Ambush site selection and ontogenetic shifts in foraging strategy in a semi-aquatic pit viper, the Eastern cottonmouth

Although habitat selection has been studied in a variety of snake taxa, little is known about habitat selection in aquatic snake species. Additionally, due to their small size and secretive nature, juvenile snakes are seldom included in habitat selection studies. The Eastern cottonmouth Agkistrodon piscivorus is a semi-aquatic pit viper known to use ambush, sit-and-wait foraging strategies. Ambush hunters are likely to select habitats that increase opportunity for successful prey capture while minimizing predation risk and maintaining appropriate thermal and hydric conditions. We characterized the foraging strategy and microhabitat use of cottonmouths at Ellenton Bay, an isolated Carolina bay freshwater wetland on the Savannah River Site in SC, USA. We measured habitat characteristics of 55 ambush sites used by 51 individual cottonmouths located during nighttime visual surveys, as well as 225 randomly selected sites within our search area. Cottonmouths exhibited an ontogenetic shift in foraging strategy with juveniles using predominately ambush foraging around the edge of the wetland while adults were most often encountered actively moving within the wetland. Principal components analysis revealed that juveniles selected foraging microhabitats that were different from random and consisted of mud substrate with sparse vegetation, whereas adults occupied a greater variety of microhabitats that did not differ from random. Concomitantly, free-ranging cottonmouths exhibited ontogenetic shifts in diet: juveniles consumed mostly salamanders, while adults ate a greater variety of prey including other snakes and birds. Our results highlight the importance of understanding how ontogenetic changes in coloration, diet and predation risk influence foraging strategy and microhabitat selection in snakes.     

Sexual size dimorphism of the tongue in a North American pitviper

Sexual dimorphisms – phenotypic dissimilarities between the sexes – are common and widespread among plants and animals, and classical examples include differences in body size, colour, shape, ornamentation and behaviour. In general, sexual dimorphisms are hypothesized to evolve by way of sexual selection acting on one sex through priority-of-access for sexual partners via mate choice and/or intra-sexual competition. In snakes, males are the mate-searching sex and one form of sexual selection involves male–male competition in locating females by following pheromone trails using their forked tongues, the structure used to sample environmental chemicals for transduction in the vomeronasal chemosensory system (VNS). Based on several lines of empirical evidence, increased tongue forking (bifurcation) in snakes (and some lizard taxa) appears to enhance chemical trail-following abilities through tropotaxis (the simultaneous comparison of stimulus intensities on two sides of the body) and thus aids in prey location and mate searching in males. We predicted that male copperheads, Agkistrodon contortrix , a North American pitviper, should have more deeply forked tongues than females owing to male–male competition for priority-of-access to widely dispersed females during the mating seasons. We examined formalin-fixed, ethanol-preserved museum specimens of adult A. contortrix for sexual size dimorphism (SSD) of the tongue. Tongue dimensions showed differences indicative of SSD, and the degree of bifurcation (i.e. mean tine length) was significantly greater in males. Various structures of the VNS and associated regions (e.g. muscles) in some vertebrate taxa show sexual dimorphism, but our study is the first to document dimorphism in the tongue of a tetrapod vertebrate.     

Genetic analysis reveals diverse kin‐directed routes to helping in the rifleman Acanthisitta chloris

The social organization of cooperatively breeding species is extremely variable, with diverse social group composition and patterns of relatedness. Species that exhibit alternative routes to helping within the same population are potentially useful systems to investigate the causes and fitness consequences of diverse evolutionary pathways to cooperative behaviour. In this study, we use microsatellite markers and field observations to describe helping behaviour and patterns of relatedness in the unusual cooperative breeding system of the rifleman Acanthisitta chloris. First, we show that rifleman helpers consist of a remarkably diverse demographic, including males and females, who may be adult or juvenile, failed breeders or nonbreeders, or even successful breeders that simultaneously feed their own brood. Adult helpers mostly helped at first‐brood nests, while first‐brood juveniles assisted their parents at second broods. Second, we show that rifleman pairs are strictly sexually monogamous, and helpers did not gain any current reproductive success through helping. Third, genotyping showed that contrary to previous assumptions, helpers were closely related to the recipients of their care and preferentially directed care towards relatives over contemporaneous nests of nonrelatives. Finally, we show that variation in helper provisioning effort was attributed to age: juvenile helpers provisioned less than adults and were less responsive to the demands of a growing brood. Overall, our results show that the diverse routes to helping in this unusual species are driven by the common theme of kinship between helper and recipients, resulting in a previously underestimated potential for helpers to gain indirect fitness benefits.     

