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The lacertid Latastia ornata was known to date only by its holotype collected in 1938 in Bafatá, central Guinea-Bissau. We report new specimens and localities from Guinea-Conakry, a new country record and major range extension of 700 km SE of the type-locality. We provide an updated diagnosis of the species, including the first genetic and osteological data, and confirm that Latastia ornata is closely related to, but distinct from, L. longicaudata based on external morphology, cranial osteology, DNA data and zoogeography.

We describe a new species of Lepidodactylus from Umboi Island, just to the west of New Britain. It is a member of the Lepidodactylus guppyi Group and can be distinguished from all other Melanesian Lepidodactylus by aspects of digital scalation, digital webbing, enlarged femoral/precloacal scales, and color pattern. It is genetically distinct from its closest congeners, and genetic and morphological data indicate that the new species is most similar among named species to Lepidodactylus guppyi from the Solomon Islands, but it diverged from this species and other close relatives approximately 8 MYA or longer at a time prior to the existence of the island that it now occupies. The new species is known from only three individuals collected on a single tree, and efforts to find more animals in what seemed good habitat nearby were unsuccessful. This duplicates the pattern of apparent rarity seen for many Lepidodactylus species. Sufficient habitat exists on Umboi Island for arboreal geckos, suggesting that the species is not actually endangered but is ecologically cryptic. However, lack of needed information leads us to assess this species’ conservation status as Data Deficient.

Tropidonotus nicobarensis Sclater, 1891 is an enigmatic snake currently referred to the natricine genus Hebius Thompson, 1913. No specimens have been recorded since the original discovery of the name-bearing type in the late 19th century, which was allegedly collected on Kamorta Island in India’s Nicobar Archipelago. Recently, a second observation of this species was reported from Havelock Island (Swarajdweep) in the nearby Andaman Archipelago. However, the snake in question is clearly conspecific with another natricine, Thamnophis saurita (Linnaeus, 1766), native to eastern North America. This discovery prompted us to examine the type specimen of Tropidonotus nicobarensis, which revealed that it too is conspecific with Thamnophis saurita. The provenance of the Tropidonotus nicobarensis type specimen and the Havelock Island snake are discussed, with the latter likely representing an introduced animal.

We describe a new species of Selvasaura lizard from the western slopes of the eastern Andes of central Peru. Among other characters, the new species differs from congeners in having keeled dorsal scales and more transverse rows of scales on dorsum. We present a phylogeny as additional evidence supporting delimitation of the new species.

The analyses of molecular genetic data (mtDNA markers 16S, ND4, CYTB, and the nuclear marker c-mos) provided evidence that the Asian cat snake taxa Boiga multomaculata and B. ochracea actually represent a single species. They form mixed clades of low intraclade genetic differentiation. This evidence for conspecificy is supported by the lack of differentiation in all examined pholidotic and morphometric characters. Therefore, we formally place Dipsas ochracea Theobald, 1868 in the synonymy of Dipsas multomaculata Boie, 1827. We provide a summary of the tangled taxonomic history of the taxa involved in this study. Also, we resurrect Dipsadomorphus stoliczkae Wall, 1909 from synonymy of B. ochracea, for specimens exhibiting 21 midbody dorsal scale rows. Boiga stoliczkae is found in the Himalayas north and west of the Brahmaputra valley. Finally, based on the detection of historical genetic lineages within the newly defined species Boiga multomaculata we recognize three subspecies: Boiga multomaculata multomaculata (Boie, 1827), Boiga multomaculata ochracea (Theobald, 1868), and Boiga multomaculata septentrionalis n. ssp. which is distributed in northern Myanmar and Assam and Nagaland, India. We designate BMNH 1946.1.2.60 (1) as neotype of Dipsas ochracea Theobald, (2) as lectotype of D. ochraceus Günther, and (3) as lectotype of Boiga ochracea walli Smith, thereby making these names objective synonyms. Finally, we designate BMNH 94.12.31.55 as lectotype of Dipsadomorphus stoliczkae Wall.

We describe Liolaemus attenboroughi sp. nov., a lizard distributed in the northwestern Patagonian Steppe of Chubut province (Argentina) previously confused with L. kingii (Bell 1843). Recent studies based on molecular evidence supports its evolutionary independence. Here we provide a morphological diagnosis of this lineage, comparisons between three molecular species delimitation methods, and an updated phylogeny of the L. kingii group. Based on current knowledge of its distribution, this new species is allopatric with geographically close species of the L. kingii group.

Snakes are ecologically and morphologically plastic organisms that exhibit extensive variations in body size and body condition in response to environmental factors. Documenting inter-population variations is important to describe species comprehensively across their distribution range and to monitor trends over time (e.g. decreasing body condition due to alteration of habitat). Thus, we analyzed the influence of population and sex on body size and body condition in three populations of nose-horned vipers (Vipera ammodytes) in Serbia. In one population, males were larger than females (F1, 39=4.802, p=0.034), but not in the two other populations (F1, 36=0.075, p=0.786; F1, 21=0.018, p=0.893). Females exhibited higher body condition (residual values from the regression of log-body mass against log-body size) than males (F1, 90=10.444, p=0.002); this sex difference was not found in one population when analyzed separately (F1, 35=1.834, p=0.184). Moreover, we found strong inter-population differences in mean body size and mean body condition (F2, 96=8.822, pF2, 90=10.319, p=0.001, respectively). While inter-population difference in body size was driven by males, inter-population difference in body condition was driven by females. These results suggest that, in this species, body size might be an important determinant of mating success in males, while body condition may play a major role in female fecundity.

The present study is aimed to validate the occurrence of two species of Diadem or Royal snakes of Genus Spalerosophis Jan, 1865 from the Poonch District of Jammu and Kashmir, India along with the presentation of eight different colour morphs and diagnostic characteristics. The eight different colour morphs include four colour morphs of adults of S. d. diadema (Schlegel, 1837), two colour morphs of adults of S. atriceps (Fischer, 1885), one colour morph of subadults of unknown parentage and one colour morph of the adult of S. diadema (Schlegel, 1837) represented by a single individual whose identification at subspecies level (ssp. cliffordii/diadema) is the subject for further investigations for want of more specimens. This solitary specimen shares many similarities with S. d. diadema, but it has less number of sub-caudal scales which point towards the likelihood of S. d. cliffordii (Schlegel, 1837). The distribution, activity, habitat and behaviour have also been reported along with the morphological, morphometric and meristic characters. Between two identified species S. atriceps is a new report from the Poonch District. Reported specimens are mapped across the study area and are depicted here in the distribution map.

Species of the agamid genus Bronchocela Kaup, 1827 in the Andaman and Nicobar Islands are reassessed based on newly collected specimens from different parts of the Nicobar Archipelago. An assessment based on morphology and distribution of the identified groups reveal two new, unnamed populations, one allied to B. cristatella and the other allied to B. danieli. These two populations are described as new species, and Bronchocela cristatella is redescribed based on Sundaic specimens. The remaining species, B. danieli and B. rubrigularis are redescribed. Distributions of all of these four species are mapped and recommendations on their conservation status are suggested.

The pit viper snake genus Trimeresurus Lacépède, 1804 sensu lato, is a diverse group of nocturnal serpents comprising over 61 species. The genus is morphologically heterogeneous and has been divided into several subgenera. We present an updated phylogeny of Asian pitvipers and propose a revised classification. Additionally, we revise the taxonomy of T. popeiorum Smith, 1937 and propose taxonomic changes with support from molecular and morphological data. We restrict T. popeiorumsensu stricto to northeastern India, Bangladesh, southern China, and northern Myanmar; populations beyond these areas require further assessment. We also synonymize T. yingjiangensisChen et al., 2019 with T. popeiorum based on overlapping morphological characters, molecular data, and distribution. The findings shed new light on the taxonomy of T. popeiorum, warranting the need for assessing the population of T. popeiorum from southeast Asia.

Wagler described Natrix sexcarinata through a specimen collected on the banks of the Amazon River in Brazil, without citing a specific location. The species was later transferred to different Neotropical genera (Chironius, Herpetodryas, Phrynonax, and Pseustes) currently allocated in the Family Colubridae. The taxonomic instability of Natrix sexcarinata is probably due to its brief original description in combination with its supposedly inaccurate illustration. Despite the holotype being lost during World War II, some authors have pointed out that its description is somewhat similar to the widespread concept of Pseustes poecilonotus. More recently, a molecular study proposed the synonymy of the genera Pseustes with Spilotes, resurrecting the Phrynonax genus to allocate Ph. poecilonotus and Ph. polylepis. However, the taxonomic positioning of Natrix sexcarinata remains unsatisfactory with respect to the cis-Andean taxa placed in Phrynonax or even Spilotes throughout the Amazon. Considering the taxonomic and nomenclatural problems mentioned above, we investigated how many Phrynonax species occur along the Amazon Basin and what would be their applicable name. We examined the external morphology (meristic, morphometric, and color pattern data) of 118 Phrynonax specimens and prepared the hemipenes of seven specimens to understand geographic, sexual, and ontogenetic variability parameters. Our result suggests the synonymy of Ph. polylepis and N. sexcarinata under the combination Phrynonax sexcarinatus. Based on available evidence, we reinforce that Phrynonax sexcarinatus is the only congener that occurs along the Amazon Basin. We have also designated a Natrix sexcarinata neotype to stabilize its complex nomenclature.

The Anolis heterodermus group comprises eight big-headed and short-legged lizard species from the highlands of the northernmost South American Andes. Recent studies revealed unknown lineages within this group that had previously been categorized as a species complex. By widely sampling and applying an integrative taxonomic framework, we (1) assessed the species diversity of the group using a molecular dataset (two mitochondrial and one nuclear markers) along with an inclusive morphological study (scalation, scale configuration and ornamentation, morphometrics, and dewlap and body colour patterns); and (2) we inferred the evolutionary relationships within this species group. Our analyses confirmed the formerly reported differentiation between populations of those high-altitude lizards, and we identified several unknown evolutionary lineages. Our results provided evidence for the existence of nine distinct, independently evolving evolutionary lineages in the heterodermus group. As a result, we described two morphologically and genetically highly distinct lineages as species new to science (A. quimbayasp. nov. and A. tequendamasp. nov.). We redescribed A. heterodermus and erected as a valid species Anolis richteri, a previously described synonym of A. heterodermus. A taxonomic key for the identification of species of the Phenacosaurus clade was presented. The identification of two additional poorly-known lineages suggested that the diversity of this group of lizards is still unknown; therefore, it is necessary to establish measurements for the group´s conservation, as well as to perform fieldwork and revision of herpetological collections to identify possible hidden diversity within the group.

A new species of Liotyphlops Peters, 1881, Liotyphlops palauophissp. nov., is described from the neighborhoods of Bogota, Colombia from a previous syntype of L. anops, and a lectotype is designated for the latter species. The new species is readily distinguished from congeners by having the frontal scale divided (vs single), and a central foramen in the parabasisphenoid (vs foramen absent). High-resolution x-ray computed tomography (HRXCT) was used to study and present data on the skull of the holotype of the new species, the lectotype of L. anops, and the holotype of L. ternetzii. Additionally, extensive study of skull characters and external morphology failed to find diagnostic characters to differentiate L. beui and L. ternetzii, and the former is here considered a junior synonym of L. ternetzii, which is also redescribed.

A new species of Dipsas Laurenti, 1768, from Central Panama is described based on molecular analyses, hemipenial morphology, and external characters. This is the sixth species of Dipsas to be described for the country; the snake has been suspected to exist since 1977 and has not been thoroughly studied until now. Additionally, morphological comparations including scale counts are done with other species within the genus, and the current geographic distribution of Dipsas temporalis (Werner, 1909), the sister species, is updated. Finally, a key to the species of Dipsas currently known from Middle America is presented.

Colour polymorphism has been previously reported in several colubrid snakes including Boiga spp. In this paper, we report colour variations within the poorly known southern Indian Boiga dightoni, provide the first molecular data for this species, from two localities (including the type locality) and compare them with data from other congeners. Additionally, we provide detailed dentition and hemipenis descriptions for B. dightoni. Molecular data for B. dightoni show very little difference (0.2–0.4% 16S; 0.9–1.2% cyt b) to the recently described Boiga whitakeri, also from southern India. We have re-examined and present new information on the pholidosis of the type specimens of B. whitakeri and reconsider its taxonomic status. On the basis of molecular data and overlapping morphological characteristics, we argue that Boiga whitakeri and Boiga dightoni are conspecific, and place B. whitakeri under the subjective synonymy of the latter. Furthermore, we show that colour polymorphism in B. dightoni is a gender-independent character and that both colour morphs are found in high as well as low elevations and partly in sympatry. A revised key to the Boiga ceylonensis complex is provided.

Animal colouration has a significant ecological role in defence, reproduction, and thermoregulation. In the case of melanism, it is a complex topic. Besides potential disadvantages such as higher risk of predation, melanistic ectotherms may have certain physiological advantages such as more efficient thermoregulation in colder climates and thus, reduced basking time. The common grass snake (Natrix natrix) is a widespread species throughout Europe and Asia. It exhibits a wide range of colour polymorphisms, from olive to dark grey, even albinistic and melanistic. Between 2016 and 2021, we conducted fieldwork in the Danube Delta Biosphere Reserve (DDBR) with the aim to document the geographic range of melanistic grass snakes. We categorised the melanistic expressions of N. natrix individuals as melanotic, completely melanistic, and partially melanistic. Melanistic snakes were encountered in all six localities visited, suggesting that the occurrence of melanistic grass snakes in the DDBR is geographically widespread. We observed both juveniles (n=2) and adults (n=11) with melanism, suggesting that individuals are born melanistic. However, the proportion of melanistic individuals in the general population of N. natrix from the DDBR is unknown. Only at Histria locality we studied the proportion of melanism in the grass snake population and 6.3% of the snakes caught were melanistic. Body size comparisons are not statistically significant because of the low sample size. The N. natrix melanistic morph’s geographical distribution in the DDBR is most likely due to an interaction of climate and habitats, which offer a thermal advantage in the face of predation pressure.

Urban habitats receive an increasing number of species due to anthropogenic activities, mainly transportations. Here, we report a new addition to the herpetofauna of Athens (Greece): a small population of the Pelasgian wall lizard (Anatololacerta pelasgiana) was found in a suburb of the Athenian metropolitan area. The species normally occurs in southwestern Anatolia and southeastern Aegean islands and this is the first record in the Greek mainland. Allochthonous species that successfully colonize cities raise new challenges to urban ecology.

