrally) towards inguinal region; this lateral band is darkest and near uniformly shaded along head (although tympanum is slightly lighter) becoming lighter and more diffusely speckled posterior to the humerus, into light tan irregular blotching in the inguinal region; the dorsal margin of this lateral band is nearly smooth and darkest brown in coloration (with respect to the rest of band), forming a clear dark line demarcating lighter dorsal and darker lateral colors extending from snout tip (although absent from unpigmented snout tip) along the canthus rostralis (above nostril) and upper eyelid, and posterior onto flanks slightly above tympanum; dark dorsolateral line fades in the inguinal region becoming prominent again on hindlimbs (although with more irregular margin) particularly on anterior edge of thigh and knee, fading into diffuse irregular tan blotching on tibia; underside of throat dark brown with lighter unpigmented mottling throughout; brown ventral coloration of throat fades posterior to the pectoral region becoming diffuse irregular tan blotching on pale cream midventer; ventral surfaces of forearms tan, darker brown towards outer edge of forearms; inner forearm with unpigmented band extending to underside of humerous; underside of hindlimbs similar in coloration to midventer, pale cream with brown irregular mottling; when viewed in posterior aspect cloacal region nearly uniform dark brown with smooth, dark dorsal margin above cloaca extending along the posterior of thighs clearly demarcating it from lighter dorsum; when in a seated position, this dark cloacal-femoral band grades into similarly colored dark band along outer edge of tarsus; small dark brown blotch just above cloaca; ventral surfaces of hands and feet dark brown with pale subarticular banding. Variation. Compared to the holotype (SAMA R64760), dorsal coloration of the head and body of the paratypes ranges from a darker brown (SAMA R64763, SAMA R64764, SAMA R71008, SJR 10843) to a lighter cream (SAMA R64761, SAMA R64762, SAMA R71007) background coloration, and whereas the holotype has a nearly uniform dorsal coloration, all of the paratypes exhibit some degree of dorsal markings. SAMA R71008, SAMA R64763, and SJR 10843 possess a faint, thin and lighter vertebral stripe. Four of the paratypes (SAMA R71007, SAMA R71008, SAMA R64762, SAMA R64761) possess a pair of discrete darker brown to black dorsal spots that are posteroventrally located just behind the projection of the sacrum. Several paratypes also have irregular darker paravertebral spots between the scapulae, which are similar in size to the posterior dorsal spots in some (SAMA R64761, SAMA R64762) or a faint speckling in others (SJR 10843, SAMA R64763). SAMA R64761 exhibits more extensive irregular dark flecking and blotching along the dorsum. Three paratypes (SAMA R64764, SAMA R64761, SAMA R71007) possess a faint, thin and darker interorbital bar. Several paratypes also exhibit faint darker banding extending from behind eyes and/or the sacral region intersecting at the midscapular region, leaving a faint x or v-shaped pattern along the dorsum. All of the paratypes possess the transverse bars on the limbs as described in the holotype. There is variation in breadth and extent of the dark lateral band that extends from the loreal region onto the flanks. In some individuals, the lateral band is darker and more filled in throughout the snout and flank region, giving appearance of dark face mask which remains solid (not fading) along the flanks. However, in others the lateral band is limited to more of a dark stripe from the tip of the snout, along the canthus rostralis, and posterior towards the inguinal region, which quickly fades ventrally into a light-brown coloration similar to the color of the dorsum. In two individuals (SAMA R64761, SAMA R71007), the lateral band is almost entirely limited to the snout along the canthus rostralis and quickly broken and fading posteriorly behind the eye, and in these individuals there is almost no darker shading below the band in the loreal and supralabial region of the snout or along the flanks. Ventral coloration of several of the paratypes is similar to that of the holotype, with a darker brown chin and throat region fading into a lighter cream midventral region and the underside of the hindlimbs with diffuse brown speckling. However, in one individual (SAMA R64762) the entire vertrum is similar in coloration to the brown throat, and in three individuals (SAMA R64761, SAMA R71007, SJR 10843) the brown pigmentation has faded, leaving the entire ventrum pale cream in coloration. In most of the paratypes the skin texture is smooth, similar to that of the holotype, however several specimens exhibit finely granular skin along the dorsum, and one indiviudal (SAMA R64761) possesses small raised dorsal ridges and minute tubercles associated with the darker brown spots and flecks along the dorsal surface of the body. The small unpigmented tubercles on the chin and throat of the holotype are also present on most of the paratypes (although varying in number and extend along the throat); however, they are absent on the juvenile (SJR 10843) and female (SAMA R64763) specimens, suggesting that it may be a sexually dimorphic character. Additionally, the unpigmented projection of the snout on the holotype is prominent on most of the paratypes except the aforementioned juvenile and female paratypes, suggesting it may also be a sexually dimorphic character. Mensural variation is presented in Table 2. All mensural ratios for the single female paratype are within the ranges found in males, possibly indicating little sexual dimorphism in morphometric characters. Color in life. Based on color images of the holotype (SAMA R64760, Figs. 2–3) and two paratypes (SAMA R64763 and SAMA R64764, Fig. 2) in life taken during the daytime, prior to preservation: Dorsal ground coloration of body and limbs of the holotype (SAMA R64760) a pale pinkish-salmon that is overlain with small darker pinkish-orange spots or streaking (streaks most prominent on the tibia) with some small irregular whitish spotting throughout; thicker (but faint) darker brown transverse bars extending across the hindlimbs and three thin (broken) darker brown transverse bars on the forearm; dorsal coloration of the head orangish, giving the appearance of a head cap (from just behind the unpigmented snout projection extending in a U-shaped pattern posterior to the eyes to the mid-scapular region); three small whitish and two small dark brown interorbital spots between the anterior margins of the eyes; margin of the upper eyelid with a thin light whitish band speckled with diffuse brown flecking. Dorsal coloration of the two paratypes different than that of the holotype. One paratype (SAMA R64763) has a more solid and uniformly darker brown coloration across the head, body, and limbs flecked with minute irregular dark brown, whitish, and reddish spots; limbs also with some suffused lighter orangishbrown highlights; a thin and slightly broken vertebral stripe is present, and a cluster of small dark-brown paravertebral spots in the midscapular region; faint darker brown transverse bars present on the limbs. The other paratype (SAMA R64764) possesses lighter tan dorsal coloration on the head and body, fading to darker brownishgray hindlimbs (forearms of similar color as hindlimbs), leaving the transverse bars on the hindlimbs poorly diagnosable; faint minute irregular dark-brown, whitish, and reddish spots or blotching on the dorsal surfaces of the body and limbs; light yellow-gold pigment faintly dusting the forelimbs and hands; two darker brown dorsal bands also apparent, one interorbital band between the midpoints of the orbits and upper eyelids, and one extending as a forward-facing V-shape connecting at the midscapular region and extending posteroventrally to the mid-dorsum. The shading of the dark-brown lateral band (from the snout tip to the flanks) also varies slightly. In all three specimens the dorsal margin of the lateral band is the darkest brown, particularly in the loreal region along the canthus rostralis; in SA

Maximum Likelihood tree (RAxML, combined mitochondrial and nuclear data) and including all sampled specimens of Lepidodactylus s.l.

Varanus samarensis Koch et al. 2010 Synonymy. Varanus cumingi samarensis Koch et al., 2010. Zootaxa 2446, 19 ��� 23. Holotype.��� ZFMK 64713, subadult male, collected by M. Gaulke 1989, San Augustin near Gandara, Samar Island, Philippines. New material examined.���Samar (1 specimen): KU 310870 (CDS Field No. 2654), adult male, collected by Cameron D. Siler, 13 October 2007, Barangay San Rafael, Municipality of Taft, Eastern Samar Province, Samar Island, Philippines; Bohol (1 specimen): KU 335263 (CDS Field No. 4768), adult male, collected by Cameron D. Siler, 0 4 August 2009, Barangay Danicop, Municipality of Sierra Bullones, Bohol Province, Bohol Island, Philippines. Other material. ���Refer to Koch et al. (2010 b). Diagnosis. ���Following the findings of Koch et al. (2010 b), Varanus samarensis can be distinguished from other members of the V. salvator Complex by a combination of (1) distinctive dorsal color pattern of 5���8 transverse bands of yellow ocelli or spots through the trunk; (2) a dark (black) head dorsally, with symmetrical yellow markings; (3) a variable black stripe in the temporal region, occasionally bordered by a bright (yellow-white) stripe ventrally; (4) the absence of a medio-dorsal stripe on the lateral surfaces of the nuchal region and trunk; (5) narial openings positioned approximately 2.5 times closer to the tip of the snout than to the ocular. Comment and justification. ���Although within the same PAIC (Brown & Diesmos 2009), the Samar���Leyte versus Mindanao Island biogeographical boundary (informally referred to as the ���Dinagat���Caraga Filter Zone���) has been evoked in several previous studies to explain the existence of deep phylogenetic splits between the northern and southern reaches of the Mindanao faunal region (Steppan et al. 2003; Hosner et al. 2013, 2014; Brown et al. 2013, 2014). We therefore do not find it surprising that Samar, Leyte, Bohol (and, presumably, smaller associated lanmasses) have a distinctive species of monitor lizard that apparently is isolated from Mindanao���s Varanus cumingi (Welton et al. 2013 a, b). Distribution. ���See Koch et al. (2010 b) for additional data about variation, ecology, and distribution., Published as part of Welton, Luke J., Travers, Scott L., Siler, Cameron D. & Brown, Rafe M., 2014, Integrative taxonomy and phylogeny-based species delimitation of Philippine water monitor lizards (Varanus salvator Complex) with descriptions of two new cryptic species, pp. 201-227 in Zootaxa 3881 (3) on page 221, DOI: 10.11646/zootaxa.3881.3.1, http://zenodo.org/record/226203, {"references":["Koch, A., Gaulke, M. & Bohme, W. (2010 b) Uravelling the underestimated diversity of Philippine water monitor lizards (Squamata: Varanus salvator complex), with the description of two new species and a new subspecies. Zootaxa, 2446, 1 - 54.","Brown, R. M. & Diesmos, A. C. (2009) Philippines, Biology, In: Gillespie, R. & Clague, D. (Eds.), Encyclopedia of Islands. University of California Press, Berkeley, pp. 723 - 732.","Steppan S. J., Zawadzki C, Heaney L. R. (2003) Molecular phylogeny of the endemic Philippine rodent Apomys (Muridae) and the dynamics of diversification in an oceanic archipelago. Biological Journal of the Linnaean Society, 80, 699 - 715.","Welton, L. J., Siler, C. D., Linkem, C. W., Diesmos, A. C., Diesmos, M. L., Sy, E. & Brown, R. M. (2013 a) Dragons in our midst: Phyloforensics of illegally traded Southeast Asian monitor lizards. Biological Conservation, 159, 7 - 15.","Welton, L. J., Siler, C. D., Oaks, J. R., Diesmos, A. C. & Brown, R. M. (2013 b) Mulilocus Phlogeny and Bayesian estimates of species boundaries reveal hidden evolutionary relationships and cryptic diversity in Southeast Asian monitor lizards. Molecular Ecology, 22 (13), 3495 - 3510."]}