A test of non‐kin social foraging in the southern flying squirrel (Glaucomys volans)

It can be challenging to understand the evolution of sociality, particularly the occurrence of co‐operation by non‐kin. Southern flying squirrels (Glaucomys volans) are an interesting example of non‐kin co‐operation because of the mutual benefits obtained by social thermoregulation during winter. Because group survival confers benefits to the entire group, flying squirrels may also follow an aggregation economy, whereby co‐operative foraging during winter is advantageous. However, the extent of such social foraging in flying squirrels is unknown. We tested for social foraging of southern flying squirrels, and also for relatedness among foraging groups. To determine the structure of foraging groups, we set up and remotely monitored feeding stations and nest cavities. All squirrels at the study site were tagged with passive integrated transponder (PIT) tags and nests and feeding stations were monitored with automated PIT‐tag recorders for a 24‐month period. Squirrels were found most often foraging alone. Squirrels that were recorded foraging together comprised unrelated individuals that were also found to share nest cavities. Squirrels were also recorded travelling farther distances between nest cavity and feeding station in the winter season than in the summer season, suggesting that, during winter, squirrels trade‐off proximity to food caches for membership in a nest group. Our data suggest that squirrels forage and cache alone in their summer home range and make solitary returns to this summer range to collect their cache during the winter months, despite exhibiting social winter nesting. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 1126–1135.     

Offspring food allocation by parents and helpers in a cooperative mammal

In cooperatively breeding species, helpers and parents commonlyface two decisions when they find a food item: first, whetherto feed the item to a young group member or to eat it themselves;and second, which offspring to feed. Little is known aboutthe factors that influence these decisions in cooperative mammals,though optimal foraging theory provides a basis for a rangeof predictions. In this article we describe pup feeding behaviorby helpers and parents in a cooperative mongoose, the meerkat(Suricata suricatta). When meerkat pups begin accompanyingthe group, they beg food from older group members, who digup dispersed prey items. As predicted, the probability of aprey item being fed to a pup shows a positive relationship with prey size and a negative relationship with pup distance.Meerkats apparently follow a "feed the nearest pup rule" andare more likely to feed the nearest pup if it is hungry. Hungrierpups beg more and follow older group members more closely.Across all age categories, females feed pups more frequentlythan males, both in terms of the relative frequency of feeds,and the proportion of prey biomass found by each individualthat is fed to pups. Females also feed female pups significantlymore than male pups, while males feed pups of both sexes equally.These sex biases in feeding contributions may result from femalegroup members benefiting more than males from higher pup survival,and in particular higher female pup survival, because femalesare the philopatric sex.     

Short‐term dynamics of nest occupancy in an allopaternal species,the tessellated darter Etheostoma olmstedi

To inquire how male size interacts with alloparental behaviour and mating success in the tessellated darter Etheostoma olmstedi, males were given a choice of nests with or without eggs; subsequent nest occupancy, takeovers and egg deposits were monitored. Subordinate males readily occupied available nests with eggs but were often evicted by dominant males, suggesting that males of all sizes compete for the opportunity to provide allopaternal care in this species.     

Social semantics: toward a genuine pluralism in the study of social behaviour

Pluralism is the coexistence of equivalent theoretical frameworks, either because they are historically entrenched or because they achieve separate insights by viewing the same process in different ways. A recent article by West et al. [Journal of Evolutionary Biology (2007) vol. 20, 415-432] attempts to classify the many equivalent frameworks that have been developed to study the evolution of social behaviour. This article addresses shortcomings in the West et al.'s article, especially with respect to multilevel selection, in a common effort to maximize the benefits of pluralism while minimizing the semantic costs.     

