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Micrelapidae new family. Type genus: Micrelaps Boettger, 1880. Type species: Micrelaps muelleri Boettger, 1880. Etymology: Boettger (Bottger ¨) did not give the etymology for the generic nomen but was almost certainly from the Latin adjective micro-, derived from the Greek mikros (small), and elaps, the Latinised form of the Greek noun´ellops or´elaps (literally sea-fish or serpent, but here in reference to the snake genus Elaps, now a synonym of Homoroselaps). Micrelapidae fam. nov. is derived from Micrelaps by the taking the stem elap- of the root word of the nomen. Content: Micrelaps muelleri Boettger, 1880, Micrelaps bicoloratus Sternfeld, 1908, Micrelaps vaillanti Mocquard, 1888, Brachyophis revoili Mocquard, 1888. Diagnosis and definition: In the crania of Micrelaps and Brachyophis we examined the ectopterygoid was laterally and medially expanded at the point of contact with the pterygoid, with this expansion not being contiguous with the ectopterygoid anterolateral and anteromedial lobes (Fig. 4, Supplementary material fig. 53, 54). The lateral expansion is a posterolaterally and somewhat ventrally directed, very prominent protuberance continuous with a ridge on the ventral surface of the pterygoid. This character state was not present in any other cranium we examined and is very likely a synapomorphy of the family. Other common cranial characters include a premaxilla adapted for a fossorial lifestyle, premaxillary transverse processes closely approaching the maxilla, a short maxilla with ascending processes abutting the prefrontal, well-developed, grooved fangs below the orbit, preceded by a diastema and 2 – 3 teeth, an ectopterygoid deeply forked into anterolateral and anteromedial lobes that articulate with maxillary ectopterygoid processes leaving a foramen in the middle, prefrontal and parietal supraorbital processes laterally bordering the frontal and almost meeting each other, a tendency towards fusion of cranial bones (especially because the supratemporal is absent, very likely fused to the quadrate in Brachyophis and to posterior chondrocranial elements in Micrelaps), and a short quadrate. Brachyophis, however, differs from the type genus in possessing a postorbital (versus postorbital absent in Micrelaps), dorsolateral adductor ridges on the parietal (versus a single sagittal ridge in Micrelaps), only a faint pseudocoronoid ridge on the dentary (versus a prominent process in Micrelaps). Scalation characters that are common in both genera include 1 nasal, 7 supralabials, 15 smooth dorsal scale rows, absence of a loreal, 2 anal shields. Ventrals range from 170 to 280 and subcaudals (paired) 16 – 32 in Micrelaps (Boulenger, 1896; De Witte and Laurent, 1947; Rasmussen, 2002; Werner et al., 2006; Spawls et al., 2018). In Brachyophis, ventrals range from 103 to 123 and subcaudals (single) 8 – 14 (De Witte and Laurent, 1947; Lanza, 1966). Brachyophis has a large, azygous occipital shield (Boulenger, 1896). Micrelaps and Brachyophis possess a rectal caecum and a short genital sinus in the female, two soft tissue traits used to cluster these two genera by Underwood and Kochva (1993). Distribution: Micrelaps spp. is distributed in eastern and northeastern Africa and western Asia. Brachyophis is limited to Somalia in north-eastern Africa. Distribution: Micrelaps spp. is distributed in eastern and northeastern Africa and western Asia. Brachyophis is limited to Somalia in north-eastern Africa. Remarks: Geniez (2018) commented that Micrelaps “could constitute a separate family within its own right, that of Micrelapsidae”. Bar et al. (2021) likewise wrote that “. The actual placement of the genus [Micrelaps] is often poorly supported within studies and inconsistent across them. We suspect it will soon be placed in its own family — as is the norm in taxonomy these days. We predict this family, containing a single genus (Micrelaps), will be called Micrelapidae.”. However, these authors did not explicitly express that they are erecting a new family for these snakes. Rather, it was a suggestion about what should/could be done. It therefore is not in accordance with Article 16.1 and Recommendation 16A of The Code (ICZN, 1999). They also did not also provide characters for the express purpose of differentiating or diagnosing “Micrelapsidae” or “ Micrelapidae ”, nor did they cite a work containing the same (again, very likely because a nomenclatural act presumably was not the intention of Geniez [2018] and it was not the intention of SM, who wrote this in Bar et al. [2021] either) and this contravenes Articles 13.1.1, 13.1.2 and Recommendation 13A of The Code. Hence, we regard the nomen “Micrelapsidae” as unavailable. The ZooBank LSID for this taxonomic action is urn:lsid:zoobank.org:pub:D8475246-AD8E-4886-AB55- 12F6F242E9C4., Published as part of Das, Sunandan, Greenbaum, Eli, Meiri, Shai, Bauer, Aaron M., Burbrink, Frank T., Raxworthy, Christopher J., Weinell, Jeffrey L., Brown, Rafe M., Brecko, Jonathan, Pauwels, Olivier S. G., Rabibisoa, Nirhy, Raselimanana, Achille P. & Merila, Juha, 2023, Ultraconserved elements-based phylogenomic systematics of the snake superfamily Elapoidea, with the description of a new Afro-Asian family, pp. 1-11 in Molecular Phylogenetics and Evolution 180 on pages 8-9, DOI: 10.1016/j.ympev.2022.107700, http://zenodo.org/record/7746501, {"references":["Sternfeld, R., 1908. Zur Schlangenfauna Ostafrikas. I. Schlangen aus Sud-Abessinien. Mitt. Zool. Mus. Berlin 4, 239 - 247.","Mocquard, F., 1888. Sur une collection de Reptiles et de Batraciens rapport´es des Pays Comalis et de Zanzibar par M. G. R´evoil. M´emoires Publies par la Soci´ete´ Philomathique a l' occasion du Centenaire de sa fondation 1788 - 1888, 109 - 134.","Boulenger, G. A., 1896. Catalogue of the Snakes in the British Museum (Natural History). Volume III, containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae, and Viperidae. Trustees of the British Museum, London.","de Witte, G. - F., Laurent, R., 1947. R´evision d' un groupe de Colubridae africains: genres Calamelaps, Miodon, Aparallactus et formes affines. M´em. Mus. R. His. Nat. Belg. 29, 1 - 134.","Rasmussen, J. B., 2002. A review of the African members of the genus Micrelaps Boettger 1880 (Serpentes Atractaspididae). Tropical Zoology 15, 71 - 87.","Werner, Y. L., Babocsay, G., Carmely, H., Thuna, M., 2006. Micrelaps in the southern Levant: variation, sexual dimorphism, and a new species (Serpentes: Atractaspididae). Zool. Middle East 38, 29 - 48.","Spawls, S., Howell, K., Hinkel, H., Menegon, M., 2018. Field Guide to East African Reptiles, second edition. Bloomsbury Wildlife, London.","Lanza, B., 1966. Il genere Brachyophis e descrizione di una nuova forma (Reptilia, Serpentes, Colubridae). Monit. Zool. Ital. 74, 30 - 48.","Underwood, G., Kochva, E., 1993. On the affinities of the burrowing asps Atractaspis (Serpentes: Atractaspididae). Zool. J. Linn. Soc. 107, 3 - 64.","Geniez, P., 2018. Snakes of Europe, North Africa and the Middle East: A Photographic Guide. Princeton University Press, Princeton and Oxford.","Bar, A., Haimovitch, G., Meiri, S., 2021. Field guide to the amphibians and reptiles of, Israel. Edition. Chimaira, Frankfurt Am Main.","ICZN (International Commission on Zoological Nomenclature), 1999. International Code of Zoological Nomenclature, fourth edition. International Trust for Zoological Nomenclature, London."]}

Micrelapidae new family. Type genus: Micrelaps Boettger, 1880. Type species: Micrelaps muelleri Boettger, 1880. Etymology: Boettger (Bottger ¨) did not give the etymology for the generic nomen but was almost certainly from the Latin adjective micro-, derived from the Greek mikros (small), and elaps, the Latinised form of the Greek noun´ellops or´elaps (literally sea-fish or serpent, but here in reference to the snake genus Elaps, now a synonym of Homoroselaps). Micrelapidae fam. nov. is derived from Micrelaps by the taking the stem elap- of the root word of the nomen. Content: Micrelaps muelleri Boettger, 1880, Micrelaps bicoloratus Sternfeld, 1908, Micrelaps vaillanti Mocquard, 1888, Brachyophis revoili Mocquard, 1888. Diagnosis and definition: In the crania of Micrelaps and Brachyophis we examined the ectopterygoid was laterally and medially expanded at the point of contact with the pterygoid, with this expansion not being contiguous with the ectopterygoid anterolateral and anteromedial lobes (Fig. 4, Supplementary material fig. 53, 54). The lateral expansion is a posterolaterally and somewhat ventrally directed, very prominent protuberance continuous with a ridge on the ventral surface of the pterygoid. This character state was not present in any other cranium we examined and is very likely a synapomorphy of the family. Other common cranial characters include a premaxilla adapted for a fossorial lifestyle, premaxillary transverse processes closely approaching the maxilla, a short maxilla with ascending processes abutting the prefrontal, well-developed, grooved fangs below the orbit, preceded by a diastema and 2 – 3 teeth, an ectopterygoid deeply forked into anterolateral and anteromedial lobes that articulate with maxillary ectopterygoid processes leaving a foramen in the middle, prefrontal and parietal supraorbital processes laterally bordering the frontal and almost meeting each other, a tendency towards fusion of cranial bones (especially because the supratemporal is absent, very likely fused to the quadrate in Brachyophis and to posterior chondrocranial elements in Micrelaps), and a short quadrate. Brachyophis, however, differs from the type genus in possessing a postorbital (versus postorbital absent in Micrelaps), dorsolateral adductor ridges on the parietal (versus a single sagittal ridge in Micrelaps), only a faint pseudocoronoid ridge on the dentary (versus a prominent process in Micrelaps). Scalation characters that are common in both genera include 1 nasal, 7 supralabials, 15 smooth dorsal scale rows, absence of a loreal, 2 anal shields. Ventrals range from 170 to 280 and subcaudals (paired) 16 – 32 in Micrelaps (Boulenger, 1896; De Witte and Laurent, 1947; Rasmussen, 2002; Werner et al., 2006; Spawls et al., 2018). In Brachyophis, ventrals range from 103 to 123 and subcaudals (single) 8 – 14 (De Witte and Laurent, 1947; Lanza, 1966). Brachyophis has a large, azygous occipital shield (Boulenger, 1896). Micrelaps and Brachyophis possess a rectal caecum and a short genital sinus in the female, two soft tissue traits used to cluster these two genera by Underwood and Kochva (1993). Distribution: Micrelaps spp. is distributed in eastern and northeastern Africa and western Asia. Brachyophis is limited to Somalia in north-eastern Africa. Distribution: Micrelaps spp. is distributed in eastern and northeastern Africa and western Asia. Brachyophis is limited to Somalia in north-eastern Africa. Remarks: Geniez (2018) commented that Micrelaps “could constitute a separate family within its own right, that of Micrelapsidae”. Bar et al. (2021) likewise wrote that “. The actual placement of the genus [Micrelaps] is often poorly supported within studies and inconsistent across them. We suspect it will soon be placed in its own family — as is the norm in taxonomy these days. We predict this family, containing a single genus (Micrelaps), will be called Micrelapidae.”. However, these authors did not explicitly express that they are erecting a new family for these snakes. Rather, it was a suggestion about what should/could be done. It therefore is not in accordance with Article 16.1 and Recommendation 16A of The Code (ICZN, 1999). They also did not also provide characters for the express purpose of differentiating or diagnosing “Micrelapsidae” or “ Micrelapidae ”, nor did they cite a work containing the same (again, very likely because a nomenclatural act presumably was not the intention of Geniez [2018] and it was not the intention of SM, who wrote this in Bar et al. [2021] either) and this contravenes Articles 13.1.1, 13.1.2 and Recommendation 13A of The Code. Hence, we regard the nomen “Micrelapsidae” as unavailable. The ZooBank LSID for this taxonomic action is urn:lsid:zoobank.org:pub:D8475246-AD8E-4886-AB55- 12F6F242E9C4.

A primary goal of biogeography is to understand how large-scale environmental processes, like climate change, affect diversification One often-invoked but seldom tested process is the “species-pump” model, in which repeated bouts of co-speciation are driven by oscillating climate-induced habitat connectivity cycles. For example, over the past three million years, the landscape of the Philippine Islands has repeatedly coalesced and fragmented due to sea-level changes associated with glacial cycles. This repeated climate-driven vicariance has been proposed as a model of speciation across evolutionary lineages codistributed throughout the islands. This model predicts speciation times that are temporally clustered around the times when interglacial rises in sea level fragmented the islands. To test this prediction, we collected comparative genomic data from 16 pairs of insular gecko populations. We analyze these data in a full-likelihood, Bayesian model-choice framework to test for shared divergence times among the pairs. Our results provide support against the species-pump model prediction in favor of an alternative interpretation, namely that each pair of gecko populations diverged independently. These results suggest the repeated bouts of climate-driven landscape fragmentation has not been an important mechanism of speciation for gekkonid lizards on the Philippine Islands.

Weinell, Jeffrey L, Barley, Anthony J, Siler, Cameron D, Orlov, Nikolai L, Ananjeva, Natalia B, Oaks, Jamie R, Burbrink, Frank T, Brown, Rafe M (2021): Phylogenetic relationships and biogeographic range evolution in cat-eyed snakes, Boiga (Serpentes: Colubridae). Zoological Journal of the Linnean Society 192 (1): 169-184, DOI: 10.1093/zoolinnean/zlaa090, URL: https://academic.oup.com/zoolinnean/article/192/1/169/5892978

Toxicodryas blandingii (Hallowell, ���1844��� 1845) (Table 1, Figs. 5���8) Dipsas Blandingii: Hallowell (���1844��� 1845:170); type locality: ��� Liberia, West Africa. ��� Triglyphodon fuscum: Dum��ril, Bibron & Dum��ril (1854:1101); type locality: ��� Grand-Bassam, sur la C��te d���Ivoire (Guin��e)��� [Ivory Coast]. Dipsas fasciata: Fischer (1856:84); type locality: ��� Peki (West-Afrika)��� [Ghana]. Dipsas valida: Fischer (1856:87); type locality: ��� Edina (Grand Bassa County, West-Afrika)��� [Liberia]. Dipsas globiceps: Fischer (1856:89); type locality: ��� Edina (Grand Bassa County, Liberia, West-Afrika).��� Toxicodryas Blandingii: Hallowell (1857:60); comb. nov. Dipsas Fischeri: Jan in Dum��ril (1859:212); no type locality provided. Triglyphodon fuscum var. obscurum: Dum��ril (1861:211); type locality: ��� C��te d���Or ��� [Ghana]. Dipsas regalis: Jan (1871:3, Livraison 38, pl. vi, fig. 2) in Jan & Sordelli (1870 ���1881); type locality: ��� C��te d���Or ��� [Ghana]. Dipsas globiceps var. tumboensis: M��ller (1885:688); type locality: ��� Tumbo-Insel ��� [Guinea]. Boiga blandingi occidentalis: Stucki-Stirn (1979:377); inferred type locality: ���Besongabang��� Cameroon. Boiga blandingi subfulva: Stucki-Stirn (1979:381); type locality: none provided, but limited to Cameroon according to book title. Toxicodryas blandingii was originally described by Hallowell (1845:170) based on a single specimen collected by his friend, ���Dr. Blanding,��� in Liberia. The dorsum and venter of the specimen was noted to have a ���light yellow��� color with a series of blotches of ���leaden colour.��� This specimen reportedly possessed 2 preoculars, 2 postoculars, 272 ventrals, 131 subcaudals, body length (i.e., SVL) of 1.22 meters, and tail length of 0.39 meters. Hallowell (1854) provided additional details of the specimen���s teeth, noted it had 17 ���rows of scales,��� and corrected the tail length to 0.37 meters. Hughes & Barry (1969), Wallach et al. (2014) and Uetz et al. (2019) stated that the type was lost, which is consistent with Malnate (1971), who did not list a type specimen from the ANSP collection. Wallach et al. (2014) noted the type was a 1.67 m specimen, slightly longer than the total length of 1.61 m reported by Hallowell (1845) in the original description, but the longer measurement is likely a typographical error in reference to the latter citation (V. Wallach, pers. comm.). A query by EG to the Philadelphia Academy of Sciences in spring 2020 resulted in location of the type specimen (ANSP 10083, Fig. 6), and a redescription of this specimen is provided below. According to Loveridge (1957:269), the name Dipsas Fischeri was proposed by Jan (in Dum��ril 1859) to combine the minor color pattern variants Dipsas fasciata, D. valida, and D. globiceps named by Fischer (1856). Pel (1852:171) coined the name Dipsas regalis, and as translated by Savage & McDiarmid (2017:73), Pel stated, ���the third species of venomous snake, Naja atropos, belongs to cobras (spectacled snakes) and reaches a length of 6 to 7 feet [1.8���2.1 m]. Its color is entirely black... As this snake in general shows much similarity to a tree snake, Dipsas regalis, which equals it in color and size, but is not venomous.��� Perhaps because of this poor description, Boulenger (1896:78) attributed the latter name to Jan & Sordelli (1870 ���1881), who provided an illustration that served as an appropriate description. Jan listed the name in the Index des Planches for Livraison 38 as Dipsas cynodon Cuv. vari��t��? (D. regalis Schlegel), but according to Savage & McDiarmid (2017:73), the attribution to Hermann Schlegel is in error because he never used the name in any publication. M��ller (1885:687) seemed to suggest that Jan illustrated his specimen (collected by Dr. M��hly from ���Goldk��ste��� [i.e., Gold Coast or modern-day Ghana]) from Basel, but Hughes & Barry (1969:1020) listed a personal communication from M.S. Hoogmoed, who noted the type of D. regalis (specimen ���Leiden 958���) was collected by Pel in February 1844 from Accra, Ghana. Hallowell (1857:60) coined the genus Toxicodryas because he noticed that his specimen of T. blandingii had a ���single channelled posterior tooth on each side... and therefore...[it] cannot belong to the genus Triglophodon [sic] of Dum. and Bibron, which has three.��� Subsequent herpetological publications in the 19 th and early 20 th centuries seemingly ignored Hallowell���s new genus and continued to recognize the taxon in either the genus Dipsas (e.g., Mocquard 1896) or more commonly, Dipsadomorphus (e.g., Boulenger 1896, 1919; de Witte 1933). Schmidt (1923) transferred the taxon to the genus Boiga in his opus on Congolese snakes, recognizing B. (Toxicodryas) blandingii and B. (Toxicodryas) pulverulenta, an action that was followed by most subsequent authors for decades. Based on the placement of African Boiga in the ��� Dipsadidae: Lycodontinae ��� by Underwood (1967), Welch (1982) seems to have been the first to recognize the genus Toxicodryas for both species of the genus, an action followed by Meirte (1992) and observed by most herpetologists in the 21 st century (e.g., Uetz et al. 2020). Boulenger (1896) included all of the above West African, 19 th- century names in the synonymy of Dipsas (Toxicodryas) blandingii. Because the dubious subspecies described by Stucki-Stirn (1979) both occur in Cameroon, where one molecular sample (CAS 253611) from Allen et al. (in press) is recovered in a well-supported clade with West African samples (Fig. 2), we confirm the taxonomic nomenclature of Wallach et al. (2014) in treating these taxa as synonyms of T. blandingii. Marques et al. (2018) noted that northwestern Angolan records from ���Chinchoxo,��� ���Piri-Dembos,��� and ���Quirimbo��� by Peters (1877), Bocage (1895), Parker (1936), and Hellmich (1957a, b) are attributable to T. pulverulenta. However, the morphometric data provided by Hellmich (1957b) for Angolan snakes are inconsistent with the size and scale rows of the latter species, and herein, we consider his records to be attributable to T. vexator sp. nov. Diagnosis. Toxicodryas blandingii, as recognized herein, is restricted to West Africa and west-central Africa (west of the confluence of the Congo and Ubangi rivers), defined by the following combination of characters: maximum SVL> 1 meter (vs. maximum SVL T. pulverulenta and T. adamanteus sp. nov.); DSRN 23���25 (vs. 19���21 in T. pulverulenta and 18���23 in T. adamanteus sp. nov.); DSRM 21���25 (vs. 19���21 in T. pulverulenta and 18���21 in T. adamanteus sp. nov.); cloacal plate usually divided (vs. divided or undivided in T. vexator sp. nov., and always undivided in T. pulverulenta and T. adamanteus sp. nov.); adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters (vs. both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots in T. pulverulenta and T. adamanteus sp. nov.); hemipenis relatively short and massive (i.e., broad), proximal third covered with spines, distal two-thirds dimpled with a flattened apex (vs. relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, and with a domed apex in T. pulverulenta and T. adamanteus sp. nov.); venom toxicity LD 50 = 2.85���3.55 mg /kg in mice (vs. venom toxicity LD 50 = 4.88 mg /kg in mice for T. vexator sp. nov.). Redescription of the holotype. ANSP 10083 (Fig. 6) adult female in poor condition, 1330 mm SVL; head triangular and distinct from neck, 1.92% of SVL (25.5 mm); right loreal missing, left loreal partially obscured by supralabials due to cranial damage, upper side tapering superiorly; body triangular; tail moderately long (400 mm; 30.1% of SVL). Supralabials ���/9, ���/4 th, 5 th, and 6 th contacting orbit; infralabials 14/13, 1 st on each side in contact behind mental, 1 st ���4 th /1 st ���4 th contacting anterior chin shields and 4 th ���7 th /4 th ���7 th contacting posterior chin shields; 2 preoculars; 3 postoculars (on left, missing on right); temporals ���/2 + 2; 2 internasals; nasal divided (on left, missing on right); frontal slightly longer than wide, only left side undamaged; dorsal scale rows 23 one head length posterior to jaw rictus, 23 at midbody, 17 one head length anterior to vent, smooth and oblique, vertebral scales broad and apically flattened; ventrals 273 (standard), 271 (Dowling); cloacal plate undivided; paired subcaudals 132. Coloration (in preservative). After approximately 176 years in preservation, specimen is faded, with creamy tan background color in dorsal and ventral views. Brown markings on posterior edge of supralabials and dorsum of head. Irregular brown and dark brown blotches and saddles on dorsum from neck to tip of tail (Fig. 6). Variation. Morphometric variation of Toxicodryas blandingii is shown in Table 1. M��ller (1885:688) provided data for a snake from Ghana with 15 infralabials and noted that most of its scales have two ���Endgruben��� [terminal pits], which likely refer to apical pits. In his description of Dipsas globiceps var. tumboensis M��ller (1885) noted his specimen from Guinea (Fig. 7) had 147 subcaudals. In snakes from West Africa (without separating by sex), Angel (1933) noted a temporal formula of 2 + 2 or 2 + 3, 21���25 scales at midbody, 240���289 ventrals, 120���147 subcaudals, either an undivided or divided cloacal plate, and a maximum total size of 2290 mm, and nearly verbatim variation was listed by Villiers (1950a), Doucet (1963), Stucki-Stirn (1979), Chippaux (2006), and Trape & Man�� (2006). However, in snakes from Ghana, Swiecicki (1965:302) noted a maximum total length of 2450 mm for a ���black form��� individual, Gauduin (1970) listed the maximum total length as 2700 mm (600 mm tail) for Cameroon, Chirio & LeBreton (2007) provided a slightly larger total length of 2740 mm for Cameroon, and Luiselli et al. (1998a) noted a maximum size of 2800 mm, presumably for Nigeria. Villiers (1951) noted an unsexed individual from Benin with 115 subcaudals. Cansdale (1965) documented 21���25 scales at midbody, and Segniagbeto et al. (2011) documented snakes from Togo with 19���24 scales at midbody and 102���159 subcaudals. In our examined specimens, temporal formula includes the variation noted by Angel (1933), but is more extensive (2 +5, 3 + 4, 3 + 3, 1 + 5, 3 + 2, or 2 + 4), and either supralabials 3���5 or 4���6 contact the eye, which is consistent with the observations of Angel (1933), Villiers (1950b), and Chippaux (2006); the latter author also noted that sometimes only 2 supralabials contact the eye. Rasmussen (1997a) noted specimens with the 4th���5th, 5th���7 th, or 4th���7 th supralabials in contact with the eye, and in general, this species has sloping and smooth scales with apical pits, and the vertebral row is greatly enlarged. The holotype, one male and one female from Liberia (Loveridge 1941; Johnsen 1962), one male from Gabon (Pauwels et al. 2002b), one male from DRC (RBINS 10888), and a juvenile from Cameroon (Werner 1897) are unusual in having an undivided cloacal plate, because all other examined specimens have a divided cloacal plate, including the type specimens of Dipsas fasciata, D. valida, and D. globiceps (Fischer 1856). Rasmussen (1997a) remarked that his specimens have either a divided or undivided cloacal plate. In his book on West and Central African snakes, including countries west of the Congo River, Chippaux (2006:154) noted the anal [cloacal plate] is sometimes entire, but more often divided. Trape & Man�� (2006) stated that the cloacal plate is almost always divided. Segniagbeto et al. (2011) noted individuals with divided or undivided cloacal plates in Togo. Combined descriptions by Fischer (1856) of the teeth of Dipsas fasciata, D. valida, and D. globiceps (all now synonyms of T. blandingii) suggest the species has 9 maxillary teeth that increase in size posteriorly, followed by two fangs (three on the right side in one specimen), and 12 mandibular teeth, which decrease in size posteriorly. In snakes from Ghana, Leeson (1950) noted 10���11 maxillary teeth, becoming larger posteriorly, and two fangs followed by a shorter fang; fourteen large palatine and pterygoid teeth, and 15 mandibular teeth (anterior ones largest). Taylor & Weyer (1958:1217) described a Liberian specimen with 9���10 maxillary teeth (on different sides) that increase in size from the 1 st to 4 th tooth, and then become subequal; two fangs occur after this series of teeth, and after a short diastema, there is a third fang with only traces of a groove. A second Liberian specimen had 10 maxillary teeth followed by three fangs, the last of which had only ���a suggestion of a groove.��� Based on specimens ranging from Guinea to Congo, Rasmussen (1997a:98) noted 10 maxillary teeth followed by three enlarged, furrowed venomous teeth, with the 3 rd fang slightly smaller than the previous two. Fischer (1856) provided detailed descriptions of the color patterns of West African Dipsas fasciata, D. valida, and D. globiceps (all now synonyms of T. blandingii), which seem to suggest he examined a subadult male that had been kept in alcohol for a long (unspecified amount) time (D. fasciata), an adult female (D. valida), and a subadult that retained juvenile coloration (D. globiceps). In his description of Dipsas globiceps var. tumboensis M��ller (1885:689) noted his specimen from Guinea (Fig. 7) had a gray-reddish dorsum with 30 black transverse bands, usually containing milky white spots. The frontal, supraoculars, and occipital scales had large black spots, the labial scales and postoculars were edged with black, and the head shields had multiple milk-white speckles. The tail was bright red with dark, irregular transverse bands. Aspects of this coloration description are highly unusual for this species (e.g., bright red tail), and more typical of T. pulverulenta, but the black edging of the labial scales, number of preoculars (3), supralabials in contact with the eye (5 and 6), ventrals (269), and subcaudals (147) clearly indicate this taxon is a synonym of T. blandingii (Fig. 7, Tables 1���2). Mocquard (1887:80) described a recently collected, unsexed subadult (���la longueur du tronc��� [trunk length] 1.1 m) from Gabon as having a dorsal color of a general tint of Burgundy with slightly darker spots on the flanks that have a dirty white spot a little above their lower edge. Mertens (1938) described an adult male from Cameroon as solid black dorsally and ventrally, with the exception of the anterior third of the venter, which was white, but the ventral scales had dark gray edges. The labial scales were gray with vertical black borders. Villiers (1950b) described the color pattern of an unsexed individual from Ivory Coast as sooty black or brownish in places on the dorsum; underside iridescent dark gray posteriorly, becoming whitish anteriorly, with the posterior edge of the ventrals edged with gray; underside of head white, and labials whitish and edged in black. Another unsexed individual from Liberia was described as bluish black above, yellow below; supralabials yellow with black edges, and posterior part of venter and underside of tail black. A third unsexed individual from Liberia had identical coloration to the latter one, except for the presence of whitish bars on the neck. Leeson (1950) noted that snakes from Ghana have a dull green or gray dorsum. Monard (1951:162) described an unsexed individual from Cameroon as a beautiful light redbrown, ���barr����� [barred] with dark brown. Taylor & Weyer (1958:1217) described a Liberian brown-phase female with pale grayish green on the ventral side of the head and neck, merging into gray with ���a greenish cast��� 12.7 cm posteriorly, and at 40.6 cm behind the head, it transitioned into plain tan to the tip of the tail. Isemonger (1962:12) remarked that this species has a ���delicate bloom on the skin.��� Cansdale (1965:43) described a highly unusual color pattern for juveniles by noting that ���the young brown form is pink with irregular chocolate markings that break up its outline very effectively and make it difficult to pick out in a tree or shrub.��� Leston & Hughes (1968:753) described an unusual specimen from Ghana as ���pale grey with darker greyish-green transverse bands, the bands irregular but more or less diamond shaped on each side. The ventrals are also grey but more glossy.��� Groves (1973:107) described the coloration of hatchlings from a captive Liberian female as ���light grey background colour with pinkish undertones; black, roughly oval, lateral blotches narrowing as they approach the midline, where many of them fail to conjoin; top of head light grey; belly dark grey.��� Rasmussen (1997a:98) noted the scales of his specimens were dull and almost dusty, a sentiment also shared by Cansdale (1965). Adult males were solid black on the dorsum and yellow on the venter (becoming black posteriorly), whereas adult females were noted to be gray, brown or yellow-brown on the dorsum and yellow-brown on the venter, sometimes without transverse bands. Hughes (2000:8) noted juvenile and subadult (approximately 1 meter in total length or less) snakes had a dorsal coloration that was ���a distinctively bright and contrasting pattern of chocolate brown blotches.��� He noted that most male specimens lose this coloration as they age, becoming increasingly melanistic, and although exceptions are possible, this melanistic progression does not seem to occur in females. Chippaux (2006:154) noted there are two dorsal color morphs: (1) uniform black or dark blue with ���reflets velout��s��� [velvety reflections] or (2) gray with darker, poorly defined transverse spots. The venter was noted as dull yellow to charcoal gray, and juvenile coloration as light brown with darker transverse ring-shaped spots. Stucki-Stirn (1979), perhaps confused by the tw, Published as part of Greenbaum, Eli, Allen, Kaitlin E., Vaughan, Eugene R., Pauwels, Olivier S. G., Wallach, Van, Kusamba, Chifundera, Muninga, Wandege M., Aris- Tote, Mwenebatu M., Mali, Franck M. 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Toxicodryas pulverulenta (Fischer, 1856) (Table 2, Figs. 8, 11) Dipsas pulverulenta: Fischer (1856:11); type locality: ��� Edina, Grand Bassa County in Liberia (West-Afrika).��� Dipsadomorphus Boueti: Chabanaud (1917a:373); type locality: none provided, but subsequently clarified to ��� Porto-Novo ��� ��� Dahomey ��� [Porto-Novo, Benin] by Chabanaud (1917b:12). As noted by Cer��aco et al. (2018), the description of this species was based on two specimens (only one syntype remains today according to Hallermann 1998 and Uetz et al. 2019), one of which was depicted in plate 4 of livraison 38 by Jan & Sordelli (1871). In a footnote to the nomenclatural history of Boiga (Toxicodryas) pulverulenta, Hughes & Barry (1969:1020) wrote ���Dr. Ladiges [then curator at ZMH] tells us that the type is still in Hamburg and is from St. Thom��, not Edina, Grand Bassa County, Liberia as stated by Loveridge (1958:269).��� The latter citation is in error (Loveridge 1957). Cer��aco et al. (2018) deduced that the information from Ladiges was based on an old label associated with the presumed remaining type specimen. Because extensive fieldwork on the island of S��o Tom�� over the last century failed to document this species, Cer��aco et al. (2018) concluded that it does not occur there, and the original type locality from Fischer (1856) ���Edina, Grand Bassa County in Liberia (West-Afrika)��� is correct. Moreover, the former authors determined that the pholidosis from this presumed type (Cer��aco et al. 2018:fig. 1) is not consistent with the extensively detailed original description, and thus, it is not a type specimen (contra Uetz et al. 2019). Although we agree with this conclusion, an examination of this specimen (ZMH R04376) by J. Hallermann (pers. comm.) found some differences in the morphometric data provided by Cer��aco et al. (2018) (e.g., SVL = 635 mm, not 880 mm), and we used these updated data in our analyses of morphological data (Table 2). Chabanaud (1917a:375), convinced that the odd configuration of subcaudals (combination of single and double subcaudals near the vent) seen in one male and one female was unique to Colubridae, named the taxon Dipsadomorphus boueti from these specimens that were donated by ���Dr Bouet,��� who collected for the Paris Museum in Ivory Coast (1909) and Dahomey (1910���1913). Unfortunately no specific type locality was provided. Chabanaud (1917b) then clarified and corrected himself by stating that both types were males (the mistaken female was a juvenile), and they were collected from ���Porto-Novo��� in Dahomey, which is modern-day Porto-Novo, the capital city of Benin. Chabanaud (1917c) subsequently realized that his specimens of D. boueti represented T. pulverulenta individuals with abnormal subcaudals, and he synonymized the former taxon with the latter (as D. pulverulentus). We agree with this taxonomic placement, because the types of D. boueti have subcaudal counts (106 and 116) and TL/SVL ratios (0.25 and 0.28) that are most consistent with T. pulverulenta (Table 2). The Porto-Novo locality is on the eastern edge of the Dahomey Gap (sensu Demenou et al. 2016:fig. 1), a renowned biogeographic barrier to at least some forest-specialist species in West Africa. Because T. pulverulenta does not seem to be a strict forest specialist (see Habitat below), and the Porto-Novo locality occurs on the eastern edge of the Dahomey Gap, it is likely that the Gap is not a barrier to dispersal for this species. We therefore hypothesize that the Niger Delta is the biogeographic barrier separating T. pulverulenta from its cryptic sister taxon T. adamanteus sp. nov. Diagnosis. A species of Toxicodryas restricted to West Africa, west of the Niger Delta, defined by the following combination of characters: maximum SVL 1 meter in T. blandingii and T. vexator sp. nov.), DSRN 19���21 (vs. 23���25 in T. blandingii and 23���29 in T. vexator sp. nov.), DSRM 19���21 (vs. 21���25 in T. blandingii and T. vexator sp. nov.); cloacal plate undivided (vs. usually divided in T. blandingii, and divided or undivided in T. vexator sp. nov.); both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots (vs. adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters in T. blandingii and T. vexator sp. nov.); hemipenis relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, with a domed apex (vs. hemipenis relatively short and massive [i.e., broad], proximal third covered with spines, distal two-thirds dimpled with a flattened apex in T. blandingii and T. vexator sp. nov.). Variation. Morphometric variation of Toxicodryas pulverulenta is shown in Table 2. Chabanaud (1917a) first noted that some individuals (including the type of Dipsadomorphus boueti) can have a few single subcaudals posterior to the cloacal plate. Bogert (1940) provided morphometric data, but he did not distinguish between populations from Liberia (T. pulverulenta) and former French Cameroon (T. adamanteus sp. nov.), except that ���the Liberia specimens are distinctly more reddish brown than the Cameroon specimens.��� However, he listed the maximum total length of a male (1112 mm) and female (1050 mm) T. pulverulenta from Liberia. In snakes from West Africa (without noting sex), including Cameroon where T. adamanteus sp. nov. occurs, Angel (1933:146) noted temporal formula variation of 2 + 2 (rarely 2 + 3 or 3 + 2 or 1 +2), 236���276 ventrals, 96���132 subcaudals, and a maximum size of 1225 mm; nearly identical data were reported by Villiers (1950a), Doucet (1963), Stucki-Stirn (1979), and Chippaux (2006). Although all of our examined specimens and most literature records noted the 3 rd to 5 th supralabial in contact with the eye (e.g., Villiers 1950b), Chippaux (2006) documented individuals with the 4 th to 6 th supralabial in contact with the eye, and sometimes, only two scales in contact with the eye, but some of these specimens might be attributable to T. adamanteus sp. nov. Segniagbeto et al. (2011) reported snakes from Togo with 165���175 ventrals, which is undoubtedly erroneous. Fischer (1856:83) noted his type specimen had seven maxillary teeth that were oriented nearly backwards, becoming larger posteriorly. These were followed by two larger teeth (i.e., fangs) that were furrowed in a ���besonderen Hauttasche��� [special skin pocket]. There were 10���12 mandibular teeth, which were slightly curved towards the back of the mouth, and increased in size posteriorly. Bogert (1940:61) remarked ��� five specimens examined show variation from eleven to thirteen anterior subequal teeth followed after a very short diastema by two larger grooved fangs and a smaller fang, the total number of teeth being fourteen to eighteen.��� Because only two of his examined specimens originated from former French Cameroon (attributable to T. adamanteus sp. nov.), at least three of these specimens were from Liberia, which are attributable to T. pulverulenta. In snakes from Ghana, Leeson (1950) noted 11���13 maxillary teeth, becoming slightly larger posteriorly, followed by 2 fangs, and sometimes a 3 rd, smaller fang; 14���16 palatine and pterygoid teeth, and 15 mandibular teeth with the anteriormost ones largest. Johnsen (1962) reported that his specimen from Liberia had 11 maxillary teeth on one side, and six on the other. Based on a specimen from Liberia, Malnate (1972) noted the species lacks posterior hypapophyses. In his original description of the species, Fischer (1856:83) described the coloration in great detail as chocolate brown above, yellow below, finely dotted everywhere. On each side, near the back, there was a large number (60��� 70) of pale red spots that lacked dark edging and extended over 4���6 scales. Usually the spots on one side alternated with those on the other; sometimes they also were opposite to each other and in this case were connected to weak ���Querbinden��� [cross-ties] by bright red connecting strips that extended over the back. Usually there was a small black spot under each of these spots on the outermost tip of the corresponding ventral shields. The innumerable fine black points, with which the whole body was sown, were grouped on the abdominal shields on each side at the point where they bent over to rise sideways, to form a black spot, which in their succession looked like a black longitudinal band, whereby the narrow belly appeared delimited from the flanks. Head was brown and without black lines. Upper lip, lower lip, throat yellow, dotted with black. G��nther (1858:173) described the dorsum of two Nigerian specimens as ���brown with a strong cast of purple��� with elliptical transverse streaks, and at mid-body, these streaks had a small yellow spot in their center. In his description of Dipsadomorphus boueti, Chabanaud (1917a) noted the types were light brownish gray and dotted with brown, whereas the supralabials and venter were yellowish gray. Leeson (1950) described animals from Ghana as brownish red on the dorsum, with ���dull brown��� heads, and ���lightly coloured��� patches on the flanks that had a black edge on the lower margins. There were more narrow light gray bands on the anterior third of the body that gradually diminished to form light gray patches on the posterior two-thirds of the body. The ventral surface of the head was white or cream, whereas the venter of the body was pinkish with numerous brown spots. Brown lines ���commence���[ed] just before the middle of the body on each side of the ventrals, and continued along the edge of the ventrals to the tip of the tail. In his description of two specimens from Ivory Coast, Villiers (1950b) noted their color pattern was pale brownish gray, the dorsal scales dotted with black, the back with alternate dark brown diamond spots (some ocellated with white). Yellowish ventral surface with two black lateral lines. Doucet (1963) noted the dorsal coloration ranged from uniform reddish or yellowish or powdery brown, sometimes with dark bars, with ventral coloration ranging from yellowish to pinkish. Chippaux (2006) noted dorsal coloration as dark beige or reddish with occasional dark gray designs or crossbars; venter pinkish with two dark lateral lines. Based on photographs of an adult male from Guinea (Fig. 11B), the base of the tongue is orangish red, and the forked tip is silvery white with black edging. Our observations of live specimens suggest the dorsal scales have an almost satiny sheen, similar to the appearance of a spider web (MOR, pers. obs.). Hemipenis. Bogert (1940) did not note whether his hemipenis description was based on specimens from Liberia (T. pulverulenta) or former French Cameroon (T. adamanteus sp. nov.). Doucet (1963:301) described the hemipenis of a snake from Ivory Coast as very different from that of B. [Toxicodryas] blandingi. Quite elongated, thorny at the base and dimpled at the apex, it is not bifid. The spines on either side of the sperm groove are very long in the middle part and decrease in size towards the apex and the base. The apex is domed, not flattened. Diet. Villiers (1950a) noted both species of Toxicodryas in West Africa feed mainly on birds. Cansdale (1954) reported an individual from Ghana with two young mice in its stomach. Johnsen (1962:121) reported an adult male from Liberia with ���a small insect-eating bird��� in its stomach. In their paper on snakes of Ghana, Leston & Hughes (1968:754) noted the presence of undigested hair and a 10-cm long rodent tail in the stomachs of two specimens, and that ���this suggests the attraction of Cocoa is for its Rodent fauna.��� Another specimen contained the foot and tail of an Agama sp. lizard (sensu Leach�� et al. 2017). B��hme (2000) noted the diet of snakes from Guinea included small mammals (shrews and rodents), adult and nestling birds, and lizards with well-developed limbs. Greene (1989) suggested that there is an ontogenetic shift in diet from mostly lizards in young individuals to mostly birds and/or mammals as adults, and some individuals can ingest a third of their body weight if they eat more than one prey item in a nest or roost. Behavior. In a paper on Guinean snakes, B��hme (2000) classified the species as relatively common and nocturnal. Reproduction. An adult female found in Ghana in early September contained eggs ���about 8 mm maximum breadth and between 30 and 35 mm long��� attached to the left and right ovaries (Leston & Hughes 1968). Habitat. Menzies (1966) documented several specimens from forest in Sierra Leone, but one animal found at Rokupr seemed to occur in a habitat labeled as mangrove and coastal savanna. Leston & Hughes (1968:754) noted that this species was very common in cocoa tree pods in Ghana. Hughes & Barry (1969) listed the habitat as forest. Leston (1970:143) encountered several snakes in Ghana in trees (including one in a rotten cocoa pod), ���amidst old files on an office shelf,��� on a 1.5-meter high cocoa tree in a greenhouse, and in savanna. Hughes (1988) noted the species only from rainforest in Ghana. R��del & Mahsberg (2000) found snakes in Ivory Coast from thick undergrowth in a swamp bordered by fields and secondary forest, and swampy primary forest. Branch & R��del (2003) captured a juvenile in a funnel trap on the ground in closed-canopy forest in Ivory Coast. In Togo, Segniagbeto et al. (2011) noted the species is most common in forest, but some specimens were found outside of it, presumably in savanna. Geographic distribution and habitat. Based on molecular data from Allen et al. (in press) and patterns of our morphometric data (Table 2), we hypothesize that this species occurs west of the Niger Delta from Guinea to Nigeria. Venom. Venom composition and effects are unknown (Weinstein et al. 2011)., Published as part of Greenbaum, Eli, Allen, Kaitlin E., Vaughan, Eugene R., Pauwels, Olivier S. G., Wallach, Van, Kusamba, Chifundera, Muninga, Wandege M., Aris- Tote, Mwenebatu M., Mali, Franck M. M., Badjedjea, Gabriel, Penner, Johannes, R��del, Mark-Oliver, Rivera, Jacqueline, Sterkhova, Viktoria, Johnson, Grant, Tapondjou, Walter P. & Brown, Rafe M., 2021, Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata Colubridae) with descriptions of two new cryptic species from Central Africa, pp. 1-44 in Zootaxa 4965 (1) on pages 24-27, DOI: 10.11646/zootaxa.4965.1.1, http://zenodo.org/record/4723024, {"references":["Fischer, J. G. (1856) Neue Schlangen des Hamburgischen Naturhistorischen Museums. Abhandlungen aus dem Gebiete der Naturwissenschaften herausgegeben von Naturwissenschaftlichen Verein in Hamburg, 3, 79 - 116 + 3 pl.","Chabanaud, P. (1917 a) Enumeration des ophidiens non encore etudies de l'Afrique occidentale, appartenant aux collections du Museum avec la description des especes et des varietes nouvelles. Bulletin du Museum d'Histoire Naturelle, Paris, 22, 362 - 382. https: // doi. org / 10.5962 / bhl. part. 17132","Chabanaud, P. (1917 b) Note complementaire sur les ophidiens de l'Afrique occidentale, avec la description d'une espece nouvelle. Bulletin du Museum d'Histoire Naturelle, Paris, 23, 7 - 14.","Ceriaco, L. M. P., Marques, M. P. & Bauer, A. M. (2018) Miscellanea Herpetologica Sanctithomae, with a provisional checklist of the terrestrial herpetofauna of Sao Tome, Principe and Annobon islands. Zootaxa, 4387 (1), 091 - 108. https: // doi. org / 10.11646 / zootaxa. 4387.1.4","Hallermann, J. (1998) Annotated catalog of the type specimens of the herpetological collection in the Zoological Museum of the University of Hamburg. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 95, 197 - 223.","Uetz, P., Cherikh, S., Shea, G., Ineich, I., Campbell, P. D., Doronin, I. V., Rosado, J., Wynn, A., Tighe, K. A., McDiarmid, R., Lee, J. L., Kohler, G., Ellis, R., Doughty, P., Raxworthy, C. J., Scheinberg, L., Resetar, A., Sabaj, M., Schneider, G., Franzen, M., Glaw, F., Bohme, W., Schweiger, S., Gemel, R., Couper, P., Amey, A., Dondorp, E., Ofer, G., Meiri, S. & Wallach, V. (2019) A global catalog of primary reptile type specimens. Zootaxa, 4695 (5), 438 - 450. https: // doi. org / 10.11646 / zootaxa. 4695.5.2","Hughes, B. & Barry, D. H. (1969) The snakes of Ghana: a checklist and key. Bulletin de l'Institut fondamental d'Afrique noire. Serie A, Sciences naturelles, 31 (3), 1004 - 1041.","Loveridge, A. (1957) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya, Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology, Harvard, 117 (2), 151 - 362 + i - xxxvi (index).","Chabanaud, P. (1917 c) Revision de quelques reptiles d'Afrique et description de trois especes nouvelles. Bulletin du Museum d'Histoire Naturelle, Paris, 23, 442 - 454.","Demenou, B. B., Pineiro, R. & Hardy, O. J. (2016) Origin and history of the Dahomey Gap separating West and Central African rain forests: Insights from the phylogeography of the legume tree Distemonanthus benthamianus. Journal of Biogeography, 43 (5), 1020 - 1031. https: // doi. org / 10.1111 / jbi. 12688","Bogert, C. M. (1940) Herpetological results of the Vernay Angola expedition. Bulletin of the American Museum of Natural History, 77, 1 - 107, pl. I.","Angel, F. (1933) Les Serpents d'Afrique Occidentale Francaise. Larose Editeur, Paris, 246 pp.","Villiers, A. (1950 a) Initiations Africaines. II. Les Serpents de l'Ouest Africain. Institut Francais d'Afrique Noire, Dakar, Senegal, 148 pp.","Doucet, J. (1963) Les serpents de la Republique de Cote d'Ivoire. Serpents venimeux. Acta Tropica, 20 (3 - 4), 297 - 340.","Stucki-Stirn, M. C. (1979) A Comparative Study of the Herpetological Fauna of the former West Comeroon [sic] / Africa: With a Classification and Synopsis of 95 Different Snakes and Description of Some New Sub-species. Snake Report 721. Herpeto- Verlag, Teuffenthal, Switzerland, 650 pp.","Chippaux, J-P. (2006) Les Serpents d'Afrique Occidentale et Centrale. 3 rd Edition. IRD Editions, Paris, 311 pp.","Villiers, A. (1950 b) Catalogues. IV. La Collection de Serpents de l'I. F. A. N. Institut Francais d'Afrique Noire, Dakar, Senegal, 155 pp.","Segniagbeto, G. H., Trape, J. F., David, P., Ohler, A., Dubois, A. & Glitho, I. A. (2011) The snake fauna of Togo: Systematics, distribution and biogeography, with remarks on selected taxonomic problems. Zoosystema, 33 (3), 325 - 360. https: // doi. org / 10.5252 / z 2011 n 3 a 4","Leeson, F. (1950) Identification of Snakes of the Gold Coast. The Crown Agents for the Colonies, London, x + 142 pp.","Johnsen, P. (1962) Notes on African snakes, mainly from northern Rhodesia and Liberia. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn, 124, 115 - 130.","Malnate, E. V. (1972) Observations on the vertebral hypapophyses and associated musculature in some snakes, with special reference to the Colubridae. Zoologische Mededelingen, 47 (18), 225 - 239.","Gunther, A. (1858) Catalogue of Colubrine Snakes in the Collection of the British Museum. Trustees of the British Museum, London, xvi + 281 pp. https: // doi. org / 10.5962 / bhl. title. 13272","Cansdale, G. (1954) Gold Coast snakes - a complete list. The Nigerian Field, 19, 118 - 132.","Leston, D. & Hughes, B. (1968) The snakes of Tafo, a forest cocoa-farm locality in Ghana. Bulletin de l'Institut Francaise d'Afrique Noire, 30 A, 737 - 770.","Leache, A. D., Grummer, J. A., Miller, M., Krishnan, S., Fujita, M. K., Bohme, W., Schmitz, A., Lebreton, M., Ineich, I., Chirio, L., Ofori-Boateng, C., Eniang, E. A., Greenbaum, E., Rodel, M. - O. & Wagner, P. (2017) Bayesian inference of species diffusion in the West African Agama agama species group (Reptilia, Agamidae). Systematics and Biodiversity, 15 (3), 192 - 203. https: // doi. org / 10.1080 / 14772000.2016.1238018","Bohme, W. (2000) Diversity of a snake community in a Guinean rain forest (Reptilia, Serpentes). In: Rheinwald, G. (Ed.), Isolated Vertebrate Communities in the Tropics. Proceedings of the 4 th International Symposium in Bonn. Bonner zoologische Monographien, 46, 69 - 78.","Greene, H. W. (1989) Ecological, evolutionary, and conservation implications of feeding biology in Old World cat snakes, Genus Boiga (Colubridae). Proceedings of the California Academy of Sciences, 46 (8), 193 - 207.","Menzies, J. I. (1966) The snakes of Sierra Leone. Copeia, 1966 (2), 169 - 179. https: // doi. org / 10.2307 / 1441123","Leston, D. (1970) Some snakes from the forest zone of Ghana. British Journal of Herpetology, 4 (6), 141 - 144.","Hughes, B. (1988) Herpetology in Ghana (West Africa). British Herpetological Society Bulletin, (25), 29 - 37.","Rodel, M. - O. & Mahsberg, D. (2000). Vorlaufige Liste der Schlangen des Tai-Nationalparks / Elfenbeinkuste und angrenzender Gebiete. Salamandra, 36 (1), 25 - 38.","Branch, W. R. & Rodel, M. - O. (2003) Herpetological survey of the Haute Dodo and Cavally forests, western Ivory Coast, Part II: Trapping results and reptiles. Salamandra, 39 (1), 21 - 38.","Weinstein, S. A., Warrell, D. A., White, J. & Keyler, D. E. (2011) \" Venomous \" Bites from Non-Venomous Snakes: A Critical Analysis of Risk and Management of \" Colubrid \" Snake Bites. Elsevier, Amsterdam, Boston, Heidelburg, London, New York, Oxford, Paris, San Diego, San Francisco, Singapore, Sydney and Tokyo, xxvii + 336 pp."]}

