Scorpion speciation in the Holy Land: Multilocus phylogeography corroborates diagnostic differences in morphology and burrowing behavior among Scorpio subspecies and justifies recognition as phylogenetic, ecological and biological species.

Scorpio Linnaeus, 1758 (family Scorpionidae Latreille, 1802) was considered monotypic for over a century, and comprised a single species, Scorpio maurus Linnaeus, 1758, with 19 subspecies, distributed from West Africa, throughout the Maghreb and the Middle East, to Iran. Two parapatric subspecies, Scorpio maurus fuscus (Ehrenberg, 1829) and Scorpio maurus palmatus (Ehrenberg, 1828), have long been recognized in the eastern Mediterranean region. We examined morphological variation, burrow architecture and genetic divergence among 39 populations across the distribution of the two subspecies to assess whether they are conspecific and, if not, how many species might be involved. Cuticle coloration, pedipalp chela digital carina condition, and selected measurements were recorded. Sixty burrows were excavated and examined for burrow structure and depth. A multilocus dataset comprising concatenated fragments of one nuclear (28S rDNA) and three mitochondrial (12S rDNA, 16S rDNA, Cytochrome c Oxidase Subunit I) loci, totaling ca. 2400 base-pairs, was produced for 41 individuals, and a single-locus dataset comprising 658 base-pairs of the COI locus for 156 individuals. Despite overlapping ranges in morphometric characters of pedipalp chela shape, the putative subspecies were easily distinguished by cuticle coloration and condition of the pedipalp chela digital carina, and were also found to differ significantly in burrow architecture and depth. Phylogeographical analyses of the COI and multilocus datasets recovered seven distinct clades. Separate analyses of mitochondrial sequences, and combined analyses of mitochondrial and nuclear sequences support most clades. The two major clades corresponded with the geographical distributions of S. m. fuscus and S. m. palmatus in the region. Specimens from these clades were genetically distinct, and exhibited different burrow structure in geographically-proximate localities, suggesting reproductive isolation. The palmatus clade included two distinct subclades of specimens from localities adjacent to the Dead Sea. Three other clades, comprising specimens from the most northeastern localities, were tentatively assigned to subspecies previously recorded in neighboring Jordan and Syria. The morphological, behavioral and genetic evidence supports previous suggestions that Scorpio maurus is a species complex and justifies the following taxonomic emendations: Scorpio fuscus (Ehrenberg, 1829), stat. nov.; Scorpio kruglovi Birula, 1910, stat. nov.; Scorpio palmatus (Ehrenberg, 1828), stat. nov.; Scorpio propinquus (Simon, 1872), stat. nov.

Testing the morphological constraint hypothesis of tail length in the sexually dimorphic Cerastes vipera and new perspectives.

The morphological constraint hypothesis (MCH) states that, in snakes, males typically have relatively longer tails than females to accommodate the hemipenes and retractor muscles. To date, most studies testing the MCH have been interspecific and results have been equivocal. We tested the MCH intraspecifically on Cerastes vipera, a species with a relatively short tail and suitable for testing the MCH. The relative tail length and length of the hemipenes pocket in Cerastes vipera were measured in preserved museum-maintained males (n = 35) and in free-ranging males and females (n = 277). Males exhibited relatively longer tails than females, which was explained fully by the length of the hemipenes pocket. The relatively short tail of C. vipera presents a constraint to the reproductive structures in males, as the length of the hemipenes pocket occupies a greater proportion in shorter- than longer-tailed individuals. This is the first report presenting these intraspecific findings in support of the MCH. Whether these relations are widespread among snake families, within Viperidae, or specifically within C. vipera warrants further studies.

Replacement of fangs in a free-ranging desert viperid, Cerastes vipera.

Venomous viperid snakes possess relatively large and fragile hollow fangs that are an integral part of the envenomation apparatus for predation. We hypothesized that fangs serve like disposable needles and predicted a high loss rate and, hence, high replacement rate in free-ranging snakes. Snakes also possess smaller rear teeth that aid in gripping and swallowing the prey. We reasoned that these teeth are less delicate than fangs and predicted that their loss would be at a slower rate than fangs. To test our predictions, we analyzed fecal samples of free-ranging Saharan sand vipers, Cerastes vipera, in the Northern Negev desert, Israel. Close to 25% of fecal samples contained fangs, averaging more than one fang per sample and, consequently, our first prediction was supported. We estimated that fangs are replaced each fourth predation, and that replacement rate under natural conditions is at a high rate of approximately every twenty days. Fecal samples contained rear teeth at the same proportion as fangs, which indicated that the rapid replacement of teeth was not limited only to fangs and, therefore, our second prediction was not supported. These findings reflect the importance of both front fangs and rear teeth in the hunting of prey in free-ranging C. vipera. This is the first quantitative report of fang and rear teeth loss in a free-ranging viperid which is based on their recovery in feces; and we believe that similar high rates of loss occur in other viperid species.

