Do drivers intentionally target wildlife on roads?

Despite frequent reliance on surveys to document public attitudes towards conservation issues (such as invasive‐species control), only rarely do researchers assess the validity of statements made by the public in response to such surveys. Therefore, how well responses match actual behaviour remains an open question. We conducted a survey asking drivers if they had seen and/or run over (intentionally or not) snakes, native frogs or invasive cane toads (Rhinella marina) on roads in the Northern Territory of Australia. To compare actual driver behaviour to the survey responses, we also carried out field experiments where we quantified the rates at which model snakes, frogs and toads (and controls) were run over on a rural highway. Our results show a discrepancy between survey responses and driver behaviour: for example, 25% of the people we surveyed indicated that they intentionally run over cane toads, yet field experiments showed that model toads were run over no more frequently than expected by chance, or than any other type of model.     

Effects of size, sex, temperature and condition on activity metabolism and defence behaviour of the viperine snake, Natrix maura

The activity metabolism (aerobic and anaerobic), performance (burst speed and endurance) and defence behaviour (static or active) of the snake Natrix maura were investigated in relation to size, sex, temperature and fasting. Conditions which reduced performance (low temperature, fasting) promoted the use of static defensive behaviour. Aerobic scope for activity decreased with increasing size, but this was compensated for by increased capacity for anaerobic metabolism. Burst speed (as lengths time^-1) was slightly lower in large snakes; endurance was much lower in small snakes. Increased use of static defence by small snakes is related to their predators, which are likely to take occasional snakes in a diet of worms; a balled, still snake may not provide a stimulus for feeding. Large snakes have predators which feed on vertebrates and may recognize them as food in any position, and so they are better off trying to escape.
Aerobic scope was not maximal at the mean selected body temperature (25 °C), but increased right up to near lethal temperatures (> 35 °C). Anaerobic metabolism (lactate production) was less temperature-sensitive, and accounted for about 80 and 50% of the energy produced during 30 and 5 min of maximal activity, respectively. Resting and active oxygen consumption were weight-dependent, VO₂∝ W^0·75, the same exponent as that of interspecific comparisons. This exponent was independent of temperature and acclimation state. Acclimation to a variable or constant temperature regime had no effect on maximal oxygen consumption, but there was a significant effect on resting metabolic rate, possibly attributable to energy conservation when inactive. There were no differences in activity metabolism, performance or defence behaviour between similar-sized snakes of different sexes.     

Sexual motivation suppresses paternal behaviour of male gerbils during their mates' postpartum oestrus

Adult male Mongolian gerbils, Meriones unguiculatus, avoid contact with their young on the day that the young are born. However, on succeeding days, fathers spend nearly as much time in contact with their offspring as do mothers. We undertook a series of studies to investigate the causes of the day-to-day change in male parental behaviour. In experiment 1, we tested males' response to pups before, after and on the day of their mates' parturition, and found that males were more parental both before and after the day of birth of their young than on that day. In experiment 2, we compared the parental behaviour of males paired either with intact or with ovariectomized dams (which do not come into postpartum oestrus) and found that males that were mated to intact females were less attentive to pups on the day of their birth than were the males mated to ovariectomized females. In experiment 3, we compared the parental responses of castrated and intact males to newborn pups and found that castrated males were more parental than intact males. In experiment 4, we compared the parental responses of males that were exposed to postpartum oestrous females but prevented from mating for 24 h. Extending the period of male sexual arousal to 24 h inhibited paternal responsiveness to neonates for 24 h. We interpret these results as being consistent with the hypothesis that on the day of birth of a litter, a male's parental behaviour is inhibited by the motivation to mate during his partner's postpartum oestrus.     

Dietary Correlates of the Origin and Radiation of Snakes

Stomach analyses of living families and of a fossil containingprey were used to address possible dietary correlates of thehistory of snakes. Aniliids, morphologically primitive amongliving snakes, feed on relatively heavy, elongate vertebrates.Large aniliids eat larger prey than do small individuals but,as in advanced snakes, they also take small items. Living boids,structurally intermediate between aniliids and advanced snakes,feed on relatively heavy prey of a much greater variety of shapesthan do aniliids. An Eocene fossil that might be a boid containsa relatively large crocodilian in its gut. These findings, previousstudies, and morphological considerations suggest that veryearly snakes used constriction and powerful jaws to feed onelongate, heavy prey. This would have permitted a shift fromfeeding often on small items to feeding rarely on heavy items,without initially requiring major changes in jaw structure relativeto a lizard-like ancestor. Subsequent morphological changescould then have allowed boids to utilize a broad range of preytypes, including many of those currently eaten by advanced snakes.More recent dietary themes include the consumption of even heavierprey by highly venomous elapids and viperids, and frequent feedingon relatively small items by some other advanced snakes.     