Toxicodryas adamanteus sp. nov. (Table 2, Figs. 8, 12–13) We hypothesize that this new, cryptic species occurs in western, central and eastern Africa, east of the Niger Delta, and it has been considered to be conspecific with Toxicodryas pulverulenta since it was first documented to occur in Angola by Peters (1877). The recognition of this new species is supported by evidence from molecular data (Fig. 2) and significant differences in subcaudal scale counts (see Results). Diagnosis. A species of Toxicodryas restricted to West, Central and East Africa, east of the Niger Delta, defined by the following combination of characters: maximum SVL 1 meter in T. blandingii and T. vexator sp. nov.), DSRN 18–23 (vs. 23–25 in T. blandingii and 23–29 in T. vexator sp. nov.), DSRM 18–21 (vs. 21–25 in T. blandingii and T. vexator sp. nov.); cloacal plate undivided (vs. usually divided in T. blandingii, and divided or undivided in T. vexator sp. nov.); both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots (vs. adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters in T. blandingii and T. vexator sp. nov.); hemipenis relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, with a domed apex (vs. hemipenis relatively short and massive [i.e., broad], proximal third covered with spines, distal twothirds dimpled with a flattened apex in T. blandingii and T. vexator sp. nov.). Holotype. UTEP 22204 (field no. ELI 2213; Figs. 8, 12–13), adult male from Npenda village, NE of Lake Tumba (00.7465° S, 18.2243° E, 311 m), Equateur Province, DRC, collected by local Twa people and brought to Eli Greenbaum, Chifundera Kusamba, Wandege M. Muninga, and Mwenebatu M. Aristote on 8 July 2013. Paratopotype. UTEP 22203 (field no. ELI 2212) adult female with same collection details as the holotype. Paratype. RBINS 2699 (formerly RBINS 9127) (field no. Paul Leloup #26), adult female from Région Tshabondo (2.690861° S, 27.341972° E), South Kivu Province, DRC, collected by Paul Leloup on 11 October 1958. Description of the holotype. Adult male, 812 mm SVL; head weakly triangular and moderately distinct from neck, 2.4% of SVL (19.3 mm); interocular distance 13.4 mm, pupil elliptical, eye diameter 5.2 mm; loreal nearrectangular, shorter (1.7 mm) than high (2.0 mm), slightly tapering superiorly and vertically; body semi-triangular; tail moderately long (27.7% of SVL). Supralabials 8/8, 3 rd –5 th /3 rd –5 th contacting orbit; infralabials 13/12, 1 st on each side in contact behind mental, 1 st –5 th /1 st –5 th contacting anterior chin shields and 5 th –7 th /5 th –7 th contacting posterior chin shields; 1/1 preocular; 2/2 postoculars; temporals 2 + 3 + 3/2 + 3 + 3; 2 internasals; nasal divided; frontal width and length equal (6.6 mm); dorsal scale rows 21 one head length posterior to jaw rictus, 19 at midbody, 15 one head length anterior to vent, smooth and oblique with apical pits visible on the neck, vertebral scales broad and apically flattened; ventrals 264 (standard), 262 (Dowling); cloacal plate undivided; paired subcaudals 117; both hemipenes everted. Coloration (in life) of the holotype. Silvery gray-brown over the entire length of the dorsum including the head, with similarly colored but darker diamonds occurring laterally over the full length of the neck, body, and tail. These diamonds are elongated vertically, have pale gray centers, and often have black spots at the top and bottom corners. Between each diamond on the body and neck is a more vaguely defined, pale gray diamond or stripe. The lateral and antero-dorsal sides of the head are speckled, as is the ventral side of the head, neck, body, and tail. The venter is otherwise white, with the speckles forming two pale stripes running down either side of the ventrals and subcaudals. The base of the tongue is orangish red, and the forked tip is silvery white with black edging. Coloration (in preservative) of the holotype. This specimen’s coloration has become slightly darker and browner than it was in life, with less pronounced patterning on the flanks, but otherwise the appearance is similar to the coloration in life. Variation. Morphometric variation of Toxicodryas adamanteus sp. nov. is shown in Table 2. We observed extensive temporal scale variation, including 1 + 1, 1 + 2, 2 + 2 (most commonly), 2 + 3, 2 + 5, 3 + 2, 3 + 3, and 4 + 3. Chabanaud (1917c) described a male from Gabon with supralabials 3–6 contacting the eye, and this specimen seems to be the size record at 1,225 mm total length (995 mm SVL, 230 mm tail length). Schmidt (1923) listed ventral counts of 251–269 in snakes from DRC, and noticed one individual that had a preocular fused to the supraocular on one side, and in two individuals, the loreal was fused to the lower portion of the preocular and thus contacted the eye. Laurent (1956) noticed that snakes from DRC (i.e., T. adamanteus sp. nov.) had more subcaudals than snakes from West Africa (i.e., T. pulverulenta). Skinner (1973) noted ventral scale counts range from 240–269, subcaudal scale counts range from 105–126 (identical to Pitman 1974), and the maximum size is about 2 meters, substantially larger than all other published records, and thus, highly doubtful. De Witte (1975) provided data for ventral scale counts of DRC snakes ranging from 235–242 (males) and 239–249 (females), and subcaudal counts of 112–120 (both sexes). Spawls & Branch (2020) listed the maximum size as “about 1.25 m ” but no specific locality or record was provided. Although most of our examined specimens and literature records noted the 3 rd to 5 th supralabial in contact with the eye, some individuals have the 4 th to 6 th supralabial in contact with the eye (e.g., Loveridge 1937) and one specimen from Gabon (CAS 258155) had only the 4 th and 5 th supralabial in contact with the eye. In snakes from Uganda, Pitman (1974) reported ventral scale counts of 240–269 and subcaudal counts of 105–126 (not distinguished by sex). In de Witte’s (1975) study of snakes from Virunga National Park in eastern DRC, he noted ventrals range from 235–242 in males and 239–249 in females; subcaudals ranged from 112–120 in both sexes. Rasmussen (1997b:106) described individuals with 3 or 4 postoculars, 7–9 supralabials, 9–13 infralabials, 236–278 ventrals, and 96–132 subcaudals (sometimes undivided). In general, he noted this species has sloping and smooth scales with apical pits, and the vertebral row is more or less enlarged. Bogert (1940:fig. 8) illustrated the maxillary teeth of a specimen (AMNH 50590) from former French Cameroon, noting “ five specimens examined show variation from eleven to thirteen anterior subequal teeth followed after a very short diastema by two larger grooved fangs and a smaller fang, the total number of teeth being fourteen to eighteen.” Because only two of his examined specimens originated from former French Cameroon, at least three of these specimens are from Liberia, which are attributable to T. pulverulenta. Schmidt (1923:103) quoted field notes of Herbert Lang for DRC snakes as “coloration, in life, reddish brown above, head darker brown. Irregular dark gray lateral bars, wider in the middle, extend from the vertebral line to the venter, tipped above and below with black. A cream-colored central spot in the broad portion of each lateral bar. Faint narrow grayish crossbars between the wider ones, disappearing posteriorly. The wider crossbars are usually alternate, sometimes confluent on the back. Venter pinkish gray, heavily dotted with brown which forms two lines at the inner edges of the ventral edges of the ventral angle. These lines are more distinct beneath the tail.” Hellmich (1957b) described several individuals from Angola with a reddish brown to reddish blue-gray dorsum, with varying degrees of contrasting rhombic patterns and spots. Pitman (1974) provided a similar description for specimens from Uganda, noting they range from pinkish-brown to reddish-brown. Hedges (1983:21) described Kenyan specimens as, “a rather pretty pattern of coffee coloured diamond shaped markings on a mushroom pink body.” Rasmussen (1997b) described coloration in this species (presumably based on examined specimens from Nigeria and Cameroon) as reddish brown on the dorsum of the body; dorsum of the head darker brown; irregular, dark gray, diamondshaped spots on the flanks that are widest at mid-body; spots edged in black on superior and inferior edges with a cream spot in the center that sometimes fuse together in the midline; faint grayish transverse bands between spots that fade and disappear posteriorly; venter pinkish-gray, with dense brown spots that form a line just inside and parallel to the keeled ventrals, most salient on the tail. Pauwels et al. (2019b) noted a specimen from Gabon with an orange tongue in life. Spawls & Branch (2020:239) noted the dorsal color can range from pinkish to brown, redbrown or pinky gray with “darker” cross-bars that enclose a “pale” spot. The dorsum and venter are “finely dusted” with brown or black specks, whereas the venter is pale pink with “dashed dark lines” on each side of the ventrals. Contrary to Pauwels et al. (2019b), they noted the tongue is pink with a white tip. Based on our photographs of DRC snakes (e.g., Fig. 12G), the tongue is orangish red, and the forked tip is silvery white with black edging, which is consistent with the coloration of the holotype in life (EG pers. obs.). An unsexed individual from Banalia, DRC (Fig. 12F) is unique in having a golden yellow mid-dorsal stripe. Diet. Pitman (1938) mentioned a juvenile from Uganda that had a small mouse in its stomach. In Kenya, Hedges (1983) noted the species eats small chameleons, geckos, and frogs. Butler & Reid (1990) described an adult female from Nigeria that contained the remains of an Agama sp. lizard (sensu Leaché et al. 2017). Spawls & Branch (2020) listed the diet as arboreal lizards and rodents. Parasites. Pitman (1938) noted a juvenile from Uganda with numerous ticks. Behavior. Pitman (1974:128) described the species as “arboreal, nocturnal... amiable disposition,” and suggested that it might rely on camouflage to avoid predation, a sentiment also suggested by Gans (1961). Chirio & LeBreton (2007) suggested the species is shy and rarely bites. Spawls & Branch (2020) noted that when threatened, it elevates the anterior part of its body, “flickers its red tongue,” hisses, and strikes. Reproduction. Schmidt (1923) described a DRC female collected in June 1914 with an unspecified number of eggs that measured 11 x 29 mm. Butler & Reid (1990) described an adult female from Nigeria (captured 21 January 1988) that contained two developing eggs (34–35 mm long, 11 mm wide). Branch (2005) noted the species lays 2– 5 eggs. Habitat. Parker (1936) noted this species is restricted to rain forest and “its outliers.” Perret (1961) listed the species from forest in Cameroon. De Witte (1962) listed the habitats for DRC snakes as equatorial forest and gallery forest, and this classification was repeated by Thys van den Audenaerde (1965). Blackwell (1967) listed the species from forest, thickets, and gardens in Nigeria. Pitman (1974:128) described this snake as a “forest species” in Uganda, but also noted one individual from a “fowl house,” and another from an epiphytic fern (Platycerium angolense) on a forest tree. Butler & Reid (1986) listed the species only from forest in Nigeria. Lawson (1993) documented the species from forest and “farmbush” in Cameroon. Rasmussen (1997b) described the species as nocturnal and restricted to forest. Chirio & LeBreton (2007) recorded the species from forests and savanna-forest mosaic in Cameroon. Spawls & Branch (2020) included forest, woodland, and forest-savanna mosaic from sea level to about 2,000 m elevation. Among the 19 snake species recorded within the garden of a villa in Yenzi ( 2.77261° S, 10.03403° E), Gamba, Nyanga Province, southwestern Gabon (forest-savanna mosaic), inhabited by one of the authors from 2004 to 2011 (OSGP, unpubl. data), T. adamanteus sp. nov. was among the most rarely encountered species. Geographic distribution and habitat. Based on molecular data from Allen et al. (in press) and patterns of our morphometric data (Table 2), we hypothesize that this species occurs east of the Niger Delta in Nigeria, Cameroon, Equatorial Guinea, Gabon, CAR, Republic of Congo, DRC, Angola, Uganda, South Sudan, and Kenya. Ullenbruch & Böhme (2017) recently listed the species from South Sudan. Venom. Based on a specimen from Cameroon, Taub (1967) described the histological morphology of the Duvernoy’s gland. Venom composition and effects are unknown (Weinstein et al. 2011). Etymology. The specific epithet adamanteus is a Latin adjective referring to the diamond-shaped marks on the flanks and dorsum of this species.

Toxicodryas vexator sp. nov. (Table 1, Figs. 1, 8���10) We hypothesize that this new, cryptic species occurs east of the confluence of the Congo and Ubangi rivers, and it has been considered to be conspecific with Toxicodryas blandingii since it was first documented to occur east of these rivers in DRC (Boulenger 1919). The recognition of this new species is supported by molecular data (Fig. 2), several significant differences in scale counts (see Results), and less toxic venom in comparison to its sister taxon T. blandingii (see Venom). Diagnosis. A species of Toxicodryas restricted to east-central and East Africa (east of the confluence of the Congo and Ubangi rivers), defined by the following combination of characters: maximum SVL> 1 meter (vs. maximum SVL T. pulverulenta and T. adamanteus sp. nov.); DSRN 23���29 (vs. 19���21 in T. pulverulenta and 18���23 in T. adamanteus sp. nov.); DSRM 21���25 (vs. 19���21 in T. pulverulenta and 18���21 in T. adamanteus sp. nov.); cloacal plate divided or undivided (vs. usually divided in T. blandingii, and always undivided in T. pulverulenta and T. adamanteus sp. nov.); adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters (vs. both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots in T. pulverulenta and T. adamanteus sp. nov.); hemipenis relatively short and massive (i.e., broad), proximal third covered with spines, distal two-thirds dimpled with a flattened apex (vs. relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, and with a domed apex in T. pulverulenta and T. adamanteus sp. nov.); venom toxicity LD 50 = 4.88 mg /kg in mice (vs. venom toxicity LD 50 = 2.85���3.55 mg /kg in mice for T. blandingii). Holotype. UTEP 22196 (field number MUSE 10341; Fig. 10), adult male collected in Mulisi, Nzovu Est, Kahuzi-Biega National Park (02.447291�� S, 28.2825378�� E, 1101 m), South Kivu Province, DRC, collected by Guillain M. Mitamba, Deo Kujirakwinja, Emmanuel Muhindo, Radar Nushili, Wandege M. Muninga, and Andrew J. Plumptre on 13 November 2015. Paratypes. UTEP 22195 (field number EBG 1362; Figs. 1B, 9C), adult female collected in the vicinity of Irangi (01.8780�� S, 28.4524�� E, 811 m), South Kivu Province, DRC, collected from a tree by a resident and brought to Maurice Luhumyo, Chifundera Kusamba, Mwenebatu M. Aristote, Wandege M. Muninga, and Eli Greenbaum on 30 August 2007; RBINS 2698 (formerly RBINS 8621) (field no. Leloup #27), adult male from Bunyakiri (2.075630�� S, 28.573194�� E, 1000 m), South Kivu Province, DRC, collected by Paul Leloup on 6 March 1958. Description of the holotype. Adult male, 1438 mm SVL; head strongly triangular and distinct from the neck, 2.1% of SVL (30.9 mm); interocular distance 21.7 mm, pupil elliptical, maximum horizontal eye diameter 7.5 mm; loreal near-rectangular, longer (3.0 mm) than high (2.4 mm), slightly tapering superiorly; body triangular; tail moderately long (32.1% of SVL). Supralabials 9/9, 4 th, 5 th, and 6 th /4 th, 5 th, and 6 th contacting orbit; infralabials 14/14, 1 st on each side in contact behind mental, 1 st ���4 th /1 st ���4 th contacting anterior chin shields, 5 th ���8 th /5 th ���8 th contacting posterior chin shields, and 1 cuneate between the 5 th and 6 th left infralabials; 2/2 preoculars; 2/2 postoculars; temporals 2 + 2/2 + 2; 2 internasals; nasal divided on both sides; frontal slightly wider (10.1 mm) than long (9.2 mm); dorsal scale rows 23 one head length posterior to jaw rictus, 21 at midbody, 15 one head length anterior to vent, smooth and oblique with apical pits visible on the neck, vertebral scales broad and apically flattened; ventrals 261 (standard), 260 (Dowling); cloacal plate undivided; paired subcaudals 136 (terminal spine missing); both hemipenes everted. Coloration (in life) of the holotype. Solid black over the entire length of the dorsum, and black ventrally except for the anterior half of the body, where the ventral scales are pale yellow with a black posterior edge that increases in thickness in correlation with increased distance from the head. The head is black dorsally and laterally except for yellow on the lower, anterior region of the 5 th ���9 th supralabials, and yellow ventrally except for thin black rims on the posterior edges of the infralabials. Coloration (in preservative) of the holotype. The specimen���s coloration in preservative is nearly the same as in life, although the ventral yellow coloration has dulled to a yellowish off-white; some scales also seem cloudy under close lighting, indicative of some stage of ecdysis in life. Variation. For years, the size record for this species was attributed to Goodman (1985), who recorded a total length of 2515 mm for an adult female collected in Uganda. However, Hedges (1983:20) measured a specimen from Kenya that was 2707 mm total length, and said it ���is reputed to exceed 3300 mm.��� Weinstein et al. (2011) noted the species may exceed 3.5 meters. Spawls & Branch (1995, 2020) listed the maximum size as ���about 2.8 m ��� (no specific location provided), but Weinstein (pers. comm.) estimated a captive specimen from Kakamega was approximately 3.0 meters total length. Schmidt (1923:104) remarked that two specimens from DRC had fused prefrontals, creating a single transverse scale. Five of his 20 DRC specimens had a divided cloacal plate, ���with indications of a groove in two others,��� and as a result, he stated that the condition of the cloacal plate is ���obviously useless��� as a diagnostic character. Laurent (1956) noted that, with the exception of DRC specimens from Mayombe (north of the Congo River in Kongo Central Province and thus attributable to T. blandingii as recognized herein) with consistently divided cloacal plates, 13 of 31 snakes from other areas of DRC had an undivided cloacal plate. Our examined specimens confirm this sentiment, because 17/ 92 specimens have a divided cloacal plate, and an additional five specimens have a partially divided cloacal plate. Hellmich (1957b) noted maximum sizes of Angolan males (1740 mm SVL; 523 mm TL) and females (1730 mm SVL; 505 mm TL), and aberrant individuals that had 11 supralabials and 9���14 infralabials. De Witte (1966) noted that the species has 17 scale rows at midbody, but this datum is so aberrant that it is likely an error (Pitman 1974; Table 1). Skinner (1973) provided ranges of scale counts (161���274 ventrals, 86���147 subcaudals) that are so aberrant that they are clearly erroneous (at least for lower counts), and noted a maximum length of about 3 meters. For Uganda snakes, Pitman (1974) listed ventral scale counts from 240���260 (males) and 240���259 (females). De Witte (1975) noted a single individual with 25 scale rows at midbody, and Broadley et al. (2003) also noted 25 scale rows as the maximum amount of variation. Based on two specimens from Lukolela, Belgian Congo (AMNH 45907) and Akenge, Belgian Congo (AMNH 12243), Bogert (1940:61) described their maxillary teeth as ���ten in number, followed after a short diastema by three enlarged, grooved fangs, the posterior one of which is smaller than the other two.��� Schmidt (1923:105) described two color phases of DRC specimens, including (1) a black phase with the anterior portion of the venter yellow, each ventral scale bordered with black on its posterior edge, the border increasing in width until the yellow color disappears on the posterior two-thirds of the venter; and (2) a brownish phase, ���with more or less distinct wide dark cross-bars, confluent anteriorly, alternate posteriorly on the vertebral line.��� Pitman (1938:211) described an unsexed individual from Uganda as ���darkish gunmetal with a purplish mottling towards the tail, belly pale yellow from head to tail but with slight brown markings on the posterior half increasing in occurrence towards and on the tail.��� A female from Uganda was described as rich chestnut dorsally and ventrally, with handsome chocolate blotches on the flanks, each one containing a small white spot. On the posterior part of the body, including most of the tail, these blotches were confluent with pale edging. The ventral aspect of the head and the first 50 ventrals were ���paly yellowish tinged green.��� The dorsum of the head was dark brown, and the posterior supralabials were dull greenish grey. The supralabials had black edging posteriorly, except for the last one. There were two elongated dark brown blotches behind the eye, which was iridescent hazel. Bogert (1940:61) provided data for a specimen (AMNH 45907; determined to be a subadult male based on photographs examined by EG) that had been described by James P. Chapin as ���olive-brown, with dark brown patches; below grayish brown.��� Hellmich (1957b) described the coloration of several individuals from Angola, including an adult male that had a blue-reddish-black dorsum with a venter that was pale yellow in the front half of the body, with an ever-widening blue-black border that eventually enclosed the entire ventral scales towards the posterior side. Another adult male had a dark black-brown dorsum, and light yellow venter on the first third of the body, but starting on the 5 th ventral, there was a dark gray-blue spot that increased in size posteriorly until it covered the entire venter towards the posterior end of the body. An adult female���s dorsum was described as clay yellow with mostly alternating dark transverse bars and a pale yellow venter. A second adult female had similar coloration to the latter specimen, but the transverse bands were only faintly visible on the posterior sixth of the body and tail. A juvenile dorsum was described as gray-brown with transverse black-brown spots that converge in the anterior part of the body, but then alternate posteriorly, with a light yellowish-white spot in the lower center of the spots. Laurent (1956:195) listed a section about possible sexual dimorphism for this species, but stated that nothing glaring appears in his data. Pitman (1974:125) described several individuals from Uganda that were similar to the descriptions above, and added that both color phases ���exhibit a handsome suede effect.��� He noted that although subadult males might retain the brown phase, black females are unknown, and the black and brown color phases have ���sexual significance.��� The dark blotches of juveniles were noted to vary from ���blackish, chocolate, reddishbrown or dark or bright chestnut,��� with interspaces ranging from light gray, pale brown or chestnut, sometimes with a pinkish tinge. Goodman (1985:56) described two adult females from Uganda as ���dingy olive-brown.��� Hughes (2000) noted that Laurent (1964) seemed to imply that his largest male specimen from Dundo, Angola was not black, but this is not clear, because the latter author only mentioned in passing that a smaller male from Andrada, Angola represented the black phase. Spawls & Branch (1995, 2020) remarked that the yellow ventral pigmentation might form a stripe in the middle of the venter. In the brown color morph, the skin between the scales is bluish gray, and especially visible when the snake inflates its body during a threat display. The eye can be yellowish or brown. The black phase snakes are usually male, whereas the brown phase ones are usually female. Based on photos of an adult male from Banalia, DRC (individual shown in Fig. 9A), the base of the tongue is bluish black, and the forked tip is silvery gray. Hemipenis. Bogert (1940:61) described the hemipenis of an individual, presumably from DRC, as ���not bifurcate, extending to the tenth caudal, the undivided sulcus deeply buried between two fleshy fold [sic]. The basal half is heavily armed with stout spines but distally the organ is calyculate, the edges of the calyces crenulated.��� Hemipenes of our examined specimens had a simple, subcylindrical shape, simple sulcus spermaticus, and spinose ornamentation with a rough apical structure, as also noted for T. blandingii. Because only two specimens with everted hemipenes were available for both T. blandingii and T. vexator sp. nov., any finer-scale differences cannot be attributed to species-level differentiation (Dowling & Savage 1960; Doucet 1963). Diet. Laurent (1964) noted a juvenile snake contained the remains of a bird. In snakes from Uganda, Pitman (1974) noted the diet included weaver finches, robber birds��� eggs, small rodents, bats, frogs, large agamids, and chameleons. Hedges (1983) described a Kenyan snake that regurgitated a Nectarinia famosa sunbird. Spawls & Branch (1995, 2020) described the diet as birds, bird eggs, arboreal lizards, frogs, arboreal rodents, and bats. Nagy et al. (2011) documented an adult female from DRC that predated a Short-palated Fruit Bat (Casinycteris argynnis) from a bat net in secondary forest. Another male specimen from DRC was captured with a nestling bushshrike (Laniarius sp.) in its stomach. Parasites. Loveridge (1937) noted a tick on the scales of a specimen (ANSP 20504) from Belgian Congo. Parasites of Ugandan snakes included internal nematodes (Kalicephalus sp.) and cestodes, ectoparasites (ticks, including Aponomma latum), and captive specimens had bacterial infections, including mouth rot (Pitman, 1974). Goodman (1985:56) documented dozens of ticks, ���probably Aponomma sp.,��� and linguatulid worms from the posterior saccular lung cavity of an adult female from Uganda. Behavior. Spawls & Branch (1995, 2020) described the snake as arboreal, sometimes climbing to heights of 30 meters in trees, but also descending to the ground to cross open spaces and roads. It is mostly nocturnal, sheltering in leaf clumps and tree hollows during the day. When threatened, it will open its mouth widely and expose the pink lining inside, inflate its body, flatten the head, lift the anterior part of its body off the ground and into C-shaped coils, and strike at perceived threats. It is adept at smelling sleeping birds in nests at night, and will make a slow, deliberate approach to attack them. Reproduction. A personal communication from ���Leakey��� (most likely Richard Leakey) to Pitman (1974) noted that a female from Kakamega (Kenya) laid 9 eggs that were approximately 20 x 40 mm on 9 January. Spawls & Branch (1995) reported an average of 7��� 14 eggs in this species (3���14 in Spawls & Branch 2020), presumably based mostly on East African data, which is substantially more than clutch sizes reported for T. blandingii by Luiselli et al. (1998a). Habitat. Laurent (1954) remarked that an adult female from Dundo, Angola was captured after it fell out of a mango tree. Laurent (1960) stated that his colleague M. Leloup often encountered the species in eastern DRC. De Witte (1962) listed the habitats for DRC snakes as equatorial forest and gallery forest, a classification that was repeated by Thys van den Audenaerde (1965). Pitman (1974) also limited the species to forest in Uganda, but he remarked that the species frequently enters houses (where it hunts for bats), and one individual was killed in a tree over 20 meters above the ground. Broadley & Cotterill (2004) stated the species inhabits forests and wooded savannas. Spawls & Branch (1995, 2020) noted the species from forest, woodland, forest-savanna mosaic, riverine woodland, and human habitations from sea level to about 2,200 m elevation. The holotype was found basking on a shrub (ca. 4 meters above ground) in a clearing created by a fallen tree in primary forest. The Irangi paratype was found in secondary forest near a road. Geographic distribution. Based on molecular data from Allen et al. (in press) and patterns of our morphometric data (Table 1), we hypothesize that this species occurs east of the Congo and Ubangi Rivers in CAR, DRC, Angola, Zambia, South Sudan, Uganda, Kenya, and Tanzania. Although Boulenger (1896) listed a specimen from ��� Zanzibar,��� the locality likely referred to mainland records (Parker et al. 1940; Loveridge 1957; Pitman 1974). Venom. Wakeman (1966) documented a 2.59 m (presumably total length) captive adult from Uganda that struck an adult rat, which died after 8.5 minutes. Pitman (1974:127) described the case of a young European who was bitten on the thumb three times in rapid succession by a small (305 mm) individual (erroneously reported as 350 mm by Goodman 1985); only ���initial smarting��� at the puncture sites occurred. Spawls (1979) described the bite of a captive 2.1 meter snake on a man, including three fang punctures on his thumb, which resulted in mild swelling of his hand. Anecdotal observations of a bite by a captive snake on a mouse resulted in death in under a minute. Hedges (1983:20) was bitten, ���and although it drew a considerable amount of blood, I felt no effects whatsoever.��� He also remarked that this species ���chews��� when biting, ostensibly to ensure the rear fangs penetrate the flesh to inject venom. Based on venom collected from a snake originating from Kenya, Weinstein & Smith (1993) estimated the LD 50 value as 4.88 mg /kg in mice. Weinstein & Smith (1993:88) also noted ���the differences in lethal potencies between the two samples of B. blandingi [compare with LD 50 = 2.85���3.55 mg /kg in T. blandingii from West Africa] demonstrate variation of secretion toxicity in this species,��� suggesting possible species-level differences in venom composition of these sister taxa (Weinstein & Kardong 1994). Immunological cross reactivity with elapid venom antisera was demonstrated, but Weinstein & Smith (1993) did not distinguish between samples originating from West Africa (i.e., T. blandingii) or East Africa (i.e., T. vexator sp. nov.). Branch (2005) suggested bites cause nausea and headaches, but fatalities are unknown. Dashevsky et al. (2018) included a Toxicodryas from Tanzania in their proteomics study, and they noted its venom is dominated by three-finger toxins. Scott Weinstein (pers. comm., October 2020) clarified that a snakebite mentioned by Weinstein & Smith (1993) and a second one that occurred one year after this publication were caused by snakes originating from Kakamega forest, Kenya. ���The bite mentioned in the paper was delivered by [an approximately] 3.0 m jet black male that was a favourite member of my collection for nearly 7 years. He typically performed the whole gamut of open mouthed posturing and mock strikes, but was also a very aggressive feeder, and although I tried to condition him to receiving food (freeze-thawed chicks and rodents) out of his enclosure, he still occasionally would ���pour��� ou, Published as part of Greenbaum, Eli, Allen, Kaitlin E., Vaughan, Eugene R., Pauwels, Olivier S. G., Wallach, Van, Kusamba, Chifundera, Muninga, Wandege M., Aris- Tote, Mwenebatu M., Mali, Franck M. M., Badjedjea, Gabriel, Penner, Johannes, R��del, Mark-Oliver, Rivera, Jacqueline, Sterkhova, Viktoria, Johnson, Grant, Tapondjou, Walter P. & Brown, Rafe M., 2021, Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata Colubridae) with descriptions of two new cryptic species from Central Africa, pp. 1-44 in Zootaxa 4965 (1) on pages 17-21, DOI: 10.11646/zootaxa.4965.1.1, http://zenodo.org/record/4723024, {"references":["Boulenger, G. A. (1919) Batraciens et reptiles recueillis par le Dr C. Christy au Congo Belge dans le districts de Stanleyville, Haut-Uele et Ituri en 1912 - 1914. Revue Zoologique Africaine, 7, 1 - 29. https: // doi. org / 10.5962 / bhl. part. 15108","Goodman, J. D. (1985) Two record size Blanding's tree snakes from Uganda. East African Natural History Society Bulletin, 1985, 56 - 57.","Hedges, N. G. (1983) Reptiles and Amphibians of East Africa. Kenya Literature Bureau, Nairobi, Kenya, xii + 139 pp.","Weinstein, S. A., Warrell, D. 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Edition Chimaira, Frankfurt am Main, 280 pp.","Bogert, C. M. (1940) Herpetological results of the Vernay Angola expedition. Bulletin of the American Museum of Natural History, 77, 1 - 107, pl. I.","Pitman, C. R. S. (1938) A guide to the snakes of Uganda. Part XI. (Conclusion). The Uganda Journal, 5, 160 - 244, pls. XV - XVIII.","Hughes, B. (2000) Polymorphism, sex, size and other aspects of the African snake, Boiga blandingi (Hallowell, 1844). Herpetological Bulletin, 74, 8 - 12.","Laurent, R. F. (1964) Subsidos para o estudo da biologia na Lunda, Museo do Dundo. Reptiles et amphibiens de l'Angola (Troisieme contribution). Publicacoes Culturais da Companhia de Diamantes de Angola, 67, 17 - 165.","Dowling, H. G. & Savage, J. M. (1960) A guide to the snake hemipenis: A survey of basic structure and systematic characteristics. Zoologica, 45 (Part 1), 17 - 28, pls. I - III.","Doucet, J. (1963) Les serpents de la Republique de Cote d'Ivoire. Serpents venimeux. Acta Tropica, 20 (3 - 4), 297 - 340.","Nagy, Z. T., Chifundera Kusamba, Z., Gembu Tungaluna, G. - C., Lotana Lokasola, A., Kolby, J. & Kielgast, J. (2011) Foraging acrobatics of Toxicodryas blandingii in the Democratic Republic of the Congo. Herpetology Notes, 4, 91 - 92.","Loveridge, A. (1937) Zoological results of the George Vanderbilt African expedition of 1934. Part VII - Reptiles and amphibians. Proceedings of the Academy of Natural Sciences of Philadelphia, 89, 265 - 296.","Luiselli, L., Akani, G. C. & Barieenee, I. F. (1998 a) Observations on habitat, reproduction and feeding of Boiga blandingi (Colubridae) in south-eastern Nigeria. Amphibia-Reptilia, 19 (4), 430 - 436. https: // doi. org / 10.1163 / 156853898 X 00098","Laurent, R. F. (1954) Reptiles et batraciens de la region de Dundo (Angola). Deuxieme note. Publicacoes Culturais da Compan- hia de Diamantes de Angola, 23, 35 - 84.","Laurent, R. F. (1960) Notes complementaires sur les cheloniens et les ophidiens du Congo Oriental. Annales du Musee Royal du Congo Belge, Serie in- 8 °, 84, 1 - 86.","de Witte, G. - F. (1962) Genera des serpents du Congo et du Ruanda-Urundi. Annales du Musee Royal de l'Afrique Centrale, Sciences Zoologique, Serie in Octavo, 104, i - viii + 1 - 203.","Thys van den Audenaerde, D. F. E. (1965) Les serpents des environs de Leopoldville. Revue de zoologie et de botanique africaines, 72 (3 - 4), 366 - 388.","Broadley, D. G. & Cotterill, F. P. D. (2004) The reptiles of southeast Katanga, an overlooked ' hot spot. ' African Journal of Herpetology, 53 (1), 35 - 61. https: // doi. org / 10.1080 / 21564574.2004.9635497","Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History). Vol. III. Containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae, and Viperidae. Trustees of the British Museum (Natural History), London, xiv + 727 pp.","Parker, H. W., Moreau, R. E. & Pakenham, R. H. W. (1940) XXV. - Notes on the herpetology of the East African Islands. Journal of Natural History, 5 (27), 309 - 314. https: // doi. org / 10.1080 / 00222934008527048","Loveridge, A. (1957) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya, Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology, Harvard, 117 (2), 151 - 362 + i - xxxvi (index).","Wakeman, B. N. (1966) Uganda's poisonous snakes: Further observations on feeding habits. The Uganda Journal, 30, 101 - 103","Spawls, S. (1979) Sun, Sand & Snakes. William Morrow and Company, Inc., New York, New York, 254 pp.","Weinstein, S. A. & Smith, L. A. (1993) Chromatographic profiles and properties of Duvernoy's secretions from some boigine and dispholidine colubrids. Herpetologica, 49 (1), 78 - 94.","Weinstein, S. A. & Kardong, K. V. (1994) Properties of Duvernoy's secretions from opisthoglyphous and aglyphous colubrid snakes. Toxicon, 32 (10), 1161 - 1185. https: // doi. org / 10.1016 / 0041 - 0101 (94) 90347 - 6","Branch, W. R. (2005) A Photographic Guide: Snakes Other Reptiles and Amphibians of East Africa. Ralph Curtis Publishing, Inc., Sanibel Island, Florida, 132 pp.","Dashevsky, D., Debono, J., Rokyta, D., Nouwens, A., Josh, P. & Fry, B. G. (2018) Three-finger toxin diversification in the venoms of cat-eye snakes (Colubridae: Boiga). Journal of Molecular Evolution, 86, 531 - 545. https: // doi. org / 10.1007 / s 00239 - 018 - 9864 - 6"]}

Toxicodryas vexator sp. nov. (Table 1, Figs. 1, 8–10) We hypothesize that this new, cryptic species occurs east of the confluence of the Congo and Ubangi rivers, and it has been considered to be conspecific with Toxicodryas blandingii since it was first documented to occur east of these rivers in DRC (Boulenger 1919). The recognition of this new species is supported by molecular data (Fig. 2), several significant differences in scale counts (see Results), and less toxic venom in comparison to its sister taxon T. blandingii (see Venom). Diagnosis. A species of Toxicodryas restricted to east-central and East Africa (east of the confluence of the Congo and Ubangi rivers), defined by the following combination of characters: maximum SVL> 1 meter (vs. maximum SVL T. pulverulenta and T. adamanteus sp. nov.); DSRN 23–29 (vs. 19–21 in T. pulverulenta and 18–23 in T. adamanteus sp. nov.); DSRM 21–25 (vs. 19–21 in T. pulverulenta and 18–21 in T. adamanteus sp. nov.); cloacal plate divided or undivided (vs. usually divided in T. blandingii, and always undivided in T. pulverulenta and T. adamanteus sp. nov.); adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters (vs. both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots in T. pulverulenta and T. adamanteus sp. nov.); hemipenis relatively short and massive (i.e., broad), proximal third covered with spines, distal two-thirds dimpled with a flattened apex (vs. relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, and with a domed apex in T. pulverulenta and T. adamanteus sp. nov.); venom toxicity LD 50 = 4.88 mg /kg in mice (vs. venom toxicity LD 50 = 2.85–3.55 mg /kg in mice for T. blandingii). Holotype. UTEP 22196 (field number MUSE 10341; Fig. 10), adult male collected in Mulisi, Nzovu Est, Kahuzi-Biega National Park (02.447291° S, 28.2825378° E, 1101 m), South Kivu Province, DRC, collected by Guillain M. Mitamba, Deo Kujirakwinja, Emmanuel Muhindo, Radar Nushili, Wandege M. Muninga, and Andrew J. Plumptre on 13 November 2015. Paratypes. UTEP 22195 (field number EBG 1362; Figs. 1B, 9C), adult female collected in the vicinity of Irangi (01.8780° S, 28.4524° E, 811 m), South Kivu Province, DRC, collected from a tree by a resident and brought to Maurice Luhumyo, Chifundera Kusamba, Mwenebatu M. Aristote, Wandege M. Muninga, and Eli Greenbaum on 30 August 2007; RBINS 2698 (formerly RBINS 8621) (field no. Leloup #27), adult male from Bunyakiri (2.075630° S, 28.573194° E, 1000 m), South Kivu Province, DRC, collected by Paul Leloup on 6 March 1958. Description of the holotype. Adult male, 1438 mm SVL; head strongly triangular and distinct from the neck, 2.1% of SVL (30.9 mm); interocular distance 21.7 mm, pupil elliptical, maximum horizontal eye diameter 7.5 mm; loreal near-rectangular, longer (3.0 mm) than high (2.4 mm), slightly tapering superiorly; body triangular; tail moderately long (32.1% of SVL). Supralabials 9/9, 4 th, 5 th, and 6 th /4 th, 5 th, and 6 th contacting orbit; infralabials 14/14, 1 st on each side in contact behind mental, 1 st –4 th /1 st –4 th contacting anterior chin shields, 5 th –8 th /5 th –8 th contacting posterior chin shields, and 1 cuneate between the 5 th and 6 th left infralabials; 2/2 preoculars; 2/2 postoculars; temporals 2 + 2/2 + 2; 2 internasals; nasal divided on both sides; frontal slightly wider (10.1 mm) than long (9.2 mm); dorsal scale rows 23 one head length posterior to jaw rictus, 21 at midbody, 15 one head length anterior to vent, smooth and oblique with apical pits visible on the neck, vertebral scales broad and apically flattened; ventrals 261 (standard), 260 (Dowling); cloacal plate undivided; paired subcaudals 136 (terminal spine missing); both hemipenes everted. Coloration (in life) of the holotype. Solid black over the entire length of the dorsum, and black ventrally except for the anterior half of the body, where the ventral scales are pale yellow with a black posterior edge that increases in thickness in correlation with increased distance from the head. The head is black dorsally and laterally except for yellow on the lower, anterior region of the 5 th –9 th supralabials, and yellow ventrally except for thin black rims on the posterior edges of the infralabials. Coloration (in preservative) of the holotype. The specimen’s coloration in preservative is nearly the same as in life, although the ventral yellow coloration has dulled to a yellowish off-white; some scales also seem cloudy under close lighting, indicative of some stage of ecdysis in life. Variation. For years, the size record for this species was attributed to Goodman (1985), who recorded a total length of 2515 mm for an adult female collected in Uganda. However, Hedges (1983:20) measured a specimen from Kenya that was 2707 mm total length, and said it “is reputed to exceed 3300 mm.” Weinstein et al. (2011) noted the species may exceed 3.5 meters. Spawls & Branch (1995, 2020) listed the maximum size as “about 2.8 m ” (no specific location provided), but Weinstein (pers. comm.) estimated a captive specimen from Kakamega was approximately 3.0 meters total length. Schmidt (1923:104) remarked that two specimens from DRC had fused prefrontals, creating a single transverse scale. Five of his 20 DRC specimens had a divided cloacal plate, “with indications of a groove in two others,” and as a result, he stated that the condition of the cloacal plate is “obviously useless” as a diagnostic character. Laurent (1956) noted that, with the exception of DRC specimens from Mayombe (north of the Congo River in Kongo Central Province and thus attributable to T. blandingii as recognized herein) with consistently divided cloacal plates, 13 of 31 snakes from other areas of DRC had an undivided cloacal plate. Our examined specimens confirm this sentiment, because 17/ 92 specimens have a divided cloacal plate, and an additional five specimens have a partially divided cloacal plate. Hellmich (1957b) noted maximum sizes of Angolan males (1740 mm SVL; 523 mm TL) and females (1730 mm SVL; 505 mm TL), and aberrant individuals that had 11 supralabials and 9–14 infralabials. De Witte (1966) noted that the species has 17 scale rows at midbody, but this datum is so aberrant that it is likely an error (Pitman 1974; Table 1). Skinner (1973) provided ranges of scale counts (161–274 ventrals, 86–147 subcaudals) that are so aberrant that they are clearly erroneous (at least for lower counts), and noted a maximum length of about 3 meters. For Uganda snakes, Pitman (1974) listed ventral scale counts from 240–260 (males) and 240–259 (females). De Witte (1975) noted a single individual with 25 scale rows at midbody, and Broadley et al. (2003) also noted 25 scale rows as the maximum amount of variation. Based on two specimens from Lukolela, Belgian Congo (AMNH 45907) and Akenge, Belgian Congo (AMNH 12243), Bogert (1940:61) described their maxillary teeth as “ten in number, followed after a short diastema by three enlarged, grooved fangs, the posterior one of which is smaller than the other two.” Schmidt (1923:105) described two color phases of DRC specimens, including (1) a black phase with the anterior portion of the venter yellow, each ventral scale bordered with black on its posterior edge, the border increasing in width until the yellow color disappears on the posterior two-thirds of the venter; and (2) a brownish phase, “with more or less distinct wide dark cross-bars, confluent anteriorly, alternate posteriorly on the vertebral line.” Pitman (1938:211) described an unsexed individual from Uganda as “darkish gunmetal with a purplish mottling towards the tail, belly pale yellow from head to tail but with slight brown markings on the posterior half increasing in occurrence towards and on the tail.” A female from Uganda was described as rich chestnut dorsally and ventrally, with handsome chocolate blotches on the flanks, each one containing a small white spot. On the posterior part of the body, including most of the tail, these blotches were confluent with pale edging. The ventral aspect of the head and the first 50 ventrals were “paly yellowish tinged green.” The dorsum of the head was dark brown, and the posterior supralabials were dull greenish grey. The supralabials had black edging posteriorly, except for the last one. There were two elongated dark brown blotches behind the eye, which was iridescent hazel. Bogert (1940:61) provided data for a specimen (AMNH 45907; determined to be a subadult male based on photographs examined by EG) that had been described by James P. Chapin as “olive-brown, with dark brown patches; below grayish brown.” Hellmich (1957b) described the coloration of several individuals from Angola, including an adult male that had a blue-reddish-black dorsum with a venter that was pale yellow in the front half of the body, with an ever-widening blue-black border that eventually enclosed the entire ventral scales towards the posterior side. Another adult male had a dark black-brown dorsum, and light yellow venter on the first third of the body, but starting on the 5 th ventral, there was a dark gray-blue spot that increased in size posteriorly until it covered the entire venter towards the posterior end of the body. An adult female’s dorsum was described as clay yellow with mostly alternating dark transverse bars and a pale yellow venter. A second adult female had similar coloration to the latter specimen, but the transverse bands were only faintly visible on the posterior sixth of the body and tail. A juvenile dorsum was described as gray-brown with transverse black-brown spots that converge in the anterior part of the body, but then alternate posteriorly, with a light yellowish-white spot in the lower center of the spots. Laurent (1956:195) listed a section about possible sexual dimorphism for this species, but stated that nothing glaring appears in his data. Pitman (1974:125) described several individuals from Uganda that were similar to the descriptions above, and added that both color phases “exhibit a handsome suede effect.” He noted that although subadult males might retain the brown phase, black females are unknown, and the black and brown color phases have “sexual significance.” The dark blotches of juveniles were noted to vary from “blackish, chocolate, reddishbrown or dark or bright chestnut,” with interspaces ranging from light gray, pale brown or chestnut, sometimes with a pinkish tinge. Goodman (1985:56) described two adult females from Uganda as “dingy olive-brown.” Hughes (2000) noted that Laurent (1964) seemed to imply that his largest male specimen from Dundo, Angola was not black, but this is not clear, because the latter author only mentioned in passing that a smaller male from Andrada, Angola represented the black phase. Spawls & Branch (1995, 2020) remarked that the yellow ventral pigmentation might form a stripe in the middle of the venter. In the brown color morph, the skin between the scales is bluish gray, and especially visible when the snake inflates its body during a threat display. The eye can be yellowish or brown. The black phase snakes are usually male, whereas the brown phase ones are usually female. Based on photos of an adult male from Banalia, DRC (individual shown in Fig. 9A), the base of the tongue is bluish black, and the forked tip is silvery gray. Hemipenis. Bogert (1940:61) described the hemipenis of an individual, presumably from DRC, as “not bifurcate, extending to the tenth caudal, the undivided sulcus deeply buried between two fleshy fold [sic]. The basal half is heavily armed with stout spines but distally the organ is calyculate, the edges of the calyces crenulated.” Hemipenes of our examined specimens had a simple, subcylindrical shape, simple sulcus spermaticus, and spinose ornamentation with a rough apical structure, as also noted for T. blandingii. Because only two specimens with everted hemipenes were available for both T. blandingii and T. vexator sp. nov., any finer-scale differences cannot be attributed to species-level differentiation (Dowling & Savage 1960; Doucet 1963). Diet. Laurent (1964) noted a juvenile snake contained the remains of a bird. In snakes from Uganda, Pitman (1974) noted the diet included weaver finches, robber birds’ eggs, small rodents, bats, frogs, large agamids, and chameleons. Hedges (1983) described a Kenyan snake that regurgitated a Nectarinia famosa sunbird. Spawls & Branch (1995, 2020) described the diet as birds, bird eggs, arboreal lizards, frogs, arboreal rodents, and bats. Nagy et al. (2011) documented an adult female from DRC that predated a Short-palated Fruit Bat (Casinycteris argynnis) from a bat net in secondary forest. Another male specimen from DRC was captured with a nestling bushshrike (Laniarius sp.) in its stomach. Parasites. Loveridge (1937) noted a tick on the scales of a specimen (ANSP 20504) from Belgian Congo. Parasites of Ugandan snakes included internal nematodes (Kalicephalus sp.) and cestodes, ectoparasites (ticks, including Aponomma latum), and captive specimens had bacterial infections, including mouth rot (Pitman, 1974). Goodman (1985:56) documented dozens of ticks, “probably Aponomma sp.,” and linguatulid worms from the posterior saccular lung cavity of an adult female from Uganda. Behavior. Spawls & Branch (1995, 2020) described the snake as arboreal, sometimes climbing to heights of 30 meters in trees, but also descending to the ground to cross open spaces and roads. It is mostly nocturnal, sheltering in leaf clumps and tree hollows during the day. When threatened, it will open its mouth widely and expose the pink lining inside, inflate its body, flatten the head, lift the anterior part of its body off the ground and into C-shaped coils, and strike at perceived threats. It is adept at smelling sleeping birds in nests at night, and will make a slow, deliberate approach to attack them. Reproduction. A personal communication from “Leakey” (most likely Richard Leakey) to Pitman (1974) noted that a female from Kakamega (Kenya) laid 9 eggs that were approximately 20 x 40 mm on 9 January. Spawls & Branch (1995) reported an average of 7– 14 eggs in this species (3–14 in Spawls & Branch 2020), presumably based mostly on East African data, which is substantially more than clutch sizes reported for T. blandingii by Luiselli et al. (1998a). Habitat. Laurent (1954) remarked that an adult female from Dundo, Angola was captured after it fell out of a mango tree. Laurent (1960) stated that his colleague M. Leloup often encountered the species in eastern DRC. De Witte (1962) listed the habitats for DRC snakes as equatorial forest and gallery forest, a classification that was repeated by Thys van den Audenaerde (1965). Pitman (1974) also limited the species to forest in Uganda, but he remarked that the species frequently enters houses (where it hunts for bats), and one individual was killed in a tree over 20 meters above the ground. Broadley & Cotterill (2004) stated the species inhabits forests and wooded savannas. Spawls & Branch (1995, 2020) noted the species from forest, woodland, forest-savanna mosaic, riverine woodland, and human habitations from sea level to about 2,200 m elevation. The holotype was found basking on a shrub (ca. 4 meters above ground) in a clearing created by a fallen tree in primary forest. The Irangi paratype was found in secondary forest near a road. Geographic distribution. Based on molecular data from Allen et al. (in press) and patterns of our morphometric data (Table 1), we hypothesize that this species occurs east of the Congo and Ubangi Rivers in CAR, DRC, Angola, Zambia, South Sudan, Uganda, Kenya, and Tanzania. Although Boulenger (1896) listed a specimen from “ Zanzibar,” the locality likely referred to mainland records (Parker et al. 1940; Loveridge 1957; Pitman 1974). Venom. Wakeman (1966) documented a 2.59 m (presumably total length) captive adult from Uganda that struck an adult rat, which died after 8.5 minutes. Pitman (1974:127) described the case of a young European who was bitten on the thumb three times in rapid succession by a small (305 mm) individual (erroneously reported as 350 mm by Goodman 1985); only “initial smarting” at the puncture sites occurred. Spawls (1979) described the bite of a captive 2.1 meter snake on a man, including three fang punctures on his thumb, which resulted in mild swelling of his hand. Anecdotal observations of a bite by a captive snake on a mouse resulted in death in under a minute. Hedges (1983:20) was bitten, “and although it drew a considerable amount of blood, I felt no effects whatsoever.” He also remarked that this species “chews” when biting, ostensibly to ensure the rear fangs penetrate the flesh to inject venom. Based on venom collected from a snake originating from Kenya, Weinstein & Smith (1993) estimated the LD 50 value as 4.88 mg /kg in mice. Weinstein & Smith (1993:88) also noted “the differences in lethal potencies between the two samples of B. blandingi [compare with LD 50 = 2.85–3.55 mg /kg in T. blandingii from West Africa] demonstrate variation of secretion toxicity in this species,” suggesting possible species-level differences in venom composition of these sister taxa (Weinstein & Kardong 1994). Immunological cross reactivity with elapid venom antisera was demonstrated, but Weinstein & Smith (1993) did not distinguish between samples originating from West Africa (i.e., T. blandingii) or East Africa (i.e., T. vexator sp. nov.). Branch (2005) suggested bites cause nausea and headaches, but fatalities are unknown. Dashevsky et al. (2018) included a Toxicodryas from Tanzania in their proteomics study, and they noted its venom is dominated by three-finger toxins. Scott Weinstein (pers. comm., October 2020) clarified that a snakebite mentioned by Weinstein & Smith (1993) and a second one that occurred one year after this publication were caused by snakes originating from Kakamega forest, Kenya. “The bite mentioned in the paper was delivered by [an approximately] 3.0 m jet black male that was a favourite member of my collection for nearly 7 years. He typically performed the whole gamut of open mouthed posturing and mock strikes, but was also a very aggressive feeder, and although I tried to condition him to receiving food (freeze-thawed chicks and rodents) out of his enclosure, he still occasionally would ‘pour’ ou