South Asian Cnemaspis are one of the most diverse clades of gekkonids in South Asia with their highest diversity in the Western Ghats and Sri Lanka. These geckos include only a few nocturnal species and are largely diurnal or cathemeral and restricted to relatively cool habitats. One of the prominently diurnal subgroups in South Asian Cnemaspis is the bangara clade, which includes six species distributed in southern India on the eastern slopes of the Western Ghats, the southern Eastern Ghats and Palghat Gap. In this paper, we describe five more species of the bangara clade from the Shevaroyan landscape, including three from Kollimalai and one each from Yercaud and Pachaimalai, all in Tamil Nadu. These new species show 4.6–19.7 % uncorrected sequence divergence on the mitochondrial ND2 gene from each other and known species of the bangara clade and are morphologically diagnosable in body size, the number of paravertebral tubercles between limb insertions, the number of dorsal tubercle rows, the number of ventral scale rows across the belly, the number of femoral and precloacal pores and poreless scales separating these series, and aspects of colouration. The discovery of these five new species adds to the growing discoveries of cool-adapted species in southern India outside the Western Ghats and highlights the role of sky-islands in diversification. The Shevaroyan landscape shows high levels of microendemism with eight species distributed in an area of < 2000 km2, and all these species restricted to much smaller areas of actual distribution. With an area of < 500 km2 respectively, the massif of Pachaimalai has a single endemic and the massifs of Yercaud and Kollimalai have three endemic Cnemaspis species each.

The gymnophthalmid Andean lizards of the genus Proctoporus (Gymnophthalmidae, Cercosaurinae) are semifossorial species that inhabit the montane forests, inter-Andean valley, and humid grasslands across the Cordillera de los Andes from Peru to Argentina. The distribution range of Proctoporus is discontinuous, with many places lacking herpetological surveys. Here, based on morphological data, we describe a new species of Proctoporus from a remote location on a mountaintop north of the Cordillera de Vilcabamba in southeastern Peru. The new species is readily distinguished from all other species of Proctoporus by the presence of three anterior infralabials, three rows of pregular scales and the absence of enlarged pregular scales. With this new species, the species diversity has increased to 19 since 18 species have previously been identified.

We present a comprehensive catalogue of the reptiles of Equatorial Guinea, consisting of 118 species belonging to 67 genera and 22 families. There are two species of Crocodylia, ten of Testudines and 106 of Squamata; this last taxon is represented by 62 species of snakes, two amphisbaenians and 42 lizards. Of these 118 species, seven are present only in Annobon, seven only in Bioko, 47 only in Río Muni, 53 occur both in Bioko and Río Muni (or Bioko, Río Muni and other islands), and four are sea turtles. Despite its high diversity, the level of endemism of Bioko is relatively low, with only four endemic species out of the 60 species reported for the island. In contrast, despite its low diversity, Annobon has the highest endemicity level, with five endemic species (and two introduced). No endemic species are known for the rest of the regions of Equatorial Guinea, which contain 100 species. We reveal several new country and species records, and point to some pending taxonomic questions to be addressed. Among the new species records, we highlight the presence of Cyclanorbis elegans, which represents the most meridional known population of the genus. Additional species are expected to be found in Equatorial Guinea as further field and taxonomic work is developed.

Bothrocophias microphthalmus (Cope, 1875) currently comprises most mid- to highland populations of the genus Bothrocophias in the eastern versant of the Andes. We describe two new species of Bothrocophias from the highlands of the Colombian Andes previously referred to as B. microphthalmus based on morphological and genetic evidence. Our phylogenetic analysis revealed that B. micropthalmus as currently recognized is paraphyletic with respect to B. hyoprora, and the two new taxa are sister lineages. These new toadheaded pitvipers can be morphologically distinguished from their congeners based on the presence of contact between the internasal scales, the number of prefoveal scales, the presence of a lacunolabial scale, the arrangement of supralabial scales, ventral scale counts, the color pattern of the dorsal and ventral surfaces of the body and tail, and hemipenial morphology. We discuss some possible taxonomic scenarios for the lineages found within the B. microphthalmus species complex but refrain from making additional taxonomic changes given our reduced sampling of the southern lineages.

Automated image classification is a thriving field of machine learning, and various successful applications dealing with biological images have recently emerged. In this work, we address the ability of these methods to identify species that are difficult to tell apart by humans due to their morphological similarity. We focus on distinguishing species of wall lizards, namely those belonging to the Podarcis hispanicus species complex, which constitutes a well-known example of cryptic morphological variation. We address two classification experiments: (1) assignment of images of the morphologically relatively distinct P. bocagei and P. lusitanicus; and (2) distinction between the overall more cryptic nine taxa that compose this complex. We used four datasets (two image perspectives and individuals of the two sexes) and three deep-learning models to address each problem. Our results suggest a high ability of the models to identify the correct species, especially when combining predictions from different perspectives and models (accuracy of 95.9% and 97.1% for females and males, respectively, in the two-class case; and of 91.2% to 93.5% for females and males, respectively, in the nine-class case). Overall, these results establish deep-learning models as an important tool for field identification and monitoring of cryptic species complexes, alleviating the burden of expert or genetic identification.

We describe a new species of kukri snake from Da Lat City and its vicinity in Lam Dong Province, southern Vietnam on the basis of morphological characters of four specimens, including two males and two females. Oligodon tuani sp. nov. differs from other congeners by a combination of the following characters: large size in adults (TL ≤ 888 mm); 19 dorsal scale rows at midbody; ventrals 173–179 in males and 187–193 in females; subcaudals 58 or 59 in males and 44 or 45 in females; presubocular present; 8 supralabials, fourth and fifth entering orbit; cloacal plate undivided; 10 maxillary teeth, posterior three enlarged; vertebral stripe present; temporal streak present but faint and interrupted; long and deeply forked hemipenes, extending to 25th subcaudal, without spines and papillae; and 13–15+3–4 dorsal blotches.

We describe the unusual case of erythrism in the Eastern Grass Snake, Natrix natrix. This colour morph is very rare and has not been reported in the literature before. Despite having observed thousands of N. natrix in the field, we personally detected this morph in only three individuals originating from Slovakia, Romania, and mainland Greece, while photos of a fourth individual from a Greek island were provided to us. In addition, a recent study with a large data set from citizen scientists was unable to produce a single reddish Eastern Grass Snake. Such colouration is likewise uncommon in the western members of Grass Snakes (N. helvetica, N. astreptophora), with two examples provided herein. Because the potential biological importance of erythristic colouration is unclear, we encourage other field herpetologists and naturalists to publish their observations of reddish Grass Snakes in the printed literature.

Podarcis wall lizards mainly feed on coleopterans, orthopterans, arachnids, and other small invertebrates. However, Aegean wall lizards, Podarcis erhardii, are widely distributed across Aegean islands and are increasingly observed eating non-traditional food items ranging from plant material to conspecific eggs and body parts. Here, we report the first documented case of P. erhardii consuming a large centipede, Scolopendra cingulata. The predator-prey relationship between these species has appeared to go both ways and may intensify on islands.

The genera Amauta Houlbert, 1918 and Divana J.Y. Miller, 1982 are revised, with discussion of diagnostic features of males and females of all taxa. Details on their nomenclatural history, biogeography, and biology are included to solve several nomenclatural issues. Lectotypes are designated for Castnia (Amauta) papilionaris affinis Rothschild, 1919 and Castnia tricolor C. Felder & R. Felder, 1874. The status of the following taxa is revised: Amauta procera (Boisduval, [1875]) stat. rest., Amauta angusta (H. Druce, 1907) stat. rest., Castnia ambatensis Houlbert, 1917 syn. nov. of Castnia papilionaris papilionaris Walker, [1865], Castnia velutina Houlbert, 1917 syn. nov. of Castnia papilionaris papilionaris Walker, [1865], and Castnia diva chiriquiensis Strand, 1913 syn. nov. of Castnia diva diva Butler, 1870. Other taxa are revised, and their taxonomic status clarified. A new genus Vadina gen. nov. is proposed for Castnia hodeei Oberthür, 1881, which is removed (comb. nov.) from Telchin Hübner, [1825], whilst Amauta stat. rest. and Divana stat. rest., subsumed into Telchin in the most recent revision of the family, are herein reinstated as valid genera.

The genus Heliobolus comprises four recognized species, all endemic to sub-Saharan Africa. Of these, only Heliobolus lugubris occurs in southern Africa, its distribution extending from Angola in the west to Mozambique in the east and reaching as far south as parts of northern South Africa. Like many of the reptile species that occur in southern Africa, Heliobolus lugubris is poorly studied, and preliminary investigation suggested that it may contain cryptic diversity. The present work focusses on the Angolan population of H. lugubris and uses an integrative taxonomic approach based on morphological, coloration and DNA sequence data. The results indicate that some of the current and historical specimens of H. lugubris from Angola do not correspond to the nominotypical form, and that differences between specimens suggest the presence of two additional species, described here as Heliobolus bivarisp. nov. from the southernmost xeric/desertic regions and plateau of Namibe Province, southwestern Angola and H. crawfordisp. nov. from the Serra da Neve inselberg north through the sub-desert coastal regions of northern Namibe, Benguela, and Kwanza Sul provinces. Nominotypical Heliobolus lugubris is confirmed to occur in Cuando Cubango Province, southeastern Angola.

Anolis riparius, new species Holotype. UCR 5579 (Fig 6) collected on 7 September 2009 by Douglas Robinson and his students; an adult male from Pacuarito river, Pacuarito 10 Km S-SW of Pérez Zeledón, 9.31390 N, - 83.77220 W, 880 masl, San José Province, Costa Rica. Paratypes (all from Costa Rica: San José Province). Adult males: SMF 82183–82184 (collected 9 October 2002 by Axel Fläschendräger) from road between Platanillo and San Isidro, 9.31056 N; - 83.77500 W, ca. 850 masl; SMF 92428 (collected 15 February 2010 by Gunther Köhler) from 1 km N Cedral 9.35789 N, - 83.56094 W, 1425 masl; UCR 7893 (collected 24 August 1979 by Douglas Robinson) from I.85 Km NE Alfombra, 9.3167 N, - 83.7722 W, ca. 930 masl; MCZ-R 18171 (collected 15 December 2007 by Steve Poe and Mason J. Ryan) from Alfombra, 12 Km S-SW of Pérez Zeledón (9.31670 N, - 83.77220 W, 930 masl); MSB 95201 (collected on 10 January 2009 by Mason J Ryan and Ian Latella) from Alfombra, 12 Km S-SW of Pérez Zeledón (9.31670 N, - 83.77220 W, 930 masl). Adult females: SMF 82185–82186 (collected 9 October 2002 by Axel Fläschendräger) from road between Platanillo and San Isidro (ca. 850 masl); SMF 83093–83094 (collected 9 October 2002 by Axel Fläschendräger) from near Platanillo (ca. 850 masl); SMF 92429 (collected 15 February 2010 by Gunther Köhler) from 1 km N Cedral (9.35789 N, - 83,56094 W, 1425 masl); UCR 5580 (same data of the holotype); UCR 15959 (collected 3 March 2001 by Mason J Ryan) from Mollejones (9.22930 N, - 83.64200 W, 800 masl); UCR 16162 (collected 2 July 2001 by Mason J Ryan) from Alfombra, 12 Km S-SW of Pérez Zeledón (9.31670 N, - 83.77220 W, 930 masl); MCZ R-186143 (collected 15 December 2007 by Steve Poe and Mason J Ryan) from Alfombra, 12 Km S-SW of Pérez Zeledón (9.31670 N, - 83.77220 W, 930 masl); MSB 95200 (collected 15 December 2007 by Steve Poe and Mason J Ryan) from Alfombra, 12 Km S-SW of Pérez Zeledón (9.31670 N, - 83.77220 W, 930 masl); UCR 16638 (collected 15 March 2002 by Franklin Aguilar) from Fila Piedras Blancas area (9.50130 N, - 83.91280 W, 900 masl) in the Zona Protectora Los Santos in the Río Savegre basin. Diagnosis. Anolis riparius sp. nov. and A. aquaticus are the only semiaquatic Middle American anoles that have a large orange-red dewlap with yellow in males (Fig. 6B). Anolis riparius sp. nov. differs from A. aquaticus in possessing smaller dorsal scales of the head and body (e.g., 15–19 scales across the snout between the second canthals in A. riparius sp. nov. versus 7–14 in A. aquaticus, Fig. 5B), and is phylogenetically distinct (see below). Anolis riparius sp. nov. is distinguished from the other Middle American semiaquatic anoles as follows: from A. robinsoni sp. nov. by male dewlap color (chocolate brown with ill-defined brick red horizontal stripes in A. robinsoni sp. nov.); from A. lionotus and A. poecilopus by male dewlap color (solid yellow-orange in A. lionotus and A. poecilopus), lack of postcloacal scales in males (usually present in A. lionotus and A. poecilopus), and possession of zero to three enlarged rows of middorsal scales (a broadband of 10–24 enlarged middorsal scale rows in A. poecilopus and A. lionotus); from A. barkeri by smaller size (maximum SVL 73 mm in A. riparius sp. nov., 91 mm in A. barkeri), presence of distinctly expanded toe pads (ratio width of expanded pads / width of distal phalanx 1.8–2.2 in A. riparius sp. nov. versus 1.4–1.6 in A. barkeri), and presence of a single row of middorsal caudal scales (double row in A. barkeri). Description of the holotype (Figure 6). Adult male as indicated by dewlap and everted hemipenes SVL 73.3 mm; HL 20.1 mm; HL/SVL 0.27; HW 10.7 mm; EOH 1.64 mm; IL 1.45 mm; IL/EOH.88; SLS-I 0.1; AGL 31.26 mm; FL 18.34 mm; T4L 15.95 mm; T4W 1.54 mm; TL 135.00 mm, tail complete; TL/SVL 1.84; TH 5.994 mm and TW 3.42 mm. 18 scales across the snout between second canthals; two scales separate the nasal opening from rostral scale; ten postrostral scales between supralabials; 12 supralabial scales from rostral scale to level of middle of eye; four scales separating supraorbital semicircles; 2 rows of scales separate subocular scales from supralabials; one slightly elongate superciliary scale; nine scales separating interparietal scale from supraorbital semicircles; four enlarged scales in supraocular disk; eight loreal scales in column just anterior to eye; 12 postmental scales posteriorly in contact with mental scale between infralabials; 15 scales in the loreal region; 13 rows of single scales in the dewlap (21–48 scales per row, rows somewhat irregular); seven enlarge middorsal scale row; 16 dorsal scales in 5% of SVL; 12 ventral scales in 5% of SVL; 220 scales around midbody; 15 expanded lamellae under fourth toe; 23 small middorsal caudal scales before the first large caudal scale (Fig. 6D). Preoccipital, enlarged infralabial, tail crest, middorsal, postcloacal scales absent; supraocular disc presents with small scales. Frontal region of head concave; scales in supraocular disc unicarinate; rostral scale with very weak cleft and overlaps mental scale; ear opening vertically ovoid; dorsal edge of ear lacks ornamentation; mental scale completely divided posteriorly and with its posterior edge convex; dewlap extends well on to chest with anterior insertion at level of anterior portion of the eye and without protruding scales at distal edge; dorsal scales strongly keeled and uniform in size; ventral scales in diagonal rows keeled and larger than dorsal scales; lateral scales homogeneous; tail is compressed and triangular in cross-section with the base taller than wide; middorsal caudal scales in single row and its size is less than three times than the adjacent scales (Fig. 6D); the longest toe of adpressed limb reaches anterior to eye and, supradigital scales keeled and multicarinate. Color in Life (based on field notes and color photographs; Fig. 6A and 6B). Dewlap yellow with orange-red horizontal streaks. Dorsal ground color olive to chestnut brown with transverse olive-green bars across dorsum down flanks extending to tail. A dirty cream-colored line extends posteriorly from the labial scales down the side of the body. Scattered small greenish spots are present on the dorsum. The dorsum and tail are marked with olive-green transverse bars. The limbs are marked with pale green to yellowish transverse bars with small punctuations. Variation. Size of reproductive males (69.6 ± 5.3, 65.8–73.3 mm) was larger than females (58.0 ± 6.0, 49.1–65.1 mm). Table 1 shows variation in the lepidosis characters of Anolis riparius sp. nov.. Individuals exhibited little variation in coloration. Although some individuals were dark, the amount and distribution of the light bands and dots were similar among all specimens. Hemipenis morphology. The completely everted hemipenis of SMF 92428 (Fig. 7B) is a stout bilobed organ. The sulcus spermaticus is bordered by well-developed sulcal lips and opens into two broad concave areas, one on each lobe. A small asulcate ridge is present. The apex is strongly calyculate, truncus with transverse folds. Etymology. The specific epithet is an adjective taken directly from Latin (riparius = of stream banks) about the riparian habitat of this species. Distribution. Anolis riparius sp. nov. is known from streams in the riparian gallery forests of the Pacuar, Guabo, and Savegre river basins (Fig. 1) from 100 to 1450 masl. This species inhabits lowland and premontane tropical wet forest life zones. Ecology. Little is known about the ecology and life history of Anolis riparius sp. nov. but it seems to share lifehistory traits with A. aquaticus and A. robinsoni sp. nov.. What follows is based mainly on field observations by MJR from Alfombra and additional unnamed streams of the Rio Guabo watershed. This species may be common in small order and headwater streams but absent from some other streams within a watershed. During the day this species may be found perched on rocks and boulders along streams and can jump into the water or crawl into interstitial cracks between rocks and boulders when pursued. At night these lizards are found sleeping on moss mats, fern fronds, and other large leafy riparian vegetation, especially near the splash zone around boulders. We have also found juveniles and adults sleeping on vertical rock walls and boulders in and next to waterfalls. We have observed these lizards sleeping from 0.5 to 2.0 meters above the ground. At Alfombra on 21 December 2007 MJR and SP found a congregation of seven gravid females on the same fern-covered boulder. One meter from the sleeping females was a cluster of 6 loosely aggregated eggs that were stuck to the moss on the same large boulder. Morphological comparisons of the two new species and Anolis aquaticus . Anolis aquaticus is the smallest species in this complex. It has fewer scales for all traits except for the number of scales between the suboculars and supralabials (Table 1 ). Anolis aquaticus has relatively larger dorsal scales, whereas A. robinsoni sp. nov. and A. riparius sp. nov. have small, granular dorsal scales that result in a higher number of total of scales around the body (more than 160, whereas A. aquaticus has 160 or fewer). The number of scales between the second canthals is close to half in number in A. aquaticus relative to the other species (Table 1, Fig. 4B, 5B, 6B), indicating the larger size of the scales in A. aquaticus. The number of small middorsal caudal scale varies between 0 to 10 in A. aquaticus (Fig. 5C) while the other species has more than 20 small middorsal caudal scales (Fig. 4D and 4D). However, the most striking feature separating one of the three species of this group is the color of the male dewlap. The red and yellow dewlap of A. aquaticus and A. riparius sp. nov. (Fig. 5A and 6B) contrasts with the dark brown dewlap of A. robinsoni sp. nov. (Fig. 4B). We did not find any difference in hemipenes morphology among A. aquaticus, A. riparius sp. nov., and A. robinsoni sp. nov. (Fig. 7).

