Skink Research Articles

Ecological divergence combined with ancient allopatry in lizard populations from a small volcanic island

Population divergence and speciation are often explained by geographical isolation, but may also be possible under high gene flow due to strong ecology-related differences in selection pressures. This study combines coalescent analyses of genetic data (11 microsatellite loci and 1 Kbp of mtDNA) and ecological modelling to examine the relative contributions of isolation and ecology to incipient speciation in the scincid lizard Chalcides sexlineatus within the volcanic island of Gran Canaria. Bayesian multispecies coalescent dating of within-island genetic divergence of northern and southern populations showed correspondence with the timing of volcanic activity in the north of the island 1.5-3.0 Ma ago. Coalescent estimates of demographic changes reveal historical size increases in northern populations, consistent with expansions from a volcanic refuge. Nevertheless, ecological divergence is also supported. First, the two morphs showed non-equivalence of ecological niches and species distribution modelling associated the northern morph with mesic habitat types and the southern morph with xeric habitat types. It seems likely that the colour morphs are associated with different antipredator strategies in the different habitats. Second, coalescent estimation of gene copy migration (based on microsatellites and mtDNA) suggest high rates from northern to southern morphs demonstrating the strength of ecology-mediated selection pressures that maintain the divergent southern morph. Together, these findings underline the complexity of the speciation process by providing evidence for the combined effects of ecological divergence and ancient divergence in allopatry.

Activity Patterns and Movements of Free-Ranging Bluetongue Lizards (Tiliqua scincoides intermediaandTiliqua multifasciata) in the Australian Wet-Dry Tropics

Abstract We used GPS-based radiotelemetry to quantify diel activity patterns and movements by two species of large scincid lizards in the Australian wet-dry tropics. From an average of 63 days tracking of 54 Northern Bluetongues (Tiliqua scincoides intermedia) and 42 days tracking of 8 Centralian Bluetongues (Tiliqua multifasciata), we obtained 53,801 and 8,139 data points, respectively, on lizard locations over a 13-month period. Lizard species, sex, and study site had relatively little effect on distances moved, at both daily and hourly levels. However, Northern Bluetongues moved further per day than did Centralian Bluetongues at a site where they were sympatric. Most movements were <20 m, with longer movements (dispersal between core areas within the wider home range) occurring primarily during midmorning and late afternoon. The only major effect of ambient temperatures on lizard activity patterns likely was that midday movements were precluded by the risk of overheating; at all other times of the day ...

Notes Taxonomy, Distribution, Geographic Variation and Natural History in Trachylepis vittata (Olivier, 1804) in the Zagros Mountains,Western Iran

The Bridled Skink, Trachylepis vittata is a banded skink finitely distributed in western Iran; in Kermanshah, Lorestan and IlamProvinces. In present study, geographic variation was analyzed in 48 specimens of T. vittata for meristic and morphometric characters in three populations from western Iran by the independent sample t -test, the one way ANOVA (LSD-Test) and the principal component analysis. Results of analyses show subtle geographic variation among populations of west Iran. We report observations on the natural history of this skink in mentioned provinces. The conservation status, taxonomy, and habitat of T. vittata is discussed. This species is not regarded as threatened.

A new species of Plestiodon (Squamata: Scincidae) from Kuchinoshima Island in the Tokara Group of the Northern Ryukyus, Japan.

A scincid lizard of the genus Plestiodon from Kuchinoshima Island in the Tokara Group of the Northern Ryukyus, Japan, has proved to be genetically and morphologically differentiated from any previously recognized species in the genus. We thus describe this island population as a new species, Plestiodon kuchinoshimensis. The new species shows characteristics of the P. latiscutatus species group, but differs from other species of this group by the combination of the following character states: postnasal absent; hatchling with five longitudinal light lines on dorsum; lateral light line on each side passing over ear opening and the sixth to eighth scale rows at midbody; dorsolateral light line beginning from behind supraoculars; patch of enlarged irregular scales on posterior femur absent; scale rows around midbody 27-32; and brownish background on the dorsal surface of the juvenile.

A new species of lowland skink (genus Lipinia Gray, 1845) from northeastern Peninsular Malaysia.

