Search

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.

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.

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.

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 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.

Müller, Rodrigo Temp, Garcia, Mauricio Silva, Fonseca, André de Oliveira (2022): A new proterochampsid (Archosauriformes: Proterochampsia) from the Late Triassic of southern Brazil and the emergence of archosaurian hind limb traits. Journal of Systematic Palaeontology 20 (1): 1-19, DOI: 10.1080/14772019.2022.2128913, URL: http://dx.doi.org/10.1080/14772019.2022.2128913

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.

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.

Preservation of soft parts (collagen fibres, blood vessels and cells) in extinct vertebrates is rare and usually limited to fossilized bone and cartilage. Well-preserved coarse collagenous fibre bundles embedded in a mineralized matrix of tendons, as well as numerous hollow, tubular structures consistent morphologically with fibril bundles, blood vessels and associated cells, were identified in ossified tendons of Late Cretaceous ornithischians from North America and Central East Asia. Detailed, high-accuracy imaging, along with spectroscopic characterization of those fibrous structures and comparison with ossified tendons of modern-day turkeys, support the proposition that physiologically driven tendon ossification is common for avians and non-avian dinosaurs. The examined soft parts were preserved through the pathway of iron-induced crosslinking and alumino-silification, documenting a variety of pathways for the preservation of soft parts, depending on the burial environment. For the first time, the structure of dinosaur fossilized tendons is analysed in detail, revealing shared histogenetic principles with modern birds and the nature of preservation.

Pinacosaurus grangeri (ZPAL MgD-II/32) The caudal tendons reveal extensive secondary remodelling, with at least three generations of secondary osteons (Fig. 3A–E). The cross-section of the flattened, elliptical tendon shows a densely secondary remodelled inner part (Fig. 3B, C) in the thickest place and more loosely arranged secondary osteons in the thinner parts of the cross-section and the periphery of the tendon (Fig. 3D). The latter is mainly composed of bundles of mineralized fibres and few vascular canals. Although, the secondary osteons mainly overlap each other in some places, between them patches of mineralized fibres are present (interfascicular spaces; Fig. 3E, F). The cross-section of the second sample from the same individual shows three circular tendons. The two bigger tendons (2.1 and 1.7 mm in diameter) are extensively secondarily remodelled (Fig. 3A) and almost nonvascular mineralized fibres are present on their periphery. In contrast, the smallest tendon (0.6 mm in diameter) is exclusively composed of mineralized fibres, weakly vascularized. The difference in the structure between the smallest and two larger tendons result from different places of sectioning: the smallest is from the terminal part of the tendon, and the bigger ones closer to the middle part. NANOSTRUCTURE OF THE TENDONS Detailed scanning electron microscopy (SEM) observation of vascular canals of fossilized tendons revealed the presence of mostly randomly oriented fibrous matrix in all three studied ornithischian samples (Fig. 4A–G). Comparative SEM images of modern-day turkey tibialis cranialis tendons (Fig. 4I–H) reveal similar fibrous structures. The observed structures exhibit a hierarchic pattern – individual fibres are gathered in bundles (or fascicles). The diameters of the bundles range from 0.7 to 2 µm (Supporting Information, Fig. S1) and the bundles are located in the walls of vascular canals of diameters varying between a dozen and 50 µm. The fibrous structures are visible only in the vascular walls exposed on the surfaces of freshly broken tendons. The EDS spectra collected from the fibrous structures and surrounding mineral matrix of all studied tendons reveal oxygen, calcium and phosphorus as the main components in all studied taxa (Fig. 4K, L), which is a typical elemental signature of modern and fossil bones. To visualize vascular canal shape and geometry, topographic microscope imaging (Fig. 5A, B) of a Homalocephale calathocercos tendon sample was applied. The results indicate that the observed fibrous structures are located deep inside the vascular canals exclusively and incorporated in their walls along their length. In addition to the surface morphology imaging, information of vascular canal topography in the tendon was collected using an atomic force microscope (AFM) in contact mode. The generated images (lock-in amplitude and lockin-phase, Fig. 5B–G) show the nanostructure of the fibres. Furthermore, AFM imaging on an individual fibre (Fig. 5G, H) reveals its striped pattern and the performed measurements allow estimation of a periodicity measuring about 24 nm (Fig. 5I, J; Supporting Information, Data S2). PRESERVATION OF SOFT PARTS All fossil tendon samples were demineralized (see Material and methods) to remove the apatite matrix and to assess whether any soft parts were preserved. No fossilized soft parts were obtained from the extract of Pinacosaurus grangeri. The demineralization of the tendons of Homalocephale calathocercos (Fig. 6A) revealed an extract consisting of individual brown- to orange-coloured tubular structures, as well as a dense network of vascular canals still attached to a translucent layer loosening from the tendon (Fig. 6C, D; Supporting Information, Fig. S2). Furthermore, the Edmontosaurus regalis sample revealed the presence of numerous rust- to brown, dark to translucent, elongated, bifurcated and H-shaped structures (Fig. 6B). Additionally, over a dozen tubular, branched structures (Fig. 6C, D) seems to be morphologically consistent with blood vessels or fibril bundle layers lining the vascular canals observed with SEM. Among them, ovate to tent-shaped cell-like structures attached to the surfaces of vessels or bundle layers were found (Fig. 6F–K). Other cell-like structures, more spindle-shaped, and projecting branching structures (Fig. 6L–N, black asterisks) were usually observed in isolation (Fig. 6L, N), sometimes attached to the fibrous matrix. The EDS spectra collected from the demineralized parts of the tendons show that their elemental composition is different from the mineral matrix of the tendons. In the case of the external sheath of a partially demineralized tendon of H. calathocercos, which detached from the tendon, a dense network of vessel-like tubes was observed (Fig. 6C, D; compare also: Supporting Information, Fig. S2A, B). In this sample, oxygen, silicon and aluminium dominate (Fig. 4E), while remains of phosphates are still present (Supporting Information, Fig. S2C). The fossilized vascular-like tubular structures with attached cell-like structures (Fig. 6F–K), as well as individual cells (Fig. 6L–N) extracted from the samples taken from E. regalis and H. calathocercos, are composed mainly of iron and sulphur with smaller addition of oxygen and silicon (Fig. 6P). ORGANIC MATTER RESIDUES Total carbon (TC), total organic carbon (TOC) and total inorganic carbon (TIC), as well as total sulphur (TS), contents were measured (see Material and methods) to document the preservation of organic matter in all studied fossil samples. Obtained results demonstrate TOC values equal to 1.28 wt% for Homalocephale calathocercos, 2.57 wt% for Pinacosaurus grangeri, and 2.81 wt% for Edmontosaurus regalis tendon samples. An elevated level of total sulphur (4.54 wt%) was also reported for the latter. Furthermore, the control sample of sediment associated with E. regalis tendons reveals the highest level of TOC, up to 6 wt% with sulphur content below 0.5 wt%. SPECTROSCOPIC STUDIES AND MASS SPECTROMETRY FTIR spectroscopic studies, intended mainly to identify organic residues in the fossil samples, were performed only on the selected tendon samples that were never glued or treated with any organicbased materials (e.g. consolidants). Therefore, the Pinacosaurus grangeri tendon sample was not included (Supporting Information, Data S3). A modern-day turkey (Meleagris gallopavo, GIUS-12-3741) was a control sample. FTIR studies of H. calathocercos, E. regalis and M. gallopavo show a band arrangement pointing to co-association of various minerals, including phosphate (PO 4) 3–, carbonate (CO 3) 2– and iron oxide (FeO 4) 5–, as well as silicon dioxide (SiO 2) moieties, suggesting the presence of carbonate apatite, goethite and silicates (Fig. 7). The results are thus consistent with the EDS spectra (Fig. 4H, L, P) for the same samples. The FTIR measurements of demineralized parts of tendon samples from H. calathocercos and E. regalis revealed also signals associated with organic residues (Fig. 5B, D), especially well visible due to several bands in the 1550–1800 cm –1 corresponding to the amide I (Fig. 7F, G). A percentage proportion of each component in the amide I band was computed as a ratio of a fractional area of the suitable peak after the band fitting and the sum of the areas of the peaks belonging to the amide I band (Byler & Susi, 1986; Jackson & Mantsch, 1995; Litvinov et al., 2012). As a result, two intensive bands were observed at 1633 and 1668 cm –1 with 64 and 36% proportion between β- sheet structures and turns (Litvinov et al., 2012) for H. calathocercos, as well as three bands at 1630, 1663 and 1690 cm –1 with 56, 25 and 19% for E. regalis. Two other marginal bands (at 1594 and 1714 cm –1) were also detected in both samples and correspond to the lipid signal, including fatty acids. The comparison of the fossilized tendons with the turkey tendon (Fig. 7E, H) strongly support the possibility of organic (proteinaceous) preservation in the fossil samples, especially based on the comparison of the amide I fingerprint region (Fig. 7F–H). Mass spectra (Supporting Information, Data S4) were collected both from the demineralized sample of H. calathocercos and E. regalis tendons across the regions of interest with a mass-to-charge ratio (m / z) range of 1–150 Da, the fingerprint region for lower mass amino acid anions (Surmik et al., 2017). There, various non-organic (Si-containing and Fe-containing), as well as organic (N-containing), species were identified. The detected ions CH 4 N + (at m / z 30.03 Da), C 2 H 3 N + (at m / z 41.03 Da), C 2 H 6 N + (at m / z 44.05 Da), C 3 H 3 N + (m / z 53.02 Da), C 3 H 6 N + (at m / z 56.04 Da), C 4 H 6 N + (at m / z 68.05 Da), C 4 H 8 N + (at m / z 70.07 Da), C 4 H 10 N + (at m / z 72.08 Da) and C 5 H 10 N + (at m / z 84.08 Da) are related to amino-acids (Surmik et al., 2017) (see also fibrous matrix (white arrow) and cell-like structures attached to the surface (yellow arrows); H, group of cells-like structures attached to the surface; I, lateral view of a cell-like structures (white arrow) attached to the fibrous surface of the vessel-like structures; J, K, examples of cells on the fibrous surface of vessels; L–N, osteocyte-like cells with branching cytoplasmic processes (white arrows) attached to the fibrous matrix (yellow arrow); O, example of isolated vessel-like tubular structure with attached cells (yellow asterisk indicate spot for EDS survey); P, AFM 3D height profile of an individual cell (rectangle in O) documenting the height difference between the cell and the surface of the vessel; the EDS spectrum revealing the elemental composition of the studied cell as iron and sulphur with addition of oxygen and silicon. Supporting Information, Data S4). The presence of the Si- and Fe-containing species correlates with the results of the EDS surveys (Fig. 6E, P) and confirms the mineralogical composition of the studied samples., Published as part of Surmik, Dawid, Słowiak-Morkovina, Justyna, Szczygielski, Tomasz, Wojtyniak, Marcin, Środek, Dorota, Dulski, Mateusz, Balin, Katarzyna, Krzykawski, Tomasz & Pawlicki, Roman, 2023, The first record of fossilized soft parts in ossified tendons and implications for the understanding of tendon mineralization, pp. 747-766 in Zoological Journal of the Linnean Society 198 (3) on pages 752-758, DOI: 10.1093/zoolinnean/zlad001, http://zenodo.org/record/8141891, {"references":["Byler DM, Susi H. 1986. Examination of the secondary structure of proteins by deconvolved FTIR spectra. Biopolymers 25: 469 - 487.","Jackson M, Mantsch HH. 1995. The use and misuse of FTIR spectroscopy in the determination of protein structure. Critical ReVieaes in Biochemistry and Molecular Biology 30: 95 - 120.","Litvinov RI, Faizullin DA, Zuev YF, Weisel JW. 2012. The α- helix to β- sheet transition in stretched and compressed hydrated fibrin clots. Biophysical Journal 103: 1020 - 1027.","Surmik D, Rothschild BM, Pawlicki R. 2017. Unusual intraosseous fossilized soft tissues from the middle triassic nothosaurus bone. The Science of Nature 104: 25."]}

Edmontosaurus regalis (UAMES 52615) Both cross-sectioned tendons show a highly porous structure built of fibrous primary matrix (Fig. 1A–C). The general structure of the sectioned tendons is homogenous. The matrix is composed of coarse collagenous fibre bundles containing numerous bone cell lacunae (Fig. 1A, C). The vascular canals forming the porous structure of the tendons are young Haversian canals (Fig. 1A–C). The cross-sectioned larger tendon shows few scattered secondary osteons in the centre of the tendon surrounded by collagen-fibre bundles (Fig. 1D). The tendon of a slightly smaller cross-section (3.5 mm diameter; Fig. 1A, B) is more porous than the larger one (6 mm; Fig. 1C, D). The different structure between the sampled tendons may result from a different location in the body., Published as part of Surmik, Dawid, Słowiak-Morkovina, Justyna, Szczygielski, Tomasz, Wojtyniak, Marcin, Środek, Dorota, Dulski, Mateusz, Balin, Katarzyna, Krzykawski, Tomasz & Pawlicki, Roman, 2023, The first record of fossilized soft parts in ossified tendons and implications for the understanding of tendon mineralization, pp. 747-766 in Zoological Journal of the Linnean Society 198 (3) on page 751, DOI: 10.1093/zoolinnean/zlad001, http://zenodo.org/record/8141891

Homalocephale calathocercos (MPC-D 100/1201) The cross-section of the tendon shows dispersed primary osteons embedded in mineralized fibrous primary matrix (Fig. 2A–F). In some cases, the structural remnant of a primary osteon encapsulates a smaller concentric ring of lamellar bone apparently separated with a cement line, appearing to be a small secondary osteon deposited in the centre of a primary osteon (Fig. 2D). These structures probably represent a first generation of secondary osteons (a short cycle of resorption of primary tissue and subsequent deposition of secondary tissue along blood vessels). The osteons are bigger and denser in the innermost part of the tendon, and become less abundant and smaller closer to the periphery (Fig. 2A, B, E). Under normal light, in both the longitudinal and cross-section, visible bone cell lacunae are numerous in the centre of the tendon, but rare to absent close to the external margin of the tendon (Fig. 2A, E). Under polarized light, the longitudinal section shows bundles of fibres running in a herringbone-like pattern along the tendon (Fig. 2F), in the cross-section the fibres form a lattice-like pattern (Fig. 2B, C). Thus, the osteons are surrounded by coarse collagenous fibre bundles. The width of the longitudinally sectioned tendon varies from 1.1 to 1.4 mm, its total length is 7 mm., Published as part of Surmik, Dawid, Słowiak-Morkovina, Justyna, Szczygielski, Tomasz, Wojtyniak, Marcin, Środek, Dorota, Dulski, Mateusz, Balin, Katarzyna, Krzykawski, Tomasz & Pawlicki, Roman, 2023, The first record of fossilized soft parts in ossified tendons and implications for the understanding of tendon mineralization, pp. 747-766 in Zoological Journal of the Linnean Society 198 (3) on page 751, DOI: 10.1093/zoolinnean/zlad001, http://zenodo.org/record/8141891

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"]}

The biological diversity of the Horn of Africa is one of the least studied in the world. Yet the Horn supports rich communities of species that are mostly endemic to the region. Here we study the diversity of Sand Snakes (Psammophis) in East Africa, their phylogeny and systematics. Previous studies have unveiled several cryptic and potentially undescribed species of Psammophis that occur in the Horn and their taxonomic status has remained unclear to this day. We used sequence data from two mitochondrial and one nuclear genes to reconstruct the phylogeny of the genus, in which we included newly obtained samples of six different Psammophis species from Somalia, Ethiopia, Eritrea, Sudan, and Egypt. Our aim was to assess the status of some of the undescribed species, examine the level of intraspecific genetic variation within individual species, improve our understanding of the species distributions, and contribute to the taxonomy of the genus. Our results confirm the existence of two undescribed species, one in eastern Somalia, which we formally describe as new, and one in southern Ethiopia that we refer to as Psammophis cf. sudanensis in accordance with previous studies. Further, we provide first genetic data for the nominotypical subspecies of P. punctulatus and confirm the species status for its subspecies P. trivirgatus. In addition, we provide new genetic data for P. tanganicus from Ethiopia and Somalia, and range extension records for P. rukwae from Eritrea and Ethiopia and for P. aegyptius from Somalia. Our findings contribute considerably to our understanding of the diversity and distribution of Psammophis in East Africa.