Varanus bangonorum sp. nov. Figs. 3, 6���9 Holotype. PNM 9798 (formerly University of Kansas Natural Biodiversity Institute [KU] 335744; Field no. RMB 17757), sub-adult male, collected by SLT and RMB, 16 July 2013, 230 m above sea level (12.787806, 120.915972; WGS- 84), Sitio Aruyan, Barangay Malisbong, Municipality of Sablayan, Occidental Mindoro Province, Mindoro Island, Philippines. Paratopotypes. KU 335742 (RMB Field No. 17720), juvenile, collected 11 July 2013. Paratypes. KU 335743 (RMB Field No. 17824), juvenile, Lake Libuao, Barangay Malisbong, Municipality of Sablayan, Occidental Mindoro Province, Mindoro Island, Philippines; PNM 9799 (formerly KU 308403), 308437 (ELR Field Nos. 843, 877), juvenile and adult female, Sitio Ulasan, Barangay Harrison, Municipality of Paluan, Occidental Mindoro Province, Mindoro Island, Philippines; KU 305151, 305161, 305163 (CDS Field Nos. 714, 715, 648), juveniles, Barangay Tinogboc, Municipality of Caluya, Antique Province, Semirara Island, Philippines. Other Material. See Specimens Examined Section Etymology. The specific epithet, bangonorum, is a plural noun derived from the name of the indigenous Bangon peoples of Mindoro Island, who reside along the rivers and central Cordillera mountain system in Oriental Mindoro Province. Anthropologists have shown the Bangon tribal group to be culturally distinct from other tribal groups inhabiting Mindoro (Tweddell 1970; Mangyan Heritage Center). Suggested common name: Bangon Monitor Lizard. Diagnosis. Varanus bangonorum can be distinguished from congeners by (1) distinct, relatively large dark spots or blotches in the gular region; (2) low counts of dorsal gular scales; (3) low counts of dorsal trunk scales; and (4) low counts of scales around the neck anterior to the gular fold. Additionally, this distinct lineage is biogeographically circumscribed on the deepwater islands of Mindoro and Semirara. Comparisons. This new species in phenotypically most similar to Varanus marmoratus, from which it can be distinguished by the presence of distinct, dark and irregularly-shaped spots or blotches in the gular region (versus speckling and transverse bands; Fig. 3), and an allopatric distribution on the islands of Mindoro and Semirara (versus Luzon and Lubang islands, and the Batanes and Babuyan island groups; Figs. 1, 2). Additionally, although the new species is phenotypically similar to V. marmoratus, it is not closely related to this species (Fig. 2, phylogeny). Varanus bangonorum can be distinguished from its closest but allopatric relative, V. palawanensis, by having generally fewer rictus���rictus scales (mean = 56 �� 4, versus 62 �� 4), fewer dorsal scales from the tympanum to the gular fold (mean = 26 �� 2, versus 32 �� 3), fewer dorsal scales from the nuchal fold to the hind limb insertion (mean = 80 �� 9, versus 96 �� 3), fewer total dorsal scales (mean = 106 �� 10, versus 128 �� 3), fewer scales around the neck anterior to the gular fold (mean = 76 �� 6, versus 87 �� 4), and by having distinct, dark irregular spots or blotches in the gular region (versus irregular speckling with anterior transverse bands). Of the remaining geographically proximate species of the V. salvator Complex, this new species can be distinguished from V. nuchalis by a distribution on Mindoro and Semirara islands (versus Visayan islands), generally fewer midbody scales (mean = 136 �� 9, versus 150 �� 7), generally more dorsal scales from the tympanum to the gular fold (mean = 26 �� 2, versus 23 �� 2), and the presence of distinct dark spots or blotches in the gular region (versus a uniform dark gular coloration). Additionally, this new species can be distinguished from the allopatric Bicol species V. dalubhasa by having distinct, irregularly shaped spots or blotches (versus variable speckling and faint anterior transverse bands) in the gular region. Lastly, V. bangonorum exhibits entirely unique haplotypes or haplotype networks, relative to all other members of the V. salvator Complex (see Welton et al. 2010 a, b). We have constrained our morphological analyses to the geographically most proximate taxa due to previous studies (Koch et al. 2007, 2010 b) demonstrating their distinctiveness relative to the remaining diversity within the V. salvator Complex. Description of holotype. A sub-adult male, hemipenes not everted; snout���vent length 343 mm; tail 601 mm; head relatively slender, length 65.9 mm, width 38.2 mm, and depth at eye 28.5 mm; snout length 34.0 mm, 51.6% head length; snout rounded anteriorly; narial openings 4.3 mm, ovular, posteriorly elevated, encircled by 9 L/ 9 R small polygonal scales; snout elevated sharply just anterior to narial region, then continuing gradually to ocular region; narial region only moderately elevated above horizon of snout; cranial table squarish, slightly wider than long. Head scales ovular to polygonal (Fig. 9), heterogenous in size with the smallest occuring on lateral margins of cranial table and directly surrounding pinneal scale, and largest occuring dorso-medially from tip of snout to pineal region; supraoculars 7 L/ 6 R, subrectangular and elongated transversely; supralabials 52, slightly larger than head scales; infralabials 52, smaller than supralabials; both supra- and infralabials increasing in size anteriorly; nuchals large, circular, generally homogenous in size throughout nuchal region except for directly posterior to cranial table; lateral nuchals small and granular, approximately �� the size of dorsals; sutures between scales larger than those on head; nuchals 25, in semi-regular transverse rows from posterior margin of tympanum to forelimb insertion; dorsal trunk scales arranged in 77 semi-regular transverse rows from fore- to hindlimb insertions, ovular, generally smaller than nuchals, and decreasing in size posteriorly to hindlimb insertion as well as laterally, with smallest scales occurring adjacent to limb insertions; midbody scales 131; axilla-groin distance 157 mm; dorsal limb scales ovular, decreasing in size distally and ventrally; fore- and hindlimbs 55 and 82 mm, respectively, 16.0% and 23.9% snout���vent length, respectively, 24.2 and 36.8 mm diameter at insertions, respectively; limb scales present in semiregular rows; scales of manus and pes smaller and more irregular in shape, ranging from circular to nearly square dorsally; supradigitals rectangular dorsally, ovular laterally, with terminal scale nearly twice as large as others; digits terminate in long, slender recurved claws; caudals ovular, small and granular anteriorly, becoming rectangular posteriorly; dorsal keel composed of paired, posteriorly elevated scales commenses 37.7 mm from hindlimb insertion, continuing to tail terminus with scales gradually decreasing in size; caudal scales moderately keeled, in regular transverse and longitudinal rows laterally, decreasing in size posteriorly. Ventrals less variable than dorsals; gular scales generally rectangular anteriorly, bcoming more ovular towards gular fold; smallest scales medially in anterior third of gular region, with largest scales just anterior to gular fold but followed posteriorly by three transverse rows of smaller, more granular gulars; scales around the neck anterior to the gular fold 67, mid-gular 76; gular scales in 70 irregular rows from tip of snout to gular fold; ventrals of fore- and hindlimbs circular to polygonal, with those of forelimbs slightly smaller than those of hindlimbs; precloacals similar to those of hindlimbs; ventral trunk scales from gular fold to hind limb insertion 83, ovular anteriorly, becoming rectangular posteriorly, present in semi-regular rows; scales decreasing in size laterally and at hindlimb insertions; ventral caudal scales rectangular, smallest just posterior to cloacal opening; scale surface slightly rounded anteriorly, becoming sharply keeled and more longitudinally elongate posteriorly. Measurements of holotype (in mm). SVL 343; tail length 601; head length 65.9, width 38.2, depth at eye 28.5; eye���naris distance 22.2; naris���snout distance 11.6; rictus���rictus scales 52; scales around tail base 91; scales around tail 1 / 3 distance from base 47; mid-body scales 131; gular scales 70; ventrals from gular fold to hind limb insertion 83; total ventral scales 153; dorsals from tympanum to gular fold 25; dorsals from gular fold to hind limb insertion 77; total dorsals 102; supralabials 52; scales around the neck anterior to gular fold 67; and enlarged supraoculars 7 L/ 6 R. Coloration of holotype in preservative. Dorsal surfaces of body, head and limbs black; head nearly uniform black, with dark brown accents dorsal to narial openings and with white pineal scale; trunk with five transverse rows of white blotches; tail with nine irregular bands comprised of diffuse white speckling; forelimbs with sparse white speckling, becoming more regular ventrally; hindlimbs more speckled than forelimbs, and with more speckling ventrally; dorsal surfaces of manus and pes with few white speckles, but surface of pes with slightly more; terminal scale on digits