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Predator evasion by white-tailed deer fawns

Despite their importance for understanding predator–prey interactions, factors that affect predator evasion behaviours of offspring of large ungulates are poorly understood. Our objective was to characterize the influence of selection and availability of escape cover and maternal presence on predator evasion by white-tailed deer, Odocoileus virginianus, fawns in the northern Great Plains, U.S.A. We observed 45 coyote, Canis latrans, chases of fawns, and we participated in 83 human chases of fawns during 2007–2009, of which, 19 and 42 chases, respectively, ended with capture of the fawn. Evasive techniques used by fawns were similar for human and coyote chases. Likelihood of a white-tailed deer fawn escaping capture, however, was influenced by deer group size and a number of antipredator behaviours, including aggressive defence by females, initial habitat and selection of escape cover, all of which were modified by the presence of parturient females. At the initiation of a chase, fawns in grasslands were more likely to escape, whereas fawns in forested cover, cultivated land or wheat were more likely to be captured by a coyote or human. Fawns fleeing to wetlands and grasslands also were less likely to be captured compared with those choosing forested cover, wheat and cultivated land. Increased probability of capture was associated with greater distance to wetland and grassland habitats and decreased distance to wheat. Use of wetland habitat as a successful antipredator strategy highlights the need for a greater understanding of the importance of habitat complexity in predator avoidance.

Animal Behaviour

Male traits, mating tactics and reproductive success in the buff-breasted sandpiper, Tryngites subruficollis

Buff-breasted sandpipers use a variety of mating tactics to acquire mates, including remaining at a single lek for most of the breeding season, attending multiple leks during the season, displaying solitarily or displaying both on leks and solitarily. We found that differences in body size, body condition, fluctuating asymmetry scores, wing coloration, territory location and behaviour (attraction, solicitation and agonistic) did not explain the observed variation in mating tactics used by males. Which males abandoned versus returned to leks was also not related to morphology or behaviour, and there was no tendency for males to join leks that were larger or smaller than the lek they abandoned. These results suggest that male desertion of leks was not dependent on a male's characteristics nor on the size of the lek he was presently attending. Males did join leks with larger males than their previous lek, perhaps to mate with females attracted to these larger 'hotshot' males. Males at both leks and solitary sites successfully mated. Lek tenure did not affect mating success, although lekking males appeared to mate more frequently than solitary males. Courtship disruption and to a lesser extent, female mimicry, were effective at preventing females from mating at leks, and may offer a partial explanation for female mating off leks. Our analysis that combined all males together within a year (regardless of mating tactic) indicated that males that attended leks for longer periods of time and that had fewer wing spots were significantly more likely to mate. Given some evidence that wing spotting declines with age, and that females inspect male underwings during courtship, the latter result suggests that female choice may play some role in determining male success. We suggest that male buff-breasted sandpipers may use alternative mating tactics more readily than males in other 'classic' lek-breeding species because: (1) unpredictable breeding conditions in this species' high arctic breeding range leads to low lek stability, which in turn hinders mate selection mechanisms mediated by male dominance and female choice; and (2) males are not constrained by morphological markings that indicate status or sex. Both characteristics may reduce the reproductive benefits associated with males adopting one mating tactic and result in a sort of scramble competition in which males switch between tactics as local conditions change.

Animal Behaviour

Certainty of paternity and paternal investment in eastern bluebirds and tree swallows

Extra-pair paternity is common in many socially monogamous passerine birds with biparental care. Thus, males often invest in offspring to which they are not related. Models of optimal parental investment predict that, under certain assumptions, males should lower their investment in response to reduced certainty of paternity. We attempted to reduce certainty of paternity experimentally in two species, the eastern bluebird, S ialia sialis , and the tree swallow, Tachycineta bicolor , by temporarily removing fertile females on two mornings during egg laying. In both species, experimental males usually attempted to copulate with the female immediately after her reappearance, suggesting that they experienced the absence of their mate as a threat to their paternity. Experimental males copulated at a significantly higher rate than control males. However, contrary to the prediction of the model, experimental males did not invest less than control males in their offspring. There was no difference between experimental and control nests in the proportion of male feeds, male and female feeding rates, nestling growth and nestling condition and size at age 14 days. We argue that females might have restored the males’ confidence in paternity after the experiment by soliciting or accepting copulations. Alternatively, males may not reduce their effort, because the fitness costs to their own offspring may outweigh the benefits for the males, at least in populations where females cannot fully compensate for reduced male investment.

