Movements and habitat use in gravid and nongravid female garter snakes (Colubridae: Thamnophis)

Two dhole ( Cuon alpinus ) packs were monitored in Mudumalai Sanctuary, southern India, during 1989-93 to look at population dynamics, movement pattern, and foraging strategy and their inter-relationship with the maintenance of social groups. Pack size fluctuated substantially (4–18 and 4–25 in the two packs) owing to dispersal and demographic factors such as females not breeding in a given year. Both packs killed a much higher proportion of chital ( Axis axis ) and sambar ( Cervus unicolor ) fawns (2 and 54.2km² for the two packs) was not correlated with pack size. Pack movement from one resource patch (consisting of resting sites and aggregations of prey species) to another was not random or based on factors such as inter-patch distance or relative prey densities. There was no difference in mean residence time of the pack across the four resource patches; the pack moved across these in a sequential manner in one direction. We conclude that dholes live in groups not because of any advantages accruing from enhanced group sizes through increased per capita yield of food, but as a consequence of the dispersion of resources.     

Adaptive maternal cannibalism in snakes (Epicrates cenchria maurus, Boidae)

In many animal species, mothers that produce stillborn offspring or undeveloped ova immediately ingest them. This cannibalism has been interpreted in two ways: (1) as a form of parental care (to prevent disease spreading to healthy littermates, and eliminate predator-attracting scent cues); or (2) to recycle otherwise wasted energy, thus facilitating maternal recovery. Our experimental study on captive Colombian rainbow boas ( Epicrates cenchria maurus ) provides the first quantitative support for this latter hypothesis. We show that by eating their non-viable offspring and undeveloped eggs, female boas can rapidly recycle a significant component of their otherwise wasted reproductive investment. Female boas that ingested non-viable progeny equivalent to half their litter mass exhibited rapid recovery of dorsal musculature and hence were able to constrict prey items more forcefully than were unfed females when tested 2 weeks after parturition. The consequent enhancement of constricting ability may influence maternal survival and foraging success. Thus, maternal cannibalism may be an effective tactic to avoid wasting reproductive resources, and to shorten the period required for recovery from pregnancy.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 767–774.     

Mechanisms and consequences of sexual conflict in garter snakes (Thamnophis sirtalis, Colubridae)

Red-sided garter snakes (Thamnophis sirtalis parietalis) courtand mate in early spring around large communal overwinteringdens in central Manitoba. Emerging females are immediately coveredby dozens or hundreds of vigorously-courting males, potentiallyimposing significant costs to the female. By manipulating numbersof courting males (both directly and by applying anticourtshippheromones), we quantified the degree to which female dispersalfrom the den is hindered by courtship. Courted females dispersedonly about half as fast as did solitary females. Blood lactatelevels were higher in mating than in courting or noncourtingsnakes of both sexes; the high levels of lactate in mating femalessupport the idea that courtship is physiologically stressfulto these animals, perhaps via constraints to female respiration.In arena trials, females that were exercised to exhaustion beforecourtship mated with smaller males than did control females.The spatial distribution of snakes around the den exhibits substantialheterogeneity, with densities often varying markedly betweenadjacent areas. Arena trials mimicking this heterogeneity showedthat unmated females avoided parts of the enclosure containingscent cues from males. Our data support the hypothesis thatcourtship in T. s. parietalis confers significant costs to females,and that female behaviors have evolved to reduce those costs.     

Feeding ecology of North American gopher snakes (Pituophis catenifer, Colubridae)