with conspicuous white spot. Lateral coloration of head consisting of vague, anterior dark brown band extending ventrally from dark brown dorsal markings to narial openings; post-orbitally, lateral surface with longitudinal white stripe extending from the posterior margin of ocular to tympanum (Fig. 9); lateral coloration of the nuchal region consists of bright white mottling, gradually increasing in dorsal extent posteriorly, and terminating at forelimb insertion; lateral portions of trunk with variable white blotches and speckling, with greatest concentration directly posterior to forelimbs, the distribution of which does not correspond to dorsal banding; terminal half of tail solid black dorsally. Ventral coloration predominately yellow-white throughout; gular region with characteristic dark blotches, becoming more prominent posteriorly; anterior portions of gular region with three triangular projections extending medially; scapular region with irregular dark blotches; trunk with irregular dark triangular projections extending medially from margins, and generally not connecting with projections from opposite side; forelimbs and hindlimbs almost uniform white at insertions, but with increasing dark mottling laterally and in distal portions; manus and pes dark gray; venter of tail similar to that of trunk, but with triangular projections connecting and forming vague bands posteriorly; terminal third of tail solid black ventrally. Variation. Our small type series exhibits some color variation but specimens are generally phenotypically similar to one another. Four specimens from Mindoro, including the holotype (KU 308437, 335742��� 44) generally appear darker dorsally, owing to a combination of darker interstitial skin between the scales and fewer whitecolored scales. The holotype (KU 335744) and an additional specimen from Mindoro (KU 308437) were the largest intact specimens examined, and both exhibit a reduction in the size and vibrancy of dorsal banding, with the latter individual having bands that are only faintly discernable. Juvenile specimens from both Mindoro (KU 335742, 335743) and Semirara (KU 305151, 305161, 305163) exhibit much more distinct dorsal bands comprised of series of yellowish to white spots or oscelli. One juvenile from Mindoro (PNM 9799; formerly KU 308403) appears to be an intermediate, with dorsal bands discernable posteriorly on the trunk and becoming less defined anteriorly. One specimen from Semirara (KU 305161) exhibits a faded reticulate pattern interspersed among the more well-defined dorsal bands, owing to clusters of grayish spots comprised of lightly colored scales and interstitial space. Specimens from Semirara (KU 305151, 305161, 305163) and one from Mindoro (KU 335742) exhibit a more well-defined light-colored longitudinal stripe laterally in the nuchal region, which extends from the ventral margin of the tympanum dorso-posteriorly over the forelimb insertions, joining with the first dorsal trunk band to form a vague ���U��� shape when viewed from dorsal perspective. Dorsal nuchal coloration varies from uniform black in the holotype (KU 335774) and three paratypes (PNM 9799, KU 308437, 335772), the presence of 4���10 variable spots or blotches (KU 305151, 308161, 335743), to a more mottled and irregular combination of stipes and blotches posteriorly (KU 305163). One specimen (KU 308437) exhibits a reduction in white speckling on the dorsal surfaces of the limbs. Ventral coloration is moderately variable. The characteristic gular blotches or spots are present in all specimens, though the number and relative size of blotches is somewhat reduced in four specimens (KU 305161, 308403, 335742, 335743). Two specimens (KU 305151, 305163) exhibit much larger and more conspicuous gular spots. Ventral trunk coloration is less variable, with only three specimens (KU 305161, 308403, 308437) exhibiting reduced transverse dark bands, which generally fail to converge medially. Ventral coloration of the limbs is nearly identical among specimens, with only slight variation in the appearance of black color encroaching on the lateral margins of both fore- and hindlimbs, but generally appearing as partial transverse bands. The holotype (KU 335774) and three paratypes (KU 308437, 305151, 305161) exhibit a more irregular pattern on the margins of the ventral limb surfaces, with dark coloration appearing as an almost reticulate pattern. Ventral coloration of the tail is nearly uniform across specimens, with only minor variation in the extent of medial extension of dark bands. In all specimens, dark bands fail to coalesce anteriorly, but posteriorly to the terminus form a distinct alternating black and white pattern (see Figures 6 and 7 for additional information on color pattern variation). Coloration in life. Based on the only available image, KU 335742 (Fig. 8 B), and field notes of the authors. Dorsal ground coloration black, with transverse crossbands of irregularly shaped creamy white to light yellow ocelli from the posterior margin of the nuchal region through the hind limb insertion. Trunk with few, larger ocelli with darkened, ground-colored centers. Additionally, dorso-lateral margins of trunk with smaller, variably distributed light yellow spots or blotches. Lateral margins of head and nuchal region with conspicuous light yellow stripe, extending from the the posterior margin of the ocular, through the tympanum, and midway into the anterior portion of the nuchal region. An additional light yellow stripe extends dorsally and posteriorly from the ventrolateral margin of the nuchal region, and extends to the dorsal region above the forelimb insertion. Snout with distinct and conspicuous bands extending ventrally from dorsal surface. Dorsal limb surfaces colored similarly to body, but with ocelli irregularly arranged and significantly smaller, lacking any dark-colored center. Forelimbs with larger ocelli dorso-medially. Ventral coloration creamy white to light yellow, with scattered black pigmentation and with ocelli of ventrolateral surfaces irregularly arranged and significantly smaller, lacking any dark-colored center (Fig. 8 b). Ecology, Distribution, and Natural History.���As with all members of the Varanus salvator Complex, V. bangonorum should be considered a habitat generalist, often thriving in a range of habitats from primary forest to more urbanized areas. Like V. dalubhasa, no detailed studies have been undertaken on this taxon specifically, but the findings of Gaulke (in Luxmoore & Groombridge 1989, 1992) are likely relevant. This taxon is most likely to be found in mangrove and riparian habitats (as their common name might suggest). Additionally, most water monitors seek shelter in rock crevices or even trees, often near water sources. Not surprisingly, Varanus bangonorum has been found on the islands of Mindoro and Semirara (Figs. 1, 2), both of which are separated by deep water channels that likely have facilitated the isolation of this species and its evolution as a distinct lineage, even during inter-glacial cycles and lowered sea levels (Voris 2000; Brown & Diesmos 2009). Again, the findings of Gaulke (1992) are relevent as far as dietary preferences are concerned, and it can reasonably be assumed that the diet of Varanus bangonorum consists primarily of invertebrates and smaller vertebrates (fish, rodents, frogs, snakes). Additionally, as with any species of water monitor, this species is attracted to a number of carion items, with a particular affinity for fish (LJW pers. obs.)., Published as part of Welton, Luke J., Travers, Scott L., Siler, Cameron D. & Brown, Rafe M., 2014, Integrative taxonomy and phylogeny-based species delimitation of Philippine water monitor lizards (Varanus salvator Complex) with descriptions of two new cryptic species, pp. 201-227 in Zootaxa 3881 (3) on pages 216-221, DOI: 10.11646/zootaxa.3881.3.1, http://zenodo.org/record/226203, {"references":["Tweddell, C. E. (1970) The identity and distribution of the Mangyan tribes of Mindoro, Philippines. Anthropological Linguistics, 12 (6), 189 - 207.","Welton, L. J., Siler, C. D. Bennet, D., Diesmos, A. C., Duya, M. R., Dugay, R., Rico, E. L., van Weerd, M. & Brown, R. M. (2010) A spectacular new Philippine monitor lizard reveals a hidden biogeographic boundary and a novel flagship species for conservation. Biology Letters, 6, 654 - 658.","Luxmoore, R. & Groombridge, B. (1989) Asian monitor lizards: a review of distribution, status, exploitation and trade in four selected species. Report to the Cites Secretariat, World Conservation Monitoring Centre, Cambridge.","Voris, H. K. (2000) Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems, time durations. Journal of Biogeography, 27, 1153 - 1167.","Brown, R. M. & Diesmos, A. C. (2009) Philippines, Biology, In: Gillespie, R. & Clague, D. (Eds.), Encyclopedia of Islands. University of California Press, Berkeley, pp. 723 - 732.","Gaulke, M. (1992) Taxonomy and biology of Philippine water monitors (Varanus salvator). The Philippine Journal of Science 121 (4), 345 - 364."]}