Chaves, Gerardo, Ryan, Mason J., Bolaños, Federico, Márquez, Cruz, Köhler, Gunther, Poe, Steven (2023): Two new species of semiaquatic Anolis (Squamata: Dactyloidae) from Costa Rica. Zootaxa 5319 (2): 249-262, DOI: 10.11646/zootaxa.5319.2.6, URL: http://dx.doi.org/10.11646/zootaxa.5319.2.6

Uropeltis tricuspida sp. nov. Figs. 4–6; Tables 2–3 urn:lsid:zoobank.org.act: 3BFE988E-B0D4-4E52-BEAF-6B2DEFD3B6C3 Uropeltis petersi (Beddome, 1878): Boulenger 1890, 1893 in part; Wall 1923, in part; Smith 1943, in part; Gans 1966, in part; Wallach et al. 2014, in part; Pyron et al. 2016, in part; Sampaio et al. 2023, in part Holotype. CAS 244452 (Fig. 5), adult female based on large size, and relatively high ventral and low subcaudal count. Collected by C. Gans, C. Rajasundaram and R. A. Bonhoure on 11 June 1990 from Sevenmalai (also spelled Sevenmallay or Sivanmalai, a tea estate), 4 km West of Munnar, Idukki District, Kerala, India, 2,100 m (ca. 10.075°N, 77.0417°E, from Google Earth and Rajendran (1985: map 3), though land here is ca. 1,600 m elevation versus the reported 2,100 m). Paratypes (n = 7). CAS 244451 (male), 244453 (female), 244454 (male), 244455 (female) and 244456 (male), same data as for the holotype. CAS 244441 and 244442 (both male) collected by C. Gans, C. Rajasundaram and R. A. Bonhoure on 10 June 1990 from Sevenmalai Estate, 2 km East of Munnar, Idukki District, Kerala, India, 2,000 m. Whole-specimen photographs of these specimens are presented in Fig. 6. Referred specimens (n = 30). CAS 244407 (male), 244408 (male), 244409 (female), 244411 (female), 244413 (male), 244414 (female), 244415 (male), 244416 (male), and 244417 (male) collected by C. Gans, C. Rajasundaram and R. A. Bonhoure on 9 June 1990 from Sevenmalai Estate, 2 km East of Munnar, 2,000 m. CAS 244421 (female), 244422 (female), 244424 (male), 244425 (female), 244426 (female), 244427 (male), 244443 (female), and 244444 (male) collected by C. Gans, C. Rajasundaram and R. A. Bonhoure on 9 June 1990 from Parvathi (Parvathy) Division, Sevenmalai Estate, 4 km East of Munnar, 2,000 m. CAS 244450 (male) collected by C. Gans, C. Rajasundaram and R. A. Bonhoure on 11 June 1990 from Parvati Division, Sevenmalai Estate, 4 km West of Munnar, 2,100 m. CAS 244447 (female) and 244436 (male) collected by C. Gans, C. Rajasundaram and R. A. Bonhoure on 10 June 1990 from Sevenmalai Estate, 2 km East of Munnar, 5,500 feet (= 1,676 m). CAS 244437 (male); CAS 244438 (female), 244439 (male), and 244440 (male) collected by C. Gans, C. Rajasundaram and R. A. Bonhoure on 10 June 1990 from Sevenmalai Estate, 2 km East of Munnar, 2,000 m. CAS 244886, 244888 and 244889 (three females) from Sevenmalai, 20 April 1972. MNHN 1897.249 (male) from Peermad, N. Travancore, 4,000 feet; BNHS 3621 (male: voucher specimen reported as UK MW 2228 by Sampaio et al. 2023) from near Munnar collected by O. V. Oommen and colleagues October 2001; BMNH 1946.1.17.8 (male), Anamallays, 4,000 feet, R.H. Beddome. Diagnosis. A species of Uropeltis that differs from all congeners except U. liura, U. maculata and U. petersi in having 17 rows of dorsal scales at midbody, nasals that make contact along the midline posterior to the rostral, and that lack a flattened or mildly longitudinally convex tail shield composed of matt scales with substantial keels— instead having a strongly longitudinally convex tail tip with scales bearing low, inconspicuous ridges. Differs from U. petersi in having more ventral scales in females (178–185 versus 156–162) and more subcaudal scales (6–10 and 10–13 versus 5–6 and 10–11 in males and females, respectively), and in having a tricuspid versus bicuspid terminal scute. Differs from U. maculata in lacking large yellow spots or blotches laterally on the anterior end of the body, in having typically two versus three small, non-subcaudal scales overlapped by each anal shield, and in having a tricuspid versus bicuspid terminal scute. Differs from U. liura in having fewer ventral scales (156–174 versus 173 to>195, respectively), in having typically two versus three small, non-subcaudal scales overlapped by each anal shield, lacking yellow dots on most of the body scales on the dorsum, and in having a tricuspid versus bicuspid terminal scute. Uropeltis tricuspida sp. nov. is distinct from these three phenotypically superficially similar congeners (and all other species) also in DNA sequence data (Sampaio et al. 2023). Description of holotype. Some meristic and morphometric data presented in Table 2. Female, based on relatively high number of ventrals, relatively short tail and relatively few subcaudals (see paratype variation section below). Good condition, preserved in a tight but flexible U-shape. Body subcylindrical, slightly wider anteriorly than posteriorly, venter slightly flattened, perhaps somewhat artefactually; tapering over c. 15 mm immediately behind head; very gently tapering posteriorly up to vent and onto tail until final c. 5 mm when tapering more strongly. Head strongly tapering in dorsal view, sides straight to slightly convex to just behind nostrils, rounded anteriorly; tapering also in lateral view, gently with straight to slightly convex edges up to level with nostrils. Rostral rounded anteriorly, wider than long in dorsal view, much deeper than long in lateral view; dorsally without longitudinal ridge or crest; at widest slightly in front of and below nostril. Rostral shorter (in dorsal view) than rostral-frontal gap. Frontal six-sided, lateralmost edges in contact with oculars diverging anteriorly. Frontal much longer, wider than rostral. Paired nasals in brief contact behind rostral. Midline contact between nasals and between prefrontals (left overlapping right in both cases) subequal in length; midline sutures not parallel with long axis, left prefrontal contacts right nasal but not vice versa. External naris small, subcircular, slightly countersunk within small depression, located towards anteroventral corner of undivided nasal shield. Nasal contacts first and second supralabials (SLs). Prefrontal (as nasal) wider than long, shorter than frontal, contacts SL2 and SL3. Supralabials four, SL1 smallest, making least contribution to margin of mouth; SL2 larger than SL1, smaller than SL3; SL4 much the largest and longest. Ocular contacts SL3 and SL4; posterodorsal margin slightly concave. Eye small but distinct, diameter less than one half and more than one third length of ocular shield, located near anteroventral corner of ocular; bulges slightly from ocular surface, pupil appears subcircular; eyes directed anterolaterally and slightly dorsally. Paired parietals each about as long as frontal, posteriorly broadly rounded. Opposite parietals in brief midline contact, left overlapping right. Parietals slightly longer than wide, wider than frontal and rostral. Each parietal contacts four scales other than head shields. Three infralabials (ILs) on each side, IL1 and IL3 subequal in length, notably shorter than IL2. Left and right first infralabials in midline contact, separating small, slightly prominent mental from first midline ventral scale. First four ventrals longer than wide, fifth and subsequent ventrals wider than long. Six maxillary and eight mandibular teeth on each side. Teeth simple, pointed, distinctly retrorse, straight to slightly recurved, evenly spaced, smallest posteriorly. Head and body scales macroscopically smooth, lacking keels, except on far posterior of body and tail. Body scales generally evenly sized on dorsum and along body except for those involved in dorsal scale row reductions. Midline ventral scales between mental and anal of even size though anterior- and posteriormost ones gradually narrow, posteriormost ventral with V-shaped posterior margin (straight to gently rounded in most preceding ventrals). Ventrals 181, at midbody approximately 1.6 times as broad as exposed part of adjacent first dorsal scale row. Dorsal scale rows 19 anteriorly (by two head lengths behind head), reducing to 17 by level with 32 nd ventral up to level with vent. At level of 52 nd (left) and 53 rd (right) ventral the fourth dorsal-row scale is noticeably enlarged (“b”) and the fifth-row scale notably small (“s”: represented in square brackets in formula below): 4+5 (32) [b4, s5: 53] 19 --------------- 17 ----------------- 17 4+5 (32) [b4, s5: 52] Dorsal scale rows 15 at base of tail. Paired anal scales (right overlying left) considerably larger than posteriormost ventrals and subcaudals. Distal margin of each anal overlaps two other scales in addition to anteriormost subcaudals. Seven subcaudals on each side. Last few dorsal scale row scales on body, but not ventrals, anals or subcaudals with multiple (typically 3 to 5, see Fig. 5) short and very low subparallel ridges, increasingly prominent posteriorly and dorsally. Very inconspicuous other than on approximately last four scales anterior to terminal scute. Tail ‘shield’ not clearly demarcated, distinctly convex both longitudinally and transversely; most closely matching Type III among Smith’s (1943) states for Uropeltis tails (also type III of Pyron et al. 2016). Terminal scute prominent, approximately as wide as long, surrounded by nine scales including last subcaudals; ventrally smooth and convex both longitudinally and transversely; dorsally somewhat flat centrally, strongly upturned posteriorly, with irregular (though subparallel, anteriorly diverging) short, tubercles, more prominent on anterior half. Posterior upturned edge of terminal scute ends in thin tricuspid subhorizontal ridge with paramedian cusps (0.5 mm apart) flanking more prominent midline cusp. In alcohol, background dorsal body colour uniform brown; edges of distal tips of dorsal body scales translucent and narrow edges of bases of scales off-white to pale dull yellow, such that most of the dorsum (the nine central dorsal scale rows; except for posteriormost c. 30 mm of specimen) bears very narrow, longitudinal (zigzag) pale lines between darker scales. Scales on dorsum mostly unblemished though some (especially anteriorly) with pale yellow flecks. Body scales slightly iridescent. Head brownish, about as dark as anterior of body; fairly uniform above except for slightly paler orange-brown tip of rostral and pale-yellow upper lips and most of SL4. Underside of head with more and more-irregular pale flecking. An irregular, lateral, pale-yellow line extends posteriorly from SL4, diminishing and becoming increasingly wavy and irregular over anteriormost quarter or so of body. Ventral aspect of specimen with more pale-yellow markings than dorsally, mostly confined to first three dorsal scale rows; ventrals mostly brown, except over anteriormost c. 25 mm where mostly with at least some pale yellow. Most scales with pale-yellow on dorsal rows 1–3 have pale blotches rather than being entirely pale. Subcaudals brown with pale-yellow lateral edges that, along with pale markings on medial edges of first dorsal scale row on tail, form a pair of tapering, gradually converging pale stripes on ventrolateral surface of tail. Anterior ends of these stripes merge with broad transverse pale-yellow band that covers both anals and most of the posteriormost ventral. Upper surface of tail brown-grey except for increasingly pale posterior end of terminal scute. Variation in paratypes. Seven more or less U-shaped specimens (Fig. 6), in generally good condition. CAS 244454 close to ecdysis when fixed, with broken spine at approximately midbody, and small piece of viscera protruding though midventral hole into coelom, with patch of outer layer of scales missing anterior and posterior of this hole. CAS 244441 slightly macerated; CAS 244442 with outer scale layer missing in some patches. Some meristic and morphometric data presented in Table 2; scale-reduction formulae presented in Appendix 1. Generally similar to holotype. Rostral slightly longer or shorter or subequal to gap between rostral and frontal. Lateralmost edges of frontal barely diverge in CAS 244456. Right prefrontal contacts left nasal in CAS 244451. Parietals generally notably slightly shorter than frontal. Parietals slightly wider than long in CAS 244456. CAS 244454 has seven rather than six maxillary teeth on each side. Midbody ventrals 1.5–1.9 times width of adjacent dorsal-scale row scales. Dorsal scale rows reduce to 16 shortly anterior to vent in CAS 244456, and 14 instead of 15 rows at base of tail in 244455 and 244456. Low longitudinal ridges on posteriormost tail scales most noticeable on four to six (CAS 244451, 244455 and 244456) to seven to nine (CAS 244441, 244442) scales anterior to terminal scute. Posterior end of terminal scute tricuspid in all paratypes except CAS 244441, in which it is asymmetrical and somewhat more bicuspid. Dorsum generally lacks pale flecks. Anterolateral pale stripe more or less well developed and extensive. Venter in some paratypes with incomplete, pale transverse bars one scale long with two darker scales between each pale bar, most notably in CAS 244455 and 244456. More pale blotches on dorsal surface of head in CAS 244456. CAS 244442 lacks pale, narrow, longitudinal zigzag lines on anterior of dorsum. Ventrolateral pale stripes on tail broken on one side in CAS 244451, 244453 (anals brown rather than pale), 244454, 244455 (anals partly brown), and 244456 (small brown blotches on anals; pale ventrolateral marks meet midventrally on posterior half of tail). Variation in referred specimens. Some meristic and morphometric data presented in Table 3. Scale-reduction formula for one aberrant specimen with many fluctuations in scale rows on right side of body presented in Appendix 1. Each anal overlaps three rather than two small scales (in addition to subcaudals) in MNHN 1897.249 and on one side in CAS 244436. Sexual dimorphism. The type and non-type material is together strongly sexually dimorphic in that (as in many other uropeltids) females have more ventrals (176–185 versus 170–178), fewer subcaudals (7–9 versus 11–13), and relative shorter tails (3.1–4.4 versus 4.4–6.3 % of total length) (Tables 2, 3). Etymology. The name is in reference to the species having typically three cusps on the posterior end of the terminal shield. For nomenclatural purposes, the species epithet is a noun in apposition. Suggested vernacular names. Three-cusped uropeltis or Three-cusped shieldtail (English). Distribution, habitat and conservation. One of the early ( According to the catalogue, nine of the CAS specimens that have associated habitat data were “dug up from very wet turf” and eight were “dug up in garden patches, wet soil”. BNHS 3621 was also dug from moist soil, while searching for caecilian amphibians in a low-intensity agricultural habitat (O.V. Oommen, pers. comm.). Although the new species thus appears to tolerate at least some human disturbance and was locally abundant, at least, in 1990, little else is known of its natural history, and it has been recorded only from a very small area. Thus, it likely qualifies as Data Deficient conservation status based on IUCN Red List criteria (IUCN Species Survival Commission 2012).