Animal Behaviour

Forced copulation results in few extrapair fertilizations in Ross's and lesser snow geese

Extrapair paternity varies from 0 to over 70% of young among various populations of birds. Comparative studies have suggested that this variation is related to nesting density, breeding synchrony and the proportion of extrapair copulations. We used minisatellite DNA fingerprinting to examine levels of extrapair paternity in Ross's geese, Chen rossi, and lesser snow geese, C. caerulescens c. (hereafter snow geese) nesting in the largest known goose colony in the world. These geese have one of the highest known percentages of extrapair copulation (46-56% of all attempted copulations), and all of these appeared to be forced. Among all successful copulations, 33 and 38% were extrapair in Ross's and snow geese, respectively. Despite the high percentage of extrapair copulations, extrapair paternity was low in both Ross's and snow geese (2-5% of young). Extrapair paternity was not related to nest density in either species. However, in snow geese, extrapair paternity was more likely to occur in nests of females that nested asynchronously, either early or late in the season. This is one of a few reported examples of a negative relationship between extrapair paternity and breeding synchrony. Extrapair young also tended to come from eggs laid later in the clutch. Although forced extrapair copulations appear to be a relatively inefficient reproductive tactic for males, they may provide a reproductive advantage for some males.

Animal Behaviour

Mate fidelity and breeding site tenacity in a monogamous sandpiper, the black turnstone

We examined the relationship between mate fidelity and breeding site tenacity during a 5-year study of the black turnstone, Arenaria melanocephala, a socially monogamous sandpiper breeding in subarctic Alaska. We tested the predictions of several hypotheses regarding the incidence of divorce and the benefits of fidelity to mate and breeding site. Interannual return rates to the breeding grounds (88% for males, 79% for females) were among the highest yet recorded for any scolopacid sandpiper, and 88% of returning birds nested on their previous year's territory. The annual divorce rate was only 11%, and mate fidelity was significantly linked to fidelity to territory but independent of sex and year. Males arrived in spring significantly earlier than their mates and interannual fidelity was influenced by the relative timing of arrival of pair members. Reunited pairs had significantly higher fledging success than new pairs formed after death or divorce. The incidence of divorce was unrelated to reproductive success the previous year, although birds nested significantly further away after failure than after a successful nesting attempt. Sightings of marked individuals suggested that members of pairs do not winter together, and breeding site tenacity provides a mechanism through which pair members can reunite. We reject the 'incompatibility' hypothesis for divorce in turnstones, and our data contradict predictions of the 'better option' hypothesis. Alternatively, we propose the 'bet-hedging' hypothesis to explain the occurrence of divorce, which transpires when an individual pairs with a new mate to avoid the cost of waiting for a previous mate to return. Such costs can include remaining unmated, if the former mate has died, or experiencing lower reproductive success because of delayed breeding.

Animal Behaviour

Ontogenetic improvement of visual function in the medaka Oryzias latipes based on an optomotor testing system for larval and adult fish

We developed a system for evaluation of visual function in larval and adult fish. Both optomotor (swimming) and optokinetic (eye movement) responses were monitored and recorded using a system of rotating stripes. The system allowed manipulation of factors such as width of the stripes used, rotation speed of the striped drum, and light illuminance levels within both the scotopic and photopic ranges. Precise control of these factors allowed quantitative measurements of visual acuity and motion detection. Using this apparatus, we tested the hypothesis that significant posthatch ontogenetic improvements in visual function occur in the medaka Oryzias latipes , and also that this species shows significant in ovo neuronal development. Significant improvements in the acuity angle alpha (ability to discriminate detail) were observed from approximately 5 degrees at hatch to 1 degree in the oldest adult stages. In addition, we measured a significant improvement in flicker fusion thresholds (motion detection skills) between larval and adult life stages within both the scotopic and photopic ranges of light illuminance. Ranges of flicker fusion thresholds ( X ±SD) at log I=1.96 (photopic) varied from 37.2±1.6 cycles/s in young adults to 18.6±1.6 cycles/s in young larvae 10 days posthatch. At log I=−2.54 (scotopic), flicker fusion thresholds varied from 5.8±0.7 cycles/s in young adults to 1.7±0.4 cycles/s in young larvae 10 days posthatch. Light sensitivity increased approximately 2.9 log units from early hatched larval stages to adults. The demonstrated ontogenetic improvements in visual function probably enable the fish to explore new resources, thereby enlarging their fundamental niche.