Causes and consequences of aggregation by neonatal tiger snakes (Notechis scutatus,Elapidae)

Although snakes traditionally have been regarded as asocial animals, recent studies have revealed complex interactions among neonatal snakes and their mothers. We noticed frequent aggregation by captive neonatal Australian elapids (tiger snakes, Notechis scutatus), and conducted simple experiments to clarify the proximate causation of, and potential consequences of, aggregative behaviour. Litters of neonates exhibited statistically significant aggregation (clustering) in empty containers, especially if the test area was subjected to rapid cooling. Aggregation was most pronounced inside shelter‐sites, and familiar shelters (i.e. containing scent cues from the litter) attracted snakes more than did novel (unscented) shelters. Snakes in larger aggregations cooled more slowly (reflecting their higher combined mass and thus, thermal inertia) and higher body temperatures facilitated neonatal locomotor performance, retreat‐site location and anti‐predator tactics. Plausibly, aggregation in neonatal tiger snakes (and other reptiles) functions to retard cooling rates, with the result that the young snakes are better able to evade or repel attacks by predators.     

Parental defence on the reef: antipredator tactics of coral‐reef fishes against egg‐eating seasnakes

On coral reefs in New Caledonia, the eggs of demersal‐spawning fishes are consumed by turtle‐headed seasnakes (Emydocephalus annulatus). Fish repel nest‐raiding snakes by a series of tactics. We recorded 232 cases (involving 22 fish species) of antipredator behaviour towards snakes on a reef near Noumea. Blennies and gobies focused their attacks on snakes entering their nests, whereas damselfish (Pomacentridae) attacked passing snakes, as well as nest‐raiders (reflecting territorial defence). Biting the snake was the most common form of attack, although damselfish and blennies also slapped snakes with the tail, or (blennies only) plugged the nest entrance with the parent fish's body. Gobies rarely defended the nest, although they sometimes bit or threw sand at the snake. A snake was more likely to flee if it was attacked before it began feeding rather than after it found the eggs (82% versus 3% repelled) and if bitten on the head rather than the body (68% versus 53%). Tail‐slaps were not effective, although plugging the burrow and throwing sand often caused snakes to flee. These strong patterns reflect phylogenetic variation in fish behaviour (e.g. damselfish detect a snake approach sooner than do substrate‐dwelling blennies and gobies) coupled with intraspecific variation in snake diets. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 415–425.     

Contextual Flexibility: Reassessing the Effects of Prey Size and Status on Prey Restraint Behaviour of Macrostomate Snakes

Contextual flexibility in prey restraint behaviour has been documented in advanced snakes (Colubroidea), but the degree of flexibility for earlier snake lineages has been largely unstudied. We document the prey restraint behaviour of five snake species belonging to three early macrostomate lineages: Loxocemidae, Erycinae and Boidae. Species from these lineages were chosen for this study because they utilize similar prey resources but exhibit different ecological habits that may have important consequences on prey restraint behaviour. Snakes (n = 27) were studied in a systematic experimental design assessing the effects of mouse size (small and large) and status (live and dead) across a total of 216 feeding trials. Loxocemus and Erycine snakes were highly flexible in their prey restraint behaviour patterns and these varied across prey category. Individuals of Boa constrictor exhibited very little contextual flexibility in feeding behaviour, confirming earlier reports. Flexibility in prey restraint behaviour corresponded with loop application pattern, whether the snake bent laterally or ventrally when forming a loop around prey. Our study is the first to show that early macrostomate snakes exhibit flexible prey restraint behaviours. Thus, our results suggest that flexibility in predatory behaviour may be more widespread across snake taxa than previously thought and we offer hypotheses for the observed interspecific differences in snake feeding behaviour.     