Varanus dalubhasa sp. nov. Figs. 3, 5���8 Holotype. PNM 9796 (formerly University of Kansas Biodiversity Institute [KU] 306603; Field No. CWL 521), adult male, collected by Charles W. Linkem and CDS, 0 8 July 2006, 18 m above sea level (14.03202, 122.34143; WGS- 84), Barangay Madlangdungan, Municipality of Calauag, Quezon Province, Luzon Island, Philippines. Paratopotypes. KU 305155 (CDS Field No. 2202), adult male; PNM 9797 (formerly KU 306601; CWL Field No. 440), juvenile; KU 306602 (CWL Field No. 520), adult male. Paratypes. KU 308216 (CDS Field No. 2298), juvenile, collected 22 February 2007, Barangay Buenavista, Municipality of Bato, Catanduanes Province, Catanduanes Island, Philippines; KU 313880 (RMB Field No. 9910), adult male, collected 0 1 July 2008, Barangay Tulay Na Lupa, Municipality of Labo, Camarines Norte Province, Luzon Island; KU 326702 and 326703 (LJW Field No. 0 0 75, 0077), adult males, salvaged 3 August 2009, Polillo Island (exact locality information unknown). Other Material. See Specimens Examined Section. Etymology. The specific epithet, dalubhasa, is derived from the Tagolog word ��� dalubhasa ��� meaning a person who has authoritative and comprehensive knowledge of a particular area, or a skilled expert in a particular subject. We choose this term in honor of Vicente ���Enteng��� Yngente of Polillo Island, whose extensive knowledge of natural history and ecology of Philippine reptiles (particularly monitor lizards) has been instrumental to our research and conservation work. Suggested common name: Enteng���s Monitor Lizard. Diagnosis. Varanus dalubhasa can be distinguished from congeners by (1) small, dark speckling and variable transverse banding through the gular region (Fig. 3); (2) number of scales around the tail at 1 / 3 distance from the base; (3) number of gular scales; (4) number of dorsal scales in nuchal region; (5) total number of dorsal scales; (6) number of scales around the neck anterior to the gular fold; and (7) phylogenetic placement sister to V. nuchalis. Additionally, this distinct lineage is biogeographically circumscribed in the Bicol Peninsula faunal sub-region, a distinct geological component of greater Luzon Island which remained inuslar until 3 Ma (Hall 2002). Comparisons. The new species in phenotypically nearly indistinguishable from Varanus marmoratus, but can generally be diagnosed by the presence of small, dark speckling in the gular region and faint anterior transverse bands (versus speckling and distinct anterior transverse bands; Fig. 3), and its allopatric distribution in the Bicol faunal sub-region (versus the remaining portions of Luzon and Lubang islands, and the Batanes and Babuyan island groups). Additionally, although V. dalubhasa is phenotypically similar to V. marmoratus, available data suggest that it is not most closely related to this species (Fig. 2). Varanus dalubhasa can be conveniently distinguished from its allopatric sister taxon, V. nuchalis (distributed in the West Visayan islands of Negros, Panay, Guimaras, Masbate, and the Romblon Province islands of Sibuyan, Tablas, and Romblon; Fig. 1, 2), by the presence of more scales around the tail at 1 / 3 distance from the base (mean = 53 �� 4, versus 46 �� 3), fewer gular scales (mean = 80 �� 3, versus 73 �� 3), and more dorsal scales from the tympanum to the gular fold (mean = 27 �� 2, versus 23 �� 2). Of the remaining, geographically proximate species of the V. salvator Complex, this new species can be distinguished from V. palawanensis by having fewer dorsal scales from the tympanum to the gular fold (mean = 27 �� 2, versus 32 �� 3), fewer dorsal scales from the gular fold to the hind limb insertion (mean = 84 �� 5, versus 96 �� 3), fewer total dorsals scales longitudinally (mean = 111 �� 5, versus 128 �� 3), and fewer scales around the neck anterior to the gular fold (mean = 78 �� 3, versus 87 �� 4). Additionally, V. dalubhasa can be distinguished from the allopatric Mindoro lineage by having variable dark speckling and transverse bands in the gular region (versus distinct spotting; Fig. 3). Lastly, V. dalubhasa is comprised of entirely distinct haplotypes or haplotype networks, relative to all other members of the V. salvator Complex (see Welton et al., 2010 a). We have constrained our morphological analyses to the geographically most proximate taxa due to previous studies (Koch et al. 2007, 2010 b) demonstrating their distinctiveness relative to the remaining diversity withing the V. salvator Complex. Description of holotype. An adult male (Fig. 5), hemipenes everted; SVL 505.0 mm; tail 657 mm; head relatively slender, length 89.1 mm, width 51.0 mm, and depth at eye 32.0 mm; snout length 47.6 mm, 53.4% head length; snout rounded anteriorly; narial openings 5.4 mm, ovular, posteriorly elevated, encircled by 9 L/ 9 R small polygonal scales; snout elevated sharply just anterior to narial region, then continuing gradually to ocular region; narial region not distinctly elevated above horizon of snout; cranial table squarish, only slightly wider than long. Head scales ovular to polygonal (Fig. 5), heterogeneous in size with the smallest occurring dorso-laterally between ocular opening and tympanum, and largest ocurring dorso-medially from tip of snout to conspicuous parietal; supraoculars 6 L/ 6 R, subrectangular and elongated transversely; supralabials 59, slightly larger than lateral head scales; infralabials 61, smaller than both supralabials and nearest rows of gular scales; both supra- and infralabials increasing in size anteriorly; nuchals large, circular, increasing in size to forelimb insertion along both dorsal and lateral surfaces; lateral nuchals less than half the size of dorsals; scales immediately posterior to cranial table quite small, granular; sutures between scales slightly larger in nuchal region than on head; nuchals 31, in semi-regular transverse rows from posterior margin of tympanum to forelimb insertion; dorsal trunk scales arranged in 88 semi-regular transverse rows from fore- to hindlimb insertions, ovular, slightly smaller than nuchals, gradually decreasing in size to hindlimb insertion as well as laterally, with smallest scales occuring laterally adjecent to limb insertions; midbody scales 138; axilla���groin distance 245 mm; dorsal limb scales ovular, decreasing in size distally to digits; fore- and hindlimbs 92 and 107 mm, respectively, 18.2% and 21.2% snout���vent length, 32.6 and 53.9 mm diameter at insertions; limb scales present in semiregular rows; scales of manus and pes smaller and more irregular in shape, ranging from circular to nearly rectangular dorsally; supradigitals rectangular dorsally, ovular laterally, with terminal scale nearly twice as large as others; digits terminate in long, slender recurved claws; caudals ovular and granular at insertion, becoming rectangular posteriorly; dorsal keel composed of paired, posteriorly elevated scales commenses 65 mm posterior to hindlimb insertion, and continues to tail terminus with paired scales gradually decreasing in size; caudal scales moderately keeled, in regular transverse and longitudinal rows laterally, decreasing in size posteriorly. Ventrals less variable than dorsals; gular scales generally rectangular anteriorly, becoming more ovular towards gular fold; smallest scales medially in anterior third of gular region, with largest scales just anterior to gular fold but followed posteriorly by three transverse rows of smaller, more granular gulars; scales around the neck anterior to the gular fold 78, mid-gular 87; gular scales in 85 irregular rows from tip of snout to gular fold; ventrals of fore- and hindlimbs circular to polygonal, with those of forelimbs slightly smaller than those of hindlimbs; precloacals homogenous in size and shape with those of hindlimbs; ventral trunk scales from gular fold to hindlimb insertion 85, ovular anteriorly, becoming rectangular posteriorly, and present in semi-regular rows; scales decreasing in size laterally and at hindlimb insertions; ventral caudal scales rectangular, smallest just posterior to cloacal opening; scale surface slightly rounded anteriorly, becoming sharply keeled and more longitudinally elongate posteriorly. Right and left hemipenes partially everted; maximum length 56.2 mm, with proximal cylindrical section extending 26.2 mm; bifurcation resulting in primary cylindrical apical portion with external flounces, and more diminutive portion resembling an ���ear��� or reduced fold; flounces present in irregular rows, totalling 15 anterior to sulcus spermaticus, 6 posteriorly; lacking notable external appearance of ossified hemibaculum. Measurements of holotype (in mm). SVL 505; tail length 657; head length 89.1, width 51.0, depth at eye 32.0; eye���naris distance 33.7; naris���snout distance 14.9; rictus���rictus scales 60; scales around tail base 103; scales around tail 1 / 3 distance form base 59; mid-body scales 138; gular scales 85; ventrals from gular fold to hind limb insertion 85; total ventral scales 170; dorsals from tympanum to gular fold 31; dorsals