Diet affects female mating behaviour in a seed‐feeding beetle

In species where males provide nuptial gifts, females can improve their nutritional status and thus increase their fecundity by mating when in need of resources. However, mating can be costly, so females should only mate to acquire resources when the need for resources is large, such as when females are nutritionally‐deprived. Two populations of the seed‐feeding beetle Callosobruchus maculatus, a species that produces relatively large nuptial gifts, are used to test whether female nutritional status affects mating behaviour. Female access to water, sugar and yeast are manipulated and the fitness consequences of these manipulations are examined together with the effects of diet on the propensity of nonvirgin females to mate. Access to water has a small but significant effect on mass loss over time, lifespan and fecundity of females, relative to unfed controls. Access to sugar (dissolved in water) improves female fecundity and lifespan above that of hydrated females but access to yeast has no positive effects on female survival or reproduction. Diet has a large effect on both receptivity of nonvirgin females to a male and how quickly they accept that male. Unfed females are both more likely to mate, and accept a mate more quickly, than females provided access to water, which are more likely to mate and accept a mate more quickly than females provided with sugar. This rank order of behaviours matches the order predicted if females increase their mating rate when nutritionally deprived (i.e. it matches the effect of diet on female fitness). The results obtained also suggest that mate choice may be condition‐dependent: females from one population (Burkina Faso) show a preference for large males when well‐fed but not when unfed, although this result is not found in a second population (South India). It is concluded that nutritionally‐deprived females are more receptive to mates than are well‐fed females, consistent with the hypothesis that females ‘forage’ for nuptial gifts, or at least more willingly accept sperm in exchange for nuptial gifts, when they are nutritionally deprived.     

Giving offspring a head start in life: field and experimental evidence for selection on maternal basking behaviour in lizards

The timing of birth is often correlated with offspring fitness in animals, but experimental studies that disentangle direct effects of parturition date and indirect effects mediated via variation in female traits are rare. In viviparous ectotherms, parturition date is largely driven by female thermal conditions, particularly maternal basking strategies. Our field and laboratory studies of a viviparous lizard (Niveoscincus ocellatus) show that earlier‐born offspring are more likely to survive through their first winter and are larger following that winter, than are later‐born conspecifics. Thus, the association between parturition date and offspring fitness is causal, rather than reflecting an underlying correlation between parturition date and maternal attributes. Survival selection on offspring confers a significant advantage for increased maternal basking in this species, mediated through fitness advantages of earlier parturition. We discuss the roles of environmentally imposed constraints and parent–offspring conflict in the evolution of maternal effects on parturition date.     

Among‐individual heterogeneity in maternal behaviour and physiology affects reproductive allocation and offspring life‐history traits in the garter snake Thamnophis elegans

Accumulating evidence suggests that within‐individual plasticity of behavioural and physiological traits is limited, resulting in stable among‐individual differences in these aspects of the phenotype. Furthermore, these traits often covary within individuals, resulting in a continuum of correlated phenotypic variation among individuals within populations and species. This heterogeneity, in turn, affects individual fitness and can have cross‐generational effects. Patterns of trait covariation, among‐individual differences, and subsequent fitness consequences have long been recognized in reptiles. Here, we provide a test of patterns of among‐individual heterogeneity in behaviour and physiology and subsequent effects on reproduction and offspring fitness in the garter snake Thamnophis elegans. We find that measures of activity levels vary among individuals and are consistent within individuals in reproductive female snakes, indicating stable behavioural phenotypes. Blood hormone and glucose concentrations are not as stable within individuals, indicating that these traits do not describe consistent physiological phenotypes. Nonetheless, the major axes of variation in maternal traits describe behavioural and physiological phenotypes that interact to predict offspring body condition and mass at birth. This differential allocation of energy to offspring, in turn, strongly influences subsequent offspring growth and survival. This pattern suggests the potential for strong selection on phenotypes defined by behaviour–physiology interactions.     

Correlated geographic variation in predation risk and antipredator behaviour within a wide‐ranging snake species (Notechis scutatus,Elapidae)

Geographic variation in antipredator behaviour within wide‐ranging species may be driven by both genetic and environmental influences. We quantified antipredator responses in neonatal (laboratory born, n = 555) and adult (field caught, n = 346) tiger snakes (Notechis scutatus) from 11 mainland and island sites in southern Australia. We used these data to test predictions from Bonnet et al.'s hypothesis that the vigour of antipredator responses in this species reflects behavioural plasticity (in turn, driven by an individual snake's exposure to predators during its lifetime) rather than by genetic variation in this trait. We used the number of predator taxa in each area as an index of predator risk. As predicted, adult snakes from predator‐rich areas had more vigorous defensive responses when handled, whereas neonatal behaviour (although also variable among populations) was unrelated to predator species richness. Adult males bit more readily than adult females (as expected from the greater predation exposure of males during mate searching) but no such sex difference was evident in neonates. Although alternative models remain possible, our data are most consistent with the hypothesis that geographic divergence in antipredator tactics within this species primarily reflects developmentally plastic responses to local predation risk.     