Studies of food relations are important to our understanding of ecology at the individual, population and community levels. Detailed documentation of the diet of large‐bodied, widespread snakes allows us to assess size‐dependent and geographical variation in feeding preferences of gape‐limited predators. Furthermore, with knowledge of the food habits of sympatric taxa we can explore possible causes of interspecific differences in trophic niches. The feeding ecology of the North American gopher snake, Pituophis catenifer, was studied based on the stomach contents of more than 2600 preserved and free‐ranging specimens, and published and unpublished dietary records. Of 1066 items, mammals (797, 74.8%), birds (86, 8.1%), bird eggs (127, 11.9%), and lizards (35, 3.3%) were the most frequently eaten prey. Gopher snakes fed upon subterranean, nocturnal and diurnal prey. The serpents are primarily diurnal, but can also be active at night. Therefore, gopher snakes captured their victims by actively searching underground tunnel systems, retreat places and perching sites during the day, or by pursuing them or seizing them while they rested at night. Gopher snakes of all sizes preyed on mammals, but only individuals larger than 40 and 42 cm in snout–vent length took bird eggs and birds, respectively, possibly due to gape constraints in smaller serpents. Specimens that ate lizards were smaller than those that consumed mammals or birds. Gopher snakes raided nests regularly, as evidenced by the high frequency of nestling mammals and birds and avian eggs eaten. Most (332) P. catenifer contained single prey, but 95 animals contained 2–35 items. Of the 321 items for which direction of ingestion was determined, 284 (88.5%) were swallowed head‐first, 35 (10.9%) were ingested tail‐first, and two (0.6%) were taken sideways. Heavier gopher snakes took heavier prey, but heavier serpents ingested prey with smaller mass relative to snake mass, evidence that the lower limit of prey mass did not increase with snake mass. Specimens from the California Province and Arid Deserts (i.e. Mojave, Sonoran and Chihuahuan Deserts) took the largest proportion of lizards, whereas individuals from the Great Basin Desert consumed a higher percentage of mammals than serpents from other areas, and P. catenifer from the Great Plains ate a greater proportion of bird eggs. Differences in prey availability among biogeographical regions and unusual circumstances of particular gopher snake populations may account for these patterns. Gopher snakes have proportionally longer heads than broadly sympatric Rhinocheilus lecontei (long‐nosed snake), Charina bottae (rubber boa) and Lampropeltis zonata (California mountain kingsnake), which perhaps explains why, contrary to the case in P. catenifer, the smaller size classes of those three species do not eat mammals. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77 , 165–183.     

How snakes eat snakes: the biomechanical challenges of ophiophagy for the California kingsnake, Lampropeltis getula californiae (Serpentes: Colubridae)

In this study we investigated how ophiophagous snakes are able to ingest prey snakes that equal or exceed their own length. We used X-ray video, standard video, dissection, and still X-rays to document the process of ophiophagy in kingsnakes (Lampropeltis getula) feeding on corn snakes (Elaphe guttata). Most kingsnakes readily accepted the prey snakes, subdued them by constriction, and swallowed them head first. In agreement with previous observations of ophiophagy, we found that the predator snake forces the vertebral column of the prey snake to bend into waves. These waves shorten the prey's body axis and allow it to fit inside the gastrointestinal (GI) tract and body cavity of the predator. Dissection of a kingsnake immediately following ingestion revealed extensive longitudinal stretching of the anterior portion of the GI tract (oesophagus and stomach), and no visible incursion of the prey into the intestine. X-ray video of ingestion showed that the primary mechanism of prey transport was the pterygoid walk, with some contribution from concertina-like compression and extension cycles of the predator's vertebral column in two out of three observations. Complete digestion was observed in only one individual, as others regurgitated before digestion was finished. X-ray stills taken every 4 days following ingestion revealed that the corn snakes were about half digested within the first 4 days, and digestion was complete within 15 days.     

Descriptions of six new species of Caryospora (Apicomplexa: Eimeriidae) from Guatemalan snakes (Serpentes: Colubridae and Viperidae)

One hundred and seventy snakes were collected in Guatemala and examined for coccidia. Of these, 8 individuals representing 6 host species were positive for Caryospora spp., 6 of which are described as new species. Sporulated oocysts of Caryospora bothriechis n. sp. from Bothriechis aurifer are spheroidal to subspheroidal, 12.7 x 12.5 (12-14 x 12-13) microm, with a length/width (L/W) ratio of 1.0; they lack a micropyle (M) or oocyst residuum (OR), but 1 large polar granule (PG) is usually present. Sporocysts are ovoidal, 9.0-7.5 (8-10 x 7-8) microm, and have a L/W ratio of 1.2, and a Stieda body (SB) and sporocyst residuum (SR). Oocysts of Caryospora coniophanis n. sp. from Coniophanes imperialis are spheroidal to subspheroidal, 18.8 x 18.1 (17-20.5 x 16-20) microm, with a L/W ratio of 1.0; they lack a M and OR, but 1 large PG is usually present. Sporocysts are ovoidal, 13.2 x 9.4 (12-15 x 8-10) microm with a L/W ratio of 1.4, and a SB, substieda body (SSB), and SR. Oocysts of Caryospora conophae n. sp. from Conophis lineatus are spheroid to subspheroidal, 20.4 x 19.5 (17-26 x 17-25) microm, with a L/W ratio of 1.0; they lack a M and OR, but 1 large PG is usually present. Sporocysts are ovoidal, 13.1 x 9.8 (11-15 x 8-11) microm with a L/W ratio of 1.3 and a SB, SSB, and SR. Oocysts of Caryospora guatemalensis n. sp. from Lampropeltis triangulum are spheroidal to subspheroidal, 23.9 x 23.2 (20-27 x 20-26) microm, with a L/W ratio of 1.0; they lack a M and OR, but 1 large PG is usually present. Sporocysts are ovoidal, 14.4 x 10.6 (13-18 x 9-13) microm, with a L/W ratio of 1.4 and a SB, SSB, and SR. Oocysts of Caryospora mayorum n. sp. from Conophis lineatus are spheroidal to subspheroidal, 25.6 x 24.4 (24-27 x 24-25) microm, with a L/W ratio of 1.0; they lack a M and OR, but 1 large PG is usually present. Sporocysts are ovoidal, 16.3 x 11.9 (16-18 x 11-13) microm, with a L/W ratio of 1.4 and a SB, SSB, and SR. Oocysts of Caryospora zacapensis n. sp. from Masticophis mentovarius are spheroidal to subspheroidal, 22.5 x 21.8 (19-25 x 18-25) microm, with a L/W ratio of 1.0; they lack a M and OR, but 1 large PG is usually present. Sporocysts are ovoidal, 14.6 x 11.4 (11-16 x 10-13) microm, with a L/W ratio of 1.3 and a SB, SSB, and SR.     