A new species of scincid lizard, Lipinia sekayuensis sp. nov. from Hutan Lipur Sekayu, Terengganu State in northeastern Peninsular Malaysia is most similar to L. surda (Boulenger) but differentiated from it and all other species of Lipinia by having the combination of an adult SVL of 42.3 mm; six supralabials; five infralabials; four supraoculars; prefrontals widely separated; two loreals; fused frontoparietals; lower eyelids bearing a large, transparent disc; 21 midbody scale rows; 56 paravertertebral scale rows; 65 ventral scale rows; enlarged, precloacal scales; 10 subdigital lamellae on the third finger; 11, 15, and seven lamellae on the third, fourth, and fifth toes, respectively; distal subdigital lamellae keeled; a median row of slightly enlarged, subcaudal scales present; a generally unicolor, dark-brown dorsum bearing nine very faint, diffuse, darker stripes; and an external ear opening replaced by a scaly, auditory depression.

Phylogenetic disassembly of species boundaries in a widespread group of Australian skinks (Scincidae: Ctenotus)

Scincid lizards in the genus Ctenotus represent one of Australia’s most species-rich vertebrate clades, with more than 100 recognized species. Formal diagnoses of many species have relied on qualitative assessments of adult color pattern, but the validity of many such species has not been tested in a phylogenetic framework. We used mitochondrial and nuclear DNA to perform the first phylogenetic analysis of species in the Ctenotus inornatus group, a complex of at least 11 nominal forms that are distributed widely across the Australian continent. Mitochondrial and nuclear gene phylogenies support the presence of multiple species in the group, but these clades largely fail to match species boundaries as currently defined. Multivariate analyses of color pattern indicate that extreme intraspecific morphological variation in this character has created a significant impediment to understanding taxonomic diversity in the group. Our results suggest that nearly all species in the C. inornatus group require substantial taxonomic revision, and several geographically widespread forms (“C. saxatilis” and “C. robustus”) appear to be polyphyletic taxa drawn from phenotypically similar but genetically distinct lineages. We describe one new species and provide redescriptions for three additional species. We synonymize names applied to a number of genetically incoherent or otherwise poorly-defined forms. The results of our study highlight an acute need for population genetic studies of species boundaries in Australian skinks, many of which are recognized by morphological traits that vary greatly within and between populations.

Analysis and Visualization of Complex Macroevolutionary Dynamics: An Example from Australian Scincid Lizards

The correlation between species diversification and morphological evolution has long been of interest in evolutionary biology. We investigated the relationship between these processes during the radiation of 250+scincid lizards that constitute Australia's most species-rich clade of terrestrial vertebrates. We generated a time-calibrated phylogenetic tree for the group that was more than 85% complete at the species level and collected multivariate morphometric data for 183 species. We reconstructed the dynamics of species diversification and trait evolution using a Bayesian statistical framework (BAMM) that simultaneously accounts for variation in evolutionary rates through time and among lineages. We extended the BAMM model to accommodate time-dependent phenotypic evolution, and we describe several new methods for summarizing and visualizing macroevolutionary rate heterogeneity on phylogenetic trees. Two major clades (Lerista, Ctenotus; >90 spp. each) are associated with high rates of species diversification relative to the background rate across Australian sphenomorphine skinks. The Lerista clade is characterized by relatively high lability of body form and has undergone repeated instances of limb reduction, but Ctenotus is characterized by an extreme deceleration in the rate of body shape evolution. We estimate that rates of phenotypic evolution decreased by more than an order of magnitude in the common ancestor of the Ctenotus clade. These results provide evidence for a modal shift in phenotypic evolutionary dynamics and demonstrate that major axes of morphological variation can be decoupled from species diversification. More generally, the Bayesian framework described here can be used to identify and characterize complex mixtures of dynamic processes on phylogenetic trees. [Bayesian; diversification; evolvability; lizard; macroevolution, punctuated equilibrium, speciation.].

Molecular insight into systematics, host associations, life cycles and geographic distribution of the nematode family Rhabdiasidae