from gular fold to hind limb insertion 88; total dorsals 119; supralabials 59; scales around the neck anterior to gular fold 78; and enlarged supraoculars 6 L/ 6 R. Coloration of holotype in preservative. Dorsal surfaces of head, body, and limbs black; aggregation of yellow-gold scales present dorsoanteriorly to narial openings, along canthal ridge just anterior to oculars, and irregularly dispersed throughout cranial table; trunk with seven transverse rows comprised of yellow-gold, indistinct ocelli; tail with nine, slightly more distinct yellow-gold bands, becoming more diffuse or speckled laterally; forelimbs with irregularly distributed yellow-gold scales, density increasing distally and along lateral surfaces; hindlimbs with irregular agreggations of ocelli consisting of only two or three yellow-gold scales (Fig. 5). Lateral coloration of head consisting of partial, near vertical yellow-gold bands at the tip of snout, posterior margin of narial opening, and two between the narial opening and the ocular; posterior band with ventral widening oriented more posteriorly; post-orbitally, a yellow-gold stipe extends to tympanum, and a yellow-gold spot consisting of four or five scales appears ventral to the anterior margin of the stripe; lateral coloration of the nuchal region consists of 2 L/ 3 R faint spots directly posterior to the tympanum; ventral margins of nuchal region with yellow-gold variable demarcation, extending more dorsally along the posterior margin and appearing to wrap around the dorsal margin of the fore-limb insertions; trunk with variable yellow-gold spotting directly posterior to forelimbs, fading to solid black posteriorly (Fig. 5). Ventral coloration predominately yellow-gold throughout; gular region with irregular spots, faint anteriorly but becoming prominent and dark along lateral margins just anterior to gular fold; scapular region with two irregular black bands; trunk with irregular black bands and spots anteriorly, becoming more broken medially and posteriorly, appearing as laterally elongated yellow-gold ovals; forelimbs with faint, irregular dark mottling; hindlimbs with near uniform venter medially, with dark mottling along lateral margins; manus and pes dark yellow-gray; venter of tail appearing anteriorly much like that of the trunk, with yellow-gold laterally elongated ovals; posterior portions of tail with more regular alternation of dark and yellow-gold bands (Fig. 5). Variation. Our small type series exhibits some color variation, but specimens are generally phenotypically similar. The holotype (KU 306603) and three paratypes (KU 305155, 306602, and 308126) have lighter interstitial coloration between the dorsal scales, yielding an overall lighter appearance than other specimens. These three specimens also exhibit much more diffuse yellow gold coloration on limbs and tail, while two specimens (KU 313880, 326702) exhibit much more distinct yellow-gold spots or ocelli throughout. Additionally, the yellow-gold coloration in the lateral portions of the nuchal region, among all specimens, and on either side of a single specimen, are present as either a single longitudinal stripe extending posteriorly from the posterior margin of the tympanum or as a series of two or three longitudinally elongated spots or blotches. Two specimens (KU 305155, 306602) exhibit two relatively well-defined yellow crossbands on the snout, with the former���s being even more pronounced than the latter, and with this specimen (KU 305155) also exhibiting a much larger proportion of yellow coloration in the supraocular and temporal regions. In contrast, two specimens (KU 308216, 313880) exhibit nearly uniform, dark dorsal head coloration, with the former being completely devoid of yellow-gold accents with the exception of the pineal scale. Ventral coloration is only slightly more variable than that of the dorsum. All specimens exhibit variable speckling in the gular region, gernerally consisting of 1���4 black scales. In the holotype (KU 306603), this speckling is much more reduced. In one specimen (KU 305155) anterior speckling in the gular region gives way to 2 broken transverse bands, and somewhat reduced speckling through the remainder of the gular region. Three specimens (KU 306602, 308216, and 326702) exhibit four, more well-defined dark transverse bands in the anterior portion of the gular regions, with speckling throughout the remainder of the gular region being relatively dense. A single specimen (KU 313880) exhibits nearly complete dark transverse bands throughout the gular region, with less prominent bands present anteriorly. Ventral trunk coloration is nearly uniform among specimens, with the only notable variation present in two individuals (KU 305155, 326702), the first of which exhibits reduced dark coloration extending medially onto the abdomen, and the second of which with darker, more distinct coloration extending medially onto the abdomen. Ventral tail coloration lacks any notable variation among specimens (see Figures 6 and 7 for further information on color pattern variation). Coloration in life. Based on the only available image, PNM 9797 (formerly KU 306601; Fig. 8 A), and field notes of the authors. Dorsal ground coloration black to dark gray, with consipcuous transverse cross bands of yellow-gold ocelli from the posterior margin of the nuchal region through the anterior third of the tail. Trunk ocelli larger with a darkened, ground-colored center. Additionally, dorsal trunk with faded reticulate pattern, owing to light brown interstitial coloration. Lateral margins of head and nuchal region with conspicuous yellow-gold stripe, extending from the the posterior margin of the ocular, through the tympanum, and midway into the nuchal region. Yellow-gold ocelli brightest at lateral margins of trunk and adjacent to hindlimb and tail insertions. Dorsal limb surfaces colored similarly to body, but with ocelli irregularly arranged and significantly smaller, lacking any dark-colored center. Dorsal ground color of limbs bright yellow, with irregular dark pigmentation. Ventral coloration bright yellow-gold, with scattered black pigmentation and with ocelli of ventro-lateral surfaces irregularly arranged and significantly smaller, lacking any dark-colored center (Fig. 8 A). Ecology, Distribution, and Natural History. As with all members of the Varanus salvator Complex, V. dalubhasa should be considered a habitat generalist, often thriving in a range of habitats from primary forest to more urbanized areas. While no detailed studies have been undertaken on this taxon specifically, the findings of Gaulke (in Luxmoore & Groombridge 1989, 1992) are likely relevant. This taxon is most likely to be found in mangrove and riparian habitats (as their common name might suggest). Additionally, most water monitors seek out shelters in the form of rock crevices or even trees, most often near water sources. In more urban areas, human habitation may be used in lieu of more ���natural��� shelters (Gaulke 1992). Not surprisingly, Varanus dalubhasa has been found throughout the Bicol Peninsula and faunal sub-region, including genetically confirmed specimen identifications from Polillo and Catanduanes islands (Figs. 1, 2). This is an area with significant agricultural (rice fields) and coastal environments, and is additionally subdivided by a number of moderately sized river drainages. The northern extent of distribution for this taxon, and potential contact zone with V. marmoratus requires further study. Although no dietary preferences have been observed for this taxon, one can reasonably assume that its diet consists primarily of invertebrates and smaller vertebrates (fish, rodents, frogs, snakes). Additionally, this species is attracted to a number of carion items, with a particular affinity for fish (LJW pers. obs.)., Published as part of Welton, Luke J., Travers, Scott L., Siler, Cameron D. & Brown, Rafe M., 2014, Integrative taxonomy and phylogeny-based species delimitation of Philippine water monitor lizards (Varanus salvator Complex) with descriptions of two new cryptic species, pp. 201-227 in Zootaxa 3881 (3) on pages 212-215, DOI: 10.11646/zootaxa.3881.3.1, http://zenodo.org/record/226203, {"references":["Hall, R. (2002) Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based reconstructions, model and animations. Journal of Asian Earth Sciences, 20, 353 - 431.","Welton, L. J., Siler, C. D. Bennet, D., Diesmos, A. C., Duya, M. R., Dugay, R., Rico, E. L., van Weerd, M. & Brown, R. M. (2010) A spectacular new Philippine monitor lizard reveals a hidden biogeographic boundary and a novel flagship species for conservation. Biology Letters, 6, 654 - 658.","Luxmoore, R. & Groombridge, B. (1989) Asian monitor lizards: a review of distribution, status, exploitation and trade in four selected species. Report to the Cites Secretariat, World Conservation Monitoring Centre, Cambridge.","Gaulke, M. (1992) Taxonomy and biology of Philippine water monitors (Varanus salvator). The Philippine Journal of Science 121 (4), 345 - 364."]}