The transvestite serpent: why do male garter snakes court (some) other males?

In large mating aggregations of red-sided garter snakes, Thamnophis sirtalis parietalis, in Manitoba, male courtship is directed not only to females, but also to other males with female-like skin lipids ('she-males'). We show that 'she-maleness' is an intrinsic property of a male rather than an artefact of lipid transfer from females, and that male-male courtship is very common in the field. She-males were distinctive in terms of appearance (they were heavier than other males and more often covered with mud), behaviour (they were inactive and rarely courted females) and performance (they were slow crawlers, ineffective courters and easily outcompeted by other males in mating trials). 'She-maleness' was not a characteristic of a particular subset of males, as envisaged in previous work; instead, it was a transitory phase that most (perhaps all) male snakes passed through soon after they first emerged from the winter den. Recently emerged males spent their first day or two relatively inactive, while restoring physiological functions (including locomotor performance and courtship ability). Experimental application of female skin lipids on to males dramatically decreased courtship levels of the recipient snakes. Thus, recently emerged males may derive two kinds of benefit from mimicking female skin lipids. First, female mimicry 'switches off' the male's own (energetically expensive) courtship at a time when that courtship would be unproductive. Second, it may disadvantage his rivals by distracting them from females, and increasing their energy expenditure. Copyright 2000 The Association for the Study of Animal Behaviour.     

Physiological and hormonal control of thermal depression in the tiger snake, Notechis scutatus

Plasma sodium concentrations in field-caught Western tiger snakes, Notechis scutatus, from semi-arid Carnac Island (CI) varied seasonally, with snakes exhibiting significant hypernatraemia during summer and normal concentrations following autumn rain. In contrast, field-caught tiger snakes from a perennial fresh-water swamp (Herdsman Lake, HL) exhibited no significant increase in plasma sodium concentrations during summer. Laboratory-induced hypernatraemia caused thermal depression in both populations; there was a weak negative relationship between plasma sodium concentration and temperature selection that was significant for CI snakes. Hypernatraemia significantly elevated circulating concentrations of the neuropeptide arginine vasotocin (AVT) in both CI and HL snakes. CI snakes injected with a physiological dosage of AVT also evidenced thermal depression. Despite the positive correlation between AVT and both plasma sodium concentration and osmolality for laboratory snakes, field samples from CI snakes indicate that circulating levels of AVT may be influenced more by plasma osmolality than sodium levels. The data suggest that, in CI snakes, chronic dehydration in the field leads to hypernatraemia which may lead to elevated levels of AVT if plasma osmolality also increases. This will in turn invoke a depression in thermal behaviour that may improve the water economy and survival of snakes on semi-arid CI. Although HL snakes do not experience seasonal dehydration, physiological changes away from the stable homeostatic state appear to prompt the same behavioural shifts, illustrating the intrinsic nature of the thermal behaviour in different populations of the same species of snake.     

DNA Double-Strand Breakage as an Endpoint to Examine Metal and Radionuclide Exposure Effects to Water Snakes on a Nuclear Industrial Site

This study examined metal levels (especially U and Ni) in the tail tissues of water snakes from contaminated (Tim's Branch) and reference areas on the Department of Energy's Savannah River Site (SRS). Home ranges of snakes were quantified to determine the ratio of the habitat that they use in relation to the contaminated areas to better estimate exposure Compared to conventional methods that do not. The exposure assessment indicated that water snakes in the contaminated areas could expect U exposure at 3–4 orders of magnitude greater than the Agency for Toxic Substances and Disease Registry's Minimum Risk Level (MRL) from ingestion of amphibians and fish. Ni and U, in addition to Se, Mn, and Cu, were related to increased DNA double-strand breakage (DDSB) in water snakes. We report burdens for each metal individually, but the results of the DDSB indicated that these metals did not behave independently, but as a suite. If we did not have a secondary endpoint (DDSB), we might have assumed from the exposure predictions and tissue burden analyses that U was the sole metal of concern to water snakes in Tim's Branch. These data also imply that these toxicants do not biomagnify at the spatial and temporal scale of this study.     