Differences in endothelin receptor types in the vasculature of Bothrops jararaca (Viperidae) and Oxyrhopus guibei (Colubridae) snakes

Endothelins (ETs) are vasoactive peptides evolutionary well conserved that exert their effects through two specific receptors (ET(A) and ET(B)) widely distributed in all vertebrates. In snakes, the presence and function of endothelins and their receptors are still scarcely described. We have recently demonstrated the presence of ET(A) and ET(B2) receptors in the snake Bothrops jararaca (Bj). In the present work we showed that distinctively from Bj, the vascular contraction induced by endothelin in Oxyrhopus guibei (Og) snake is mediated only by ET(A) receptors. Selective ET(B) agonists (SRTX-c and IRL(1620)) and antagonists (IRL(1038) and BQ(788)) were ineffective in Og preparations of isolated aorta. We also showed that ET-1 response on Og arterial blood pressure was monophasic hypertensive as opposed to biphasic (hypotension followed by hypertension) in Bj. Furthermore, we characterized the relaxing properties of endothelin receptor ET(B1) in pre-contracted aorta preparations. We showed that IRL(1620) induced relaxation of pre-contracted Bj aorta but was ineffective in relaxing Og preparations. IRL(1620) relaxing effect on Bj aorta was abolished by l-NAME, indicating involvement of NO release, and was reduced by selective ET(B) antagonists. Our findings suggest that Og snake has a more primitive spectrum of ET receptors (only ET(A) receptor) than Bj (presence of ET(A), ET(B1) and ET(B2) receptors).     

Morphological specializations of the yolk sac for yolk processing in embryonic corn snakes (Pantherophis guttatus: Colubridae)

Non‐avian reptiles commonly are assumed to be like birds in their overall patterns of development. However, colubrid corn snakes (Pantherophis guttatus ) have mechanisms of yolk cellularization and processing that are entirely different from the avian pattern. In birds, a vascular “yolk sac” surrounds and digests the liquid yolk. In contrast, in corn snakes, the yolk material is converted into vascularized cords of yolk‐filled cells. In this study, we used stereomicroscopy, histology, and scanning electron microscopy to analyze this unusual developmental pattern in corn snakes. Our observations reveal that the yolk sac cavity is invaded by endodermal cells that proliferate, absorb yolk spheres, and form aggregates of interconnected cells within the liquid yolk mass. As development proceeds, small blood vessels arise from the yolk sac omphalopleure, penetrate into the yolk mass, and become tightly encased in the endodermal cells. The entire vitellus ultimately becomes converted into a mass of vascularized, “spaghetti‐like” strands of yolk‐laden cells. The resulting arrangement allows yolk to be digested intracellularly and yolk products to be transported to the developing embryo. Indirect evidence for this pattern in other species raises the possibility that it is ancestral for squamates and quite possibly Reptilia in general.     