Lygodactylus regulus sp. nov. Prince Dwarf Gecko (Figs. 6���8) Holotype. MVZ 266138, Adult male, Mozambique, Zamb��zia Province, Mt. Namuli (15 �� 23 ' 15 "S, 37 ��04' 24 "E, 1281 m a.s.l.), collected by D. M. Portik, 10 August 2011 (Fig. 6). Paratypes. Three specimens: MVZ 266137, adult male, Mozambique, Zamb��zia Province, Mt. Namuli (15 �� 22 ��� 28 ���S, 37 ��03��� 54 ���E, 1421 m a.s.l.), collected by D. M. Portik, 10 August 2011 (Fig. 7); PEM R 14922, adult female, Mozambique, Zamb��zia Province, Mt. Namuli, Ridge leading to Marakuni Ridge (1537 AC, 1800 m a.s.l.), collected by P. Ryan, 30 November 1998; PEM R 20277, adult female, Mozambique, Zamb��zia Province, Mt. Namuli, Muretha Plateau (15 �� 23 ���04.5���S, 37 ��03��� 13.6 ���E, 1878 m a.s.l.), collected by J. Bayliss, 27 May 2007 (Fig. 8). Etymology. The specific epithet is derived from Latin regulus = petty king, prince. The closest relative of this species is Lygodactylus rex, the King Dwarf Gecko or Regal Dwarf Gecko, which exhibits a larger body size and inhabits nearby Mt. Mulanje, Malawi, an overall larger massif than Mt. Namuli, Mozambique. Given the smaller body size, smaller size of the massif inhabited, and close relationship with the King Dwarf Gecko, we recommend the common name Prince Dwarf Gecko for this species. Diagnosis. Lygodactylus regulus sp. nov. differs from all other species in the genus Lygodactylus, except for L. angularis, L. picturatus, and L. rex, in its larger body size (males 38���39 mm SVL, females 38���40 mm SVL). Lygodactylus regulus sp. nov. differs from both L. angularis and L. picturatus by having a mental with shallow lateral fissures (mental entire in L. angularis and L. picturatus), and by having a conspicuous ocellus above the shoulder, a trait shared with L. rex. From the latter it can be distinguished by having an overall smaller body size (L. regulus sp. nov.: males 38���39 mm SVL, females 38���40 mm SVL; L. rex: males 47���55 mm SVL, females 39���46 mm SVL), often having a condition of three postmentals rather than two postmentals (L. regulus sp. nov.: 75 %, n = 4; L. rex: 37.5 %, n = 16), males possessing a greater number of precloacal pores (L. regulus sp. nov.: 12���13; L. rex: 8���11), having more variability in the condition of contact of first infralabial with the postmental or postpostmentals (L. regulus sp. nov.: ranging from contact with postmental only to L. rex: ranging from p -distance values between L. regulus sp. nov. and L. rex being 12.9 % (ND 2), 1.2 % (MXRA 5), and 3.0% (RAG- 1). Finally, the two species are geographically well separated by over 160 km. Description of holotype. MVZ 266138; Adult male; SVL: 38.9 mm; TrunkL: 18.5 mm; CrusL: 5.9 mm; TaiIL: 49.2 mm; TaiIW: 4.7 mm; HeadL: 11.2 mm; HeadW: 7.0 mm; HeadH: 4.5 mm; EarL: 0.9 mm; ForeaL: 5.3 mm; OrbD: 1.9 mm; NarEye: 3.5 mm; SnEye: 4.7 mm; EyeEar: 3.2 mm; Internar: 2.0 mm; Interorb: 4.1 mm. Build moderate; head very broad and distinct from neck. Snout long and broad, the distance from the tip to anterior border of the eye slightly greater than the interorbital distance anteriorly, greater than the distance between eye and ear opening. Snout covered with heterogeneous flattened granular scales, medium-sized on crown of head and middle of snout, becoming smaller laterally above eye, and larger anteriorly on snout, reaching largest size in loreal region. Scales on the snout larger than those on the occiput, which are, in turn, much larger than the small convex dorsal granules. Canthus rostralis not prominent. Rostral pentagonal, wider than high. Nostril oval and directed slightly posteriorly, bordered by rostral, first supralabial, supranasal, and two postnasals; supranasals separated by a single small internasal granule. Spiny superciliaries present. Mental heptaganol with shallow lateral fissures, followed by two large postmentals. Scales behind postmentals enlarged, decreasing in size towards base of throat, then increasing in size on chest and belly. Supralabials 8 - 8; infralabials 8 - 8. Gulars imbricate, 26 between posteriormost infralabials. Dorsal granular scales roughly homogeneous, slightly larger on ventrolateral portions of trunk. Limbs well developed but short, pentadactyl, 1 st digit greatly reduced and lacking claw. Distal portions of remaining digits expanded, bearing claw. Six pairs of lamellae under fourth finger; 8 enlarged subdigital scales occur proximally to manual lamellae; six pairs of lamellae beneath fourth toe. Granular scales on dorsal surface of limbs roughly homogeneous and similar size to those on trunk; scales on ventral surface of thighs subhexagonal, subimbricate, and same size as those on trunk; scales on ventral surface of arms subhexagonal, subimbricate, and smaller in size than those on trunk. Precloacal pores 12, arranged in a chevron roughly 6 scale rows anterior to cloaca posteriorly. Three rows of 4���6 enlarged, glandular scales (��-glands) present along the ventral surface of thighs. Tail shape cylindrical, caudal scales dorsally semi-flattened, imbricate, largely homogeneous; subcaudal scales large, flattened, subhexagonal, subimbricate, and arranged in a single broad row. Further measurements are recorded in Tables 4 and 5. Coloration in ethanol. Above, a more-or-less uniform cinnamon brown background coloration, with a broad continuous kingfisher rufous lateral stripe extending from the shoulder to the base of the tail (Fig. 6 B). Anterior to the shoulder the stripe breaks and continues irregularly to the posterior border of the eye, narrowing substantially in width. Just above the shoulder is a large black patch with a small white spot in the center. Dorsum of head anterior to crown is uniform burnt umber, throat is sulphur yellow with three pairs of irregular black stripes approximately two granules wide which converge from the labials towards the chest, with the two most distal pairs meeting and the most proximal pair remaining separated by three granules. Dorsal surfaces of limbs cinnamon brown, with irregular lighter markings. Body venter smoky white with no markings or speckling. Gular region suffused with pale sulphur yellow and bearing medium neutral gray markings (Fig. 6 C, yellow lacking in paratype MVZ 266137 ��� Fig. 7 D). Dorsum of tail cinnamon brown with lighter chevron pattern (kingfisher rufous to pale pinkish buff), laterally exhibiting 19 cinnamon brown rectangular markings separated by kingfisher rufous background with minute pale pinkish buff spots, ventrally smoky white suffused with grey posteriorly. Measurements recorded to the nearest 0.1 millimeter. Note: Paratype PEM R 20277 has a damaged head and several measurements were unable to be recorded. Museum abbreviations: MVZ, Museum of Vertebrate Zoology, University of California, Berkeley; PEM, Port Elizabeth Museum. Coloration in life. Recently euthanized specimens prior to fixation (Figs. 6 A, 7 A) are smoke gray to cinnamon-drab in background coloration with darker brown to black markings and a paler thick dorsolateral stripe. In the holotype this stripe contains a series of well-defined circular to oval markings ranging from cinnamon-drab anteriorly to cream white posteriorly. The ocellus anterior to the forelimb insertion is bright white with a black border. In paratype MVZ 266137 the pale dorsolateral stripe extends to the posterior border of the orbit. Paratype PEM R 20277 (Fig. 8) exhibits a much more mottled pattern than seen in other members of the type series or in L. rex. This is dominated by brick red and dusky brown to smoky white dorsal markings, including a series of light lateral blotches. The ocellus near the limb insertion is bright white with a blackish surround. Venter of throat and chest bright white; abdomen, thighs and tail venter yellow (between pale greenish yellow and sulphur yellow) with area of ��-glands trogon yellow (based on MVZ 266137, Fig. 7 B). The iris is crimson. Paratype variation. MVZ 266137: Nostril bordered by rostral, first supralabial, supranasal, and one postnasal; mental hexaganol with shallow lateral fissures, followed by three postmentals (two large, one small); supralabials 7 - 7; infralabials 7 - 7; precloacal pores 13; gulars between posteriormost infralabials 28. Color in ethanol: Broad kingfisher rufous lateral stripe extends unbroken from base of tail to posterior border of eye, narrowing significantly in width approaching eye. Dorsum of anterior part of head is uniform burnt umber, with 3 to 4 minute cream spots along canthus rostralis. Throat is white with three pairs of irregular black stripes that converge from the infralabials towards the chest. Lateral dark rectangles on tail less conspicuous, but a line of minute pale pinkish buff spots still discernable. PEM R 14922: Five pairs of lamellae under 4 th finger; mental with shallow lateral fissures, followed by three postmentals (two large, one small); supralabials 7 - 7; infralabials 8 - 8. PEM R 20277: Five pairs of lamellae under 4 th toe; mental with shallow lateral fissures, followed by three postmentals (two large, one small). Additional differences in morphological measurements are recorded in Tables 4 and 5. Size. Largest male (MVZ 266138) 38.8 + 49.2 = 88.0 mm; largest female (PEM R 14922) 40.7 + 38.0 = 78.7 mm (tail truncated). Distribution and habitat. The holotype (MVZ 266138) was collected from the rafters of a village hut, similar to the structure appearing in the foreground of Fig. 9 A. The immediate surrounding vegetation had been cleared for agricultural plots, however a riparian forest strip was approximately 40 m west of the plot. This limited riparian forest followed the flowing Nanchili drainage below Nanchili Falls, and during a visual survey no specimens were located in this habitat. The paratype, MVZ 266137, was found basking on wooden debris adjacent to a local residence, surrounded by cultivated plots. The paratype PEM R 14922 was collected on rock in an open grassy area west of Ukalini Forest on the southern side of Mt. Namuli (15 �� 22 ���S, 37 ��03���E, 1800 m a.s.l.) (Fig. 9 B, C). This area consists of a large granite whaleback with a thin layer of peaty soil that dries seasonally, thereby exfoliating and exposing the rock below (similar to Fig. 9 C). Branch and Ryan (2001) report another individual (presumably L. regulus sp. nov.) was seen on a dead tree stump in similar habitat bordering the forest edge at the southern flank of Peseni, a peak 4 km southwest of Mt. Namuli (15 �� 23 ���S, 37 ��02���E). Another paratype (PEM R 20277) was collected on the Muretha plateau on an exfoliating rock flake (15 �� 23 ���04.5���S, 37 ��03��� 13.6 ���E, 1878 m a.s.l.). Based on the limited number of specimens and observations, Lygodactylus regulus sp. nov. has been found in open grassy habitat between 1281���1878 m a.s.l. and utilizes a variety of basking sites, including exposed rock, tree stumps, woody debris, and artificial habitat. The exposed granite domes forming the main peaks throughout the Namuli massif may also serve as suitable habitat, provided some retreat is available. Lygodactylus regulus sp. nov. has not been recorded in Ukalini Forest, which consists of Afromontane forest vegetation, despite concerted attempts to locate it. All reports indicate this species does not require montane forest or mid-elevation forest habitat, and is often found adjacent to forest habitats. This contrasts with L. rex which is known only from midelevation forest habitats (Lukubula Forest Station, 825 m a.s.l.; Ruo Gorge riverine forest along Lujeri River, 995 m a.s.l.), except for a population on Lichenya Hut on Mt. Mulanje (1858 m a.s.l.). There are no other records of the species on the summit of Mt Mulanje, indicating that the Lichenya Hut population is possibly introduced as the hut is the largest on the mountain and was constructed in the 1920 ���s for use as a ���hill station��� allowing families from the tea estates to escape the summer heat (Eastwood 1988). The species has only been observed on trees or buildings, and has not been observed on rocks. However, a series of L. rex eggs were found in a crumbling rock crack sheltered by forest trees in Ruo Gorge. The broader Namuli massif covers an area of roughly 200 km 2 above 1200 m, with an estimated 300 ha of higher elevation grasslands (Timberlake et al. 2009), and it is likely that L. regulus sp. nov. is dispersed throughout this habitat. A survey at the nearby lower elevation town of Guru�� produced only L. capensis, which seems to replace L. regulus sp. nov. at elevations below 800 m (Portik et al. 2013)., Published as part of Portik, Daniel M., Travers, Scott L., Bauer, Aaron M. & Branch, William R., 2013, A new species of Lygodactylus (Squamata: Gekkonidae) endemic to Mount Namuli, an isolated ' sky island' of northern Mozambique, pp. 415-435 in Zootaxa 3710 (5) on pages 423-431, DOI: 10.11646/zootaxa.3710.5.2, http://zenodo.org/record/248924