Animal Behaviour

Scent-marking in lone wolves and newly formed pairs

Scent-marking was studied in wolves (Canis lupus) along 133 km of tracks in northern Minnesota during winters of 1975 to 1976 and 1976 to 1977 and in two captive packs and four captive pairs for various periods. Lone wolves, which possess neither mates nor territories, rarely marked by raised-leg urination and defaecated and urinated less along roads and trails, where territorial pairs and packs generally marked. Newly formed pairs marked the most, eventually decreasing their rates to those of established packs. Generally, wolves that scent-marked also bred, whereas non-marking wolves usually did not breed. Scent-marking apparently is important to the success of courtship in new pairs and to reproductive synchrony in established pairs, as well as serving a territorial function.

Animal Behaviour

Group foraging by a stream minnow: shoals or aggregations?

The importance of social attraction in the formation of foraging groups was examined for a stream-dwelling cyprinid, the rosyside dace, Clinostomus funduloides. Dace arrivals and departures at natural foraging sites were monitored and tested for (1) tendency of dace to travel in groups, and (2) dependency of arrival and departure rates on group size. Dace usually entered and departed foraging sites independently of each other. Group size usually affected neither arrival rate nor departure probability. Thus, attraction among dace appeared weak; foraging groups most often resulted from dace aggregating in preferred foraging sites. The strongest evidence of social attraction was during autumn, when dace departure probability often decreased with increasing group size, possibly in response to increased threat of predation by a seasonally occurring predator. Dace also rarely avoided conspecifics, except when an aggressive individual defended a foraging site. Otherwise, there was little evidence of exploitative competition among dace for drifting prey or of foraging benefits in groups, because group size usually did not affect individual feeding rates. These results suggest that the benefits of group foraging demonstrated under laboratory conditions in other studies may not always apply to field conditions.

Animal Behaviour

DNA fingerprint similarity between female and juvenile brown-headed cowbirds trapped together

This DNA fingerprinting study investigates whether females of the brood parasite brown-headed cowbird, Molothrus ater, associate with their own juvenile offspring at feeding sites more often than would be expected by chance. Cowbirds lay their eggs in the nests of a variety of host species and, as far as is known, leave them to the care of foster parents. Using baited walk-in funnel traps, 36 adult female-juvenile pairs (or trios) of cowbirds were trapped. Blood samples were collected from these individuals to conduct DNA fingerprinting analyses, calculate similarity indices, and to compare S-values for the 11 comparisons of juveniles and the females with which they were caught with S-values of random pairings of juveniles and the females in adjacent gel lanes with which they were not caught. Overall band-sharing was significantly higher for the individuals trapped together than for the random pairings. These associations between juvenile cowbirds and their mothers could occur as a result of female cowbirds monitoring the development of their young in the nests where they have laid. Alternatively, nestling cowbirds in the nest could become familiar visually and locally with a female parent that is frequently in their territory and could follow her when she departs for feeding grounds. In either case these data suggest that adult cowbirds associate with juveniles, in some cases their own offspring, and that offspring may learn to function as cowbirds in part from this association.