Uropeltis petersi (Beddome, 1878) Figs. 1–3; Table 1 Lectotype. BMNH 1946.1.16.75 (formerly BMNH 78.1.11.3: Fig. 1) by present designation. Male from Anamallays, 4,000 feet; provided by R. H. Beddome. Paralectotype (n = 1). BMNH 1946.1.17.9 (male; Fig. 2) from Anamallays, 4,000 feet; provided by R. H. Beddome. Referred material (n = 8). BMNH 1946.1.17.7 (female; Fig. 3) from Anamallays, 4,000 feet; provided by R.H. Beddome; MNHN 1895.80 (3 specimens; 2 female, 1 male) from Anamallays forest, 4,000 feet; MCZ 6201 (4 specimens; 3 female, 1 male) from Anamallays, 4,700 feet. Revised diagnosis. A species of Uropeltis that differs from all congeners except U. liura, U. maculata and U. tricuspida sp. nov. in having 17 rows of dorsal scales at midbody, nasals that make contact along the midline posterior to the rostral, and that lack a flattened or mildly longitudinally convex tail shield composed of matt scales with substantial keels—instead having a strongly longitudinally convex tail tip with scales bearing low, inconspicuous ridges. Differs from U. liura in having fewer ventral (156–163 versus>172) and subcaudal scales (5– 6 and 10–11 versus 7–9 and 11–13 in males and females, respectively), in having typically two versus three small, non-subcaudal scales overlapped by each anal shield, and in lacking yellow dots on most of the body scales on the dorsum. Differs from U. tricuspida sp. nov. in having fewer ventral scales in females (156–162 versus 163–174) and fewer subcaudal scales (5–6 and 10–11 versus 6–10 and 10–13 in males and females, respectively). Differs from U. maculata in having fewer subcaudal scales (5–6 in females and 10–11 in males versus 7–9 and 11–13, respectively), fewer and less sexually dimorphic ventrals (156–163 and 157–163 versus 163–174 and 154–163 in females and males, respectively) and in lacking large yellow spots or blotches laterally on the anterior end of the body. Uropeltis petersi is distinct from these three phenotypically superficially similar congeners (and all other species) also in DNA sequence data (Sampaio et al. 2023). Remarks. Beddome (1878) did not specify the number of type specimens used for the description of Silybura petersi but, given that the number of ventral scales was reported as “155 to 158” and the number of subcaudal scales as “ten to twelve pairs”, it is clear that there was more than one type. McDiarmid et al. (1999) report four types (1946.1.17.7–9 and 1946.1.16.75), but Pyron et al. (2016) report “putatively” six—the four reported by McDiarmid et al. (1999) plus MNHN 1895.80a–b that were also reported as types by Gans (1966) and Wallach et al. (2014). Several of these specimens can be ruled out as potential types. BMNH 1946.1.17.7 and two of the MNHN specimens have only five or six subcaudals on each side. Additionally, BMNH 1946.1.17.8 (Fig. 4) is clearly not one of the types given that it has 183 ventrals (many more than the range of 155–158 reported by Beddome 1878 or Beddome 1886). The latter specimen is here referred to the new species described below. Of the remaining specimens, I select BMNH 1946.1.16.75 (previously 78.1.11.3) as the lectotype because it is the only unambiguously Beddome specimen that is associated with a precise accession date (11 January 1878), and it was accessioned on that date under the name Uropeltis petersi in the same year that the name was first published. BMNH 1946.1.17.9 is considered the sole paralectotype, being the only other BMNH Beddome specimen that has a ventral and subcaudal count within the range reported in the original description. In his characterisation of Uropeltis petersi, Beddome (1886) extended the reported range in numbers of ventrals from 155–158 to 155–160, and also added that females had 6 subcaudals (and males 10–12)—the additional, female specimen(s) that this characterisation is based on might have been or included BMNH 1946.1.17.7 that was subsequently considered a type specimen by other workers. Boulenger (1893: 48; plate VI, fig. 1) documented 11 specimens of U. petersi, with four reported as types (presumably the BMNH types reported by Gans 1966, McDiarmid et al. 1999 and Pyron et al. 2016), and the remaining seven specimens likely being the four subsequently sent to Harvard (MCZ 6201) and three sent to Paris (MNHN 1895.80). According to Boulenger (1893), the four BMNH specimens were from Beddome, and the other seven from W. Davison. Constable (1949: 122) briefly reported on the four topotypic (“Anaimalai Hills at 4,700 feet ”) specimens sent from the British Museum to Harvard (MCZ 6201). Data for all of the known specimens are presented here in Table 1. I concur with Pyron et al. (2016) that U. petersi has seemingly not been collected or new field sightings reported in the scientific literature since before 1900. The precise location of the previously collected or any extant populations are currently unknown, other than likely being in the region of 1,200 m elevation in the Anamalai hills.

Gower, David J. (2023): On the taxonomy of Uropeltis petersi (Beddome, 1878) (Serpentes: Uropeltidae) and description of a new, closely related species from the Western Ghats of India. Zootaxa 5319 (1): 103-119, DOI: 10.11646/zootaxa.5319.1.7, URL: http://dx.doi.org/10.11646/zootaxa.5319.1.7

Lee Grismer, Evan S. H. Quah L., Syafiq, Muhamad Fatihah, Rujirawan, Attapol, Aowphol, Anchalee, Ahmad, Amirrudin B., Anuar, M.S. Shahrul (2023): Comments on the taxonomic status of Cyrtodactylus zebraicus Taylor, 1962 (Squamata: Gekkonidae) from Northern Peninsular Malaysia. Zootaxa 5318 (4): 489-503, DOI: 10.11646/zootaxa.5318.4.3, URL: http://dx.doi.org/10.11646/zootaxa.5318.4.3