Portik, Daniel M., Travers, Scott L., Bauer, Aaron M., Branch, William R. (2013): A new species of Lygodactylus (Squamata: Gekkonidae) endemic to Mount Namuli, an isolated ' sky island' of northern Mozambique. Zootaxa 3710 (5): 415-435, DOI: http://dx.doi.org/10.11646/zootaxa.3710.5.2

Oedipina nica sp. nov. Nicaraguan Highland Worm Salamander Figure 1 Oedipina cyclocauda (in part). K��hler 1999, 2001. Holotype. MVZ 263774, an adult male, from El Gobiado, Reserva Natural Cerro Datanl��-El Diablo, 13 ��09���N, 85 �� 52 ���W, 1420 m above sea level (a.s.l.), Dept. Jinotega, Nicaragua, collected 30 August 2009 by S. M. Rovito, T. J. Papenfuss, J. Sunyer, and L. A. Obando (original field number SMR 842). Paratypes (5). UF 156445, from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156446 ��� 47, from Reserva Natural Cerro Kilamb��, 13 �� 35 ���N, 85 �� 43 ���W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156453 ��� 54, from Reserva Natural Macizos de Pe��as Blancas, 13 �� 17 ���N, 85 �� 43 ���W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Referred specimens (12). SMF 78736, from Camp El Hielo, Reserva Natural Cerro Kilamb��, 1490 m a.s.l., Dept. Jinotega, Nicaragua; SMF 78737, 78739��� 40, from Camp 2, Reserva Natural Cerro Kilamb��, 13 �� 35.25 'N, 85 �� 41.50 'W, 1360 m a.s.l., Dept. Jinotega, Nicaragua; UF 156443 ��� 44, from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156448 ��� 50, from Reserva Natural Cerro Kilamb��, 13 �� 35 ���N, 85 �� 43 ���W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156451 ��� 52, two tails dropped by escaped adults from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1600 m a.s.l., Dept. Jinotega, Nicaragua; UF 156455 from Reserva Natural Macizos de Pe��as Blancas, 13 �� 17 ���N, 85 �� 43 ���W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Diagnosis. A slender, small-sized (largest specimen 48.5 mm SL) species assigned to the genus Oedipina based on the presence of a high number (19���20) of costal grooves between the short limbs, very narrow manus and pes, long tail (tail length ca 2 x SL), overall morphological similarity to other Oedipina species, and molecular phylogenetic evidence. This species is a member of the subgenus Oeditriton (based on molecular data; Fig. 2) and is distinguished from the only other members of that clade (O. kasios and O. quadra) by its more slender habitus and fewer vomerine teeth. Further distinguished from other Nicaraguan members of the genus Oedipina as follows: from O. collaris by being much smaller and more slender, having short limbs and narrow manus and pes, and having a short, rather bluntly tipped snout; from O. cyclocauda by being smaller and more slender without nearly pad-like feet; from O. pseudouniformis by being more slender with shorter, less robust limbs and narrower manus and pes, and by having almost uniformly dark brownishblack coloration with no distinct light pigmentation on the upper parts of the proximal segment of the limbs. Description of the holotype. The male holotype is 39.2 mm SL and is judged to be an adult by the presence of small papillae in the anterior margins of the vent and a small mental gland behind the anterior tip of the mandible. The small head is rounded in dorsal profile, and somewhat flattened in lateral profile with a relatively blunt snout and distinct nasolabial protuberances. Snout to posterior angle of vent 9.8 times head width and 6.5 times head length. Nostrils are small but conspicuous under magnification. The small eyes are inconspicuous and do not extend beyond the lateral margins of the head. The suborbital groove does not intercept the lip line. The skin of the dorsal surface of the head behind the eyes is depressed in a pattern that suggests the presence of a frontoparietal fenestra. There is a single slightly enlarged premaxillary tooth that is located distinctly anterior to the maxillary teeth. Maxillary teeth total 41. The small vomerine teeth are in a short, fairly straight series and total 11. There are 20 costal grooves between the small limbs with a limb interval of 13 intercostal folds. Hands and feet are tiny, narrow, and elongated. Digit I is fused with digit II and digit IV is fused with digit III on the forelimbs, with only a part of a phalangeal segment of digit III free between digits II���III on the forelimbs. Digit I is fused with digit II and digit V is fused with digit IV on the hind limbs; both outer digits are extremely short on the hind limbs. The triangular tip of digit III of the hind limb is distinct from the tiny rounded tips of digits II and IV, but less than a full phalangeal segment of that digit is free of the others. Subdigital pads are nearly imperceptible. Digits on forelimbs in order of decreasing length are III���II���IV���I; digits on hind limbs are III���II���IV���(V���I). The tail is round in cross section and tapers gradually to a narrow tip; the tail is almost twice SL. Measurements (in millimeters) of the holotype. Head width 4.0; snout to gular fold (head length) 6.0; head depth at posterior angle of jaw 1.8; eyelid width 1.1; eyelid length 1.5; eye to nostril 0.6; anterior rim of eye to snout 1.2; horizontal orbital diameter 1.0; interorbital distance 1.7; distance separating eyelids 0.7; nostril diameter 0.2; snout projecting beyond mandible 0.4; distance from eye to distal end of postorbital groove 1.9; snout to posterior angle of vent (SL) 39.2; snout to anterior angle of vent 37.0; snout to forelimb 9.4; axilla to groin 26.0; limb interval 13; shoulder width 2.7; tail length 77.0; tail width at base 2.4; tail depth at base 2.4; forelimb length (to tip of longest digit) 3.7; hind limb length (to tip of longest digit) 4.5; forelimb foot width 0.6; hind limb foot width 0.9; free length of longest digit 0.2. Coloration of the holotype in alcohol. Dorsal and lateral surfaces of the head, body, and tail are black. Costal grooves on the body and tail are the same color as the body or slightly lighter dark gray, but at the deepest part of the groove pigment is lacking so the grooves are prominent. Ventral surfaces of the head, body, and tail are slightly paler black than those dorsal surfaces. Dorsal and ventral surfaces of the limbs are the same as for the body. Tiny white speckles are most visible behind the eye and over the shoulder, then in an irregular, narrow line along the dorsolateral part of the body to the hind limbs, but they are few in number and obscure. White spots are a little larger in the gular and chest areas ventrally. The nasolabial protuberances are unpigmented and contrast with the surrounding tissue. Variation. Variation based on study of the holotype and three male (UF 156445, 156447, 156453), one female (UF 156446), and one juvenile (UF 156454) paratypes: the four males, judged to be mature based on having inconspicuous mental glands and anterior vent margins bearing fine pinnae, range from 38.5���44.5 SL; the female is 48.5 SL. Tails are present in the holotype, three male paratypes, and the juvenile and all taper gradually to slender tips. SL is from 0.86 (adult) to 0.52 (holotype) times tail length. SL in males is 9.8���11.8 times head width and 9.9 in the single female. Limbs are relatively short; number of costal folds between adpressed limbs is 13���14 in males and 13 in the female. There are 19���20 costal grooves between the limbs in the males and 20 in the female. Feet are narrow; 0.8���1.1. The digits are joined to their neighbors, with only the tips of the longest toes free. Males have 1���2 premaxillary teeth, the female has 4. Maxillary teeth range from 41���47 in males; 48 in the female. Males have 9���16 vomerine teeth; the female 18. If one adds the referred specimens, the size range of the adult males increases to 35.3���48.5 SL, and that of females from 37.5���48.5 SL. These specimens are consistent in measurements, tooth and costal groove counts, and coloration with the type series. SMF 78738, from Reserva Natural Cerro Kilamb�� (44.0 SL), fits within the range of the new species in all measurements (see discussion). Variation of coloration based on color photographs taken in life during daytime. Coloration is in general uniformly dark brownish-black with no distinguishing features. The upper side of the proximal segment of the limbs is the same color. The costal grooves are very evident because of the lack of pigment at their centers. Details for particular individuals follow. Adult male paratype (UF 156447; Fig. 1 b): Dorsum of body and tail blackish brown becoming a faded lighter brown on the dorsal and lateral surfaces of head extending to the chin. A lighter brown speckling is faintly noticeable dorsally from the head becoming imperceptible towards the base of the tail. Ventral surfaces of the body are blackish brown, similar in coloration to the tail. Limbs faded brown similar to color of head. Skin creases such as costal grooves, gular fold, and those around the paratoid region lack pigmentation and appear light grey. Iris dark rust brown. Adult male (UF 156450): Dorsum of body and tail blackish brown becoming faded lighter brown on the anterior and lateral surfaces of head. Gold speckling mixed with some white is located on the dorsal surface of the head and body. Speckling is most prominent on the top of the head becoming sparser posteriad, not extending beyond base of tail. Dorsolaterally above the anterior limbs a slight concentration of tiny white speckles forms a short, irregular line. Below the extent of the speckling ventrolaterally coloration is same as that of tail. Limb coloration is same as that of anterior and lateral surface of head. Iris dark rust brown. Juvenile paratype (UF 156454; Fig. 1 d): Dorsal and lateral surfaces of body, head, tail, and limbs are a uniform jet black. There is a faint overlay of tiny dirty white flecks along the dorsum of head, body, and tail giving a somewhat ashy appearance. This flecking is most concentrated dorsolaterally giving the impression of an irregular line extending from behind the jaw to the hind limb. Flecking is also present around the mouth and chest area. Larger white speckles are mixed sparsely and intermittently along the head, body, tail, and limbs. Habitat and distribution. This species is known to occur from 1360���1660 m elevation at three isolated localities in the Lower Montane Moist Forest formation (Holdridge 1967) in north-central Nicaragua (Fig. 3). The holotype was collected in a small patch of secondary cloud forest (Fig. 4 a���b) mostly surrounded by cattle pastures and agricultural land. It was found while raking during the daytime inside a moderately large rotten log. The paratypes and UF referred specimens from Reserva Natural Cerro Kilamb�� were collected in undisturbed, primary cloud forest (Fig. 4 c) during both diurnal and nocturnal surveys. All were found underneath logs, rocks, and other debris in areas along the mountain ridge where the substrate comprised a spongy mass of plant roots and mosses. The SMF referred specimens from Reserva Natural Cerro Kilamb�� were also collected in undisturbed, primary cloud forest during diurnal surveys. Most of them were found inside dead basal ramifications of giant ferns. The paratypes from Reserva Natural Macizos de Pe��as Blancas were collected in undisturbed, primary cloud forest (Fig. 4 d). One was uncovered during the daytime beneath a fallen log, and two others were found active on the forest floor around the campsite at 2130 h. Those found active within the camp were likely dislodged or disturbed by raking and uprooting logs that occurred during set-up of the campsite earlier the same day. Phylogenetic relationships. Phylogenetic analysis of sequence data from two mitochondrial genes (16 S and cyt b) confirms the monophyly of this taxon (ML bootstrap support = 99, Bayesian posterior probability = 1.0) and its position as sister species to Oedipina kasios in the Oeditriton clade (Fig. 2). A sister-species relationship between O. kasio s and O. nica is strongly supported (ML bootstrap support = 93, Bayesian posterior probability = 0.99) as is the monophyly of an Oeditriton clade containing O. kasio s, O. nica, and O. quadra (ML bootstrap support = 96, Bayesian posterior probability = 1.0). Oeditriton is the sister clade to the remaining Oedipina, which are divided into the subgenera Oedipina and Oedopinola (Fig. 2), which supports previous phylogenetic hypotheses for Oedipina (Garc��a-Par��s & Wake 2000; McCranie et al. 2008). All wellsupported phylogenetic relationships are congruent between the ML and Bayesian results. Etymology. The specific name nica is a short name for Nicaraguan (Nicarag��ense in Spanish) and is a colloquial word used in Central America to denote Nicaraguan people. The name alludes to the country of origin of the type specimens. Conservation status. Using the criteria established by IUCN for evaluating threatened species, Oedipina nica should be classified as Endangered (EN B 2 ab[iii]), due to its limited distribution (known only from three isolated highland forest areas with a total extent of less than 500 km 2) and the continued loss of habitat at these localities., Published as part of Sunyer, Javier, Wake, David B., Townsend, Josiah H., Travers, Scott L., Rovito, Sean M., Papenfuss, Theodore J., Obando, Lenin A. & K��hler, Gunther, 2010, A new species of worm salamander (Caudata: Plethodontidae: Oedipina) in the subgenus Oeditriton from the highlands of northern Nicaragua, pp. 29-39 in Zootaxa 2613 on pages 32-35, DOI: 10.5281/zenodo.197858, {"references":["Kohler, G. (1999) The amphibians and reptiles of Nicaragua - a distributional checklist with keys. Courier Forschungsinstut Senckenberg, 213, 1 - 212.","Kohler, G. (2001) Anfibios y Reptiles de Nicaragua. Herpeton. Offenbach, Germany.","Holdridge, L. R. (1967) Life Zone Ecology. Tropical Science Center. San Jose, Costa Rica.","McCranie, J. R., Vieites, D. R. & Wake, D. B. (2008) Description of a new divergent lineage and three new species of Honduran salamanders of the genus Oedipina (Caudata, Plethodontidae). Zootaxa, 1930, 1 - 17."]}