Limnonectes beloncioi, new species urn:lsid:zoobank.org:act: 4CFCEF76-99D6-40DC-9B53- 58DECB1027CF Mindoro Fanged Frog Figures 1, 6, 7 Rana macrodon blythi Boulenger, 1920 (partim). Rana acanthi Taylor, 1923; Taylor and Elbel, 1958. Rana macrodon acanthi (Inger, 1954). Rana macrodon macrocephala (Inger, 1954) (partim: three Mindoro specimens [USNM] provisionally referred by Inger’s ‘‘tentative identification’’ [Inger, 1954: 129]). Rana magna acanthi (Inger, 1958). Limnonectes (Limnonectes) acanthi Dubois, 1987 (partim). Limnonectes cf. acanthi Evans et al., 2003; Setiadi et al., 2010; Diesmos et al., 2015. Holotype.— PNM 9870 (adult male; formerly KU 303343; Field collector No. RMB 4957), Philippines, Mindoro Island, Oriental Mindoro Province, Municipality of Bongabong, Barangay Carmundo, Sitio Paypay-Ama, Paypay-Ama River, 12.7354 ° N, 121.4141 ° E, 100 m above sea level, WGS 84, R. M. Brown, A. C. Diesmos, C. D. Siler, and E. L. B. Rico, 13 March 2005. Paratypes (Paratopotypes).— KU 302084, 302087–88 (adult females), 302085–86, 302089 (adult males), 303343 (juvenile of undetermined sex), 303369–78 (10 subadults of undetermined sex), bearing the same data as the holotype. Other paratypes.— Mindoro Island, Oriental Mindoro Province, Municipality of Bongabong, Barangay Formon: KU 302090–91 (adult females), 302093, 302095, 302097, 302100 (3 adult males, 1 female), 302109–11 (3 adult males), C. D. Siler, 12 March 2005; Municipality of Victoria, Barangay Loyal: KU 302112–18 (2 adult males, 2 adult females, 3 juveniles of undetermined sex), C. D. Siler, 13 March 2005; Barangay Loyal, Sitio Panguisan, Panguisan River: KU 303470–78 (4 adult females, 5 subadults of undetermined sex), R. M. Brown, A. C. Diesmos, and C. D. Siler, 14 March 2005; Municipality of Gloria, Barangay Malamig: KU 302108 (adult female), 303344 (juvenile), J. B. Fernandez and R. M. Brown, 17 March 2006; KU 303346–54 (2 adult males, 2 females, 5 juveniles), R. M. Brown, C. D. Siler, and A. C. Diesmos, 13 March 2005; Sitio Balogbog, Cueba Simbahan: KU 303379–80 (2 subadults of undetermined sex), R. M. Brown, C. D. Siler, and E. L. B. Rico, 12 March 2005; Sitio Pastohan, Tanguisian Falls: KU 303381–402 (22 subadults of undetermined sex), A. C. Diesmos and E. L. B. Rico, 11 March 2005; Occidental Mindoro Province, Municipality of Calintaan, Barangay New Dagupan: KU 303266, 303345 (subadults), R. M. Brown, 8 March 2005; Municipality of Magsaysay, Barangay Nicolas, Sitio Banban: KU 303404–30 (1 adult female, 25 subadults and juveniles /metamorphs of undetermined sex), C. D. Siler and R. M. Brown, 9 March 2005; KU 304131–32 (adult male and subadult of undetermined sex), R. M. Brown; Municipality of Sablayan, Barangay Batong Buhay, Sitio Batulai, Mt. Siburan: KU 303430–52 (5 adult males, 6 adult females, 12 subadults of undetermined sex), E. L. B. Rico, 14 February 2006; KU 305450–51, 306637 (adult female, 2 subadult females), E. L. B. Rico, 19 February 2006; Barangay Malisbong, Sitio Aruyan: KU 335863–83 (11 females, 10 males), S. N. Travers, C. H. Oliveros, and R. M. Brown, 6 July 2013; Barangay Burgos, Sitio Posoy, Posoy River: KU 303453–69 (adult male, adult female, 15 juveniles of undetermined sex), R. M. Brown, 8 March 2005; Municipality of Paluan, Barangay Harrison, Sitio Ulasan, local name ‘‘ Matingaram’ ’: KU 308307, 308309, 308313–18, 308321–23, 308327, 308360, 308362–63, 308367–68, 308370–71, 308385, 308391, 308393, 308422, 308457, 308462, 308464–65, 308469, 308472 (15 adult females, 15 adult males), E. L. B. Rico, 4 January 2007; Municipality of Puerto Galera, Barangay San Isidro, Sitio Minolo, Ponderosa Golf Resort, adult female, J. A. McGuire and V. Yngente, 15 January 1996: TNHC 54920; Municipality of San Teodoro, Barangay Villaflor, Tamaraw Falls, approximately km 15 from Puerto Galera on Calapan-to-Puerto Galera road, 8 subadult males, 1 immature female, 3 adult males, J. A. McGuire and V. Yngente, 17 January 1996: TNHC 54921–29, 55023, 55025, 55029, 55033; same locality, 10 adult males, 11 adult females, R. I. Crombie and V. Yngente, 8 March 1995: USNM 556073–94; Municipality of Baco, Barangay Lantuyan, near Cabinuangang River: 6 adult males, R. I. Crombie and V. Yngente, 2 July 1991: USNM 508558–63; 5 adult males, 3 adult females, 1 immature specimen of undetermined sex, R. I. Crombie and V. Yngente, 7 March 1995: USNM 508564–72; Municipality of Tarogin, ca. 30 km S of Calapan Town, Mt. Halcon SE slope: CAS-SU 22146 (adult female), Q. Alcala and party, 1 April 1963; CAS-SU 22145 (adult female), same data, 31 March 1962; CAS-SU 22147–49 (adult male, 2 adult females), 1 April 1963; CAS-SU 22150 (adult female), S. Magusara and C. Batal, 14 April 1963; CAS-SU 22576 (adult male), Q. Alcala and party, 13 March 1963; CAS-SU 22577, 23508 (adult male and female), 31 March 1963; CAS-SU 23499, 23501, 23525 (adult females), 23505, 23514–15, 23519–20 (adult males), CAS-SU 23485, 23487, 23496–97, 23512–13, 23522 (subadult males), 23489, 23498, 23502 (subadult females), Q. Alcala and party, 10 March 1963; Municipality of Tarogin, Mt. Halcon: CAS-SU 22240 (juvenile), Q. Alcala and party, 14 April 1963; CAS-SU 22288– 22295, 23500, 23510–11, 23517–18, 23521 (juveniles), Q. Alcala and party, 1–20 April 1963; E side of Mt. Halcon, SE slope of Barawanan Peak, 830 m: CAS-SU 22151 (adult female), M. Pinero and party; Semirara Island, Oriental Mindoro Province, Municipality of Caluya, Barangay Tinogboc: KU 302105–07 (2 adult males, 1 adult female), C. D. Siler, 16 November 2000. Referred specimens.— Mindoro Island, Oriental Mindoro Province, Municipality of Baco, Mt. Baco, Alangsa River: USNM 508534–57; Occidental Mindoro Province, Municipality of Paluan, Barangay Harrison, Sitio Ulasan, local name ‘‘ Matingaram’ ’: KU 308308, 308310–12, 308319–20, 308324–26, 308361, 308364–66, 308369, 308372–76, 308386–90, 308392, 308394, 308416–21, 308423, 308430, 308451–52, 308456, 308461, 308463, 308467–68, 308470–87, 308500, 308528, 308538, 308561–69, 308586, 308589, 308590–92; Municipality of Paluan, Barangay 1, Sitio Ipol: KU 308593, 308597, 308599. Diagnosis and comparisons.— Limnonectes beloncioi is a medium-sized fanged frog, assigned to the genus Limnonectes (family Dicroglossidae), on the basis of its prominent, sexually dimorphic odontoid processes characteristic of the genus among other osteological synapomorphies (Inger, 1954, 1966; Emerson and Berrigan, 1993). The new species can be distinguished from all other known congeners based on a combination of its single-pulse/note advertisement call (vs. dual-pulses/note in L. acanthi from Palawan Island faunal region and PAIC landmasses), its phylogenetic position (sister to L. acanthi from Palawan PAIC; Evans et al., 2003; Setiadi et al., 2011), and its geographic distribution on Mindoro and Semirara Islands (vs. Palawan PAIC); it is the only species of Limnonectes known to occur on Mindoro and Semirara Islands and associated small satellite islands and, therefore, has no sympatric congeners. The new species is morphologically similar to its closest relative, L. acanthi; however, it may be distinguished from this allopatric congener by its male advertisement call. The note pulse substructure of L. beloncioi is singular (1 pulse per note vs. 2 pulses per note in L. acanthi), and the new species has a slower note repetition rate (11.5–12.8 notes per second vs. 14.2–15.4 notes per second), and has a higher dominant frequency (2,067.2–2,799.3 Hz vs. 1,335.1–1,679.6 Hz in L. acanthi). With the exception of the morphologically indistinguishable L. acanthi, the new species can be distinguished from all other Philippine species of Limnonectes (L. diuatus, L. ferneri, L. leytensis, L. macrocephalus, L. magnus, L. micrixalus, L. palavanensis, L. parvus, L. visayanus, and L. woodworthi) by a combination of body size, fang (odontoid) length, snout shape, relative lengths of the first and second finger, dorsal skin rugosity, restriction of white-tipped dermal asperities to the sacral region (not aggregated in radial clusters), the presence of irregular, elongate, discontinuous dorsolateral ridges (absence of a continuous dorsolateral fold), complete interdigital webbing of the foot, and the absence of a dark inverted ‘‘V’’-shaped mark on the dorsum. We provide morphological comparisons below, based on our data, in conjunction with or with consideration of the descriptions of Stejneger (1910), Taylor (1923), Inger (1954), Brown and Alcala (1977), and Siler et al. (2009). Limnonectes beloncioi differs from L. diuatus and L. ferneri by its rounded snout in lateral aspect (vs. posteroventrally sloping), its Finger I> Finger II relative finger lengths (vs. approximately equivalent length), and by restriction of white-tipped dorsal asperities to dorsal sacral region, and not distributed in radial clusters (vs. asperities not posteriorly restricted in L. diuatus; and densely distributed across entire of dorsum, and concentrated in radial sacral clusters in L. ferneri); and the presence of irregular dorsolateral ridges (absent in L. diuatus and L. ferneri); from L. leytensis, the new species can be distinguished by its larger adult body size (SVL range 54.2–83.1 vs. 25.8.2–34.0 mm in L. leytensis), rounded snout (vs. snout moderately pointed in lateral aspect in L. leytensis), Finger I> Finger II (equivalent length in L. leytensis), complete webbing (webbing incomplete/reduced in L. leytensis), and the absence of an inverted ‘‘V’’-shaped mark on the anterior dorsum (vs. present in L. leytensis); from a large male specimen of L. macrocephalus (the species endemic to the Luzon PAIC landmasses of Luzon, Polillo, Catanduanes, and Marinduque), the new species can be distinguished by the observation that it attains a considerably smaller maximal adult male body size (SVL range 54.2– 83.1 vs. 78.9–144.6 mm in L. macrocephalus); the new species also possesses a fully exposed tympanum (vs. dorsal and/or posterior edge of tympanum hidden beneath overlapping supratympanic dermal ridge skin in L. macrocephalus) and lacks sexual size dimorphism (Table 2), whereas L. macrocephalus exhibits reverse sexual size dimorphism (males larger); from a large adult male specimen of L. magnus, the new species can similarly be distinguished by its smaller adult body size (SVL range 54.2–83.1 vs. 66.3–164.4 mm in L. magnus), rugose middorsal skin texture (vs. smooth to shagreened in L. magnus), fully exposed tympanum (vs. dorsal and/or posterior edge of tympanum hidden beneath overlapping supratympanic dermal ridge skin in L. magnus), rounded snout (vs. pointed in L. magnus), and by the absence of reverse sexual size dimorphism (present in L. magnus); from L. parvus, L. micrixalus, and L. palavanensis, the new species can be distinguished by its larger adult body size (SVL 54.2–83.1 mm; vs. 24.2–35.5 in L. parvus; 28.1–30.2 in L. micrixalus; 30.0–37.6 in L. palavanensis), rugose (vs. smooth) dorsal skin, Finger I> Finger II (equivalent length), the presence (vs. absence) of white-tipped sacral asperities and the presence of irregular, discontinuous dorsolateral ridges (vs. asperities absent, dorsolateral folds continuous), complete webbing (vs. reduced), and by the absence (vs. presence) of an inverted ‘‘V’’-shaped middorsal marking (vs. present); the new species is additionally distinguished from these species by the absence of sexual size dimorphism (vs. females larger in L. parvus, L. micrixalus, and L. palavanensis); from L. visayanus, L. beloncioi is readily diagnosed by its rounded snout (vs. pointed in L. visayanus), a tendency towards longer adult male fangs (2.6–5.6 vs. 1.8–3.0 mm in L. visayanus), by the presence (vs. absence) of white-tipped sacral asperities; finally, the new species can be distinguished from L. woodworthi by its longer male fangs (2.6–5.6 vs. 1.3– 1.6 mm in L. woodworthi), moderately rugose middorsal skin (vs. smooth in L. woodworthi), rounded snout (vs. snout moderately pointed in L. woodworthi), the presence (vs. absence) of white-tipped sacral asperities, and by the absence (vs. presence) of continuous dorsolateral folds. Description of holotype.— A mature male, specimen in excellent condition; small portion of liver preserved separately for genetic material; habitus robust; head broader than body, its length 98.3% of its width, 39.0% of SVL; snout tip rounded in dorsal and lateral aspect (Fig. 6); supralabial region markedly swollen, increasingly protuberant towards angle of jaw; interorbital region and dorsal rostrum nearly flat; eye diameter 62.0% snout length, 97.5% eye–nares distance, 1.4 ⅹ eye–tympanum distance; pupil horizontally sub-elliptical with discontinuous posterior margin; canthus rostralis distinct, slightly medially bowed in dorsal aspect; loreal region concave; nostrils oriented dorsolaterally, narial openings visible in dorsal view; internarial region slightly convex; tympanum exposed, annulus slightly distinct, diameter 55.7% of eye diameter; supratympanic fold thick, strongly protuberant, moderately rugose, extending from posterior corner of eye, extending horizontally over (concealing) dorsoposterior corner and posterior margin of tympanic annulus, turning ventrally at nearly right angle, to end in supra-axillary region, where it is discontinuous with postrictal tubercular swelling at angle of jaw. Tongue elongate, tapered anteriorly, with narrow anterior attachment and laterally expanded, free, bilobed posterior margin at rest (anterior edge when tongue projected); choanae situated at anterolateral edge of palate, subcircular, their anterolateral edge partially concealed by palatal shelf of maxilla in ventral view; choanae widely separated by distance five or six times greater than diameter of single choana, each located just anterolaterally to (in contact with) lateral tip of dentigerous process of vomer; dentigerous process of vomer distinct, with four or five conical teeth on each side; dentigerous process angled anterolaterally (rostrally), approximately at 45 ° inclination, with closest (posterior) points separated by distance approximately equal to one choana, their most distant (anterior) ends separated by distance equal to three choanae; enlarged odontoid ‘‘fangs’’ large, recurved, unsheathed by oral mucosa for> distal half their length, situated on either side of mandibular symphysis/medial bulge, their tips sharply pointed, total length 4.6 mm (perpendicular distance from ventral edge of mandible), inclined dorsoposteriorly, tips 2.9 mm perpendicular from dorsal mandible surface; maxillary fang ‘‘sockets’’ anteromedial to choanae, large, round, similar in size to one choana; vocal apertures large, elongate, surrounded by extensive mucosal invaginations, situated at posteroventral margin of buccal floor, just medial to angle of jaw. Hand length 46.8% foot length; foot 93.4% tibia length; tibia length 58.3% SVL; fingers laterally, irregularly ovoid in cross section, due to presence of slight lateral dermal flange, extending from base of each digit, on either side, to proximal margins of terminal finger discs; terminal discs not expanded beyond widths of penultimate phalanges (Fig. 6), their relative descending lengths: III> I> II ¼ IV; subarticular tubercles prominent, their ventral surfaces convex and velvety in texture; one subarticular tubercle below Fingers I and II, two tubercles under Fingers III and IV; terminal discs and subarticular tubercles with gray, velvety, thickened surfaces; distal margins of tubercle more distinct than their proximal margins, and supernumerary tubercles absent, but articular surfaces of fingers between subarticular tubercles of digits, and at base of all digits covered medially with thickened tubercular surface; palmar surface with large, elongate, thenar tubercle (ventromedial surface of Finger I), enlarged, flattened, squarish medial ‘‘inner’’ (base of Finger III) metacarpal tubercle, and small, ovoid, convex outer (base of Finger IV) metacarpal tubercle; surface of these palmar structures, intervening, and surrounding surfaces all covered with similar, thickened, velvety (matte) tubercular dermis layer; nuptial excrescences or pads, asperities, and webbing absent; forearm musculature not hypertrophied. Tarsus folds and flaps absent; terminal discs of toes slightly expanded, with distinct circummarginal grooves; plantar surfaces of foot with well-developed, prominently protruding (ventrally), rounded subarticular tubercles (Fig. 6); plantar surfaces of foot smooth, with velvety-textured subarticular tubercles; relative lengths of toes: I Skin of dorsal surfaces of trunk and head smooth to slight shagreened texture, bearing low but clearly evident fleshy dermal tubercle clusters in supra- and post-tympanic regions, and tuberculate dorsolateral ridges, immediately following supratympanic region, and continuing posteriorly to the scapular region approximately to the points of forearm insertion (Fig. 6); similar dorsolateral tubercular ridges extend from this point, along more lateral (flanks) and slightly medial (dorsal) surfaces, and extend back to the sacral region; on posterior half of trunk and sacral region, dermal tubercles present mid-dorsally, consisting of single, raised tubercles or short, raised tubercular ridges; in sacral region, some tubercles capped with round, weakly keratinized dermal asperities; not arranged in clusters, or rows, but lightly dispersed in sacral region and upper one-third of dorsal surface of thigh and supra-cloacal region; ventral surfaces of head smooth; lateral and ventral surfaces of limbs smooth; remaining dorsal surfaces of limbs smooth to lightly shagreened, with occasional low tubercles; tarsus smooth on dorsolateral surface; cloacal region rugose (wrinkled), with smooth laterally and ventrally surrounding skin. Coloration of holotype in preservative.— Dominant dorsal color on head, body, and forelimbs uniform Dark Grayish-Brown (Köhler, 2012; color 284) Dark Grayish-Olive (275) with irregular, diffuse, Dusky Brown markings (285) concentrated on occiput and sacral regions; lateral head surfaces Drab (19) with diffuse Warm Sepia (40) m

Limnonectes beloncioi, new species urn:lsid:zoobank.org:act: 4CFCEF76-99D6-40DC-9B53- 58DECB1027CF Mindoro Fanged Frog Figures 1, 6, 7 Rana macrodon blythi Boulenger, 1920 (partim). Rana acanthi Taylor, 1923; Taylor and Elbel, 1958. Rana macrodon acanthi (Inger, 1954). Rana macrodon macrocephala (Inger, 1954) (partim: three Mindoro specimens [USNM] provisionally referred by Inger’s ‘‘tentative identification’’ [Inger, 1954: 129]). Rana magna acanthi (Inger, 1958). Limnonectes (Limnonectes) acanthi Dubois, 1987 (partim). Limnonectes cf. acanthi Evans et al., 2003; Setiadi et al., 2010; Diesmos et al., 2015. Holotype.— PNM 9870 (adult male; formerly KU 303343; Field collector No. RMB 4957), Philippines, Mindoro Island, Oriental Mindoro Province, Municipality of Bongabong, Barangay Carmundo, Sitio Paypay-Ama, Paypay-Ama River, 12.7354 ° N, 121.4141 ° E, 100 m above sea level, WGS 84, R. M. Brown, A. C. Diesmos, C. D. Siler, and E. L. B. Rico, 13 March 2005. Paratypes (Paratopotypes).— KU 302084, 302087–88 (adult females), 302085–86, 302089 (adult males), 303343 (juvenile of undetermined sex), 303369–78 (10 subadults of undetermined sex), bearing the same data as the holotype. Other paratypes.— Mindoro Island, Oriental Mindoro Province, Municipality of Bongabong, Barangay Formon: KU 302090–91 (adult females), 302093, 302095, 302097, 302100 (3 adult males, 1 female), 302109–11 (3 adult males), C. D. Siler, 12 March 2005; Municipality of Victoria, Barangay Loyal: KU 302112–18 (2 adult males, 2 adult females, 3 juveniles of undetermined sex), C. D. Siler, 13 March 2005; Barangay Loyal, Sitio Panguisan, Panguisan River: KU 303470–78 (4 adult females, 5 subadults of undetermined sex), R. M. Brown, A. C. Diesmos, and C. D. Siler, 14 March 2005; Municipality of Gloria, Barangay Malamig: KU 302108 (adult female), 303344 (juvenile), J. B. Fernandez and R. M. Brown, 17 March 2006; KU 303346–54 (2 adult males, 2 females, 5 juveniles), R. M. Brown, C. D. Siler, and A. C. Diesmos, 13 March 2005; Sitio Balogbog, Cueba Simbahan: KU 303379–80 (2 subadults of undetermined sex), R. M. Brown, C. D. Siler, and E. L. B. Rico, 12 March 2005; Sitio Pastohan, Tanguisian Falls: KU 303381–402 (22 subadults of undetermined sex), A. C. Diesmos and E. L. B. Rico, 11 March 2005; Occidental Mindoro Province, Municipality of Calintaan, Barangay New Dagupan: KU 303266, 303345 (subadults), R. M. Brown, 8 March 2005; Municipality of Magsaysay, Barangay Nicolas, Sitio Banban: KU 303404–30 (1 adult female, 25 subadults and juveniles /metamorphs of undetermined sex), C. D. Siler and R. M. Brown, 9 March 2005; KU 304131–32 (adult male and subadult of undetermined sex), R. M. Brown; Municipality of Sablayan, Barangay Batong Buhay, Sitio Batulai, Mt. Siburan: KU 303430–52 (5 adult males, 6 adult females, 12 subadults of undetermined sex), E. L. B. Rico, 14 February 2006; KU 305450–51, 306637 (adult female, 2 subadult females), E. L. B. Rico, 19 February 2006; Barangay Malisbong, Sitio Aruyan: KU 335863–83 (11 females, 10 males), S. N. Travers, C. H. Oliveros, and R. M. Brown, 6 July 2013; Barangay Burgos, Sitio Posoy, Posoy River: KU 303453–69 (adult male, adult female, 15 juveniles of undetermined sex), R. M. Brown, 8 March 2005; Municipality of Paluan, Barangay Harrison, Sitio Ulasan, local name ‘‘ Matingaram’ ’: KU 308307, 308309, 308313–18, 308321–23, 308327, 308360, 308362–63, 308367–68, 308370–71, 308385, 308391, 308393, 308422, 308457, 308462, 308464–65, 308469, 308472 (15 adult females, 15 adult males), E. L. B. Rico, 4 January 2007; Municipality of Puerto Galera, Barangay San Isidro, Sitio Minolo, Ponderosa Golf Resort, adult female, J. A. McGuire and V. Yngente, 15 January 1996: TNHC 54920; Municipality of San Teodoro, Barangay Villaflor, Tamaraw Falls, approximately km 15 from Puerto Galera on Calapan-to-Puerto Galera road, 8 subadult males, 1 immature female, 3 adult males, J. A. McGuire and V. Yngente, 17 January 1996: TNHC 54921–29, 55023, 55025, 55029, 55033; same locality, 10 adult males, 11 adult females, R. I. Crombie and V. Yngente, 8 March 1995: USNM 556073–94; Municipality of Baco, Barangay Lantuyan, near Cabinuangang River: 6 adult males, R. I. Crombie and V. Yngente, 2 July 1991: USNM 508558–63; 5 adult males, 3 adult females, 1 immature specimen of undetermined sex, R. I. Crombie and V. Yngente, 7 March 1995: USNM 508564–72; Municipality of Tarogin, ca. 30 km S of Calapan Town, Mt. Halcon SE slope: CAS-SU 22146 (adult female), Q. Alcala and party, 1 April 1963; CAS-SU 22145 (adult female), same data, 31 March 1962; CAS-SU 22147–49 (adult male, 2 adult females), 1 April 1963; CAS-SU 22150 (adult female), S. Magusara and C. Batal, 14 April 1963; CAS-SU 22576 (adult male), Q. Alcala and party, 13 March 1963; CAS-SU 22577, 23508 (adult male and female), 31 March 1963; CAS-SU 23499, 23501, 23525 (adult females), 23505, 23514–15, 23519–20 (adult males), CAS-SU 23485, 23487, 23496–97, 23512–13, 23522 (subadult males), 23489, 23498, 23502 (subadult females), Q. Alcala and party, 10 March 1963; Municipality of Tarogin, Mt. Halcon: CAS-SU 22240 (juvenile), Q. Alcala and party, 14 April 1963; CAS-SU 22288– 22295, 23500, 23510–11, 23517–18, 23521 (juveniles), Q. Alcala and party, 1–20 April 1963; E side of Mt. Halcon, SE slope of Barawanan Peak, 830 m: CAS-SU 22151 (adult female), M. Pinero and party; Semirara Island, Oriental Mindoro Province, Municipality of Caluya, Barangay Tinogboc: KU 302105–07 (2 adult males, 1 adult female), C. D. Siler, 16 November 2000. Referred specimens.— Mindoro Island, Oriental Mindoro Province, Municipality of Baco, Mt. Baco, Alangsa River: USNM 508534–57; Occidental Mindoro Province, Municipality of Paluan, Barangay Harrison, Sitio Ulasan, local name ‘‘ Matingaram’ ’: KU 308308, 308310–12, 308319–20, 308324–26, 308361, 308364–66, 308369, 308372–76, 308386–90, 308392, 308394, 308416–21, 308423, 308430, 308451–52, 308456, 308461, 308463, 308467–68, 308470–87, 308500, 308528, 308538, 308561–69, 308586, 308589, 308590–92; Municipality of Paluan, Barangay 1, Sitio Ipol: KU 308593, 308597, 308599. Diagnosis and comparisons.— Limnonectes beloncioi is a medium-sized fanged frog, assigned to the genus Limnonectes (family Dicroglossidae), on the basis of its prominent, sexually dimorphic odontoid processes characteristic of the genus among other osteological synapomorphies (Inger, 1954, 1966; Emerson and Berrigan, 1993). The new species can be distinguished from all other known congeners based on a combination of its single-pulse/note advertisement call (vs. dual-pulses/note in L. acanthi from Palawan Island faunal region and PAIC landmasses), its phylogenetic position (sister to L. acanthi from Palawan PAIC; Evans et al., 2003; Setiadi et al., 2011), and its geographic distribution on Mindoro and Semirara Islands (vs. Palawan PAIC); it is the only species of Limnonectes known to occur on Mindoro and Semirara Islands and associated small satellite islands and, therefore, has no sympatric congeners. The new species is morphologically similar to its closest relative, L. acanthi; however, it may be distinguished from this allopatric congener by its male advertisement call. The note pulse substructure of L. beloncioi is singular (1 pulse per note vs. 2 pulses per note in L. acanthi), and the new species has a slower note repetition rate (11.5–12.8 notes per second vs. 14.2–15.4 notes per second), and has a higher dominant frequency (2,067.2–2,799.3 Hz vs. 1,335.1–1,679.6 Hz in L. acanthi). With the exception of the morphologically indistinguishable L. acanthi, the new species can be distinguished from all other Philippine species of Limnonectes (L. diuatus, L. ferneri, L. leytensis, L. macrocephalus, L. magnus, L. micrixalus, L. palavanensis, L. parvus, L. visayanus, and L. woodworthi) by a combination of body size, fang (odontoid) length, snout shape, relative lengths of the first and second finger, dorsal skin rugosity, restriction of white-tipped dermal asperities to the sacral region (not aggregated in radial clusters), the presence of irregular, elongate, discontinuous dorsolateral ridges (absence of a continuous dorsolateral fold), complete interdigital webbing of the foot, and the absence of a dark inverted ‘‘V’’-shaped mark on the dorsum. We provide morphological comparisons below, based on our data, in conjunction with or with consideration of the descriptions of Stejneger (1910), Taylor (1923), Inger (1954), Brown and Alcala (1977), and Siler et al. (2009). Limnonectes beloncioi differs from L. diuatus and L. ferneri by its rounded snout in lateral aspect (vs. posteroventrally sloping), its Finger I> Finger II relative finger lengths (vs. approximately equivalent length), and by restriction of white-tipped dorsal asperities to dorsal sacral region, and not distributed in radial clusters (vs. asperities not posteriorly restricted in L. diuatus; and densely distributed across entire of dorsum, and concentrated in radial sacral clusters in L. ferneri); and the presence of irregular dorsolateral ridges (absent in L. diuatus and L. ferneri); from L. leytensis, the new species can be distinguished by its larger adult body size (SVL range 54.2–83.1 vs. 25.8.2–34.0 mm in L. leytensis), rounded snout (vs. snout moderately pointed in lateral aspect in L. leytensis), Finger I> Finger II (equivalent length in L. leytensis), complete webbing (webbing incomplete/reduced in L. leytensis), and the absence of an inverted ‘‘V’’-shaped mark on the anterior dorsum (vs. present in L. leytensis); from a large male specimen of L. macrocephalus (the species endemic to the Luzon PAIC landmasses of Luzon, Polillo, Catanduanes, and Marinduque), the new species can be distinguished by the observation that it attains a considerably smaller maximal adult male body size (SVL range 54.2– 83.1 vs. 78.9–144.6 mm in L. macrocephalus); the new species also possesses a fully exposed tympanum (vs. dorsal and/or posterior edge of tympanum hidden beneath overlapping supratympanic dermal ridge skin in L. macrocephalus) and lacks sexual size dimorphism (Table 2), whereas L. macrocephalus exhibits reverse sexual size dimorphism (males larger); from a large adult male specimen of L. magnus, the new species can similarly be distinguished by its smaller adult body size (SVL range 54.2–83.1 vs. 66.3–164.4 mm in L. magnus), rugose middorsal skin texture (vs. smooth to shagreened in L. magnus), fully exposed tympanum (vs. dorsal and/or posterior edge of tympanum hidden beneath overlapping supratympanic dermal ridge skin in L. magnus), rounded snout (vs. pointed in L. magnus), and by the absence of reverse sexual size dimorphism (present in L. magnus); from L. parvus, L. micrixalus, and L. palavanensis, the new species can be distinguished by its larger adult body size (SVL 54.2–83.1 mm; vs. 24.2–35.5 in L. parvus; 28.1–30.2 in L. micrixalus; 30.0–37.6 in L. palavanensis), rugose (vs. smooth) dorsal skin, Finger I> Finger II (equivalent length), the presence (vs. absence) of white-tipped sacral asperities and the presence of irregular, discontinuous dorsolateral ridges (vs. asperities absent, dorsolateral folds continuous), complete webbing (vs. reduced), and by the absence (vs. presence) of an inverted ‘‘V’’-shaped middorsal marking (vs. present); the new species is additionally distinguished from these species by the absence of sexual size dimorphism (vs. females larger in L. parvus, L. micrixalus, and L. palavanensis); from L. visayanus, L. beloncioi is readily diagnosed by its rounded snout (vs. pointed in L. visayanus), a tendency towards longer adult male fangs (2.6–5.6 vs. 1.8–3.0 mm in L. visayanus), by the presence (vs. absence) of white-tipped sacral asperities; finally, the new species can be distinguished from L. woodworthi by its longer male fangs (2.6–5.6 vs. 1.3– 1.6 mm in L. woodworthi), moderately rugose middorsal skin (vs. smooth in L. woodworthi), rounded snout (vs. snout moderately pointed in L. woodworthi), the presence (vs. absence) of white-tipped sacral asperities, and by the absence (vs. presence) of continuous dorsolateral folds. Description of holotype.— A mature male, specimen in excellent condition; small portion of liver preserved separately for genetic material; habitus robust; head broader than body, its length 98.3% of its width, 39.0% of SVL; snout tip rounded in dorsal and lateral aspect (Fig. 6); supralabial region markedly swollen, increasingly protuberant towards angle of jaw; interorbital region and dorsal rostrum nearly flat; eye diameter 62.0% snout length, 97.5% eye–nares distance, 1.4 ⅹ eye–tympanum distance; pupil horizontally sub-elliptical with discontinuous posterior margin; canthus rostralis distinct, slightly medially bowed in dorsal aspect; loreal region concave; nostrils oriented dorsolaterally, narial openings visible in dorsal view; internarial region slightly convex; tympanum exposed, annulus slightly distinct, diameter 55.7% of eye diameter; supratympanic fold thick, strongly protuberant, moderately rugose, extending from posterior corner of eye, extending horizontally over (concealing) dorsoposterior corner and posterior margin of tympanic annulus, turning ventrally at nearly right angle, to end in supra-axillary region, where it is discontinuous with postrictal tubercular swelling at angle of jaw. Tongue elongate, tapered anteriorly, with narrow anterior attachment and laterally expanded, free, bilobed posterior margin at rest (anterior edge when tongue projected); choanae situated at anterolateral edge of palate, subcircular, their anterolateral edge partially concealed by palatal shelf of maxilla in ventral view; choanae widely separated by distance five or six times greater than diameter of single choana, each located just anterolaterally to (in contact with) lateral tip of dentigerous process of vomer; dentigerous process of vomer distinct, with four or five conical teeth on each side; dentigerous process angled anterolaterally (rostrally), approximately at 45 ° inclination, with closest (posterior) points separated by distance approximately equal to one choana, their most distant (anterior) ends separated by distance equal to three choanae; enlarged odontoid ‘‘fangs’’ large, recurved, unsheathed by oral mucosa for> distal half their length, situated on either side of mandibular symphysis/medial bulge, their tips sharply pointed, total length 4.6 mm (perpendicular distance from ventral edge of mandible), inclined dorsoposteriorly, tips 2.9 mm perpendicular from dorsal mandible surface; maxillary fang ‘‘sockets’’ anteromedial to choanae, large, round, similar in size to one choana; vocal apertures large, elongate, surrounded by extensive mucosal invaginations, situated at posteroventral margin of buccal floor, just medial to angle of jaw. Hand length 46.8% foot length; foot 93.4% tibia length; tibia length 58.3% SVL; fingers laterally, irregularly ovoid in cross section, due to presence of slight lateral dermal flange, extending from base of each digit, on either side, to proximal margins of terminal finger discs; terminal discs not expanded beyond widths of penultimate phalanges (Fig. 6), their relative descending lengths: III> I> II ¼ IV; subarticular tubercles prominent, their ventral surfaces convex and velvety in texture; one subarticular tubercle below Fingers I and II, two tubercles under Fingers III and IV; terminal discs and subarticular tubercles with gray, velvety, thickened surfaces; distal margins of tubercle more distinct than their proximal margins, and supernumerary tubercles absent, but articular surfaces of fingers between subarticular tubercles of digits, and at base of all digits covered medially with thickened tubercular surface; palmar surface with large, elongate, thenar tubercle (ventromedial surface of Finger I), enlarged, flattened, squarish medial ‘‘inner’’ (base of Finger III) metacarpal tubercle, and small, ovoid, convex outer (base of Finger IV) metacarpal tubercle; surface of these palmar structures, intervening, and surrounding surfaces all covered with similar, thickened, velvety (matte) tubercular dermis layer; nuptial excrescences or pads, asperities, and webbing absent; forearm musculature not hypertrophied. Tarsus folds and flaps absent; terminal discs of toes slightly expanded, with distinct circummarginal grooves; plantar surfaces of foot with well-developed, prominently protruding (ventrally), rounded subarticular tubercles (Fig. 6); plantar surfaces of foot smooth, with velvety-textured subarticular tubercles; relative lengths of toes: I Coloration of holotype in preservative.— Dominant dorsal color on head, body, and forelimbs uniform Dark Grayish-Brown (Köhler, 2012; color 284) Dark Grayish-Olive (275) with irregular, diffuse, Dusky Brown markings (285) concentrated on occiput and sacral regions; lateral head surfaces Drab (19) with diffuse Warm Sepia (40) m, Published as part of Herr, Mark W., Goyes Vallejos, Johana, Meneses, Camila G., Abraham, Robin K., Otterholt, Rayanna, Siler, Cameron D., Rico, Edmund Leo B. & Brown, Rafe M., 2021, A New, Morphologically Cryptic Species of Fanged Frog, Genus Limnonectes (Amphibia: Anura: Dicroglossidae), from Mindoro Island, Central Philippines, pp. 188-210 in Ichthyology & Herpetology 109 (1) on pages 197-203, DOI: 10.1643/h2020095, http://zenodo.org/record/7846717, {"references":["Taylor, E. H. 1923. Additions to the herpetological fauna of the Philippine Islands, III. Philippine Journal of Science 22: 515 - 555.","Inger, R. F. 1954. Systematics and zoogeography of Philippine Amphibia. Fieldiana Zoology 33: 182 - 531.","Evans, B. J., R. M. Brown, J. A. McGuire, J. Supriatna, N. Andayani, A. C. Diesmos, D. J. Melnick, and D. C. Cannatella. 2003. Phylogenetics of fanged frogs: testing biogeographical hypotheses at the interface of the Asian and Australian faunal zones. Systematic Biology 52: 794 - 819.","Diesmos, A. C., J. L. Watters, N. A. Huron, D. R. Davis, A. C. Alcala, R. I. Crombie, L. E. Afuang, G. Gee-Das, R. V. Sison, M. B. Sanguila, M. L. Penrod, M. J. Labonte, C. S. Davey, E. A. Leone... C. D. Siler. 2015. Amphibians of the Philippines, part I: checklist of the species. Proceedings of the California Academy of Sciences 62: 457 - 539.","Inger, R. F. 1966. The systematics and zoogeography of the Amphibia of Borneo. Fieldiana 52: 1 - 402.","Emerson, S. B., and D. Berrigan. 1993. Systematics of Southeast Asian ranids: multiple origins of voicelessness in the subgenus Limnonectes (Fitzinger). Herpetologica 49: 22 - 31.","Setiadi, M. I., J. A. McGuire, R. M. Brown, M. Zubairi, D. T. Iskandar, N. Andayani, J. Supriatna, and B. J. Evans. 2011. Adaptive radiation and ecological opportunity in Sulawesi and Philippine fanged frog (Limnonectes) communities. American Naturalist 178: 221 - 240.","Stejneger, L. 1910. Description of a new frog from the Philippine Islands. Smithsonian Miscellaneous Collections 52: 437 - 439.","Brown, W. C., and A. C. Alcala. 1977. A new frog of the genus Rana from the Philippines. Proceedings of the Biological Society of Washington 90: 669 - 675.","Siler, C. D., J. McVay, A. C. Diesmos, and R. M. Brown. 2009. A new species of fanged frog (Dicroglossidae; genus Limnonectes) from southern Mindanao Island, Philippines. Herpetologica 65: 105 - 114.","Kohler, G. 2012. Color Catalog for Field Biologists. Herpeton, Offenbach, Germany.","Taylor, E. H. 1928. Amphibians, lizards, and snakes of the Philippines, p. 214 - 241. In: Distribution of Life in the Philippines. R. Dickerson (ed.). Philippine Bureau of Science, Monograph 21, Manila, Philippines.","Brown, W. C., and A. C. Alcala. 1970. The zoogeography of the Philippine Islands, a fringing archipelago. Proceedings of the California Academy of Sciences 38: 105 - 130.","Brown, R. M., and S. I. Guttman. 2002. Phylogenetic systematics of the Rana signata complex of Philippine and Bornean stream frogs: reconsideration of Huxley's modification of Wallace's Line at the Oriental-Australian faunal zone interface. Biological Journal of the Linnean Society 76: 393 - 461.","Lizuka, H., H. Tamura, T. Pokavanich, M. C. D. Rubio- Paringit, K. Nadaoka, and M. D. Fortes. 2009. Highly skewed tidal circulation pattern and water quality in Puerto Galera Bay, Mindoro Island, Philippines. Coastal Engineering Journal 51: 341 - 361.","Wells, K. D. 2007. The Ecology and Behavior of Amphibians. University of Chicago Press, Chicago.","Lopez, V. B. 1976. The Mangyans of Mindoro: An Ethnohistory. University of the Philippines Press, Quezon City, Philippines.","Kikuchi, Y. 1984. Mindoro highlanders: the life of swidden agriculturalists. New Day Publishers, Quezon City, Philippines.","Schult, V. 1991. Mindoro: A Social History of a Philippine Island in the 20 th Century. Divine World Publications, Manila, Philippines.","Scheffers, B., R. Corlett, A. C. Diesmos, and W. Laurance. 2012. Local demand drives a bushmeat industry in a Philippine forest preserve. Tropical Conservation Science 5: 133 - 141.","IUCN (International Union for the Conservation of Nature). 2020. The IUCN Red List of Threatened Species. Version 2020 - 2. https: // www. iucnredlist. org. Downloaded on 09 July 2020.","Gonzalez, J. C. T., C. A. A. Layusa, L. E. Afuang, M. R. M. Duya, L. R. Heaney, D. S. Balete, D. G. E. Tabaranza, C. P. Espanola, W. A. C. Van den Ven, A. C. Diesmos, R. M. Causaren, M. L. L. Diesmos, R. T. Lagat, E. Y. Sy... P. Ong. 2018. Review and update of the 2004 National List of Threatened Terrestrial Fauna of the Philippines. Sylvatrop, The Technical Journal of Philippine Ecosystems and Natural Resources 28: 73 - 144."]}

Herr, Mark W., Goyes Vallejos, Johana, Meneses, Camila G., Abraham, Robin K., Otterholt, Rayanna, Siler, Cameron D., Rico, Edmund Leo B., Brown, Rafe M. (2021): A New, Morphologically Cryptic Species of Fanged Frog, Genus Limnonectes (Amphibia: Anura: Dicroglossidae), from Mindoro Island, Central Philippines. Ichthyology & Herpetology 109 (1): 188-210, DOI: 10.1643/h2020095, URL: http://dx.doi.org/10.1643/h2020095

Chan, Kin Onn, Abraham, Robin K., Grismer, L. Lee, Brown, Rafe M. (2020): A systematic review of the Pulchrana picturata complex, with the description of a new species from Peninsular Malaysia, Sumatra, and southern Thailand. Raffles Bulletin of Zoology 68: 880-890, DOI: 10.26107/RBZ-2020-0096