Snake modulates constriction in response to prey's heartbeat

Many species of snakes use constriction-the act of applying pressure via loops of their trunk-to subdue and kill their prey. Constriction is costly and snakes must therefore constrict their prey just long enough to ensure death. However, it remains unknown how snakes determine when their prey is dead. Here, we demonstrate that boas (Boa constrictor) have the remarkable ability to detect a heartbeat in their prey and, based on this signal, modify the pressure and duration of constriction accordingly. We monitored pressure generated by snakes as they struck and constricted warm cadaveric rats instrumented with a simulated heart. Snakes responded to the beating heart by constricting longer and with greater total pressure than when constricting rats with no heartbeat. When the heart was stopped midway through the constriction, snakes abandoned constriction shortly after the heartbeat ceased. Furthermore, snakes naive to live prey also responded to the simulated heart, suggesting that this behaviour is at least partly innate. These results are an example of how snakes integrate physiological cues from their prey to modulate a complex and ancient behavioural pattern.     

Antipredator behaviour of invasive geckos in response to chemical cues from snakes

Antipredator behaviours and the ability to appropriately assess predation risk contribute to increased fitness. Predator avoidance can be costly; however, so we expect prey to most strongly avoid predators that pose the greatest risk (i.e., prey should show threat sensitivity). For invasive species, effectively assessing the relative risk posed by predators in the new environment may help them establish in new environments. We examined the antipredator behaviour of introduced Asian house geckos, Hemidactylus frenatus (Schlegel), by determining if they avoided shelters scented with the chemical cues of native predatory snakes (spotted pythons, Antaresia maculosa [Peters]; brown tree snakes, Boiga irregularis [Merrem]; common tree snakes, Dendrelaphis punctulata [Grey]; and carpet pythons, Morelia spilota [Lacépède]). We also tested if Asian house geckos collected from vegetation vs. anthropogenic substrates (buildings) responded differently to the chemical cues of predatory snakes. Asian house geckos did not show a generalised antipredator response, that is, they did not respond to the chemical cues of all snakes in the same way. Asian house geckos avoided the chemical cues of carpet pythons more strongly than those of other snake species, providing some support for the threat‐sensitivity hypothesis. There was no difference in the antipredator behaviour of Asian house geckos collected from buildings vs. natural vegetation, suggesting that individuals that have invaded natural habitats have not changed their antipredator behaviour compared to urban individuals. Overall, we found some evidence indicating Asian house geckos are threat‐sensitive to some Australian predacious snakes.     

Why don't small snakes bask? Juvenile broad‐headed snakes trade thermal benefits for safety

Previous studies have suggested that most small Australian elapid snakes are nocturnal and rarely bask in the open because of the risk of predation by diurnal predatory birds. Because the physiology and behaviour of reptiles is temperature dependent, staying in refuges by day can entail high thermoregulatory costs, particularly for juveniles that must grow rapidly to maximise their chances of survival. We investigated whether the risk of predation deters juveniles of the endangered broad-headed snake ( Hoplocephalus bungaroides ) from basking, and if so, whether there are thermal costs associated with refuge use. To estimate avian attack rates on snakes, we placed 900 plasticine snake replicas in sunny locations and underneath small stones on three sandstone plateaus for 72 h. At the same time we quantified the thermal benefits of basking vs refuge use. On sunny days, juveniles could maintain preferred body temperatures for 4.7 h by basking but only for 2.0 h if they remained inside refuges. Our predation experiment showed that basking has high costs for juvenile snakes. Predators attacked a significantly higher proportion of exposed models (13.3%) than models under rocks (1.6%). Birds were the major predators of exposed models (75% of attacks), and avian predation did not vary across the landscape. By trading heat for safety, juvenile H. bungaroides decreased the potential time period that they could maintain preferred body temperatures by 57%. Thermal costs of refuge use may therefore contribute to the slow growth and late maturation of this endangered species. Our results support the hypothesis that nocturnal activity in elapid snakes has evolved to minimise the risk of avian predation.     