Molecular systematics of New World lampropeltinine snakes (Colubridae): implications for biogeography and evolution of food habits

We used mitochondrial gene sequences to infer phylogenetic relationships among North American snakes of the colubrid tribe Lampropeltini (Arizona, Bogertophis, Cemophora , New World Elaphe, Lampropellis, Pituophis, Rhinocheilus, Senticolis, Stilosoma) , and assessed the implications of our findings for the biogeography and evolution of food habits among these serpents. The maximum likelihood phylogeny identified Rhinocheilus as the sister taxon to all other lampropeltinines, and supported the monophyly of Lampropeltis (including Stilosoma) , New World Elaphe , and Pituophis , but not that of Bogertophis. This phylogeny also suggested a sister group relationship between Cemophora and Lampropeltis , and between New World Elaphe and Pituophis , and strongly supported that Sentkolis belongs within Lampropeltini, thus contradicting previous suggestions that Senticolis is not a lampropeltinine. Using a method for approximating ancestral areas of clades, we determined that western North America was most likely the ancestral area of lampropeltinines. Our survey of published studies, combined with unpublished data, indicated that lampropeltinines as a group feed mainly on mammals, less frequently on lizards, birds, and bird eggs, and only rarely on squamate eggs, snakes, anurans, and insects. Some individual species indeed emphasize mammals in their diets, but others most frequently eat lizards, squamate eggs, bird eggs, or snakes, whereas others take two prey types with similar frequency. Our reconstruction of the evolution of food habits among lampropeltinines suggests that a diet emphasizing lizards is ancestral, and therefore diets that mosdy consist of mammals, squamate and bird eggs, and snakes are derived within the clade. In at least some species, smaller individuals prey mostly on lizards and larger ones add mammals to their diets.     

Understanding the formation of ancient intertropical disjunct distributions using Asian and Neotropical hinged-teeth snakes (Sibynophis and Scaphiodontophis: Serpentes: Colubridae)

Numerous taxa show ancient intertropical disjunct distributions. Many can be explained by well-known processes of historical vicariance, such as the breakup of Gondwanaland. Others, such as Asian–Neotropical divergences are not as well understood. To clarify the phylogenetic position and understand biogeographic and temporal origins of the geographically disjunct and morphologically unique genera of hinged-teeth snakes, Scaphiodontophis (n = 1) and Sibynophis (n = 9; Colubridae), we inferred a time-calibrated phylogeny with additional 107 taxa representing the superfamily Colubroidea using four genes (c-mos, cyt-b, ND2, RAG-1; 3085 bp). We used this tree to estimate ancestral areas for the group. The results show that Scaphiodontophis is sister to Sibynophis, both originated in the late Eocene/Oligocene in Asia and likely dispersed through Beringia to the New World, but unlike other snake groups left no extant species in temperate North America. Current recognition of Scaphiodontophiinae renders Colubrinae paraphyletic, and we resurrect the previously named subfamily Sibynophiinae to encompass both genera and use the tribes Sibynophiini (Sibynophis) and Scaphiodontophiini (Scaphiodontophis) to highlight the geographically distinct areas occupied by these taxa. These results suggest that intercontinental dispersal with extinction in intermediate areas can explain puzzling patterns of ancient intertropical disjunct distributions.     

Not so rare snakes: a revision of the Geophis sieboldi group (Colubridae: Dipsadinae) in lower Central America and Colombia

The Geophis sieboldi species group is composed of 16 currently recognized species distributed from Mexico to Colombia. Within this group, snakes of populations referred to the Geophis brachycephalus complex of lower Central America and Colombia display a remarkable polychromatism and the systematic status of these and other populations is problematic. The present study provides an analysis, including multivariate techniques, of variation in scalation, coloration, relative tail length and hemipenes to clarify the specific allocation of the populations belonging to this clade. Our results confirm the validity of three previously described taxa, namely G. brachycephalus , G. nigroalbus and G. talamancae , with reassignments of several populations previously referred to G. brachycephalus . In addition we recognize as a new species a suite of western Panama Geophis previously of uncertain status. We further provide a review of all other members of the G. sieboldi group in lower Central America and Colombia based on material obtained since the last revision of the group. Basic synonymies, diagnostics and known distributions are included for the treated taxa. Dietary guild, possible venomous coral snake mimicry and distributional anomalies for the group are discussed.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 561–599.     