Rhabdiasidae Railliet, 1915 is a globally distributed group of up to 100 known species of nematodes parasitic in amphibians and reptiles. This work presents the results of a molecular phylogenetic analysis of 36 species of Rhabdiasidae from reptiles and amphibians from six continents. New DNA sequences encompassing partial 18S rDNA, ITS1, 5.8S rDNA, ITS2 and partial 28S rDNA regions of nuclear ribosomal DNA were obtained from 27 species and pre-existing sequences for nine species were incorporated. The broad taxonomic, host and geographical coverage of the specimens allowed us to address long-standing questions in rhabdiasid systematics, evolution, geographic distribution, and patterns of host association. Our analysis demonstrated that rhabdiasids parasitic in snakes are an independent genus sister to the rest of the Rhabdiasidae, a status supported by life cycle data. Based on the combined evidence of molecular phylogeny, morphology and life cycle characteristics, a new genus Serpentirhabdias gen. nov. with the type species Serpentirhabdias elaphe (Sharpilo, 1976) comb. nov. is established. The phylogeny supports the monophyly of Entomelas Travassos, 1930, Pneumonema Johnston, 1916 and the largest genus of the family, Rhabdias Stiles and Hassall, 1905. DNA sequence comparisons demonstrate the presence of more than one species in the previously monotypic Pneumonema from Australian scincid lizards. The distribution of some morphological characters in the genus Rhabdias shows little consistency within the phylogenetic tree topology, in particular the apical structures widely used in rhabdiasid systematics. Our data suggest that some of the characters, while valuable for species differentiation, are not appropriate for differentiation among higher taxa and are of limited phylogenetic utility. Rhabdias is the only genus with a cosmopolitan distribution, but some of the lineages within Rhabdias are distributed on a single continent or a group of adjacent zoogeographical regions. Serpentirhabdias, Entomelas and Pneumonema show rather strict specificity to their host groups. The evolution of the Rhabdiasidae clearly included multiple host switching events among different orders and families of amphibians as well as switching between amphibians and squamatan reptiles. Only a few smaller lineages of Rhabdias demonstrate relatively strict associations with a certain group of hosts.

Does Invasion Success Reflect Superior Cognitive Ability? A Case Study of Two Congeneric Lizard Species (Lampropholis, Scincidae)

A species' intelligence may reliably predict its invasive potential. If this is true, then we might expect invasive species to be better at learning novel tasks than non-invasive congeners. To test this hypothesis, we exposed two sympatric species of Australian scincid lizards, Lampropholis delicata (invasive) and L. guichenoti (non-invasive) to standardized maze-learning tasks. Both species rapidly decreased the time they needed to find a food reward, but latencies were always higher for L. delicata than L. guichenoti. More detailed analysis showed that neither species actually learned the position of the food reward; they were as likely to turn the wrong way at the end of the study as at the beginning. Instead, their times decreased because they spent less time immobile in later trials; and L. guichenoti arrived at the reward sooner because they exhibited “freezing” (immobility) less than L. delicata. Hence, our data confirm that the species differ in their performance in this standardized test, but neither the decreasing time to find the reward, nor the interspecific disparity in those times, are reflective of cognitive abilities. Behavioural differences may well explain why one species is invasive and one is not, but those differences do not necessarily involve cognitive ability.

Weight watching in burrows: variation in body condition in pygmy bluetongue lizards

The pygmy bluetongue lizard, Tiliqua adelaidensis, is an endangered scincid lizard that occurs in remnants of natural grasslands in the mid-north of South Australia. We assessed the factors affecting body condition of male and female lizards, using body size residuals as an index, over five sampling years. We included sex, phenotypic (patterned or plain morphs), temporal (sampling year and activity period within year), and climatic factors in our analyses. The results indicated that sampling year and activity period within the year were the two most important factors influencing variation in body condition of both male and female lizards over the period of the study. There were similar trends when we considered males and females separately. However, when prepartum and postpartum females were analysed separately, sampling year did not affect their body condition. None of the analyses showed any significant effect of phenotype on body condition. Winter–spring rainfall was positively correlated with body condition of females in the prepartum period, but showed no significant effect on mean body condition of any other grouping of adult lizards, nor a consistent direction of correlation among the different subsets of adult lizards that we considered. The substantial annual variations in the body condition of lizards, although of uncertain cause, provide important information for conservation managers who monitor persisting populations of this endangered species.