“Then they prayed,they did nothing else,they just prayed for the boy and he was well”: A qualitative investigation into the perceptions and behaviours surrounding snakebite and its management in rural communities of Kitui county,Kenya

IntroductionHuman-snake interactions are common in tropical regions where subsistence-farming and livestock-herding activities predominate alongside proliferation of snakes. Local beliefs and perceptions about snakes and snakebites influence human behaviour. Understanding these beliefs and perceptions can inform the development of resources to drive behaviour change and to minimise the risk of injury to both humans and snakes. This qualitative study, conducted between May and July 2019, sought to explore the beliefs and perceptions regarding snakes and snakebites, and methods of prevention and management among members of the community in Kitui County, Kenya.MethodsSemi-structured interviews were used to collect qualitative data from 23 participants, recruited using a stratified purposeful sampling strategy in four selected sub-counties of Kitui county. Interview data was anonymised and coded and a thematic analysis was conducted using NVivo 12.ResultsPeople from Kitui county mostly had negative perceptions about snakes. There was a generalised awareness of the need to prevent snakebite, predominantly through keeping snakes away from homes/compounds. However, implementation was limited by financial constraints. Participants also identified logistic and financial obstacles to early hospital presentation following a snakebite, and they expressed a strong preference of having their snakebites treated in a hospital over consulting traditional healers. There was a universal recognition of the benefit of early intervention with a specific appreciation of the utility of the black stone. Furthermore, the removal of a snake’s “teeth” was an expected treatment outcome for some community members, with the failure to do so perceived as causing poor wound healing or persistence of symptoms. Some religious groups held views which differed from most participants.ConclusionThere is a need to explore and clarify common misconceptions about snakes and first aid treatment of snakebites, encourage learning about the true nature of snakes, and highlight beneficial uses of snakes. A change in the epistemological conception of community education material by enhancing the value and use of local forms of knowledge, and the employment of art techniques to transmit this knowledge, could improve community perception and methods of snakebite prevention. Patient expectations should be appropriately managed by discussing possible outcomes, incorporating follow-up visits and addressing long-term complications of snakebites.     

Antipredatory function of head shape for vipers and their mimics

Most research into the adaptive significance of warning signals has focused on the colouration and patterns of prey animals. However, behaviour, odour and body shape can also have signal functions and thereby reduce predators' willingness to attack defended prey. European vipers all have a distinctive triangular head shape; and they are all venomous. Several non-venomous snakes, including the subfamily Natricinae, commonly flatten their heads (also known as head triangulation) when disturbed. The adaptive significance of this potential behavioural mimicry has never been investigated.We experimentally tested if the triangular head shape typical of vipers offers protection against predation. We compared the predation pressure of free-ranging predators on artificial snakes with triangular-shaped heads against the pressure on replicas with narrow heads. Snakes of both head types had either zigzag patterned bodies, typical of European vipers, or plain (patternless) bodies. Plain snakes with narrower Colubrid-like heads suffered significantly higher predation by raptors than snakes with triangular-shaped heads. Head shape did not, however, have an additive effect on survival in zigzag-patterned snakes, suggesting that species which differ from vipers in colouration and pattern would benefit most from behavioural mimicry. Our results demonstrate that the triangular head shape typical of vipers can act as a warning signal to predators. We suggest that head-shape mimicry may be a more common phenomenon among more diverse taxa than is currently recognised.     

Once Bitten,Twice Shy: Does Previous Experience Influence Behavioural Decisions of Snakes in Encounters with Predators?

Injuries are common in animals of diverse taxa and are usually attributed to encounters with predators. Although often non‐lethal, injuries nevertheless represent effects of predators that can have negative consequences for demography and fitness (e.g. reproductive costs). However, encounters with predators also represent experience through which animals can learn and positively adapt their future behaviour, potentially mitigating, at least partly, the negative effects of prior exposure to predators. I predicted that injured grass snakes (Natrix natrix), which presumably had been handled previously by a predator, would be more likely to move before capture than uninjured snakes. This prediction was borne out. Snakes with injuries also had lower body condition than uninjured snakes, although the effect was non‐significant. Snakes that had been previously captured also were significantly more likely to move before capture than snakes that had never been caught before. These results provide strong evidence for the role of experience and learning in modifying the antipredator behaviour of snakes.     