Yushchenko, Platon V., Grismer, L. Lee, Bragin, Andrey M., Dac, Le Xuan, Poyarkov, Nikolay A. (2023): A new species of the genus Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from a lowland monsoon forest in Southern Vietnam. Zootaxa 5306 (5): 537-550, DOI: 10.11646/zootaxa.5306.5.2, URL: http://dx.doi.org/10.11646/zootaxa.5306.5.2

Hemiphyllodactylus cattien sp. nov. Cat Tien Slender Gecko Figs. 3 & 4, Table 2 ZooBank registration: urn:lsid:zoobank.org:act: 4B7A2B89-BB4C-4F22-B0CC-F4E07B354327 Holotype. Adult female (VRTC NAP-10586; formerly ZMMU Re-17039) collected along the trail to Lake Bau Sau, Nam Cat Tien sector, Cat Tien National Park, Dong Nai Province, southern Vietnam (GPS data 11.4458° N, 107.3864° E WGS; 140 m a.s.l.) on 14 February 2021 at 18:00 h by P.V. Yushchenko and N.A. Poyarkov. Diagnosis. Hemiphyllodactylus cattien sp. nov. can putatively be distinguished from all other congeners by possessing the unique combination of having a maximum SVL of 35.1 mm; trunk slightly elongate and gracile (AG/ SVL 0.52); seven chin scales; enlarged postmentals; five circumnasal scales; three intersupranasals (= postrostrals); nine supralabials; nine infralabials; 20 longitudinal rows of dorsal scales and 11 longitudinal rows of ventral scales at midbody contained within one eye diameter; 4444 digital formulas on hands and 5555 formulas on feet; five subdigital lamellae on the first finger; six subdigital lamellae on the first toe; 19 perforated precloacal scales that extend part-way onto the proximal femoral region; one cloacal spur on each side; no platelike subcaudal scales; a weak dark-colored postorbital stripe extending to at base of neck; no dorsolateral light-colored spots or dark-clored dorsolateral stripe on trunk; no dark-colored ventrolateral stripe on trunk; no dark-clored paravertebral markings on trunk; no light-colored postsacral marking bearing anteriorly projecting arms; and caecum and gonads darkly pigmented. Description of holotype (Table 2). Adult female SVL 35.1 mm (Fig. 4a); head moderate, (HL/SVL 0.24, HW/ SVL 0.16) triangular in dorsal profile (Fig. 4c), depressed, distinct from neck; lores flat to slightly convex; rostrum moderate in length (NE/HL 0.32); prefrontal region weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate (NE/HL 0.32), rounded in dorsal profile; eye large (ED/HL 0.26); ear opening elliptical, small (Fig. 4b); eye-ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by large supranasals (Fig. 4c); three equally sized intersupranasals (= postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (= circumnasals); nine (R,L) rectangular supralabials tapering to below posterior margin of eye; nine (R,L) rectangular infralabials not tapering to below posterior margin of eye; scales of rostrum, lores, top of head, and occiput small, raised, those of rostrum largest; dorsal superciliaries flat, mostly square, subimbricate, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by two postmentals (Fig. 4d); postmentals in contact with first infralabial and bordered laterally by a slightly smaller chin shield; five lateral chin shields; gular scales small, subimbricate, grading posteriorly into slightly larger, subimbricate throat and even larger pectoral scales which grade into slightly larger, subimbricate ventrals. Body moderate in stature, trunk not noticeably elongate (AG/SVL 0.52), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 20 dorsal scales at midbody contained within one eye diameter; ventral scales flat, subimbricate, larger than dorsal scales, 11 ventral scales contained within one eye diameter; precloacal scales larger than abdominal scales; 19 perforated, slightly enlarged femoroprecloacal scales in an angular series, scale at apex has a pit; single enlarged tubercle (spur) on lateral margin of tail base (Fig. 4e); forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales slightly raised, subimbricate; all digits except digit I well developed; digit I vestigial, clawless; distal subdigital lamellae of digits II–V divided, angular and U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 4444 (R,L); five transversely expanded lamellae on digit I; claws on digits II—V well-developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; hind limbs short, more robust than forelimbs, covered with slightly raised, juxtaposed scales dorsally and by larger, flat subimbricate scales anteriorly and ventrally; plantar scales slightly raised, subimbricate; all digits except digit I well developed; digit I vestigial, clawless; distal subdigital lamellae of digits II–V divided, angular and U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 5555 (R,L); six transversely expanded lamellae on digit I; claws on digits II–V well developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; tail regenerated, 13.7 mm in length. Raw morphometric data (in mm) are TL 13.7 (regenerated), AG 18.4 HL 8.3, HW 5.9, SN 4.0, NE 2.7, ED 2.2, and SW 1.3. Coloration in life. Dorsal ground color of head and body nearly unicolor wheat-gray with dark-brown and white stippling (Fig. 3); dark-colored, diffuse stripe, bordered dorsally by a white band, extends from rostral scale to forelimb insertion (Fig. 3); another dark-colored U-shaped stripe can be distinguished below the eye; dark-colored, diffuse markings on the head, limbs and trunk are almost indistinguishable in color; sacral region with two pairs of more or less distinct light-colored spots; gular, pectoral and abdominal regions beige with dark stippling; stippling dense on underside of limbs; the regenerated tail is totally dark-gray bearing scattered black and white scales (Fig. 3). Coloration after six months in preservative is much the same but not nearly as distinct and the overall coloration has become dark-grey (Fig. 4). Distribution and Natural history. Hemiphyllodactylus cattien sp. nov. is known only from the type locality in Cat Tien National Park (NP), Dong Nai Province, southern Vietnam (Fig. 1). The holotype was collected at sundown (18:00 h) ca. 1.5 m above the ground on the trunk of a tree, in primary dipterocarp forest along the trail to Bau Sau Lake (Fig. 5). Two specimens were recorded sitting close to one another (ca. 10 cm) on the tree trunk; they tried to escape our presence by ascending the tree and only a single specimen was collected. Despite numerous subsequent attempts to collect additional specimens of the new species and intensive surveys along the Bau Sau Lake trail by P.V. Yushchenko and A.M. Bragin from 2021–2023, no other individuals were recorded. The occurrence of Hemiphyllodactylus cattien sp. nov. in other parts of the Cat Tien NP is expected, including the Cat Loc and Tay Cat Tien sectors covering the lowland and hilly areas of the adjacent parts of Lam Dong and Binh Phuoc provinces of southern Vietnam. Etymology. The name of the new species is a noun in apposition and is therefore invariable; the species name is given in reference to its type locality, the Cat Tien National Park in Dong Nai Province of southern Vietnam. We recommend the names “Cat Tien Slender Gecko” and “Thạch sùng dẹp Cát Tiên” as common names of the new species in English and Vietnamese, respectively. nov cattien sp. dalatensis arakuensis banaensis bonkowskii aurantiacus flaviventris maximum SVL35.145.93937.9514839.2Chin scales77–108–11 10–14 6–75–77–10Postmentals distinctly enlarged (1) or not (0);11 0 0 11/Circumnasal scales5,5 3 /2–4 3 3 5Intersupranasals32–32–33–6 4–11 2–43Supralabials910–119–1210–139–128–109–12Infralabials99–109–128–129–118–118–11Dorsals20 16 13–16 11–17 17–20 24–27 16–18 Ventrals11 8–9 7–9 8–129–12 13–15 8–101st finger lamellae5 3–4 4 or 53,45551st toe lamellae6 3–4 5 or 64,554–55Digits 2–5 finger lamellae44444555/4565 2222 2222 3(4)4(5)4(5)434444444Digits 2–5 toe lamellae55555666/5665 2(3)333 2232(3) 455545544555Continuous femoral/precloacal pores19 (perf)23–2512//1416–2518–2119Precloacal and femoral pore series separate (1) or continuous (0);00 1 1 00/Cloacals on each side1/1 2–3 2–3 1–3111 or 2Subcaudals enlarged, plate–like (1) or not (0)0000000Dark postorbital stripe present (1) or absent (0)1111111Light postocular or trunk spots (1) or absent (0)0 1 0 1 1 1 1 Dark dorsolateral stripe present (1) or not (000 1 0 1 0/Dark dorsal transverse blotches (1) or not (0)00/ 1 1 0/Longitudinal series of white (1) or yellow or red (0) dorsal spots000 1 0 1 0Postsacral mark lacking anteriorly projecting arms (1) or arms present (0)1 0 1 0 0 1 0 Gonads and cecum pigmented (1) or not (0)/01100/ ......continued on the next page Comparisons. Morphologically, Hemiphyllodactylus cattien sp. nov. differs from its close relative H. indosobrinus by having fewer dorsal scales (20 versus 30), supralabials (nine versus 15), infralabials (nine versus 13), and a narrower head (HW/SVL = 0.16 versus 0.18). It differs from its close relative H. flaviventris by having more dorsal scales (20 versus 16—18), 19 perforated femoroprecloacal scales in the female versus nine precloacal pores in male, and more elongate trunk (AG/SVL = 0.52 versus 0.47—0.50) and a narrower (HW/SVL = 0.15 versus 0.17—0.18) and shorter head (HL/SVL = 0.24 versus 0.26—0.27). See Table 2 for potential diagnostic characters separating H. cattien sp. nov. from all other species from Vietnam and South Asian species within clade 5. We are well-aware that additional samples of the new species coupled with statistical analyses of the characters may indicate that some of these putatively diagnostic differences may prove not to be diagnostic but that the same probability exists that other diagnostic differences may be revealed.