Body size as a primary determinant of ecomorphological diversification and the evolution of mimicry in the lampropeltinine snakes (Serpentes: Colubridae)

Evolutionary correlations between functionally related character suites are expected as a consequence of coadaptation due to physiological relationships between traits. However, significant correlations may also exist between putatively unrelated characters due to shared relationships between those traits and underlying variables, such as body size. Although such patterns are often dismissed as simple body size scaling, this presumption may overlook important evolutionary patterns of diversification. If body size is the primary determinant of potential diversity in multiple unrelated characters, the observed differentiation of species may be governed by variability in body size, and any biotic or abiotic constraints on the diversification thereof. Here, we demonstrate that traits related to both predatory specialization (gape and diet preference) and predatory avoidance (the development of Batesian mimicry) are phylogenetically correlated in the North American snake tribe Lampropeltini. This is apparently due to shared relationships between those traits and adult body size, suggesting that size is the primary determinant of ecomorphological differentiation in the lampropeltinines. Diversification in body size is apparently not linked to climatic or environmental factors, and may have been driven by interspecific interactions such as competition. Additionally, we find the presence of a ‘key zone’ for the development of both rattle‐ and coral snake mimicry; only small snakes feeding primarily on ectothermic prey develop mimetic colour patterns, in or near the range of venomous model species.     

The biogeographical history of Asian keelback snakes of the genus Hebius (Squamata: Colubridae: Natricinae) in the Ryukyu Archipelago,Japan

The islands of the central and southern parts of the Ryukyu Archipelago and Taiwan accommodate suites of allopatric endemics for many terrestrial vertebrates. These subregions, which were isolated as three independent islands since the Late Miocene period, became connected to the continent by the Early Pleistocene land bridge (EPL). Many endemics are considered to have emerged through vicariant events accompanied with the collapse of EPL, but this is not well corroborated. We conducted divergence dating analyses for samples of four allopatric keelback snakes from 20 islands using a published timeframe for natricine snakes. Phylogenetic analyses recognised the existence of eight subclades for the four species. Four of the five subclades in the Central Ryukyus have emerged after the initial isolation of three subregions but before the EPL collapse, indicating iterative divergences within the super‐island and stagnation of diverged entities. In contrast, the Southern Ryukyus was subjected to multiple incursions from both Central Ryukyus and Taiwan, irrespective of the EPL connection. Consequently, among all seven nodes, only two were consistent with the assumption of vicariance. These findings suggest that dispersals and subsequent nonvicariant divergences have contributed to the formation of the current phylogeographical pattern in this continental island chain.     

Species boundaries in Malagasy snakes of the genus Madagascarophis (Serpentes: Colubridae sensu lato) assessed by nuclear and mitochondrial markers

We studied mitochondrial divergence in 27 individuals of colubrid snakes of the genus Madagascarophis Mertens from most of its distribution area in Madagascar. Combined analyses of 16S rRNA and cytochrome b sequences identified six major clades which only partly agreed with previously proposed classifications. Analysis of nuclear DNA sequences of the c-mos gene as well as of ISSR fingerprints revealed consistent differences only among three clades which we consider as distinct species: a widespread Madagascarophis colubrinus (Schlegel), with M. citrinus (Boettger) as a junior synonym, a southern M. meridionalis Domergue, and a presumably undescribed species from the extreme north of Madagascar. The species M. ocellatus Domergue was not available for our study. Within M. colubrinus there are two populations from the north-west, each showing two divergent haplotypes with pairwise divergences of up to 5.2% in the cytochrome b gene. Maximum divergence in this gene within M. colubrinus was 7.1%. These high values emphasise that caution needs to be applied before genetic distance values are used for species delimitation. Phylogeographically, most of the genetic variation in M. colubrinus is found in northern Madagascar, indicating that the species might have originated in this region. Later one haplotype clade colonised western and eastern Madagascar, with a putative secondary introgression into north-western populations.     

The strange occurrence of male cannibalism on adult females in snakes

Cannibalism is widespread in the animal kingdom. The most common, well-known cases are found in some arthropod systems where the larger females occasionally consume the smaller males before, during, or after mating (i.e., sexual cannibalism). The killing and consumption of females by males are, on the other hand, seemingly much rarer and theoretically more difficult to explain, at least among adults, because the consumption of females would result in the loss of potential mates for males. Here, we use observations submitted to a community science reporting system to describe three instances of adult males feeding on adult females in the Montpellier snake, Malpolon monspessulanus, in the south of France. We discuss our observations in the theoretical framework of cannibalism to provide potential explanations for these observations and briefly review empirical findings of cannibalism in vertebrates, which confirms that this sex-reversed form of sexual cannibalism is rare in nature.     