Liopeltis tiomanica sp. nov. Liopeltis tricolor (part): J.L. Grismer et al. 2004: 275; Grismer 2011: 203. Liopeltis tricolour [sic] (part): Grismer et al. 2006: 178. Holotype. LSUHC 5037, adult female (Figs. 2���4), Peninsular Malaysia, Pahang State, Pulau Tioman (= Tioman Island), Gunung Kajang trail, 2.77900��N 104.15703��E (Datum WGS84), 618 m elev., coll. 11 August 2002 by Jesse L. Grismer, Perry L. Wood, Jr., and L. Lee Grismer. Etymology. The specific epithet refers to the new species��� type and only known locality on the island of Pulau Tioman. The specific epithet is feminine, in agreement with the gender of the genus (Poyarkov et al. 2019). Diagnosis. A species of Liopeltis having the combination of the nasal scale fused with the internasal scale; preoculars two; ventrals 161; distinct black lateral cephalic stripe extending from rostral through eye to approximately 30 mm behind neck before fading; four longitudinal stripes on dorsum; and venter immaculate, without stripes. Description of holotype (Figs. 2���3). Adult female; head slightly distinct from neck, triangular in dorsal view; snout weakly pointed in lateral view, projecting beyond lower jaw; eye large, pupil round; body slender; tail thin. Rostral wide, clearly visible from above; nasal partially divided, portion anterior to nostril continuous with internasal, portion posterior to nostril with horizontal suture; loreal absent; two preoculars, upper rectangular, approximately 25% size of lower, trapezoidal preocular; two postoculars, squarish or rounded, upper scale slightly larger than lower scale; eight supralabials, 4 th and 5 th in contact with eye, 2 nd��� 4 th in contact with lower preocular, 5 th (right side) and 5 th��� 6 th (left side) in contact with lower postocular, 7 th largest; eight infralabials, 5 th largest; one elongate anterior temporal; two subequal posterior temporals, shorter than anterior temporal; two internasals; two prefrontals; single frontal, approximately 63% length of parietals, with one supraocular on each side; two large, elongate parietals; anterior pair and posterior pairs of chin shields subequal in length, anterior pair broader; dorsal scales smooth, in rows 15:15:13, apical pits absent; 161 ventrals; cloacal plate divided; 119 paired subcaudals. Scale counts are summarized in Table 2. *Data from Taylor (1922: 164, ���No. 940, Bureau of Science Collection;��� specimen destroyed during World War II). Color of holotype in life (Fig. 2). Dorsal surface of head bronze-brown (Grayish Horn Color 268); chin, supralabials, and infralabials cream (Cream White 52); black stripe on lateral side of head and neck separates dorsal and ventral coloration, stripe slightly wider than one dorsal scale, extending from rostral though eye to approximately 30 mm posterior to neck, gradually fading; dorsum bronze (Grayish Horn Color 268) with four longitudinal brown (Olive-Brown 278) stripes, two dorsal stripes extending from neck to tail, two lateral stripes extending from area posterior of black head stripe to level of cloacal plate; venter uniform pale cream color (Cream White 52) with small dark spots on ventral scales. Color of holotype in preservative (Fig. 3). Dorsum faded to grayish green (Olive Horn Color 16); dorsal stripes faded to light brown (Clay Color 18). Molecular data. The standard deviation of split frequencies among the four Bayesian runs was 0.003717 and the Estimated Sample Sizes (ESS) of parameters were ��� 3,390. The holotype of Liopeltis tiomanica sp. nov. was recovered with strong support (Bayesian posterior probability 1.00) to be the sister taxon of L. philippina (Fig. 4). The L. tiomanica sp. nov. + L. philippina clade was recovered with strong support (Bayesian posterior probability 1.00) to be nested within a clade containing L. tricolor and L. stoliczkae, although the relationships among L. tricolor, L. stoliczkae, and the L. tiomanica sp. nov. + L. philippina clade were not resolved (Fig. 4). Liopeltis tiomanica sp. nov. had uncorrected p -distances in cyt b of 9.6% from L. philippina and 12.9���13.3% from L. tricolor. Distribution, natural history, and conservation. Liopeltis tiomanica sp. nov. is known only from the holotype specimen collected from Pulau Tioman (Fig. 1). The specimen was found in the afternoon while perched on a small tree branch 2 m above the ground in primary hill dipterocarp forest (J. Grismer et al. 2004, Fig. 5). Comparisons. Liopeltis tiomanica sp. nov. differs from all other species of Liopeltis, except its sister taxon L. philippina (Fig. 4), by having the nasal fused with the internasal and four longitudinal stripes on the dorsum (Weinell et al. 2019). Liopeltis tiomanica sp. nov. differs from L. philippina by having 161 ventral scales (vs. 139���150 in L. philippina); two preocular scales (vs. one in L. philippina); and a distinct black lateral cephalic stripe (vs. indistinct in L. philippina; Fig. 5). Liopeltis tiomanica sp. nov. further differs from the sympatric (and possibly syntopic) L. tricolor by having the prefrontal in contact only with the second supralabial (vs. second and third supralabials in L. tricolor); and immaculate ventral scales (vs. three gray ventral stripes in most L. tricolor). Scalation differences among these three species are summarized in Tables 2 and 3., Published as part of Som, Hannah E., Grismer, L. Lee, Grismer, Jesse L., Wood, Perry L., Quah, Evan S. H., Brown, Rafe M., Diesmos, Arvin C., Weinell, Jeffrey L. & Stuart, Bryan L., 2020, A new Liopeltis Fitzinger, 1843 (Squamata: Colubridae) from Pulau Tioman, Peninsular Malaysia, pp. 472-484 in Zootaxa 4766 (3) on pages 475-477, DOI: 10.11646/zootaxa.4766.3.6, http://zenodo.org/record/3765613, {"references":["Grismer, J. L., Grismer, L. L., Das, I., Yaakob, N. S., Liat, L. B., Leong, T. M., Youmans, T. M. & Kaiser, H. (2004) Species diversity and checklist of the herpetofauna of Pulau Tioman, Peninsular Malaysia, with a preliminary overview of habitat utilization. Asiatic Herpetological Research, 10, 247 - 279.","Grismer, L. L. (2011) Field Guide to the Amphibians and Reptiles of the Seribuat Archipelago, Peninsular Malaysia. Edition Chimaira, Frankfurt, 258 pp.","Grismer, L. L., Youmans, T. M., Wood, Jr. P. L. & Grismer, J. L. (2006) Checklist of the herpetofauna of the Seribuat archipelago, West Malaysia with comments on biogeography, natural history, and adaptive types. Raffles Bulletin of Zoology, 54 (1), 157 - 180.","Poyarkov, N. A., Nguyen, T. V. & Vogel, G. (2019) A new species of the genus Liopeltis Fitzinger 1843 from Vietnam (Squamata: Colubridae). Journal of Natural History, 53, 1647 - 1672. https: // doi. org / 10.1080 / 00222933.2019.1656784","Taylor, E. H. (1922) The Snakes of the Philippine Islands. Bureau of Printing, Manila, Philippines, 312 pp. https: // doi. org / 10.5962 / bhl. title. 55346","Weinell, J. L., Hooper, E., Leviton, A. E. & Brown, R. M. (2019) Illustrated key to the snakes of the Philippines. Proceedings of the California Academy of Sciences, 66 (1), 1 - 49."]}

Som, Hannah E., Grismer, L. Lee, Grismer, Jesse L., Wood, Perry L., Quah, Evan S. H., Brown, Rafe M., Diesmos, Arvin C., Weinell, Jeffrey L., Stuart, Bryan L. (2020): A new Liopeltis Fitzinger, 1843 (Squamata: Colubridae) from Pulau Tioman, Peninsular Malaysia. Zootaxa 4766 (3): 472-484, DOI: 10.11646/zootaxa.4766.3.6

Liopeltis tiomanica sp. nov. Liopeltis tricolor (part): J.L. Grismer et al. 2004: 275; Grismer 2011: 203. Liopeltis tricolour [sic] (part): Grismer et al. 2006: 178. Holotype. LSUHC 5037, adult female (Figs. 2–4), Peninsular Malaysia, Pahang State, Pulau Tioman (= Tioman Island), Gunung Kajang trail, 2.77900°N 104.15703°E (Datum WGS84), 618 m elev., coll. 11 August 2002 by Jesse L. Grismer, Perry L. Wood, Jr., and L. Lee Grismer. Etymology. The specific epithet refers to the new species’ type and only known locality on the island of Pulau Tioman. The specific epithet is feminine, in agreement with the gender of the genus (Poyarkov et al. 2019). Diagnosis. A species of Liopeltis having the combination of the nasal scale fused with the internasal scale; preoculars two; ventrals 161; distinct black lateral cephalic stripe extending from rostral through eye to approximately 30 mm behind neck before fading; four longitudinal stripes on dorsum; and venter immaculate, without stripes. Description of holotype (Figs. 2–3). Adult female; head slightly distinct from neck, triangular in dorsal view; snout weakly pointed in lateral view, projecting beyond lower jaw; eye large, pupil round; body slender; tail thin. Rostral wide, clearly visible from above; nasal partially divided, portion anterior to nostril continuous with internasal, portion posterior to nostril with horizontal suture; loreal absent; two preoculars, upper rectangular, approximately 25% size of lower, trapezoidal preocular; two postoculars, squarish or rounded, upper scale slightly larger than lower scale; eight supralabials, 4 th and 5 th in contact with eye, 2 nd– 4 th in contact with lower preocular, 5 th (right side) and 5 th– 6 th (left side) in contact with lower postocular, 7 th largest; eight infralabials, 5 th largest; one elongate anterior temporal; two subequal posterior temporals, shorter than anterior temporal; two internasals; two prefrontals; single frontal, approximately 63% length of parietals, with one supraocular on each side; two large, elongate parietals; anterior pair and posterior pairs of chin shields subequal in length, anterior pair broader; dorsal scales smooth, in rows 15:15:13, apical pits absent; 161 ventrals; cloacal plate divided; 119 paired subcaudals. Scale counts are summarized in Table 2. *Data from Taylor (1922: 164, “No. 940, Bureau of Science Collection;” specimen destroyed during World War II). Color of holotype in life (Fig. 2). Dorsal surface of head bronze-brown (Grayish Horn Color 268); chin, supralabials, and infralabials cream (Cream White 52); black stripe on lateral side of head and neck separates dorsal and ventral coloration, stripe slightly wider than one dorsal scale, extending from rostral though eye to approximately 30 mm posterior to neck, gradually fading; dorsum bronze (Grayish Horn Color 268) with four longitudinal brown (Olive-Brown 278) stripes, two dorsal stripes extending from neck to tail, two lateral stripes extending from area posterior of black head stripe to level of cloacal plate; venter uniform pale cream color (Cream White 52) with small dark spots on ventral scales. Color of holotype in preservative (Fig. 3). Dorsum faded to grayish green (Olive Horn Color 16); dorsal stripes faded to light brown (Clay Color 18). Molecular data. The standard deviation of split frequencies among the four Bayesian runs was 0.003717 and the Estimated Sample Sizes (ESS) of parameters were ≥ 3,390. The holotype of Liopeltis tiomanica sp. nov. was recovered with strong support (Bayesian posterior probability 1.00) to be the sister taxon of L. philippina (Fig. 4). The L. tiomanica sp. nov. + L. philippina clade was recovered with strong support (Bayesian posterior probability 1.00) to be nested within a clade containing L. tricolor and L. stoliczkae, although the relationships among L. tricolor, L. stoliczkae, and the L. tiomanica sp. nov. + L. philippina clade were not resolved (Fig. 4). Liopeltis tiomanica sp. nov. had uncorrected p -distances in cyt b of 9.6% from L. philippina and 12.9–13.3% from L. tricolor. Distribution, natural history, and conservation. Liopeltis tiomanica sp. nov. is known only from the holotype specimen collected from Pulau Tioman (Fig. 1). The specimen was found in the afternoon while perched on a small tree branch 2 m above the ground in primary hill dipterocarp forest (J. Grismer et al. 2004, Fig. 5). Comparisons. Liopeltis tiomanica sp. nov. differs from all other species of Liopeltis, except its sister taxon L. philippina (Fig. 4), by having the nasal fused with the internasal and four longitudinal stripes on the dorsum (Weinell et al. 2019). Liopeltis tiomanica sp. nov. differs from L. philippina by having 161 ventral scales (vs. 139–150 in L. philippina); two preocular scales (vs. one in L. philippina); and a distinct black lateral cephalic stripe (vs. indistinct in L. philippina; Fig. 5). Liopeltis tiomanica sp. nov. further differs from the sympatric (and possibly syntopic) L. tricolor by having the prefrontal in contact only with the second supralabial (vs. second and third supralabials in L. tricolor); and immaculate ventral scales (vs. three gray ventral stripes in most L. tricolor). Scalation differences among these three species are summarized in Tables 2 and 3.

Ptychozoon popaense sp. nov. Mt. Popa Parachute Gecko Fig. 2 Holotype. LSUHC 13508, adult male, collected by Myint Kyaw Thura on 27 May 2017 from Yay Su camp, 16 km northeast of Kyauk-pa-taung Town, Kyauk-pa-taung Township, Mt. Popa, Mandalay Region, Myanmar (20.88331°N, 95.26638°E, 629 m above sea level). Paratopotype. Adult female LSUHC 13507 bears the same collection data as the holotype. Diagnosis. Ptychozoon popaense s p. nov. can be differentiated from all other species of Ptychozoon based on having the unique combination of separated supranasals; presence of an infra-auricular flap; dorsal and caudal tubercles absent; 32–36 enlarged ventral scales; enlarged femorals absent; 21 enlarged, pore-bearing precloacals in the male and 12 dimpled precloacal scales in the female; 13 or 14 transversely enlarged subdigital lamellae under the fourth toe; 28 or 29 scales across the caudal flap; distal caudal lobes not fused into a long caudal flap; bases of 15–19 lobes fused anterior to the caudal flap; caudal lobes angling slightly posteriorly; thick, dark, postorbital stripe present; four dark, dorsal, body bands between limb insertions; irregularly shaped, with white, vertebral markings; and adults not having distinct immaculate black and white caudal bands dorsally (Table 4). Description of holotype. Adult male SVL 86.2 mm; head short (HL/SVL 0.26), wide (HW/SVL 0.20), depressed (HD/HL 0.43), distinct from neck; snout rounded at tip in dorsal profile; interorbital region flat; lores rounded; rostral scale large, rectangular, without dorsomedial groove, in contact posteriorly with two supranasals, one postnasal, and laterally with nostrils and first supralabials; supralabials (9R,10L), supralabial eight in midorbital position; infralabials (9R,10L); nostrils elliptical with long axis oriented dorsoventrally, occupying anterior portion of nasal scale, bordered anteriorly by rostral, dorsally by supranasal, posteriorly by four postnasals of varying sizes, and ventrally by first supralabial; scales on rostrum granular slightly larger than granular scales on top of head and occiput; no ridge of tubercles along mandibles; eyes large (ED/HL 0.24), less than snout length and EAO distance; pupil vertically elliptical, crenelated; supraciliaries elongate, posteriormost spinose; auricular opening rounded, lacking enlarged lobes; tympanum deeply sunk; infra-auricular flap broad, rounded, extending from below corner of mouth to base of neck, measuring 3.5 mm at its widest point; dorsal scales of infra-auricular flap large, subimbricate proximally, small juxtaposed distally, minute and granular ventrally; infra-auricular flap on right side deeply notched; mental triangular, wider than deep, bordered laterally by first infralabials and posteriorly by paired, rectangular postmentals contacting medially for 70% of their length; one row of enlarged sublabials bordering infralabials, anteriormost largest; gular scales small, rounded, grading abruptly into larger imbricating throat and ventral scales. Body dorsoventrally depressed, relatively stout (AG/SVL 0.47); axilla-groin cutaneous expansion (flap) 5.5 mm at midpoint of body and bearing enlarged, juxtaposed, rectangular scales dorsally and minute, juxtaposed, subrectangular scales ventrally; dorsal body scales minute, flat, rounded, juxtaposed, largest mid-dorsally; 36 transverse rows of large, smooth, flat, subimbricate ventrals, ventrals much larger than dorsals, decreasing in size laterally; 21 elevated, enlarged, pore-bearing, precloacal scales; seven rows of enlarged, post-precloacal scales; scales immediately anterior to vent granular. Limbs short, robust (FL/SVL 0.11; TBL/SVL 0.16); dorsal scales of forelimbs, flat, juxtaposed, slightly larger than dorsal body scales and juxtaposed ventral forelimb scales; anterior and posterior margins of forelimbs, posterior margins of hind limbs, and anterior margins of forelegs bearing wide, cutaneous flaps extending to bases of digits I and V, bearing subimbricate, moderately larger scales dorsally and smaller imbricate scales ventrally wide; wide predigital notch in preantebrachial flap; palmar scales smooth, rounded; digits fully webbed, relatively short, dorsoventrally compressed; undivided transverse subdigital lamellae number 12 (I), 12 (II), 15 (III), 15 (IV), 12 (V), distalmost lamellae V-shaped; claws arise from within the dorsal surface of digital pads; first digit lacks a claw; dorsal scales of hind limbs, flat, juxtaposed, smaller than dorsal body scales and flat, subimbricate scales of thigh; posterior margins of thigh and foreleg, and anterior margin of foreleg have wide, cutaneous flaps bearing subimbricate, moderately large scales dorsally and much smaller subimbricate scales ventrally; pretibial flap not contacting base of digit V; post-tibial flap contacts base of digit I; plantar scales smooth, subimbricate; digits fully webbed; transverse subdigital lamellae number 11 (I), 13 (II), 15 (III), 13 (IV), 14 (V), distalmost lamellae Vshaped; claws arise from within the dorsal surface of digital pads; and first digit lacks a claw. Tail original, flattened, moderate in length (TaL/SVL 1.04); two median rows of transversely widened, smooth subcaudals; postcloacal scales on heimpenial swelling large, flat, imbricate; dorsal caudals flat, juxtaposed, larger than dorsal body scales, occurring in whorls; 6–8 smaller scales between larger scales delimiting whorls; tail width and caudal lobes slightly decrease posteriorly; 25 caudal lobes on each side slightly angled posteriorly; 19 lobes fused at their bases, prior to grading posteriorly into short, straight-edged, terminal tail flap terminal tail flap short (8.6 mm), not wider than tail. Coloration in life (Fig. 2). Dorsal ground color of head, body and limbs light-green, that of tail light-brown; top of head bearing three elongate, grey markings extending onto nape; wide, grey preorbital stripe extends through eye terminating on nape, becoming progressively darker and thicker posteriorly; labials much lighter than ground color of head, bearing thin dark lines; four wide, sinuous, grey bands between limb insertions bearing diffuse anterior margins and black, well-defined posterior margins and extending onto flanks; a series of irregularly shaped, poorly defined, immaculate, white vertebral blotches extending from nape to sacrum; dorsal surface of limbs bearing grey, diffuse, irregularly shaped, banded to reticulate patterns; seven dark caudal bands becoming darker posteriorly separated by six lighter bands, posterior two lightest; terminal flap nearly black; ventral surfaces of head, body, and limbs beige, generally immaculate; subcaudal region distinctly banded. All colors have faded to varying shades of grey in preservative. Variation. The female paratopotype is nearly identical to the holotype in all aspects of coloration and pattern except that the terminal flap is white as opposed to being nearly black and the original tail was broken off at the time of collection. Meristic differences are presented in Table 5. Comparisons. Ptychozoon popaense sp. nov. is most closely related to P. lionotum sensu lato but differs from it by having 15–19 fused caudal lobes as opposed to 2–11; lacking as opposed to having a thick, dark, postorbital stripe; and having as opposed to lacking irregularly shaped, white, vertebral markings. From P. bannaense it differs by having 21 as opposed to 17 pore-bearing precloacal scales in males; 13 or 14 as opposed to 16 or 17 enlarged, transverse, subdigital lamellae on the fourth toe; slight as opposed to extreme posteriorly angled caudal lobes; minimal as opposed to extreme reduction in the width of the caudal lobes toward the end of the tail; and having as opposed to lacking irregularly shaped, white, vertebral markings. Combinations of other characters differentiating P. popaense sp. nov. from the other more distantly related species are presented in Table 4. Distribution. Ptychozoon popaense is know only from the type locality from Yay Su camp, 16 km northeast of Kyauk-pa-taung Town, Kyauk-pa-taung Township, Mt. Popa, Mandalay Region, Myanmar (Fig. 1). Etymology. The gender of the genus Ptychozoon is neutral, therefore the specific epithet popaense is the adjective used here in reference to the type locality, Mt. Popa, Mandalay Region. Natural history. The type material was collected in deciduous dipterocap forest (Fig. 3). Both specimens were found on the walls of a small wooden building while foraging for insects during the evening at approximately 1900 hrs.

Ptychozoon popaense sp. nov. Mt. Popa Parachute Gecko Fig. 2 Holotype. LSUHC 13508, adult male, collected by Myint Kyaw Thura on 27 May 2017 from Yay Su camp, 16 km northeast of Kyauk-pa-taung Town, Kyauk-pa-taung Township, Mt. Popa, Mandalay Region, Myanmar (20.88331��N, 95.26638��E, 629 m above sea level). Paratopotype. Adult female LSUHC 13507 bears the same collection data as the holotype. Diagnosis. Ptychozoon popaense s p. nov. can be differentiated from all other species of Ptychozoon based on having the unique combination of separated supranasals; presence of an infra-auricular flap; dorsal and caudal tubercles absent; 32���36 enlarged ventral scales; enlarged femorals absent; 21 enlarged, pore-bearing precloacals in the male and 12 dimpled precloacal scales in the female; 13 or 14 transversely enlarged subdigital lamellae under the fourth toe; 28 or 29 scales across the caudal flap; distal caudal lobes not fused into a long caudal flap; bases of 15���19 lobes fused anterior to the caudal flap; caudal lobes angling slightly posteriorly; thick, dark, postorbital stripe present; four dark, dorsal, body bands between limb insertions; irregularly shaped, with white, vertebral markings; and adults not having distinct immaculate black and white caudal bands dorsally (Table 4). Description of holotype. Adult male SVL 86.2 mm; head short (HL/SVL 0.26), wide (HW/SVL 0.20), depressed (HD/HL 0.43), distinct from neck; snout rounded at tip in dorsal profile; interorbital region flat; lores rounded; rostral scale large, rectangular, without dorsomedial groove, in contact posteriorly with two supranasals, one postnasal, and laterally with nostrils and first supralabials; supralabials (9R,10L), supralabial eight in midorbital position; infralabials (9R,10L); nostrils elliptical with long axis oriented dorsoventrally, occupying anterior portion of nasal scale, bordered anteriorly by rostral, dorsally by supranasal, posteriorly by four postnasals of varying sizes, and ventrally by first supralabial; scales on rostrum granular slightly larger than granular scales on top of head and occiput; no ridge of tubercles along mandibles; eyes large (ED/HL 0.24), less than snout length and EAO distance; pupil vertically elliptical, crenelated; supraciliaries elongate, posteriormost spinose; auricular opening rounded, lacking enlarged lobes; tympanum deeply sunk; infra-auricular flap broad, rounded, extending from below corner of mouth to base of neck, measuring 3.5 mm at its widest point; dorsal scales of infra-auricular flap large, subimbricate proximally, small juxtaposed distally, minute and granular ventrally; infra-auricular flap on right side deeply notched; mental triangular, wider than deep, bordered laterally by first infralabials and posteriorly by paired, rectangular postmentals contacting medially for 70% of their length; one row of enlarged sublabials bordering infralabials, anteriormost largest; gular scales small, rounded, grading abruptly into larger imbricating throat and ventral scales. Body dorsoventrally depressed, relatively stout (AG/SVL 0.47); axilla-groin cutaneous expansion (flap) 5.5 mm at midpoint of body and bearing enlarged, juxtaposed, rectangular scales dorsally and minute, juxtaposed, subrectangular scales ventrally; dorsal body scales minute, flat, rounded, juxtaposed, largest mid-dorsally; 36 transverse rows of large, smooth, flat, subimbricate ventrals, ventrals much larger than dorsals, decreasing in size laterally; 21 elevated, enlarged, pore-bearing, precloacal scales; seven rows of enlarged, post-precloacal scales; scales immediately anterior to vent granular. Limbs short, robust (FL/SVL 0.11; TBL/SVL 0.16); dorsal scales of forelimbs, flat, juxtaposed, slightly larger than dorsal body scales and juxtaposed ventral forelimb scales; anterior and posterior margins of forelimbs, posterior margins of hind limbs, and anterior margins of forelegs bearing wide, cutaneous flaps extending to bases of digits I and V, bearing subimbricate, moderately larger scales dorsally and smaller imbricate scales ventrally wide; wide predigital notch in preantebrachial flap; palmar scales smooth, rounded; digits fully webbed, relatively short, dorsoventrally compressed; undivided transverse subdigital lamellae number 12 (I), 12 (II), 15 (III), 15 (IV), 12 (V), distalmost lamellae V-shaped; claws arise from within the dorsal surface of digital pads; first digit lacks a claw; dorsal scales of hind limbs, flat, juxtaposed, smaller than dorsal body scales and flat, subimbricate scales of thigh; posterior margins of thigh and foreleg, and anterior margin of foreleg have wide, cutaneous flaps bearing subimbricate, moderately large scales dorsally and much smaller subimbricate scales ventrally; pretibial flap not contacting base of digit V; post-tibial flap contacts base of digit I; plantar scales smooth, subimbricate; digits fully webbed; transverse subdigital lamellae number 11 (I), 13 (II), 15 (III), 13 (IV), 14 (V), distalmost lamellae Vshaped; claws arise from within the dorsal surface of digital pads; and first digit lacks a claw. Tail original, flattened, moderate in length (TaL/SVL 1.04); two median rows of transversely widened, smooth subcaudals; postcloacal scales on heimpenial swelling large, flat, imbricate; dorsal caudals flat, juxtaposed, larger than dorsal body scales, occurring in whorls; 6���8 smaller scales between larger scales delimiting whorls; tail width and caudal lobes slightly decrease posteriorly; 25 caudal lobes on each side slightly angled posteriorly; 19 lobes fused at their bases, prior to grading posteriorly into short, straight-edged, terminal tail flap terminal tail flap short (8.6 mm), not wider than tail. Coloration in life (Fig. 2). Dorsal ground color of head, body and limbs light-green, that of tail light-brown; top of head bearing three elongate, grey markings extending onto nape; wide, grey preorbital stripe extends through eye terminating on nape, becoming progressively darker and thicker posteriorly; labials much lighter than ground color of head, bearing thin dark lines; four wide, sinuous, grey bands between limb insertions bearing diffuse anterior margins and black, well-defined posterior margins and extending onto flanks; a series of irregularly shaped, poorly defined, immaculate, white vertebral blotches extending from nape to sacrum; dorsal surface of limbs bearing grey, diffuse, irregularly shaped, banded to reticulate patterns; seven dark caudal bands becoming darker posteriorly separated by six lighter bands, posterior two lightest; terminal flap nearly black; ventral surfaces of head, body, and limbs beige, generally immaculate; subcaudal region distinctly banded. All colors have faded to varying shades of grey in preservative. Variation. The female paratopotype is nearly identical to the holotype in all aspects of coloration and pattern except that the terminal flap is white as opposed to being nearly black and the original tail was broken off at the time of collection. Meristic differences are presented in Table 5. Comparisons. Ptychozoon popaense sp. nov. is most closely related to P. lionotum sensu lato but differs from it by having 15���19 fused caudal lobes as opposed to 2���11; lacking as opposed to having a thick, dark, postorbital stripe; and having as opposed to lacking irregularly shaped, white, vertebral markings. From P. bannaense it differs by having 21 as opposed to 17 pore-bearing precloacal scales in males; 13 or 14 as opposed to 16 or 17 enlarged, transverse, subdigital lamellae on the fourth toe; slight as opposed to extreme posteriorly angled caudal lobes; minimal as opposed to extreme reduction in the width of the caudal lobes toward the end of the tail; and having as opposed to lacking irregularly shaped, white, vertebral markings. Combinations of other characters differentiating P. popaense sp. nov. from the other more distantly related species are presented in Table 4. Distribution. Ptychozoon popaense is know only from the type locality from Yay Su camp, 16 km northeast of Kyauk-pa-taung Town, Kyauk-pa-taung Township, Mt. Popa, Mandalay Region, Myanmar (Fig. 1). Etymology. The gender of the genus Ptychozoon is neutral, therefore the specific epithet popaense is the adjective used here in reference to the type locality, Mt. Popa, Mandalay Region. Natural history. The type material was collected in deciduous dipterocap forest (Fig. 3). Both specimens were found on the walls of a small wooden building while foraging for insects during the evening at approximately 1900 hrs., Published as part of Grismer, L. Lee, Wood, Perry L., Thura, Myint Kyaw, Grismer, Marta S., Brown, Rafe M. & Stuart, Bryan L., 2018, Geographically structured genetic variation in Ptychozoon lionotum (Squamata: Gekkonidae) and a new species from an isolated volcano in Myanmar, pp. 202-214 in Zootaxa 4514 (2) on pages 208-211, DOI: 10.11646/zootaxa.4514.2.4, http://zenodo.org/record/2607928

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

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 (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, Published as part of Travers, Scott L., Richards, Stephen J., Broadhead, Taylor S. & Brown, Rafe M., 2018, A new miniature Melanesian Forest Frog (Ceratobatrachidae: CORNufeR) from New Britain Island, constituting the first record of the subgenus BATRAchylOdes from outside of the Solomon Archipelago, pp. 23-44 in Zootaxa 4370 (1) on pages 26-35, DOI: 10.11646/zootaxa.4370.1.2, http://zenodo.org/record/1138506, {"references":["Brown, R. M., Siler, C. D., Richards, S. J., Diesmos, A. C. & Cannatella, D. C. (2015) Multilocus phylogeny and a new classification for Southeast Asian and Melanesian forest frogs (family Ceratobatrachidae). Zoological Journal of the Linnaean Society, 174, 130 - 168. https: // doi. org / 10.1111 / zoj. 12232","Brown, W. C. & Parker F. (1970) New frogs of the genus Batrachylodes (Ranidae) from the Solomon Islands. Breviora, 346, 1 - 31.","Allison, A. & Kraus, F. (2001) New species of Platymantis (Anura: Ranidae) from New Ireland. Copeia, 2001, 194 - 202. https: // doi. org / 10.1643 / 0045 - 8511 (2001) 001 [0194: NSOPAR] 2.0. CO; 2","Kraus, F. & Allison, A. (2009) New species of frogs from Papua New Guinea. Bishop Museum Occasional Papers, 104, 1 - 36.","Kraus, F. (2008) Remarkable case of anuran sexual size dimorphism: Platymantis rhipiphalcus is a junior synonym of Platymantis boulengeri. Journal of Herpetology, 42, 637 - 644. https: // doi. org / 10.1670 / 07 - 238 Rl. 1"]}

Siler, Cameron D., Davis, Drew R., Freitas, Elyse S., Huron, Nicholas A., Geheber, Aaron D., Watters, Jessa L., Penrod, Michelle L., Papeș, Monica, Amrein, Andrew, Anwar, Alyssa, Cooper, Dontae, Hein, Tucker, Manning, Annalisa, Patel, Neeral, Pinaroc, Lauren, Diesmos, Arvin C., Diesmos, Mae L., Oliveros, Carl H., Brown, Rafe M. (2016): Additions to Philippine Slender Skinks of the Brachymeles bonitae Complex (Reptilia: Squamata: Scincidae) II: a new species from the northern Philippines. Zootaxa 4132 (1): 15-29, DOI: http://doi.org/10.11646/zootaxa.4132.1.2

Brachymeles dalawangdaliri sp. nov. (Figs. 2, 3) Brachymeles bonitae Dum��ril & Bibron 1839; Taylor 1917; Brown 1956: 5; Brown & Rabor 1967: 526; Brown & Alcala 1970; Brown & Alcala 1980: 20; Davis et al. 2014; Geheber et al. 2016; Siler et al. 2016. Holotype. CAS 137149, adult male, collected among rotting leaves in early secondary growth forest on 24 May 1972, in Barrio Dubduban, Municipality of San Augustin, Romblon Province, Tablas Island, Philippines (12.58333 �� N, 122.10000�� E; WGS 84; 500���800 ft. elev.), by L.C. Alcala. Paratypes. Five adult females (CAS 137148, 137150, 137151, 137153, 137154) and one adult male (CAS 137152) were collected in Barrio Dubduban, Municipality of San Augustin, Romblon Province, Tablas Island, Philippines (12.58333 �� N, 122.10000�� E; WGS 84), by L.C. Alcala. CAS 137148 was collected in a rotting coconut trunk on 23 May 1972 (150���250 ft. elev.), CAS 137150���137152 were collected in humus under a rotting log in original forest on 1 June 1972 (450 ft. elev.), and CAS 137153, 137154 were collected in humus under a rotting log in original forest on 2 June 1972 (550 ft. elev.). Diagnosis. Following recent taxonomic revisions of Brachymeles (Siler et al. 2011; Davis et al. 2014) the new species is assigned to the B. bonitae Complex based on the following suite of morphological characters: (1) limbs present, (2) non-pentadactyl, (3) fore-limbs with 0���3 fingers. (4) hind limbs with 0���2 toes, (5) paravertebral scale rows ��� 91, (6) presacral vertebrae 47���53, (7) supraoculars four, (8) enlarged, differentiated nuchals present, (9) longitudinal rows of dark spots around the body absent, and (10) auricular opening absent. Brachymeles dalawangdaliri sp. nov. can be distinguished from congeners by the following combination of characters: (1) body size small (SVL 66.0��� 80.9 mm), (2) fore-limbs bidactyl, (3) hind limbs digitless, unidactyl, or bidactyl, (4) limb length short, (5) supralabials six or seven, (6) infralabials six or seven, (7) supraciliaries five, (8) supraoculars four, (9) midbody scale rows 24 or 25, (10) axilla���groin scale rows 80���83, (11) paravertebral scale rows 97���101, (12) prefrontal contact absent, (13) frontoparietal contact present or absent, (14) enlarged chin shields in two pairs, (15) nuchals enlarged, (16) auricular opening absent, (17) presacral vertebrae 49, and (18) uniform body color (Tables 1, 2). Comparisons. Brachymeles dalawangdaliri sp. nov. can be distinguished from B. ilocandia, B. isangdaliri, B. mapalonggaon, and B. tridactylus by having bidactyl fore-limbs and 0���2 digits on the hind limbs (versus being digitless [B. ilocandia, B. mapalanggaon], unidactyl [B. isangdaliri], or tridactyl [B. tridactylus]); from B. bonitae by having fewer number of presacral vertebrae (49 versus 53), fewer number of axilla���groin scale rows (80���83 versus 83���90), fewer number of paravertebral scale rows (97���101 versus 103���110), a greater number of infralabials (6, 7 versus 5), and by the absence (versus presence) of a 3 rd chin shield and fused mental; from B. ilocandia by having a longer fore-limb (1.4���2.2 mm versus 1.1���1.4) and hind limb length (1.9���2.2 mm versus 1.6��� 1.9), fewer number of presacral vertebrae (49 versus 50���53), and by the absence (versus presence) of a 3 rd chin shield pair; from B. isangdaliri by having a longer snout���vent length (66.0��� 80.9 mm versus 59.5), a longer forelimb length (1.4���2.2 mm versus 1.3), a greater number of presacral vertebrae (49 versus 47), greater number of midbody scale rows (24, 25 versus 22), a greater number of axilla���groin scale rows (80���83 versus 73), and fewer number of supraciliaries (5 versus 6); from B. ligtas by having a longer fore-limb length (1.4���2.2 mm versus 1.2��� 1.4), fewer number of presacral vertebrae (49 versus 50), a greater number of midbody scale rows (24, 25 versus 22), a greater number of axilla���groin scale rows (80���83 versus 74���76), a greater number of paravertebral scale rows (97���101 versus 91���93), and by the absence (versus presence) of a 3 rd chin shield pair; from B. mapalanggaon by having a longer fore-limb (1.4���2.2 mm versus 0.8 ���1.0) and hind limb length (1.9���2.2 mm versus 1.2���1.6), fewer number of presacral vertebrae (49 versus 51), and a greater number of midbody scale rows (24, 25 versus 22, 23); from B. tridactylus by having a shorter hind limb length (1.9���2.2 mm versus 2.6���3.6), a greater number of presacral vertebrae (49 versus 47), a greater number of axilla���groin scale rows (80���83 versus 72���79), and by the absence (versus presence) of a 3 rd chin shield pair. Brachymeles dalawangdaliri sp. nov. can be distinguished from all limbless species of Brachymeles by having limbs, and from all pentadactyl species of Brachymeles by having bidactyl fore-limbs and digitless, unidactyl, or bidactyl hind limbs. Description of holotype. Details of the head scalation are shown in Figure 2. Adult, male, body small, slender, SVL 74.9 mm; head weakly differentiated from neck, nearly as wide as body, HW 7.9 % SVL, 107.3 % HL; HL 37.1 % SnFa; SnFa 19.7 % SVL; snout narrow, sharply rounded in dorsal and lateral profile, SNL 60.2 % HL; ear completely hidden by scales; eyes small, ED 17.9 % HL, 40.5 % END, pupil subcircular; body slightly depressed, nearly uniform in thickness, MBW 100.0% MBH; scales smooth, glossy, imbricate; longitudinal scale rows at midbody 24; paravertebral scale rows 97; axilla���groin scale rows 80; limbs short, diminutive, bluntly rounded, with digits reduced to two small digit growths on fore-limbs, single digit growth on hind limb; finger and toe lamellae absent; FLL 3.6 % AGD, 2.8 % SVL; HLL 3.9 % AGD, 3.0% SVL; tail not as wide as body, TW 72.6 % MBW, autotomized posterior to cloaca, length unavailable. Rostral projecting onto dorsal snout to level in line with posterior edge of nasal, roughly equal in width and height, in contact with frontonasal; frontonasal wider than long; nostril small, ovoid, in center of single trapezoidal nasal, longer axis directed posterodorsally and anteroventrally; supranasals present; postnasals absent; prefrontals moderately separated; frontal roughly hexagonal-shaped, its anterior margin in moderate contact with frontonasal, in contact with first two anterior supraoculars, 4 �� wider than anterior supraocular; supraoculars four; frontoparietals moderate in size, in moderate medial contact, frontoparietals in contact with supraoculars two and three; interparietal moderate in size, its length roughly equal to 2 �� midline length of frontoparietal, longer than wide, diamond-shaped, wider anteriorly, pineal eyespot not visible; parietals wider than frontoparietals, in moderate contact behind interparietal; head scales damaged in nuchal region; enlarged nuchals present; loreals two, anterior loreal about as long as and slightly higher than posterior loreal; preoculars two; presubocular one; supraciliaries five, the anteriormost contacting prefrontal and separating posterior loreal from first supraocular, posteriormost extending to midline edge of fourth supraocular; subocular scale row single, incomplete, in contact with supralabials; lower eyelid with one row of scales; supralabials seven, first twice the width of others, third, fourth and fifth subocular; infralabials seven (Figs. 2, 3). Mental wider than long, in contact with first infralabial on left side of head, fused with first infralabial on right side of head; postmental single, enlarged, its width equal to width of mental; followed by two pairs of enlarged chin shields, first pair moderately separated, second pair slightly smaller than first, broadly separated by three medial scales (Figs. 2, 3). Scales on limbs smaller than body scales. Variation. Differences in digit number and scalation patterns were observed between the holotype and paratype series. The holotype (CAS 137149) and three paratypes (CAS 137148, 137153, 137154) possessed seven supralabials and seven infralabials, with three other paratypes (CAS 137150���137152) having six supralabials and six infralabials. Unlike the holotype and majority of paratypes, the frontoparietal scales were observed to contact medially in two specimens (CAS 137153, 137154). Observed digit number variation included body plans with two fingers and one toe (CAS 137149 [holotype], 137151, 137153, 137154), two fingers and two toes (CAS 137148, 137150), and two fingers and no toes (CAS 137152). Coloration of holotype in life. Coloration in life is unrecorded; however, because Brachymeles specimens do not change significantly during preservation (Siler & Brown 2010; Siler et al. 2011 a, 2012 a; Davis et al. 2014), we suspect that the preserved coloration and patterns are much like those in life. Coloration of holotype in preservative. The background color on the dorsal, lateral and ventral sides of the body and tail is a solid Prout���s Brown (Color 47; K��hler 2012). The ventral side of the head is the same solid color as the ventral side of the body (Fig. 3; Prout���s Brown; Color 47; K��hler 2012). On the dorsal and lateral sides of the head, a single Fuscous splotch can be seen just above each orbit (Fig. 3; Color 283; K��hler 2012). Etymology. The name of the new species presented in this paper is derived from the Tagalog (Filipino) terms ���dalawa,��� meaning two, and ���daliri,��� meaning finger, referencing one of the defining characteristics of the new species, the two digits on the fore-limbs. Suggested common name: Tablas Slender Skink. Distribution. Brachymeles dalawangdaliri sp. nov. is known only from Tablas Island, Romblon Island Group, Philippines (Fig. 1). TABLE ��. Summary of meristic anđ mensural characters among species of the Brachymeles bonitae Complex. Sample size, bođy length anđ total length among males anđ females, anđ general geographical đistribution (PAIC = Pleistocene Aggregate Islanđ Complexes, sensu Brown & Diesmos [2002]) are incluđeđ for reference (SVL, TotL, FLL, HLL given as range over mean �� stanđarđ đeviation; all bođy proportions given as percentage over mean �� stanđarđ đeviation). bonitae dalawangdaliri sp. nov. ilocandia isangdaliri ligtas mapalanggaon tridactylus (3 m, 1 f) (2 m, 5 f) (9 f) (1 f) (3 m, 2 f) (3 m, 6 f) (12 m, 9 f) Range Luzon & Polillo Tablas Islanđ Luzon & Luzon Islanđ Lubang Islanđ Masbate Islanđ West Visayan islanđs Camiguin Norte PAIC islanđs SVL (f) 69.4 69.8 = 80.9 65.7 = 77.6 59.5 60.7 = 69.2 61.7 = 75.8 59.9 = 82.3 (73.6 �� 4.4) (73.7 �� 3.7) (65.0 �� 6.0) (67.2 �� 5.4) (71.4 �� 6.9) SVL (m) 69.7 = 78.4 66.0= 74.9 N /A N/A 69.4 = 79.6 65.1 = 72.7 60.7 = 77.6 (72.8 �� 4.8) (70.4 �� 6.3) (74.5 �� 5.1) (68.4 �� 3.9) (69.0 �� 6.0) TotL (f) N/A N/A 122.3 =146.0 106.1 119.4 120.2 133.6 1 (134.1 �� 8.0) TotL (m) 122.0 N /A N/A N/A 160.6 112.6 = 118.6 120.9 = 154.1 (115.6 �� 4.3) (136.0 �� 9.8) / SVL 73 N/A 81 = 90 78 97 = 102 67 = 84 85 = 112 (85 �� 3) (99 �� 4) (78 �� 9) (95 �� 10) FLL 1.0= 1.7 1.4 = 2.2 1.1 = 1.4 1.3 1.2 = 1.4 0.8 =1.0 1.5 = 2.5 (1.3 �� 0.3) (1.9 �� 0.3) (1.3 �� 0.1) (1.3 �� 0.1) (0.9 �� 0.1) (2.0 �� 0.3) FLL / SVL 1 = 2 2 = 3 1 = 2 2 2 = 2 1 = 2 2 = 4 (2 �� 0) (3 �� 0) (2 �� 0) (2 �� 0) (1 �� 0) (3 �� 0) HLL 1.5 = 2.3 1.9 = 2.2 1.6 = 1.9 2.2 1.6 =2.0 1.2 = 1.6 2.6 = 3.6 (1.9 �� 0.3) (2.1 �� 0.1) (1.8 �� 0.1) (1.8 �� 0.1) (1.4 �� 0.1) (3.1 �� 0.3) HLL / SVL 2 = 3 2 = 3 2 = 3 4 2 = 3 2 = 2 3 = 6 (3 �� 0) (3 �� 0) (2 �� 0) (3 �� 0) (2 �� 0) (5 �� 1) TABLE 2. Summary of meristic anđ qualitative điagnostic characters (present, absent) among species of the Brachymeles bonitae Complex. The pairs of enlargeđ scales posterior to the postmental scale are abbreviateđ as chin shielđ pairs with reference to the 1 st, 2 nđ, anđ 3 rđ pairs (when present). In cases of scale count variation within species, numbers of inđiviđuals showing specific counts are given in parentheses. bonitae dalawangdaliri sp. nov. ilocandia isangdaliri ligtas mapalanggaon tridactylus (3 m, 1 f) (2 m, 5 f) (9 f) (1 f) (3 m, 2 f) (3 m, 6 f) (12 m, 9 f) Number of đigits (fingers/toes) 0/0 1 2 /0= 2 0/0 1 / 1 2 /0 0/0 3 / 3 2 / 1 1 PSV 53 49 50 = 53 47 50 51 47 MBSR 21 = 24 24, 25 22 = 24 22 22 22, 23 22 = 24 AGSR 83 = 90 80 = 83 80 = 82 73 74 = 76 80 = 84 72 = 79 PVSR 103 = 110 97 = 101 97 = 100 97 91 = 93 99 = 102 90 = 98 6 6 (3) 6 6 6 6 6 (13) 7 (4) 7 (8) 5 6 (3) 5 (3) 6 6 5 (8) 6 (13) 7 (4) 6 (6) 6 (1) 7 (8) 5 5 5 6 5 5 5 4 4 4 4 4 4 4 Prefrontal contact Absent Absent Absent or Point Absent Absent Absent Absent contact Frontoparietal contact Absent Present or Absent Absent Absent Present Point contact or Absent Absent chin shielđ pair contact Absent Absent Absent Absent Present or Absent Absent Present or Absent chin shielđ pair Present Absent Present Absent Present Present or Absent Present Mental/ 1 st IFL fusion Present Absent Present or Absent Absent Absent Present or Absent Absent Enlargeđ nuchals Present Present Present Present Present Present Present Longituđinal rows of đark Absent Absent Absent Absent Absent Absent Absent spots Observeđ for two inđiviđuals. Natural history. Presumably, Brachymeles dalawangdaliri sp. nov. once occurred in primary growth forest; however, little to no primary forest remains on Tablas Island. This species has never been observed in high densities, a pattern consistent with many members of the B. bonitae Complex. Furthermore, no recent collections of this species have been made, with the last documented observation of B. dalawangdaliri sp. nov. in the wild occurring more than four decades ago. Brachymeles dalawangdaliri sp. nov. may be found in sympatry with B. talinis (Siler & Brown 2010). Little is known about the ecology and distribution of Brachymeles dalawangdaliri sp. nov. on Tablas Island. However, this species is endemic to an island with complete or near complete deforestation and no recent observations of wild populations. Therefore, in evaluating B. dalawangdaliri sp. nov. against the IUCN criteria for classification, we find that it qualifies for the status of Vulnerable (VU) based on the following criteria: VU B 2 ab(iii,iv); D 2 (IUCN 2015)., Published as part of Davis, Drew R., Geheber, Aaron D., Watters, Jessa L., Penrod, Michelle L., Feller, Kathryn D., Ashford, Alissa, Kouri, Josh, Nguyen, Daniel, Shauberger, Kathryn, Sheatsley, Kyra, Winfrey, Claire, Wong, Rachel, Sanguila, Marites B., Brown, Rafe M. & Siler, Cameron D., 2016, Additions to Philippine Slender Skinks of the Brachymeles bonitae Complex (Reptilia: Squamata: Scincidae) III: a new species from Tablas Island in Zootaxa 4132 (1), DOI: 10.11646/zootaxa.4132.1.3, http://zenodo.org/record/258936, {"references":["Dumeril, A. M. C. & Bibron, G. (1839) Erpetologie General ou, Histoire Naturelle Complete des Reptiles. Librairie Encylopedique de Roret, Paris, 488 pp.","Taylor, E. H. (1917) Brachymeles, a genus of Philippine lizards. Philippine Journal of Science, 12, 267 - 279.","Brown, W. C. & Rabor, D. S. (1967) Review of the genus Bracheymeles (Scincidae), with descriptions of new species and subspecies. Proceedings of the California Academy of Sciences, Fourth Series, 15, 525 - 548.","Brown, W. C. & Alcala, A. C. (1970) The zoogeography of the herpetofauna of the Philippine Islands, a fringing archipelago. Proceedings of the California Academy of Sciences, Series 4, 18, 105 - 130.","Brown, W. C. & Alcala, A. C. (1980) Philippine Lizards of the Family Scincidae. Philippine University Press, Dumaguete City, 264 pp.","Siler, C. D., Fuiten, A. M., Jones, R. M., Alcala, A. C. & Brown, R. M. (2011 a) Phylogeny-based species delimitation in Philippine slender skinks (Reptilia: Squamata: Scincidae) II: taxonomic revision of Brachymeles samarensis and description of five new species. Herpetological Monographs, 25, 76 - 112. http: // dx. doi. org / 10.1655 / herpmonographs-d- 10 - 00013.1","Siler, C. D., Jones, R. M., Diesmos, A. C., Diesmos, M. L. & Brown, R. M. (2012 a) Phylogeny-based species delimitation in Philippine slender skinks (Reptilia: Squamata: Scincidae) III: taxonomic revision of the Brachymeles gracilis Complex, with descriptions of three new species. Herpetological Monographs, 26, 135 - 172. http: // dx. doi. org / 10.1655 / herpmonographs-d- 11 - 00006.1","Kohler, G. (2012) Color Catalog for Field Biologists. Herpeton, Offenbach, 49 pp.","Brown, R. M. & Diesmos, A. C. (2002) Application of lineage-based species concepts to oceanic island frog populations: the effects of differing taxonomic philosophies on the estimation of Philippine biodiversity. Silliman Journal, 42, 133 - 162."]}