Cyrtodactylus zebraicus Taylor, 1962 (Figs. 3–5; Table 2 & 3) Cyrtodactylus peguensis (in part): Das & Norsham 2007: 77; Grismer 2008: 30 Cyrtodactylys peguensis zebraicus: Ulber 1989: 4 & 5; Denzer & Manthey 1991: 314; Manthey & Grossmann 1997: 226; Das 2010: 213 Referred specimens. USMHC 2635, collected by Evan S.H. Quah and M.S. Shahrul Anuar on 7 November 2019 at approximately 21:00 h from Perlis State Park, Perlis, Malaysia (approximately 6.698222N, 100.191611E; 144 m elevation). USMHC 2639 bears the same collection data as USMHC 2635 but was collected on 9 November 2019. UMTZC 2413, collected by Muhamad Fatihah Syafiq, Baizul Hafsyam Badli-Sham and Amirrudin B. Ahmad on 5 October 2022 at 20:25 h from Bukit Ayer, Perlis, Malaysia (6.542000N, 100.168000E; 52 m elevation). Description of specimen USMHC 2635. Adult male with 62.0 mm SVL; head moderate in length (HL/SVL 0.29), wide (HW/HL 0.58), slightly flattened (HD/HL 0.40), distinct from neck, and triangular in dorsal profile; lores concave anteriorly, inflated posteriorly; frontal region flattened, prefrontal region slightly concave, canthus rostralis rounded; snout short (ES/HL 0.40), rounded in rostral region, eye to snout distance slightly less than head depth; eye large (ED/HL 0.22), eyeball slightly protuberant, eye diameter less than the eye to ear distance, pupil vertical; ear opening elliptical, obliquely oriented, moderate in size (EL/HL 0.09); rostral large, subrectangular, height 1.5 mm, shorter than wide, 2.7 mm, medially divided by a dorsal furrow, reaching to approximately halfway down rostral height, bordered posteriorly by supranasals and internasal, laterally by first supralabials and nostrils; external nares at anterior angle of snout, directed lateroposteriorly, bordered anteriorly by rostral, dorsally by two large supranasals, posteriorly by two smaller postnasals, ventrally by first supralabial; internarial distance narrow; anterior pair of supranasals subrectangular, separated by one small internasal, bordered anteriorly by rostral, laterally by nostrils, posteriorly by posterior supranasal and three smaller scales; one internasal, subhexagonal, vertically arranged, slightly protruding rostral, bordered posteriorly by two small scales; 9/8 (right/left) supralabials extending to below midpoint of eye, 15/12 to below the posterior margin of the eyeball, subrectangular anteriorly, elliptical shape posteriorly; 6/7 infralabials extending to below midpoint of eye, 9/11 to below the posterior margin of the eyeball, larger than supralabials, tapering smoothly posteriorly; scales of frontonasal, prefrontal and lores, small, relatively raised, domed, slightly larger than granular scales on top of head and occiput; scales on occiput intermixed with scattered, slightly larger, more rounded, dome to subconical tubercles, more prominent tubercles between occiput and above ear opening; dorsal supraciliaries crenulated, not elongate or keeled; mental large, triangular, 2.0 mm in width, 1.7 mm in length, bordered laterally by first infralabials and posteriorly by large, right and left trapezoidal postmentals that contact medially for 50% of their length posterior to mental; one row of slightly enlarged chin shields extending posteriorly to sixth (right/left) infralabials; and gular and throat scales small, granular, grading posteriorly into larger, flat, smooth, imbricate, pectoral and ventral scales. Body relatively short, somewhat stout (AG/SVL 0.46), lacking ventrolateral folds; scales on dorsum small, mostly homogenous, granular, interspersed with larger, irregularly arranged, slightly prominent trihedral keeled tubercles; tubercles extending from occiput beyond to the base of the original portion of tail; tubercles on occiput, nape and anterior of body at level above shoulder smaller, subconical; those mid-dorsally and on the posterior section of the body larger, being more dense, slightly more prominently keeled, and more regularly arranged in sacral region and tail base; tubercles on flanks sparse; approximately 15 longitudinal rows of dorsal tubercles; approximately 22 paravertebral tubercles; 30 flat, imbricate, smooth ventral scales, those near midline larger than those laterally and dorsal scales; femoral are not enlarged; precloacal scales smooth, approximately twice the size of femoral scales; femoral pores absent; seven precloacal scales; six precloacal pores; four rows of enlarged postprecloacal scales; and precloacal groove or depression absent. Limbs moderately slender;forelimbs relatively short(FL/SVL 0.16); scales on dorsal surface domed to subconical, granular, slightly larger than those on body, interspersed with sparsely enlarged, subconical and trihedrally keeled tubercles; dorsal scales of wrist and palm flat, smooth, round, imbricate; ventral scales of palm flat, weakly rounded, slightly raised, not imbricate, smaller than those on body; 14/14 (right/left) total subdigital lamellae on fourth finger, 4/4 proximal subdigital lamellae rectangular with rounded to weakly rounded corners, broadly expanded proximal to joint inflection on fourth finger, 10/10 distal subdigital lamellae, slightly expanded immediately distal to joint, becoming gradually more expanded near the claw; digits well-developed, relatively long, inflected at basal interphalangeal joints; digits slightly narrower distal to inflections; no interdigital webbing; claw well-developed, relatively short, claw base sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.17); dorsal scales domed to subconical, granular, interspersed with enlarged subconical and trihedrally keeled tubercles, and anterior part of thigh covered by flat, slightly larger, imbricate scales; ventral scales of femora flat, smooth, imbricate, smaller than those on body; small postfemoral scales form an abrupt union with large, flat ventral scales of posteroventral margin of thigh; ventral scales of tibia flat, imbricate; dorsal scales of plantar surface relatively smooth, rounded, imbricate; ventral scales of plantar surface low flat, weakly rounded; 16/16 (right/left) total subdigital lamellae on fourth toe, 5/5 proximal subdigital lamellae, rectangular with rounded to weaky rounded corners, broadly expanded proximal to joint inflection on fourth toe, 11/11 distal subdigital lamellae, slightly expanded immediately distal to joint, becoming gradually more expanded near the claw; digits well-developed, relatively long, inflected at basal, interphalangeal joints; and claw well-developed, relatively short, claw base sheathed by a dorsal and ventral scale. Tail 41.4 mm in length, regenerated, shorter than SVL (TL/SVL 0.67), moderate in proportions, cylindrical, wide anteriorly, 5.3 mm in width at base, tapering to a tip, covered with small, flat, imbricating scales on the dorsal surface but slightly larger, flat, imbricating scales on ventral surface; dorsal scales of original portion of tail base granular, round, becoming larger, flatter, subimbricate posteriorly on regenerated portions of tail; those on tail base bearing trihedrally keeled tubercles forming paravertebral rows, no caudal furrow; base of tail forming hemipenial swelling; 2/2 (right/left) postcloacal tubercles on the enlarged smooth hemipenial swelling; and postcloacal tubercles approximately equal size. Coloration in life (Fig. 2). Dorsal ground color of head, body, and limbs light-brown; top of head bearing large, dark-brown blotches edged in yellow, forming a light-colored reticulated pattern; superciliary scales yellow; supralabials and infralabials off-white bearing darker markings; wide, discontinuous, dark-brown nuchal loop edged in yellow, extending from posterior margin of one orbit, across occiput and nape to posterior margin of the other orbit; five dark-brown paravertebral blotches on the dorsum starting from shoulder region on top of the forelimbs until the sacral band, all blotches edged in yellow but the second and third pair of paravertebral blotches not divided medially along the vertebral column; all yellow outlines of the paravertebral blotches extend along the flanks towards the venter to form a series of narrow, yellow bands along the flanks; sacral band between hindlimbs composed of paravertebral blotches; postsacral band composed of confluent blotches; regenerated tail dark-grey with lighter grey and orange-brown speckling; dorsal portion of forelimbs and hindlimbs bearing irregularly shaped yellow banding and reticulations; all ventral surfaces generally greyish white, immaculate, except for ventral surface of knee and elbow region which bear some dark mottling. Coloration in preservation (Fig. 2). The overall color pattern is similar to that in life except that it is faded. Ground color of head, body, and limbs light-brown; nuchal, body and sacral bands dark brown but slightly lighter than in life; the light-yellow reticulated pattern on the dorsal surfaces of the head, body and limbs faded to off-white; and all ventral surfaces light-beige. Variation (Fig. 2, 3 & 5). Morphometric, meristic and color pattern characters of the other referred specimens of C. zebraicus from Peninsular Malaysia are presented in Table 1. All the specimens are generally similar in most aspects of morphology with subtle variation in color pattern (Figs. 2 & 3). In USMHC 2639 the right 4TLU is damaged and approximately 2/3 of tail is original: bearing four longitudinal rows of dorsal tubercles, two transverse rows of dorsal tubercles extend from tail base to anterior margin of first light caudal band, 4.7 mm from tail base, approximately 8.3% of tail; a single enlarged median row of transverse scales on subcaudal region starting from second light caudal band on original portion of tail and ends at the start of the regenerated portion of the tail. The color pattern of USMHC 2639 is also slightly lighter both in life and preservation and the original portion of the tail bears light and dark caudal bands that completely encircle the tail. Similarly, the tail that is original in UMTZC 2413 bear light and dark caudal bands that completely encircle the tail. ......continued on the next page TABLE 1 ( Continued) Distribution in Peninsular Malaysia (Fig. 1). Cyrtodactylus zebraicus is currently only known from around the Nakawan Range in Perlis state but the species may range more widely in Peninsular Malaysia. However, given what is known about its habits in Thailand (Grismer et al. 2021), it is expected to be confined to only lowland areas in the extreme north at the Malaysian-Thai border, in the states of Kedah, Perak and Kelantan. The extralimital distribution of the species outside of Peninsular Malaysia is in peninsular Thailand, south of the Isthmus of Kra (Grismer et al. 2021). Natural history (Fig. 4). In Peninsular Malaysia, C. zebraicus is a habitat generalist and has been found in lowland, semi-deciduous forest (Sharma et al. 1996, 2001) associated with karstic areas along the Nakawan Range. Specimens USMHC 2635 and 2639 were both observed at night approximately 1m above the ground on the trunks of small trees, in an open area near the forest edge. It was raining the night USM 2635 was found and the gecko was perched on a dry section of the tree trunk and partially protected by low vegetation. In comparison, UMTZC 2413 was found at night crawling in a grassy area bordering the main road that cuts through the forested area. The weather was clear the night UMTZC 2413 was found but it had been raining for a few days beforehand. Conservation status. At present, there is only limited ecological knowledge for the species in Peninsular Malaysia. Anecdotal observations indicate that Peninsular Malaysian specimens are similar to Thai populations in habits, and it is a habitat generalist that can survive in habitats with moderate human disturbance (Grismer et al. 2021). In addition, the species occurs in protected areas managed by the Perlis Forestry Department such as Bukit Ayer. As such, we believe that the future of C. zebraicus in the country is secure and we recommend listing it as a species of Least Concern (LC). Comparison and comments on taxonomy. Cyrtodactylus zebraicus can be easily differentiated from all other species of Cyrtodactylus known in Peninsular Malaysia based on a combination of color pattern and scale characters. From the agamensis group in Peninsular Malaysia composed of C. jarakensis, C. metropolis, C. pantiensis, C. payacolus, C. semenanjungensis and C. tiomanensis (Grismer et al. 2021), C. zebraicus differs by having fewer paravertebral tubercles (22–27 vs. 32–40), fewer ventral scales (30–36 vs. 36–61), fewer subdigital lamellae on fourth toe (13–17 vs. 17–24) and the color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. dark speckling with absence of white reticulum) (Grismer 2011; Grismer et al. 2014a). From C. aurensis of the philipinicus group (Grismer et al. 2021), C. zebraicus differs by its smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 95.0– 99.4 mm SVL), fewer ventral scales (30–36 vs. 45–51), fewer subdigital lamellae on fourth toe (13–17 vs. 18–23), precloacal groove (absent vs. present and deep), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. lightly colored reticulate pattern) and patterning on body (dense bold banding or blotched pattern on body, and bold light and dark banding on original tail vs. narrow light bands on body and tail) (Grismer 2011). From members of the pulchellus group consisting of C. astrum, C. australotitiwangsaensis, C. bintangrendah, C. bintangtinggi, C. dayangbuntingensis, C. evanquahi, C. hidupselamanya, C. jelawangensis, C. langkawiensis, C. lenggongensis, C. macrotuberculatus, C. pulchellus, C. sharkari, C. timur and C. trilatofasciatus (Grismer et al. 2021), C. zebraicus can be easily differentiated on the basis of lacking a deep precloacal groove, smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 99.0– 122.2 mm SVL), contiguous series of enlarged pore-bearing femoral and precloacal scales (absent vs. present), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. generally immaculate with absence of white reticulum) and patterning on body (dense bold banding or blotched pattern on body vs. 3–7 bold, dark bands on the body edged in white) (Grismer 2011; Quah et al. 2019; Wood et al. 2020). From members of the brevipalmatus group composed of C. cf. brevipalmatus and C. elok (Grismer et al. 2021), C. zebraicus differs by webbing at the base of digits (absent vs. present), caudal fringe of spiny tubercles along tail (absent vs. present), fewer ventral scales (30–36 vs. 36–47), fewer subdigital lamellae on fourth toe (13–17 vs. 19–21), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. mottled with absence of white reticulum) and patterning on body (dense bold banding or blotched pattern on body vs. color similar to tree bark with usually faint bands and mottling) (Grismer 2011; Grismer et al. 2022). From C. durio of the lateralis group (Grismer et al. 2021), C. zebraicus differs by its smaller adult length in both sexes (50.0– 65.7 mm SVL vs. 79.3 mm SVL in adult male), elongate, spinose tubercles in ventrolateral body fold and on ventrolateral margin of tail (absent vs. present), fewer ventral scales (30–36 vs. 59), fewer subdigital lamellae on fourth toe (13–17 vs. 22), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. irregular mottling) and patterning on body (dense bold banding or blotched pattern on body vs. squarish blotches on body) (Grismer 2011). From C. leegrismeri of the condorensis group (Grismer et al. 2021), C. zebraicus differs in its smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 90.0–92.0 mm SVL), fewer subdigital lamellae on fourth toe (13–17 vs. 18–21), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. weak reticulum) and patterning on body (dense bold banding or blotched pattern on body, and bold light and dark banding on original tail vs. dark, indistinct symmetrical blotches on body and tail) (Grismer 2011). From members of the sworderi group consisting of C. guakanthanensis, C. gunungsenyumensis, C. quadrivirgatus, C. sworderi and C. tebuensis (Grismer et al. 2021), C. zebraicus differs by its smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 71.0– 82.2 mm SVL), fewer paravertebral tubercles (22–27 vs. 32–40) and color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. mottled with absence of white reticulum) (Grismer 2011; Grismer et al. 2016). From members of the darmandvillei group composed of C. batucolus and C. seribuatensis (Grismer et al. 2021), C. zebraicus differs by its smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 70.1–75.2 mm SVL), fewer ventral scales (30–36 vs. 28–42), fewer subdigital lamellae on fourth toe (13–17 vs. 17–22), contiguous series of enlarged pore-bearing femoral and precloacal scales (absent vs. present), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. dark speckling with absence of white reticulum) and patterning on body (dense bold banding or blotched pattern on body vs. dark blotches on body, bordered by diffuse, light colored bands or spots) (Grismer 2011). From populations of the C. consobrinus complex of the malayanus group (Grismer et al. 2021), C. zebraicus differs by its smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 117.4–125.0 mm SVL), fewer ventral scales (30–36 vs. 58–65), fewer subdigital lamellae on fourth toe (13–17 vs. 23–28), enlarged pore-bearing femoral scales in males (absent vs. present), and patterning of body (dense bold banding or blotched pattern on body vs. narrow, relatively widely spaced lighter bands) (Grismer 2011).

Dendrelaphis in Vietnam is provided below: 1a. Vertebral scale row distinctly enlarged, two supralabials touching the eye........................................ 2 1b. Vertebral scale row feebly enlarged, one supralabial touching the eye............................................ 3 2a. Ventrolateral black stripe present................................................................. D. pictus 2b. Ventrolateral black stripe absent............................................................ D. ngansonensis 3a. Dorsal scale rows 15 on midbody............................................................. D. subocularis 3b. Dorsal scale rows 13 on midbody........................................................... D. binhi sp. nov., Published as part of Nguyen, Sang Ngoc, Nguyen, Vu Dang Hoang, Le, Manh Van, Nguyen, Luan Thanh, Vo, Thi-Dieu-Hien, Vo, Ba Dinh, Che, Jing & Murphy, Robert W., 2023, A new snake of the genus Dendrelaphis Boulenger, 1890 (Squamata: Colubridae) from the coastal area of southern Vietnam, pp. 130-144 in Zootaxa 5318 (1) on page 142, DOI: 10.11646/zootaxa.5318.1.6, http://zenodo.org/record/8158335