Accidental altruism in insular pit-vipers (Gloydius shedaoensis, Viperidae)

Darwinian theory predicts that organisms will display traits that benefit themselves rather than other individuals; exceptions to this rule usually are explicable by kin selection. Our studies on an insular population of venomous snakes in north-eastern China reveal a different situation. Only one species of snake (Gloydius shedaoensis, Viperidae) occurs on the island of Shedao, and displays altruism between size (age) classes. First, small snakes frequently kill prey items larger than they can swallow themselves. This behaviour enhances rates of feeding of larger conspecifics, which scavenge the birds' carcasses. Second, large snakes kill raptorial birds (sparrowhawks Accipiter nisus) that pose little or no threat to themselves. This behaviour reduces predation risk for smaller snakes. These effects are presumably accidental consequences of the high venom toxicity of the pit-vipers, which enable them to kill inedible prey and non-threatening predators at little cost. Nonetheless, this accidental altruism may have significant ecological consequences. For example, these behaviours may contribute to the remarkably high population densities of snakes on Shedao.     

Coral snake mimicry: live snakes not avoided by a mammalian predator

The occurrence of coral snake coloration among unrelated venomous and non-venomous New World snake species has often been explained in terms of warning coloration and mimicry. The idea that snake predators would avoid coral snakes in nature seems widely established and is postulated in many discussions on coral snake mimicry. However, the few workers that have tested a potential aposematic function of the conspicuous colour pattern focused exclusively on behaviour of snake predators towards coloured abstract models. Here we report on behaviour of temporarily caged, wild coatis (Nasua narica) when confronted with co-occurring live snakes, among which were two species of venomous coral snakes. Five different types of responses have been observed, ranging from avoidance to predation, yet none of the coatis avoided either of the two coral snake species or other species resembling these. As in earlier studies coatis appeared to avoid coral snake models, our findings show that results from studies with abstract snake models cannot unconditionally serve as evidence for an aposematic function of coral snake coloration.     

Can Wall Lizards Combine Chemical and Visual Cues to Discriminate Predatory from Non‐Predatory Snakes Inside Refuges?

The ability to use multiple cues in assessing predation risk is especially important to prey animals exposed to multiple predators. Wall lizards, Podarcis muralis, respond to predatory attacks from birds in the open by hiding inside rock crevices, where they may encounter saurophagous ambush smooth snakes. Lizards should avoid refuges with these snakes, but in refuges lizards can also find non‐saurophagous viperine snakes, which lizards do not need to avoid. We investigated in the laboratory whether wall lizards used different predator cues to detect and discriminate between snake species within refuges. We simulated predatory attacks in the open to lizards, and compared their refuge use, and the variation in the responses after a repeated attack, between predator‐free refuges and refuges containing visual, chemical, or visual and chemical cues of saurophagous or non‐saurophagous snakes. Time to enter a refuge was not influenced by potential risk inside the refuge. In contrast, in a successive second attack, lizards sought cover faster and tended to increase time spent hidden in the refuge. This suggests a case of predator facilitation because persistent predators in the open may force lizards to hide faster and for longer in hazardous refuges. However, after hiding, lizards spent less time in refuges with both chemical and visual cues of snakes, or with chemical cues alone, than in predator‐free refuges or in refuges with snake visual cues alone, but there were no differences in response to the two snake species. Therefore, lizards could be overestimating predation risk inside refuges. We discuss which selection pressures might explain this lack of discrimination of predatory from similar non‐predatory snakes.     

Are natural threats superior threats?

Threat superiority effects describe the reaction time advantage for locating threatening objects in a visual search paradigm, compared to locating visually similar non-threatening objects. They are widely reported for threats of both natural (snakes and spiders) and man-made (guns and knives) origins. Across two experiments, the current study contrasts threat superiority effects for natural and man-made targets. When targets are not depicted held, snakes and spiders tended to exhibit larger threat superiority effects, and were searched for with additional caution, than were guns and knives. When snakes and spiders were depicted held and weapons wielded, systematic differences between the natural and man-made threats disappeared. This means the advantage for threats of natural origin observed when all targets were depicted not held may be attributable to differences in animation – snakes and spiders are alive and may strike at any time if in your vicinity, whereas a weapon can only inflict harm if wielded. From these data there is no evidence that evolved visual sensitivities to the basic shapes of venomous animals support faster detection and response times to these animals than can occur to targets such as guns and knives, whose shapes must be learned. The selection pressures that led to the evolution of such sensitivities (observable even in infancy) may therefore lie in protecting young children and babies from envenomation, before they even have the cognitive capacity to understand the dangers that snakes and spiders pose.