Pseudogekko isapa sp. nov. (Fig. 2) Holotype. PNM 9816 (ACD Field No. 8920), adult male, collected at 21:30 hrs in 2014, in Mt. Guiting-Guiting Natural Park, Municipality of Magdiwang, Barangay Talaba, Romblon Province, Sibuyan Island, Philippines (12.401873��N, 122.539747��E; WGS 84; 479 m elev.), by ACD and party. Paratype. One adult female (CAS 139713) collected on 22 May 1972, on a small vine in secondary forest, from Dubduban Barrio, Municipality of San Agustin, Romblon Province, Tablas Island, Philippines (12.58332��N, 122.1��E; WGS 84; ~ 17 m elev.), by L. C. Alcala and party. Diagnosis. Pseudogekko isapa sp. nov. can be distinguished from congeners by the following combination of characters: (1) body elongate (SVL 62.1, 63.4 mm), (2) axilla���groin distance long (32.1, 33.0 mm), (3) relative head length long (19% SVL), (4) relative snout length long (62, 64% head length), (5) Finger-III scansors 13, (6) Toe-IV scansors 17, (7) paravertebrals 240, 246, (8) ventral scales 135, 141, (9) supralabials 20, 21, (10) infralabials 17, 19, (11) circumorbitals 50, 54, (12) precloacal pores 15 in males, none in females, (13) femoral pores absent, (14) conspicuous head spotting present, (15) conspicuous dorsolateral spotting absent, (16) body striping absent, and (17) interorbital banding present (Tables 1, 2; Fig. 2). Comparisons. Characters distinguishing Pseudogekko isapa sp. nov. from all other species of the Pseudogekko compresicorpus Complex sensu Siler et al. (2014b) are summarized in Tables 1, 2. Pseudogekko isapa sp. nov. most closely resembles P. compresicorpus; however, it differs from this species by having a longer snout���vent length (SVL 62.1���63.4 mm versus 54.9���59.7 mm), wider body width (midbody width 8.5���8.9 mm versus 5.4���6.7 mm), greater relative head length (head length 19% SVL versus 16���18%), fewer Finger-III (13 versus 15���17) and Toe-IV (17 versus 18 or 19) scansors, more (17���19 versus 13���16), more circumorbitals (50���54 versus 39���45), more paravertebrals (240���246 versus 226���234), more ventrals (135���141 versus 127���130), and more enlarged precloacal pore-bearing scales (15 versus 10���14; Fig. 2). From the remaining members of the P. compresicorpus Complex, P. isapa sp. nov. can be distinguished from P. pungkaypinit by having a smaller body size (SVL 62.1���63.4 mm versus 66.6���76.8 mm), smaller axilla���groin distance (32.1���33.0 mm versus 37.2���41.2), longer relative snout length (62���64% head length versus 54���60%), fewer Finger-III scansors (13 versus 15���17), fewer paravertebrals (240���246 versus 265���280), fewer enlarged pore-bearing precloacals (15 versus 17���20), and the absence (versus presence) of body striping; and from P. chavacano and P. ditoy by having a larger body size (SVL 62.1���63.4 mm versus 54.7���55.9 [P. chavacano] or 49.4��� 52.6 [P. d i t o y]), larger axilla���groin distance (32.1���33.0 mm versus 26.7���30.0 [P. chavacano] or 25.1���29.7 [P. ditoy]), longer relative snout length (62���64% head length versus 55���58% [P. chavacano] or 57���59% [P. ditoy]), fewer Finger-III scansors (13 versus 15 or 16 [P. chavacano] or 14 or 15 [P. d i t oy]) and enlarged pore-bearing precloacals (15 versus 16 [P. chavacano] or 18 [P. ditoy]), and more circumorbitals (50���54 versus 46 [P. chavacano] or 40���43 [P. d i t oy]), paravertebrals (240���246 versus 195���197 [P. chavacano] or 180���185 [P. ditoy]), and ventrals (135���141 versus 122 or 123 [P. chavacano] or 111���118 [P. ditoy]; Tables 1, 2). Pseudogekko isapa sp. nov. can be distinguished from members of the P. brevipes Complex (P. brevipes, P. atiorum), by having a larger body size (SVL 62.1���63.4 mm versus 41.1���52.5 [P. a t i o r u m] or 34.5���42.4 [P. brevipes]), more supralabials (20 or 21 versus 15���17 [P. a ti o ru m] or 13 or 14 [P. brevipes]), infralabials (17���19 versus 12���15 [P. a t io ru m] or 14 or 15 [P. brevipes]), circumorbitals (50���54 versus 35���38 [P. atiorum] or 33���35 [P. brevipes]), and ventrals (135���141 versus 119���129 [P. atiorum] or 96���117 [P. brevipes]). From P. smaragdinus, P. isapa sp. nov. can be distinguished by having fewer Finger-III scansors (13 versus 15��� 18) and lacking pore-bearing femorals (15 versus 32���41), more supralabials (20 or 21 versus 16���19), circumorbitals (50���54 versus 33���35), and ventrals scales (135���141 versus 124���130), a tendency towards having fewer Toe-IV scansors (17 versus 16���22), a tendancy towards having more infralabials (17 or 19 versus 14���17), and by the presence of brownish tan (versus yellow to orange [undisturbed] or neon green [disturbed]) body coloration, and absence (versus presence) of enlarged femoral pore-bearing scales in males. Description of holotype. Adult male, well preserved; small incision in the sternal region (portion of liver removed for genetic a sample). Body moderately large, elongate, slender; limbs well developed, moderately slender; tail autotomized, absent; margins of limbs smooth, cutaneous flaps or dermal folds absent. Head size moderate, differentiated from neck, characterized by only slightly hypertrophied temporal and adductor musculature; snout broadly rounded in dorsal view and sharply rounded in lateral view; head width 118.8% midbody width, 85.6% head length; head length 19.0% SVL; snout length 74.2% head width, 63.6% head length; dorsal surfaces of head relatively homogeneous, with slightly to moderately pronounced concave postnasal, internasal, prefrontal, and interorbital concavities; auricular opening small, teardrop-shaped, beneath temporal swellings on either side of head; tympanum deeply sunken; orbit large; eye large, pupil vertical, margin wavy; limbs and digits relatively short; thighs moderately thicker compared to brachium; tibia length 8.2% SVL, 53.1% femur length. Rostral size moderately large, margin oval in anterodorsal view, nearly as broad as high, sutured anterolaterally with anteriormost enlarged supralabials, projecting onto dorsal surface of head to point in line anteriormost edge of nasal; nasal surrounded by first labial, rostral, one enlarged postnasal, and two enlarged supranasals; medial supranasals separated by three moderately enlarged median scales; enlarged supranasals larger than postnasals, anteriormost supranasal largest. Differentiated supralabials bordered dorsally by one row of slightly enlarged snout scales; differentiated infralabials bordered ventrally by undifferentiated scales or one row of slightly enlarged scales around the lower jaw; mentals and postrictals undifferentiated; gulars small, round, nonimbricate, juxtaposed; dorsal cephalic scales fairly homogeneous in size, shape, disposition, and distribution; cephalic scalation slightly convex, round to oval scales; posterior head scales granular, slightly raised into flat, plateaued dorsal surfaces. Axilla���groin distance 50.6% SVL; dorsal body scales round, slightly raised into flat, dorsal plateaued, juxtaposed, relatively homogeneous in size; each dorsal scale surrounded by 4���6 interstitial granules, many forming a ���Star of David��� pattern around dorsal scale (Siler et al. 2014b: Fig. 5); dorsals gradually transition to imbricate ventrals along lateral body surface; scales on dorsal surfaces of limbs more imbricate than dorsals; scales on dorsal surfaces of hands and feet similar to dorsal limb scales, heavily imbricate; ventral body scales flat, cycloid, strongly imbricate, much larger than lateral or dorsal body scales, relatively homogeneous in size. Enlarged pore-bearing scales in continuous precloacal series, arranged in a widely obtuse, ���V���-formation (Fig. 2); patch of slightly enlarged scales posterior to precloacal series, roughly 6 or 7 scale rows in width, 3 or 4 scale rows in length, forming an oval patch just posterior to precloacal series (Fig. 2). Digits moderately expanded and covered on palmar and plantar surfaces by bowed, unnotched, undivided scansors (Fig. 2); digits with minute vestiges of interdigital webbing on hands, moderate interdigital webbing on feet; subdigital scansors of fingers (left/right) I (10,9), II (11,11), III (13,13), IV (18,17), V (11,11); subdigital scansors of toes (left/right) I (10,9), II (12,11), III (16,17), IV (17,17), V (11,11); subdigital scansors of hands and feet bordered proximally (on palmar and plantar surfaces) by 1���3 slightly enlarged scales that form a nearcontinuous series with enlarged scansors; all digits clawed, but first (inner) claw greatly reduced; remaining terminal claw-bearing phalanges compressed, with moderately sized recurved claws. Coloration of holotype in preservative. Background dorsal trunk coloration light brownish tan with intermittent to dense small dark brown speckles, continuing down to base of tail; dorsal region of head with similar color pattern, except for pronounced dark brown area above orbits; dorsal and lateral regions of limbs with same color pattern as trunk; supralabials and infralabials with similar pattern as head, but dark brown spots more diffuse; ventral side of trunk with same background coloration, dark brown speckling pattern significantly reduced; ventral side of head with sparse dark brown speckling only; ventral surfaces of the limbs, and palms of hands and feet lighter cream colored with medium to dark brown speckles scattered sparsely throughout; ventral surfaces of digits darker tan to grey with irregular, sparse dark brown speckles; skin between scansors dark brown. Coloration in life unrecorded. Measurements and scale counts of holotype in mm. SVL 63.4; axilla���groin distance 32.1; total length N/A; tail length N/A; head length 11.8; head width 10.1; head depth 7.0; eye diameter 3.9; snout length 7.5; eye���nares distance 5.9; internarial distance 2.6; interorbital distance 7.0; midbody width 8.5; femur length 9.8; tibia length 5.2; Toe-I length 2.3; Toe-IV length 4.1; tail width 5.3; tail depth 4.7; supralabials 20; infralabials 19; circumorbitals 50; paravertebral scales 240; ventral scales 135; Finger-III scansors 13; Toe-IV scansors 17. Distribution, ecology and natural history. Pseudogekko isapa sp. nov. is known from the islands of Tablas and Sibuyan in the Romblon Island Group (Fig. 1). This species also may occur on Romblon Island itself, and eventually may be found on several of the smaller islands scattered across the province, such as Banton and Simara islands just off of the north coast of Tablas Island. Besides P. atiorum, P. isapa sp. nov. is the only other species outside of the Luzon and Mindanao PAICs; all other species are distributed across the northern, eastern, and southern islands (Siler et al. 2014a, b; Davis et al. 2015a). Like most members of the genus, P. isapa sp. nov. appears to be primarily a forest obligate species; the holotype was collected within high quality secondary forest in Mt. Guiting- Guiting Natural Park on Sibuyan Island. This species��� poor representation in museum collections globally is a testament to the secretive nature of these species in their native habitat (Siler et al. 2014a, b). Due to the absence of data allowing for confirmation of potential declines in population size as a result of habitat destruction, and the lack of available information about the species��� natural history, we do not find that it qualifies for formal threatened status under IUCN criteria (IUCN 2015). Therefore, we recommend that the species be considered Data Deficient at the current time, but strongly recommend that immediate and comprehensive survey efforts be made to study the health of wild populations on islands throughout the Romblon Island Group (Fig. 1). With at least a very large portion of its range dominated by Mt. Guiting-Guiting Natural Park, the majority of its hypothesized distribution is very well protected. Etymology. We derive the new species epithet from the Tagalog (Filipino) phrase isa pa, meaning ���another��� or ���one more��� in reference to our surprise at the discovery of yet another distinctive new species of Pseudogekko immediately following our review of the genus (Siler et al. 2014a), including what we thought was a comprehensive review of the P. compresicorpus (Siler et al. 2014b) and P. brevipes complexes (Davis et al. 2015a). Suggested common name: Romblon Province False Gecko. isapa sp. nov. (1 m, 1 f) SVL 63.4, 62.1 Axilla���groin distance 32.1, 33.0 Total length ��� Midbody width 8.5, 9.9 Head length 11.8, 11.7 Head width 10.1, 9.4 Head height 7.0, 7.2 Snout length 7.5, 7.2 Eye diameter 3.9, 4.0 Eye���nares distance 5.9, 6.0 Internarial distance 2.6, 2.4 Interorbital distance 7.0, 6.7 Femur length 9.8, 10.1 Tibia length 5.2, 5.5, Published as part of Siler, Cameron D., Davis, Drew R., Diesmos, Arvin C., Guinto, Faith, Whitsett, Collin & Brown, Rafe M., 2016, A new species of Pseudogekko (Squamata: Gekkonidae) from the Romblon Island Group, Central Philippines, pp. 248-260 in Zootaxa 4139 (2) on pages 249-255, DOI: 10.11646/zootaxa.4139.2.8, http://zenodo.org/record/272017, {"references":["Siler, C. D., Welton, L. J., Davis, D. R., Watters, J. L., Davey, C. S., Diesmos, A. C., Diesmos, M. L. & Brown, R. M. (2014 b) Taxonomic revision of the Pseudogekko compresicorpus Complex (Reptilia: Squamata: Gekkonidae), with descriptions of three new species. Herpetological Monographs, 28, 110 - 139. http: // dx. doi. org / 10.1655 / herpmonographs-d- 14 - 00005","Siler, C. D., Dececchi, T. A., Merkord, C. L., Davis, D. R., Christiani, T. J. & Brown, R. M. (2014 a) Cryptic diversity and population genetic structure in the rare, endemic, forest-obligate, slender geckos of the Philippines. Molecular Phylogenetics and Evolution, 70, 204 - 209. http: // dx. doi. org / 10.1016 / j. ympev. 2013.09.014","Davis, D. R., Watters, J. L., Kohler, G., Whitsett, C., Huron, N. A., Brown, R. M., Diesmos, A. C. & Siler, C. D. (2015 a) Redescription of the rare Philippine slender gecko Pseudogekko brevipes (Reptilia: Squamata: Gekkonidae) and description of a new species. Zootaxa, 4020, 357 - 374. http: // dx. doi. org / 10.11646 / zootaxa. 4020.2.7"]}

Geheber, Aaron D., Davis, Drew R., Watters, Jessa L., Penrod, Michelle L., Feller, Kathryn D., Davey, Conner S., Ellsworth, Elyse D., Flanagan, Rachel L., Heitz, Brendan B., Moore, Tana, Nguyen, Marie D. C., Roberts, Austyn, Sutton, John, Sanguila, Marites B., Linkem, Charles W., Brown, Rafe M., Siler, Cameron D. (2016): Additions to Philippine Slender Skinks of the Brachymeles bonitae Complex (Reptilia: Squamata: Scincidae) I: a new species from Lubang Island. Zootaxa 4132 (1): 1-14, DOI: http://doi.org/10.11646/zootaxa.4132.1.1

Brachymeles ligtas sp. nov. (Figs. 2, 3) Brachymeles bonitae Duméril & Bibron 1839; Taylor 1917; Brown 1956: 5; Brown & Rabor 1967: 526; Brown & Alcala 1970; Brown & Alcala 1980: 20; Davis et al. 2014. Holotype. PNM 9818 (CDS Field No. 3886, formerly KU 320472), adult female, collected on 26 April 2009 (14:00 h) in Sitio Dangay, Barangay Vigo, Municipality of Lubang, Occidental Mindoro Province, Lubang Island, Philippines (13.79995° N, 120.163930° E; WGS 84; 45 m elev. elevation), by J. Fernandez and CDS. Paratypes (Paratopotypes). Four paratopotypes were also collected in Sitio Dangay, Barangay Vigo, Municipality of Lubang, Occidental Mindoro Province, Lubang Island, Philippines (13.79995° N, 120.163930° E; WGS 84; 45 m elev.), by J. Fernandez and CDS. One adult male (KU 320470) was collected on 24 April 2009 at 14:00 h, two adult females (KU 320471, 320473) were collected on 26 April 2009 at 14:00 h, and one juvenile (KU 320474) was collected on 29 April 2009 at 14:00 h. Paratypes. One adult male (KU 307755) was collected on 8 December 2005 in Barangay Vigo, Municipality of Lubang, Occidental Mindoro Province, Lubang Island, Philippines (13.826552° N, 120.120514° E; WGS 84; 27 m elev.), by RMB, CDS, and CWL. Diagnosis. Following recent taxonomic revisions of Brachymeles (Siler et al. 2011; Davis et al. 2014) the new species is assigned to the B. bonitae Complex based on the following suite of morphological characters: (1) limbs present, (2) non-pentadactyl, (3) fore-limbs with 0–3 fingers, (4) hind limbs with 0–2 toes, (5) paravertebral scale rows ≥ 91, (6) presacral vertebrae 47–53, (7) supraoculars four, (8) enlarged, differentiated nuchals present, (9) longitudinal rows of dark spots around the body absent, and (10) auricular opening absent. Brachymeles ligtas sp. nov. can be distinguished from congeners by the following combination of characters: (1) body size small (SVL 60.7–79.6 mm), (2) fore-limbs bidactyl, (3) hind limbs digitless, (4) limb length short, (5) supralabials six, (6) infralabials six, (7) supraciliaries five, (8) supraoculars four, (9) midbody scale rows 22, (10) axilla–groin scale rows 74–76, (11) paravertebral scale rows 91–93, (12) prefrontal contact absent, (13) frontoparietal contact present, (14) enlarged chin shields in three pairs, (15) nuchals enlarged, (16) auricular opening absent, (17) presacral vertebrae 50, and (18) uniform body color (Tables 1, 2). Comparisons. Brachymeles ligtas sp. nov. can be distinguished from other species in the B. bonitae Complex (B. bonitae, B. isangdaliri, B. mapalanggaon, B. tridactylus), by the number of presacral vertebrae (50 versus 53 [B. bonitae], 51 [B. mapalanggaon], 47 [B. isangdaliri, B. tridactylus]), and by having bidactyl fore-limbs and digitless hind limbs (versus bidactyl fore-limbs and unidactyl hind limbs [B. bonitae], digitless [B. bonitae, B. mapalanggaon], unidactyl [B. isangdaliri], or tridactyl [B. tridactylus]; Table 2); further, from B. bonitae by having a greater number of infralabials (6 versus 5) and absence (versus presence) of a fused mental and first chin shield; from B. isangdaliri by having fewer supraciliaries (5 versus 6) and the presence (versus absence) of a third chin shield pair; from B. mapalanggaon by having a longer fore-limb length (1.2–1.4 mm versus 0.8–1.0) and a longer hind limb length (1.6–2.0 mm versus 1.2–1.6); from B. tridactylus by having a shorter fore-limb length (1.2–1.4 mm versus 1.5–2.5); from B. isangdaliri and B. tridactylus by having a greater number of presacral vertebrae (50 versus 47 [B. isangdaliri, B. tridactylus]) and a shorter hind limb length (1.6–2.0 mm versus 2.2 [B. isangdaliri] or 2.6–3.6 [B. tridactylus]); from B. bonitae and B. mapalanggaon by having fewer presacral vertebrae (50 versus 53 [B. bonitae] or 51 [B. mapalanggaon]), fewer axilla–groin scale rows (74–76 versus 83–90 [B. bonitae] or 80–84 [B. mapalanggaon]); from B. bonitae, B. isangdaliri, and B. mapalanggaon by having fewer paravertebral scale rows (91–93 versus 103–110 [B. bonitae], 97 [B. isangdaliri], or 99–102 [B. mapalanggaon]); from B. bonitae, B. isangdaliri, and B. tridactylus by the presence of frontoparietals in contact (versus not in contact). Finally, Brachymeles ligtas sp. nov. can be distinguished from all limbless species of Brachymeles by having limbs, and from all pentadactyl species of Brachymeles by having bidactyl fore-limbs and digitless hind limbs. Description of holotype. Adult female, body small, slender, SVL 79.6 mm; head weakly differentiated from neck (Fig. 2, 3), nearly as wide as body, HW 5.7% SVL, 96.0% HL; HL 34.2% SnFa; SnFa 17.3% SVL; snout rounded in dorsal and lateral profile, SNL 56.0% HL; ear completely hidden by scales; eyes small, ED 21.3% HL, 50.9% END, pupil subcircular; body slightly depressed, nearly uniform in thickness, MBW 124.7% MBH; scales smooth, glossy, imbricate; longitudinal scale rows at midbody 22; paravertebral scale rows 93; axilla–groin scale rows 76; limbs short, diminutive, bluntly rounded, with digits reduced to two small digit growths on fore-limbs; hind limb digits absent; finger lamellae absent; FLL 2.2% AGD, 1.7% SVL; HLL 3.1% AGD, 2.5% SVL; tail not as wide as body, gradually tapered towards end, TW 88.9% MBW, TL 101.7% SVL. Rostral projecting onto dorsal snout to level in line with midline of nasal, wider than long, in contact with frontonasal; frontonasal wider than long; nostril ovoid, in anteroventral corner of single trapezoidal nasal, longer axis directed posterodorsally and anteroventrally; supranasals present; postnasals absent; prefrontals moderately separated; frontal roughly diamond-shaped, its anterior margin in moderate contact with frontonasal, in contact with first two anterior supraoculars, 5 × wider than anterior supraocular; supraoculars four; frontoparietals moderate, in narrow contact, each contacts supraoculars two and three; interparietal moderate, its length roughly equal to 1.5× midline length of frontoparietal, longer than wide, diamond-shaped, wider anteriorly, pineal eyespot ovoid, visible in center; parietals broader than frontoparietals, in moderate contact behind interparietal; enlarged nuchals present; loreals two, anterior loreal longer and slightly higher than posterior loreal; preocular one; presubocular one; supraciliaries five, the anteriormost contacting prefrontal and separating posterior loreal from first supraocular, posteriormost extending to midline of fourth supraocular; subocular scale row complete, in contact with supralabials; lower eyelid with one row of scales; supralabials six, first twice the width of others, third, fourth and fifth subocular; infralabials six (Fig. 2). Frontoparietal contact Absent Absent Present Point contact or Absent Absent 1 st chin shield pair contact Absent Absent Present or Absent Absent Present or Absent Mental wider than long, in contact with first infralabials; postmental single, enlarged, its width greater than width of mental; followed by three pairs of enlarged chin shields, first and second pairs moderately separated by single medial scale, second pair largest followed by first pair, third pair smallest, separated by four medial scales (Fig. 2). Scales on limbs smaller than body scales. Variation. All specimens examined in this series match the holotype closely except one individual (KU 307755) that has the first chin shield pair in contact. Coloration of holotype in life. (Fig. 3) The dorsal, and lateral portions of the trunk and tail are solid pink-like Beige (Color 254; Köhler 2012). Just above the orbit, a single splotch of Pratt’s Payne’s Gray (Color 293; Köhler 2012) can be seen on the dorsal and lateral portions of the head, as well as Cinnamon-Drab on the snout (Color 50; Köhler 2012). Coloration of holotype in preservative. The dorsal, lateral, and ventral portions of the trunk and tail are a solid Prout’s Brown (Color 47; Köhler 2012). Just above the orbit, a single splotch of Fuscous (Color 283; Köhler 2012) can be seen on the dorsal and lateral portions of the head. The ventral portion of the head is the same background color as the trunk (Prout’s Brown; Color 47; Köhler 2012). Etymology. The specific epithet is derived from the Tagalog (Filipino) term "nakaligtas," meaning "survivor" and " ligtas," meaning "salvation." We name this species in honor of the people of Lubang Island who endured nearly three decades of violence and guerrilla warfare, from 1945 to 1974, led by the Imperial Japanese Army intelligence officer Hiroo Onoda, and four Japanese soldiers. After being driven into the jungle of Lubang Island by allied forces near the end of World War II, Onoda resisted surrender for 29 years believing the war was not yet over. Onoda would finally surrender in 1974, allowing the communities of Lubang to move on from the hardships faced during this time period, including the loss of over 30 lives and injuries to dozens more. Suggested common name: Lubang Slender Skink. Distribution. Brachymeles ligtas sp. nov. is currently known only from Lubang Island (Fig. 1) and we expect it is found on several smaller nearby landmasses. Natural history. Brachymeles ligtas sp. nov. likely once occurred in low- to mid-elevation primary forest habitats. As most primary forest on Lubang Island has been destroyed, the recent observations of this species have occurred in secondary growth forest habitats. In contrast to the other members of the B. bonitae Complex, this species appears to be relatively common in secondary growth forest fragments on the island. To date, no other congeners have been documented on Lubang Island. We have evaluated this species against the International Union for Conservation of Nature (IUCN) criteria for classification and find that it does not qualify for Critically Endangered, Endangered, Vulnerable, or Near Threatened status. Although B. ligtas sp. nov. is known from a single island only, the species appears relatively common in secondary growth forest on Lubang, and until additional data are presented to support otherwise, we classify this species as Least Concern (LC; IUCN 2015).

Brachymeles ligtas sp. nov. (Figs. 2, 3) Brachymeles bonitae Dum��ril & Bibron 1839; Taylor 1917; Brown 1956: 5; Brown & Rabor 1967: 526; Brown & Alcala 1970; Brown & Alcala 1980: 20; Davis et al. 2014. Holotype. PNM 9818 (CDS Field No. 3886, formerly KU 320472), adult female, collected on 26 April 2009 (14:00 h) in Sitio Dangay, Barangay Vigo, Municipality of Lubang, Occidental Mindoro Province, Lubang Island, Philippines (13.79995�� N, 120.163930�� E; WGS 84; 45 m elev. elevation), by J. Fernandez and CDS. Paratypes (Paratopotypes). Four paratopotypes were also collected in Sitio Dangay, Barangay Vigo, Municipality of Lubang, Occidental Mindoro Province, Lubang Island, Philippines (13.79995�� N, 120.163930�� E; WGS 84; 45 m elev.), by J. Fernandez and CDS. One adult male (KU 320470) was collected on 24 April 2009 at 14:00 h, two adult females (KU 320471, 320473) were collected on 26 April 2009 at 14:00 h, and one juvenile (KU 320474) was collected on 29 April 2009 at 14:00 h. Paratypes. One adult male (KU 307755) was collected on 8 December 2005 in Barangay Vigo, Municipality of Lubang, Occidental Mindoro Province, Lubang Island, Philippines (13.826552�� N, 120.120514�� E; WGS 84; 27 m elev.), by RMB, CDS, and CWL. Diagnosis. Following recent taxonomic revisions of Brachymeles (Siler et al. 2011; Davis et al. 2014) the new species is assigned to the B. bonitae Complex based on the following suite of morphological characters: (1) limbs present, (2) non-pentadactyl, (3) fore-limbs with 0���3 fingers, (4) hind limbs with 0���2 toes, (5) paravertebral scale rows ��� 91, (6) presacral vertebrae 47���53, (7) supraoculars four, (8) enlarged, differentiated nuchals present, (9) longitudinal rows of dark spots around the body absent, and (10) auricular opening absent. Brachymeles ligtas sp. nov. can be distinguished from congeners by the following combination of characters: (1) body size small (SVL 60.7���79.6 mm), (2) fore-limbs bidactyl, (3) hind limbs digitless, (4) limb length short, (5) supralabials six, (6) infralabials six, (7) supraciliaries five, (8) supraoculars four, (9) midbody scale rows 22, (10) axilla���groin scale rows 74���76, (11) paravertebral scale rows 91���93, (12) prefrontal contact absent, (13) frontoparietal contact present, (14) enlarged chin shields in three pairs, (15) nuchals enlarged, (16) auricular opening absent, (17) presacral vertebrae 50, and (18) uniform body color (Tables 1, 2). Comparisons. Brachymeles ligtas sp. nov. can be distinguished from other species in the B. bonitae Complex (B. bonitae, B. isangdaliri, B. mapalanggaon, B. tridactylus), by the number of presacral vertebrae (50 versus 53 [B. bonitae], 51 [B. mapalanggaon], 47 [B. isangdaliri, B. tridactylus]), and by having bidactyl fore-limbs and digitless hind limbs (versus bidactyl fore-limbs and unidactyl hind limbs [B. bonitae], digitless [B. bonitae, B. mapalanggaon], unidactyl [B. isangdaliri], or tridactyl [B. tridactylus]; Table 2); further, from B. bonitae by having a greater number of infralabials (6 versus 5) and absence (versus presence) of a fused mental and first chin shield; from B. isangdaliri by having fewer supraciliaries (5 versus 6) and the presence (versus absence) of a third chin shield pair; from B. mapalanggaon by having a longer fore-limb length (1.2���1.4 mm versus 0.8���1.0) and a longer hind limb length (1.6���2.0 mm versus 1.2���1.6); from B. tridactylus by having a shorter fore-limb length (1.2���1.4 mm versus 1.5���2.5); from B. isangdaliri and B. tridactylus by having a greater number of presacral vertebrae (50 versus 47 [B. isangdaliri, B. tridactylus]) and a shorter hind limb length (1.6���2.0 mm versus 2.2 [B. isangdaliri] or 2.6���3.6 [B. tridactylus]); from B. bonitae and B. mapalanggaon by having fewer presacral vertebrae (50 versus 53 [B. bonitae] or 51 [B. mapalanggaon]), fewer axilla���groin scale rows (74���76 versus 83���90 [B. bonitae] or 80���84 [B. mapalanggaon]); from B. bonitae, B. isangdaliri, and B. mapalanggaon by having fewer paravertebral scale rows (91���93 versus 103���110 [B. bonitae], 97 [B. isangdaliri], or 99���102 [B. mapalanggaon]); from B. bonitae, B. isangdaliri, and B. tridactylus by the presence of frontoparietals in contact (versus not in contact). Finally, Brachymeles ligtas sp. nov. can be distinguished from all limbless species of Brachymeles by having limbs, and from all pentadactyl species of Brachymeles by having bidactyl fore-limbs and digitless hind limbs. Description of holotype. Adult female, body small, slender, SVL 79.6 mm; head weakly differentiated from neck (Fig. 2, 3), nearly as wide as body, HW 5.7% SVL, 96.0% HL; HL 34.2% SnFa; SnFa 17.3% SVL; snout rounded in dorsal and lateral profile, SNL 56.0% HL; ear completely hidden by scales; eyes small, ED 21.3% HL, 50.9% END, pupil subcircular; body slightly depressed, nearly uniform in thickness, MBW 124.7% MBH; scales smooth, glossy, imbricate; longitudinal scale rows at midbody 22; paravertebral scale rows 93; axilla���groin scale rows 76; limbs short, diminutive, bluntly rounded, with digits reduced to two small digit growths on fore-limbs; hind limb digits absent; finger lamellae absent; FLL 2.2% AGD, 1.7% SVL; HLL 3.1% AGD, 2.5% SVL; tail not as wide as body, gradually tapered towards end, TW 88.9% MBW, TL 101.7% SVL. Rostral projecting onto dorsal snout to level in line with midline of nasal, wider than long, in contact with frontonasal; frontonasal wider than long; nostril ovoid, in anteroventral corner of single trapezoidal nasal, longer axis directed posterodorsally and anteroventrally; supranasals present; postnasals absent; prefrontals moderately separated; frontal roughly diamond-shaped, its anterior margin in moderate contact with frontonasal, in contact with first two anterior supraoculars, 5 �� wider than anterior supraocular; supraoculars four; frontoparietals moderate, in narrow contact, each contacts supraoculars two and three; interparietal moderate, its length roughly equal to 1.5�� midline length of frontoparietal, longer than wide, diamond-shaped, wider anteriorly, pineal eyespot ovoid, visible in center; parietals broader than frontoparietals, in moderate contact behind interparietal; enlarged nuchals present; loreals two, anterior loreal longer and slightly higher than posterior loreal; preocular one; presubocular one; supraciliaries five, the anteriormost contacting prefrontal and separating posterior loreal from first supraocular, posteriormost extending to midline of fourth supraocular; subocular scale row complete, in contact with supralabials; lower eyelid with one row of scales; supralabials six, first twice the width of others, third, fourth and fifth subocular; infralabials six (Fig. 2). Frontoparietal contact Absent Absent Present Point contact or Absent Absent 1 st chin shield pair contact Absent Absent Present or Absent Absent Present or Absent Mental wider than long, in contact with first infralabials; postmental single, enlarged, its width greater than width of mental; followed by three pairs of enlarged chin shields, first and second pairs moderately separated by single medial scale, second pair largest followed by first pair, third pair smallest, separated by four medial scales (Fig. 2). Scales on limbs smaller than body scales. Variation. All specimens examined in this series match the holotype closely except one individual (KU 307755) that has the first chin shield pair in contact. Coloration of holotype in life. (Fig. 3) The dorsal, and lateral portions of the trunk and tail are solid pink-like Beige (Color 254; K��hler 2012). Just above the orbit, a single splotch of Pratt���s Payne���s Gray (Color 293; K��hler 2012) can be seen on the dorsal and lateral portions of the head, as well as Cinnamon-Drab on the snout (Color 50; K��hler 2012). Coloration of holotype in preservative. The dorsal, lateral, and ventral portions of the trunk and tail are a solid Prout���s Brown (Color 47; K��hler 2012). Just above the orbit, a single splotch of Fuscous (Color 283; K��hler 2012) can be seen on the dorsal and lateral portions of the head. The ventral portion of the head is the same background color as the trunk (Prout���s Brown; Color 47; K��hler 2012). Etymology. The specific epithet is derived from the Tagalog (Filipino) term "nakaligtas," meaning "survivor" and " ligtas," meaning "salvation." We name this species in honor of the people of Lubang Island who endured nearly three decades of violence and guerrilla warfare, from 1945 to 1974, led by the Imperial Japanese Army intelligence officer Hiroo Onoda, and four Japanese soldiers. After being driven into the jungle of Lubang Island by allied forces near the end of World War II, Onoda resisted surrender for 29 years believing the war was not yet over. Onoda would finally surrender in 1974, allowing the communities of Lubang to move on from the hardships faced during this time period, including the loss of over 30 lives and injuries to dozens more. Suggested common name: Lubang Slender Skink. Distribution. Brachymeles ligtas sp. nov. is currently known only from Lubang Island (Fig. 1) and we expect it is found on several smaller nearby landmasses. Natural history. Brachymeles ligtas sp. nov. likely once occurred in low- to mid-elevation primary forest habitats. As most primary forest on Lubang Island has been destroyed, the recent observations of this species have occurred in secondary growth forest habitats. In contrast to the other members of the B. bonitae Complex, this species appears to be relatively common in secondary growth forest fragments on the island. To date, no other congeners have been documented on Lubang Island. We have evaluated this species against the International Union for Conservation of Nature (IUCN) criteria for classification and find that it does not qualify for Critically Endangered, Endangered, Vulnerable, or Near Threatened status. Although B. ligtas sp. nov. is known from a single island only, the species appears relatively common in secondary growth forest on Lubang, and until additional data are presented to support otherwise, we classify this species as Least Concern (LC; IUCN 2015)., Published as part of Geheber, Aaron D., Davis, Drew R., Watters, Jessa L., Penrod, Michelle L., Feller, Kathryn D., Davey, Conner S., Ellsworth, Elyse D., Flanagan, Rachel L., Heitz, Brendan B., Moore, Tana, Nguyen, Marie D. C., Roberts, Austyn, Sutton, John, Sanguila, Marites B., Linkem, Charles W., Brown, Rafe M. & Siler, Cameron D., 2016, Additions to Philippine Slender Skinks of the Brachymeles bonitae Complex (Reptilia: Squamata: Scincidae) I: a new species from Lubang Island, pp. 1-14 in Zootaxa 4132 (1) on pages 4-8, DOI: 10.11646/zootaxa.4132.1.1, http://zenodo.org/record/271882, {"references":["Taylor, E. H. (1917) Brachymeles, a genus of Philippine lizards. Philippine Journal of Science, 12, 267 - 279.","Brown, W. C. (1956) A revision of the genus Brachymeles (Scincidae), with descriptions of new species and subspecies. Breviora, 54, 1 - 19.","Brown, W. C. & Rabor, D. S. (1967) Review of the genus Bracheymeles (Scincidae), with descriptions of new species and subspecies. Proceedings of the California Academy of Sciences, Series 4, 15, 525 - 548. Avaliable from: http: // researcharchive. calacademy. org / research / herpetology / philippine _ collection / phil _ pdfs / casp _ 34 _ 525. pdf (Accessed 20 Jun. 2016)","Brown, W. C. & Alcala, A. C. (1980) Philippine Lizards of the Family Scincidae. Philippine University Press, Dumaguete City, Philippines, 264 pp.","Kohler, G. (2012) Color Catalog for Field Biologists. Herpeton, Offenbach, 49 pp."]}

Pseudogekko isapa sp. nov. (Fig. 2) Holotype. PNM 9816 (ACD Field No. 8920), adult male, collected at 21:30 hrs in 2014, in Mt. Guiting-Guiting Natural Park, Municipality of Magdiwang, Barangay Talaba, Romblon Province, Sibuyan Island, Philippines (12.401873°N, 122.539747°E; WGS 84; 479 m elev.), by ACD and party. Paratype. One adult female (CAS 139713) collected on 22 May 1972, on a small vine in secondary forest, from Dubduban Barrio, Municipality of San Agustin, Romblon Province, Tablas Island, Philippines (12.58332°N, 122.1°E; WGS 84; ~ 17 m elev.), by L. C. Alcala and party. Diagnosis. Pseudogekko isapa sp. nov. can be distinguished from congeners by the following combination of characters: (1) body elongate (SVL 62.1, 63.4 mm), (2) axilla–groin distance long (32.1, 33.0 mm), (3) relative head length long (19% SVL), (4) relative snout length long (62, 64% head length), (5) Finger-III scansors 13, (6) Toe-IV scansors 17, (7) paravertebrals 240, 246, (8) ventral scales 135, 141, (9) supralabials 20, 21, (10) infralabials 17, 19, (11) circumorbitals 50, 54, (12) precloacal pores 15 in males, none in females, (13) femoral pores absent, (14) conspicuous head spotting present, (15) conspicuous dorsolateral spotting absent, (16) body striping absent, and (17) interorbital banding present (Tables 1, 2; Fig. 2). Comparisons. Characters distinguishing Pseudogekko isapa sp. nov. from all other species of the Pseudogekko compresicorpus Complex sensu Siler et al. (2014b) are summarized in Tables 1, 2. Pseudogekko isapa sp. nov. most closely resembles P. compresicorpus; however, it differs from this species by having a longer snout–vent length (SVL 62.1–63.4 mm versus 54.9–59.7 mm), wider body width (midbody width 8.5–8.9 mm versus 5.4–6.7 mm), greater relative head length (head length 19% SVL versus 16–18%), fewer Finger-III (13 versus 15–17) and Toe-IV (17 versus 18 or 19) scansors, more (17–19 versus 13–16), more circumorbitals (50–54 versus 39–45), more paravertebrals (240–246 versus 226–234), more ventrals (135–141 versus 127–130), and more enlarged precloacal pore-bearing scales (15 versus 10–14; Fig. 2). From the remaining members of the P. compresicorpus Complex, P. isapa sp. nov. can be distinguished from P. pungkaypinit by having a smaller body size (SVL 62.1–63.4 mm versus 66.6–76.8 mm), smaller axilla–groin distance (32.1–33.0 mm versus 37.2–41.2), longer relative snout length (62–64% head length versus 54–60%), fewer Finger-III scansors (13 versus 15–17), fewer paravertebrals (240–246 versus 265–280), fewer enlarged pore-bearing precloacals (15 versus 17–20), and the absence (versus presence) of body striping; and from P. chavacano and P. ditoy by having a larger body size (SVL 62.1–63.4 mm versus 54.7–55.9 [P. chavacano] or 49.4– 52.6 [P. d i t o y]), larger axilla–groin distance (32.1–33.0 mm versus 26.7–30.0 [P. chavacano] or 25.1–29.7 [P. ditoy]), longer relative snout length (62–64% head length versus 55–58% [P. chavacano] or 57–59% [P. ditoy]), fewer Finger-III scansors (13 versus 15 or 16 [P. chavacano] or 14 or 15 [P. d i t oy]) and enlarged pore-bearing precloacals (15 versus 16 [P. chavacano] or 18 [P. ditoy]), and more circumorbitals (50–54 versus 46 [P. chavacano] or 40–43 [P. d i t oy]), paravertebrals (240–246 versus 195–197 [P. chavacano] or 180–185 [P. ditoy]), and ventrals (135–141 versus 122 or 123 [P. chavacano] or 111–118 [P. ditoy]; Tables 1, 2). Pseudogekko isapa sp. nov. can be distinguished from members of the P. brevipes Complex (P. brevipes, P. atiorum), by having a larger body size (SVL 62.1–63.4 mm versus 41.1–52.5 [P. a t i o r u m] or 34.5–42.4 [P. brevipes]), more supralabials (20 or 21 versus 15–17 [P. a ti o ru m] or 13 or 14 [P. brevipes]), infralabials (17–19 versus 12–15 [P. a t io ru m] or 14 or 15 [P. brevipes]), circumorbitals (50–54 versus 35–38 [P. atiorum] or 33–35 [P. brevipes]), and ventrals (135–141 versus 119–129 [P. atiorum] or 96–117 [P. brevipes]). From P. smaragdinus, P. isapa sp. nov. can be distinguished by having fewer Finger-III scansors (13 versus 15– 18) and lacking pore-bearing femorals (15 versus 32–41), more supralabials (20 or 21 versus 16–19), circumorbitals (50–54 versus 33–35), and ventrals scales (135–141 versus 124–130), a tendency towards having fewer Toe-IV scansors (17 versus 16–22), a tendancy towards having more infralabials (17 or 19 versus 14–17), and by the presence of brownish tan (versus yellow to orange [undisturbed] or neon green [disturbed]) body coloration, and absence (versus presence) of enlarged femoral pore-bearing scales in males. Description of holotype. Adult male, well preserved; small incision in the sternal region (portion of liver removed for genetic a sample). Body moderately large, elongate, slender; limbs well developed, moderately slender; tail autotomized, absent; margins of limbs smooth, cutaneous flaps or dermal folds absent. Head size moderate, differentiated from neck, characterized by only slightly hypertrophied temporal and adductor musculature; snout broadly rounded in dorsal view and sharply rounded in lateral view; head width 118.8% midbody width, 85.6% head length; head length 19.0% SVL; snout length 74.2% head width, 63.6% head length; dorsal surfaces of head relatively homogeneous, with slightly to moderately pronounced concave postnasal, internasal, prefrontal, and interorbital concavities; auricular opening small, teardrop-shaped, beneath temporal swellings on either side of head; tympanum deeply sunken; orbit large; eye large, pupil vertical, margin wavy; limbs and digits relatively short; thighs moderately thicker compared to brachium; tibia length 8.2% SVL, 53.1% femur length. Rostral size moderately large, margin oval in anterodorsal view, nearly as broad as high, sutured anterolaterally with anteriormost enlarged supralabials, projecting onto dorsal surface of head to point in line anteriormost edge of nasal; nasal surrounded by first labial, rostral, one enlarged postnasal, and two enlarged supranasals; medial supranasals separated by three moderately enlarged median scales; enlarged supranasals larger than postnasals, anteriormost supranasal largest. Differentiated supralabials bordered dorsally by one row of slightly enlarged snout scales; differentiated infralabials bordered ventrally by undifferentiated scales or one row of slightly enlarged scales around the lower jaw; mentals and postrictals undifferentiated; gulars small, round, nonimbricate, juxtaposed; dorsal cephalic scales fairly homogeneous in size, shape, disposition, and distribution; cephalic scalation slightly convex, round to oval scales; posterior head scales granular, slightly raised into flat, plateaued dorsal surfaces. Axilla–groin distance 50.6% SVL; dorsal body scales round, slightly raised into flat, dorsal plateaued, juxtaposed, relatively homogeneous in size; each dorsal scale surrounded by 4–6 interstitial granules, many forming a “Star of David” pattern around dorsal scale (Siler et al. 2014b: Fig. 5); dorsals gradually transition to imbricate ventrals along lateral body surface; scales on dorsal surfaces of limbs more imbricate than dorsals; scales on dorsal surfaces of hands and feet similar to dorsal limb scales, heavily imbricate; ventral body scales flat, cycloid, strongly imbricate, much larger than lateral or dorsal body scales, relatively homogeneous in size. Enlarged pore-bearing scales in continuous precloacal series, arranged in a widely obtuse, “V”-formation (Fig. 2); patch of slightly enlarged scales posterior to precloacal series, roughly 6 or 7 scale rows in width, 3 or 4 scale rows in length, forming an oval patch just posterior to precloacal series (Fig. 2). Digits moderately expanded and covered on palmar and plantar surfaces by bowed, unnotched, undivided scansors (Fig. 2); digits with minute vestiges of interdigital webbing on hands, moderate interdigital webbing on feet; subdigital scansors of fingers (left/right) I (10,9), II (11,11), III (13,13), IV (18,17), V (11,11); subdigital scansors of toes (left/right) I (10,9), II (12,11), III (16,17), IV (17,17), V (11,11); subdigital scansors of hands and feet bordered proximally (on palmar and plantar surfaces) by 1–3 slightly enlarged scales that form a nearcontinuous series with enlarged scansors; all digits clawed, but first (inner) claw greatly reduced; remaining terminal claw-bearing phalanges compressed, with moderately sized recurved claws. Coloration of holotype in preservative. Background dorsal trunk coloration light brownish tan with intermittent to dense small dark brown speckles, continuing down to base of tail; dorsal region of head with similar color pattern, except for pronounced dark brown area above orbits; dorsal and lateral regions of limbs with same color pattern as trunk; supralabials and infralabials with similar pattern as head, but dark brown spots more diffuse; ventral side of trunk with same background coloration, dark brown speckling pattern significantly reduced; ventral side of head with sparse dark brown speckling only; ventral surfaces of the limbs, and palms of hands and feet lighter cream colored with medium to dark brown speckles scattered sparsely throughout; ventral surfaces of digits darker tan to grey with irregular, sparse dark brown speckles; skin between scansors dark brown. Coloration in life unrecorded. Measurements and scale counts of holotype in mm. SVL 63.4; axilla–groin distance 32.1; total length N/A; tail length N/A; head length 11.8; head width 10.1; head depth 7.0; eye diameter 3.9; snout length 7.5; eye–nares distance 5.9; internarial distance 2.6; interorbital distance 7.0; midbody width 8.5; femur length 9.8; tibia length 5.2; Toe-I length 2.3; Toe-IV length 4.1; tail width 5.3; tail depth 4.7; supralabials 20; infralabials 19; circumorbitals 50; paravertebral scales 240; ventral scales 135; Finger-III scansors 13; Toe-IV scansors 17. Distribution, ecology and natural history. Pseudogekko isapa sp. nov. is known from the islands of Tablas and Sibuyan in the Romblon Island Group (Fig. 1). This species also may occur on Romblon Island itself, and eventually may be found on several of the smaller islands scattered across the province, such as Banton and Simara islands just off of the north coast of Tablas Island. Besides P. atiorum, P. isapa sp. nov. is the only other species outside of the Luzon and Mindanao PAICs; all other species are distributed across the northern, eastern, and southern islands (Siler et al. 2014a, b; Davis et al. 2015a). Like most members of the genus, P. isapa sp. nov. appears to be primarily a forest obligate species; the holotype was collected within high quality secondary forest in Mt. Guiting- Guiting Natural Park on Sibuyan Island. This species’ poor representation in museum collections globally is a testament to the secretive nature of these species in their native habitat (Siler et al. 2014a, b). Due to the absence of data allowing for confirmation of potential declines in population size as a result of habitat destruction, and the lack of available information about the species’ natural history, we do not find that it qualifies for formal threatened status under IUCN criteria (IUCN 2015). Therefore, we recommend that the species be considered Data Deficient at the current time, but strongly recommend that immediate and comprehensive survey efforts be made to study the health of wild populations on islands throughout the Romblon Island Group (Fig. 1). With at least a very large portion of its range dominated by Mt. Guiting-Guiting Natural Park, the majority of its hypothesized distribution is very well protected. Etymology. We derive the new species epithet from the Tagalog (Filipino) phrase isa pa, meaning “another” or “one more” in reference to our surprise at the discovery of yet another distinctive new species of Pseudogekko immediately following our review of the genus (Siler et al. 2014a), including what we thought was a comprehensive review of the P. compresicorpus (Siler et al. 2014b) and P. brevipes complexes (Davis et al. 2015a). Suggested common name: Romblon Province False Gecko. isapa sp. nov. (1 m, 1 f) SVL 63.4, 62.1 Axilla–groin distance 32.1, 33.0 Total length – Midbody width 8.5, 9.9 Head length 11.8, 11.7 Head width 10.1, 9.4 Head height 7.0, 7.2 Snout length 7.5, 7.2 Eye diameter 3.9, 4.0 Eye–nares distance 5.9, 6.0 Internarial distance 2.6, 2.4 Interorbital distance 7.0, 6.7 Femur length 9.8, 10.1 Tibia length 5.2, 5.5