Cyrtodactylus arndti sp. nov. urn:lsid:zoobank.org:act: 83F60753-8E51-49D9-97E7-BD870AC772F3 Figs 6–9; Table 3 Diagnosis The new species of Cyrtodactylus is distinguished from remaining congeners of the C. irregularis species group by a combination of the following characters: SVL: 73.4–80.9 mm; dorsal pattern with 6 or 7 irregularly shaped bands; moderately broad nuchal band; original tail with irregular transverse bands; subcaudals transversely enlarged; 5–11 enlarged femoral scales; males with 0–2 pitted femoral pores, those absence in females; males with 6 precloacal pores, females with 6 pitted precloacal pores, pore-bearing scales arranged in a single ˄-shaped series; ventral scales 26–38; dorsal tubercles in 17–20 irregular longitudinal rows; precloacal groove absent; supralabials 8–13; infralabials 8–12; number of subdigital lamellae on fourth finger 15–20 and on fourth toe 17–22. Etymology We name this species in honor of our colleague, Prof. Dr. Hartmut Arndt, Institute of Zoology, University of Cologne, Germany, in recognition of his support for biodiversity research in Vietnam. As common names, we suggest Arndt’s Bent-toed Gecko (English) and Thằn lằn ngón arndt (Vietnamese). Type material (Figs 6–9) Holotype VIETNAM • ♂; Binh Dinh Province, Van Canh District, near Hiep Ha Village; 13°39.858′ N, 108°53.355′ E; 270 m a.s.l.; 13 Aug. 2016; D.T. Do and T.V. Nguyen; Field No. BD.2016.141; IEBR R.4930. Paratypes VIETNAM • 1 ♂; Binh Dinh Province, Quy Nhon District, Quy Nhon City; 13°41.718′ N, 109°10.277′ E; 140 m a.s.l; 8 Aug. 2016; D.T. Do and T.V. Nguyen; Field No. BD.2016.1; IEBR R.5219 • 2 ♂♂, 1 ♀; Binh Dinh Province, Van Canh District, near Dak Dum Village; 13°38.365′ N, 108°57.863′ E; 150 m a.s.l.; 11 Aug. 2016; D.T. Do and T.V. Nguyen; Field No. BD.2016.86, BD2016.87, BD.2016.88; IEBR R.4931 to IEBR.4933 • 1 ♀; Binh Dinh Province, Van Canh District, near Hiep Ha Village; 13°39.858′ N, 108°53.355′ E; 270 m a.s.l.; D.T. Do and T.V. Nguyen; Field No. BD.2016.86; IEBR R.5077 • 2 ♂♂; same collection data as for preceding; Field No. BD.2016.142, BD.2016.143; ZFMK 103910, ZFMK 103911. Description of holotype Adult male; snout-vent length 74.4 mm; tail regenerated, 98.15 mm in length; body slender, elongate (AG/SVL ratio 0.39); head distinct from neck, elongate, depressed (HL/SVL 0.29, HW/HL 0.67, HH/ HL 0.37); loreal region concave; snout long, blunt in dorsal profile (SE/HL 0.37), longer than diameter of orbit (OD/SE 0.61); scales on snout small, round or oval, granular, lager than scales on occiput; orbit large (OD/HL 0.22); pupils vertical; ear opening small, oval (ED/HL 0.06); rostral almost twice as wide as high with an inverse Y-shapted structure, surrounded by first supralabial, naris, nasorostral on each side, and internasal; nostril opening small and oval, surrounding by rostral, nasorostral, 2 supranasals and one postnasal; mental scale triangular, wider than high (ML/MW 0.74); two enlarged, triangular postmentals; supralabials 12/12; infralabials 11/13. Dorsal scales granular, dorsal tubercles round, keeled, conical, in 20 irregular rows at midbody; tubercles on occiput small; each tubercle surrounded by 9 or 10 granular scales; ventral scales smooth, round, midventral scales approximately 3–4 times as large as dorsal scales, slightly imbricate laterally, in 37 longitudinal rows at midbody between ventrolateral folds; precloacal groove absent; enlarged femoral scales 7 or 8 on each thigh, about twice the size of surrounding scales; enlarged precloacal scales 17, arranged in a rhombus; femoral pores absent; precloacal pores 6, arranged in ˄-shaped series. Fore and hindlimbs moderately slender (ForeaL/SVL 0.15 mm, CrusL/SVL 0.16); forelimbs dorsally covered by several slightly enlarged tubercles; dorsal surface of hindlimbs bearing well-developed tubercles; two postcloacal tubercles on each side on the hemipenal swellings; phalanges without webbing; each claw sheathed by two scales, the ventral sheath larger than the upper scale; number of subdigital lamellae on first finger 12/13, on first toe 13/13, on fourth finger 18/18, on fourth toe 22/22. Coloration in preservative Dorsal surface of head, body and limbs light-brown with some dark-brown pattern, without light bordering; occiput marbled with small, irregular dark-brown banded; rostral, mental, first three supralabials and first infralabials greyish brown, remaining infralabials light beige, some with greyish brown speckles or frames; nuchal band discontinuous, consisting of two stripes extending from the orbits to the neck, ending by a dark blotch on each side and a third blotch medially; dorsum with 7 irregular bands, the first two interrupted; dorsolateral region covered with round or elongate dark-brown blotches; tail with 6 dark-brown bands, fade ventrally, some small dark-brown spots arranged in a line along the lateral side of tail, tail tip dark-brown; dorsal surface of limbs with 6 or 7 irregular dark-brown bands; phalanges brown with beige knuckles; dorsal tubercles white or dark-brown depending on position; tubercles on dorsal surface of limbs and tail light-brown; venter greyish brown. Sexual dimorphism and variation The female (IEBR R.4933) differs from the males by the absence of hemipenal swellings. All male specimens have 6 precloacal pores but the females has 6 pitted scales only. Three males (IEBR R.4930, ZFMK 103910, ZFMK 103911) and the female lack femoral pores (IEBR R.5077, IEBR R.4933). For further morphological characters see Table 3. Comparisons The new species can be distinguished from all other member of Cyrtodactylus irregularis group from Vietnam by morphological characteristics (see Table 4). Cyrtodactylus arndti sp. nov. differs from C. badenensis by having the presence of enlarged femoral scales (5–11 vs absent in C. badenensis), the presence of precloacal pores in males (6 vs absent in C. badenensis), the presence of pitted precloacal pores in females (6 vs absent in C. badenensis), and different dorsal pattern (irregular bands vs banded in C. badenensis); differs from C. bidoupimontis by having the presence of pitted precloacal pores in females (6 vs absent in C. bidoupimontis), having moderately broad nuchal band, continuous or discontinuous band (vs well developed, widened posteriorly in C. bidoupimontis), different dorsal pattern (6 or 7 irregular transverse bands, colouration dark-brown on light-brown background vs 4–5 dark irregular transverse dorsal bands, usually with light borders), and the presence of transversely enlarged subcaudals vs absent in C. bidoupimontis; differs from C. bugiamapensis by having fewer precloacal pores in males (6 vs 7–11 in C. bugiamapensis), different dorsal color pattern (irregular, dark-brown transverse bands vs unclear transversal bands formed by irregular roundish to oblong, dark brown spots in C. bugiamapensis), moderately broad nuchal band, continuous or discontinuous bands (vs dark nuchal band, which can be medially divided, narrow, U-shape in C. bugiamapensis), and the presence of transversely enlarged subcaudals vs absent in C. bugiamapensis; differs from C. buchardi by having a lagger size (SVL of 73.4–80.9 vs 60–65 in C. buchardi), the presence of enlarged femoral scales (5–11 vs absent in C. buchardi), more subdigital lamellae on first finger (15–20 vs 14 in C. buchardi), more subdigital lamellae on first toe (17–22 vs 12 in C. buchardi) and the presence of transversely enlarged subcaudals vs absent in C. buchardi); differs from C. caovansungi by having a smaller size (SVL of 73.4–80.9 vs 90.4–94.0 mm), fewer femoral pores (0–2 vs 6 in C. caovansungi), fewer precloacal pores in males (6 vs 9 in C. caovansungi), the presence of pitted precloacal pores in females (6 vs absent in C. caovansungi), and fewer subdigital lamellae under the fourth toe (17–22 vs 23–25 in C. caovansungi); differs from C. cattienensis by having a larger size (73.4–80.9 mm vs 43.5–69.0 mm in C. cattienensis), the presence of pitted precloacal pores in females (6 vs absent in C. cattienensis), and the presence of transversely enlarged subcaudals (vs absent in C. cattienensis); differs from C. chungi by having a larger size (73.4–80.9 mm vs 66.6– 68.5 mm in C. chungi), fewer precloacal pores in males (6 vs 7 in C. chungi), fewer enlarged precloacal scales (17–23 vs 41–45 in C. chungi), and the presence of transversely enlarged subcaudals (vs absent in C. chungi); differs from C. cryptus by having fewer ventral scale rows (26–38 vs 47–50 in C. cryptus), the presence of enlarged femoral scales (5–11 vs absent in C. cryptus), fewer precloacal pores in males (6 vs 9–11 in C. cryptus), the presence of pitted precloacal pores in females (6 vs absent in C. cryptus), and the presence of transversely enlarged subcaudals (vs absent in C. cryptus); differs from C. cucdongensis by having a larger size (SVL 73.4–80.9 mm vs 55.8–65.9 mm in C. cucdongensis), fewer ventral scale rows (26–38 vs 41–44 in C. cucdongensis), more subdigital lamellae under the first toe (11–14 vs 8–11 in C. cucdongensis), and the presence of transversely enlarged subcaudals (vs absent in C. cucdongensis); differs from C. culaochamensis by having fewer ventral scale rows (26–38 vs 45– 50 in C. culaochamensis), the presence of enlarged femoral scales (5–11 vs absent in C. culaochamensis), fewer precloacal pores in males (6 vs 7–8 in C. culaochamensis), and the presence of pitted precloacal pores in females (6 vs absent in C. culaochamensis); differs from C. dati by having fewer ventral scale rows (26–38 vs 42–48 in C. dati), fewer femoral pores (0–2 vs 3–4 on each side in C. dati), different dorsal color pattern (irregular bands vs blotches in C. dati), and the presence of transversely enlarged subcaudals (vs absent in C. dati); differs from C. gialaiensis by having a larger size (73.4–80.9 mm vs 50.1–62.8mm in C. gialaiensis), the presence of enlarged femoral scales (5–11 vs absent in C. gialaiensis), fewer precloacal pores in males (6 vs 9–10 in C. gialaiensis), and more subdigital lamellae under the fourth finger (15–20 vs 14–15 in C. gialaiensis); differs from C. huynhi by having fewer ventral scale rows (26–38 vs 43–46 in C. huynhi), more enlarged femoral scales (5–11 vs 3–5 in C. huynhi), fewer precloacal pores in males (6 vs 7–9 in C. huynhi), and the presence of transversely enlarged subcaudals (vs absent in C. huynhi); differs from C. irregularis by having the presence of precloacal pores in males (1–2 vs absent in C. irregularis), different dorsal color pattern (irregular bands vs blotched in C. irregularis), and the presence of transversely enlarged subcaudals (vs absent in C. irregularis); differs from C. kingsadai by having a smaller size (SVL 73.4–80.9 mm vs 83.0–94.0 mm in C. kingsadai), fewer ventral scale rows (26–38 vs 39–46 in C. kingsadai), and fewer precloacal pores in males (6 vs 7–9 in C. kingsadai); differs from C. orlovi by having a larger size in males (SVL 73.4–80.9 mm vs 61.0– 68.2 mm in C. orlovi), the presence of pitted precloacal pores in females (6 vs absent in C. orlovi), and the presence of transversely enlarged subcaudals(vs absent in C. orlovi); differs from C.phnomchiensis by having fewer ventral scale rows (26–38 vs 45–54 in C. phnomchiensis), more precloacal pores in males (6 vs 4–5 in C. phnomchiensis), fewer AG/SVL ratio (0.34–0.43 vs 0.45–0.48 in C. phnomchiensis), and the presence of transversely enlarged subcaudals (vs absent in C. phnomchiensis); differs from C. phumyensis by having a larger size (SVL 73.4–80.9 mm vs 63.6–66.8 mm in C. phumyensis), and the presence of transversely enlarged subcaudals (vs absent in C. phumyensis); differs from C. phuocbinhensis by having a larger size (SVL 73.4–80.9 mm vs 46.0– 60.4 mm in C. phuocbinhensis), fewer ventral scale rows (26–38 vs 43–47 in C. phuocbinhensis), fewer precloacal pores in males (6 vs 7 in C. phuocbinhensis), the presence of pitted precloacal pores in females (6 vs absent in C. phuocbinhensis); differs from C. pseudoquadrivirgatus by having fewer ventral scale rows (26–38 vs 41–57 in C.pseudoquadrivirgatus), the presence of enlarged femoral scales (5–11 vs absent in C. pseudoquadrivirgatus), different dorsal color pattern (irregular bands vs blotched in C. pseudoquadrivirgatus), and the presence of transversely enlarged subcaudals (vs absent in C. pseudoquadrivirgatus); differs from C. raglai by having a smaller size (SVL 73.4–80.9 mm vs 95–111.7 mm in C. raglai), more precloacal pores in males (6 vs 5 in C. raglai) and the presence of pitted precloacal pores in females (6 vs absent in C. raglai); differs from C. sangi by having a larger size (SVL 73.4–80.9 mm vs 49.9–56.3 mm in C. sangi), more enlarged femoral scales (5–11 vs 4 in C. sangi), fewer precloacal pores in males (6 vs 7 in C. sangi), and more pitted precloacal pores in females (6 vs 4 in C. sangi); differs from C. takouensis by having more enlarged femoral scales (5–11 vs 3–5 in C. takouensis), more precloacal pores in males (6 vs 3–4), the presence of pitted precloacal pores in females (6 vs absent in C. takouensis), and the different dorsal color pattern (irregular bands vs banded in C. takouensis); differs from C. taynguyenensis by having more ventral scale rows (26–38 vs 42–49 in C. taynguyenensis), the presence of enlarged femoral scales (5–11 vs absent in C. taynguyenensis), the different dorsal color pattern (irregular bands vs blotched in C. taynguyenensis), and the presence of transversely enlarged subcaudals (vs absent in C. taynguyenensis); differs from C. yangbayensis by having the presence of pitted precloacal pores in females (6 vs absent in C. yangbayensis), more subdigital lamellae under the fourth toe (17–22 vs 15–17), and fewer subdigital lamellae on first toe (11–14 vs 18–20 in C. yangbayensis); differs from C. ziegleri by having a smaller size (SVL 73.4–80.9 mm vs 84.6–93.0 mm in C. ziegleri), the presence of transversely enlarged subcaudals (vs absent in C. ziegleri), and dark-brown transverse bands of the tail narrower than the light-brown interspaces (vs dark transverse bands wider than the light interspaces in C. ziegleri). Differs from C. chumuensis sp. nov. by having a larger size (SVL 73.4–80.9 mm vs maximum 67.5 mm in C. chumuensis), fewer ventral scale rows (26–38 vs 43–45 in C. chumuensis), and the presence of transversely enlarged subcaudals (vs absent in C. chumuensis). Distribution Cyrtodactylus arndti sp. nov. is currently known only from the Van Canh District, Binh Dinh Province, Vietnam (Fig. 1). Natural history Specimens were found at night between 19:00 and 22:00, on trees or on granite rock, along rocky streams, about 0.6–1.5 m above the ground, at elevations between 150 and 300 m a.s.l. The surrounding habitat was evergreen forest of medium and small hardwoods mixed with shrubs and vines (Fig. 9). The humidity was approximately 40–62% and the air temperature ranged from 28.9 to 33.1 ° C. Other reptiles species found at the sites included Acanthosaura coronata (Günther, 1861) Dixonius vietnamensis Das, 2004, Gekko gecko (Linnaeus, 1758), Gekko sp., Eutropis multifasciata (Kuhl, 1820), Boiga jaspidea (Duméril, Bibron & Duméril, 1854), Psammodynastes pulverulentus (Boie, 1827), and Trimeresurus sp., Published as part of Ngo, Hanh Thi, Hormann, Helene, Pham, Cuong The, Phung, Trung My, Ostrowski, Sabrina, Nguyen, Truong Quang & Ziegler, Thomas, 2023, The discovery of two new species in the Cyrtodactylus irregularis group highlights that hidden diversity remains in the largest clade of the mega-diverse genus Cyrtodactylus, pp. 70-100 in European Journal of Taxonomy 875 (1) on pages 84-94, DOI: 10.5852/ejt.2023.875.2141, http://zenodo.org/record/8064396, {"references":["Ziegler T., Rosler H., Herrmann H. - W. & Vu T. N. 2002. Cyrtodactylus phongnhakebangensis sp. n., ein neuer Bogenfingergecko aus dem annamitischen Karstwaldmassiv, Vietnam. Herpetofauna 24 (141): 11 - 25.","Nguyen S. N., Orlov N. L. & Darevsky I. S. 2006. Descriptions of two new species of the genus Cyrtodactylus Gray, 1827 (Squamata: Sauria: Gekkonidae) from southern Vietnam. Russian Journal of Herpetology 13 (3): 215 - 226. https: // doi. org / 10.30906 / 1026 - 2296 - 2006 - 13 - 3 - 215 - 226","Nguyen S. N., Yang J-X., Le T-N. T., Nguyen L. T., Orlov N., Hoang C. V., Nguyen T. Q., Jin J-Q., Rao D-Q., Hoang T. N., Che J., Murphy R. W. & Zhang Y-P. 2014. DNA barcoding of Vietnamese bent-toed geckos (Squamata: Gekkonidae: Cyrtodactylus) and the description of a new species. Zootaxa 3784 (1): 48 - 66. https: // doi. org / 10.11646 / zootaxa. 3784.1.2","Nguyen S. N., Zhou W-W., Le T-N. T., Tran A-D. T., Jin J-Q., Vo B. D., Nguyen L. T., Nguyen T. T., Nguyen T. Q., Hoang D. D., Orlov N. L., Che J., Murphy R. W. & Zhang Y-P. 2017 a. Cytonuclear discordance, cryptic diversity, complex histories, and conservation needs in Vietnamese bent-toed geckos of the Cyrtodactylus irregularis species complex. Russian Journal of Herpetology 24 (2): 133 - 154.","Nguyen S. N., Nguyen L. T., Nguyen V. D. H. & Murphy R. W. 2017 b. Genetic diversity, phylogeny, and altitudinal distribution of geckos genus Cyrtodactylus on Hon Ba Mountain, Khanh Hoa Province, with note on the subcaudals of C. yangbayensis. Journal of Biotechnology 15 (3 A): 55 - 62.","Heidrich A., Rosler H., Vu N. T., Bohme W. & Ziegler T. 2007. Another new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha-Ke Bang National Park, central Truong Son, Vietnam. Zootaxa 1445: 35 - 48. https: // doi. org / 10.11646 / zootaxa. 1445.1.3","Orlov N. L., Nguyen T. Q., Nazarov A. R., Ananjeva B. N. & Nguyen S. N. 2007. A new species of the genus Cyrtodactylus Gray, 1827 and redescription of Cyrtodactylus paradoxus (Darevsky et Szczerbak, 1997) [Squamata: Sauria: Gekkonidae] from South Vietnam. Russian Journal of Herpetology 14: 145 - 152.","Nazarov R. A., Orlov N. L., Nguyen N. S. & Ho T. C. 2008. Taxonomy of naked-toe geckos Cyrtodactylus irregularis complex of South Vietnam and description of a new species from Chu Yang Sin Natural Park (Krong Bong District, Dac Lac Province), Vietnam. Russian Journal of Herpetology 15: 141 - 156.","Nazarov R., Poyarkov N. A., Orlov N. L., Phung T. M., Nguyen T. T., Hoang D. M. & Ziegler T. 2012. Two new cryptic species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from southern Vietnam. Zootaxa 3302 (1): 1 - 24. https: // doi. org / 10.11646 / zootaxa. 3302.1.1","Ngo T. V. & Bauer A. M. 2008. Descriptions of two new species of Cyrtodactylus Gray 1827 (Squamata: Gekkonidae) endemic to southern Vietnam. Zootaxa 1715: 27 - 42. https: // doi. org / 10.11646 / zootaxa. 1715.1.2","Ngo T. V., Grismer L. L. & Grismer J. L. 2010. A new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) in Phu Quoc National Park, Kien Giang Biosphere Reserve, Southwestern Vietnam. Zootaxa 2604 (1): 37 - 51. https: // doi. org / 10.11646 / zootaxa. 2604.1.3","Rosler H., Vu T. N., Nguyen T. Q., Ngo V. T. & Ziegler T. 2008. A new Cyrtodactylus (Squamata: Gekkonidae) from central Vietnam. Hamadryad 32: 125 - 140 -","Ngo T. V. & Chan K. O. 2010. A new species of Cyrtodactylus Gray, 1826 (Squamata: Gekkonidae) from Khanh Hoa province, southern Vietnam. Zootaxa 2504 (1): 47 - 60. https: // doi. org / 10.11646 / zootaxa. 2504.1.4","Ngo T. V. & Chan O. K. 2011. A new krastic cave-dwelling Cyrtodactylus Gray (Squamata: Gekkonidae) from Northern Vietnam. Zootaxa 3125 (1): 51 - 63. https: // doi. org / 10.11646 / zootaxa. 1909.1.4","Ngo T. V. & Grismer L. L. 2012. A new species of Cyrtodactylus Gray (Squamata: Gekkonidae) from Tho Chu Island, southwestern Vietnam. Zootaxa 3228 (1): 48 - 60. https: // doi. org / 10.11646 / zootaxa. 3228.1.2","Ziegler T., Nazarov R., Orlov N., Nguyen T. Q., Vu T. N., Dang K. N., Dinh T. H. & Schmitz, A. 2010. A third new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha-Ke Bang National Park, Truong Son Range, Vietnam. Zootaxa 2413 (1): 20 - 36. https: // doi. org / 10.11646 / zootaxa. 2413.1.2","Ziegler T., Phung T. M., Le M. D. & Nguyen T. Q. 2013. A new Cyrtodactylus (Squamata: Gekkonidae) from Phu Yen Province, southern Vietnam. Zootaxa 3686 (4): 432 - 446. https: // doi. org / 10.11646 / zootaxa. 3686.4.2","Luu V. Q., Nguyen T. Q., Do Q. H. & Ziegler T. 2011. A new Cyrtodactylus (Squamata: Gekkonidae) from Huong Son limestone forest, Hanoi, northern Vietnam. Zootaxa 3129: 39 - 50. https: // doi. org / 10.11646 / zootaxa. 3129.1.3","Luu V. Q., Dung T. V., Nguyen T. Q., Le M. D. & Ziegler T. 2017. A new species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from Gia Lai Province, Central Highlands of Vietnam. Zootaxa 4362 (3): 385 - 404. https: // doi. org / 10.11646 / zootaxa. 4362.3.4","Phung T. M., van Schingen M., Ziegler T. & Nguyen T. Q. 2014. A third new Cyrtodactylus (Squamata: Gekkonidae) from Ba Den Mountain, Tay Ninh Province, southern Vietnam. Zootaxa 3764 (3): 347 - 363. https: // doi. org / 10.11646 / zootaxa. 3764.3.5","Le D. T., Nguyen T. Q., Le M. D. & Ziegler T. 2016. A new species of Cyrtodactylus (Squamata: Gekkonidae) from Ninh Binh Province, Vietnam. Zootaxa 4162 (2): 268 - 282. https: // doi. org / 10.11646 / zootaxa. 4162.2.4","Ostrowski S., Le M. D., Ngo H. T., Pham C. T., Nguyen T. Q. & Ziegler T. 2020. A new species of Cyrtodactylus (Squamata: Gekkonidae) from southern Vietnam. Zootaxa 4789 (1): 171 - 203. https: // doi. org / 10.11646 / zootaxa. 4789.1.5","Ostrowski S., Le M. D., Ngo H. T., Pham C. T., Phung T. M., Nguyen T. Q. & Ziegler T. 2021. A new Cyrtodactylus (Squamata: Gekkonidae) from Binh Thuan Province, southern Vietnam. European Journal of Taxonomy 731 (1): 47 - 70. https: // doi. org / 10.5852. ejt. 2021.731.1023","Do Q. H., Phung T. M., Ngo H. T., Le M. D., Ziegler T., Pham C. T. & Nguyen T. 2021. A new species of Cyrtodactylus (Squamata: Gekkonidae) from Ninh Thuan Province, southern Vietnam. Zootaxa 4999 (1): 58 - 76. https: // doi. org / 10.11646 / zootaxa. 4999.1.4"]}