Effects of nest temperature and moisture on phenotypic traits of hatchling snakes (Tropidonophis mairii, Colubridae) from tropical Australia

Previous research on developmentally plastic responses by reptile embryos has paid relatively little attention to tropical species, or to possible interactions between the effects of thermal and hydric regimes. In the present study, eggs of keelback snakes ( Tropidonophis mairii ), from a tropical area with strong temporal and spatial variation in soil temperatures and moisture levels, were incubated. The phenotypic traits of hatchling snakes (body size, shape, muscular strength) were affected by moisture content of the incubation medium (vermiculite plus 100% vs. 50% water by mass), by mean incubation temperatures (25.7 vs. 27.9 °C) and by diel thermal variation (diel range 6.0 vs. 8.4 °C). Interactions between these factors were negligible. Cooler, more thermostable, moister conditions resulted in larger offspring, a trait under strong selection in this population. Thermal and hydric conditions covary in potential nest-sites (e.g. deeper nests are more thermostable as well as moister). This covariation may influence the evolution of reaction norms for embryogenesis. For example, if moister nests enhance offspring fitness and are cooler, then selection will favour the ability to develop in cool as well as moist conditions. Thus, the evolution of optimal incubation conditions with respect to one variable (e.g. temperature) may be driven by patterns of association with another variable (e.g. soil moisture) among natural nest-sites. Perhaps for this reason, the thermal optimum for incubation is surprisingly low in this tropical species.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 159–168.     

Food habits, growth rates, and reproductive biology of grass snakes, Natrix natrix (Colubridae) in the Italian Alps

A five-year mark-recapture study at Sella Nevea, a montane (1100 m a.s.1.) site in the Carnic Alps, provided information on diets, growth rates, and reproductive output in an Italian population of the wide-ranging grass snake, Natrix natrix. Our snakes resembled a previously-studied population in lowland Sweden in terms of body size at sexual maturation in females (70 cm) and mean adult female body length (82 cm). However, growth rates were lower in our population, and sexual maturation was delayed (6–8 years, versus 4–5 years in Sweden), perhaps because of the cool climate and relatively brief growing period each year. Females produced a single clutch of 4–24 eggs in late July each year. Larger females produced larger clutches, but clutch size relative to maternal size was lower than in Swedish grass snakes. Hatchling sizes and Relative Clutch Masses (RCMs) did not shift with increasing female size. RCMs may provide a useful index of 'costs of reproduction' in this population, because females with high RCMs were very emaciated after oviposition, and hence may experience a greater risk of mortality, as well as a high energy expenditure. Prolonged incubation gave rise to longer, thinner hatchlings, but the low environmental temperatures at the study site may favour early hatching (and hence, result in a shorter fatter hatchling emerging from the egg, with more of its energy stores unused). Compared to sympatric viviparous snakes ( Coronella austriaca and Vipera berus ), the oviparous grass snakes can achieve a much higher reproductive output owing to a larger clutch size and more frequent reproduction (annual, rather than biennial or triennial). The abundant prey resource used by grass snakes (amphibians) may also enable them to recoup energy more rapidly after reproduction; dietary composition shifts ontogenetically in both sexes, with the largest prey (mice and adult toads) taken primarily by large female snakes.     

Molecular systematics of neotropical cat-eyed snakes: a test of the monophyly of Leptodeirini (Colubridae: Dipsadinae) with implications for character evolution and biogeography

The Leptodeirini has been presumed to be a monophyletic assemblage based on albumin immunological data and morphology and consisted of neotropical cat-eyed snakes ( Leptodeira ), blunt-headed vine snakes ( Imantodes ), nightsnakes ( Eridiphas , Hypsiglena , and Pseudoleptodeira ), and the cloud forest snake ( Cryophis ). In the present study, approximately 1.4 kb of mitochondrial DNA sequence data (from cob and nad4 ) were collected to test the monophyly of the Leptodeirini. These data were analysed using maximum parsimony, maximum likelihood, and Bayesian methods. None of the results supported the monophyly of the Leptodeirini. There was strong support for a clade containing Imantodes and Leptodeira , and another clade containing the nightsnakes, with the latter placed closer to Cryophis and other dipsadine genera ( Sibon , Dipsas , and Atractus ). This partial reassessment of the Dipsadinae infers the group to have an ancestral condition of being rear-fanged, mildly venomous, and feeding on ectothermal vertebrates, with a more derived radiation that has lost the rear-fanged, venomous condition and has a diet specialized on invertebrates. A brief discussion on the biogeography of the dipsadines shows the group to be much older (Palaeocene), consistent with earlier views, as opposed to a more recent (Miocene) hypothesis, as has been recently suggested.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 483–500.