Davis, Drew R., Watters, Jessa L., Köhler, Gunther, Whitsett, Collin, Huron, Nicholas A., Brown, Rafe M., Diesmos, Arvin C., Siler, Cameron D. (2015): Redescription of the rare Philippine false gecko Pseudogekko brevipes (Reptilia: Squamata: Gekkonidae) and description of a new species. Zootaxa 4020 (2): 357-374, DOI: http://dx.doi.org/10.11646/zootaxa.4020.2.7

Pseudogekko brevipes (Boettger, 1897) (Figs. 1���3) Lepidodactylus brevipes Boettger, 1897: 161, holotype male (Senckenberg Research Institute and Natural History Museum [SMF] 8988) from "Samar Island"; Taylor, 1922: 74; Brown, 1964: 6; Wermuth, 1965: 97; Kluge, 1967: 29; Kluge, 1968: 332. Pseudogekko brevipes (Boettger): Kluge, 1968: 332; Brown & Alcala, 1978: 118; Kluge, 1993: 30; R��sler, 2000: 106; Gaulke et al., 2003: 111; Siler et al., 2014 c. Diagnosis. Pseudogekko brevipes can be distinguished from congeners by the following combination of characters: (1) body size moderate (SVL 34.5���42.4 mm); (2) axilla���groin distance moderate (17.8���29.8 mm); (3) head length moderate (6.2���9.5 mm); (4) snout length long (3.8���4.4 mm); (5) Toe-IV scansors 15; (6) paravertebrals 211���218; (7) ventrals 96���117; (8) supralabials 13 or 14; (9) infralabials 14 or 15; (10) circumorbitals 33���35; (11) precloacal pores 12; (12) femoral pores absent; (13) conspicuous dorsolateral spotting present; (14) limb spotting absent; (15) tail banding absent; (16) body striping absent; (17) interorbital banding present (Tables 1, 2; Figs. 2, 3). Comparisons. Characters distinguishing Pseudogekko brevipes from all other species of Pseudogekko are summarized in Tables 1���3. Pseudogekko brevipes most closely resembles the new species described below; however, it differs from this species by having a shorter snout���vent length in both males (39.0 mm vs. 41.1���52.5) and females (34.5���42.4 mm vs. 44.8���48.7), shorter eye���nares distance (3.0��� 3.4 mm vs. 3.5���4.5), a tendency towards having both a shorter snout length (3.8���4.4 mm vs. 4.3���5.8) and internares distance (1.2���1.6 mm vs. 1.5��� 1.9), fewer supralabials (13 or 14 vs. 15���17), circumorbitals (33���35 vs. 35���38), paravertebrals (211���218 vs. 226��� 240), ventrals (96���117 vs. 119���129), and precloacal pores (12 vs. 13���15; Fig. 3). Pseudogekko brevipes can be distinguished from P. chavacano by having a shorter snout���vent length (34.5��� 42.4 mm vs. 54.7, 55.9), total length (72.0��� 87.5 mm vs. 95.8), midbody width (4.5���5.8 mm vs. 6.4, 6.7), head length (6.2���9.5 mm vs. 10.4), head width (5.6���6.5 mm vs. 7.8, 8.5), and snout length (3.8���4.4 vs. 5.8, 6.0), fewer Finger-III scansors (12 vs. 15, 16), Toe-IV scansors (15 vs. 17���20), supralabials (13, 14 vs. 15, 16), infralabials (14, 15 vs. 16, 17), circumorbitals (33���35 vs. 46), ventrals (96���117 vs. 122, 123), and precloacal pores (12 vs. 16), greater number of paravertebrals (211���218 vs. 195���197), absence (vs. presence) of limb spotting and tail banding, and presence (vs. absence) of interorbital banding; from P. compresicorpus by having a shorter snout���vent length (34.5���42.4 mm vs. 54.9���59.7), total length (72.0��� 87.5 mm vs. 105.9���117.3), head width (5.6���6.5 mm vs. 7.5��� 10.1), and snout length (3.8���4.4 mm vs. 5.3���7.4), fewer Finger-III scansors (12 vs. 15���17), Toe-IV scansors (15 vs. 18 or 19), supralabials (13 or 14 vs. 16���20), circumorbitals (33���35 vs. 39���45), and ventrals (96���117 vs. 127���130), absence (vs. presence) of limb spotting, and presence (vs. absence) of interorbital banding; from P. d i t oy by having a shorter snout���vent length (34.5���42.4 mm vs. 49.4���52.6), midbody width (4.5���5.8 mm vs. 6.3���7.3), head width (5.6���6.5 mm vs. 7.7���7.9), and snout length (3.8���4.4 mm vs. 5.4���5.7), fewer Finger-III scansors (12 vs. 14 or 15), Toe-IV scansors (15 vs. 16 or 17), supralabials (13 or 14 vs. 17���20), infralabials (14 or 15 vs. 16 or 17), circumorbitals (33���35 vs. 40���43), and precloacal pores (12 vs. 18), greater number of paravertebrals (211���218 vs. 180���185), presence (vs. absence) of dorsolateral spotting and interorbital banding; from P. pungkaypinit by having a shorter snout���vent length (34.5���42.4 mm vs. 66.6���76.8), axilla���groin distance (17.8���29.8 mm vs. 37.2���41.2), total length (72.0��� 87.5 mm vs. 125.3���141.2), midbody width (4.5���5.8 mm vs. 7.7���9.1), head length (6.2���9.5 mm vs. 11.4���13.6), head width (5.6���6.5 mm vs. 9.3���11.2), and snout length (3.8���4.4 mm vs. 6.7���7.5), fewer Finger-III scansors (12 vs. 15���17), Toe-IV scansors (15 vs. 17���21), supralabials (13 or 14 vs. 16���20), infralabials (14 or 15 vs. 17���19), circumorbitals (33���35 vs. 50���55), paravertebrals (211���218 vs. 265���280), ventrals (96���117 vs. 125��� 155), and precloacal pores (12 vs. 17���20), presence (vs. absence) of dorsolateral spotting and interorbital banding, and absence (vs. presence) of body striping; from P. smaragdinus by having a shorter snout���vent length (34.5���42.4 mm vs. 50.2���64.3), total length (72.0��� 87.5 mm vs. 103.6���129.7), midbody width (4.5���5.8 mm vs. 6.2���10.4), head width (5.6���6.5 mm vs. 7.1���9.8), and snout length (3.8���4.4 mm vs. 4.9���6.5), fewer Finger-III scansors (12 vs. 15��� 18), Toe-IV scansors (15 vs. 16���22), supralabials (13, 14 vs. 16���19), ventrals (96���117 vs. 124���130), and precloacal pores (12 vs. 32���41), absence (vs. presence) of femoral pores, conspicuous limb spotting, and tail banding, and presence (vs. absence) of interorbital banding. Redescription (based on the holotype [Boettger, 1897] and three referred specimens). Measurements and scale counts of the holotype are provided below in brackets. Body small, slender, SVL 39.0 mm (male) [39.0], 34.5���42.4 mm (female); limbs well developed, moderately slender; tail slender; margins of limbs smooth, lacking cutaneous flaps or dermal folds; trunk lacking ventrolateral cutaneous fold. Head size moderate, slightly differentiated from neck, characterized by only slightly hypertrophied temporal and adductor musculature; snout rounded in dorsal and lateral aspect; HW 113.2 ���128.0% MBW [115.6 %], 58.2��� 93.7 % HL [58.2 %]; HL 16.7���24.4 % SVL [24.4 %]; SNL 65.5���76.2 % HW [76.2 %], 44.4���62.3 % HL [56.5 %]; dorsal surfaces of head relatively homogeneous, with only slightly pronounced concave postnasal, internasal, prefrontal, and interorbital concavities; auricular opening small, ovoid, angled slightly anteroventrally and posterodorsally from beneath temporal swellings on either side of head; tympanum deeply sunken; orbit large; eye large, pupil vertical, margin wavy; limbs and digits relatively short; thighs moderately thicker compared to upper arms; tibia length 9.5���13.5 % SVL [13.5 %], 61.9���81.8 % femur length [81.8 %]. Rostral small, margin in wide heart shape in anterodorsal view, nearly as broad as high, sutured anterolaterally with anteriormost enlarged supralabials, projecting onto dorsal surface of head to point in line with anteriormost edge of nasal; nasal surrounded by first labial, rostral, one enlarged postnasal, and two enlarged supranasals; anteriormost supranasals separated by two or three small median scales; enlarged supranasals moderately larger than postnasals; circumorbitals 33���35 [35]. Total number of differentiated supralabials 13 or 14 [13], bordered dorsally by one row of differentiated, slightly enlarged snout scales; total number of differentiated infralabials 14 or 15 [14], bordered ventrally by two or three rows of slightly enlarged scales; chin scales slightly enlarged; postrictal scales undifferentiated; gulars small, round, nonimbricate, juxtaposed. Dorsal cephalic scales fairly homogeneous in size, shape, disposition, and distribution; cephalic scalation slightly convex, round to oval scales; postnasal, prefrontal, internasal, and interorbital depressions; undifferentiated posterior head scales granular, slightly convex. Axilla���groin distance 48.9���70.2 % SVL [55.6 %]; undifferentiated dorsal body scales roughly round, moderately convex, juxtaposed, relatively homogeneous in size, some with slightly raised edge, irregular in placement on individual scales across body; dorsals gradually transition to imbricate ventrals along lateral body surface; paravertebral scales between midpoints of limb insertions 211���218 [218]; ventrals between midpoints of limb insertions 96���117 [117]; scales on dorsal surfaces of limbs more imbricate than body dorsal scales; scales on dorsal surfaces of hands and feet similar to dorsal limb scales, heavily imbricate; ventral body scales flat, cycloid, strongly imbricate, much larger than lateral or dorsal body scales, relatively homogeneous in size. Twelve enlarged scales in continuous precloacal series (pore-bearing series in males), arranged in a widely obtuse, ���V���-formation; patch of slightly enlarged scales posterior to precloacal series, roughly 4���6 scale rows in width, three or four scale rows in length, forming an oval patch just posterior to precloacal series. Digits moderately expanded and covered on palmar and plantar surfaces by bowed, unnotched, undivided scansors; digits with minute vestiges of interdigital webbing; mite eggs visible between distal subdigital scansors of some digits on hands and feet on all specimens examined; subdigital scansors of hands and feet bordered proximally (on palmar and plantar surfaces) by two or three slightly enlarged scales that form a near-continuous series with enlarged scansors; all digits clawed, but first (inner) claw greatly reduced; remaining terminal clawbearing phalanges compressed, with moderately sized recurved claws. Tail relatively long compared to body, TL 33.0��� 45.1 mm [33.0], 84.6���108.6 % SVL [84.6 %]; round, not heavily depressed; TH 83.2���100 % TW [86.4 %]; caudals similar in size to dorsals, subcaudals similar in size to ventral scales. Coloration in life. ���Above gray-brown, with darker clouded markings and on either side a dorsolateral row of about ten small, round white dots; lips and underside of body dirty white or with brownish markings or fine brown dots.��� (Taylor, 1922, p. 75). Distribution, ecology and natural history. Pseudogekko brevipes is known only from a handful of specimens from the eastern Visayan islands of Bohol, Leyte, and Samar. Unlike the new species described below, P. brevipes possesses a distribution that may overlap with those of several other species in the genus (P. ditoy and P. pungkaypinit) in the eastern Philippines (Siler et al., 2014 b, c). As far as modern observations are concerned, only a single genetic record of this rare false gecko has been reported from the Municipality of Sogud on southern Leyte Island (Siler et al., 2014 b), and the holotype (SMF 8988) represents the only specimen ever recorded on Samar Island. Over the last decade, we have conducted several major biodiversity survey expeditions to multiple sites on Bohol, Leyte, and Samar islands; however, with the exception of the single genetic sample, we have yet to observe wild populations of real P. brevipes. Although at present there exists a general paucity of information about the ecology and natural history of P. brevipes, we believe this species to be an obligate primary forest dweller, and due to threats of habitat loss we expect that this species will likely qualify for a threatened category in the near future. Therefore, we recommend classification of P. brevipes as Vulnerable, based on the following criteria: VU B 1 ab(iii); D 2 (IUCN, 2013). In addition, we recommend that continued biodiversity surveys of Bohol, Leyte, and Samar islands be conducted to increase our understanding of this species��� distribution and instraspecific diversity., Published as part of Davis, Drew R., Watters, Jessa L., K��hler, Gunther, Whitsett, Collin, Huron, Nicholas A., Brown, Rafe M., Diesmos, Arvin C. & Siler, Cameron D., 2015, Redescription of the rare Philippine false gecko Pseudogekko brevipes (Reptilia: Squamata: Gekkonidae) and description of a new species, pp. 357-374 in Zootaxa 4020 (2) on pages 359-363, DOI: 10.11646/zootaxa.4020.2.7, http://zenodo.org/record/242048, {"references":["Boettger, O. (1897) Neue Reptilien und Batrachier von den Philippinen. Zoologischer Anzeiger, 20, 161 - 166.","Taylor, E. H. (1922) The lizards of the Philippine Islands. Philippine Bureau of Science, 17, 1 - 269.","Brown, W. C. (1964) The status of Pseudogekko shebae and observations on the geckos of the Solomon Islands. Brevoria, 204, 1 - 8.","Wermuth, H. (1965) Liste der rezenten amphibien and reptilien: Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich, 80, 1 - 246.","Kluge, A. G. (1967) Higher taxonomic categories of gekkonid lizards and their evolution. Bulletin of the American Museum of Natural History, 135, 1 - 60.","Kluge, A. G. (1968) Phylogenetic relationships of the gekkonid lizard genera Lepidodactylus Litzinger, Hemiphyllodactylus Bleeker, and Pseudogekko Taylor. The Philippine Journal of Science, 95, 331 - 352.","Brown, W. C. & Alcala, A. C. (1978) Philippine Lizards of the Family Gekkonidae. Silliman University Press, Damaguete City, 146 pp.","Kluge, A. G. (1993) Gekkonoid Lizard Taxonomy. International Gecko Society, San Diego, 245 pp.","Rosler, H. (2000) Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28 - 153.","Gaulke, M., Demegillo, A., Reiter, J. & Tacud, B. (2003) Additions to the herpetofauna of Panay Island, Philippines. Salamandra, 39, 111 - 122.","Siler, C. D., Welton, L. J., Davis, D. R., Watters, J. L., Davey, C. S., Diesmos, A. C., Diesmos, M. L. & Brown, R. M. (2014 c) Taxonomic revision of the Pseudogekko compresicorpus Complex (Reptilia: Squamata: Gekkonidae), with descriptions of three new species. Herpetological Monographs, 28, 110 - 139. http: // dx. doi. org / 10.1655 / herpmonographs-d- 14 - 00005","Siler, C. D., Dececchi, T. A., Merkord, C. L., Davis, D. R., Christiani, T. J. & Brown, R. M. (2014 b) Cryptic diversity and population genetic structure in the rare, endemic, forest-obligate, slender geckos of the Philippines. Molecular Phylogenetics and Evolution, 70, 204 - 209. http: // dx. doi. org / 10.1016 / j. ympev. 2013.09.014"]}

Pseudogekko atiorum sp. nov. (Figs. 1, 3��� 6) Holotype. PNM 9518 (CDS Field No. 954, formerly KU 302818), adult female, collected at 22:00 h on 21 February 2005, on Mt. Talinis, Barangay Bongbong, Municipality of Valencia, Negros Oriental Province, Negros Island, Philippines (N: 9.26000��, E: 123.20327 ��; WGS 84; 1100 m elev.), by CDS. Paratypes. Two adult females (CAS-SUR 19372, 21122) collected from Municipality of Cauayan, Negros Occidental Province, Negros Island, Philippines by A.C. Alcala and Q. Alcala. One adult female (CAS 138097) collected by Q. Alcala and two adult males (CAS 145793, 147491) collected by Q. Alcala and M. Pinero, respectively, from Municipality of Pamplona, Negros Oriental Province, Negros Island, Philippines. One adult female (CAS 134269) collected by A.C. Alcala and one adult male (CAS-SUR 26778) collected by D. Empeso from Municipality of Siaton, Negros Oriental Province, Negros Island, Philippines. Four adult males (CAS 128956, 128959, 128963, 128971) collected from the Municipality of Sibulan, Negros Oriental Province, Negros Island, Philippines by L.C. Alcala. Two adult females (CAS 134292, TNHC 62478) collected by A.C Alcala and RMB, respectively, and one juvenile female (KU 327770) collected by RMB from Municipality of Valencia, Barangay Bongbong, Negros Oriental Province, Negros Island, Philippines. One adult male (CAS 145710) collected from Municipality of San Juan, Siquijor Province, Negros Island, Philippines by L.C. Alcala. Other Referred Specimens. Five specimens (CAS 89812, 147523, CAS-SUR 24588���24590) collected from Municipality of Cauayan, Negros Occidental Province, Negros Island, Philippines. Three specimens (CAS 134298, 137789, CAS-SUR 26776) collected from Negros Occidental Province, Negros Island, Philippines. Twenty specimens (CAS 137653, 137672, 137790, 138096, 138134, 139258, 145791, 145792, 145804 ���6, 145875, 145876, 145976, 173760, 185991, 200539, 200540, CAS-SUR 24323, 24829) collected from Municipality of Pamplona, Negros Oriental Province, Negros Island, Philippines. One specimen (CAS-SUR 26775) collected from Municipality of Siaton, Negros Oriental Province, Negros Island, Philippines. Thirty specimens (CAS 128955, 128957, 128958, 128960���62, 128964���70, 128972���78, 131796, 132609, 142039��� 46) collected from Municipality of Sibulan, Negros Oriental Province, Negros Island, Philippines. Thirteen specimens (CAS 133055, 137791, 200538, CAS-SUR 21803, 21839, 21840, 21856, 21857, 21872, 22218 ���20, 24385) collected from Negros Oriental Province, Negros Island, Philippines. Diagnosis. Pseudogekko atiorum can be distinguished from congeners by the following combination of characters: (1) body size moderate (SVL 41.1���52.5 mm); (2) axilla���groin distance moderate (19.4���28.4 mm); (3) head length moderate (7.4���9.4 mm); (4) snout length long (4.3���5.8 mm); (5) Toe-IV scansors 14���17; (6) paravertebral scales 226���240; (7) ventral scales 119���129; (8) supralabials 15���17; (9) infralabials 12���15; (10) circumorbitals 35���38; (11) precloacal pores 13���15; (12) femoral pores absent; (13) dominant body coloration dark brown; (14) conspicuous head spotting present, cream; (15) conspicuous dorsolateral spotting present, cream; (16) conspicuous limb spotting absent; (17) tail banding absent; (18) body striping absent; (19) interorbital banding present, light brown; (20) iris ring coloration absent (Tables 1, 2; Figs. 3���6). Comparisons. Characters distinguishing Pseudogekko atiorum from all other species of Pseudogekko are summarized in Tables 1���3. Pseudogekko atiorum most closely resembles P. b rev i pe s; however, it differs from this species by having a longer snout���vent length in both males (41.1���52.5 mm vs. 39.0) and females (44.8���48.7 mm vs. 34.5���42.4), longer eye���nares distance (3.5���4.5 mm vs. 3.0��� 3.4), a tendency towards having both a longer snout length (4.3���5.8 mm vs. 3.8���4.4) and internares distance (1.5���1.9 mm vs. 1.2���1.6), and a greater number of supralabials (15���17 vs. 13 or 14), circumorbitals (35���38 vs. 33���35), paravertebrals (226���240 vs. 211���218), ventrals (119���129 vs. 96���117), and precloacal pores (13���15 vs. 12; Figs. 3���5). Pseudogekko atiorum can be distinguished from P. chavacano by having a shorter snout���vent length (41.1��� 52.5 mm vs. 54.7���55.9), longer head length (7.4���9.4 mm vs. 10.4), a tendency towards a shorter snout length (4.3��� 5.8 mm vs. 5.8 ���6.0), fewer infralabials (12���15 vs. 16 or 17), circumorbitals (35���38 vs. 46), and precloacal pores (13���15 vs. 16), greater number of paravertebrals (226���240 vs. 195���197), cream colored (vs. neon green) head and dorsolateral spotting, absence (vs. presence) of limb spotting and tail banding, and presence of interorbital banding (vs. absence); from P. compresicorpus by having a shorter snout���vent length (41.1���52.5 mm vs. 54.9���59.7), a tendency towards both a shorter axilla���groin distance (19.4���28.4 mm vs. 27.0��� 32.6) and head length (7.4���9.4 mm vs. 9.2���11.3), fewer Toe-IV scansors (14���17 vs. 18 or 19) and circumorbitals (35���38 vs. 39���45), cream colored (vs. neon green) head and dorsolateral spotting, absence (vs. presence) of limb spotting and a light blue iris ring, and presence (vs. absence) of interorbital banding; from P. di t o y by having fewer infralabials (12���15 vs. 16 or 17) and circumorbitals (35���38 vs. 40���43), greater number of paravertebrals (226���240 vs. 180���185) and ventrals (119���129 vs. 111���118), fewer precloacal pores (13���15 vs. 18), and the presence (vs. absence) of head spotting, dorsolateral spotting, and interorbital banding; from P. pungkaypinit by having a shorter snout���vent length (41.1���52.5 mm vs. 66.6���75.3), axilla���groin distance (19.4���28.4 mm vs. 37.2���41.2), total length (80.2���113.5 mm vs. 125.3���141.2), midbody width (4.8���7.7 mm vs. 7.7���9.1), head length (7.4���9.4 mm vs. 11.4���13.6), head width (5.9���8.3 mm vs. 9.3���11.2), and snout length (4.3���5.8 mm vs. 6.7���7.5), fewer Finger-III scansors (12���14 vs. 15���17), infralabials (12���15 vs. 17���19), circumorbitals (35���38 vs. 50���55), paravertebrals (226���240 vs. 265���280), and precloacal pores (13���15 vs. 17���20), a tendency towards having fewer ventrals (119���129 vs. 125���155), the presence (vs. absence) of head and dorsolateral spotting and interorbital banding, and the absence (vs. presence) of body striping; from P. smaragdinus by a tendency towards having a shorter snout���vent length (41.1���52.5 mm vs. 50.2���64.3) and axilla��� groin distance (19.4���28.4 mm vs. 26.2���35.5), fewer Finger-III scansors (12���14 vs. 15���18), paravertebrals (226��� 240 vs. 241���252), and enlarged pores (13���15 vs. 32���41), a tendency towards having fewer Toe-IV scansors (14���17 vs. 16���22) and supralabials (15���17 vs. 16���19), greater number of circumorbitals (35���38 vs. 33���35), presence of dark brown (vs. yellow to orange [undisturbed] or neon green [disturbed]) body coloration, cream colored (vs. large black and small white) head and dorsolateral spotting, presence (vs. absence) of interorbital banding, and absence (vs. presence) of limb spotting and tail banding. Description of holotype. Adult female in excellent condition; small incision in the sternal region (portion of liver removed for genetic sample). Body small, slender, SVL 44.8 mm; limbs well developed, moderately slender; tail slender; margins of limbs smooth, cutaneous flaps or dermal folds absent. Head size moderate, slightly differentiated from neck, characterized by only slightly hypertrophied temporal and adductor musculature; snout broadly rounded in dorsal view and bluntly rounded in lateral view; HW 96.9 % MBW, 83.6 % HL; HL 18.4 % SVL; SNL 70.7 % HW, 59.2 % HL; dorsal surfaces of head relatively homogeneous, with slightly to moderately pronounced concave postnasal, internasal, prefrontal, and interorbital concavities; auricular opening small, nearly round, beneath temporal swellings on either side of head; tympanum deeply sunken; orbit large; eye large, pupil vertical, margin wavy; limbs and digits relatively short; thighs moderately thicker compared to upper arms; TBL 8.0% SVL, 57.9 % FL. Rostral size moderately large, margin in wide heart shape in anterodorsal view, nearly as broad as high, sutured anterolaterally with anteriormost enlarged supralabials, projecting onto dorsal surface of head to point in line with anteriormost edge of nasal; nasal surrounded by first labial, rostral, one enlarged postnasal, and two enlarged supranasals; medial supranasals separated by 3 small median scales; enlarged supranasals moderately larger than postnasals; circumorbitals 36 in total. Total number of differentiated supralabials 15, bordered dorsally by one row of differentiated, slightly enlarged snout scales; total number of differentiated infralabials 12, bordered ventrally by 2 or 3 rows of slightly enlarged scales; chin scales enlarged; postrictal scales undifferentiated; gulars small, round, nonimbricate. Dorsal cephalic scales fairly homogeneous in size, shape, disposition, and distribution; cephalic scalation slightly convex, round to oval scales; postnasal, prefrontal, internasal, and interorbital depressions; undifferentiated posterior head scales granular, slightly convex. Axilla���groin distance 55.1 % SVL; undifferentiated dorsal body scales round to tubular, moderately to sharply convex, juxtaposed, relatively homogeneous in size, many with raised edge, irregular in placement on individual scales across body; each dorsal scale surrounded by 4���6 interstitial granules; dorsals gradually transition to imbricate ventrals along lateral body surface; paravertebrals between midpoints of limb insertions 232; ventrals between midpoints of limb insertions 128; scales on dorsal surfaces of limbs more imbricate than dorsals; scales on dorsal surfaces of hands and feet similar to dorsal limb scales, heavily imbricate; ventral body scales flat, cycloid, strongly imbricate, much larger than lateral or dorsal body scales, relatively homogeneous in size. Fourteen enlarged scales in continuous precloacal series (pore-bearing series in males), arranged in a widely obtuse, ���V���-formation; patch of slightly enlarged scales posterior to precloacal series, roughly three scale rows in width, 3���4 scale rows in length, forming an oval patch just posterior to precloacal series. Digits moderately expanded and covered on palmar and plantar surfaces by bowed, unnotched, undivided scansors; digits with minute vestiges of interdigital webbing; subdigital scansors of fingers (left/right) I (9,8), II (10,10), III (13,13), IV (14,14), V (9,9); subdigital scansors of toes (left/right) I (9,9), II (10,10), III (13,13), IV (15,15), V (12,12); mite eggs visible between distal subdigital scansors of digits on hands and feet; subdigital scansors of hands and feet bordered proximally (on palmar and plantar surfaces) by two or three slightly enlarged scales that form a near-continuous series with enlarged scansors; all digits clawed, but first (inner) claw greatly reduced; remaining terminal claw-bearing phalanges compressed, with moderately sized recurved claws. brevipes atiorum chavacano compresicorpus ditoy pungkaypinit smaragdinus sp. A (1 m, 3 f) (8 m, 7 f) (1 m, 1 f) (3 m, 5 f) (1 m, 1 f) (4 m, 2 f) (16 m, 17 f) (1 m, 1 f) individuals showing specific counts are given in parentheses. Characters summarized for adult specimens only. When coloration in life has not been documented, characters are listed as unkown. Enlarged pore series 12 (precloacal) 13���15 16 (precloacal) 10���14 (precloacal) 18 (precloacal) 17���20 (precloacal) 32���41 15 (precloacal) (precloacofemoral) (precloacal) Femoral pores ��� ��� ��� ��� ��� ��� + ��� ������continued on the next page Tail relatively long compared to body, TL 44.6 mm, 99.4 % SVL; round, not heavily depressed; TH 87.8 % TW; caudals similar in size to dorsals, subcaudals similar in size to ventrals. Coloration of holotype in preservative. Background dorsal trunk coloration light tan with intermittent small cream and dark brown speckles, continuing down the tail; dorsal region of the head with similar color pattern, except for a pronounced dark brown area above the orbits; a ridge of raised, dark brown scales present in the postnasal region; dorsal and lateral regions of the limbs with the same color pattern as the trunk; supralabials with the same pattern as the head, whereas the infralabials are a solid cream with few medium brown speckles; ventral side of the trunk has a cream background color, with the speckling pattern of the lateral side of the body wrapping around to approximately halfway to the midpoint of the venter; ventral side of the head has only sparse medium brown speckling; ventral region of the tail is solid cream with occasional light brown speckles that increase in intensity towards the tail tip; ventral regions of the limbs, hands, and feet are cream with medium to dark brown speckles scattered sparsely throughout; the space between the scansors is a medium brown. Coloration of holotype in life. Dorsal ground color of head, trunk, and tail chocolate brown; gradual transition to lighter coloration towards tip of the snout; light mottling posterior to eyes and along neck; dorsal limb surfaces colored as trunk, but with randomly distributed light blotches; tail lighter colored than trunk, but with series of longitudinally elongate, paravertebral cream blotches. Measurements and scale counts of holotype in mm. SVL 44.8; axilla���groin distance 24.7; total length 89.4; tail length 44.6; head length 8.3; head width 6.9; head height 4.7; eye diameter 3.1; snout length 4.9; eye���nares distance 4.0; internarial distance 1.6; interorbital distance 4.0; midbody width 7.1; femur length 6.2; tibia length 3.6; Toe-I length 2.1; Toe-IV length 3.9; tail width 2.5; tail height 2.2; supralabials 15; infralabials 12; circumorbitals 36; paravertebral scales 232; ventral scales 128; Finger-III scansors 13; Toe-IV scansors 15. Distribution, ecology and natural history. Pseudogekko atiorum is known only from Negros and Siquijor islands, although it is possible that it will eventually be discovered on other islands in the Western Visayan Faunal Region (i.e., Cebu, Panay; Fig. 1; Ferner et al., 2001). Pseudogekko atiorum represents the only species of Philippine false gecko distributed in the western Visayan islands, with the remaining diversity in the genus restricted to the northern, eastern, and sounthern islands of the archipelago (Siler et al., 2014 b, c). Although believed to be primarily an obligate forest species, at least one individual has been collected recently on the leaves of a banana plant at mid-elevation on Mt. Talinis in southeast Negros Island (PNM 9518). This individual was found within a small patch of disturbed forest where an agricultural plot had been established, surrounded by pristine secondary growth forest. Among all members of the genus, P. a t i or um is the best represented in museum collections, with 87 individuals housed at CAS and KU alone. Although historical collections reported this species to be quite common, recent expeditions over the last decade resulted in few observations of this Western Visayan gecko in the wild. Due to the absence of data allowing for confirmation of potential declines in population size and health resulting from widespread habitat destruction, and the lack of available information about the species ecology and intraspecific diversity, we do not find this species qualifies for Critically Endangered, Endangered, Vulnerable, or Near Threatened status under the IUCN criteria for classification (IUCN, 2013). Therefore, we recommend that the species be considered Least Concern at the current time, but strongly recommend that immediate efforts be made to study the health of wild populations of this species on Negros Island and, especially, Siquijor Island where no original forest persists (CDS, RMB, personal observations) but a small patch of protected secondary forest may protect this species. Etymology. We derive the new species name in honor of the Ati people, a Negrito ethnic group believed to be among the first to colonize the central Philippines. Similar to this new species of Pseudogekko, Inati (the native language of the Ati), is infrequently encountered and possibly threatened, and is estimated to be spoken by fewer than 2,000 people. The specific epithet is a noun of masculine gender. Suggested common name: Central Visayan False Gecko., Published as part of Davis, Drew R., Watters, Jessa L., K��hler, Gunther, Whitsett, Collin, Huron, Nicholas A., Brown, Rafe M., Diesmos, Arvin C. & Siler, Cameron D., 2015, Redescription of the rare Philippine false gecko Pseudogekko brevipes (Reptilia: Squamata: Gekkonidae) and description of a new species, pp. 357-374 in Zootaxa 4020 (2) on pages 363-371, DOI: 10.11646/zootaxa.4020.2.7, http://zenodo.org/record/242048, {"references":["Ferner, J. W., Brown, R. M., Sison, R. V. & Kennedy, R. S. (2001) The amphibians and reptiles of Panay Island, Philippines. Asiatic Herpetological Research, 9, 34 - 70.","Siler, C. D., Dececchi, T. A., Merkord, C. L., Davis, D. R., Christiani, T. J. & Brown, R. M. (2014 b) Cryptic diversity and population genetic structure in the rare, endemic, forest-obligate, slender geckos of the Philippines. Molecular Phylogenetics and Evolution, 70, 204 - 209. http: // dx. doi. org / 10.1016 / j. ympev. 2013.09.014","Siler, C. D., Welton, L. J., Davis, D. R., Watters, J. L., Davey, C. S., Diesmos, A. C., Diesmos, M. L. & Brown, R. M. (2014 c) Taxonomic revision of the Pseudogekko compresicorpus Complex (Reptilia: Squamata: Gekkonidae), with descriptions of three new species. Herpetological Monographs, 28, 110 - 139. http: // dx. doi. org / 10.1655 / herpmonographs-d- 14 - 00005"]}

Platymantis quezoni, sp. nov. ��� Platymantis sp. 27 ��� Brown et al. 2015 Figs. 2���4 Holotype. PNM 9817 (RMB 4067; formerly KU 339542), adult male, collected on limestone outcrop in original forest (19:00 hr) on 27 November 2001 in Barangay Malinao Ilaya, Municipality of Atimonan, Quezon Province, Luzon Island, Philippines (N 13.989 ��, E 121.818 ��; 275 m elevation; datum: WGS 85), by MLLD and G. V. A. Gee. Paratypes (paratopotypes). Sixteen males: TNHC 61989 ��� 94, bearing the same data as the holotype; PNM 9910 ��� 12, collected on 19 October 2013 by LADL, AL, MLLD, and Jason Fernandez; PNM 9913���9916, collected on 15 December 2014 by LADL and AL; KU 328735 and 328736, bearing the same data as the holotype; KU 339543, collected 9 January 2011; three females (PNM 9907 ���09, collected on 19 October 2013 by LADL, AL, MLLD and Jason Fernandez; one juvenile (KU 339544, bearing the same data as the holotype), all collected at the type locality. Diagnosis. Platymantis quezoni, sp. nov., can be distinguished from other Philippine congeners by the following suite of characters: body size 22.1���33.9 mm SVL for males, 33.8���39.7 mm for females; terminal discs of fingers and toes slightly expanded (Fig. 2 A,B); Finger I longer than Finger II; fingers and toes lacking dermal flanges along sides of digits, subarticular tubercles large and highly protuberant (Fig. 2 A,B); skin smooth, but with scattered and raised dermal tubercles (Figs. 3, 4); dorsolateral folds, mid-dorsal ridges, and other cephalic protuberances absent; color pattern medium to dark brown with black spots (Fig. 3); advertisement call a series of invariant low-frequency stridulated one-syllable barks: ���Iyohk, iyohk, iyohk������ (Fig. 6) and its restriction to terrestrial microhabitats in forest covering limestone karst formations (Fig. 5). Diagnostic character state differences are summarized in Table 1 and we provide detailed comparisons, emphasizing diagnostic differences between the new species and its close relatives below. Comparisons. We assign the new species to the subgenus Lupacolus Brown et al. 2015 on the basis of its placement (as ��� Platymantis sp. 27 ���) in Clade R of the genus Platymantis (Brown et al. 2015: Fig. 2). The subgenus Lupacolus is largely equivalent of the former P. dorsalis Group of Brown et al. (1997 a) and Alcala & Brown (1999), and contains the named taxa P. cagayensis, P. dorsalis, P. indeprensus, P. mimulus, P. naomiae, P. paengi, P. pseudodorsalis, P. spelaeus, and P. t a y l o r i but does not include P. biak, P. levigatus, P. pygmaeus (placed in subgenus Lahatnanguri; Clade O of Brown et al. 2015: Fig. 2) or P. corrugatus (subgenus Platymantis, clade name Tagomukhus; Clade N of Brown et al. 2015: Fig. 2). Members of the genus Platymantis, subgenus Lupacolus, all possess the following combination of characters: body size moderate (~ 25���45 mm SVL in males); digital discs non-expanded to minimally expanded; first finger longer than second; fingers and toes lacking dermal flanges along sides of digits; subarticular tubercles large and highly protuberant (Brown et al. 1997 a, c; Alcala & Brown 1999; Fig. 3). In overall phenotypic similarity Platymantis quezoni most resembles the related species P. paengi, P. spelaeus, (all three belong to the subgenus Lupacolus), the unrelated P. biak, P. insulatus (both of the subgenus Lahatnanguri), and P. bayani (subgenus Tahananpuno). All six taxa share the general mottled limestonecamouflaged color pattern and all exclusively inhabit limestone karst microhabitats. Platymantis quezoni shares with the other members of the subgenus Lupacolus (P. paengi, P. spelaeus) minimally expanded terminal digital discs, whereas these two species of subgenus Lahatnanguri (P. biak, P. insulatus), and all members of the subgenus Tahananpuno (including P. bayani) have widely expanded terminal discs of the fingers and toes (Brown & Alcala, 1970; Brown et al. 1997 a, c; Siler et al. 2009, 2010). Additionally, P. bayani, P. insulatus, and P. spel aeus further differ by their much larger body sizes, which do not overlap the size range of P. quezoni (Table 1). Finally, the advertisement call of the new species (a rapid series of loud, harsh, single-syllable barks) differs from that of P. biak (a much slower, low amplitude series of tonal notes), P. insulatus (a long clicking train of brief atonal pulses), P. spelaeus (a four-syllable, multinote complex call) and P. paengi (tonal, frequency sweep, amplitude modulated call). Platymantis quezoni is distinguished from the remaining members of the subgenus Tahananpuno (P. diesmosi, P. luzonensis, P. negrosensis, P. guentheri, and P. rabori) by the presence of minimally expanded (vs. widely expanded) terminal digital discs of the fingers and toes. The terrestrial limestone karst microhabitat of P. quezoni further distinguishes the new species from all of these arboreal tree frog taxa except P. d i e s m o s i (a terrestrial species calling exclusively from cliff ledges on Mt. Malino; Brown & Gonzales 2007) and its advertisement call likewise distinguishes P. quezoni from the tree frogs with amplitude modulated calls (P. luzonensis, P. negrosensis, and P. rabori) and P. guentheri, an arboreal species with a tonal, frequency sweep call. The new species is distinguished from all species of Philippine cloud frogs (formerly the P. ha z e l a e Group species) of the subgenus Tirahanulap (P. hazelae, P. isarog, P. lawtoni, P. montanus, P. panayensis, P. polillensis, P. sierramadrensis, and P. subterrestris) by its larger body size (vs. much smaller in these species, typically ~ 20���28 mm in males), by unadorned digits (vs. digits with wide dermal fringes), by strongly protuberant (vs. ventrally flattened) subarticular tubercles, by possessing a longer first finger than the second (vs. second longest) and its terrestrial limestone karst microhabitat (vs. shrub-layer vegetation of montane forests). Species of the subgenus Tirahanulap all possess tonal, constant frequency calls with complex harmonic structure, contrasting sharply with the loud, harsh barking calls of P. quezoni. From remaining members of the subgenus Lupacolus (previously referred to as the Platymantis dorsalis Group) (P. cagayensis, P. dorsalis, P. indeprensus, P. mimulus, P. naomiae, P. pseudodorsalis, and P. tayl ori), P. quezoni is distinguished by from all species except P. levigatus by the absence of dorsal folds and/or ridges, and limitation of dermal tubercles to its eyelids (vs. presence dermal folds/ridges and widely distributed tubercles over most dorsal body surfaces), and from all Lupacolus except P. paengi and P. s p e l a e us, by its terrestrial limestone forest microhabitat (vs. much more generalist inhabitant of leaf litter of forest floor and/or herb-layer vegetation in P. cagayensis, P. dorsalis, P. indeprensus, P. mimulus, P. naomiae, P. pseudodorsalis, and P. t a yl or i). Other than P. dorsalis and P. pseudodorsalis (which possess frequency sweep calls), these species possess complex, multisyllable call structure, which contrasts markedly with the simpler, single-syllable barking calls of P. quezoni. The new species differs from Platymantis corrugatus (subgenus Platymantis, clade name Tagomukhus) by its smooth to shagreened skin texture (vs. dorsal skin with elongate dermal folds), by its lateral head coloration similar to other dorsolateral surfaces (vs. presence of a dark ���mask��� coloration on the lateral side of the head of P. corrugatus) and by its exclusive limestone karst microhabitat (vs. a variety of terrestrial forest types). The advertisement call of P. quezoni is surprisingly quite similar to that of P. corrugatus (a loud series of single ���quacks���) except that P. corrugatus is generally crepuscular whereas P. qu e z o ni calls beyond sunset and well into night-time hours if precipitation is sufficient. The final set of comparisons for the recognition of P. quezoni is to the group of species comprising a recently discovered clade for which the subgenus Lahatnanguri was established (Brown et al. 2015). The new species differs from tree frogs of this clade by its terrestrial limestone karst microhabitat and possession of minimallyexpanded terminal discs of the fingers and toes (vs. widely expanded discs in the arboreal species P. banahao, P. cornutus), from the forest floor leaf litter-inhabiting, miniature species P. pygmaeus (SVL 13���16 mm in males) by its much larger body size (22���34 mm) and limestone karst microhabitat, and from the semi-aquatic species P. levigatus by its light brown, mottled dorsal coloration (vs. dark, more uniformly colored dorsum), by its drier, limestone karst microhabitats (vs. moist, riparian environments). Finally, P. quezoni differs from the limestone karst frogs of the subgenus Lahatnanguri by its minimally-expanded terminal discs of the fingers and toes (vs. markedly expanded in P. biak, and widely expanded in P. insulatus), the presence of distinct transverse banding patterns of the thigh and tibia (vs. absent or vague in P. insulatus), and its simple, single-syllable barking calls advertisement call (vs. atonal clicking pulses in P. insulatus and the slow series of low, tonal notes in P. biak.) Description of holotype. A mature male (Fig. 3); habitus moderate; head as wide as body, head length 43.9 % snout���vent length; snout protruding moderately beyond lower jaw, snout tip moderately pointed in dorsal and lateral aspects; snout 39.1 % head length; eyes moderate and anterolaterally protruding, beyond width of head in dorsal aspect; eye diameter 15.1 % snout���vent length, 34.4 % head length; lips flared, labial region slightly swollen (Fig. 4 A); interorbital region flat; eye diameter 87.9 % snout length, eye���naris distance 74.5 % eye diameter; pupil horizontally elliptical; canthus rostralis strait, only sightly medially bowed; loreal region moderately concave, sloping ventrolaterally to labia; nostrils oriented laterally; internarial region slightly concave; tympanic annulus distinct, 60.1 % eye diameter; supratympanic fold moderately protuberant, swollen and thick, extending from posterior corner of eye across entire dorsal margin of tympanum, along postero-dorsal edge of annulus, and to supra-axillary region. Tongue elongate, wider posteriorly, with shallow posterior notch, a single small anterior lingual papilla, and narrow anterior attachment; choanae situated at anterolateral edge of palate, irregularly ovoid, separated by distance six greater than diameter of single choana, partially concealed in ventral aspect by maxillary shelf; dentigerous process of vomer distinct, with four teeth visible on left process, six on right; dentigerous process angled slightly anterolaterally, approximately at 20���30 �� incline with closest (posterior) points separated by distance equal to 2 X diameter of single choana, their most distant (anterior) ends separated adjacent choana by a distance equivalent to one choana. Hands and feet large; manus length 52.6 % pes length; limbs well developed; tibia length 63.4 % snout���vent length, 111.9 % pes length; fingers slender, long; terminal discs moderately expanded (Fig. 2 A) with circummarginal grooves evident around distal edge of terminal discs; dorsal surfaces of terminal phalanges with small, cutaneous supra-articular flaps between ultimate and penultimate phalanges; relative lengths of fingers II Measurements of holotype (in mm). Snout���vent length 33.7; head length 14.8; head width 13.9; snout length 5.8; upper arm length 5.9; forearm length 7.5; femur length 18.1; tibia length 21.7; tarsus length 10.9; pes length 19.4; manus length 10.2; Finger I length 5.9; Finger III length 7.7; Toe IV length 12.6; eye���narial distance 3.8; interorbital distance 3.2; internarial distance 3.0; eye diameter 5.1; horizontal tympanic annulus diameter 3.1; Finger I disc width 1.0; Finger III disc width 1.2; Toe IV disc width 1.3; Finger III penultimate phalange width 0.8; Toe IV penultimate phalange width 0.7. Coloration of holotype in preservative. Dorsal surfaces of head, body, and limbs dark brown with black blotches; anterior portions of head darker greenish brown; limbs banded with black bars; interorbital bar present but faint: a thin, irregular, black line; lateral surfaces of head darker than dorsum, nearly greenish brown with black markings; black bar extends across dorsal half of canthus rostralis to anterior edge of eye, around margin of the palpebrum, and across suptratympanic fold; tympanum medium brownish-green with black covering dorsal-most half; lips with alternating dark green and black vertical labial bars, bordered anteriorly by black lower edge of canthal and subocular regions; three bold black bars radiate out of orbit dorsally, and extend ventrally to labial region to form the most conspicuous labial bars; transverse limb bars of irregular widths, with an overall darker coloration (compared to dorsum), due to a greater proportion of surface covered with black markings; upper arm almost devoid of black blotches, forearms each with two transverse black bars, thighs with five, tibia with three, tarsus with two; anterior flank coloration similar to dorsum on left, but predominantly black on right due to aggregation of black blotches, both sides progressively fading to less strongly patterned ventral color: dark brown blotches on throat, sternal region tan, inguinal region tan with steel bluish markings; ventral surfaces of limbs solid dark reddish brown; ventral surfaces of hands and feed dark reddish brown with light cream to tan subarticular tubercles; dorsal surfaces of hands are light brown, with alternating bands black bars on Fingers II���IV, and light tan on Finger I; dorsal surfaces of feet with solid redish brown above Toes I���III, boldly alternating light brown and black on Fingers IV and V. Coloration in life. Dorsal surfaces of head, body, and limbs variably mottled shades of dark brown and greenish-gray, a few scattered bright gold spots, and with bold black markings scattered throughout but congregating irregularly into the following characteristics: distinct black suprascapular bar, interorbital bar, transverse limb bars (on thigh, tibia, shank, and upper arm and forearm), a dark bar from the nares, along the canthus, aligned with the enlarged pupil at night (Fig. 3), and posteriorly across the dorsal-most half of the tympanum; upper lips with distinct black labial bars; pupil bordered by bright yellow ciliary ring, iris silver; flanks with enlarged black blotches and interrupted by reduced dark brown of dorsal surfaces, grading ventrally across flank to cream ground color with black blotches on anterior portion of flanks; dorsal surfaces of digits mostly black, with small brown spots. Ventrum very dark gray with black blotches on throat and sternum; liver nearly bluish gray through translucent ventral skin of chest and groin; ventral surfaces of legs nearly solid black, fading to dark purplish gray on ventral tibia and shank; ventral surfaces of hands and feet dark purplish gray with lighter gray subarticular tubercles and distal digital discs. Variation. Overall color pattern consistently mottled across the type series, typical of limestone karst inhabiting Platymantis (Brown & Alcala, 1970, 1982a; Siler et al., 2009, 2010), with dorsal ground color light tannish yellow or mustard (e.g., KU 328735; Fig. 3) to dark greenish-gray with black blotches (the holotype: PNM 9817; Fig. 3). The degree and concentration of dorsal black markings ranges, from specimens nearly covered dorsally in black markings (KU 339543, TNHC 61993), to individuals predominantly light gray to tan, with few black markings (KU 61989, 61991, 61992; Fig. 3 B). The suprascapular bar ranges from a distinct w-shaped pattern (the holotype: PNM 9817; TNHC 61990; KU 328736) to a few irregular, disconnected blotches (TNHC 61991) to an inverted v-shaped formation (KU 339543). Ventral surfaces less variable, predominantly as described for the holotype, but a few paratypes (KU 328735, 328736, TNHC 61990, 61994) possess darker throats, owing to aggregation of black blotches as in the holotype. Summaries of univariate morphological variation in the series are presented in Table 2. Females (PNM 9907 ��� 0 9, 9914) are considerably larger and more robust (Table 2), and the single juvenile (KU 339544; SVL 24.3 mm) was more brightly patterned in life, with bolder and contrasting light tan versus black blotches. Distribution. Platymantis quezoni is known only from the type locality but may be more wide spread in scattered karst outcrops of northern portions of the Bicol Peninsula (Restificar et al. 2006; MGB 2010), especially on limestone karst formations in the nearby Bondoc Peninsul, Published as part of Brown, Rafe M., Layola, Louise Abigail De, Ii, Antonio Lorenzo, Diesmos, Mae Lowe L. & Diesmos, Arvin C., 2015, A new species of limestone karst inhabiting forest frog, genus Platymantis (Amphibia: Anura: Ceratobatrachidae: subgenus Lupacolus) from southern Luzon Island, Philippines, pp. 191-210 in Zootaxa 4048 (2) on pages 194-204, DOI: 10.11646/zootaxa.4048.2.3, http://zenodo.org/record/239892, {"references":["Brown, W. C., Brown, R. M. & Alcala, A. C. (1997 a) Species of the hazelae group of Platymantis from the Philippines, with descriptions of two new species. Proceedings of the California Academy of Sciences, 49, 405 - 421.","Brown W. C., Alcala, A. C. & Diesmos A. C. (1999) Four new species of the genus Platymantis (Amphibia: Ranidae) from Luzon Island, Philippines. Proceedings of the California Academy of Sciences, 51, 449 - 460.","Brown, W. C., Alcala, A. C., Diesmos, A. C. & Alcala, E. (1997 c) Species of the guentheri group of Platymantis from the Philippines with descriptions of four new species. Proceedings of the California Academy of Sciences, 50, 1 - 20.","Brown, W. C. & Alcala, A. C. (1970) A new species of the genus Platymantis (Ranidae) with a list of the amphibians known from South Gigante Island, Philippines. Occasional Papers of the California Academy of Sciences, 84, 1 - 7.","Siler, C. D., Alcala, A. C., Diesmos, A. C. & Brown, R. M. (2009) A new species of limestone-forest frog, genus Platymantis (Amphibia: Anura: Ceratobatrachidae) from eastern Samar Island, Philippines. Herpetologica, 65, 92 - 104. http: // dx. doi. org / 10.1655 / 08 - 040 R. 1","Brown, R. M & Gonzales, J. C. (2007) A new forest frog of the genus Platymantis (Amphibia; Anura; Ranidae) from the Bicol Peninsula of Luzon Island, Philippines. Copeia, 2007, 251 - 266. http: // dx. doi. org / 10.1643 / 0045 - 8511 (2007) 7 [251: ANFFOT] 2.0. CO; 2","Brown, W. C. & Alcala, A. C. (1982 a) A new cave Platymantis (Amphibia: Ranidae) from the Philippine Islands. Proceedings of the Biological Society of Washington, 95, 386 - 391.","Restificar, S. D. F., Day, M. J. & Urich, P. B. (2006) Protection of karst in the Philippines. Acta Carsologica, 35, 121 - 130. http: // dx. doi. org / 10.3986 / ac. v 35 i 1.248","Mines and Geosciences Bureau (MGB) (2010) Geology of the Philippines. Mines and Geosciences Bureau, & Department of the Environment and Natural Resources, Quezon City, 311 pp.","Brown, W. C., Alcala, A. C. & Diesmos, A. C. (1997 b) A new species of the genus Platymantis (Amphibia: Ranidae) from Luzon Island, Philippines. Proceedings of the Biological Society of Washington, 110, 18 - 23."]}