Cyrtodactylus chumuensis sp. nov. urn:lsid:zoobank.org:act: F6A019B5-B1ED-4976-8C1D-803CC0D91DD9 Figs 3–5; Table 2 Diagnosis The new species can be distinguished from remaining congeners of the irregularis species group by a combination of the following characters: maximum SVL 67.5 mm; dorsal pattern with 6 irregularly shaped and short longitudinal stripes on the neck; nuchal band thin, interrupted, reaching the posterior margin of the orbits; the absence of transversely enlarged median subcaudal scales; 4 or 5 enlarged femoral scales on each thigh, 17–19 distinctly enlarged precloacal scales; males with 0 or 1 femoral pore on each thigh, 6 or 7 precloacal pores in a continuous series, ˄-shaped; ventral scales 43–45; dorsal tubercles in 20 irregular longitudinal rows; precloacal groove absent; internasal scales 2; supralabials 8–14; infralabials 9–11; number of subdigital lamellae on fourth finger 16–19 and on fourth toe 19–21. Etymology The new species is named after its type locality, Chu Mu Mountain in Dak Lak Province. We propose the following common names: Chu Mu Bent-toed Gecko (English), Thạch sùng ngón chư mư (Vietnamese). Type material (Figs 3–5) Holotype VIETNAM • ♂; Dak Lak Province, M'Drak District, Ea M’Doal Commune, Chu Mu Mountain; 12°41.330′ N, 108°55.450′ E; 500 m a.s.l.; 20 Jun. 2014; T.M. Phung leg.; Field No. PMT01; IEBR R.4928. Paratypes VIETNAM • 1 ♂ (subadult); Dak Lak Province, M'Drak District, Ea M’Doal Commune, Chu Mu Mountain; 12°41.321′ N, 108°55.382′ E; 400 m a.s.l.; 20 Jun. 2014; T.M. Phung leg.; Field No. PMT02; IEBR R.4929. Description of holotype Adult male; snout-vent length SVL 67.5 mm; tail regenerated 51.4 mm in length (regenerated portion 43.7 mm); body slender, elongate (AG/SVL 0.4); head distinct from neck, elongate (HL/SVL 0.26), relatively wide (HW/HL 0.71) and depressed (HH/HL 0.36); loreal region concave; snout long, blunt in dorsal profile (SE/HL 0.44), longer than diameter of orbit (OD/SE 0.56); scales on snout small, round or oval, granular, larger than scales on occiput; orbit large (OD/HL 0.24), pupils vertical; ear opening small, oval (ED/HL 0.06); rostral wider than high, indented medially the top, in contact with first supralabial, naris and nasorostral on each side, two internasal scales; nostril opening small, oval, surrounded by rostral, nasorostral, two supranasals and one or two postnasals; mental scale triangular, wider than high (ML/MW 0.74); two enlarged, triangular postmentals; supralabials 8/9; infralabials 9/9. Dorsal scales granular, dorsal tubercles round, keeled, conical, in 20 irregular rows at midbody; tubercles on occiput small; tubercles surrounded by 10–12 granular scales; ventral scales smooth, round, midventral scales three times as large as dorsal granular scales, in 45 longitudinal rows at midbody between ventrolateral folds; precloacal groove absent; enlarged femoral scales 4 or 5 on each thigh, the third bearing a femoral pore (Fig. 4C); enlarged precloacal scales 17, arranged in a rhombus; precloacal pores 6, arranged in a ˄-shaped series. Fore and hindlimbs moderately slender (ForeaL/SVL 0.14 mm, CrusL/SVL 0.17); forelimbs dorsally covered by several slightly enlarged tubercles; dorsal hindlimbs with well-developed tubercles; two postcloacal tubercles on each side on the hemipenal swellings; phalanges without webbing; each claw sheathed by two scales, the ventral sheath larger than the upper; number of subdigital lamellae on first finger 11/11, on first toe 10/11, on fourth finger 17/17, on fourth toe 19/19. Coloration in preservative Dorsal surface of head, body and limbs light-brown with some dark-brown bands, pattern without light bordering; occiput marbled with small, irregular dark-brown blotches; rostral, mental and infralabials creamy white, supralabials dark-beige with short greyish brown vertical stripes; neck bands dark-brown, extending in two thin stripes along lateral sides of the snout to the orbits, broader on the neck, interrupted on the left side; two dark-brown longitudinal stripes, disconnected from the neck band extending to shoulders, one dark-brown blotch next to each stripe; dorsal pattern consisting of 6 irregular bands, each formed by two triangularly shaped blotches, shifted medially along the body axis; dorsolateral region covered with small, irregular dark-brown blotches arranged in a longitudinal row from neck to groin; a blurry dark-brown transverse band on dorsal surface of original part of the tail, regenerated part greyish beige and speckled with very small light-greyish brown blotches; dorsal surface of limbs with 3 or 4 dark-brown, blurry bands; phalanges brown with creamy white knuckles; tubercles white or dark-brown depending on position on pattern or background; venter creamy white; ventral tail greyish beige without bands. For coloration of the paratype in life that closely resembles the holotype in life see Fig. 3. Variation The paratype is a subadult and therefore differs greatly in size. Its original tail showed some dark-brown irregular bands, although broken at the base. The number of precloacal pores is 7 and it lacks femoral pores. For more morphological characters see Table 2. Comparisons The new species can be distinguished from all other member of Cyrtodactylus irregularis group from Vietnam by morphological characteristics (see Table 2). Cyrtodactylus chumuensis sp. nov. differs from C. badenensis by having more ventral scale rows (43– 45 vs 25–29 in C. badenensis), the presence of enlarged femoral scales (4–5 vs absent in C. badenensis), the presence of precloacal pores in males (6–7 vs absent in C. badenensis), and the absence of transversely enlarged subcaudals (vs present in C. badenensis); differs from C. bidoupimontis by having a smaller size (SVL 67.5 mm vs 74.0– 86.3 mm in C. bidoupimontis), fewer enlarged femoral scales (4 or 5 vs 8–10 in C. bidoupimontis), a different dorsal color pattern (irregularly banded with longitudinal stripes on the neck vs transversal bands with light borders in C. bidoupimontis), and a thin discontinuous nuchal band (vs well developed, widened posteriorly in C. bidoupimontis); differs from C. bugiamapensis by having fewer enlarged femoral scales (4 or 5 vs 6–10 in C. bugiamapensis) and the different dorsal color pattern (irregularly banded with longitudinal stripes on the neck vs unclear transversal bands formed by irregular round to oblong, dark-brown spots in C. bugiamapensis); differs from C. buchardi by having more ventral scale rows (30 vs 43–45 in C. buchardi), the presence of enlarged femoral scales (4–5 vs absent in C. buchardi), more subdigital lamellae under the fourth finger (16–19 vs 14 in C. buchardi), more subdigital lamellae under the fourth toe (17–21 vs 12 in C. buchardi); differs from C. cattienensis by having more ventral scale rows (43–45 vs 28–42 in C. cattienensis), more subdigital lamellae under the fourth finger (16–19 vs 12–16 in C. cattienensis), and different dorsal color pattern (irregularly banded with longitudinal stripes on the neck vs irregular dark-brown banded, first band on the shoulder x-shaped C. cattienensis); differs from C. caovansungi by having a smaller size (SVL 67.5 mm vs 90.4–94 mm in C. caovansungi), fewer enlarged femoral scales (4 or 5 vs 8 in C. caovansungi), fewer femoral pores on each thigh in males (0–1 vs 6 in C. caovansungi), fewer precloacal pores in males (6 or 7 vs 9 in C. caovansungi), fewer lamellae under the fourth finger (16–19 vs 22 in C. caovansungi), fewer lamellae under the fourth toe (17–21 vs 23–25 in C. caovansungi), and the absence of transversely enlarged subcaudal plates (vs present in C. caovansungi); differs from C. chungi by having more ventral scale rows (43–45 vs 30 or 31 in C. chungi), more dorsal tubercle rows (20 vs 18 in C. chungi), different dorsal color pattern (irregularly banded with longitudinal stripes on the neck vs irregular transversal bands with a closed nuchal band), and a thin, discontinuous nuchal band (vs continuous nuchal band in C. chungi); differs from C. cryptus by having fewer ventral scale rows (43–45 vs 47–50 in C. cryptus), the presence of enlarged femoral scales (vs absent in C. cryptus), fewer precloacal pores in males (6 or 7 vs 9–11 in C. cryptus), a thin, discontinuous nuchal band (vs well developed, widened posteriorly in C. cryptus), and different dorsal color pattern (irregularly banded with short, longitudinal stripes on the neck vs irregular transverse bands in C. cryptus); differs from C. cucdongensis by having more dorsal tubercle rows (20 vs 16–19 in C. cucdongensis), fewer enlarged femoral scales (4 or 5 vs 5–9 in C. cucdongensis), more enlarged precloacal scales (20–21 vs 6–13), and a different dorsal colour pattern (irregularly banded with short, longitudinal stripes on the neck vs irregular dark brown transverse bands); differs from C. culaochamensis by having a smaller size (SVL 67.5 mm vs 69.8–79.8 mm in C. culaochamensis), the presence of enlarged femoral scales (vs absent in C. culaochamensis), fewer lamellae under the first finger (11 vs 13 or 14 in C. culaochamensis), and fewer lamellae under the first toe (10 or 11 vs 13–15 in C. culaochamensis); differs from C. dati by having fewer femoral pores in males (0–2 vs 3 or 4 each side in C. dati), fewer lamellae under the first toe (10 or 11 vs 12 or 13 in C. dati), the presence of blotches on head (vs absent in C. dati), and different dorsal color pattern (irregularly banded with longitudinal stripes on the neck vs irregular dark blotches); differs from C. gialaiensis by the presence of enlarged femoral scales (vs absent in C. gialaiensis), fewer precloacal pores in males (6 or 7 vs 9 or 10 in C. gialaiensis), and more subdigital lamellae under the fourth finger (16–19 vs 14 or 15 in C. gialaiensis) as well as under the fourth toe (17–21 vs 15–17 in C. gialaiensis); differs from C. huynhi by having more dorsal tubercle rows in males (20 vs 16–18 in C. huynhi), fewer lamellae under first finger (11 vs 12–15 in C. huynhi), fewer lamellae under first toe (10 or 11 vs 13–17 in C. huynhi), and a thin discontinuous nuchal band (vs well developed, widened posteriorly in C. huynhi); differs from C. irregularis by having a smaller size (SVL 67.5 mm vs 72.0–86.0 mm in C. irregularis), more ventral scale rows (43–45 vs 37–42 in C. irregularis), fewer enlarged femoral scales (4 or 5 vs 7 or 8 in C. irregularis), and different dorsal color pattern (irregularly banded with short longitudinal stripes on the neck vs blotched in C. irregularis); differs from C. kingsadai by having a smaller size (SVL 67.5 mm vs 83.0–94.0 in C. kingsadai), fewer enlarged femoral scales (4 or 5 vs 9–12 in C. kingsadai), the absence of transversely enlarged subcaudal plates (vs present in C. kingsadai), and more internasals (2 vs 1 in C. kingsadai); differs from C. orlovi by having more ventral scale rows (43–45 vs 36–39 in C. orlovi); a thin, discontinuous nuchal band (vs continuous nuchal band in C. orlovi), and different banded pattern ranges (6 vs 3–5 in C. orlovi); differs from C. phnomchiensis by having a smaller size (SVL 67.5 vs 76.1–80.7 mm in C. phnomchiensis), more precloacal pores in males (6 or 7 vs 5 in C. phnomchiensis), and different dorsal color pattern (irregularly banded vs banded in C. phnomchiensis); differs from C. phuocbinhensis by having a larger size (SVL 67.5 mm vs 46.0– 60.4 mm in C. phuocbinhensis), different dorsal color pattern (irregularly banded vs stripes or blotches in C. phuocbinhensis), and dark-brown transverse banded of the tail than light-brown interspaces (vs dark transverse banded wider than light interspaces in C. phuocbinhensis); differs from C. phumyensis by having more ventral scale rows (43–45 vs 33–41 in C. phumyensis), fewer enlarged femoral scales (4 or 5 vs 5–7 in C. phumyensis), more dorsal tubercle row (20 vs 18 or 19 in C. phumyensis), fewer enlarged precloacal scales (17–19 vs 21–41 in C. phumyensis), and different dorsal color pattern (irregularly banded with short longitudinal stripes on the neck vs anteriorly irregularly spotted and posteriorly banded in C. phumyensis); differs C. pseudoquadrivirgatus by the presence of enlarged femoral scales on each thigh (vs absent in C. pseudoquadrivirgatus), the presence of precloacal pores in males (0–2 vs absent in C. pseudoquadrivirgatus), and more enlarged precloacal scales (17–19 vs 1–12 in C. pseudoquadrivirgatus); differs C. raglai by having a smaller size (SVL 67.5 mm vs 87.5–111.7 mm in C. raglai), more ventral scale rows (43–45 vs 36–39 in C. raglai), fewer enlarged femoral scales (4 or 5 vs 9 or 10 in C. raglai), fewer precloacal pores in males (0–2 vs 5 in C. raglai), and the absence of transversely enlarged subcaudal plates (vs present in C. raglai); differs from C. sangi by having a larger size (SVL 67.5 mm vs 49.9–56.3 mm in C. sangi) and more ventral scale rows (43–45 vs 37 in C. sangi); differs from C. takouensis by having a smaller size (SVL 67.5 mm vs 74.7–81.1 mm in C. takouensis), more ventral scale rows (43–45 vs 39–40 in C. takouensis), more precloacal pores in males (6 or 7 vs 3 or 4 in C. takouensis), the absence of transversely enlarged subcaudal plates (vs present in C. takouensis), a thin discontinuous nuchal band (vs well developed, widened posteriorly in C. takouensis), and different dorsal color pattern (irregularly banded vs banded in C. takouensis); differs from C. taynguyenensis by the presence of enlarged femoral scales on each thigh (vs absent in C. taynguyenensis); irregularly banded of the tail (vs banded in C. taynguyenensis), and different dorsal color pattern (irregularly banded vs blotched in C. taynguyensis); differs from C. yangbayensis by having a smaller size (SVL 67.5 vs 78.5–92.3 mm in C. yangbayensis), more subdigital lamellae under the fourth toe (17–21 vs 15–17 in C. yangbayensis), fewer subdigital lamellae under the first toe (10–11 vs 18–20 in C. yangbayensis), and the absence of transversely enlarged subcaudal plates (vs present in C. yangbayensis); differs from C. ziegleri by having a smaller size (SVL 67.5 vs 84.6–93.0 mm in C. ziegleri), more ventral scale rows (43–45 vs 33–39 in C. ziegleri), and fewer enlarged femoral scales (4 or 5 vs 8–10 C. ziegleri). Distribution Cyrtodactylus chumuensis sp. nov. is currently known only from the Chu Mu Mountain, M'Drak District, Dak Lak Province, Vietnam (Fig. 1). Natural history Specimens were found at night between 19:00 and 22:00, on granite rock, along a rocky stream, approximately 0.5–1.0 m above the ground, at elevations between 400 and 500 m a.s.l. The surrounding habitat was evergreen forest of medium and small hardwoods mixed with shrubs and vines (Fig. 5). The humidity was approximately 50–71% and the air temperature ranged from 27.5 to 32.1°C. Other reptile species found at the sites included Gekko gecko (Linnaeus, 1758), Hemidactylus platyurus (Schneider, 1792), Ahaetulla prasina (Boie, 1827), Lycodon sp., and Oligodon sp., Published as part of Ngo, Hanh Thi, Hormann, Helene, Pham, Cuong The, Phung, Trung My, Ostrowski, Sabrina, Nguyen, Truong Quang & Ziegler, Thomas, 2023, The discovery of two new species in the Cyrtodactylus irregularis group highlights that hidden diversity remains in the largest clade of the mega-diverse genus Cyrtodactylus, pp. 70-100 in European Journal of Taxonomy 875 (1) on pages 78-84, DOI: 10.5852/ejt.2023.875.2141, http://zenodo.org/record/8064396