Supplmentary methods, data, and Figures for Oliver et al. 2017

Observed (red bars) versus simulated numbers (blue) of taxa occurring in marginal habitats in regions of Continental and Arc geology: A) the East Papuan Composite Terrane (EPCT) coded as Arc; and B) with the EPCT coded as Continental. The EPCT region is not strictly an island arc, rather a microcontinent, however geological models suggest a history of isolation similar to island arcs, and that land in this region has only recently accreted onto the present day landmass of New Guinea.

Bayesian chronogram estimated from nuclear data, using the uncorrelated lognormal model, two partitions and two deep gekkotan calibrations. Lepidodactylus s.l. highlighted in red, Gekko Group in green. Timeframe in millions of years ago.

Supplmentary methods, data, and Figures for Oliver et al. 2017

Bayesian chronogram for gekkotan lizards estimated using a five nuclear gene alignment from Gamble et al. 2015 and 4 fossil calibrations (indicated with red circles). Clade comprising of Lepidodactylus s.l and the Gekko group highlighted in green, and show in more detail, with secondary priors used in downstream analyses also highlighted with black circles. Three gecko Clades with overlapping distributions east of the Sunda Shelf are also highlighted in blue.

Supplmentary methods, data, and Figures for Oliver et al. 2017

Bayesian chronogram estimated from nuclear data, using the uncorrelated lognormal model, two partitions and two deep gekkotan calibrations. Lepidodactylus s.l. highlighted in red, Gekko Group in green. Timeframe in millions of years ago.

Observed (red bars) versus simulated numbers (blue) of taxa occurring in marginal habitats in regions of Continental and Arc geology: A) the East Papuan Composite Terrane (EPCT) coded as Arc; and B) with the EPCT coded as Continental. The EPCT region is not strictly an island arc, rather a microcontinent, however geological models suggest a history of isolation similar to island arcs, and that land in this region has only recently accreted onto the present day landmass of New Guinea.

Bayesian chronogram for gekkotan lizards estimated using a five nuclear gene alignment from Gamble et al. 2015 and 4 fossil calibrations (indicated with red circles). Clade comprising of Lepidodactylus s.l and the Gekko group highlighted in green, and show in more detail, with secondary priors used in downstream analyses also highlighted with black circles. Three gecko Clades with overlapping distributions east of the Sunda Shelf are also highlighted in blue.

Ancestral state estimation of patterns of evolutionary transition between major types of habitats within Lepidodactylus s.l.: (A) four-state habitat state model and (B) two-state habitat model dichotomised by predicted biotic diversity and complexity (i.e Lowland Rainforest v. all other habitat). Branch labels values are posterior probability of ancestral state estimation from BEAST using simple Lewis model.

Ancestral state estimation of patterns of evolutionary transition between major types of habitats within Lepidodactylus s.l.: (A) four-state habitat state model and (B) two-state habitat model dichotomised by predicted biotic diversity and complexity (i.e Lowland Rainforest v. all other habitat). Branch labels values are posterior probability of ancestral state estimation from BEAST using simple Lewis model.

BioGeobears ancestral area estimations. From left to right: 3-state for geological basement: Continental (C), Island Arc (A) and Oceanic (O); 2-state: non-Sundaic taxa (E) vs. Sunda Shelf (W) taxa; and 7-state bioregions: Sunda Shelf (su), Indian Ocean (IO), Maluku/Sulawesi (MS), Philippines (PS), New Guinea (NG), eastern Melanesia (EM) and Pacific Ocean (PO). Ancestral states estimated from biogeographic models with highest AIC values are shown.Note: For 3-state geological basement analysis presented here the Philippine regions of the Zamboanga Peninsula and Palawan Island (Asian continental fragments) were coded as Continental, while East Papuan Composite Terrance was coded as Island Arc

BioGeobears ancestral area estimations. From left to right: 3-state for geological basement: Continental (C), Island Arc (A) and Oceanic (O); 2-state: non-Sundaic taxa (E) vs. Sunda Shelf (W) taxa; and 7-state bioregions: Sunda Shelf (su), Indian Ocean (IO), Maluku/Sulawesi (MS), Philippines (PS), New Guinea (NG), eastern Melanesia (EM) and Pacific Ocean (PO). Ancestral states estimated from biogeographic models with highest AIC values are shown.Note: For 3-state geological basement analysis presented here the Philippine regions of the Zamboanga Peninsula and Palawan Island (Asian continental fragments) were coded as Continental, while East Papuan Composite Terrance was coded as Island Arc