Pseudogekko atiorum sp. nov. (Figs. 1, 3– 6) Holotype. PNM 9518 (CDS Field No. 954, formerly KU 302818), adult female, collected at 22:00 h on 21 February 2005, on Mt. Talinis, Barangay Bongbong, Municipality of Valencia, Negros Oriental Province, Negros Island, Philippines (N: 9.26000°, E: 123.20327 °; WGS 84; 1100 m elev.), by CDS. Paratypes. Two adult females (CAS-SUR 19372, 21122) collected from Municipality of Cauayan, Negros Occidental Province, Negros Island, Philippines by A.C. Alcala and Q. Alcala. One adult female (CAS 138097) collected by Q. Alcala and two adult males (CAS 145793, 147491) collected by Q. Alcala and M. Pinero, respectively, from Municipality of Pamplona, Negros Oriental Province, Negros Island, Philippines. One adult female (CAS 134269) collected by A.C. Alcala and one adult male (CAS-SUR 26778) collected by D. Empeso from Municipality of Siaton, Negros Oriental Province, Negros Island, Philippines. Four adult males (CAS 128956, 128959, 128963, 128971) collected from the Municipality of Sibulan, Negros Oriental Province, Negros Island, Philippines by L.C. Alcala. Two adult females (CAS 134292, TNHC 62478) collected by A.C Alcala and RMB, respectively, and one juvenile female (KU 327770) collected by RMB from Municipality of Valencia, Barangay Bongbong, Negros Oriental Province, Negros Island, Philippines. One adult male (CAS 145710) collected from Municipality of San Juan, Siquijor Province, Negros Island, Philippines by L.C. Alcala. Other Referred Specimens. Five specimens (CAS 89812, 147523, CAS-SUR 24588–24590) collected from Municipality of Cauayan, Negros Occidental Province, Negros Island, Philippines. Three specimens (CAS 134298, 137789, CAS-SUR 26776) collected from Negros Occidental Province, Negros Island, Philippines. Twenty specimens (CAS 137653, 137672, 137790, 138096, 138134, 139258, 145791, 145792, 145804 –6, 145875, 145876, 145976, 173760, 185991, 200539, 200540, CAS-SUR 24323, 24829) collected from Municipality of Pamplona, Negros Oriental Province, Negros Island, Philippines. One specimen (CAS-SUR 26775) collected from Municipality of Siaton, Negros Oriental Province, Negros Island, Philippines. Thirty specimens (CAS 128955, 128957, 128958, 128960–62, 128964–70, 128972–78, 131796, 132609, 142039– 46) collected from Municipality of Sibulan, Negros Oriental Province, Negros Island, Philippines. Thirteen specimens (CAS 133055, 137791, 200538, CAS-SUR 21803, 21839, 21840, 21856, 21857, 21872, 22218 –20, 24385) collected from Negros Oriental Province, Negros Island, Philippines. Diagnosis. Pseudogekko atiorum can be distinguished from congeners by the following combination of characters: (1) body size moderate (SVL 41.1–52.5 mm); (2) axilla–groin distance moderate (19.4–28.4 mm); (3) head length moderate (7.4–9.4 mm); (4) snout length long (4.3–5.8 mm); (5) Toe-IV scansors 14–17; (6) paravertebral scales 226–240; (7) ventral scales 119–129; (8) supralabials 15–17; (9) infralabials 12–15; (10) circumorbitals 35–38; (11) precloacal pores 13–15; (12) femoral pores absent; (13) dominant body coloration dark brown; (14) conspicuous head spotting present, cream; (15) conspicuous dorsolateral spotting present, cream; (16) conspicuous limb spotting absent; (17) tail banding absent; (18) body striping absent; (19) interorbital banding present, light brown; (20) iris ring coloration absent (Tables 1, 2; Figs. 3–6). Comparisons. Characters distinguishing Pseudogekko atiorum from all other species of Pseudogekko are summarized in Tables 1–3. Pseudogekko atiorum most closely resembles P. b rev i pe s; however, it differs from this species by having a longer snout–vent length in both males (41.1–52.5 mm vs. 39.0) and females (44.8–48.7 mm vs. 34.5–42.4), longer eye–nares distance (3.5–4.5 mm vs. 3.0– 3.4), a tendency towards having both a longer snout length (4.3–5.8 mm vs. 3.8–4.4) and internares distance (1.5–1.9 mm vs. 1.2–1.6), and a greater number of supralabials (15–17 vs. 13 or 14), circumorbitals (35–38 vs. 33–35), paravertebrals (226–240 vs. 211–218), ventrals (119–129 vs. 96–117), and precloacal pores (13–15 vs. 12; Figs. 3–5). Pseudogekko atiorum can be distinguished from P. chavacano by having a shorter snout–vent length (41.1– 52.5 mm vs. 54.7–55.9), longer head length (7.4–9.4 mm vs. 10.4), a tendency towards a shorter snout length (4.3– 5.8 mm vs. 5.8 –6.0), fewer infralabials (12–15 vs. 16 or 17), circumorbitals (35–38 vs. 46), and precloacal pores (13–15 vs. 16), greater number of paravertebrals (226–240 vs. 195–197), cream colored (vs. neon green) head and dorsolateral spotting, absence (vs. presence) of limb spotting and tail banding, and presence of interorbital banding (vs. absence); from P. compresicorpus by having a shorter snout–vent length (41.1–52.5 mm vs. 54.9–59.7), a tendency towards both a shorter axilla–groin distance (19.4–28.4 mm vs. 27.0– 32.6) and head length (7.4–9.4 mm vs. 9.2–11.3), fewer Toe-IV scansors (14–17 vs. 18 or 19) and circumorbitals (35–38 vs. 39–45), cream colored (vs. neon green) head and dorsolateral spotting, absence (vs. presence) of limb spotting and a light blue iris ring, and presence (vs. absence) of interorbital banding; from P. di t o y by having fewer infralabials (12–15 vs. 16 or 17) and circumorbitals (35–38 vs. 40–43), greater number of paravertebrals (226–240 vs. 180–185) and ventrals (119–129 vs. 111–118), fewer precloacal pores (13–15 vs. 18), and the presence (vs. absence) of head spotting, dorsolateral spotting, and interorbital banding; from P. pungkaypinit by having a shorter snout–vent length (41.1–52.5 mm vs. 66.6–75.3), axilla–groin distance (19.4–28.4 mm vs. 37.2–41.2), total length (80.2–113.5 mm vs. 125.3–141.2), midbody width (4.8–7.7 mm vs. 7.7–9.1), head length (7.4–9.4 mm vs. 11.4–13.6), head width (5.9–8.3 mm vs. 9.3–11.2), and snout length (4.3–5.8 mm vs. 6.7–7.5), fewer Finger-III scansors (12–14 vs. 15–17), infralabials (12–15 vs. 17–19), circumorbitals (35–38 vs. 50–55), paravertebrals (226–240 vs. 265–280), and precloacal pores (13–15 vs. 17–20), a tendency towards having fewer ventrals (119–129 vs. 125–155), the presence (vs. absence) of head and dorsolateral spotting and interorbital banding, and the absence (vs. presence) of body striping; from P. smaragdinus by a tendency towards having a shorter snout–vent length (41.1–52.5 mm vs. 50.2–64.3) and axilla– groin distance (19.4–28.4 mm vs. 26.2–35.5), fewer Finger-III scansors (12–14 vs. 15–18), paravertebrals (226– 240 vs. 241–252), and enlarged pores (13–15 vs. 32–41), a tendency towards having fewer Toe-IV scansors (14–17 vs. 16–22) and supralabials (15–17 vs. 16–19), greater number of circumorbitals (35–38 vs. 33–35), presence of dark brown (vs. yellow to orange [undisturbed] or neon green [disturbed]) body coloration, cream colored (vs. large black and small white) head and dorsolateral spotting, presence (vs. absence) of interorbital banding, and absence (vs. presence) of limb spotting and tail banding. Description of holotype. Adult female in excellent condition; small incision in the sternal region (portion of liver removed for genetic sample). Body small, slender, SVL 44.8 mm; limbs well developed, moderately slender; tail slender; margins of limbs smooth, cutaneous flaps or dermal folds absent. Head size moderate, slightly differentiated from neck, characterized by only slightly hypertrophied temporal and adductor musculature; snout broadly rounded in dorsal view and bluntly rounded in lateral view; HW 96.9 % MBW, 83.6 % HL; HL 18.4 % SVL; SNL 70.7 % HW, 59.2 % HL; dorsal surfaces of head relatively homogeneous, with slightly to moderately pronounced concave postnasal, internasal, prefrontal, and interorbital concavities; auricular opening small, nearly round, beneath temporal swellings on either side of head; tympanum deeply sunken; orbit large; eye large, pupil vertical, margin wavy; limbs and digits relatively short; thighs moderately thicker compared to upper arms; TBL 8.0% SVL, 57.9 % FL. Rostral size moderately large, margin in wide heart shape in anterodorsal view, nearly as broad as high, sutured anterolaterally with anteriormost enlarged supralabials, projecting onto dorsal surface of head to point in line with anteriormost edge of nasal; nasal surrounded by first labial, rostral, one enlarged postnasal, and two enlarged supranasals; medial supranasals separated by 3 small median scales; enlarged supranasals moderately larger than postnasals; circumorbitals 36 in total. Total number of differentiated supralabials 15, bordered dorsally by one row of differentiated, slightly enlarged snout scales; total number of differentiated infralabials 12, bordered ventrally by 2 or 3 rows of slightly enlarged scales; chin scales enlarged; postrictal scales undifferentiated; gulars small, round, nonimbricate. Dorsal cephalic scales fairly homogeneous in size, shape, disposition, and distribution; cephalic scalation slightly convex, round to oval scales; postnasal, prefrontal, internasal, and interorbital depressions; undifferentiated posterior head scales granular, slightly convex. Axilla–groin distance 55.1 % SVL; undifferentiated dorsal body scales round to tubular, moderately to sharply convex, juxtaposed, relatively homogeneous in size, many with raised edge, irregular in placement on individual scales across body; each dorsal scale surrounded by 4–6 interstitial granules; dorsals gradually transition to imbricate ventrals along lateral body surface; paravertebrals between midpoints of limb insertions 232; ventrals between midpoints of limb insertions 128; scales on dorsal surfaces of limbs more imbricate than dorsals; scales on dorsal surfaces of hands and feet similar to dorsal limb scales, heavily imbricate; ventral body scales flat, cycloid, strongly imbricate, much larger than lateral or dorsal body scales, relatively homogeneous in size. Fourteen enlarged scales in continuous precloacal series (pore-bearing series in males), arranged in a widely obtuse, “V”-formation; patch of slightly enlarged scales posterior to precloacal series, roughly three scale rows in width, 3–4 scale rows in length, forming an oval patch just posterior to precloacal series. Digits moderately expanded and covered on palmar and plantar surfaces by bowed, unnotched, undivided scansors; digits with minute vestiges of interdigital webbing; subdigital scansors of fingers (left/right) I (9,8), II (10,10), III (13,13), IV (14,14), V (9,9); subdigital scansors of toes (left/right) I (9,9), II (10,10), III (13,13), IV (15,15), V (12,12); mite eggs visible between distal subdigital scansors of digits on hands and feet; subdigital scansors of hands and feet bordered proximally (on palmar and plantar surfaces) by two or three slightly enlarged scales that form a near-continuous series with enlarged scansors; all digits clawed, but first (inner) claw greatly reduced; remaining terminal claw-bearing phalanges compressed, with moderately sized recurved claws. brevipes atiorum chavacano compresicorpus ditoy pungkaypinit smaragdinus sp. A (1 m, 3 f) (8 m, 7 f) (1 m, 1 f) (3 m, 5 f) (1 m, 1 f) (4 m, 2 f) (16 m, 17 f) (1 m, 1 f) individuals showing specific counts are given in parentheses. Characters summarized for adult specimens only. When coloration in life has not been documented, characters are listed as unkown. Enlarged pore series 12 (precloacal) 13–15 16 (precloacal) 10–14 (precloacal) 18 (precloacal) 17–20 (precloacal) 32–41 15 (precloacal) (precloacofemoral) (precloacal) Femoral pores – – – – – – + – ……continued on the next page Tail relatively long compared to body, TL 44.6 mm, 99.4 % SVL; round, not heavily depressed; TH 87.8 % TW; caudals similar in size to dorsals, subcaudals similar in size to ventrals. Coloration of holotype in preservative. Background dorsal trunk coloration light tan with intermittent small cream and dark brown speckles, continuing down the tail; dorsal region of the head with similar color pattern, except for a pronounced dark brown area above the orbits; a ridge of raised, dark brown scales present in the postnasal region; dorsal and lateral regions of the limbs with the same color pattern as the trunk; supralabials with the same pattern as the head, whereas the infralabials are a solid cream with few medium brown speckles; ventral side of the trunk has a cream background color, with the speckling pattern of the lateral side of the body wrapping around to approximately halfway to the midpoint of the venter; ventral side of the head has only sparse medium brown speckling; ventral region of the tail is solid cream with occasional light brown speckles that increase in intensity towards the tail tip; ventral regions of the limbs, hands, and feet are cream with medium to dark brown speckles scattered sparsely throughout; the space between the scansors is a medium brown. Coloration of holotype in life. Dorsal ground color of head, trunk, and tail chocolate brown; gradual transition to lighter coloration towards tip of the snout; light mottling posterior to eyes and along neck; dorsal limb surfaces colored as trunk, but with randomly distributed light blotches; tail lighter colored than trunk, but with series of longitudinally elongate, paravertebral cream blotches. Measurements and scale counts of holotype in mm. SVL 44.8; axilla–groin distance 24.7; total length 89.4; tail length 44.6; head length 8.3; head width 6.9; head height 4.7; eye diameter 3.1; snout length 4.9; eye–nares distance 4.0; internarial distance 1.6; interorbital distance 4.0; midbody width 7.1; femur length 6.2; tibia length 3.6; Toe-I length 2.1; Toe-IV length 3.9; tail width 2.5; tail height 2.2; supralabials 15; infralabials 12; circumorbitals 36; paravertebral scales 232; ventral scales 128; Finger-III scansors 13; Toe-IV scansors 15. Distribution, ecology and natural history. Pseudogekko atiorum is known only from Negros and Siquijor islands, although it is possible that it will eventually be discovered on other islands in the Western Visayan Faunal Region (i.e., Cebu, Panay; Fig. 1; Ferner et al., 2001). Pseudogekko atiorum represents the only species of Philippine false gecko distributed in the western Visayan islands, with the remaining diversity in the genus restricted to the northern, eastern, and sounthern islands of the archipelago (Siler et al., 2014 b, c). Although believed to be primarily an obligate forest species, at least one individual has been collected recently on the leaves of a banana plant at mid-elevation on Mt. Talinis in southeast Negros Island (PNM 9518). This individual was found within a small patch of disturbed forest where an agricultural plot had been established, surrounded by pristine secondary growth forest. Among all members of the genus, P. a t i or um is the best represented in museum collections, with 87 individuals housed at CAS and KU alone. Although historical collections reported this species to be quite common, recent expeditions over the last decade resulted in few observations of this Western Visayan gecko in the wild. Due to the absence of data allowing for confirmation of potential declines in population size and health resulting from widespread habitat destruction, and the lack of available information about the species ecology and intraspecific diversity, we do not find this species qualifies for Critically Endangered, Endangered, Vulnerable, or Near Threatened status under the IUCN criteria for classification (IUCN, 2013). Therefore, we recommend that the species be considered Least Concern at the current time, but strongly recommend that immediate efforts be made to study the health of wild populations of this species on Negros Island and, especially, Siquijor Island where no original forest persists (CDS, RMB, personal observations) but a small patch of protected secondary forest may protect this species. Etymology. We derive the new species name in honor of the Ati people, a Negrito ethnic group believed to be among the first to colonize the central Philippines. Similar to this new species of Pseudogekko, Inati (the native language of the Ati), is infrequently encountered and possibly threatened, and is estimated to be spoken by fewer than 2,000 people. The specific epithet is a noun of masculine gender. Suggested common name: Central Visayan False Gecko.

Brown, Rafe M., Layola, Louise Abigail De, Ii, Antonio Lorenzo, Diesmos, Mae Lowe L., Diesmos, Arvin C. (2015): A new species of limestone karst inhabiting forest frog, genus Platymantis (Amphibia: Anura: Ceratobatrachidae: subgenus Lupacolus) from southern Luzon Island, Philippines. Zootaxa 4048 (2): 191-210, DOI: http://dx.doi.org/10.11646/zootaxa.4048.2.3

ALCALUS GEN. NOV. Type species Micrixalus mariae Inger 1954. Diagnosis Members of the genus Alcalus can be distinguished from many members of the clade Anurajen (species of the genera Cornufer and Platymantis) by having (1) an intermediate body size (Al. baluensis: males 20– 25 mm Snout-Vent Length (SVL), females 26–31; Al. mariae: males 32–37 mm SVL, females 35–43); (2) a broad head (vs. slender to moderately broad); (3) a coarsely textured, shagreened, or ‘wrinkled’ skin appearance in all species (vs. smooth, tuberculate, or with longitudinal dorsolateral dermal ridges); (4) widely expanded, terminally squared, spatulate toe discs (vs. non- or minimally expanded, terminally rounded); (5) semi-aquatic microhabitat preferences (vs. preferences for terrestrial or arboreal microhabitats in most species); and by the (6) presence of nuptial pads in males (vs. absence); (7) absence of vocal sacs (vs. presence of median subgular vocal sacs); (8) absence of supernumerary tubercles on hand (vs. presence in most species); (9) presence of elongate subarticular tubercles (vs. presence, round); (10) absence of outer metatarsal tubercles on plantar surface of feet (vs. presence in most species); and (11) presence of extensive, usually full, interdigital webbing of the feet (vs. absence). Phylogenetic definition Alcalus (NCN) is a maximum crown-clade name that refers to the crown clade (C) originating in the last common ancestor of Al. mariae and all extant species that share a more recent common ancestor with Al. mariae than with Cornufer vitiensis or Platymantis corrugatus. It can also be conceived of as the largest crown clade containing Al. mariae, but not Co. vitiensis or Pl. corrugatus. Content Southeast Asian (Sunda Shelf and Palawan Island) species formerly placed in Ingerana (Table 3): Al. mariae, Al. baluensis, Al. rajae, and presumably Al. sariba (Shelford, 1905), which was not sampled (Table 3). Comment It is not surprising that the montane, semi-aquatic, Southeast Asian island archipelago species formerly referred to Ingerana comprise a monophyletic group, unrelated to the ecologically dissimilar and biogeographically disjunct mainland species of Ingerana (as presently understood, from Andaman Islands, Bhutan, China, north eastern India, Myanmar, and Nepal). Erection of a new genus to accommodate these taxa is undertaken here with reference to the phylogenetic placement of the type species of ‘true’ Ingerana (I. tenasserimensis), which in our phylogeny is more closely related (but with weak support) to the Dicroglossidae than to the Ceratobatrachidae (Fig. 2). The placement of Alcalus as the sister group of the clade Anurajen (containing genera Platymantis and Cornufer) has been confirmed elsewhere (Bossuyt et al., 2006; Frost et al., 2006; Wiens et al., 2009; Pyron & Wiens, 2011), although taxon sampling was not as extensive. The phylogenetic relationships and possible additional generic subdivision of the nonceratobatrachid (perhaps dicroglossid) species referred to Ingerana remain unstudied. Etymology A masculine noun honouring our long-term collaborator, friend, and mentor Angel C. Alcala for his numerous contributions to the systematics, ecology, conservation, and developmental biology of Southeast Asian amphibians. Suggested common name: Alcala’s dwarf mountain frogs.

Brown, Rafe M., Siler, Cameron D., Richards, Stephen J., Diesmos, Arvin C., Cannatella, David C. (2015): Multilocus phylogeny and a new classification for Southeast Asian and Melanesian forest frogs (family Ceratobatrachidae). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 174 (1): 130-168, DOI: 10.1111/zoj.12232, URL: http://dx.doi.org/10.1111/zoj.12232

CERATOBATRACHINAE BOULENGER, 1884 Diagnosis Other species of Ceratobatrachinae differ from Alcalus by having (1) a relatively narrow head (vs. wide); and (2) smooth skin, with or without dermal tubercles and/ or dermal ridges (vs. coarsely textured, shagreened, or ‘wrinkled’ in appearance); (3) rounded terminal toe discs (vs. spatulate toe discs with squarish terminal shape); (4) absence of nuptial pads in males (vs. presence); (5) presence of a medial subgular vocal sac in most species (vs. absence); (6) presence of palmar supernumerary tubercles (vs. absence); and (7) presence of rounded subarticular tubercles and outer metatarsal tubercles on plantar surface (vs. presence, elongate). Finally, most species of Ceratobatrachinae (except Cornufer guppyi, species of the subgenus Potamorana, and Platymantis levigatus) differ from Alcalus by the absence of interdigital webbing on the feet (vs. presence); and by having terrestrial or arboreal microhabitat preferences (vs. semiaquatic). Morphological synapomorphies for Ceratobatrachinae have not been identified. Content The genera Cornufer and Platymantis (see below). Comment We have not converted Ceratobatrachinae (Clade D) to a clade name, but have instead coined a new unranked clade name, Anurajen. If Ceratobatrachinae were converted to a clade name, and if the family Ceratobatrachidae (Node B) were re-ranked as Ceratobatrachinae in the future, then the phylogenetic name Ceratobatrachinae and the ranked name Ceratobatrachinae would refer to different clades, which would cause confusion. Therefore, we define a new clade name below denoting the same clade as the subfamily Ceratobatrachinae., Published as part of Brown, Rafe M., Siler, Cameron D., Richards, Stephen J., Diesmos, Arvin C. & Cannatella, David C., 2015, Multilocus phylogeny and a new classification for Southeast Asian and Melanesian forest frogs (family Ceratobatrachidae), pp. 130-168 in Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) (Zool. J. Linn. Soc.) 174 (1) on page 143, DOI: 10.1111/zoj.12232, http://zenodo.org/record/5337577, {"references":["Boulenger GA. 1884. Diagnoses of new reptiles and batrachians from the Solomon Islands, collected and presented to the British Museum by H. B. Guppy, Esq., M. B., H. M. S. ' Lark'. Proceedings of the Zoological Society of London 1884: 210 - 213."]}

ALCALINAE SUBFAM. NOV. Type genus Alcalus (see account below). Diagnosis The diagnosis for Alcalinae is the same as for the genus Alcalus, below. Phylogenetic definition We have not defined Alcalinae as a phylogenetic name because it would be redundant with Alcalus; it adds no new information about phylogenetic relationships. However, we name the ranked subfamily Alcalinae, even though it is also redundant in content with Alcalus, to provide a coordinate name for its sister-taxon Ceratobatrachinae. Content One genus, Alcalus, which includes the species Al. mariae, Al. baluensis, and Al. rajae. We anticipate that Ingerana sariba eventually will be transferred to the new genus as well. Etymology See Etymology section for the genus Alcalus below.

ALCALUS GEN. NOV. Type species Micrixalus mariae Inger 1954. Diagnosis Members of the genus Alcalus can be distinguished from many members of the clade Anurajen (species of the genera Cornufer and Platymantis) by having (1) an intermediate body size (Al. baluensis: males 20– 25 mm Snout-Vent Length (SVL), females 26–31; Al. mariae: males 32–37 mm SVL, females 35–43); (2) a broad head (vs. slender to moderately broad); (3) a coarsely textured, shagreened, or ‘wrinkled’ skin appearance in all species (vs. smooth, tuberculate, or with longitudinal dorsolateral dermal ridges); (4) widely expanded, terminally squared, spatulate toe discs (vs. non- or minimally expanded, terminally rounded); (5) semi-aquatic microhabitat preferences (vs. preferences for terrestrial or arboreal microhabitats in most species); and by the (6) presence of nuptial pads in males (vs. absence); (7) absence of vocal sacs (vs. presence of median subgular vocal sacs); (8) absence of supernumerary tubercles on hand (vs. presence in most species); (9) presence of elongate subarticular tubercles (vs. presence, round); (10) absence of outer metatarsal tubercles on plantar surface of feet (vs. presence in most species); and (11) presence of extensive, usually full, interdigital webbing of the feet (vs. absence). Phylogenetic definition Alcalus (NCN) is a maximum crown-clade name that refers to the crown clade (C) originating in the last common ancestor of Al. mariae and all extant species that share a more recent common ancestor with Al. mariae than with Cornufer vitiensis or Platymantis corrugatus. It can also be conceived of as the largest crown clade containing Al. mariae, but not Co. vitiensis or Pl. corrugatus. Content Southeast Asian (Sunda Shelf and Palawan Island) species formerly placed in Ingerana (Table 3): Al. mariae, Al. baluensis, Al. rajae, and presumably Al. sariba (Shelford, 1905), which was not sampled (Table 3). Comment It is not surprising that the montane, semi-aquatic, Southeast Asian island archipelago species formerly referred to Ingerana comprise a monophyletic group, unrelated to the ecologically dissimilar and biogeographically disjunct mainland species of Ingerana (as presently understood, from Andaman Islands, Bhutan, China, north eastern India, Myanmar, and Nepal). Erection of a new genus to accommodate these taxa is undertaken here with reference to the phylogenetic placement of the type species of ‘true’ Ingerana (I. tenasserimensis), which in our phylogeny is more closely related (but with weak support) to the Dicroglossidae than to the Ceratobatrachidae (Fig. 2). The placement of Alcalus as the sister group of the clade Anurajen (containing genera Platymantis and Cornufer) has been confirmed elsewhere (Bossuyt et al., 2006; Frost et al., 2006; Wiens et al., 2009; Pyron & Wiens, 2011), although taxon sampling was not as extensive. The phylogenetic relationships and possible additional generic subdivision of the nonceratobatrachid (perhaps dicroglossid) species referred to Ingerana remain unstudied. Etymology A masculine noun honouring our long-term collaborator, friend, and mentor Angel C. Alcala for his numerous contributions to the systematics, ecology, conservation, and developmental biology of Southeast Asian amphibians. Suggested common name: Alcala’s dwarf mountain frogs., Published as part of Brown, Rafe M., Siler, Cameron D., Richards, Stephen J., Diesmos, Arvin C. & Cannatella, David C., 2015, Multilocus phylogeny and a new classification for Southeast Asian and Melanesian forest frogs (family Ceratobatrachidae), pp. 130-168 in Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) (Zool. J. Linn. Soc.) 174 (1) on page 142, DOI: 10.1111/zoj.12232, http://zenodo.org/record/5337577, {"references":["Inger RF. 1954. Systematics and zoogeography of Philippine Amphibia. Fieldiana 33: 181 - 531.","Bossuyt F, Brown RM, Hillis DM, Cannatella DC, Milinkovitch MC. 2006. Late Cretaceous diversification resulted in continent-scale regionalism in the cosmopolitan frog family Ranidae. Systematic Biology 55: 579 - 594.","Frost DR, Grant T, Faivovich J, Bain RH, Haas A, Haddad CFB, de Sa RO, Channing A, Wilkinson M, Donnellan SC, Raxworthy CJ, Campbell JA, Blotto BL, Moler PE, Drewes RC, Nussbaum RA, Lynch JD, Green DM, Wheeler WC. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1 - 370.","Wiens JJ, Sukumaran J, Pyron RA, Brown RM. 2009. Evolutionary and biogeographic origins of high tropical diversity in Old World frogs (Ranidae). Evolution 64: 1217 - 1231.","Pyron RA, Wiens JJ. 2011. A large-scale phylogeny of Amphibia including over 2,800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution 61: 543 - 583."]}

ALCALINAE SUBFAM. NOV. Type genus Alcalus (see account below). Diagnosis The diagnosis for Alcalinae is the same as for the genus Alcalus, below. Phylogenetic definition We have not defined Alcalinae as a phylogenetic name because it would be redundant with Alcalus; it adds no new information about phylogenetic relationships. However, we name the ranked subfamily Alcalinae, even though it is also redundant in content with Alcalus, to provide a coordinate name for its sister-taxon Ceratobatrachinae. Content One genus, Alcalus, which includes the species Al. mariae, Al. baluensis, and Al. rajae. We anticipate that Ingerana sariba eventually will be transferred to the new genus as well. Etymology See Etymology section for the genus Alcalus below., Published as part of Brown, Rafe M., Siler, Cameron D., Richards, Stephen J., Diesmos, Arvin C. & Cannatella, David C., 2015, Multilocus phylogeny and a new classification for Southeast Asian and Melanesian forest frogs (family Ceratobatrachidae), pp. 130-168 in Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) (Zool. J. Linn. Soc.) 174 (1) on page 142, DOI: 10.1111/zoj.12232, http://zenodo.org/record/5337577

SUBGENUS PLATYMANTIS GÜNTHER, 1858 Diagnosis The subgenus Platymantis (currently a single recognized species, Pl. corrugatus) differs from other species of Platymantis by having (1) elongate longitudinal dermal ridges along the dorsal body surfaces (vs. dorsum smooth or tuberculate); (2) distinctive ‘quaaack’ advertisement calls (vs. frequency sweeps, pure tones, or complex calls); and (3) distinctive dark lateral head coloration (of varying shades; vs. lateral head pigment undifferentiated from surrounding coloration). The diagnostic dark lateral head coloration forms a dark ‘facemask’ that we consider a synapomorphy of this clade. Additionally, members of this subgenus can be diagnosed from species of arboreal variable Philippine forest frogs of the genera Lahatnanguri (Platymantis banahao, Pl. cornutus, and Pl. insulatus), and all members of the subgenera Tahananpuno and Tirahanulap, by the absence of expanded digital tips of fingers and toes (vs. presence of some degree of terminal digital expansion of fingers and toes), and by having a terrestrial microhabitat preference (normally calling beneath leaf litter) and a crepuscular (vs. nocturnal) calling activity pattern (Table 3). Content The allopatric populations of (1) the Luzon and West Visayan faunal regions, (2) the Camiguin Norte lineage, (3) the populations from the Mindanao faunal region islands, and (4) the Mindoro Island populations, all currently referred to Pl. corrugatus (Table 3). The subgenus Platymantis is equivalent in content to the unranked taxon Tagomukhus. Conversion of the name Platymantis would result in two different clades, ranked as genus and subgenus, with the same name, Platymantis. Therefore we define a NCN, denoting the same clade as subgenus Platymantis., Published as part of Brown, Rafe M., Siler, Cameron D., Richards, Stephen J., Diesmos, Arvin C. & Cannatella, David C., 2015, Multilocus phylogeny and a new classification for Southeast Asian and Melanesian forest frogs (family Ceratobatrachidae), pp. 130-168 in Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) (Zool. J. Linn. Soc.) 174 (1) on page 144, DOI: 10.1111/zoj.12232, http://zenodo.org/record/5337577, {"references":["Gunther ACLG. 1858. Neue Batrachier in der Sammlung des britischen Museums. Archiv fur Naturgeschichte, Berlin 24: 319 - 328."]}

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."]}

Siler, Cameron D., Linkem, Charles W., Cobb, Kerry, Watters, Jessa L., Cummings, Sean T., Diesmos, Arvin C., Brown, Rafe M. (2014): Taxonomic revision of the semi-aquatic skink Parvoscincus leucospilos (Reptilia: Squamata: Scincidae), with description of three new species. Zootaxa 3847 (3): 388-412, DOI: http://dx.doi.org/10.11646/zootaxa.3847.3.4

Parvoscincus tikbalangi sp. nov. (Figs. 3–5) Parvoscincus leucospilos: Linkem, Diesmos, Brown, 2011 (part); Linkem & Brown, 2013 (part); Brown et al. 2013 b. Holotype. PNM 9795 (ACD Field No. 1989, formerly KU 327785), adult male, collected on 15 February 2005, in Sitio Apaya, Barangay Dibuluan, Municipality of San Mariano, Isabela Province, Luzon Island, Philippines (N: 17.029 °; E: 122.1928 °; WGS- 84; 600 m in elevation), by ACD. Paratypes. KU 320522, 327786 collected on 5 February 2005 in Sitio Apaya, Barangay Dibuluan, Municipality of San Mariano, Isabela Province (same coordinates), by ACD. KU 327787 – 96 collected on 24–26 April 2005 in Barangay Del Pilar, Municipality of San Mariano, Isabela Province (N: 122.104 °, E: 16.8592 °), by ACD. Diagnosis. Parvoscincus tikbalangi can be distinguished from congeners by the following combination of characters: (1) body size medium (SVL 41.5–54.7 mm); (2) Toe-IV lamellae 14–16; (3) supralabials seven; (4) infralabials seven; (5) midbody scale rows 28–32; (6) paravertebral scale rows 58–63; (7) prefrontals separated from first supraocular; (8) frontoparietals fused; (9) head pigmentation moderately mottled; (10) upper arm pigmentation present, patchy; (11) subcaudal pigmentation absent; (12) dorsal white spots faint; (13) dorsal white bands 9–12; (14) lateral body coloration present, tan; (15) tail dorsolaterally compressed; and (16) semi-aquatic (Tables 2, 3). Comparisons. Characters distinguishing Parvoscincus tikbalangi from all species of Parvoscincus are summarized in Tables 2 and 3. Parvoscincus tikbalangi most closely resembles P. duwendorum, P. leucospilos, and P. manananggalae, but differs from P. duwendorum by having seven infralabials (vs. eight); from P. duwendorum and P. manananggalae by having Toe-IV lamellae 13–16 (vs. 12 [P. duwendorum], 17 [P. manananggalae]); from P. duwendorum by having a greater number of midbody scale rows (28–32 vs. 26) and fewer dorsal white bands (9–12 vs. 15); from P. leucospilos and P. manananggalae by having a tendency towards fewer midbody scale rows (28–32 vs. 30–34 [P. leucospilos], 32–33 [P. manananggalae]), fewer paravertebral scale rows (58–63 vs. 61–67 [P. leucospilos], 61–69 [P. manananggalae]), faint dorsal white spots (vs. large and well-defined), and by the presence of tan lateral coloration (vs. presence and bright reddish-orange [P. leucospilos], absence [P. manananggalae]); from P. leucospilos by having seven supralabials (vs. six or seven), moderate head pigmentation mottling (vs. heavy), the presence (vs. absence) of dark pigmentation on the upper arm surface; and from P. manananggalae by the absence (vs. presence) of dark subcaudal pigmentation. Description of holotype. Details of the head scalation are shown in Figure 4. An adult male Parvoscincus, SVL 53.5 mm, with clawed, pentadactyl limbs. Head distinct from neck, characterized by enlarged jaw adductor musculature in temporal region. Snout rounded in dorsal aspect, rounded in lateral aspect; rostral wide forming a nearly straight margin with nasals and frontonasal; frontonasal wider than long, in contact with nasals, rostral, anterior loreal, and prefrontal scales; prefrontals in broad medial contact, in contact with anterior and posterior loreals, frontal, frontonasal, and first supraciliary, separated from first supraocular; frontal greatly longer than wide, in contact with two supraoculars on right, two on left, and first supraciliary, rounded anteriorly, rounded posteriorly; four enlarged supraoculars, first largest; single, large frontoparietal, in contact with supraoculars II–IV; interparietal arrowhead-shaped; parietals in narrow medial contact, in narrow contact with fourth supraocular, in moderate contact with postsupraocular, and secondary temporal; primary temporals two, ventral largest, overlapping dorsal; secondary temporals two, large, dorsal largest, ventral overlapping dorsal; tertiary temporals two, dorsal largest, ventral overlapping dorsal; auricular opening large. Nasal pierced in center by large naris, surrounded anteriorly by rostral, dorsally by frontonasal, posteriorly by anterior loreal, and ventrally by first and second supralabial; anterior loreal one, narrow, posterior loreal roughly equal in size; preoculars two; supralabials seven, fifth subocular; lower eyelid scaly and semi-transparent, nonscaled “window” absent; ear large, moderately sunk. Infralabials seven, decreasing in size posteriorly in series; mental small, forming straight suture with single, large postmental and first infralabials; enlarged chin shields in three pairs; gular scales slightly smaller than ventrals. Body elongate, cylindrical, slender, with 30 equal sized midbody scales, limbs overlapping when adpressed; paravertebral scales 62, imbricate, smooth, without striations, keels or pits. Tail elongate, dorsolaterally compressed, longer than body (TL [85.0] / SVL [53.5] 1.59); subcaudal scales equal to lateral scales for basal half of tail, enlarged for distal half of original tail. Precloacal region with series of enlarged scales between pelvic region and cloaca, more elongate than ventral scales; medial precloacal scales larger; hemipenes everted. Forelimbs smaller than hind limbs, pentadactyl; forelimb scales slightly smaller in size than body scales, imbricate and smooth, reducing slightly in size distally; lamellae becoming slightly keeled distally on each digit; relative digit length I Coloration of holotype in preservative. The background dorsal coloration is dark brown, with a mottled pattern of many different shades of brown. Two faint light brown stripes run longitudinally along the length of the body, with faint light brown spots distributed between. Rows of cream spots are distributed on the dorsolateral surface of the body. The posterior half of the tail is solid medium brown in coloration. The lateral surfaces of the body are heavily mottled dark brown and cream, with small cream spots distributed throughout. The limbs are heavily mottled dark brown and cream, with the exception of a large cream spot on the upper forelimbs, proximate to the body insertion. The ventral surface of the limbs is solid cream in color, with the exception of very faint light brown mottling present on the hands and feet. The ventral portion of the body is solid cream in color to the tip of the tail, which has a medium brown mottled pattern. The dorsal surface of the head is mottled light and medium brown. This mottled color pattern is slightly darker between the eyes. The lateral surface of the head has a slightly lighter brown mottled color pattern. There is a row of six distinct cream spots on the supralabials, which are paired with another six cream spots on the infralabials. Coloration of holotype in life (Differences from preserved specimens; Fig. 5). The cream mottled color pattern and indistinct cream spots present on the lateral surface of the head transition into pale bluish-gray mottled coloration. The lateroventral surfaces of the axilla–groin region of the body and the tail are colored pale orangecream. Within the axilla–groin region and again on the tail, the coloration transitions from more orange in coloration anteriorly to more cream posteriorly. The surface around the upper forelimbs, proximate to the body, is colored light orange-cream. Measurements and scale counts of holotype in mm. SVL 53.5; AGD 26.7; TotL 138.5; TL 85.0; HL 12.2; HW 7.2; SnFa 18.2; ED 3.1; SNL 3.8; IND 1.8; FLL 5.0; HLL 6.4; MBSR 30; PVSR 62; ToeIVlam 16; SL 7; IFL 7; SO 4. Variation. We observed the following variation in the degree of contact between head scales: prefrontals in medial contact (KU 320522, 327785–93, 327795, 327796) or separated (KU 327794). Scale pigmentation was observed to vary among the examined series: dark cloacal scale pigmentation was present (KU 320522, 327785, 327791) or absent (KU 327786 –90, 327792– 96). Distribution, ecology and natural history. Parvoscincus tikbalangi is known only from northeastern Luzon Island in the Isabela Province (Fig. 1). Samples were found in the Barangays Del Pilar and Dibuluan in the Municipality of San Mariano, in the Sierra Madre Mountain range. This semi-aquatic species of forest skink occurs in primary- and secondary-growth forests, in riparian microhabitats. At this time we are unable to appropriately evaluate this species against the IUCN criteria for classification due to the lack of available information about its distribution and natural history. We therefore classify this species Data Deficient DD (IUCN, 2010) pending future studies on this unique semi-aquatic forest skink. Etymology. The specific epithet is a patronym derivation of the name “Tikbalang,” a mythological part man, part horse mountain forest creature from Filipino folklore. Tikbalang is said to jump down from trees to dispatch its unsuspecting victims by decapitation. Suggested common name: Sierra Madres Aquatic Skink.