Dendrelaphis binhi sp. nov. Holotype. ITBCZ 6663, adult male, collected from Tuy Phong, Binh Thuan Province, Vietnam; coordinates 11 o 12’36.2”N, 108 o 41’43.2”E; elevation 34 m a.s.l. by S.N. Nguyen and L. T. Nguyen, on 23 July 2018 (Figs. 3 & 4). Paratypes. Seven specimens: ITBCZ 6664, ITBCZ 6666 (adult females), ITBCZ 6665 (subadult male), collected by the same collectors, on the same date and site as the holotype; ITBCZ 6681 (adult male), collected from Tuy Phong, Binh Thuan Province, Vietnam, coordinates 11 o 13’03.3”N, 108 o 39’10.2”E, elevation 39 m a.s.l. by S.N. Nguyen and L. T. Nguyen, on 26 July 2018; ITBCZ 5944-6 (adult females), collected from Thuan Nam, Ninh Thuan Province, Vietnam; coordinates 11 o 19’18.4”N, 108 o 56’46.6”E; elevation 105 m a.s.l. by S.N. Nguyen and D.H. V. Nguyen, on 13 April 2017 (Figs. 3–5). Diagnosis. Dendrelaphis binhi sp. nov. is distinguished from all of its congeners by a combination of the following morphological characters: medium sized bronzeback snake (largest TL 936 mm in female); one long supralabial entering orbit; body scale smooth, 13 rows on neck and midbody, 9–11 before vent; vertebral scale row slightly enlarged; ventrals 154–158 in males and 161–170 in females; subcaudals 105 or 106 in males and 95–102 in females; 18 or 19 maxillary teeth, posterior teeth shorter than the others; hemipenis spinose, reaching 10 th or 11 th SC, with a tiny papilla; cloacal plate divided; dorsum bright brown with a white stripe along the lower flank. Description of holotype. Adult male; head elongate (HL/HW = 1.99), distinct from neck; body elongate, SVL 495 mm; tail long (TaL/TL = 0.305), 217 mm in length; eye quite large with round pupil, eye diameter shorter than snout length (ED/SnL: 0.76) (Table 3). Head scalation. Dorsal head scales smooth, abutting along midline rather than overlapping; rostral broader than high, visible from above, touching nasals, internasals and 1 st supralabial on both sides; supralabials 8/8, first fours small, the 5 th bordering eye on both sides, the 6 th small and triangle shaped, the 7 th and 8 th distinct larger than the 6 th and the first fours; 1/1 loreal, much longer than high (2.0 mm length vs. 0.8 mm height), touching nasal, internasal, prefrontal, preocular and the 2 nd and 3 rd supralabials; 1/1 preocular; 2/2 postoculars; 2+2/2+2 temporals; suture between internasals as long as that between prefrontals; frontal bell-shaped, widened anteriorly and much narrowed posteriorly (5.6 mm length, 3.6 mm width); parietals longer than frontal (6.5 mm vs. 5.6 mm length), bordered laterally by upper temporals and postocular and anteriorly by frontal and supraoculars; 10/10 infralabials, first pair medially in contact with each other, first five in contact with anterior chin shield, the 6 th largest and touching posterior chin shield; posterior chin shields slightly longer than anterior ones (5.1 mm vs. 4.9 mm) (Fig. 4A). Body scalation. Dorsal scales smooth, in 13-13-9 rows; lateral scales, except for the outer row, disposed obliquely; vertebral scale row slightly enlarged, narrower than the outer row of scales; scale row reductions from 13 to 11 at ventrals 89 and 90 and from 11 to 9 at ventrals 96 and 99; ventrals 158, with a suture-like lateral keel; cloacal plate divided; subcaudals 106, all paired; terminal caudal scale forming a pointed cap. Dentition. Maxillary teeth 18, continuously, the posterior teeth slightly smaller than the others. Hemipenis. Hemipenis unforked, reaching 11 th SC, spinose throughout, except for the apical naked area. The spines in the terminal half of the organ are largest. Sulcus spermaticus single, extending to tip, the sulcus lip prominent. On the apical naked area, there is a tiny lobule with small spines (Fig. 4B&C). Coloration. In life, dorsal and lateral parts of the body and tail bronze-brown, laterally this color extends to the upper half of the second scale row; upper part of head olive, lip and lower parts of the head and neck yellow; iris orange on the upper part and black on anterior, posterior, and lower parts of the eye; concealed parts of lateral scales light blue; a black temporal streak extending from eye to neck; a bright vertebral stripe on the anterior part of dorsum; two lateral stripes: a white one on the lower half of the second scale row and the upper half of the outer row and the second stripe is pale brown occurring on the lower half of the outer row and the edge of ventral scale; anterior part of the body with a series of black blotches at the suture between the lateral edge of ventral and the outer row, these blotches are less distinct or absent on the middle and posterior parts of the body; ventral white. In preservation, color faded but the pattern remained, all yellow parts becoming white. Variation. Paratypes ITBCZ 6665 and ITBCZ 5945 have 7 and 9 supralabials on the right side, respectively; paratypes ITBCZ 5945 and ITBCZ 5946 have 11 dorsal scale rows before vent; paratype ITBCZ 5946 has a short suture on the lower margin of the 5 th supralabial on both sides; paratype ITBCZ 5945 has the upper half of the second dorsal row black forming a black longitudinal line (Fig. 3E); lip and lower parts of head and neck yellow in all specimens collected in July in Tuy Phong, Binh Thuan Province but white in all snakes collected in April in Thuan Nam, Ninh Thuan Province. Table 3 summarizes variation in size and scalation of type series. Sexual dimorphism. Male has fewer ventral scales (V 154–158 in 3 males, 161–170 in 5 females) and slightly longer relative tail (TaL/TL 0.305–0.309 in 2 adult males; 0.277–0.297 in 5 adult females). Etymology. We name the new species in honor of Dr. Ngô V&abreve;n Bὶnh, a Vietnamese herpetologist who passed away suddenly in 2023 at the age of 49 by a serious stroke. We recommend “Binh’s Bronzeback Snake” and “Rắn leo cây bὶnh” as the common English and Vietnamese names of the new species, respectively. Distribution. The new species is currently known from Thuan Nam, Ninh Thuan Province and Tuy Phong, Binh Thuan Province, Vietnam (Fig. 1). Field notes. All specimens were collected at night, between 7:00 PM and 11:00 PM, while sleeping on brushes at about 1–2 m above ground. The habitat in Thuan Nam, Ninh Thuan Province is a coastal hill with brushes and granite outcrops (Fig. 5B) and the habitat in Tuy Phong, Binh Thuan Province is an agricultural area with scattered large brushes. The holotype was collected on a large brush group together with paratypes ITBCZ 6664 and ITBCZ 6665 and with two other individuals of Ahaetulla fusca (Dumeril, Bibron & Dumeril). Gravid paratype ITBCZ 5946 has three eggs. Comparisons. Dendrelaphis binhi sp. nov. differs morphologically from all congeners in Indochina as follows: from D. biloreatus (Wall, 1908) by having fewer ventrals (154–170 vs. 187–199), fewer subcaudals (95–106 vs. 139–145), one (vs. two) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Smith 1943); from D. caudolineatus by having fewer ventrals in males (154–158 vs. 171–180), fewer ventrals in females (161–170 vs. 171–185), one (vs. two) supralabial entering the eye, and absence (vs. presence) of black longitudinal stripes along the entire length of the body (Rooijen & Vogel 2012); from D. cyanochloris (Wall, 1921) by having fewer midbody scale rows (13 vs. 15), fewer ventrals (154–170 vs. 181–206), fewer subcaudals (95–106 vs. 135–157), one (vs. two) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Rooijen & Vogel 2008; Jiang et al. 2020); from D. formosus by having fewer midbody scale rows (13 vs. 15), fewer ventrals (154–170 vs. 172–194), fewer subcaudals (95–106 vs. 142–162), and one (vs. two) supralabial entering the eye (Jiang et al. 2020); from D. haasi by having fewer midbody scale rows (13 vs. 15), one (vs. two) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Das 2015); from D. kopsteini Vogel & Van Rooijen, 2007 by having fewer midbody scale rows (13 vs. 15), one (vs. two) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Vogel & Rooijen 2007); from D. ngansonensis by having fewer subcaudals (95–106 vs. 117–153), one (vs. two or three) supralabial entering the eye, vertebral scale row slightly enlarged (vs. strongly enlarged), fewer maxillary teeth (18 or 19 vs. 24 or 25), and shorter hemipenis (extending to SC 10 or 11 vs. SC 18) (Ziegler & Vogel 1999; Jiang et al. 2020); from D. nigroserratus Vogel, Rooijen & Hauser, 2012 by having fewer midbody scale rows (13 vs. 15), fewer ventrals (154–170 vs. 197– 204), fewer subcaudals (95–106 vs. 148–152), one (vs. three) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Vogel et al. 2012); from D. pictus by having fewer midbody scale rows (13 vs. 15), fewer subcaudals (95–106 vs. 113–148), fewer maxillary teeth (18 or 19 vs. 23–28), one (vs. two or three) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Smith 1943; Vogel & Rooijen 2008); from D. striatus by having fewer midbody scale rows (13 vs. 15), one (vs. two or three) supralabial entering the eye, and absence (vs. presence) of black oblique bars laterally on the body (Ziegler & Vogel 1999; Vogel & Rooijen 2007); from D. subocularis by having fewer dorsal scale rows at neck (13 vs. 15) and at midbody (13 vs. 15), and fewer maxillary teeth (18 or 19 vs. 21–23) (Smith 1943; Rooijen & Vogel 2010); from D. tristis by having fewer midbody scale rows (13 vs. 15), fewer subcaudals (95–106 vs. 108–145), one (vs. two) supralabial entering the eye, and longer hemipenis (extending to SC 10 or 11 vs. SC 8) (Smith 1943); from D. vogeli by having fewer midbody scale rows (13 vs. 15), one (vs. two) supralabial entering the eye, fewer ventrals (154–170 vs. 193–197), fewer subcaudals (95–106 vs. 130–135), and vertebral scale row slightly enlarged (vs. strongly enlarged) (Jiang et al. 2020); and from D. walli Vogel & Rooijen, 2011 by having fewer ventrals (154–170 vs. 193–197), fewer subcaudals (95–106 vs. 130–135), one (vs. two) supralabial entering the eye, and vertebral scale row slightly enlarged (vs. strongly enlarged) (Vogel & Rooijen 2011)., Published as part of Nguyen, Sang Ngoc, Nguyen, Vu Dang Hoang, Le, Manh Van, Nguyen, Luan Thanh, Vo, Thi-Dieu-Hien, Vo, Ba Dinh, Che, Jing & Murphy, Robert W., 2023, A new snake of the genus Dendrelaphis Boulenger, 1890 (Squamata: Colubridae) from the coastal area of southern Vietnam, pp. 130-144 in Zootaxa 5318 (1) on pages 136-139, DOI: 10.11646/zootaxa.5318.1.6, http://zenodo.org/record/8158335, {"references":["Smith, M. A. (1943) The fauna of British India, including Ceylon and Burma: Reptilia and Amphibia. Taylor and Francis, London, 583 pp.","Rooijen, J. V. & Vogel, G. (2012) A revision of the taxonomy of Dendrelaphis caudolineatus (Gray, 1834) (Serpentes: Colubridae). Zootaxa, 3272 (1), 1 - 25. https: // doi. org / 10.11646 / zootaxa. 3272.1.1","Rooijen, J. V. & Vogel, G. (2008) A new species of Dendrelaphis (Serpentes: Colubridae) from Java, Indonesia. The Raffles Bulletin of Zoology, 56, 189 - 197.","Jiang, K., Ren, J. L., Guo, J. P., Wang, Z., Ding, L. & Li, J. T. (2020) A new species of the genus Dendrelaphis (Squamata: Colubridae) from Yunnan Province, China, with discussion of the occurrence of D. cyanochloris (Wall, 1921) in China. Zootaxa, 4743 (1), 1 - 20. https: // doi. org / 10.11646 / zootaxa. 4743.1.1","Das, I. (2015) A Field Guide to the Reptiles of Southeast Asia. Bloomsbury Publishing, London, 376 pp.","Vogel, G. & Rooijen, J. V. (2007) A new species of Dendrelaphis (Serpentes: Colubridae) from Southeast Asia. Zootaxa, 1394 (1), 25 - 45. https: // doi. org / 10.11646 / zootaxa. 1394.1.2","Ziegler, T. & Vogel, G. (1999) On the knowledge and specific status of Dendrelaphis ngansonensis (Bourret, 1935) (Reptilia: Serpentes: Colubridae). Russian Journal of Herpelotogy, 6, 199 - 208.","Vogel, G., Rooijen, J. V. & Hauser, S. (2012) A new species of Dendrelaphis Boulenger, 1890 (Squamata: Colubridae) from Thailand and Myanmar. Zootaxa, 3392 (1), 35 - 46. https: // doi. org / 10.11646 / zootaxa. 3392.1.2","Rooijen, J. & Vogel, G. (2010) On the discovery and origin of a Javan population of the Indochinese colubrid snake Dendrelaphis subocularis (Boulenger, 1888): a multivariate study. Contributions to Zoology, 79, 85 - 92. https: // doi. org / 10.1163 / 18759866 - 07903001","Vogel, G. & Rooijen, J. V. (2011) Description of a new species of the genus Dendrelaphis Boulenger, 1890 from Myanmar (Squamata: Serpentes: Colubridae). Bonn Zoological Bulletin, 60, 17 - 24."]}

Reptiles are considered an important element of Saudi Arabia's wildlife to be conserved as a priority. However, the status and distribution of the Kingdom's reptile fauna is not well understood, thus hindering the conservation initiatives. Better understanding of the taxonomy and distribution of the Kingdom's reptile fauna is important for implementing effective conservation measures.Here, we provide the new distribution record of Pristurus minimus (Arnold, 1977; common name, Arnold's Gecko) from southern Saudi Arabia. The species was recorded from the Uruq Bani Ma'arid Protected Area (UBM) of southern Saudi Arabia. Four individuals were captured from different survey sites across the UBM and their species identification was confirmed through a mixture of physical and genetic examination. These results increase the number of species from the Pristurus genus to five for Saudi Arabia and improve the understanding of the Kingdom's reptile fauna and its distribution.

Colombophis portai Hoffstetter & Rage, 1977 (Figs 3-5) Colombophis portai Hoffstetter & Rage, 1977: 174. REVISED DIAGNOSIS. — Fossil snake with pre-cloacal vertebrae medium to large in size (cl = c. 5-11 mm); the neural arch is depressed, with a shallow median notch in the posterior border; neural spine is reduced to a small tubercle with a circular or triangular outline from dorsal view, and is restricted to the posterior end of the neural arch; prezygapophyses are anterolaterally oriented and highly inclined above the horizontal plane, reaching the level of the zygosphene roof; prezygapophyzeal process is short; presence of paracotylar and parazygantral foramina is variable; synapophyses are undivided; haemal keel is broad with a single or divergent apophysis in the posterior end; subcentral foramina are usually placed close to the sagittal plane, usually small but can be enlarged or absent; the centrum length is longer than the neural arch width (cl> naw); the neural arch is longer than width (pr-po> naw) and the length is proportional to its high (pr-po = c. h). REFERRED MATERIAL. — VPPLT-0067, an anterior-trunk vertebra (Fig. 3C); VPPLT-0068 (Fig. 3D), an anterior-trunk vertebra; VPPLT-0070, a posterior-trunk vertebra; VPPLT-0430, a mid-trunk vertebra; VPPLT-0799, a mid-trunk vertebra; VPPLT-0845, a fragmentary mid-trunk vertebra; VPPLT-0869, a posterior-trunk vertebra; VPPLT-0871, a fragmentary mid-trunk vertebra; VPPLT-1006, a mid-trunk vertebra (Fig. 3A, B); VPPLT-1160, a fragmentary posterior-trunk vertebra; VPPLT-1166, a posterior-trunk vertebra; VPPLT-1253, a posterior-trunk vertebra; VPPLT-1551, two associated mid-trunk vertebrae; VPPLT-1564, nine pre-cloacal articulated vertebrae together with seven associated and badly preserved fragments of rock matrix with ribs and vertebral fragments embedded (Fig. 5); VPPLT-1731, a fragmentary mid-trunk vertebra; VPPLT-1734, a mid-trunk vertebra; VPPLT-1735, four mid-trunk vertebrae fragments associated with a posterior-trunk vertebra (Fig. 3F); VPPLT-1738, a mid-trunk vertebra; VPPLT-1739, an anterior-trunk vertebra (Fig. 3E); VPPLT-1740, five mid-trunk vertebrae fragments associated with a posterior-trunk vertebrae and a post-cloacal vertebrae (Fig. 4)., Published as part of Alfonso-Rojas, Andrés, Vanegas, Rubén D., Mariño-Morejón, Elizabeth & Cadena, Edwin-Alberto, 2023, Newly discovered fossils provide novel insights on the biology of the South American Miocene snake Colombophis Hoffstetter & Rage, 1977, pp. 377-399 in Geodiversitas 45 (13) on page 382, DOI: 10.5252/geodiversitas2023v45a13, http://zenodo.org/record/8157175, {"references":["HOFFSTETTER R. & RAGE J. C. 1977. - Le gisement de vertebres miocenes de la Venta (Colombie) et sa faune de serpents. Annales de Paleontologie (Vertebres) 63 (2): 161 - 190."]}