Evolution and adaptation: a tribute to Richard Essex Barwick

Richard (Dick) Essex Barwick was an influential researcher whose interests spanned many branches of biology and zoology, including herpetology, vertebrate morphology, palaeontology and evolutionary biology. Although born and raised in New Zealand, he spent much of his career in Australia and made considerable contributions to Australian zoology. Born in Christchurch in 1929, Dick spent many hours in his youthwatchingandstudyinganimalsthatwerelocaltotheregion. After earning his bachelor’s degree at Victorian University College (later Victorian University of Wellington (VUW)), he thencompletedanMSc(Honours)thatinvestigatedthelifecycles and ecology of scincid lizards. This work marked the beginning of Dick’s career in herpetology, although this was only one area in which he applied his tireless efforts. After completing his MSc in the mid-1950s, Dick was appointed a junior lecturer at VUW, which sparked his second major area of investigation. He was one of four adventurers (along with Colin Bull, Barry McKelvey and Peter Webb) who initiated studies of the biology and geology of the dry valleys of the Antarctic. Dick was the biologist and artist on three Antarctic summer trips and developed an affinity for all things Antarctic as a result (Fig. 1). The VUW Antarctic Expedition named Barwick Valley in the dry region (77 � 21 0 0 00 S, 161 � 10 0 0 00 E) in his honour.

Cranial Osteology of the Bridled Skink, Trachylepis vittata (Olivier, 1804), and Its Comparison with Trachylepis aurata transcaucasica (Chernov, 1926) (Sauria: Scincidae)

In spite of high diversity of the Iranian Plateau herpetofauna, especially the lizards, and numerous studies done on the morphometric characters of lizards, information on the detailed characters of their skeleton remains insufficient. Herein, the adult cranial osteology of the bridled skink, Trachylepis vittata (Olivier, 1804), is described in detail for the first time based on cleared and double-stained skeletal elements and compared with Trachylepis aurata transcaucasica (Chernov, 1926). In this study, characters of the skull and mandibular elements were examined. The existence of slight differences between the two studied species according to bone and skull characters, are explained. The skull of both species, as typical representatives of the Scincidae, bear teeth on the premaxilla, maxilla, and pterygoids. The mandibular teeth are present on the dentaries. Some obvious differences that show off are: different number of pleurodont maxillary teeth as T. vittata and T. a. transcaucasica have 20 and 21–26 maxillary pleurodont teeth, respectively, narrower tip of the nasal processes of the premaxilla in T. vittata than that of T. a. transcaucasica , different ornamentation of the nasal processes of the frontals as are more projected to the nasals in T. vittata than those of T. a. transcaucasica , slightly higher degree of serration on the posterior margin of the frontals in T. vittata than that of T. a. transcaucasica , and fine differences in form, size and special manner of articulation in bones such as: the lacrimals, prefrontals, postfrontals, palatines, pterygoids, dentary and prearticular.

Lizard Wears Shades. A Spectacled Sphenomorphus (Squamata: Scincidae), from the Sacred Forests of Mawphlang, Meghalaya, North-east India.

A new species of lygosomatine scincid lizard is described from the sacred forests of Mawphlang, in Meghalaya, northeastern India. Sphenomorphus apalpebratus sp. nov. possesses a spectacle or brille, an unusual feature within the Scincidae, and a first for the paraphyletic genus Sphenomorphus. The new species is compared with other members of the genus to which it is here assigned, as well as to members of the lygosomatine genera Lipinia and Scincella from mainland India, the Andaman and Nicobar Islands, and southeast Asia, to which it also bears resemblance. The new taxon is diagnosable in exhibiting the following combination of characters: small body size (SVL to 42.0 mm); moveable eyelids absent; auricular opening scaleless, situated in a shallow depression; dorsal scales show a line of demarcation along posterior edge of ventral pes; midbody scale rows 27-28; longitudinal scale rows between parietals and base of tail 62-64; lamellae under toe IV 8-9; supraoculars five; supralabials 5-6; infralabials 4-5; subcaudals 92; and dorsum golden brown, except at dorsal margin of lateral line, which is lighter, with four faintly spotted lines, two along each side of vertebral row of scales, that extend to tail base. The new species differs from its congeners in the lack of moveable eyelids, a character shared with several distantly related scincid genera.

Recent cryptic extinction of squamate reptiles on Yoronjima Island of the Ryukyu Archipelago, Japan, inferred from garbage dump remains

We report recent skeletal remains of squamate reptiles screened from an old garbage dump deposit found on Yoronjima Island, a small island of the Ryukyu Archipelago, in the subtropical northwestern Pacific. Identified remains include at least three species of terrestrial snakes, one sea snake, and seven species of lizards. Among these, the Ryukyu Short-legged Skink, Ateuchosaurus pellopleurus ; Kuroiwa’s Leopard Gecko, Goniurosaurus kuroiwae ; and one unidentified Gekko species constitute entirely new findings. For a further two snake species and one lizard species, the evidence presented here is the first specimen-based documentation that these species existed on this island, where they are now evidently absent. The present absence of these species on Yoronjima is mainly attributable to predation by the Japanese Weasel, Mustela itatsi , introduced in the mid-1950s. Our results demonstrate a prominent case of recent, sizable deterioration of insular herpetofaunal diversity, which has never been documented with certainty in the Ryukyu Archipelago, and suggest that human influence should be taken into account in biodiversity research in this area.