Fine‐scale spatiotemporal patterns of genetic variation reflect budding dispersal coupled with strong natal philopatry in a cooperatively breeding mammal

The relatedness structure of animal populations is thought to be a critically important factor underlying the evolution of mating systems and social behaviours. While previous work has shown that population structure is shaped by many biological processes, few studies have investigated how these factors vary over time. Consequently, we explored the fine‐scale spatiotemporal genetic structure of an intensively studied population of cooperatively breeding banded mongooses (Mungos mungo) over a 10‐year period. Overall population structure was strong (average F_ST = 0.129) but groups with spatially overlapping territories were not more genetically similar to one another than noncontiguous groups. Instead, genetic differentiation was associated with historical group‐fission (budding) events, with new groups diverging from their parent groups over time. Within groups, relatedness was high within but not between the sexes, although the latter increased over time since group formation due to group founders being replaced by philopatric young. This trend was not mirrored by a decrease in average offspring heterozygosity over time, suggesting that close inbreeding may often be avoided, even when immigration into established groups is virtually absent and opportunities for extra‐group matings are rare. Fine‐scale spatiotemporal population structure could have important implications in social species, where relatedness between interacting individuals is a vital component in the evolution of patterns of inbreeding avoidance, reproductive skew and kin‐selected helping and harming.     

Dispersal in Kin Coalition Throughout the Non‐Breeding Season to Facilitate Fine‐Scale Genetic Structure in the Breeding Season: Evidence From a Small Passerine

Evidence shows that social cooperation among kin may evolve even in birds with extensive dispersal. In such cases, maintaining kinship during dispersal is essential to the subsequent expression of kin cooperation. This hypothesis has not been examined for most bird species. We addressed it in the ground tit (Parus humilis), a passerine where kin frequently interact in terms of cooperative polygamy and extra‐pair mating despite fast annual turnover of the breeding population. Pedigree and genotype data showed that while groups varied in composition throughout the non‐breeding season due to continual individual emigration and immigration, they always contained kin coalitions consisting of either local or immigrant individuals of different age and sexes. The first‐order kin coalitions, according to the information from local individuals, stemmed from single‐family lineages (siblings and their parents), and the lower‐order ones from neighbouring, related family lineages that merged after fledging. It was probable that immigrants had formed kin coalitions in similar ways before dispersing. Groups broke up in the breeding season. Pairing between unrelated individuals from different coalitions within a group was more likely, whereas related individuals from the same coalition tended to nest near each other. The resulting fine‐scale population genetic structure is expected to facilitate breeding interactions among kin. Our findings give clues to understanding the evolution of social cooperation in relation to dispersal.     

Maternal predator‐exposure affects offspring size at birth but not telomere length in a live‐bearing fish

The perception of predation risk could affect prey phenotype both within and between generations (via parental effects). The response to predation risk could involve modifications in physiology, morphology, and behavior and can ultimately affect long‐term fitness. Among the possible modifications mediated by the exposure to predation risk, telomere length could be a proxy for investigating the response to predation risk both within and between generations, as telomeres can be significantly affected by environmental stress. Maternal exposure to the perception of predation risk can affect a variety of offspring traits but the effect on offspring telomere length has never been experimentally tested. Using a live‐bearing fish, the guppy (Poecilia reticulata), we tested if the perceived risk of predation could affect the telomere length of adult females directly and that of their offspring with a balanced experimental setup that allowed us to control for both maternal and paternal contribution. We exposed female guppies to the perception of predation risk during gestation using a combination of both visual and chemical cues and we then measured female telomere length after the exposure period. Maternal effects mediated by the exposure to predation risk were measured on offspring telomere length and body size at birth. Contrary to our predictions, we did not find a significant effect of predation‐exposure neither on female nor on offspring telomere length, but females exposed to predation risk produced smaller offspring at birth. We discuss the possible explanations for our findings and advocate for further research on telomere dynamics in ectotherms.     

Levels of extra‐pair paternity are associated with parental care in penduline tits (Remizidae)

In most passerine birds, individuals attempt to maximize their fitness by providing parental care while also mating outside their pair bond. A sex‐specific trade‐off between these two behaviours is predicted to occur, as the fitness benefits of extra‐pair mating differ between the sexes. We use nest observations and parentage analysis to reveal a negative association between male care and the incidence of extra‐pair paternity across three species of penduline tit (Remizidae). This provides evidence of a trade‐off between these two behaviours, possibly due to the devaluing of paternal care by extra‐pair offspring.     