Ngo, Hanh Thi, Hormann, Helene, Pham, Cuong The, Phung, Trung My, Ostrowski, Sabrina, Nguyen, Truong Quang, Ziegler, Thomas (2023): The discovery of two new species in the Cyrtodactylus irregularis group highlights that hidden diversity remains in the largest clade of the mega-diverse genus Cyrtodactylus. European Journal of Taxonomy 875 (1): 70-100, DOI: https://doi.org/10.5852/ejt.2023.875.2141, URL: http://dx.doi.org/10.5852/ejt.2023.875.2141

Khandekar, Akshay, Gaikwad, Sunil M., Pal, Saunak, Thackeray, Tejas, Agarwal, Ishan (2023): An expanded description of Cnemaspis gracilis (Beddome 1870) (Squamata: Gekkonidae) based on recent material. Zootaxa 5301 (5): 540-560, DOI: 10.11646/zootaxa.5301.5.2, URL: http://dx.doi.org/10.11646/zootaxa.5301.5.2

Cnemaspis gracilis (Beddome, 1870) Gymnodactylus gracilis Beddome, 1870 (Figures 2–6; Table 2–5) Lectotype. BMNH 74.4.29.393, adult male, (designated by Manamendra-Arachchi et al. 2007), from “Palghat Hills” (Palakkad district, Kerala, India), collected by Colonel R. H. Beddome. Paralectotypes. BMNH 74.4.29.394, adult female, BMNH 74.4.29.395 adult female, BMNH 74.4.29.396 subadult female, BMNH 74.4.29.397 adult male, BMNH 74.4.29.398 subadult male, (designated by Manamendra-Arachchi et al. 2007) from “Palghat hills”, Kerala, India, collected by Colonel R. H. Beddome. Referred material. BNHS 3128 (CESL 606), adult male and BNHS 3129 (CESL 607), adult female, from Chennathanair RF (10.86823° N, 76.62161° E; ca. 330, m asl.), Palakkad district, Kerala, India; collected by Saunak Pal on 3 rd July 2012; NRC-AA-1248 (AK 133), NRC-AA-1249 (AK 134), NRC-AA-1250 (AK 135), NRC-AA-1252 (AK 138), NRC-AA-1253, (AK 139), NRC-AA-1254 (AK 141), adult males, NRC-AA-1255 (AK 144), subadult male, NRC-AA-1251 (AK 137), adult female, from Valparai town (10.3321°N, 76.9595°E; ca. 1100 m asl.), Coimbatore district, Tamil Nadu, India, collected by Akshay Khandekar, Ishan Agarwal, and Swapnil Pawar on 12 th November 2017; CES G 385, adult female (tissue + photograph only), from Chittur College campus (10.68730°N, 76.72260° E; ca. ~ 100 m asl.), Chittur, Palakkad district, Kerala, India; collected by Ishan Agarwal on 28 th May 2012. NRC-AA-1277 (AK 862), adult male, NRC-AA-1278 (AK 863), adult female, NRC-AA-1279 (AK 864), juvenile, from Timber market, Sambhaji Nagar, Kolhapur city (16.68558°N, 74.22174° E; ca. ~ 600 m asl.), Kolhapur district, Maharashtra, India; collected by Akshay Khandekar and Vivek Kuber on 6 th March 2023. Diagnosis. A small-sized Cnemaspis, snout to vent length up to 33 mm (n =19). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with a fairly regularly arranged rows of enlarged, strongly keeled, conical tubercles; last one or two rows of enlarged, weakly keeled, spine-like tubercles on flank; 10–12 rows of dorsal tubercles at mid-body, 10–14 tubercles in paravertebral rows; ventral scales smooth, subcircular, subimbricate, subequal from chest to vent, 24–29 (rarely 30, n =1/13) scales across belly at mid-body, 100–121 longitudinal scales from mental to cloaca; subdigital scansors smooth, mostly unpaired, unnotched; 10–12 lamellae under digit I of manus and pes; 15–18 lamellae under digit IV of manus and 18–24 lamellae under digit IV of pes; males with 3–5 femoral pores on each thigh separated by 6–13 poreless scales from series of two (rarely 4 n =1/8) precloacal pores, precloacal pores separated medially by 2–4 poreless scales; tail with enlarged, strongly keeled, pointed, and spine-like tubercles forming whorls; median row of subcaudals smooth, roughly oval, and distinctly enlarged. Dorsum brown to red or orange, mottled with numerous small light grey spots and fine black spots, light grey vertebral blotches forming a chain from occiput to tail base; a single central black dorsal ocellus on neck and smaller one on occiput separated by a light blotch, two or three indistinct pairs of streaks on throat; original tail in males with about 8–10 alternating dark and light grey bands, regenerated tail yellow to orange. Comparison with members of C. gracilis clade. Cnemaspis gracilis can be easily distinguished from all members of the clade by a combination of the following differing or non-overlapping characters: small-sized Cnemaspis with maximum SVL 32.9 mm (versus medium-sized Cnemaspis, SVL up to 41 mm in C. salimalii and C. thackerayi); 10–14 tubercles in paravertebral rows (versus 14–18 tubercles in paravertebral rows in C. agayagangai, 15–17 tubercles in paravertebral rows in C. fantastica, only a few irregularly arranged tubercles in paravertebral region in C. mundanthuraiensis, 16–18 tubercles in paravertebral rows in C. salimalii); 10–12 rows of dorsal tubercles at mid-body (versus eight or nine rows of dorsal tubercles at mid-body in C. jackieii, 6–8 rows of dorsal tubercles at mid-body in C. mundanthuraiensis); spine-like tubercles present on flanks (versus spine-like tubercles absent on flanks in C. agarwali, C. jackieii, C. shevaroyensis, and C. thackerayi); 24–29 (rarely 30) ventral scales across belly at mid-body (versus 30–32 in C. rudhira, 30–33 ventral scales across belly at mid-body in C. salimalii, 21–24 in C. shevaroyensis, and 22–25 in C. thackerayi); males with femoral pores, single (rarely 2) precloacal pore on each side which are separated medially by 2–4 poreless scales (versus males with femoral pores, two precloacal pores on each side which are separated medially by a single poreless scale in C. agarwali, males with femoral pores, continuous series of precloacal pores in C. pachaimalaiensis, males with femoral pores, two (rarely 3) precloacal pore on each side which are separated medially by single (rarely 2) poreless scales in C. rudhira), femoral pores separated by 6–13 poreless scales from precloacal pores on either side in males (versus femoral pores separated by 1–6 poreless scales from precloacal pores on either side in males in C. thackerayi); single central dorsal ocellus each on occiput and between forelimb insertions (versus a single dorsal ocellus present on occiput and neck, two pairs on either side just anterior and sometimes posterior to forelimb insertions in C. agarwali; a single central dorsal ocellus each on occiput and neck, ocellus on neck flanked anteriorly on each side by a slightly larger ocellus in C. agayagangai; a single central ocellus on neck, flanked posteriorly by a pair of much larger squarish blotches and anteriorly by a pair of subequal squarish blotches, indistinct spot on occiput in C. fantastica; a large central black dorsal ocellus on neck flanked anteriorly and posteriorly on each side by elongate dark ocelli, smaller ocellus on occiput flanked on each side by a smaller ocellus in C. pachaimalaiensis; a single dorsal ocellus present on occiput and neck, two pairs on either side just anterior and posterior to forelimb insertions in C. shevaroyensis). Description of topotype (based on male BNHS 3128 (CESL 606)). Adult male in good state of preservation except extreme tail tip missing, a 7.2 mm longitudinal cut on lower right pectoral region for tissue collection (Fig. 2A–E). SVL 31.0 mm, head short (HL/SVL 0.26), wide (HW/HL 0.66), not strongly depressed (HD/HL 0.43), distinct from neck. Loreal region marginally inflated, canthus rostralis not distinct. Snout half of head length (ES/HL 0.49), marginally more than 2.5 times eye diameter (ES/ED 2.56); scales on snout and canthus rostralis subcircular, subequal, and mostly smooth; much larger than those on forehead and interorbital region; scales on forehead slightly smaller, weakly keeled, elongated, and weakly conical; scales on interorbital region even smaller, granular and weakly keeled; scales on occipital and temporal region heterogeneous, slightly enlarged, weakly keeled, conical tubercles intermixed with smaller, weakly keeled and weakly conical granules (Fig. 3A). Eye small (ED/HL 0.19) with round pupil; supraciliaries short, larger anteriorly; seven interorbital scale rows across narrowest point of frontal bone; 30–32 scale rows between left and right supraciliaries at mid-orbit (Fig. 3A, C). Ear-opening deep, oval, small (EL/HL 0.07); eye to ear distance greater than diameter of eye (EE/ED 1.56; Fig. 3C). Rostral more than two times wider (1.43 mm) than high (0.6 mm), incompletely divided dorsally by a strongly developed rostral groove and internasal scale for more than half of its height; a single enlarged supranasal on each side, slightly larger than postnasals, separated from each other by a slightly smaller, elongated internasal scale; two postnasals, upper postnasal slightly larger than lower; rostral in contact with supralabial I, nostril, internasal, supranasal, and lower postnasal on either side; nostrils oval, surrounded by two postnasals, supranasal, and rostral on either side; two rows of scales separate orbit from supralabials (Fig. 3C). Mental enlarged, subtriangular, slightly wider (1.72 mm) than high (1.20 mm); two pairs of postmentals, inner pair roughly rectangular, much shorter (0.74 mm) than mental, in strong contact with each other below mental; inner pair bordered by mental, infralabial I, outer postmental, and a single enlarged chin shield on left and two on right side; outer postmentals roughly rectangular, smaller (0.50 mm) than inner pair, bordered by inner postmentals, infralabial I and II, and four enlarged chin shields on either side and; two enlarged gular scales between left and right outer postmentals; all chin scales bordering postmentals flat, subcircular, smooth, and smaller than outermost postmentals; scales on rest of throat, even smaller, flattened, subequal, and smooth (Fig. 3B). Infralabials bordered below by a row or two of slightly enlarged, much elongated scales, decreasing in size posteriorly. Nine supralabials up to angle of jaw, and six at midorbital position on either side; supralabial I largest, rest gradually decreasing in size posteriorly; eight infralabials up to angle of jaw, and five at midorbital position on either side; infralabial I largest, rest gradually decreasing in size posteriorly (Fig. 3C). Body relatively slender (BW/AGL 0.46), trunk less than half of SVL (AGL/SVL 0.36) without ventrolateral folds; spine-like scales on flank present (Fig. 4A–C). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with a fairly regularly arranged row of enlarged, strongly keeled, conical tubercles; tubercles in approximately 11 longitudinal rows at mid-body including spine-like scales at lower flank; 10 tubercles in paravertebral row from above forelimb insertion to the hind limb insertion (Fig. 4A, C). Ventral scales much larger than granular scales on dorsum smooth, subcircular, subimbricate, subequal from chest to vent; mid-body scale rows across belly 26; 113 scales from mental to anterior border of cloaca (Fig. 4B). Scales on base of neck similar to those on belly, marginally smaller; gular region with still smaller, subequal, smooth, flattened scales, those bordering postmentals enlarged, smooth, subcircular, and flattened (Fig. 4B). Five femoral pores on either thigh, separated by 11 poreless scales on left and 13 on right side from two precloacal pores, precloacal pores separated medially by a three poreless scales (Fig. 3D). Scales on palms and soles granular, smooth, subcircular, subimbricate and flattened at base; scales on dorsal aspects of limbs heterogeneous in shape and size; mixture of small granular, weakly keeled, imbricate scales which are twice the size of granules on the body dorsum, largest on anterolateral aspect of the hands and feet; posterolateral aspect of limbs with small weakly keeled to smooth granular scales; scales on lower arm and shank small, subimbricate, and keeled; ventral aspect of forelimbs with small, smooth, subimbricate scales, larger on lower arm than upper arm; ventral aspect of hindlimb with enlarged, smooth, flattened, subimbricate scales, scales on shank slightly larger than body ventrals (Fig. 2A, B). Forelimbs and hindlimbs moderately long, slender (LAL/SVL 0.14; CL/SVL 0.18); digits long, with strong, recurved claw, distinctly inflected, distal portions laterally compressed conspicuously. Digits with unpaired lamellae except basal one or two paired on some digits, separated into a basal and narrower distal series by single enlarged lamella at inflection; basal lamellae series: (1-3-3-5-5 right manus, 1-5-6-6-6 right pes), (2-3-3-5-4 left manus, Fig. 3E; 2-5-7-6-6 left pes, Fig. 3F); distal lamellae series: (10-11-13- 12-11 right manus, 10-11-13-14-13 right pes), (10-11-12-13-12 left manus, Fig. 3E; 9-11-14-14-13 left pes, Fig. 3F). Relative length of digits (measurements in mm in parentheses): IV (2.8)> III (2.7)> V (2.5) II> (2.4)> I (1.9) (left manus); IV (4.0)> V (3.6)> III (3.4)> II (3.2)> I (1.8) (left pes). Tail original, almost entire, subcylindrical, slender, marginally longer than snout-vent length (TL/SVL 1.10; Fig. 2C–E). Dorsal scales on tail base weakly keeled, granular, similar in size and shape to granular scales on mid-body dorsum, gradually becoming larger, flattened, imbricate posteriorly, intermixed with enlarged, strongly keeled, distinctly pointed, conical tubercles; enlarged tubercles on the tail forming whorls; six tubercles each on first five whorls, four in 6–11th whorls, only paravertebral tubercles on 12–15 th whorl, rest of the tail tip lacking enlarged tubercles (Fig. 2C, E). Scales on ventral aspect of tail much larger than those on dorsal aspect, subimbricate, smooth; median series distinctly larger than rest, roughly oval; scales on tail base marginally larger than those on mid-body ventrals, smooth, imbricate; a single enlarged, smooth and conical postcloacal spur on each side (Fig. 2D). Colouration in life (Fig. 5A). Dorsum of head, body, limbs and tail base reddish. Head mottled with grey and black, yellow and dark bands on labials, two dark postorbital streaks. A single central ocellus on neck, much smaller spot on occiput. Dorsum mottled with numerous grey spots and fine black spots with a vertebral chain light elongate blotches. Dorsum of limbs mottled with yellow bands, digits with alternating dark and light bands. Tail with eight black and eight light grey bands. Venter off-white with black speckles, three indistinct pairs of black streaks on throat. Variation and additional information. Mensural, meristic and additional character state data for the topotypes and specimens from additional localities are given in Tables 3–5 respectively. There are seven adult males, single subadult male, and three adult females and a juvenile ranging in size from 19.6–32.4 mm (Fig. 6B). All other specimens resemble described male topotype except as follows: Upper postmentals separated from each other below mental by enlarged median chin shield in NRC-AA-1254; upper postmentals bordered by mental, infralabial I, outer postmental in all topotypes, and additionally by single enlarged chin scale on each side and median chin scale in NRC-AA-1248, NRC-AA- 1253, NRC-AA-1254, NRC-AA-1278 and NRC-AA-1279, one chin scale on left and three on right side in NRC-AA-1251, single median scale on either side in BNHS 3129. Outer postmental bordered by inner postmental, infralabials I & II in all topotypes, and additionally, five chin scales on left and four on right side in NRC-AA- 1250, NRC-AA-1251, NRC-AA-1277, three chin scales on left and four on right side in NRC-AA-1253, NRC-AA-1254; four chin scales on left and three on right side in NRC-AA-1278, NRC-AA-1279, outer postmental separated from each other medially by three enlarged chin scales in NRC-AA-1248, NRC-AA-1251, NRC-AA-1253, NRC-AA-1254, NRC-AA-1278, NRC-AA-1279, and by single enlarged median chin scale in BNHS 3129. Snout region completely damaged in NRC-AA-1255. five specimens — NRC-AA-1249, NRC-AA-1253, and NRC-AA-1255, NRC-AA-1277, NRC-AA-1278 with original and complete tails, slightly longer than body (TL/SVL 1.18, 1.14, 1.31, 127, and 120 respectively); NRC-AA-1254 with complete but only tail tip regenerated, almost equal to the body (TL/SVL 1.04); rest of the topotypes with original but incomplete tails. Dorsal colouration varies from tan or light grey to red and orange; females, subadult and juvenile overall duller than adult males, original tail distinctly banded in males and faintly in females; regenerated portion of the tail orangish in NRC-AA-1254; only a single female topotype BNHS 3129 with multiple ocelli on body and neck (Figs. 5A–F, 6). ......Continued on the next page TABLE 5. (Continued) Distribution and Natural history. Cnemaspis gracilis is now known with surety from around its type locality (Palghat Hills) in Palakkad district, Kerala and Valparai town in Coimbatore district, Tamil Nadu (Fig. 1), besides the presumably introduced population from the Timber Market, Sambhaji Nagar, Kolhapur, Kolhapur District, Maharashtra. At Palakkad, CESG385 was spotted resting on a low shrub (Cnemaspis gracilis was observed to be common and recorded along the ghat road from Monkey falls, Pollachi to Valparai town covering the elevation gradient of 400–1100 m asl. Individuals were seen active during the day time on building walls, on rocks, road side rock cuttings and under cement culvert. In Valparai town, individuals were also seen resting on mossy walls at night. At Kolhapur, Cnemaspis gracilis was only observed in the vicinity of Timber Market. The species was seen active in high abundance (>25 individuals/hr) on trees, building walls, and timber piles during the daytime., Published as part of Khandekar, Akshay, Gaikwad, Sunil M., Pal, Saunak, Thackeray, Tejas & Agarwal, Ishan, 2023, An expanded description of Cnemaspis gracilis (Beddome 1870) (Squamata: Gekkonidae) based on recent material, pp. 540-560 in Zootaxa 5301 (5) on pages 544-556, DOI: 10.11646/zootaxa.5301.5.2, http://zenodo.org/record/8036203, {"references":["Beddome, R. H. (1870) Descriptions of some new lizards from the Madras Presidency. Madras Monthly Journal of Medical Science, 1, 30 - 35.","Manamendra-Arachchi, K., Batuwita, S. & Pethiagoda, R. (2007) A taxonomic revision of the Sri Lankan day-geckos (Reptilia: Gekkonidae: Cnemaspis), with description of new species from Sri Lanka and southern India. Zeylanica, 7, 9 - 122."]}