Parvoscincus leucospilos (Peters 1872) (Figs. 3���5) Lygosoma (Hinulia) leucospilos Peters, 1872: 684. Sphenomorphus leucospilos Brown and Alcala, 1980: 172; Bauer et al., 1995; Linkem et al., 2011; McLeod et al., 2011. Parvoscincus leucospilos Linkem, Diesmos & Brown, 2011; Linkem & Brown 2013 (part) Type designation. Based on ICZN articles 74.1 and 74.7 and in accordance with recommendation 74 D we designate ZMB 7467 as the lectotype of Parvoscincus leucospilos. Specimen CAS 64232 is designated a paralectotype. Diagnosis. Parvoscincus leucospilos can be distinguished from congeners by the following combination of characters: (1) body size medium (SVL 42.6���54.4 mm); (2) Toe-IV lamellae 15���17; (3) supralabials six or seven; (4) infralabials 6���9; (5) midbody scale rows 30���34; (6) paravertebral scale rows 61���67; (7) prefrontals in medial contact; (8) prefrontals contact first supraocular; (9) frontoparietals fused; (10) head pigmentation heavily mottled; (11) upper arm pigmentation absent; (12) subcaudal pigmentation absent; (13) dorsal white spots large, welldefined; (14) dorsal white bands 9���13; (15) lateral body coloration bright reddish-orange; (16) tail dorsolaterally compressed; and (17) semi-aquatic (Tables 2, 3). Comparisons. Characters distinguishing Parvoscincus leucospilos from all species of Parvoscincus are summarized in Tables 2 and 3. Parvoscincus leucospilos most closely resembles P. duwendorum, P. manananggalae, and P. tikbalangi. However, P. leucospilos differs from these three taxa by having head pigmentation heavily mottled (vs. moderately mottled; Figs. 4, 5), and by the absence (vs. presence) of dark pigmentation on the upper arm surface. Parvoscincus leucospilos further differs from P. duwendorum and P. manananggalae by having prefrontals in medial contact (vs. separated); from P. duwendorum and P. tikbalangi by having dorsal white spots large and well-defined (vs. faint); from P. manananggalae and P. tikbalangi by the presence of bright reddish-orange lateral coloration (vs. absence of coloration [P. manananggalae], presence, reduced, tan [P. tikbalangi]); from P. duwendorum by having a greater number of Toe-IV lamellae (15���17 vs. 12), a greater number of paravertebral scale rows (61���67 vs. 60), a greater number of midbody scale rows (30���34 vs. 26), a tendency towards a greater number of longitudinal ventral scale rows (41���47 vs. 41), and fewer dorsal white bands (9���13 vs. 15); from P. manananggalae by the absence of subcaudal dark pigmentation (vs. presence); and from P. tikbalangi by having a tendency towards a greater number of paravertebral scale rows (61���67 vs. 58���63) and a tendency towards a greater number of midbody scale rows (30���34 vs. 28���32). Redescription (based on examination of paralectotype, CAS 64232, and 10 recently collected specimens, Appendix 2). Details of the head scalation of an adult male (KU 313870) are shown in Figure 4, and scientific illustrations of the dorsal and lateral views of the head of the paralectotype are shown in Figure 6. A medium sized Parvoscincus, SVL 42.6���54.4 mm, with clawed, pentadactyl limbs. Head distinct from next, with enlarged jaw adductor musculature in the temporal region. Snout pointed in dorsal aspect, rounded in lateral aspect; rostral wide forming a nearly perpendicular margin with frontonasal, slightly rounded margin with nasals; frontonasal wider than long, in contact with nasals, rostral, anterior loreal, and prefrontal scales; prefrontals in broad medial contact, in contact with anterior and posterior loreals, frontal, frontonasal, and first supraocular, and in some specimens, in point contact with first supraciliary; frontal greatly longer than wide, in contact with two supraoculars on right, two on left, rounded anteriorly, pointed posteriorly; four enlarged supraoculars, first largest; single, large frontoparietal, in contact with supraoculars II���IV or III and IV; interparietal arrowhead-shaped; parietals in moderate to broad medial contact, in contact with fourth supraocular, postsupraoculars, and primary and secondary temporals; primary temporals two, ventral largest, overlapping dorsal; secondary temporals two, large, dorsal largest, ventral overlapping dorsal; tertiary temporals two, dorsal largest, ventral overlapping dorsal; auricular opening large. Nasal pierced in center by large naris, surrounded anteriorly by rostral, dorsally by frontonasal, posteriorly by anterior loreal, and ventrally by first supralabial; anterior loreal one, posterior loreal roughly equal in size; preoculars two; supralabials six or seven; lower eyelid scaly, semi-transparent, nonscaled ���window��� absent; ear large, moderately sunk. Infralabials 6���9, decreasing in size posteriorly in series; mental small, forming straight suture with single, large postmental and first infralabials; enlarged chin shields in three pairs; gular scales slightly smaller than ventrals. Body elongate, cylindrical, slender, with 30���34 equal sized midbody scales, limbs overlapping when adpressed; paravertebral scales 61���67, imbricate, smooth, without striations, keels or pits. Tail elongate, dorsolaterally compressed, longer than body (TL/SVL 0.99���1.57 [1.26 �� 0.18]); subcaudal scales similar to lateral scales for basal half of tail, enlarged for distal half of original tail. Precloacal region with series of enlarged scales between pelvic region and cloaca, more elongate than ventral scales; medial precloacal scales larger. Forelimbs smaller than hind limbs, pentadactyl; forelimb scales slightly smaller in size than body scales, imbricate and smooth, reducing slightly in size closer to manus; lamellae becoming slightly keeled distally on each digit; relative digit length I Coloration in preservative (Figs. 3, 4). The dorsal background color is medium brown, with pronounced mottling of various shades of brown. Three longitudinal rows of cream spots, ringed with darker brown, start just behind the head and run the length of the body with faint light brown longitudinal stripes between. Spotting and striping fade into a solid medium brown color approximately halfway down the tail. On the lateral surface of the body, spots merge to become indistinct perpendicular cream stripes, with mottled medium brown between. This pattern fades indistinctly into the solid cream present on the ventral surface. The distinctive forelimb spot is pronounced and continues nearly to the first joint. Otherwise, the mottled pattern present on the dorsal surface of the body continues down the dorsal surface of the fore- and hind limbs. The ventral surface of the limbs is a solid cream, with the exception of very faint light brown mottling present on the hands and feet. The solid ventral cream color of the body extends to about halfway down the length of the tail, then transitions to thin medium brown perpendicular stripes. The stripes become increasingly close together as they approach the tail tip, which is a solid medium brown. The dorsal surface of the head is mottled the same as the dorsal surface of the body, with various shades of brown. Three small cream spots are arranged in a triangle on the head just posterior to the eyes. A series of distinct cream spots are visible on the labial scales, one row on the supralabial series and one on the infralabial series. A slightly lighter brown mottling occurs just below the eye and is present between the spots. The ventral surface of the head and body are solid cream in color. Coloration in life ( Differences from preserved specimens; Fig. 5). On the lateral surfaces of the body, the dark brown, mottled ground coloration is replaced with a vivid reddish-orange ground color, with randomly distributed small cream spots. The non-mottled patch on the upper forelimb proximate to the body is also a solid reddishorange color. The cream and brown mottling on the lateral surface of the head and the body anterior to the forelimb insertion is replaced by a pale blue and brown mottled pattern. Measurements and scale counts of paralectotype in mm. SVL 44.2; AGD 23.2; TotL 101.0; TL 56.8; HL 6.1; HW 5.4; SnFa 16.5; ED 2.2; SNL 3.7; IND 1.0; FLL 4.3; HLL 6.2; MBSR 31; PVSR 63; ToeIVlam 17; SL 7; IFL 7; SO 4. Distribution, ecology and natural history. Parvoscincus leucospilos is known from south-central and southeastern Luzon Island (Fig. 1). No mention of the specific locality or province on Luzon Island was given for the types (Peters, 1872). Although we are certain of our identification of P. leucospilos, we are unable to pinpoint the type locality on the basis of available specimens. Specimens are from Mt. Labo, Camarines Norte Province (KU 313870), Mt. Palali, Barangay Maddiangat, Municipality of Quezon, Nueva Vizcaya Province (KU 325806 ��� 13), Barangay Kabayunan, Municipality of Dona Remedios Trinidad, Bulucan Province (KU 329388 ��� 92) and Mt. Banahao, Municipality of Tayabas, Quezon Province (TNHC 62682). The known elevational range of this species is 200���800 meters above sea level, based on Brown et al. (2013 b). Parvoscincus leucospilos occurs in primary- and secondary-growth forest habitats in riparian microhabitats; this semi-aquatic species frequently dives into cold, rapidly running water of montane streams when disturbed. The species is primarily found active during the day and has also been found sleeping on the edge of running water in forest debris at night. Parvoscincus leucospilos can be found in sympatry with Parvoscincus abstrusus, P. arvindiesmosi, P. beyeri, P. jimmymcguirei, P. laterimaculatus, and P. st eerei. We have evaluated this species against the IUCN criteria for classification, and find that it does not qualify for Critically Endangered, Endangered, Vulnerable, or Near Threatened status. Parvoscincus leucospilos is quite abundant at sampled localities provided that some vegetation cover borders the streams where it has been collected and observed. We therefore classify this species as Least Concern LC (IUCN, 2010). Etymology. The specific epithet ��� leucospilos ��� apparently refers to the distinctive coloration of this species, consisiting of prominent rows of white (from the Latin adjective ��� leucos ���) spots (Latin, noun, ��� spilos ���) down the dorsal surfaces of the body., Published as part of Siler, Cameron D., Linkem, Charles W., Cobb, Kerry, Watters, Jessa L., Cummings, Sean T., Diesmos, Arvin C. & Brown, Rafe M., 2014, Taxonomic revision of the semi-aquatic skink Parvoscincus leucospilos (Reptilia: Squamata: Scincidae), with description of three new species, pp. 388-412 in Zootaxa 3847 (3) on pages 393-397, DOI: 10.11646/zootaxa.3847.3.4, http://zenodo.org/record/226283, {"references":["Peters, W. C. H. (1872) Uber eine neue von Hrn. Dr. A. B. Meyer auf Luzon entdeckte Art von Eidechsen (Lygosoma (Hinulia) leucospilos) und eine von demselben in Nordcelebes gefundene neue Schlangengattung (Allophis nigricaudus). Monatsberichte der Koniglich preussischen Akademie der Wissenschaften zu Berlin, 1872, 684 - 687.","Brown, W. C. & Alcala, A. C. (1980) Philippine Lizards of the Family Scincidae. Silliman University Press, Dumaguete City, Philippines, 264 pp.","Linkem, C. W., Diesmos, A. C. & Brown, R. M. (2011) Molecular systematics of the Philippine forest skinks (Squamata: Scincidae: Sphenomorphus): testing morphological hypotheses of interspecific relationships. Zoological Journal of the Linnean Society, 163, 1217 - 1243. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.2011.00747. x","McLeod, D. S., Siler, C. D., Diesmos, A. C., Diesmos, M. L. D., Garcia, V. S., Arkonceo, A. O., Balaquit, K. L., Uy, C. C., Villaseran, M. M., Yarra, E. C. & Brown, R. M. (2011) Amphibians and Reptiles of Luzon Island, V: the herpetofauna of Angat Dam Watershed, Bulacan Province, Luzon Island, Philippines. Asian Herpetological Research, 2011, 177 - 198. http: // dx. doi. org / 10.3724 / sp. j. 1245.2011.00177","Brown, R. M., Siler, C. D., Oliveros, C. H., Welton, L. J., Rock, A., Swab, J., Van Weerd, M., van Beijnen, J., Jose, E., Rodriguez, D., Jose, E. & Diesmos, A. C. (2013 b) The amphibians and reptiles of Luzon Island, Philippines, VIII: the herpetofauna of Cagayan and Isabela Provinces, northern Sierra Madre Mountain Range. Zookeys, 266, 1 - 120. http: // dx. doi. org / 10.3897 / zookeys. 266.3982"]}

Parvoscincus duwendorum sp. nov. (Figs. 3–5) Parvoscincus leucospilos Linkem, Diesmos, Brown, 2011 (part); Linkem & Brown, 2013 (part); Brown et al. 2012. Holotype. PNM 9793 (RMB Field No. 14261, formerly KU 329929), subadult, collected on 18 June 2011, on Mt. Pao, Barangay Adams, Municipality of Adams, Ilocos Norte Province, Luzon Island, Philippines (N: 18.438 °, E: 120.878 °; WGS- 84; 750 m in elevation), by RMB. Diagnosis. Parvoscincus duwendorum can be distinguished from congeners by the following combination of characters: (1) adult body size presumed medium (subadult SVL 33.5 mm); (2) Toe-IV lamellae 12; (3) supralabials seven; (4) infralabials eight; (5) midbody scale rows 26; (6) paravertebral scale rows 60; (7) prefrontals separated; (8) prefrontals contact first supraocular; (9) frontoparietals fused; (10) head pigmentation moderately mottled; (11) upper arm pigmentation present, patchy; (12) cloacal scale dark pigmentation absent; (13) subcaudal pigmentation absent; (14) dorsal white spots faint; (15) dorsal white bands 15; (16) tail dorsolaterally compressed; and (17) semi-aquatic (Tables 2, 3). Comparisons. Characters distinguishing Parvoscincus duwendorum from all species of Parvoscincus are summarized in Tables 2 and 3. Parvoscincus duwendorum most closely resembles P. manananggalae, P. leucospilos, and P. tikbalangi. However, P. duwendorum differs from these three taxa by having Toe-IV lamellae 12 (vs. 17 [P. manananggalae], 15–17 [P. leucospilos], 14–16 [P. tikbalangi]), fewer midbody scale rows (26 vs. 32–33 [P. manananggalae], 30–34 [P. leucospilos], 28–32 [P. tikbalangi]), a greater number of dorsal white spot rows (15 vs. 9–13 [P. leucospilos], 9–12 [P. manananggalae, P. tikbalangi]), and the absence of precloacal scale dark pigmentation (vs. absence or presence [P. leucospilos, P. tikbalangi], presence [P. manananggalae]). Parvoscincus duwendorum further differs from P. leucospilos and P. manananggalae by having fewer paravertebral scale rows (60 vs. 61–67 [P. leucospilos], 61–69 [P. manananggalae]); from P. manananggalae and P. tikbalangi by having infralabials eight (vs. six or seven [P. manananggalae], seven [P. tikbalangi]); from P. leucospilos by having head pigmentation moderately mottled (vs. heavily mottled), and the presence of dark pigmentation on the upper arm surface (vs. absence); from P. manananggalae by having fewer longitudinal ventral scale rows (41 vs. 43–49), and the absence (vs. presence) of subcaudal dark pigmentation; from P. leucospilos and P. tikbalangi by having prefrontals separated (vs. in medial contact [P. leucospilos], separated or in medial contact [P. tikbalangi]); and from P. tikbalangi by having prefrontals and first supraoculars in contact (vs. separated). TABLE 2. Distribution of diagnostic characters (+ present; 0 absent) among species in the genus Parvoscincus. Parvoscincus palawanensis does not have prefrontal scales. (A) Brown et al. 2010, ) Linkem and Brown, 2013, (C) Brown and Alcala 1980, (D) Ferner et al. 1997. Characters included for comparison are: snout–vent length (SVL), midbody scale-row count (MBSR), paravertebral scale-row count (PVSR), Toe-IV lamellae count (ToeIVlam), supralabial count (SL), infralabial count (IFL), number of anterior loreals, contact between profrontal scales, and microhabitat preference. leucospilos duwendorum manananggalae tikbalangi (6 m 1, 5 f) (1 juv.) (3 m, 2 f) (7 m, 1 f) Paralectotype (CAS 64232) presumed male following visual inspection of external body; authors avoided creating incision on rare type specimen. Description of holotype. Details of the head scalation are shown in Figure 4. A subadult Parvoscincus, SVL 33.5 mm, with clawed, pentadactyl limbs. Head distinct from neck, characterized by enlarged jaw adductor musculature in temporal region. Snout sharply pointed in dorsal aspect, rounded in lateral aspect; rostral wide forming a nearly perpendicular margin with nasals and frontonasal; frontonasal equally wide as long, in contact with nasals, rostral, anterior loreal, and prefrontal scales; prefrontals broadly separated, in contact with anterior and posterior loreals, frontal, frontonasal, first supraciliary, and first supraocular; frontal greatly longer than wide, in contact with two supraoculars on right, two on left, rounded anteriorly, sharply rounded posteriorly; four enlarged supraoculars, first largest; single, large frontoparietals, in contact with supraoculars II–IV; interparietal arrowhead; parietals in moderate medial contact, in contact with fourth supraocular, postsupraocular, and secondary temporal; primary temporals two, ventral largest, overlapping dorsal; secondary temporals two, large, dorsal largest, ventral overlapping dorsal; tertiary temporals two, dorsal largest, ventral overlapping dorsal; auricular opening large. Nasal pierced in center by large naris, surrounded anteriorly by rostral, dorsally by frontonasal, posteriorly by anterior loreal, and ventrally by first supralabial; anterior loreal one, posterior loreal equal in size; preoculars two; supralabials seven, fifth subocular; lower eyelid scaly and semi-transparent, nonscaled “window” absent; ear large, moderately sunk. Infralabials eight, decreasing in size posteriorly in series; mental small, forming straight suture with single, large postmental and first infralabials; enlarged chin shields in three pairs; gular scales slightly smaller than ventrals. Body elongate, cylindrical, slender, with 26 equal sized midbody scales, limbs overlapping when adpressed; paravertebral scales 60, imbricate, smooth, without striations, keels or pits. Tail elongate, dorsolaterally compressed, longer than body (TL [50.0] / SVL [33.5] 1.49); subcaudal scales nondifferentiated for basal half of tail, enlarged for distal half of original tail. Precloacal region with series of enlarged scales between pelvic region and cloaca, more elongate than ventral scales; medial precloacal scales larger. FIGURE 6. Illustration of head of the examined Parvoscincus leucospilos paralectotype (CAS 64232) in dorsal and lateral views. Taxonomically diagnostic head scales are labeled as follows: C, chin shield; F, frontal; FN, frontonasal; FP, frontoparietal; IL, infralabial; IP, interparietal; L, loreal; M, mental; N, nasal; P, parietal; PF, prefrontal; PM, postmental; PO, preocular; PSO, presubocular; PoSO, postsupraoculars; R, rostral; SC, supraciliary; SL, supralabial; SO, supraocular; T 1, primary temporal; T 2, secondary temporal; and T 3, tertiary temporal. Roman numerals indicate scales in the supraocular series. Illustrations by STC and CDS. Forelimbs smaller than hind limbs, pentadactyl; forelimb scales slightly smaller in size than body scales, imbricate and smooth, reducing slightly in size closer to manus; lamellae becoming slightly keeled distally on each digit; relative digit length with lamellae V Coloration of holotype in preservative. The dorsal background color is medium brown, with tan mottling throughout. Two parallel light brown stripes run longitudinally down the body, starting at the base of the head and merging to one just past the start of the tail. Between these two stripes is a paravertebral row of darker brown spots. The venter color is a solid cream, without mottling. Along the lateral surface of the body, the dorsal and ventral color patterns merges in a scalloped pattern. There is a distinct cream spot on the forelimbs, just proximate to the body. The dorsal fore- and hind limbs possess the same mottling pattern present on the dorsal surface of the body. The ventral surfaces of the limbs are a solid cream color. The dorsal surface of the hands and feet are a mottled medium brown, with the exception of the cream-colored Finger I and Finger II on the forelimbs and Toe V on the hind limbs. The dorsal and dorsolateral surfaces of the head have a light and medium brown mottled pattern. This coloration transitions sharply into a cream and light brown mottled pattern ventrally. The labial scales are mottled light brown and cream. The ventral surface of the head is solid cream in color. Color in life —Coloration in life did not differ substantially from the preserved holotype (RMB, personal observations). Measurements and scale counts of holotype in mm. SVL 33.5; AGD 17.5; TotL 83.5; TL 50.0; HL 4.0; HW 7.7; SnFa 12.4; ED 1.5; SNL 2.7; IND 1.3; FLL 3.3; HLL 4.0; MBSR 26; PVSR 60; ToeIVlam 12; SL 7; IFL 8; SO 4. Distribution, ecology and natural history. Parvoscincus duwendorum is known from a single specimen found at 750 elevation on Mt. Pao in the Ilocos Norte Province of northwestern Luzon Island (Fig. 1). Although the species is recognized currently to occur in secondary-growth forest habitats, it is presumed the species also inhabits suitable primary forest. This species is semi-aquatic and can be found near riparian habitat. Parvoscincus duwendorum can be found in sympatry with Parvoscincus igorotorum, P. jimmymcguirei, and P. steerei. The new species was also observed on Mt. Cagua at 300–500 m (Brown et al 2013 b) but specimens eluded biologists by jumping in rapidly running montane streams. At this time we are unable to appropriately evaluate this species against the IUCN criteria for classification due to the lack of available information about its distribution and natural history. We therefore classify this species Data Deficient DD (IUCN, 2010) pending future studies on this unique semi-aquatic forest skink. Etymology. The specific epithet is a plaural derivation of the Filipino folklore term “Duwende,” which is chosen here to celebrate the country’s rich tradition of mythological forest animals and spirits. Duwendes are believed to be little fairy-like forest creatures, such as goblins, pixies, and elves, and believed to live in trees, termite mounds, and burrows in hillsides. The may bring bad or good fortune to humans and are often considered to be mischievous in nature. Suggested common name: Cordillera Aquatic Skink.

Parvoscincus tikbalangi sp. nov. (Figs. 3���5) Parvoscincus leucospilos: Linkem, Diesmos, Brown, 2011 (part); Linkem & Brown, 2013 (part); Brown et al. 2013 b. Holotype. PNM 9795 (ACD Field No. 1989, formerly KU 327785), adult male, collected on 15 February 2005, in Sitio Apaya, Barangay Dibuluan, Municipality of San Mariano, Isabela Province, Luzon Island, Philippines (N: 17.029 ��; E: 122.1928 ��; WGS- 84; 600 m in elevation), by ACD. Paratypes. KU 320522, 327786 collected on 5 February 2005 in Sitio Apaya, Barangay Dibuluan, Municipality of San Mariano, Isabela Province (same coordinates), by ACD. KU 327787 ��� 96 collected on 24���26 April 2005 in Barangay Del Pilar, Municipality of San Mariano, Isabela Province (N: 122.104 ��, E: 16.8592 ��), by ACD. Diagnosis. Parvoscincus tikbalangi can be distinguished from congeners by the following combination of characters: (1) body size medium (SVL 41.5���54.7 mm); (2) Toe-IV lamellae 14���16; (3) supralabials seven; (4) infralabials seven; (5) midbody scale rows 28���32; (6) paravertebral scale rows 58���63; (7) prefrontals separated from first supraocular; (8) frontoparietals fused; (9) head pigmentation moderately mottled; (10) upper arm pigmentation present, patchy; (11) subcaudal pigmentation absent; (12) dorsal white spots faint; (13) dorsal white bands 9���12; (14) lateral body coloration present, tan; (15) tail dorsolaterally compressed; and (16) semi-aquatic (Tables 2, 3). Comparisons. Characters distinguishing Parvoscincus tikbalangi from all species of Parvoscincus are summarized in Tables 2 and 3. Parvoscincus tikbalangi most closely resembles P. duwendorum, P. leucospilos, and P. manananggalae, but differs from P. duwendorum by having seven infralabials (vs. eight); from P. duwendorum and P. manananggalae by having Toe-IV lamellae 13���16 (vs. 12 [P. duwendorum], 17 [P. manananggalae]); from P. duwendorum by having a greater number of midbody scale rows (28���32 vs. 26) and fewer dorsal white bands (9���12 vs. 15); from P. leucospilos and P. manananggalae by having a tendency towards fewer midbody scale rows (28���32 vs. 30���34 [P. leucospilos], 32���33 [P. manananggalae]), fewer paravertebral scale rows (58���63 vs. 61���67 [P. leucospilos], 61���69 [P. manananggalae]), faint dorsal white spots (vs. large and well-defined), and by the presence of tan lateral coloration (vs. presence and bright reddish-orange [P. leucospilos], absence [P. manananggalae]); from P. leucospilos by having seven supralabials (vs. six or seven), moderate head pigmentation mottling (vs. heavy), the presence (vs. absence) of dark pigmentation on the upper arm surface; and from P. manananggalae by the absence (vs. presence) of dark subcaudal pigmentation. Description of holotype. Details of the head scalation are shown in Figure 4. An adult male Parvoscincus, SVL 53.5 mm, with clawed, pentadactyl limbs. Head distinct from neck, characterized by enlarged jaw adductor musculature in temporal region. Snout rounded in dorsal aspect, rounded in lateral aspect; rostral wide forming a nearly straight margin with nasals and frontonasal; frontonasal wider than long, in contact with nasals, rostral, anterior loreal, and prefrontal scales; prefrontals in broad medial contact, in contact with anterior and posterior loreals, frontal, frontonasal, and first supraciliary, separated from first supraocular; frontal greatly longer than wide, in contact with two supraoculars on right, two on left, and first supraciliary, rounded anteriorly, rounded posteriorly; four enlarged supraoculars, first largest; single, large frontoparietal, in contact with supraoculars II���IV; interparietal arrowhead-shaped; parietals in narrow medial contact, in narrow contact with fourth supraocular, in moderate contact with postsupraocular, and secondary temporal; primary temporals two, ventral largest, overlapping dorsal; secondary temporals two, large, dorsal largest, ventral overlapping dorsal; tertiary temporals two, dorsal largest, ventral overlapping dorsal; auricular opening large. Nasal pierced in center by large naris, surrounded anteriorly by rostral, dorsally by frontonasal, posteriorly by anterior loreal, and ventrally by first and second supralabial; anterior loreal one, narrow, posterior loreal roughly equal in size; preoculars two; supralabials seven, fifth subocular; lower eyelid scaly and semi-transparent, nonscaled ���window��� absent; ear large, moderately sunk. Infralabials seven, decreasing in size posteriorly in series; mental small, forming straight suture with single, large postmental and first infralabials; enlarged chin shields in three pairs; gular scales slightly smaller than ventrals. Body elongate, cylindrical, slender, with 30 equal sized midbody scales, limbs overlapping when adpressed; paravertebral scales 62, imbricate, smooth, without striations, keels or pits. Tail elongate, dorsolaterally compressed, longer than body (TL [85.0] / SVL [53.5] 1.59); subcaudal scales equal to lateral scales for basal half of tail, enlarged for distal half of original tail. Precloacal region with series of enlarged scales between pelvic region and cloaca, more elongate than ventral scales; medial precloacal scales larger; hemipenes everted. Forelimbs smaller than hind limbs, pentadactyl; forelimb scales slightly smaller in size than body scales, imbricate and smooth, reducing slightly in size distally; lamellae becoming slightly keeled distally on each digit; relative digit length I Coloration of holotype in preservative. The background dorsal coloration is dark brown, with a mottled pattern of many different shades of brown. Two faint light brown stripes run longitudinally along the length of the body, with faint light brown spots distributed between. Rows of cream spots are distributed on the dorsolateral surface of the body. The posterior half of the tail is solid medium brown in coloration. The lateral surfaces of the body are heavily mottled dark brown and cream, with small cream spots distributed throughout. The limbs are heavily mottled dark brown and cream, with the exception of a large cream spot on the upper forelimbs, proximate to the body insertion. The ventral surface of the limbs is solid cream in color, with the exception of very faint light brown mottling present on the hands and feet. The ventral portion of the body is solid cream in color to the tip of the tail, which has a medium brown mottled pattern. The dorsal surface of the head is mottled light and medium brown. This mottled color pattern is slightly darker between the eyes. The lateral surface of the head has a slightly lighter brown mottled color pattern. There is a row of six distinct cream spots on the supralabials, which are paired with another six cream spots on the infralabials. Coloration of holotype in life (Differences from preserved specimens; Fig. 5). The cream mottled color pattern and indistinct cream spots present on the lateral surface of the head transition into pale bluish-gray mottled coloration. The lateroventral surfaces of the axilla���groin region of the body and the tail are colored pale orangecream. Within the axilla���groin region and again on the tail, the coloration transitions from more orange in coloration anteriorly to more cream posteriorly. The surface around the upper forelimbs, proximate to the body, is colored light orange-cream. Measurements and scale counts of holotype in mm. SVL 53.5; AGD 26.7; TotL 138.5; TL 85.0; HL 12.2; HW 7.2; SnFa 18.2; ED 3.1; SNL 3.8; IND 1.8; FLL 5.0; HLL 6.4; MBSR 30; PVSR 62; ToeIVlam 16; SL 7; IFL 7; SO 4. Variation. We observed the following variation in the degree of contact between head scales: prefrontals in medial contact (KU 320522, 327785���93, 327795, 327796) or separated (KU 327794). Scale pigmentation was observed to vary among the examined series: dark cloacal scale pigmentation was present (KU 320522, 327785, 327791) or absent (KU 327786 ���90, 327792��� 96). Distribution, ecology and natural history. Parvoscincus tikbalangi is known only from northeastern Luzon Island in the Isabela Province (Fig. 1). Samples were found in the Barangays Del Pilar and Dibuluan in the Municipality of San Mariano, in the Sierra Madre Mountain range. This semi-aquatic species of forest skink occurs in primary- and secondary-growth forests, in riparian microhabitats. At this time we are unable to appropriately evaluate this species against the IUCN criteria for classification due to the lack of available information about its distribution and natural history. We therefore classify this species Data Deficient DD (IUCN, 2010) pending future studies on this unique semi-aquatic forest skink. Etymology. The specific epithet is a patronym derivation of the name ���Tikbalang,��� a mythological part man, part horse mountain forest creature from Filipino folklore. Tikbalang is said to jump down from trees to dispatch its unsuspecting victims by decapitation. Suggested common name: Sierra Madres Aquatic Skink., Published as part of Siler, Cameron D., Linkem, Charles W., Cobb, Kerry, Watters, Jessa L., Cummings, Sean T., Diesmos, Arvin C. & Brown, Rafe M., 2014, Taxonomic revision of the semi-aquatic skink Parvoscincus leucospilos (Reptilia: Squamata: Scincidae), with description of three new species, pp. 388-412 in Zootaxa 3847 (3) on pages 404-406, DOI: 10.11646/zootaxa.3847.3.4, http://zenodo.org/record/226283, {"references":["Linkem, C. W., Diesmos, A. C. & Brown, R. M. (2011) Molecular systematics of the Philippine forest skinks (Squamata: Scincidae: Sphenomorphus): testing morphological hypotheses of interspecific relationships. Zoological Journal of the Linnean Society, 163, 1217 - 1243. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.2011.00747. x","Brown, R. M., Siler, C. D., Oliveros, C. H., Welton, L. J., Rock, A., Swab, J., Van Weerd, M., van Beijnen, J., Jose, E., Rodriguez, D., Jose, E. & Diesmos, A. C. (2013 b) The amphibians and reptiles of Luzon Island, Philippines, VIII: the herpetofauna of Cagayan and Isabela Provinces, northern Sierra Madre Mountain Range. Zookeys, 266, 1 - 120. http: // dx. doi. org / 10.3897 / zookeys. 266.3982"]}

Parvoscincus manananggalae sp. nov. (Figs. 3���5) Sphenomorphus leucospilos: Linkem, Diesmos, Brown, 2011 (part); Linkem & Brown, 2013 (part); Brown et al., 2000; Siler et al., 2011 d. Holotype. PNM 9794 (RMB Field No. 10719, formerly KU 323928), adult male, collected on rocks near a rapid flowing stream during the day on 21 June 2009, in Barangay Lipimental, Municipality of San Luis, Aurora Province, Luzon Island, Philippines (N: 15.653; E: 121.507; WGS- 84; 515 m in elevation), by RMB, CDS, L. Welton. Paratypes. KU 323920 ���27, 323929��� 30 collected 21 June 2009; CMNH 5792 (from Brown et al. 2000). Diagnosis. Parvoscincus manananggalae can be distinguished from congeners by the following combination of characters: (1) body size medium (SVL 47.3���55.9 mm); (2) Toe-IV lamellae 17; (3) supralabials seven; (4) infralabials six or seven; (5) midbody scale rows 32���33; (6) paravertebral scale rows 61���69; (7) prefrontals separated; (8) prefrontals contact first supraocular; (9) frontoparietals fused; (10) head pigmentation moderately mottled; (11) upper arm pigmentation present, patchy; (12) cloacal scale dark pigmentation present; (13) subcaudal pigmentation present; (14) dorsal white spots large, well-defined; (15) dorsal white bands 9���12; (16) bright lateral body coloration absent; (17) tail dorsolaterally compressed; and (18) semi-aquatic (Tables 2, 3). Comparisons. Characters distinguishing Parvoscincus manananggalae from all species of Parvoscincus are summarized in Tables 2 and 3. Parvoscincus manananggalae most closely resembles P. duwendorum, P. leucospilos, and P. tikbalangi. However, P. manananggalae differs from these three taxa by the presence of subcaudal dark pigmentation (vs. absence). Parvoscincus manananggalae further differs from P. duwendorum and P. tikbalangi by having dorsal white spots large and well-defined (vs. faint), Toe-IV lamellae 17 (vs. 12 [P. duwendorum], 14���16 [P. tikbalangi]); from P. leucospilos and P. tikbalangi by the absence of bright lateral coloration (vs. presence and bright reddish-orange [P. leucospilos], presence, reduced, tan [P. tikbalangi]); from P. duwendorum by having a greater number of paravertebral scale rows (61���69 vs. 60), a greater number of longitudinal ventral scale rows (43���49 vs. 41), a greater number of midbody scale rows (32���33 vs. 26), dorsal white bands 9���12 (vs. 15), and the presence of cloacal scale dark pigmentation (vs. absence); from P. leucospilos by having prefrontals separated (vs. in medial contact), head pigmentation moderately mottled (vs. heavily mottled), and the presence of dark pigmentation on the upper arm surface (vs. absence); and from P. tikbalangi by having a tendency towards a greater number of midbody scale rows (32���33 vs. 28���32) and fewer paravertebral scale rows (61���69 vs. 58���63), and the presence of contact between prefrontal and first supraocular scales (vs. absence). Description of holotype. Details of the head scalation are shown in Figure 4. An adult male Parvoscincus, SVL 55.9 mm, with clawed, pentadactyl limbs; head distinct from neck, characterized by enlarged jaw adductor musculature in temporal region. Snout pointed in dorsal aspect, rounded in lateral aspect; rostral wide forming a nearly perpendicular margin with frontonasal, rounded margin with nasals; frontonasal equally wide as long, in contact with nasals, rostral, anterior loreal, and prefrontal scales; prefrontals in point medial contact, in contact with anterior and posterior loreals, frontal, frontonasal, first supraciliary, and first supraocular; frontal greatly longer than wide, in contact with two supraoculars on right, two on left, rounded anteriorly, rounded posteriorly; four enlarged supraoculars, first largest; single, large frontoparietal, in contact with supraoculars II���IV; interparietal arrowhead-shaped; parietals in narrow medial contact, in narrow contact with fourth supraocular, in moderate contact with postsupraocular, and secondary temporal; primary temporals two, ventral largest, overlapping dorsal; secondary temporals two, large, dorsal largest, ventral overlapping dorsal; tertiary temporals two, dorsal largest, ventral overlapping dorsal; auricular opening large. Nasal pierced in center by large naris, surrounded anteriorly by rostral, dorsally by frontonasal, posteriorly by anterior loreal, and ventrally by first and second supralabial; anterior loreal one, narrow, sharply curved contact with nasal, posterior loreal lower in height, 2 x width of anterior loreal; preoculars two; supralabials seven, fifth subocular; lower eyelid scaly and semi-transparent, nonscaled ���window��� absent; ear large, moderately sunk. Infralabials seven, decreasing in size posteriorly in series; mental small, forming straight suture with single, large postmental and first infralabials; enlarged chin shields in three pairs; gular scales slightly smaller than ventrals. Body elongate, cylindrical, slender, with 32 equal sized midbody scales, limbs overlapping when adpressed; paravertebral scales 68, imbricate, smooth, without striations, keels or pits. Tail elongate, dorsolaterally compressed, longer than body (TL [67.0] / SVL [55.9] 1.20); subcaudal scales nondifferentiated for basal half of tail, enlarged for distal half of original tail. Precloacal region with series of enlarged scales between pelvic region and cloaca, more elongate than ventral scales; medial precloacal scales larger; left hemipene everted. Forelimbs smaller than hind limbs, pentadactyl; forelimb scales slightly smaller in size than body scales, imbricate and smooth, reducing slightly in size distally; lamellae becoming slightly keeled distally on each digit; relative digit length I Coloration of holotype in preservative. The dorsal background color is dark brown, with extensive mottling of several shades of brown. Three dorsal longitudinal rows of cream spots start posterior to the head and continue to the anteriormost portions of the tail, where the pattern becomes mottled more darkly brown. Lighter brown mottling is present in between the three dorsal rows of spots. The dorsal mottling pattern is more broken up with small cream spots on the lateral surface of the body, until eventually becoming a solid cream on the ventral side of the body. Similar small cream spots are present and sporadically distributed across the dorsal surface of both sets of limbs, continuing distal to the distinct large cream band on the forelimb, proximate to the body. The ventral surfaces of the feet are heavily mottled brown, except for Fingers I and II and Toe V. The solid cream ventral body coloration is broken up at the cloaca by four heavily pigmented enlarged precloacal scales. The ventral surface of the tail is mottled medium and dark brown, except for the area just posterior to the cloaca, which is solid cream in color. The mottled pattern on the dorsal surface of the head is a lighter brown than the dorsal surface of the body. This mottling is slightly darker around the orbits. There are two small cream spots just anterior to the eyes. There is a transition to a lighter mottled pattern on the lateral surface of the head starting just ventral to the eyes that continues to the venter. The supra- and infralabial series have four distinct tan spots, surrounded on either side by one indistinct spot. The ventral surface of the head is solid cream. Coloration of holotype in life (Differences from preserved specimens; Fig. 5). The cream mottling and indistinct cream spots present on the lateroventral surface of the head, body, and tail is replaced by light pale bluish-gray coloration. Bright lateral coloration is absent in this species. Measurements and scale counts of holotype in mm. SVL 55.9; AGD 29.3; TotL 122.9; TL 67.0; HL 12.2; HW 7.3; SnFa 18.1; ED 2.7; SNL 4.2; IND 2.1; FLL 5.5; HLL 6.6; MBSR 32; PVSR 68; ToeIVlam 17; SL 7; IFL 7; SO 4. Variation. Scale pigmentation was observed to vary among the examined series: dark cloacal scale pigmentation was present (KU 323920, 323922, 323923, 323925, 323926, 323928���30, 325810) or absent (KU 323921, 323924, 323927, 325807, 328808, 325811��� 13). Distribution, ecology and natural history. Parvoscincus manananggalae is known only from east-central Luzon Island, in Aurora Province near the Barangay of L. Pimentel in the San Luis Mountains and from Mt. Palali in Nueva Vizcaya Province (Fig. 1). This semi-aquatic species occurs in primary- and secondary-growth forest habitats, and occurs in sympatry with Parvoscincus agtorum, P. palaliensis, P. steerei, and P. tagapayo. We have evaluated this species against the IUCN criteria for classification, and find that it does not qualify for Critically Endangered, Endangered, Vulnerable, or Near Threatened status. Parvoscincus manananggalae has a relatively broad geographic distribution on Luzon Island and is quite abundant at all sampled localities. We therefore classify this species as Least Concern LC (IUCN, 2010). Etymology. The specific epithet is a feminine noun, formed from the name ���Manananggal,��� a female, blood sucking, vampire-like creature who flies like a bat at night to hunt humans, after separating from her lower extremities (derived from the Tagalog word Tanggal, to separate). Manananggal can be repelled by garlic and even killed by heavily salting her legs once she has left to hunt for the night. Suggested common name:Aurora Aquatic Skink., Published as part of Siler, Cameron D., Linkem, Charles W., Cobb, Kerry, Watters, Jessa L., Cummings, Sean T., Diesmos, Arvin C. & Brown, Rafe M., 2014, Taxonomic revision of the semi-aquatic skink Parvoscincus leucospilos (Reptilia: Squamata: Scincidae), with description of three new species, pp. 388-412 in Zootaxa 3847 (3) on pages 402-404, DOI: 10.11646/zootaxa.3847.3.4, http://zenodo.org/record/226283, {"references":["Linkem, C. W., Diesmos, A. C. & Brown, R. M. (2011) Molecular systematics of the Philippine forest skinks (Squamata: Scincidae: Sphenomorphus): testing morphological hypotheses of interspecific relationships. Zoological Journal of the Linnean Society, 163, 1217 - 1243. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.2011.00747. x","Brown, R. M., McGuire, J. A., Ferner, J. W., Icarangal, N. Jr. & Kennedy, R. S. (2000) Amphibians and reptiles of Luzon Island, II: preliminary report on the herpetofauna of Aurora Memorial National Park, Philippines. Hamadryad, 25, 175 - 195.","Siler, C. D., Welton, L. J., Siler, J. M., Brown, J., Bucol, A., Diesmos, A. C. & Brown, R. M. (2011 d) Amphibians and Reptiles, Luzon Island, Aurora Province and Aurora Memorial National Park, Northern Philippines, New island distribution records. Check List, 7, 182 - 195."]}

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."]}

Parvoscincus duwendorum sp. nov. (Figs. 3���5) Parvoscincus leucospilos Linkem, Diesmos, Brown, 2011 (part); Linkem & Brown, 2013 (part); Brown et al. 2012. Holotype. PNM 9793 (RMB Field No. 14261, formerly KU 329929), subadult, collected on 18 June 2011, on Mt. Pao, Barangay Adams, Municipality of Adams, Ilocos Norte Province, Luzon Island, Philippines (N: 18.438 ��, E: 120.878 ��; WGS- 84; 750 m in elevation), by RMB. Diagnosis. Parvoscincus duwendorum can be distinguished from congeners by the following combination of characters: (1) adult body size presumed medium (subadult SVL 33.5 mm); (2) Toe-IV lamellae 12; (3) supralabials seven; (4) infralabials eight; (5) midbody scale rows 26; (6) paravertebral scale rows 60; (7) prefrontals separated; (8) prefrontals contact first supraocular; (9) frontoparietals fused; (10) head pigmentation moderately mottled; (11) upper arm pigmentation present, patchy; (12) cloacal scale dark pigmentation absent; (13) subcaudal pigmentation absent; (14) dorsal white spots faint; (15) dorsal white bands 15; (16) tail dorsolaterally compressed; and (17) semi-aquatic (Tables 2, 3). Comparisons. Characters distinguishing Parvoscincus duwendorum from all species of Parvoscincus are summarized in Tables 2 and 3. Parvoscincus duwendorum most closely resembles P. manananggalae, P. leucospilos, and P. tikbalangi. However, P. duwendorum differs from these three taxa by having Toe-IV lamellae 12 (vs. 17 [P. manananggalae], 15���17 [P. leucospilos], 14���16 [P. tikbalangi]), fewer midbody scale rows (26 vs. 32���33 [P. manananggalae], 30���34 [P. leucospilos], 28���32 [P. tikbalangi]), a greater number of dorsal white spot rows (15 vs. 9���13 [P. leucospilos], 9���12 [P. manananggalae, P. tikbalangi]), and the absence of precloacal scale dark pigmentation (vs. absence or presence [P. leucospilos, P. tikbalangi], presence [P. manananggalae]). Parvoscincus duwendorum further differs from P. leucospilos and P. manananggalae by having fewer paravertebral scale rows (60 vs. 61���67 [P. leucospilos], 61���69 [P. manananggalae]); from P. manananggalae and P. tikbalangi by having infralabials eight (vs. six or seven [P. manananggalae], seven [P. tikbalangi]); from P. leucospilos by having head pigmentation moderately mottled (vs. heavily mottled), and the presence of dark pigmentation on the upper arm surface (vs. absence); from P. manananggalae by having fewer longitudinal ventral scale rows (41 vs. 43���49), and the absence (vs. presence) of subcaudal dark pigmentation; from P. leucospilos and P. tikbalangi by having prefrontals separated (vs. in medial contact [P. leucospilos], separated or in medial contact [P. tikbalangi]); and from P. tikbalangi by having prefrontals and first supraoculars in contact (vs. separated). TABLE 2. Distribution of diagnostic characters (+ present; 0 absent) among species in the genus Parvoscincus. Parvoscincus palawanensis does not have prefrontal scales. (A) Brown et al. 2010, ) Linkem and Brown, 2013, (C) Brown and Alcala 1980, (D) Ferner et al. 1997. Characters included for comparison are: snout���vent length (SVL), midbody scale-row count (MBSR), paravertebral scale-row count (PVSR), Toe-IV lamellae count (ToeIVlam), supralabial count (SL), infralabial count (IFL), number of anterior loreals, contact between profrontal scales, and microhabitat preference. leucospilos duwendorum manananggalae tikbalangi (6 m 1, 5 f) (1 juv.) (3 m, 2 f) (7 m, 1 f) Paralectotype (CAS 64232) presumed male following visual inspection of external body; authors avoided creating incision on rare type specimen. Description of holotype. Details of the head scalation are shown in Figure 4. A subadult Parvoscincus, SVL 33.5 mm, with clawed, pentadactyl limbs. Head distinct from neck, characterized by enlarged jaw adductor musculature in temporal region. Snout sharply pointed in dorsal aspect, rounded in lateral aspect; rostral wide forming a nearly perpendicular margin with nasals and frontonasal; frontonasal equally wide as long, in contact with nasals, rostral, anterior loreal, and prefrontal scales; prefrontals broadly separated, in contact with anterior and posterior loreals, frontal, frontonasal, first supraciliary, and first supraocular; frontal greatly longer than wide, in contact with two supraoculars on right, two on left, rounded anteriorly, sharply rounded posteriorly; four enlarged supraoculars, first largest; single, large frontoparietals, in contact with supraoculars II���IV; interparietal arrowhead; parietals in moderate medial contact, in contact with fourth supraocular, postsupraocular, and secondary temporal; primary temporals two, ventral largest, overlapping dorsal; secondary temporals two, large, dorsal largest, ventral overlapping dorsal; tertiary temporals two, dorsal largest, ventral overlapping dorsal; auricular opening large. Nasal pierced in center by large naris, surrounded anteriorly by rostral, dorsally by frontonasal, posteriorly by anterior loreal, and ventrally by first supralabial; anterior loreal one, posterior loreal equal in size; preoculars two; supralabials seven, fifth subocular; lower eyelid scaly and semi-transparent, nonscaled ���window��� absent; ear large, moderately sunk. Infralabials eight, decreasing in size posteriorly in series; mental small, forming straight suture with single, large postmental and first infralabials; enlarged chin shields in three pairs; gular scales slightly smaller than ventrals. Body elongate, cylindrical, slender, with 26 equal sized midbody scales, limbs overlapping when adpressed; paravertebral scales 60, imbricate, smooth, without striations, keels or pits. Tail elongate, dorsolaterally compressed, longer than body (TL [50.0] / SVL [33.5] 1.49); subcaudal scales nondifferentiated for basal half of tail, enlarged for distal half of original tail. Precloacal region with series of enlarged scales between pelvic region and cloaca, more elongate than ventral scales; medial precloacal scales larger. FIGURE 6. Illustration of head of the examined Parvoscincus leucospilos paralectotype (CAS 64232) in dorsal and lateral views. Taxonomically diagnostic head scales are labeled as follows: C, chin shield; F, frontal; FN, frontonasal; FP, frontoparietal; IL, infralabial; IP, interparietal; L, loreal; M, mental; N, nasal; P, parietal; PF, prefrontal; PM, postmental; PO, preocular; PSO, presubocular; PoSO, postsupraoculars; R, rostral; SC, supraciliary; SL, supralabial; SO, supraocular; T 1, primary temporal; T 2, secondary temporal; and T 3, tertiary temporal. Roman numerals indicate scales in the supraocular series. Illustrations by STC and CDS. Forelimbs smaller than hind limbs, pentadactyl; forelimb scales slightly smaller in size than body scales, imbricate and smooth, reducing slightly in size closer to manus; lamellae becoming slightly keeled distally on each digit; relative digit length with lamellae V Coloration of holotype in preservative. The dorsal background color is medium brown, with tan mottling throughout. Two parallel light brown stripes run longitudinally down the body, starting at the base of the head and merging to one just past the start of the tail. Between these two stripes is a paravertebral row of darker brown spots. The venter color is a solid cream, without mottling. Along the lateral surface of the body, the dorsal and ventral color patterns merges in a scalloped pattern. There is a distinct cream spot on the forelimbs, just proximate to the body. The dorsal fore- and hind limbs possess the same mottling pattern present on the dorsal surface of the body. The ventral surfaces of the limbs are a solid cream color. The dorsal surface of the hands and feet are a mottled medium brown, with the exception of the cream-colored Finger I and Finger II on the forelimbs and Toe V on the hind limbs. The dorsal and dorsolateral surfaces of the head have a light and medium brown mottled pattern. This coloration transitions sharply into a cream and light brown mottled pattern ventrally. The labial scales are mottled light brown and cream. The ventral surface of the head is solid cream in color. Color in life ���Coloration in life did not differ substantially from the preserved holotype (RMB, personal observations). Measurements and scale counts of holotype in mm. SVL 33.5; AGD 17.5; TotL 83.5; TL 50.0; HL 4.0; HW 7.7; SnFa 12.4; ED 1.5; SNL 2.7; IND 1.3; FLL 3.3; HLL 4.0; MBSR 26; PVSR 60; ToeIVlam 12; SL 7; IFL 8; SO 4. Distribution, ecology and natural history. Parvoscincus duwendorum is known from a single specimen found at 750 elevation on Mt. Pao in the Ilocos Norte Province of northwestern Luzon Island (Fig. 1). Although the species is recognized currently to occur in secondary-growth forest habitats, it is presumed the species also inhabits suitable primary forest. This species is semi-aquatic and can be found near riparian habitat. Parvoscincus duwendorum can be found in sympatry with Parvoscincus igorotorum, P. jimmymcguirei, and P. steerei. The new species was also observed on Mt. Cagua at 300���500 m (Brown et al 2013 b) but specimens eluded biologists by jumping in rapidly running montane streams. At this time we are unable to appropriately evaluate this species against the IUCN criteria for classification due to the lack of available information about its distribution and natural history. We therefore classify this species Data Deficient DD (IUCN, 2010) pending future studies on this unique semi-aquatic forest skink. Etymology. The specific epithet is a plaural derivation of the Filipino folklore term ���Duwende,��� which is chosen here to celebrate the country���s rich tradition of mythological forest animals and spirits. Duwendes are believed to be little fairy-like forest creatures, such as goblins, pixies, and elves, and believed to live in trees, termite mounds, and burrows in hillsides. The may bring bad or good fortune to humans and are often considered to be mischievous in nature. Suggested common name: Cordillera Aquatic Skink., Published as part of Siler, Cameron D., Linkem, Charles W., Cobb, Kerry, Watters, Jessa L., Cummings, Sean T., Diesmos, Arvin C. & Brown, Rafe M., 2014, Taxonomic revision of the semi-aquatic skink Parvoscincus leucospilos (Reptilia: Squamata: Scincidae), with description of three new species, pp. 388-412 in Zootaxa 3847 (3) on pages 397-402, DOI: 10.11646/zootaxa.3847.3.4, http://zenodo.org/record/226283, {"references":["Linkem, C. W., Diesmos, A. C. & Brown, R. M. (2011) Molecular systematics of the Philippine forest skinks (Squamata: Scincidae: Sphenomorphus): testing morphological hypotheses of interspecific relationships. Zoological Journal of the Linnean Society, 163, 1217 - 1243. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.2011.00747. x","Brown, W. C. & Alcala, A. C. (1980) Philippine Lizards of the Family Scincidae. Silliman University Press, Dumaguete City, Philippines, 264 pp.","Brown, R. M., Siler, C. D., Oliveros, C. H., Welton, L. J., Rock, A., Swab, J., Van Weerd, M., van Beijnen, J., Jose, E., Rodriguez, D., Jose, E. & Diesmos, A. C. (2013 b) The amphibians and reptiles of Luzon Island, Philippines, VIII: the herpetofauna of Cagayan and Isabela Provinces, northern Sierra Madre Mountain Range. Zookeys, 266, 1 - 120. http: // dx. doi. org / 10.3897 / zookeys. 266.3982"]}