First experiences with free‐ranging giant day geckos (Phelsuma madagascariensis grandis,Gray 1870) in the masoala rainforest exhibit in Zurich Zoo,Switzerland

Sixty giant day geckos (Phelsuma madagascariensis grandis) have been introduced into the Masoala rainforest exhibit of Zurich Zoo. Eighteen geckos were monitored during a 3‐month study using the telemetry method. In general, the geckos were able to adapt well under these semi‐natural conditions of the exhibit. The geckos accessed to nearly all locations in the exhibit; some of the animals remained stationary, whereas others covered dozens of meters within a week. They preferred four species of plants as well as the artificial structures supporting the walls and roof of the exhibit, and displayed natural behaviors. Animals were observed feeding on all kind of arthropods and nectar substances, and also on food items dropped from feeding stations for other species. Deliberate feeding targeted at the geckos was finally reduced to a calcium‐rich squash along the visitor's main path. Reproduction was observed frequently within the exhibit. These findings suggest that the geckos will be able to build up a self‐sustaining population and contribute to a nearly‐authentic experience for zoo visitors. Zoo Biol 0:1–7, 2006. © 2006 Wiley‐Liss, Inc.     

The impact of paternity on male–infant association in a primate with low paternity certainty

In multimale groups where females mate promiscuously, male–infant associations have rarely been studied. However, recent studies have shown that males selectively support their offspring during agonistic conflicts with other juveniles and that father's presence accelerates offspring maturation. Furthermore, it was shown that males invest in unrelated infants to enhance future mating success with the infant's mother. Hence, infant care might provide fitness gain for males. Here, we investigate male–infant associations in rhesus macaques (Macaca mulatta), a primate with low paternity certainty as females mate with multiple partners and males ensure paternity less efficiently through mate‐guarding. We combined behavioural data with genetic paternity analyses of one cohort of the semi‐free‐ranging population of Cayo Santiago (Puerto Rico) and recorded affiliative and aggressive interactions between focal subjects and adult males from birth to sexual maturation (0–4 years) of focal subjects. Our results revealed that 9.6% of all interactions of focal subjects involved an adult male and 94% of all male–infant interactions were affiliative, indicating the rareness of male–infant aggression. Second and most interestingly, sires were more likely to affiliate with their offspring than nonsires with unrelated infants. This preference was independent of mother's proximity and emphasized during early infancy. Male–infant affiliation rose with infant age and was pronounced between adult males and male rather than female focal subjects. Overall, our results suggest that male–infant affiliation is also an important component in structuring primate societies and affiliation directed towards own offspring presumably represent low‐cost paternal care.     

Natural history and behaviour of Dunatothrips aneurae Mound (Thysanoptera: Phlaeothripidae), a phyllode‐gluing thrips with facultative pleometrosis

Thrips of the genus Dunatothrips (Thysanoptera: Phlaeothripidae) construct domiciles by tying phyllodes of Australian Acacia trees together with silk‐like glue. Females often co‐found domiciles (pleometrosis), an apparently cooperative behaviour that potentially provides insights into social evolution. However, little is known about their basic natural history, limiting the scope for testable predictions. Here, we address this crucial prerequisite step by investigating some key outstanding questions in the most common species, D. aneurae, on its host, Acacia aneura. We detail distribution in space and time, mating, dispersal, domicile building and defence. Dunatothrips aneurae was distributed in loosely reproductively synchronized patches, and tended to prefer east‐facing, terminal phyllodes on thin‐phyllode A. aneura varieties. Mature domiciles contained middens, concentrated areas of waste, suggesting active maintenance of domiciles and the potential for the division of labour. We observed inbreeding and outbreeding. Dunatothrips aneurae males engaged in short, truncated matings with sisters before dispersing locally, mating with females in nearby immature domiciles; longer distance dispersal, although it must happen, is still undocumented. Males and females mated multiply. Lone females required male presence to initiate domiciles, constructed them without male help and lost wings on nesting by abscission. Silk production occurred well before egg laying. Aggression or defence appeared to be entirely absent. Taken together, these observations suggest that research into co‐founding behaviour should focus on: (1) local crowding; (2) lack of aggression; and (3) potential division of labour with respect to egg production, silk production and domicile maintenance. These results should provide a springboard for questions on the potential evolution of cooperation in this species. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 802‐816.