Grass Snake Natrix natrix (Linneaus 1758) Distribution (Figure 9). Included records from Artportalen (N=200): all reports from Gästrikland, Hälsingland, and coastal Medelpad (where widespread). Reports from other areas have been included only if documentation was available. This species and black morph Vipera berus are widely confused, even among naturalists. Occurs throughout the Southern Boreal. Widespread and fairly common in Gästrikland and coastal Hälsingland, local and scarce in interior Hälsingland and coastal Medelpad. There is a disjunct well-documented record at 480 m altitude in central Härjedalen, in the Northern Boreal region (Figure 9). This is by far the highest known record, whereas the 2 nd to 4 th highest in North Sweden were all made at 180 m or lower. For the former extraordinary record there is no obvious suspicion of anthropogenic spread. However, well-documented records north of the present range (filled black circles in the map) can all be suspected to have anthropogenic origin, and none represents permanent presence or a reproducing population. Although the species is an excellent swimmer there are just two records on offshore islands off the Baltic coast of North Sweden (Limön and Eggegrund, both in Gästrikland). For the period 1900–1950, Gislén & Kauri (1959) listed many records in the Middle and Northern Boreal, far north and west of the known present range. Several were well documented, but it is not known whether those occurrences were relicts from a previously wider natural distribution, or the result of anthropogenic activities such as long-distance transport of hay and manure (Elmberg 1995). As late as the late 1960’s there were confirmed records in several places around Umeå (Västerbotten, Middle Boreal; Stefan Ericsson personal communication), an area where the species has not been documented since. During the last 50 years, though, there have not been any indications of changes in distribution. Habitat and movements. Found in open places providing shelter, such as tall field layer vegetation, cairns, stone walls, ditches, and heaps of plant debris. Typical habitats are shores of ponds and lakes, but also edges of fields and pastures. It is not known if habitat use varies over the annual activity period, but adult females must find suitable oviposition sites in early summer. This likely forces them to move considerable distances, probably through less typical habitats. All known oviposition sites in North Sweden were in composts, manure heaps or livestock fodder stacks (Löwenborg 2009; Mattias Hagman & Simon Kärvemo, personal communication). Although movements have not been studied in North Sweden, Natrix natrix are frequently seen crossing roads before and after oviposition (adult females) and after hatching in late summer (juveniles). This mobility sadly leads to many being killed by cars. Nothing is known about hibernation habits in North Sweden, but communal hibernation at suitable south-facing sites is likely, as farther south in Sweden. Abundance estimates and trends. There are not any data about abundance in North Sweden, nor any indications of changes in abundance over the last 50 years., Published as part of Elmberg, Johan, 2023, Amphibians and reptiles in North Sweden: distribution, habitat affinities, and abundance (Classes: Amphibia and Reptilia), pp. 301-335 in Zootaxa 5301 (3) on pages 319-322, DOI: 10.11646/zootaxa.5301.3.1, http://zenodo.org/record/8030434, {"references":["Gislen T. & Kauri H. (1959) Zoogeography of the Swedish amphibians and reptiles, with notes on their growth and ecology. Acta Vertebratica, 1 (3), 197 - 397.","Elmberg, J. (1995) Groddjurens och kraldjurens utbredning i Norrland. Natur i Norr, 15 (2), 57 - 82. [in Swedish]","Lowenborg, K. (2009) Reproductive ecology of the grass snake (Natrix natrix): testing the maternal manipulation hypothesis. M. Sc. Thesis, Stockholm University, Stockholm."]}

Adder Vipera berus (Linneaus 1758) Distribution (Figure 10). Included records from Artportalen (N=1250): all reports have been included. Any confusion between melanistic Vipera berus and Natrix natrix would not affect the distribution pattern. Common and widespread in the Southern and Middle Boreal. Widespread and locally common in the Northern Boreal, where often found in areas of varied topography with south-facing slopes that offer early snowmelt and dependable hibernacula (Andersson 2003; Figure 20). Scattered observations have been made in the Subalpine and Low-Alpine zones (Curry-Lindahl 1975; Frislid & Semb-Johansson 1981; Elmberg 1995). The highest reported occurrences show a slightly decreasing altitude from south to north: 940 m in Härjedalen (Flatruet), 900 m in Jämtland (Oviksfjällen), 720 m in Lycksele lappmark (Kraipe), but 820 m in Lule lappmark (Aktse). Most of these concern south-facing sites in the upper Subalpine zone, but also above treeline in the Low-Alpine zone. In the upper reaches of the river Lilla Luleälv (Lule lappmark) there are many records in the Aktse and Tarradalen areas (e.g., Cederberg 1974), suggesting widespread occurrence in the Subalpine zone there. Widespread and locally abundant on many offshore islands along much of the Baltic coast of North Sweden (Figure 10), as is the case across the sea in Finland (Terhivuo 1981). An illustrative example is the Holmön archipelago in Västerbotten, where this species is abundant on islands> 10 km from the mainland. This implies high dispersal capacity over brackish water. There are no indications of large-scale changes in distribution over the last 50 years. Habitat and movements. Males typically emerge from hibernation two weeks before females. Mating occurs near the hibernaculum, after which snakes disperse to summer habitats. The latter are largely the same as those of Zootoca vivipara: forest edges, forest clearings, stony slopes, rock outcrops, shrubbery, and shores of lakes, rivers, and the sea (Figure 12). In the vast interior of North Sweden summer habitats also include margins of bogs and mires (Figure 14). Areas providing a combination of basking sites and hiding places are favored. Anthropogenic habitats are widely used, too, for example clearings under powerlines, clear-cuts, edges of fields and meadows, and stone walls (Figures 13, 19). Very rarely found in closed forest, but sometimes in mature pine forest with natural gaps and clearings. It also occurs regularly in dense Juniperus communis thickets and in tall grass, where prey animals are likely to be found. There are not any telemetry data available from North Sweden, but judging from field observations Vipera berus appears to be more mobile in summer than are Zootoca vivipara and Anguis fragilis. However, Andersson (2003) reported from the Northern Boreal that pregnant females may remain close to the hibernaculum throughout summer. The distance from summer habitats to hibernation sites can be up to 2 km (Andersson 2003), a journey during which Vipera berus can be found in other habitats. Hibernation usually occurs in open south-facing locations with abundant crevices and underground access. Typical sites are talus slopes, stony moraine ridges (cf.Andersson 2003) and screes in the forest landscape (Figure 20), but also cairns if big and deep enough. Good hibernation sites are always well-drained and can attract individuals from a large surrounding area. There are reports from North Sweden that a single hibernaculum can host hundreds of individuals (Frislid & Semb-Johansson 1981). Abundance estimates and trends. There are not any published abundance data, but estimates based on extensive field work in the Umeå area (Västerbotten) 1975–1994 run in the neighborhood of 50– 100 adults /km 2 (Elmberg, unpublished). A mark-recapture study on the far offshore island Stora Fjäderägg (Västerbotten; 63 o 48’N, 21 o 00’E; area ca 170 hectares) suggests an average abundance of ca. 0.6 adults/hectare (60/km 2) (Stefan Andersson, unpublished). However, studies by Stefan Andersson on the mainland suggest that abundance estimates for larger areas are particularly difficult to make in this species; adults occur within a certain radius of hibernacula, but between the latter there may be large areas of good summer habitat more or less devoid of adders. There are not any true monitoring data for any period of the last 50 years, but recurrent visits to known sites of occurrence in southern Västerbotten indicate a steady and significant drop in abundance from the late 1970’s to the present day (Elmberg, unpublished)., Published as part of Elmberg, Johan, 2023, Amphibians and reptiles in North Sweden: distribution, habitat affinities, and abundance (Classes: Amphibia and Reptilia), pp. 301-335 in Zootaxa 5301 (3) on pages 322-324, DOI: 10.11646/zootaxa.5301.3.1, http://zenodo.org/record/8030434, {"references":["Curry-Lindahl, K. (1975) Groddjur och kraldjur i farg. Alla Europas arter. AWE / Gebers, Stockholm, 198 pp. [in Swedish]","Frislid, R. & Semb-Johansson, A. (Eds.) (1981) Norges Dyr 3. Fisker, amfibier, krypdyr. J. W. Cappelens Forlag, Oslo, pp 362 - 408. [in Norwegian]","Elmberg, J. (1995) Groddjurens och kraldjurens utbredning i Norrland. Natur i Norr, 15 (2), 57 - 82. [in Swedish]","Cederberg, B. (1974) Inventering av vegetationen pa Ritokkammen Sareks Nationalpark. Meddelanden, Vaxtbiologiska Institutionen, Uppsala universitet, Uppsala, 1974, 5. [in Swedish]","Terhivuo, J. (1981) Provisional atlas and population status of the Finnish amphibian and reptile species with special reference to their ranges in northern Europe. Annales Zoologici Fennici, 18, 139 - 164."]}

Elmberg, Johan (2023): Amphibians and reptiles in North Sweden: distribution, habitat affinities, and abundance (Classes: Amphibia and Reptilia). Zootaxa 5301 (3): 301-335, DOI: 10.11646/zootaxa.5301.3.1, URL: http://dx.doi.org/10.11646/zootaxa.5301.3.1