Phylogeny and divergence times of Australian Sphenomorphus group skinks (Scincidae, Squamata)

The Australian Sphenomorphus group is a diverse clade (c. 250 species) of scincid lizards (skinks) incorporating more than half of the Australian scincid fauna. Previous phylogenetic analyses of mitochondrial and nuclear gene sequence data for selected Australian Sphenomorphus group scincids have provided support for several morphologically- and ecologically-distinct clades; however, the relationships among these clades are only incompletely resolved. This paper presents a new phylogenetic analysis of the Australian Sphenomorphus group, based on nucleotide sequences for three regions of the mitochondrial genome and four nuclear genes (5645 aligned sites in total). Phylogenies generated using standard concatenation and multi-species coalescent approaches are generally similar, and nearly all conflicting nodes are weakly supported. Monophyly of a number of genera and other (informal) supra-specific taxa, including Calyptotis, Ctenotus, Eremiascincus, Hemiergis, Lerista, Notoscincus, the ‘Eulamprus’ quoyii group, and the ‘Glaphyromorphus’ crassicaudis group is well supported in all analyses. There is significant support for a clade including the ‘Eulamprus’ tenuis group, Gnypetoscincus, and Nangura, and a clade comprising Coeranoscincus, Coggeria, Ophioscincus, and Saiphos receives at least marginal support (posterior probabilities above 0.93). All analyses indicate that Anomalopus is polyphyletic, although topology tests suggest that support for this result is equivocal. Divergence times inferred using relaxed molecular clock methods are consistent with an Oligocene origin of the Australian Sphenomorphus group following trans-oceanic dispersal from Asia. Age estimates for clades distributed primarily in arid habitats (c. 9–17million years) are generally younger than those for clades occurring in mesic environments (c. 15–22million years); however, arid-zone clades are substantially more diverse, including nearly 80% of all Australian Sphenomorphus group scincids. This pattern conforms well with reconstructions of Australian palaeo-environments during the Cenozoic, which indicate a progressive shift from widespread mesic habitats in the Early Miocene (prior to 16Mya) to predominantly arid environments 4–2Mya.

Comparative cytogenetics of two species of ground skinks: Scincella assata and S. cherriei (Squamata: Scincidae: Lygosominae) from Chiapas, Mexico

Standard karyotypes of two species of the genus Scincella , S. assata and S. cherriei , both from Chiapas State, Mexico, were described for the first time. The diploid chromosome number was 28 in S. assata , whereas 30 in S. cherriei . The karyotypes of the two species, while differing in the number of microchromosomes, 14-15 in S. assata and 16-17 in S. cherriei, share four pairs of large metacentric, two pairs of medium-sized metacentric, and one particular pair (number 7) of chromosomes. Female S. assata carries chromosome pair 7 composed of two identical medium-sized subtelocentric chromosomes. This chromosome pair is heteromorphic in males of both species, i.e., one component of the pair is similar to the homomorphic chromosomes 7 of the S. assata female, while the other is nearly one-half the size of its counterpart and resembles a microchromosome. The homology of such externally different elements is deducted from the presence of an asymmetric bivalent in spermatocytes at diplotene-diakinesis. Female S. cherriei was not available. We suspect that the two Scincella species possess an XY sex determination system, as previously reported for the North American congeneric species, S. lateralis .

Distribution of Ablepharus kitaibelii fitzingeri Mertens, 1952 (Squamata: Scincidae) in Hungary

Ablepharus kitaibelii fitzingeri is the northernmost subspecies of the snake-eyed skink (the only representative of the genus in Europe), and an endemic species of the Carpathian Basin. Despite of its unique status in the Hungarian herpetofauna and its strict protection, no recent summary has been provided on its distribution in the country. In the last attempt, Dely (1978) presented 40 localities of the subspecies. Here we give an updated distribution of the subspecies (100 localities), based on records from the literature, collection of the Hungarian Natural History Museum (HNHM), field surveys made by the authors and observations of field researchers. The current conservation status of A. k. fitzingeri in Hungary is discussed and an updated distribution map is provided.