Colombophis sp. REFERRED MATERIAL. — More than 20 vertebral fragments (see Appendix 3), most of them correspond to partially complete or fragmentary precloacal vertebrae. REMARKS Listed specimens were assigned to Colombophis as they exhibit the following features: medium to large size (cl = c. 8 mm, po-po = c. 11 mm); slightly depressed neural arch with a shallow median notch on the posterior border; the neural spine is reduced and restricted to the posterior end of the neural arch; moderately inclined prezygapophyses which usually reach the zygosphene roof level; short prezygapophyseal process; paracotilar foramina present in most vertebrae; weakly divided synapophyses; broad haemal keel with small to absent subcentral foramina close to the sagittal plane and bear one or two small apophyses with a tubercular shape placed on the ventral margin on anteriorly to the condyle. Lateral foramina located near the base of the neural canal anteriorly to the neural arch constriction. DESCRIPTIONS Anterior-trunk vertebrae These vertebrae are shorter (cl Colombophis spp. (C. portai, C. spinosus, and C. sp.). Despite being broken on most of the specimens, the hypapophyses are postero-ventrally oriented and located behind the sub-central foramina also, hypapophyses have a circular (Fig. 3D) or a flattened shovel-like shape (Figs 3E; 6E) in posterior view. Mid-trunk vertebrae These vertebrae are the largest and most distinctive of Colombophis spp. Prezygapophyses are anterolaterally oriented, and usually reach the zygosphene roof level. Ventrally, the haemal keel is broad, usually with a pair of small sub-central foramina placed anterior to the coronal plane (Figs 3A, B; 4A, B; 5I, J; 6 A-C). The posterior end of the haemal keel usually presents a tubercular or bifid structure. Neural arch is broad and slightly depressed, visible from posterior region view. The zygantra possesses a deep foramen inside. Neural spines are restricted to the posterior region of the neural arch, being poorly developed (tubercle like) in C. portai (Figs 3A, B; 4 A-D; 5G, H), but well developed in C. spinosus where the anterior edge of this structure rarely reach the coronal plane of the vertebra (Fig. 6 A-D). Posterior-trunk vertebrae These vertebrae are easily recognized by the presence of subcentral paramedian lymphatic fossae, which create a notched A1 A2 A3 A4 A5 section between the cotyle and the synapophyses (Figs 3F; 4E, F; 6F). Pre-condylar constriction of the centrum is strong compared to anterior or mid-trunk vertebrae, which also makes the haemal keel less broad. Prezigapophyses are less inclined and do not reach the zygosphene level. In some specimens of C. portai, the posterior end of the haemal keel extends ventrally creating a structure similar to a hypapophysis (Fig. 4E,F) Ribs Ribs are only present in VPPLT-1564 specimen, unfortunately they are badly preserved. However, they seem to be fully ossified and slender, apparently longer than the vertebral centrum (Fig. 5C, D). The articular facet is smooth without a clear division on the articular facets, which corresponds to undivided synapophyses. A1 A2 A3 A4 A5 Post-cloacal vertebrae A potential post-cloacal vertebra was found associated in the C. portai specimen VPPLT-1740.(Fig.7A) This vertebra is considerably more vaulted than the associated precloacal vertebrae(Fig.4). Posteroventral blade like structures are present,resembling paired haemapophyses. The subcentral foramina faces antero-ventrally near to the base of haemapophyses. Prezygapophyses are short and more laterally oriented, with little or no inclination. The neural spine is a relatively high lamina, considerably different from the neural spine shape of Colombophis spp. Ontogenetic variation Most of the specimens belong to adults as they are fully ossified and present a medium to large size. However, VPPLT-1006 specimen (Fig. 3A, B), may represent a juvenile or sub adult individual based on its small size and the ovoid shape of the cotyle (Hsiou et al. 2010), it also lacks a developed prezygapophyseal processes, it has a narrow zygosphene and a relatively large neural canal as it happens in extant and ancient snake neonates (LaDuke 1991; Xing et al. 2018). Parazygantral foramina Parazygantral foramina are present in both species of Colombophis, but only in two specimens (VPPLT-1728 and VPPLT-1740), as well as in the holotype MNHN-VIV-6 (Figs 4A, B; 6A, B; 7). These structures appear as a single foramen with or without small pits aside, which are interconnected internally with the foramen, (Fig. 8). Pits may appear also in specimens with no clearly defined foramen (Figs 6C; 8E, F). The internal structure of the vertebrae reveals that the dorso-posterior region of the vertebra (postzygapophyses and neural arch) is highly vascularized, where the parazygantral foramina and its associated pits are interconnected with the zygantral and lateral foramina. This feature is particularly visible in VPPLT-1728 specimen, where the internal structure is infilled with a denser material that creates a perfect endocast of the vertebra (Fig. 8G, H)., Published as part of Alfonso-Rojas, Andrés, Vanegas, Rubén D., Mariño-Morejón, Elizabeth & Cadena, Edwin-Alberto, 2023, Newly discovered fossils provide novel insights on the biology of the South American Miocene snake Colombophis Hoffstetter & Rage, 1977, pp. 377-399 in Geodiversitas 45 (13) on pages 382-385, DOI: 10.5252/geodiversitas2023v45a13, http://zenodo.org/record/8157175, {"references":["HOFFSTETTER R. & RAGE J. C. 1977. - Le gisement de vertebres miocenes de la Venta (Colombie) et sa faune de serpents. Annales de Paleontologie (Vertebres) 63 (2): 161 - 190.","HSIOU A. S., ALBINO A. M. & FERIGOLO J. 2010. - Reappraisal of the south American Miocene snakes of the genus Colombophis, with description of a new species. Acta Palaeontologica Polonica 55 (3): 365 - 379. https: // doi. org / 10.4202 / app. 2009.1111","LADUKE T. C. 1991. - The fossil snakes of pit 91, Rancho La Brea, California. Contributions in Science 424: 1 - 28. https: // doi. org / 10.5962 / p. 226807","XING L., CALDWELL M. W., CHEN R., NYDAM R. L., PALCI A., SIMOES T. R., MCKELLAR R. C., LEE M. S. Y., LIU Y., SHI H., WANG K. & BAI M. 2018. - A mid-Cretaceous embryonic-toneonate snake in amber from Myanmar. Science Advances 4 (7): https: // doi. org / 10.1126 / sciadv. aat 5042"]}

Alfonso-Rojas, Andrés, Vanegas, Rubén D., Mariño-Morejón, Elizabeth, Cadena, Edwin-Alberto (2023): Newly discovered fossils provide novel insights on the biology of the South American Miocene snake Colombophis Hoffstetter & Rage, 1977. Geodiversitas 45 (13): 377-399, DOI: 10.5252/geodiversitas2023v45a13

Colombophis spinosus Hsiou, Albino & Ferigolo, 2010 (Fig. 6) Colombophis spinosus Hsiou, Albino & Ferigolo, 2010: 371. REVISED DIAGNOSIS. — Fossil snake with similar characteristics of C. portai but differs as most vertebrae are significantly shorter than high (pr-po, Published as part of Alfonso-Rojas, Andrés, Vanegas, Rubén D., Mariño-Morejón, Elizabeth & Cadena, Edwin-Alberto, 2023, Newly discovered fossils provide novel insights on the biology of the South American Miocene snake Colombophis Hoffstetter & Rage, 1977, pp. 377-399 in Geodiversitas 45 (13) on page 382, DOI: 10.5252/geodiversitas2023v45a13, http://zenodo.org/record/8157175, {"references":["HSIOU A. S., ALBINO A. M. & FERIGOLO J. 2010. - Reappraisal of the south American Miocene snakes of the genus Colombophis, with description of a new species. Acta Palaeontologica Polonica 55 (3): 365 - 379. https: // doi. org / 10.4202 / app. 2009.1111"]}

We examined the mitochondrial identity of Aegean Natrix natrix moreotica representing different morphotypes, with a focus on new material from Milos and Skyros. We found no correlation between distinct morphotypes and mitochondrial identity. Our results support that grass snake populations are polyphenetic and that southern subspecies, including island populations, show a higher variability than northern ones.

Varanus sp. Figure 2. Material. Two isolated teeth (NMB Hüe.1 and NMB Hüe.2). Description. Both teeth are incomplete, with NMB Hüe.1 missing the tip and tooth base, and NMB Hüe.2 being slightly abraded and also lacking the tooth base (Fig. 2). They are rather small, unicuspid, labiolingually compressed, and posteriorly curved; the curvature is more pronounced in NMB Hüe.1. Asharp carina runs throughout both the mesial and distal margins of both teeth, being especially prominent in NMB Hüe.1. In that specimen, both mesial and distal carinae are distinctly serrated, comprising prominent denticles; these denticles are fairly regular in size (about 0.09 mm /denticle) and gently rounded in profile, separated by interdenticle grooves. The serration of the distal carina of NMB Hüe.1 is prominent throughout its preserved length, while that of the mesial carina is more prominent at mid-height on the tooth and diminishes apically and especially basally. In NMB Hüe.2, the portions where the two carinae are preserved are eroded. Plicidentine (the infolding of dentine in the pulp cavity; Kearney & Rieppel, 2006) is not documented in either tooth, although it could have been present on the non-preserved tooth bases. The pulp cavity of both teeth is distinctive in being labiolingually compressed. Surrounding this slot-like pulp cavity, the basal surface of the tooth is covered by scallop-shaped erosional features reminiscent of Howship’s lacunae, which mark the action of osteoclasts that have destroyed dental tissue in preparation for tooth shedding (LeBlanc et al., 2023). In NMB Hüe.1, where the pulp cavity is more well exposed and visible, measurements on these scallop-shaped erosional features reveal typical diameters of around 30–40 μm (mean range around 20–50 μm), values that are consistent with our interpretation of them as Howship’s lacunae., Published as part of Georgalis, Georgios L., Mennecart, Bastien & Smith, Krister T., 2023, First fossil record of Varanus (Reptilia, Squamata) from Switzerland and the earliest occurrences of the genus in Europe, pp. 1-10 in Swiss Journal of Geosciences 116 (9) on page 3, DOI: 10.1186/s00015-023-00440-5, http://zenodo.org/record/8136616, {"references":["Kearney, M., & Rieppel, O. (2006). An investigation into the occurrence of plicidentine in the teeth of squamate reptiles. Copeia, 2006, 337 - 350.","LeBlanc, A. R. H., Palci, A., Anthwal, N., Tucker, A. S., Araujo, R., Pereira, M. F. C., & Caldwell, M. W. (2023). A conserved tooth resorption mechanism in modern and fossil snakes. Nature Communications, 14, 742."]}

Georgalis, Georgios L., Mennecart, Bastien, Smith, Krister T. (2023): First fossil record of Varanus (Reptilia, Squamata) from Switzerland and the earliest occurrences of the genus in Europe. Swiss Journal of Geosciences 116 (9): 1-10, DOI: 10.1186/s00015-023-00440-5, URL: http://dx.doi.org/10.1186/s00015-023-00440-5

4.3. Lygodactylus verticillatus Lygodactylus verticillatus from Madagascar and from Europa Island do not differ in scalation (Boettger 1913; Pasteur 1965). The slight difference in the number of precloacal pores mentioned by Pasteur (1965) was not supported by this study. Both populations show a very similar variation in coloration and in patterning. Thus, L. verticillatus from Madagascar and Europa cannot be distinguished by their external appearance or by the number of precloacal pores. The most basal node within L. verticillatus separates specimens from the southern Malagasy localities Sakaraha and Isalo from all other samples. Sakaraha lies approximately 135 km from Toliara on the main road to the Isalo National Park. The remaining sister subgroup consists of specimens exclusively from the coastal regions of Madagascar, approximately from Morondava southwards to Toliara, and the insular population. We cannot exclude an inland origin of the L. verticillatus from Europa Island. However, based on our morphological and genetic dataset, it seems more likely that the nearest relatives of the insular L. verticillatus are conspecifics from populations in the coastal regions of south-western Madagascar. We found no relevant morphological differences, and only weak genetic differences between the Malagasy and the insular population, and only weak genetic variation within the monophyletic lineage of Europa Island. This agrees with the hypothesis of a relatively recent colonization as suggested by Pasteur (1965). Europa Island is 355 km west-northwest from Toliara or 300 km southwest from Cap Saint-Vincent close to Morombe, 529 km east-northeast from Inhambane (Mozambique) and approximately 600 km south of Juan de Nova (Caceres 2003; Fricke et al. 2013). It is nearly circular (6 × 7 km) with a total area of 30 km 2 (Caceres 2003) and represents the largest island of the Îles Éparses. Europa Island belongs to the group of ‘modern’ isolated seamounts in the Channel, possibly developed during Oligocene to Miocene times (Courgeon et al. 2016). Lygodactylus verticillatus could have reached Europa Island recently by natural transoceanic dispersal or by human-mediated dispersal. While a natural dispersal would be supported by flows westward from Madagascar to mainland Africa (Ali & Huber 2010), a much simpler and thus more parsimonious alternative explanation is ship-borne dispersal. Lygodactylus verticillatus could have arrived at Europa Island e.g., with ships of Malagasy fishermen seasonally searching for nesting chelonians, or with ships of European (French) settlers who started their settlement from Toliara and lived on the island from approximately 1860 to the 1920s (Fricke et al. 2013). The amount of genetic variation found within Europa (three haplotypes differing by up to two mutations in 16S) would be in agreement with either (i) ship-borne dispersal or (ii) natural dispersal of multiple individuals, or (iii) natural dispersal of a single individual at a somewhat deeper point in time, with subsequent in-situ genetic diversification. Support for either of these hypotheses could come from a denser sampling of Malagasy populations: the presence of haplotypes identical to those on Europa in Malagasy populations would allow to reject the third hypothesis, and given that simultaneous or repeated natural dispersal to such a small islet is rather unlikely, ship-borne introduction would in this case remain as the most probable hypothesis. Only if the presence of the Europa haplotypes in Madagascar could be reasonably excluded, the third hypothesis would be supported. Lygodactylus verticillatus is adapted to dry climatic conditions. In south-western Madagascar, the species inhabits shrubs and trees in semi-arid areas as well as fences and walls in human settlements (Puente et al. 2009). On Europa Island, it is widely distributed and occurs in most natural habitats on rocks, shrubs, trees and in a dry euphorbia forest as well as in human settlements (Brygoo 1966; Sanchez et al. 2015, 2019). The conditions for L. verticillatus in the coastal regions of south-western Madagascar and on Europa are very similar, facilitating the survival on the island., Published as part of Röll, Beate, Sanchez, Mickaël, Gippner, Sven, Bauer, Aaron M., Travers, Scott L., Glaw, Frank, Hawlitschek, Oliver & Vences, Miguel, 2023, Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean, pp. 232-250 in Zootaxa 5311 (2) on pages 245-246, DOI: 10.11646/zootaxa.5311.2.4, http://zenodo.org/record/8094276, {"references":["Boettger, O. (1913) Reptilien und Amphibien von Madagascar, den Inseln und dem Festland Ostfarikas. In: Voeltzkow, A. (Ed.), Reise in Ostafrika in den Jahren 1903 - 1905. Wissenschaftliche Ergebnisse. Vol. 3. Systematische Arbeiten. Schweizerbartsche Verlagsbuchhandlung, Nagele und Sprosser, Stuttgart, pp. 269 - 375.","Pasteur, G. (1965) Recherches sur l'evolution des lygodactyles, lezards afromalgaches actuels. Travaux de l'Institut Scientifique Cherifien, Serie Zoologie, 29, 1 - 132.","Caceres, S. (2003) Etude prealable pour le classement en Reserve Naturelle des Iles Eparses. Memoire de DESS Sciences et Gestion de l'Environnement Tropical. Univ. Reunion (Laboratoire ECOMAR) & DIREN Reunion, 136 pp. Available from: http: // poupin. joseph. free. fr / pdf / caceres- 2003 - iles-eparses-classement. pdf (accessed 13 June 2023)","Fricke, R., Durville, P., Bernardi, G., Borsa, P., Mou-Tham, G. & Chabanet, P. (2013) Checklist of the shore fishes of Europa Island, Mozambique Channel, southwestern Indian Ocean, including 302 new records. Stuttgarter Beitrage zur Naturkunde, A, Neue Serie, 6, 247 - 276.","Courgeon, S., Jorry, S. J., Camoin, G. F., BouDagher-Fadel, M. K., Jouet, G., Revillon, S., Bachelery, P., Pelleter, E., Borgomano, J., Poli, E. & Droxler, A. W., (2016) Growth and demise of Cenozoic isolated carbonate platforms: New insights from the Mozambique Channel seamounts (SW Indian Ocean). Marine Geology, 380, 90 - 105. https: // doi. org / 10.1016 / j. margeo. 2016.07.006","Ali, J. R. & Huber, M. (2010) Mammalian biodiversity on Madagascar controlled by ocean currents. Nature, 463, 653 - 656. https: // doi. org / 10.1038 / nature 08706","Puente, M., Glaw, F., Vieites, D. R. & Vences, M. (2009) Review of the systematics, morphology and distribution of Malagasy dwarf geckos, genera Lygodactylus and Microscalabotes (Squamata: Gekkonidae). Zootaxa, 2103, 1 - 76. https: // doi. org / 10.11646 / zootaxa. 2103.1.1","Brygoo, E. R. (1966) Note sur les reptiles terrestres recoltes a Europa en avril 1964. Memoires du Museum national d'histoire naturelle, Serie A, 41, 29 - 32.","Sanchez, M. & Probst, J. - M. (2015) L'herpetofaune terrestre de l'ile d'Europa (Ocean Indien, Canal du Mozambique): synthese des connaissances et nouvelles donnees sur la repartition de l'ecologie des especes en vue de leur conservation. Bulletin de la Societe Herpetologique de France, 145, 63 - 76."]}