Hemidactylus paucifasciatus sp. nov. (Figures 2–6; Table 3) urn:lsid:zoobank.org:act: 967860CD-F420-432A-ABAF-4D8B4A3E6952 Hemidactylus sp., Dutta et al. (2009); Debata, 2017 Holotype. ZSI-R-28357, adult male, from Gadachandi Temple, Anandapur, (21.2273° N, 86.2549° E; ca. 115 m asl.), Keonjhar District, Odisha State, India, collected by Pratyush P. Mohapatra & Rakesh Kumar Mohalik on 18 April 2019. Paratypes. NCBS-BH668, adult male, from Bamanghati, Bangiriposi (22.1714° N, 86.4925° E; ca. 341 m asl.), Mayurbhanj District, Odisha State, India, collected by Pratyush P. Mohapatra & Sushil Kumar Dutta on 29 November 2014; NCBS-BH669, adult female, same locality as holotype except collected by Rakesh Kumar Mohalik on 24 February 2019; ZSI-CZRC-7117, sub-adult female, same locality and collection data as holotype. Referred specimen. ZSI-R-28526, sub-adult male, from Chakratirtha, Kathakata, Anandapur (21.2569° N, 86.2500° E; ca. 197 m asl), Keonjhar District, Odisha State, India, collected by Rakesh Kumar Mohalik on 22 June 2022. Etymology. The species epithet is derived from a combination of the Latin adjectives pauci (English: few) and fasciatus (English: banded), referring to the lesser number of dorsal bands in this species. Suggested Common Name. Few-banded termite hill gecko Diagnosis. A large-sized Hemidactylus, SVL up to 118 mm (n =4). Dorsal pholidosis heterogeneous, composed of subcircular granular scales intermixed with enlarged, fairly regularly arranged, strongly keeled, conical tubercles in 16–18 longitudinal rows, extending from occiput to tail base, that are heterogeneous in shape and size; enlarged tubercles on the two most medial parasagittal rows smaller, 21–24 tubercles in paravertebral rows; those on dorsolateral aspect of flank largest, gradually decreasing in size downwards, last two rows on flank marginally larger or equal to medial parasagittal rows, weakly keeled. Ventrolateral folds indistinct; about 29–31 scale rows across belly. Digits with enlarged scansors, lamellae in straight transverse series, all divided except the apical and 1–3 basal lamellae, nine or ten lamellae beneath first digit and 11–13 beneath fourth digit of manus and pes. Males with 12 or 13 femoral pores on each side separated by three or four poreless scales. Tail with 6–9 much enlarged, strongly keeled, conical tubercles forming whorls; median row of subcaudal plates large, covering almost entire portion of the tail; single postcloacal spur of unequal size on each side that are smooth and much smaller than dorsal tubercles at mid-body. Dorsal coloration rusty-brown with three (rarely four) transversely arranged light bands, edged with black or dark brown, between the occiput and sacrum; head lighter than body in life, with a narrow lighted collar edged by dark brown that is continuous with the post-ocular marking. Comparison with members of the prashadi group. The new species was phylogenetically recovered as part of the prashadi group and can be morphologically assigned to the group based on its large size and prominent dorsal tuberculation.We therefore restrict comparisons to other members of the prashadi group. Hemidactylus paucifasciatus sp. nov. can be easily distinguished from other members of the prashadi group by having heterogeneous dorsal scales, composed of subcircular granular scales intermixed with enlarged, fairly regularly arranged, strongly keeled, conical tubercles, 16–18 DTR (versus homogenous dorsal pholidosis of imbricate scales, lacking enlarged tubercles in H. scabriceps (Annandale); dorsal pholidosis heterogenous, composed of granular scales intermixed with enlarged, rounded, weakly-keeled tubercles, 18–20 DTR in H. aaronbaueri; dorsal pholidosis heterogenous, composed of granular scales intermixed with enlarged, fairly regularly arranged, strongly keeled, conical tubercles — 18–20 DTR in H. acanthopholis Mirza & Sanap; 18–20 DTR in H. kangerensis; 12–14 DTR in H. kimbulae; 20–24 DTR in H. maculatus; 22–24 DTR in H. paaragowli Srikanthan, Swamy, Mohan & Pal; 14–16 DTR in H. prashadi; 19 or 20 DTR in H. triedrus); males with 12 or 13 FP and three or four SBFP (versus 19 FP and six SBFP in H. aaronbaueri; 19–21 FP and 13 or 14 SBFP in H. acanthopholis; 15–19 FP and 2–4 SBFP in H. depressus Gray; 24–30 FP and 2–4 SBFP in H. easai Das, Pal, Siddarth, Palot, Deepak & Narayanan; 23–28 FP and 1–3 SBFP in H. graniticolus Agarwal, Giri & Bauer; 14 or 15 FP and 11 SBPP in H. hegdei Pal & Mirza; 22–24 FP and 3–6 SBFP in H. hunae; 18–21 FP and four SBFP in H. kangerensis; 21–24 FP and 5–7 SBFP in H. kimbulae; 21–25 FP and two or three SBFP in H. kolliensis Agarwal, Bauer, Giri & Khandekar; 16–19 FP and 5–9 SBFP in H. maculatus; 10–12 FP and 16–18 SBFP in H. paaragowli; 17–20 FP and 1–3 SBFP in H. pieresii Kelaart; 17–20 FP and three SBFP in H. prashadi; 16–18 FP and 13–15 SBFP in H. sirumalaiensis Khandekar, Thackeray, Pawar & Agarwal; 17 or 18 FP and five SBFP in H. siva; 20–23 FP and 3–6 SBFP in H. sushilduttai; 17 or 18 FP and seven SBFP in H. tamhiniensis; 7–9 FP and 1–3 SBFP in H. triedrus; 17–22 FP and ten or 11 SBFP in H. vanam Chaitanya, Lajmi & Giri; seven or eight FP and three SBFP in H. whitakeri; FP absent, only precloacal pores present in H. scabriceps); 29–31 MVSR (versus 41–43 MVSR in H. aaronbaueri; 35–40 MVSR in H. acanthopholis; 28–36 MVSR in H. depressus; 41 or 42 MVSR in H. easai; 40–46 MVSR in H. graniticolus; 32–34 MVSR in H. hegdei and H. kolliensis; 36–39 MVSR in H. kimbulae; 28–34 MVSR in H. maculatus; 33–39 MVSR in H. paaragowli; 32–39 MVSR in H. pieresii; 36–38 MVSR in H. sirumalaiensis; 54–59 MVSR in H. tamhiniensis; and 34–40 MVSR in H. vanam). Additionally, Hemidactylus paucifasciatus sp. nov. can be easily distinguished from all three described members of the H. triedrus clade by its large SVL, up to 118 mm (versus, medium SVL H. sahgali, H. triedrus, and H. whitakeri). Description of holotype. The holotype (ZSI-R-28357) is in good condition except for an 8.7 mm long incision in the sternal region for tissue collection, a minor fold of skin (not the ventrolateral fold) from axilla to groin on left side, and the tail is bent towards the right (Fig. 2). Adult male, SVL 117.7 mm; head short (HL/SVL 0.23), slightly elongate (HW/HL 0.88), not strongly depressed (HD/HL 0.55), distinct from neck. Loreal region slightly inflated, canthus rostralis indistinct (Fig. 2C). Snout short (ES/HL 0.45); much longer than eye diameter (ED/ES 0.52); scales on snout, canthus rostralis, forehead and inter-orbital region heterogenous, mostly granular and conical; scales on the snout and canthus rostralis much larger than those on occiput, forehead and inter-orbital regions, roughly oval (Fig. 2A). Eye small (ED/HL 0.23); pupil vertical with crenulated margins; supraciliaries small, pointed, gradually increasing in size towards front of the orbit (Fig. 2C). Ear opening oval (greatest diameter 2.5 mm); eye to ear distance slightly greater than diameter of eye (EE/ED 0.59). Rostral wider than deep (RW/RL 1.80), partially divided dorsally by a weakly developed rostral groove; single enlarged internasal between two slightly larger supranasals, three postnasals on each side, marginally smaller than supranasal; rostral in contact with nasal, supralabial I, internasal and supranasals on either side; nostrils small (1.1 mm), oval; nasal scale surrounded by rostral, supranasal, three postnasals, and supralabial I on either side; two rows of scales separate orbit from supralabials on each side. Mental subtriangular; two pairs of well-developed postmentals, the inner pair slightly shorter (3.4 mm) than the mental (4.1 mm), and in strong contact with each other (1.2 mm) below mental, outer pair slightly shorter (2.3 mm) than the inner pair and separated from each other by inner pair (Fig. 2B). Inner postmentals bordered by mental, infralabial I, outer postmental and five small gular scales on either side; outer postmentals bordered by infralabial I & II, inner postmental, and six gular scales increasing in size laterally, two outer-most of which are much enlarged and continues as two rows of enlarged scales below the infralabials, decreasing in size posteriorly (Fig. 2B). Labials large, decreasing in size posteriorly; supralabials (on both sides) to mid-orbital position eight, and 11 up to angle of jaw; seven infralabials (on both sides) to mid-orbital position and nine up to angle of jaw; (Fig. 2C). Body relatively stout (BW/SVL 0.23), not elongate (AGL/SVL 0.44), ventrolateral folds indistinct (Fig. 4). Dorsal pholidosis heterogeneous, composed of subcircular granular scales intermixed with enlarged, fairly regularly arranged, strongly keeled, conical tubercles in 16 longitudinal rows, extending from occiput to tail, that are heterogeneous in shape and size; enlarged tubercles on the two most medial parasagittal rows smaller, those on dorsolateral aspect of flank largest, gradually decreasing in size downwards, last two rows on flank marginally larger or equal to medial parasagittal rows, weakly keeled (Fig. 4); each enlarged tubercle surrounded by a rosette of 14–16 small granules with 1–3 granules between two longitudinally adjacent enlarged tubercles (6–8 between parasagittal rows at mid-body); enlarged tubercles on nape and shoulder marginally smaller than parasagittal rows, those on occiput still smaller, weakly keeled, conical; tubercles on temporal region slightly pronounced than those on occiput, strongly conical (Figs. 2A & 4A). Ventral scales much larger than dorsal granular scales, smooth, imbricate, subequal, slightly larger on precloacal and femoral region than on chest and abdominal region (Fig. 2B); 30 mid-body scale rows across belly; 88 scales from posterior margin of ear-opening to the lowest enlarged scale row above cloaca; gular region with small, flattened, granular scales, becoming slightly larger and imbricate on lateral aspect (Fig. 2B). Scales on palm and sole smooth, imbricate, subcircular; enlarged tubercles on dorsal aspect of upper arm much smaller than those on mid-body dorsum, weakly keeled, slightly conical, subimbricate; dorsal aspect of forearm with smaller, granular scales, intermixed with a few enlarged, keeled, conical tubercles, those on anterior aspect smooth, flat, imbricate; scales on dorsal part of thigh and shank granular, intermixed with enlarged, keeled, conical tubercles, which are larger on thigh compared to shank; anterior aspect of thigh with flatter scales, posterior aspect with granular scales (Fig. 2). Precloacal-femoral pores in slightly enlarged row of scales, separated medially by a diastema of three poreless scales, 13 (right) and 12 (left) (Fig. 3D). Fore and hind limbs relatively short, stout; forearm short (FL/SVL 0.12); tibia short (CL/SVL 0.14); digits moderately short, strongly clawed; all digits of manus and digits I–IV of pes indistinctly webbed; terminal phalanx of all digits curved, arising angularly from distal portion of expanded lamellar pad, half or more than half the length of the associated toepad; scansors beneath each toe in a straight transverse series, divided except for a single distal and three basal scansors on digit I and one in other digits: 11-12-12-13-12 (left manus), 11-12-12-13-13 (right manus; Fig. 3E), 10-13-14-13-13 (left pes), 10-13-14-13-13 (right pes; Fig. 3F). Relative length of digits (measurements in mm in parentheses): I (5.1) Tail depressed, ~ 2/3 rd of its length regenerated, flat beneath, verticillate, with well-defined median furrow; tail slightly shorter than snout-vent length (TL/SVL ratio 0.80) (Fig. 2). Dorsal scales at tail base granular, similar in size and shape to those on mid-body dorsum, gradually becoming larger, flatter, pointed, weakly keeled, subimbricate posteriorly and smooth laterally, intermixed with series of 6–9 much enlarged, strongly keeled, conical tubercles; nine enlarged tubercles in first segment, eight in second, six in third and fourth segments. Regenerated portion of tail covered above with much larger, flattened, smooth, subimbricate scales, decreasing in size posteriorly (Fig. 2A). Ventral scales at tail base smooth, imbricate, slightly larger than ventrals at mid-body; rest (including regenerated portion) of the tail with large plate like subcaudal scales (median row) covering almost entire portion of the tail, first three subcaudal scales divided, rest entire, roughly rectangular; median row bordered laterally by one or two rows of large, smooth, imbricate scales (Fig. 2B). A single, smooth postcloacal spur on either side, much smaller than dorsal tubercles at mid-body (Fig. 2B). Colouration in life. (Fig. 5A) Dorsum rusty brown, head and limbs lighter than body; a thick, dark-brown postocular stripe speckled with orange behind the eye that extends till the lateral sides of occiput and joins a thinner dark brown collar; postocular streak flanked by narrow white bands and dark brown collar is anteriorly edged with a white band. Iris dark brown with golden mottling towards the upper and posterior sides. Labials cream-white, speckled with dark-brown spots below orbit, loreal and prefrontal region speckled with light orange; rostrum dark brown. Dorsum with a series of broad (covering 3–5 tubercle rows), off-white transverse bands bordered with fine dark-brown edges; one across the shoulder with dark shades at the centre, a thinner wavy band at mid-body, an incomplete band originating from the right flank is directed upwards on the left side approaching the previous band, one over the anterior sacral region, and one on the original portion of the tail. The lateral sides of head, axilla, flanks, groin and base of the tail whitish, which is continuous with the venter. The tubercles of the body are darker at the tip. Limbs and phalanges light gray-brown. The regenerated portion of the tail is the same rusty-brown as the body without any bands. Colour in preservative. (Fig. 2) Overall colour faded in preservative to light beige; orbital and frontal region on head even lighter. The dark-brown margins of the transverse bands have faded to brown and the band on the tail base has only retained the anterior brown margins. The darker markings on the enlarged tubercles on the body are faded and appear indistinct after preservation. Crown of head similar in colour with that of dorsum, except slightly darker pigmentation on snout. Variation and additional information from paratype series. Mensural and meristic data for the type series is given in Table 3. There are three specimens (an adult male, female and a sub-adult female) ranging in SVL from 67.6 mm to 108.3 mm. All paratypes resemble the holotype except as follows: supranasals separated by two much smaller internasals in ZSI-CZRC-7117 and NCBS-BH669, supranasals are in strong contact with each other and a much smaller internasal scale above rostral in NCBS-BH668; six gular scales bordering inner postmental on each side in NCBS-BH668, five gular scales on left and six on right bordering inner postmental in ZSI-CZRC-7117 and NCBS-BH669; inner postmental is in contact with infralabial I & II on left and infralabial I on right side in ZSI-CZRC-7117. Seven gular scales bordering outer postmental on each side in ZSI-CZRC-7117, six gular scales on left and five on right side bordering outer postmental in NCBS-BH669; outer postmental divided on left side and in contact with infralabial II on each side in ZSI-CZRC-7117. Paratype NCBS-BH668 with entire but partly regenerated tail, marginally shorter than body (TL / SVL 0.89); ZSI-CZRC-7117 with incomplete tail and NCBS-BH669 is without tail. All paratypes closely agree with the holotype in colouration except the subadult specimen ZSI-CZRC-7117 which is much darker with prominent bands on body and tail. A faint mid-dorsal line may be visible. Variations are also marked in the mid-body banding in the holotype and paratypes; in contrast to the holotype which has a thinner off-white band followed by an incomplete band at the mid body, in ZSI-CZRC-7117 and NCBS-BH668 the mid body band is single, whereas in NCBS-BH669 the band is laterally dilated forming an incomplete “X” shape; the colouration of the regenerated tail in life is light brown, without any enlarged tubercles (Figs. 5B, C & 6). Distribution. (Fig. 1A) The type locality of Hemidactylus paucifasciatus sp. nov. is adjacent to the Hadagarh Wildlife Sanctuary (WLS), and based on our present understanding, the species occurs in three Protected Areas, namely Hadagarh WLS, Kuldiha WLS and Similipal Tiger Reserve, across the Keonjhar, Balasore and Mayurbhanj districts of Odisha. As per the Wildlife Management Plan of Hadagarh WLS (Anonymous, 2021), the landscape with miscellaneous vegetation falls within the Boula-Nuasahi hilly region in Garhjat Hills. This region exhibits extensive occurrence of quartzite and quartz schists on both sides of the Salandi River. The granite outcrops are commonly seen to the North of Sajanapal in the valleys and low-lying hills, which are the preferred microhabitat of the species. The area is also characterized by chromite deposits in association with ultrabasic rocks (peridolite and serpentinties), which are subjected to rapid habitat destruction due to mining activities. Natural history. (Fig. 7) Hemidactylus paucifasciatus sp. nov. was observed on rocks (rupicolous) and on trees (arboreal) and juveniles were more often found on the ground than adults. This species was observed on Mangifera indica L and Terminalia alata Heyne ex Roth trees in several instances inside Hadagarh Wildlife sanctuary. On 16 April 2020 two juveniles were observed near Pitanau beat house of Hadagarh WLS. A common krait scavenging on a road-killed Hemidactylus sp. reported from Kuldiha WLS (Debata 2017) corresponds to a sub-adult H. paucifasciatus sp. nov. This species also enters human habitation in the forest fringe villages and is seen in forest rest houses that are infrequently used by visitors. They slough loose skin easily when handled, especially on the head, a typical defensive behaviour related to regional integumentary loss observed in numerous geckos that has been reported in two members of the acanthopholis group within the H. prashadi clade (Bauer et al. 1989, 2006; Bauer & Russell 1992; Chaitanya et al. 2018; Khandekar et al. 2020). Prey comprises mostly of insects; individuals were seen feeding on moths, crickets, cockroaches, termites and beetles. The other sympa

Viviparous Lizard Zootoca vivipara (Jacquin 1787) Distribution (Figure 7). Included records from Artportalen (N=775): all reports have been included, as there are not any confusion species. Widespread and common in the Southern, Middle, and Northern Boreal. Widespread but scarce in the Subalpine zone. Locally occurring above treeline in the Low-Alpine zone in favorable microclimates. As expected, the highest known occurrences in the Scandic Mountain range are gradually lower towards the north: 1000–1050 m altitude in Härjedalen (Sånfjället and Flatruet), 740 m in Pite lappmark (west of Vuoggatjålme), and 690 m in Lule lappmark (Vastenjaure). There is just one record from a truly far offshore island (Stora Fjäderägg, Västerbotten; Figure 7; Elmberg 1995). Although common in seashore habitats on the mainland along the entire Baltic coast of North Sweden, there are surprisingly few records even from nearshore islands. An exception may be the archipelago in southern Norrbotten, where the species occurs on some outer islands (e.g., Stor-Räbben and Vargön, green offshore area in Figure 7; Stefan Andersson, personal communication). For North Sweden as a whole, this indicates a limited dispersal capacity over brackish water. There are no indications of changes in distribution over the last 50 years. Habitat and movements. Found in almost any habitat offering a combination of basking sites and protective low vegetation. Favored natural habitats are forest edges and clearings, stony slopes, rock outcrops, sandy areas, and shores of lakes, rivers and the sea (Figures 12, 14). It often occurs among Juniperus communis, Calluna vulgaris, Empetrum nigrum and other plants typical of dry sun-exposed conditions. Closed forest, tall grass, and wet habitats are avoided. Anthropogenic habitats are widely used, for example clearings under powerlines, clear-cuts, edges of fields and meadows, stone walls, cairns, and roadsides (Figures 13, 19). There have not been any dedicated studies of this species in North Sweden. As far as known, it spends the entire annual activity period in the habitats mentioned above. Daily and annual movements are not known but appear very limited. Subterranean hibernation sites are found in or close to the summer habitat, usually in south-facing situations. Abundance estimates and trends. There are not any published abundance data, but estimates based on extensive field work in the Umeå area (Västerbotten) 1975–1994 run in the neighborhood of> 500 adults /km 2 in representative landscapes near the coast (Elmberg, unpublished). There are no indications of changes in abundance over the last 50 years., Published as part of Elmberg, Johan, 2023, Amphibians and reptiles in North Sweden: distribution, habitat affinities, and abundance (Classes: Amphibia and Reptilia), pp. 301-335 in Zootaxa 5301 (3) on page 316, DOI: 10.11646/zootaxa.5301.3.1, http://zenodo.org/record/8030434, {"references":["Elmberg, J. (1995) Groddjurens och kraldjurens utbredning i Norrland. Natur i Norr, 15 (2), 57 - 82. [in Swedish]"]}

Slow Worm Anguis fragilis Linneaus 1758 Distribution (Figure 8). Included records from Artportalen (N=360): as there are not any confusion species all reports have been included. Common and widespread in the Southern Boreal and the coastal southern part of the Middle Boreal. From Medelpad and northwards all records have been made within 60 km of the Baltic coast. The vast majority of records is from coastal areas or low altitudes in river valleys. Nevertheless, there are several records from above 300 m altitude in Hälsingland and Ångermanland, and a photo-documented occurrence at 425 m in Medelpad (15 km SW of Stöde). Although most common near the coast, the total lack of records from offshore islands in North Sweden indicates poor dispersal capacity over brackish water. There are no indications of changes in distribution during the last 50 years. Note, though, that the northernmost occurrence presented in the map (Figure 8; lower Byske River valley and the adjacent Tåme area to the north (Västerbotten)) became publicly known as late as 1989, despite the species having been known locally since at least the 1920’s (Södermark 1989). This occurrence has long been regarded as a disjunct population (Elmberg 1995) and perhaps the result of anthropogenic spread, but recent records around Skellefteå have gradually closed the previously supposed 70+ km distribution gap. Habitat and movements. In North Sweden this is the only reptile regularly encountered in closed forest, particularly in mesic stands with scattered deciduous trees and protective undergrowth. However, the most widely used habitats are fairly open, yet with denser undergrowth than those preferred by Vipera berus and Zootoca vivipara: forest edges, natural grasslands, shores, and rock outcrops. Most sightings of Anguis fragilis are made in anthropogenic habitats such as clear-cuts, fields, meadows, roadsides, and near recreational buildings (Figures 13, 19). Although it is the only reptile in North Sweden frequently seen active in cloudy weather, its general habits are seclusive. Shelter is typically found under flat stones, haystacks, compost piles, sheet metal, woodpiles, and tarps, in other words often under man-made objects. This habit may facilitate inadvertent anthropogenic spread. Daily and annual movements have not been studied, including any seasonal variation in habitat affinity. Subterranean hibernation sites are most likely found in or very close to the summer habitat, but nothing is known about hibernation habits in North Sweden. Abundance estimates and trends. There are not any published abundance data, but estimates based on extensive field work in the Umeå area (Västerbotten) 1975–1994 run in the neighborhood of> 200 adults /km 2 in landscapes with mixed habitat near the coast (Elmberg, unpublished, Stefan Andersson, personal communication). There are no indications of changes in abundance over the last 50 years., Published as part of Elmberg, Johan, 2023, Amphibians and reptiles in North Sweden: distribution, habitat affinities, and abundance (Classes: Amphibia and Reptilia), pp. 301-335 in Zootaxa 5301 (3) on page 319, DOI: 10.11646/zootaxa.5301.3.1, http://zenodo.org/record/8030434, {"references":["Sodermark, H. (1989) Var gar kopparodlans nordgrans i Sverige? Fauna och Flora, 84, 224 - 228. [in Swedish]","Elmberg, J. (1995) Groddjurens och kraldjurens utbredning i Norrland. Natur i Norr, 15 (2), 57 - 82. [in Swedish]"]}