Current Population Status of Diurnal Lizards of Karachi, Pakistan

Five districts of Karachi (East, West, South, Central and Malir) contain resident populations of lizards. During the present studies the current status of diurnal lizards of Karachi was investigated. The faunistic studies were based on seven species from three districts East, South and Malir of Karachi. 182 lizards were captured belonging to seven species, six genera and four families; Agamidae, Scincidae, Lacertidae and Varanidae. Among the 182 captured lizards were 56 Indian Garden Lizards, Calotes versicolor ; 20 Indian Spiny-tailed Lizards, Uromastyx hardwickii ; 72 Bronze Grass Skinks, Mabuya macularia ; 8 Striped Grass Skinks, Mabuya dissimilis ; 15 Indian Fringe-toad Sand Lizards, Acanthodactylus cantoris ; 10 Long-tailed Desert Lacertas, Mesalina watsonana , and 1 Indian Monitor Varanus bengalensis . The results of the present study indicate that the current status of diurnal lizards, i.e., Indian Garden Lizard ( Calotes versicolor ), Bronze Grass Skink ( Mabuya macularia ), Striped Grass Skink ( Mabuya dissimilis ), Indian Fringe-toad sand Lizard ( Acanthodactylus cantoris ), and Long-tailed Desert Lacerta ( Mesalina watsonana ) are common in three selected districts of Karachi, while the Indian Spiny-tailed Lizard ( Uromastyx hardwickii ) and Indian Monitor lizard ( Varanus bengalensis ) have been recorded as threatened species. Uromastyx hardwickii is an endangered species and is already included in the IUCN Red List and CITES Appendix. Internationally, 25% of reptiles are listed as threatened (IUCN, 2002). In Pakistan some lizard species are threatened. Present studies also recorded Uromastyx hardwickii and Varanus bengalensis as threatened species in the three districts of Karachi. The Brilliant Agama ( Trapelus agilis isolepis ) was not found in the three districts of Karachi. Either it is threatened or has gone extinct or has been shifted to other habitats due to destruction of habitat and increase of human population.

DIEL PATTERNS OF ACTIVITY AND OF SELECTED AMBIENT TEMPERATURE OF THE SAND-SWIMMING LIZARD SPHENOPS SEPSOIDES (REPTILIA: SCINCIDAE)

ABSTRACT Diel patterns of activity and temperature selection were determined for two age classes of the sand-swimming skink, Sphenops sepsoides, under laboratory conditions. Activity levels were the highest between 1500 and 2100 h, and lowest between 2100 and 0600 h. Selected ambient temperatures averaged 29.7°C but varied during the diel cycle with the highest temperatures recorded at 1800 h and the lowest at 1200 h. Selected ambient temperatures were consistently about 1°C higher for juvenile than for adult individuals. The period that lizards were most active in corresponded to the period of highest selected ambient temperature. These patterns do not match those of nocturnal geckos in the field, which are the most active after dark but select high body temperatures during the day. However, these results are consistent with observations of activity and selected body temperatures of Eremiascincus fasciolatus, another sand-swimming skink. Field observations on S. sepsoides suggest that sub-surface activit...

THE DIET OF VARANUS GRISEUS IN THE SOUTHERN COASTAL PLAIN OF ISRAEL (REPTILIA: SAURIA)

ABSTRACT Fecal pellets were collected and analyzed microscopically during a two-year study of the desert monitor (Varanus griseus) in a sand-dune area south of Holon. V. griseus was found to be a predator-scavenger carnivore type, eating almost any animal of appropriate size. Although 92.2% of the fecal pellets contained invertebrates (mainly arthropods), the higher frequency of occurrence of the smaller species among the invertebrates is probably due to secondary predation. The main prey items included rodents (mostly gerbils, Gerbillus sp., and jirds, Meriones sp.), bird eggs (presumably chukar partridge, Alectoris chukar eggs), birds (mainly partridge chicks), skinks (Chalcides sp.), fringe-toed lizards (Acanthodactylus sp.), chameleons (Chamaeleo chamaeleon), elegant geckos (Stenodactylus sthenodactylus), tortoises (Testudo graeca) and snakes (including the venomous Vipera xanthina palaestinae). Less frequent were: hedgehogs (Hemiechinus auritus), leverets (Lepus europaeus), Sanis pygmy shrew (Suncus ...