4.1. Lygodactylus pakenhami At present, the taxon ‘ pakenhami’ is considered a subspecies of L. grotei in most publications. In our study, L. grotei and L. g. pakenhami are genetically distinct lineages, confirming previous results (Gippner et al. 2021). Lygodactylus g. pakenhami from Pemba Island differs greatly in 16S and ND2 pairwise distances from mainland individuals considered as L. grotei. While L. grotei and L. g. pakenhami as far as known cannot be distinguished in scalation, they clearly differ in coloration. Furthermore, hatchlings of L. grotei and L. g. pakenhami are easily identifiable and distinguishable by the coloration of their trunks and especially of their tails. The reciprocal monophyly suggested by mitochondrial DNA, conspicuously different coloration—both in adults and hatchlings—and the high genetic distance suggest that these taxa most likely represent two distinct species. This is also in agreement with differences in the RAG1 sequences (although these data are not fully conclusive due to low sample sizes: one sequence available each for grotei and pakenhami). Therefore, we elevate L. g. pakenhami to full species status, as Lygodactylus pakenhami Loveridge, 1941, being aware that this taxonomic hypothesis requires further testing, especially from nuclear-encoded DNA data sets, ideally at the phylogenomic level. Lygodactylus pakenhami is endemic on Pemba Island, the northernmost and second largest island of the Zanzibar Archipelago. Pemba lies off the continental shelf and is surrounded by water 500 to 850 m deep (Moreau & Pakenham 1941). Geological evidence indicates that Pemba Island was separated from the African mainland by faulting that produced the Pemba Channel, possibly during the late Miocene or early Pliocene, 6 million years ago (Stockley 1942; Clarke & Burgess 2000). Corresponding to its long period of isolation, Pemba is characterized by a remarkable number of endemic species, including plants, mammals and reptiles (R̂dder et al. 2010). The simplest explanation for the existence of L. pakenhami on Pemba is vicariance. Its nearest extant relative, L. grotei, is widely distributed in south-eastern Tanzania and northern Mozambique, including the coastal regions. Presumably, a population of an ancestor of L. grotei and L. pakenhami already existed on Pemba Island before its separation from the mainland. Thereafter, there was no further genetic exchange between the continental and the insular population, so that the latter evolved isolated from its continental congener. While a natural dispersal from Tanzania after the separation of Pemba Island cannot be excluded, it is considered as unlikely because the geographic position of Pemba suggests that the East African Coastal Current or a similar paleocurrent conveyed any drifting material into the open ocean very rapidly (Moreau & Pakenham 1940; Hawlitschek et al. 2016a). A recent, human-mediated transportation is unlikely, as after such a short period of isolation we would expect the sharing of mitochondrial haplotypes between mainland and island populations, which we did not detect. Although only two samples yielding DNA sequences of L. grotei have precise georeferenced information, one of these was collected on the mainland directly opposite Pemba Island. However, as is always the case with inferences of allopatric occurrence, we cannot fully exclude that undiscovered populations of L. pakenhami may occur on the African mainland, which would invalidate our biogeographic hypotheses but not our taxonomic conclusions. In contrast to Pemba Island, Zanzibar and Mafia islands lie on the continental shelf and are separated from the mainland only by shallow waters with an average depth of 30 to 35 m, rarely 50 m (Moreau & Pakenham 1941). Geological data of coastal eastern Africa point to a land connection of Zanzibar and Mafia islands with the African mainland up to the end of the Pleistocene, probably only 10,000 –18,000 years ago (Stockley 1942; Clarke & Burgess 2000). Zanzibar and Mafia are inhabited by L. grotei, L. picturatus and L. viscatus, all conspecific with continental populations. Presumably, the distribution of these three species on the islands is probably best explained by recent vicariant events. In contrast to L. pakenhami on Pemba Island, the species on Zanzibar and Mafia have been isolated on the islands for a relatively short time, about 10,000 years. Additionally, both natural dispersal over water after the separation of Zanzibar and Mafia from the mainland and/or a human-mediated transportation due to long-standing trading in the recent past cannot be excluded., Published as part of Röll, Beate, Sanchez, Mickaël, Gippner, Sven, Bauer, Aaron M., Travers, Scott L., Glaw, Frank, Hawlitschek, Oliver & Vences, Miguel, 2023, Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean, pp. 232-250 in Zootaxa 5311 (2) on page 244, DOI: 10.11646/zootaxa.5311.2.4, http://zenodo.org/record/8094276, {"references":["Gippner, S., Travers, S. L., Scherz, M. D., Colston, T. J., Lyra, M. L., Ashwini, V. M., Multzsch, M., Nielson, S. V., Rancilhac, L., Glaw, F., Bauer, A. M. & Vences, M. (2021) A comprehensive phylogeny of dwarf geckos of the genus Lygodactylus, with insights into their systematics and morphological variation. Molecular Phylogenetics and Evolution, 165, 107311. https: // doi. org / 10.1016 / j. ympev. 2021.107311","Loveridge, A. (1941) New geckos (Phelsuma and Lygodactylus), snake (Lepotyphlops) and frog (Phrynobarachits) from Pemba Island, East Africa. Proceedings of the Biological Society, Washington, 54 (8), 175 - 178.","Stockley, G. M. (1942) The geology of the Zanzibar Protectorate and its relation to the East African mainland. Geological Magazine, 79 (4), 233 - 240. https: // doi. org / 10.1017 / S 0016756800073921","Clarke, G. P. & Burgess, N. D. (2000) Geology and geomorphology. In: Burgess, N. D. & Clarke, G. P. (Eds.), Coastal Forests of Eastern Africa. IUCN, Gland, pp. 29 - 40.","Moreau, R. E. & Pakenham, R. H. W. (1940) The land vertebrates of Pemba, Zanzibar, and Mafia: a zoogeographical study. Proceedings of the Zoological Society, London, 110 (A), 97 - 128. https: // doi. org / 10.1111 / j. 1469 - 7998.1941. tb 08463. x","Hawlitschek, O., Garrido, S. R. & Glaw, G. (2016 a). How marine currents influenced the widespread natural overseas dispersal of reptiles in the Western Indian Ocean. Journal of Biogeography, 44, 1426 - 1440. https: // doi. org / 10.1111 / jbi. 12940"]}

4.2. Lygodactylus insularis Lygodactylus insularis is an endemic species on Juan de Nova, a very small island with a total area of approximately 5 km 2, 280 km from mainland Africa (East Mozambique) and 120 km to the nearest point on the Malagasy coast (Caceres 2003; Lambs et al. 2016). Pasteur (1965) already included L. insularis in the L. capensis group and assumed that L. insularis originated from either L. c. capensis or L. c. grotei. Lygodactylus capensis and L. insularis share the same number of precloacal pores and a superficially similar coloration of the adults. However, they strongly differ in the status of the subcaudals and in the appearance of their hatchlings. It is thus impossible to discern the relationships of L. insularis, L. capensis and L. grotei by means of scalation or coloration alone. Our molecular phylogenetic analyses reveal that L. insularis is embedded within the L. capensis group, clearly indicating its African origin and, in so far, supporting Pasteur (1965). While the single gene as well as the multigene analyses fully support closer relationships between L. insularis and L. capensis than between L. insularis and L. grotei, the position of L. insularis within the clade formed by L. insularis, L. nyaneka, L. capensis sensu stricto and six L. aff. capensis lineages (provisionally named as “groups”; here, the last two of these are referred to as L. capensis sensu lato) is not clearly resolved. For a taxonomic revision of the L. capensis complex—one is stated to be in progress by Travers et al. unpublished (in Marques et al. 2020)—we strongly recommend a broader incorporation of genetic markers to resolve the conflicting topologies since our multigene dataset consists of only few markers especially for L. nyaneka and for the numerous groups forming L. capensis sensu lato. The clade including L. grotei and L. pakenhami is defined as the sister clade of L. insularis, L. nyaneka and L. capensis sensu lato. Lygodactylus insularis is recovered as a monophyletic group based on mtDNA, probably resulting from a single dispersal event. Furthermore, the species based on the sampling herein exhibits little intraspecific genetic variability, possibly due to the tiny size of the island resulting in a panmictic population. Members of the genus Lygodactylus successfully crossed the Mozambique Channel from mainland Africa back to Madagascar one or perhaps two times (R̂ll et al. 2010; Mezzasalma et al. 2017; Gippner et al. 2021). The ancestor of L. insularis probably began at least one further traverse in eastward direction ending on Juan de Nova, an island in the middle of the channel. Juan de Nova developed as the uplifted top of a seamount that probably was formed during the Cretaceous along the Davie Fracture (Riaux-Gobin & Witkowski 2012). Ali & Hedges (2022) suggest that it may have remained emerged as a low elevation atoll since the Early Palaeocene. In such a scenario, the dispersal of the ancestors of L. insularis may have taken place as early as 22.3 (confidence interval 15.6–30.2) mya, the stem age of the L. capensis group estimated in the Lygodactylus timetree of Gippner et al. (2021). Results of palaeogeographic reconstructions and palaeo-oceanographic modelling suggest strong surface currents in the Mozambique Channel flowing from northeast Mozambique and Tanzania eastward towards Madagascar during the Palaeogene (66 to 23 mya), which would support this scenario (Ali & Huber 2010). While the main contemporary direction of oceanic surface currents is westward, the situation in the Mozambique Channel is very complex with numerous eddies and possible countercurrents (Hawlitschek et al. 2016a). This also suggests that dispersal could have occurred during a more recent period of mainly westward marine currents. An alternative hypothesis, dispersal via short-lived Cenozoic land-bridges between mainland Africa and Madagascar (e.g., Mazza et al. 2019; Masters et al. 2020), is disputed and has been ruled out based on current geological reconstructions (e.g., Ali & Huber 2010; Ali & Hedges 2022). It also must be emphasized that despite a reasonable sampling of Eastern Africa, especially Mozambique (with L. capensis group sequences from 14 localities available so far), it is possible that a still undiscovered species representing the sister lineage of L. insularis may exist in this region. A somewhat similar situation was uncovered in the frogs of the Ptychadena mascariensis complex in which the sister lineage of the Malagasy species has only been recorded from a small area of Malawi and could have easily been overlooked without intensive sampling efforts (Zimkus et al. 2017). If a closer relative of L. insularis was found on the African mainland, it would lead to younger age estimates of the colonization event of Juan de Nova by these geckos., Published as part of Röll, Beate, Sanchez, Mickaël, Gippner, Sven, Bauer, Aaron M., Travers, Scott L., Glaw, Frank, Hawlitschek, Oliver & Vences, Miguel, 2023, Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean, pp. 232-250 in Zootaxa 5311 (2) on pages 244-245, DOI: 10.11646/zootaxa.5311.2.4, http://zenodo.org/record/8094276, {"references":["Caceres, S. (2003) Etude prealable pour le classement en Reserve Naturelle des Iles Eparses. Memoire de DESS Sciences et Gestion de l'Environnement Tropical. Univ. Reunion (Laboratoire ECOMAR) & DIREN Reunion, 136 pp. Available from: http: // poupin. joseph. free. fr / pdf / caceres- 2003 - iles-eparses-classement. pdf (accessed 13 June 2023)","Lambs, L., Mangion, P., Mougin, E. & Fromard, F. (2016) Water cycle and salinity dynamics in the mangrove forests of Europa and Juan de Nova Islands, southwest Indian Ocean. Rapid Communications in Mass Spectrometry, 30, 311 - 320. https: // doi. org / 10.1002 / rcm. 7435","Pasteur, G. (1965) Recherches sur l'evolution des lygodactyles, lezards afromalgaches actuels. Travaux de l'Institut Scientifique Cherifien, Serie Zoologie, 29, 1 - 132.","Marques, M. P., Ceriaco, L. M. P., Buehler, M. D., Bandeira, S. A., Janota, J. M. & Bauer, A. M. (2020) A revision of the dwarf geckos, genus Lygodactylus (Squamata: Gekkonidae), from Angola, with description of three new species. Zootaxa, 4853 (3), 301 - 352. https: // doi. org / 10.11646 / zootaxa. 4853.3.1","Mezzasalma, M., Andreone, F., Aprea, G., Glaw, F., Odierna, G. & Guarino, F. M. (2017) Molecular phylogeny, biogeography and chromosome evolution of Malagasy geckos of the genus Lygodactylus (Squamata, Gekkonidae). Zoological Scripta, 46, 42 - 54. https: // doi. org / 10.1111 / zsc. 12188","Gippner, S., Travers, S. L., Scherz, M. D., Colston, T. J., Lyra, M. L., Ashwini, V. M., Multzsch, M., Nielson, S. V., Rancilhac, L., Glaw, F., Bauer, A. M. & Vences, M. (2021) A comprehensive phylogeny of dwarf geckos of the genus Lygodactylus, with insights into their systematics and morphological variation. Molecular Phylogenetics and Evolution, 165, 107311. https: // doi. org / 10.1016 / j. ympev. 2021.107311","Riaux-Gobin, C. & Witkowski, A. (2012) Small-sized and discoid species of the genus Cocconeiopsis (Bacillariophyta) on Holothuria atra (Juan de Nova, Mozambique Channel). Phytotaxa, 54, 43 - 58. https: // doi. org / 10.11646 / phytotaxa. 54.1.5","Ali, J. R. & Hedges, S. B. (2022) A review of geological evidence bearing on proposed Cenozoic land connections between Madagascar and Africa and its relevance to biogeography. Earth-Science Reviews, 232, 104103. https: // doi. org / 10.1016 / j. earscirev. 2022.104103","Ali, J. R. & Huber, M. (2010) Mammalian biodiversity on Madagascar controlled by ocean currents. Nature, 463, 653 - 656. https: // doi. org / 10.1038 / nature 08706","Hawlitschek, O., Garrido, S. R. & Glaw, G. (2016 a). How marine currents influenced the widespread natural overseas dispersal of reptiles in the Western Indian Ocean. Journal of Biogeography, 44, 1426 - 1440. https: // doi. org / 10.1111 / jbi. 12940","Mazza, P. P. A., Buccianti, A. & Savorelli, A. (2019) Grasping at straws: a re-evaluation of sweepstakes colonisation of islands by mammals. Biological Reviews, 94, 1364 - 1380. https: // doi. org / 10.1111 / brv. 12506","Masters, J. C., Genin, F., Zhang, Y., Pellen, R., Huck, T., Mazza, P. P. A., Rabineau, M., Doucoure & M., Aslanian, D. (2020) Biogeographic mechanisms involved in the colonization of Madagascar by African vertebrates: Rifting, rafting and runways. Journal of Biogeography, 48, 492 - 510. https: // doi. org / 10.1111 / jbi. 14032","Zimkus, B. M., Lawson, L. P., Barej, M. F., Barratt, C. D., Channing, A., Dash, K. M., Dehling, J. M., Du Preez, L., Gehring, P. - S., Greenbaum, E., Gvozdik, V., Harvey, J., Kielgast, J., Kusamba, C., Nagy, Z. T., Pabijan, M., Penner, J., R ˆ del, M. - O., Vences, M. & L ˆ tters, S. (2017) Leapfrogging into new territory: How Mascarene ridged frogs diversified across Africa and Madagascar to maintain their ecological niche. Molecular Phylogenetics and Evolution, 106, 254 - 269. https: // doi. org / 10.1016 / j. ympev. 2016.09.018"]}