Relative tail length correlates with body condition in male but not in female crowned leafnose snakes (Lytorhynchus diadema).

Reproductive success is the ultimate measure of individual quality; however, it is difficult to determine in free-living animals. Therefore, indirect measures that are related to reproduction are generally employed. In snakes, males typically possess longer tails than females and this sexual size dimorphism in tail length (TL) has generally been attributed to the importance of the tail in mating and reproduction. Thus, intra-sexual differences in tail length, specifically within males, were hypothesized to reflect individual quality. We used a body condition index (BCI) as a measure of quality in snakes and predicted that tail length would be correlated with BCI in males. We tested our prediction by determining BCI in the free-ranging adult male and female crowned leafnose snake (Lytorhynchus diadema), a colubrid species that inhabits mainly desert sand dunes. The relative TL was correlated positively and significantly to BCI in males (F1,131 = 11.05; r2adj = 0.07; P < 0.01) but not in females, thus supporting our prediction. This is the first time that the relationship between TL and body condition was tested in a free-ranging species. In addition, sexual size dimorphism of TL increased intra-specifically with body size, which was also found in interspecific analyses following Rensch's rule.

Sexual dichromatisation and sexual differences in hunting behavior and dietary intake in a free-ranging small viperid snake, Cerastes vipera.

The tip of the tail in female Cerastes vipera, a small viperid snake, is black and conspicuous, whereas that of the male is not. We tested the hypothesis, albeit indirectly, that this sexual dimorphic chromatisation is related to caudal luring, a feeding mimicry hunting strategy. C. vipera can hunt nocturnally-active lizards only via sit-and-wait ambush and, consequently, we predicted that females would use caudal luring more often than males and that the proportion of nocturnal prey items in the diet of females would be higher than in males. Our hypothesis was supported as: 1) only females demonstrated caudal luring towards nocturnally-active lizards and more than 85% did so, whereas none of the males demonstrated such behavior; and 2) females consumed a significantly higher proportion (15/40 vs 4/27) of nocturnally-active lizards than did males. We concluded that sexual dichromatisation in C. vipera is associated with hunting strategy that results in different hunting behavior and different dietary intake between sexes. These novel findings: 1) provide a functional explanation for the black tail of female C. vipera; and 2) suggest different evolutionary driving forces between sexes and, consequently, different ecological impacts of male and female C. vipera on lizard populations.

Seasonal biotic and abiotic factors affecting hunting strategy in free-living Saharan sand vipers, Cerastes vipera.

Sit-and-wait ambushing and active hunting are two strategies used by predators to capture prey. In snakes, hunting strategy is conserved phylogenetically; most species employ only one strategy. Active hunters encounter and capture more prey but invest more energy in hunting and have higher risks of being predated. This trade-off is important to small predators. The small Cerastes vipera employs both modes of hunting, which is unlike most viperids which use only sit-and wait ambushing. This species hibernates in October and emerges in April. Energy intake should be high prior to hibernation to overcome the non-feeding hibernation period and for reproduction on their emergence. We predicted that more individuals would hunt actively towards hibernation and an abiotic factor would trigger this response. Furthermore, since more energy is required for active hunting, we predicted that snakes in good body condition would use active hunting to a greater extent than snakes in poor body condition. To test our predictions, we tracked free-living snakes year round and determined their hunting strategy, estimated their body condition index (BCI), and calculated circannual parameters of day length as environmental cues known to affect animal behaviour. Two novel findings emerged in this study, namely, hunting strategy was affected significantly by 1) the circannual change in day length and 2) by BCI. The proportion of active hunters increased from 5% in April to over 30% in October and BCI of active foragers was higher than that of sit-and-wait foragers and, therefore, our predictions were supported. The entrainment between the proportion of active hunting and the abiotic factor is indicative of an adaptive function for choosing a hunting strategy. A trend was evident among life stages. When all life stages were present (September-October), the proportion of active foragers increased with age: 0.0% among neonates, 18.2% among juveniles and 31.4% among adults. We concluded that vulnerable small neonates used sit-and-wait ambush not only as a hunting strategy but also as a hiding technique.