Mohapatra, Pratyush P., Agarwal, Ishan, Mohalik, Rakesh Kumar, Dutta, Sushil K., Khandekar, Akshay (2023): Hemidactylus paucifasciatus (Squamata: Gekkonidae), a new species of large-bodied, tuberculate gecko from Northern Odisha, India. Zootaxa 5301 (3): 365-382, DOI: 10.11646/zootaxa.5301.3.3, URL: http://dx.doi.org/10.11646/zootaxa.5301.3.3

Cyrtopodion aravallense (Gill, 1997) is a poorly known species, known only from the holotype and original description. I re-examined the holotype housed in the museum of Bombay Natural History Society, Mumbai and find some discrepancies between the original description and the holotype. Here, I redescribe the holotype of C. aravallense in greater detail and provide a revised diagnosis of the species. It is distinguished from all its congeners in having enlarged, regularly arranged transverse rows of 15 trihedral tubercles; 25–26 midbody scale rows across belly; 102 midventral scales; males with 6 precloacal pores, 7–8 femoral pores on each side separated by 3–5 poreless scales between precloacal and femoral pores.

The Caatinga is the largest seasonal dry tropical forest in South America and it has been historically neglected in terms of its biodiversity. Regarding lizards, different studies led to the current knowledge of diversity and endemism in Caatinga, but detailed syntheses are scarce in the literature. We present the most detailed and up-to-date synthesis of knowledge about Caatinga lizards by providing a detailed (i) list of species; (ii) taxonomic richness patterns; (iii) knowledge gaps and spatial biases; and (iv) detailed distribution maps of all species that contain at least one occurrence record within Caatinga limits. We created a distribution database using occurrences of lizards in Caatinga based on scientific collections, field collection, and literature. We produce up-to-date distribution maps, calculate the Extent of Occurrence and provide the environmental and bioclimatic profile for each species recorded. We draw taxonomic richness and sampling gap maps. Our database has 20,538 records of occurrence of lizards of the Caatinga. We recorded 93 lizard species (13 families), 52.7% of which are endemic. Forthy-four percent of the species present restricted distributions. We identified that 53% of the Caatinga area (or 70% of the municipalities) has no record of occurrence of lizards. The data presented are an important step towards synthesizing in detail the accumulated knowledge about Caatinga lizards and is crucial for accurate strategies for the conservation planning. It directs actions to advance our knowledge on Caatinga lizards: to concentrate inventories in sample void areas; continuous update of the species occurrence database, advance in the generation of autoecology data for species.

We redescribe and illustrate the type specimens of ten taxa of sea snakes of the genera Hydrophis Latreille in Sonnini & Latreille, 1801 and Laticauda Laurenti, 1768 in the collections of the Zoological Survey of India. The specimens comprise holotypes and syntypes of ten synonymous nominal taxa that represent seven valid nominal taxa. We here clarify that one specimen ZSI 8278 is a syntype of Hydrophis dayanus Stoliczka, 1872, not holotype as previously stated. In one case, four holotypes of four nominal taxa are synonyms of the same taxon—Hydrophis cyanocinctus Daudin, 1803. Many of these type specimens are herein first depicted in photographs in a publication.

This study presents a comprehensive morphological comparison along with molecular phylogeny of the genus Gloydius based on five mitochondrial genes (12S, 16S, COI, cytb, and ND4). The specimens collected from Jiuzhaigou National Nature Reserve are shown to be a new species, Gloydius lateralissp. nov. Zhang, Shi, Jiang & Shi based on a combination of morphological and molecular accounts. G. lateralissp. nov. differs from other congeneric species by a series of diagnostic morphological characteristics and forms a strongly supported monophyletic group. The new species is phylogenetically closely related to G. swild, another recently described species from Heishui, Aba, Sichuan.

The genus Anadia (family Gymnophthalmidae) consists of 19 species. It has remained almost taxonomically stable for decades, scarcely observed, in addition to being one of the less sampled gymnophthalmid genera with respect to molecular phylogenies. New Anadia species are discovered at a relatively low pace, and few specimens are found in the field, probably due to the arboreal habits of many low and mid elevation species. We describe here a new species of Anadia from the cloud forests of northwestern Colombia: the new species is easily diagnosed by the combination of shape and imbrication of dorsal scales, very large body size, the largest within its group, and large and non-overlapping number of longitudinal scale rows around midbody. We also tested the phylogenetic position of the recently described and geographically close A. antioquensis. A phylogenetic analysis based on four genomic regions recovered the new species as sister to A. buenaventura, whereas A. antioquensis was reassigned to the genus Riama. The new species is currently known from only three specimens, collected throughout eight years within less than 5 ha of the Mesenia-Paramillo Nature Reserve. Although its apparent rarity may be due to secretive habits, the species is provisionally declared vulnerable, while new information is available. To stimulate further research on this genus, we also compiled and present here the comparable information on the distribution and morphology of Anadia species. Altogether, the results stress the urge for a new review of the genus with the help of molecular data.

Cyrtodactylus monilatussp. nov. is described from Si Sawat District, Kanchanaburi Province, in western Thailand. The new species superficially resembles C. zebraicus Taylor, 1962 from southern Thailand. However, differences between the new species from C. zebraicus and other congeners were supported by an integrative taxonomic analysis of molecular and morphological data. Phylogenetic analyses based on the mitochondrial NADH dehydrogenase subunit 2 (ND2) gene showed that the new species is a member of the C. oldhami group and closely related to Cyrtodactylus sp. MT468911 from Thong Pha Phum National Park, Thong Pha Phum District, Kanchanaburi Province. Uncorrected pairwise genetic divergences (p-distances) between the new species and its congeners, including C. zebraicus, ranged from 7.7–17.7%. Cyrtodactylus monilatussp. nov. can also be distinguished from all members of the C. oldhami group by having a unique combination of morphological characters, including a snout to vent length of 53.7–63.3 mm in adult males and 58.6–75.8 mm in adult females; 22–34 paravertebral tubercles; 34–42 ventral scales; 30–39 enlarged contiguous femoroprecloacal scales; femoral pores and precloacal pores absent in both sexes; four or five rows of postprecloacal scales; enlarged median subcaudal scales absent; weak ventrolateral folds present; 4–7 rows of paired, paravertebral, dark-brown blotches edged in yellow or yellowish white; and two rows of small, diffuse, yellow or yellowish white spots on flanks. The new species occurs in a narrow range of forest at mid to low elevations associated with karst landscapes in the Tenasserim mountain range.

Emydopoidea is one of the major dicynodont subclades and includes some purported fossorial taxa. Various cranial and postcranial adaptations for fossoriality have long been recognized in cistecephalid emydopoids, but anatomical variation of their braincases remains poorly understood. Here, using laboratory and synchrotron X-ray tomography, we provide detailed anatomical descriptions of the basicranial axis of three emydopoids (Myosaurus, Kawingasaurus and a Malawian cistecephalid DMMM-PK-16-1) and compare them to the basal dicynodont Pristerodon. Cistecephalids show the presence of divergent crests on the posterior aspect of the opisthotic and a nuchal crest on their occipital plate, contrasting with the featureless occipital plate of other dicynodonts. These depressions and crests increase the attachment area of the atlanto-occipital muscles, suggesting that cistecephalids were capable of powerful movements of the head during digging. Additionally, Kawingasaurus has a pneumatized braincase and highly co-ossified basicranium, which is probably linked to the auditory system. We corroborate the hypothesis that cistecephalids, in addition to being forelimb diggers, were likely head-lift diggers, and we highlight some derived adaptations consistent with a quasi-obligate fossorial lifestyle. Furthermore, new basicranial phylogenetic characters and a re-evaluation of emydopoid relationships are proposed. We recovered Rastodon as a basal emydopoid, Thliptosaurus as a non-kingoriid emydopoid and novel interrelationships among cistecephalids.

Secondarily marine tetrapod lineages have independently evolved osmoregulatory adaptations for life in salt water but inferring physiological changes in extinct marine tetrapods is difficult. The Mesozoic crocodylomorph clade Thalattosuchia is unique in having both direct evidence from natural endocasts and several proposed osteological correlates for salt exocrine glands. Here, we investigate salt gland evolution in thalattosuchians by creating endocranial reconstructions from CT scans of eight taxa (one basal thalattosuchian, one teleosauroid, two basal metriorhynchoids and four metriorhynchids) and four outgroups (three extant crocodylians and the basal crocodyliform Protosuchus) to identify salt gland osteological correlates. All metriorhynchoids show dorsolateral nasal cavity expansions corresponding to the location of nasal salt glands in natural casts, but smaller expansions in teleosauroids correspond more with the cartilaginous nasal capsule. The different sizes of these expansions suggest the following evolutionary sequence: (1) plesiomorphically small glands present in semi-aquatic teleosauroids draining through the nasal vestibule; (2) moderately sized glands in the basalmost metriorhynchoid Pelagosaurus; and (3) hypertrophied glands in the clade comprising Eoneustes and metriorhynchids, with a pre-orbital fenestra providing a novel exit for salt drainage. The large gland size inferred from basal metriorhynchoids indicates advanced osmoregulation occurred while metriorhynchoids were semi-aquatic. This pattern does not precisely fit into current models of physiological evolution in marine tetrapods and suggests a unique sequence of changes as thalattosuchians transitioned from land to sea.

We report the rediscovery of Oligodon melaneus 112 years after its original description and document the third, and only non-type, specimen for the species. The new specimen was found 267 km east of the type locality (Tindharia, West Bengal state) from Assam state, India. We designate a lectotype for the species, and provide an extended description of a freshly collected male specimen. Phylogenetic analyses of 16s and cytb mitochondrial genes provide support for O. melaneus being closely related to the widespread South Asian endemic O. arnensis.

Recent phylogenetic analyses differ in their interpretations of the origin and interrelationships of snakes, resulting in polarized views of snake ecology, habit and acquisition of features associated with wide-gaped feeding (macrostomy). Here, we report a new specimen of the Late Cretaceous nest predator Sanajeh indicus that helps to resolve the origin of macrostomy. The new specimen preserves an ossified upper temporal bar and a posteriorly expanded otooccipital region that lacks a free-ending supratemporal bone and retains a lizard-like palatomaxillary arch that allows limited movements during swallowing. Phylogenetic analyses of a large-scale total evidence dataset resolve Sanajeh near the base of Pan-Serpentes, as the sister group of Najash, Dinilysia and crown-group Serpentes. The Cretaceous Tetrapodophis and Coniophis represent the earliest-diverging members of Pan-Serpentes. The Cretaceous hindlimbed pachyophiids and Cenozoic Australian ‘madtsoiids’ are inside crown Alethinophidia, whereas mosasaurs are recovered invariably within anguimorphs. Our results suggest that the wide-gape condition in mosasaurs and snakes might have evolved independently, as functionally distinct mechanisms of prey ingestion. The intermediate morphology preserved in Sanajeh indicates that ingestion of large prey items (macrophagy) preceded wide-gaped, unilateral feeding (macrostomy), which appeared 35 Myr later, in the common ancestor of pachyophiids, Cenozoic Australian ‘madtsoiids’ and alethinophidians.

The gekkonid diversity of the under-explored dry zones in Peninsular India is not yet completely inventoried, as evidenced by frequent new species descriptions. Here, we describe two new species of Hemidactylus from the dry rocky zones of Deccan Plateau viz. Hemidactylus mahonyi sp. nov. from Sandur Hills and Hemidactylus srikanthani sp. nov. from Devarayana Durga Hills, Karnataka State of Peninsular India. The new species belong to the morphologically cryptic H. murrayi Clade but differ from each other and the known congeners by a set of morphological characters and possess non-overlapping distribution ranges. Furthermore, the new species are also divergent (> 6.4% uncorrected p-distance) from their nearest related congeners, and from one another with respect to the ND2 mitochondrial gene. Recently, other new species of reptiles have been described from the two aforesaid type localities, indicating that further bio-inventories in the dry zones of peninsular India will yield more such novelties.

Mexico is a megadiverse country with high endemicity in its herpetofauna. We examine how species richness, proportion of state and country endemic species, and proportion of species in a category of conservation concern using listings in the International Union for Conservation of Nature (IUCN) Red List and the Secretaría del Medio Ambiente y Recursos Naturales (SEMARNAT) in 27 of 32 Mexican states are related to environmental and human demographic and socioeconomic variables. Amphibian and reptile species richness were positively related to latitude range and number of physiographic regions and negatively related to latitude. The proportion of state endemic amphibian species in a state was negatively related to latitude whereas no variables influenced the proportion in reptiles. The proportion of country endemics in a state was positively related to human population density and the number of physiographic regions and negatively related to per capita gross domestic product (GDP) and latitude range for amphibians; it was positively related to human population density and elevation range and negatively related to latitude range for reptiles. The proportion of amphibian species in an IUCN category of concern in a state was positively related to human population density and negatively related to latitude; for reptiles, it was negatively related to human population density. The proportion of SEMARNAT-listed species in a state was positively related to human population density for both amphibians and reptiles and negatively related to latitude range for amphibians. Our analyses found that larger macroecological patterns (e.g., latitudinal species gradient, heterogeneity-richness relationships) and human population density play important roles in determining the richness and conservation status of Mexican amphibians and reptiles.

Here we provide the first phylogenetic analysis that include Afrogecko ansorgii and a detailed morphological comparison with other species of leaf-toed geckos. For this purpose, we used two mitochondrial (16S, ND2) and four nuclear (RAG1, RAG2, CMOS, PDC) genes to produce a robust phylogenetic reconstruction. This allowed us to show that A. ansorgii is not related as previously believed to circum-Indian Ocean leaf-toed geckos and is rather more closely related to other Malagasy leaf-toed geckos. Additionally, we explore and compare osteological variation in A. ansorgii skulls through High Resolution X-ray Computed Tomography with previously published material. This allowed us to describe herein a new genus, Bauerius gen. nov., and additionally provide a detailed redescription of the species (including the first description of male material), supplementing the limited original description and type series, which consisted of only two females.

Osteological knowledge of the sauropod dinosaur Ligabuesaurus leanzai is increased by the description of new postcranial elements assigned to the holotype MCF-PVPH-233. Furthermore, a newly referred specimen, MCF-PVPH-228, is recognized after a detailed revision of the abundant sauropod material collected from the Lohan Cura Formation outcrops in the Cerro de los Leones locality (southern Neuquén Basin, Patagonia, Argentina). Recent laboratory preparation and fieldwork allowed us to recognize several new morphological features of the pectoral and pelvic girdles and the cervical and caudal anatomy. Thus, a new diagnosis of Ligabuesaurus is proposed that includes new autapomorphies and a unique combination of features. A phylogenetic analysis based on this new material recovers Ligabuesaurus as a non-titanosaurian somphospondylan, more derived than Sauroposeidon. Therefore, we discuss the palaeobiogeographical implications for the diversification and distribution of South American somphospondylans, especially in the Neuquén Basin, which are closely related to the early stages of evolution of Titanosauria. In this context, Ligabuesaurus represents one of the more complete Early Cretaceous Titanosauriformes and the earliest non-titanosaurian somphospondylan of South America. Finally, the new information on Ligabuesaurus contributes not only to reconstruction of the sauropod faunal composition of south-western Gondwana, but also sheds light on the early stages and emergence of titanosaurians.