Animal Behaviour

Weather as a proximate explanation for fission–fusion dynamics in female northern long-eared bats

Fission–fusion dynamics appear common among temperate bats where females form roost groups that change in size and composition, as females switch roosts almost daily. One hypothesis for frequent roost switching is that females move to find suitable thermal conditions as ambient conditions change. Tests of this hypothesis have, however, been conducted mostly at roosts in artificial structures where microclimate is relatively stable. The goal of our study was to determine whether roost switching and roost use by northern long-eared bats, Myotis septentrionalis , that roost in trees are related to ambient conditions. We used generalized linear fixed effects models to explore the influence of roost characteristics and changes in ambient conditions on the likelihood of roost switching. We used canonical correlation analyses to examine the relationship between ambient conditions and roost characteristics. Roost switching was indeed linked to ambient conditions together with characteristics of roosts on the previous day; the best descriptors of roost switching differed between the two geographical regions we analysed. In Nova Scotia, females were less likely to switch roosts when it rained, particularly if they were in roosts below surrounding canopy whereas they were more likely to switch roosts when they were in roosts of high decay. Females roosted in shorter trees in earlier decay classes on warm days, as well as on windy and rainy days. In Kentucky, females were more likely to switch roosts at high temperatures, particularly when they were in roosts in high decay. Females roosted in shorter, decayed trees on warm days, and in less decayed trees with small diameter on windy and rainy days. Our results suggest bats switch roosts in response to changes in ambient conditions to select suitable roosting conditions, which may explain some of the proximate factors shaping fission–fusion dynamics of bats.

Animal Behaviour

Birds choose long-term partners years before breeding

Pair bonds can provide social benefits to long-term monogamous species alongside their benefits for reproduction. However, little is known about when these bonds form, in particular how long they are present before breeding. Previous studies of pair formation in long-term monogamous birds have been rather data-limited, but for many migratory birds they report pair formation on the wintering grounds. We provide the first systematic investigation of prebreeding association patterns of long-term monogamous pairs by examining entire life histories based on tracking data of migratory whooping cranes, Grus americana. We found that a substantial portion (62%) of breeding pairs started associating at least 12 months before first breeding, with 16 of 58 breeding pairs beginning to associate over 2 years before first breeding. For most pairs, these associations with future breeding partners also became unique and distinguishable from association patterns with nonpartner individuals 12 months before first breeding. In addition, 60% of pair associations began before at least one partner had reached nominal sexual maturity. Most pairs began associating in the late spring upon arrival at the summer grounds, while associations beginning at other times of the year were rare. Patterns in the associations of pairs prior to breeding can point to the potential benefits of prebreeding relationships, for instance providing support in competitive interactions or increasing partner familiarity.

Animal Behaviour

Migratory hummingbirds make their own rules: The decision to resume migration along a barrier

Knowing how naïve migrants respond to intrinsic and extrinsic factors experienced en route will allow a more thorough understanding of the endogenous migratory programme. To understand how inexperienced individuals respond to ecological features, we tracked the migratory departures of juvenile ruby-throated hummingbirds, Archilochus colubris , one of the smallest (∼ 3 g) and least-studied migrants, along the Gulf of Mexico during southbound migration using an international automated radiotelemetry system. The recent miniaturization of radiotags provides a novel method to track one of the smallest migratory birds, rendering the first information on departure decisions of known hummingbirds in relation to an ecological barrier. Using weather conditions and individual attributes, we also determined which factors influenced the time and direction of departure from a coastal stopover site. Most migrants (83%) departed in the morning, and daily departure time was only influenced by stopover duration, the amount of time spent at a stopover site. The majority (77%) of departure orientations paralleled the coastline, and we found little influence of any factor on departure direction. Our results reveal that (1) juvenile hummingbirds departing coastal Alabama move in a direction indicative of a circum-Gulf path during southbound migration and (2) departure decisions support a fly-and-forage strategy in which hummingbirds likely take advantage of resources along the coast while moving towards their destination.

Gulf of Mexico

Effects of rearing treatment on the behavior of captive whooping cranes (Grus americana)

Small founder populations of whooping cranes are managed to maximize egg production for the purpose of reintroducing young to the wild. This results in an excessive number of hatched chicks that cannot be naturally reared by parents. Hand-rearing techniques have been developed to raise the additional hatches. However, hand rearing may affect the behavior of the birds and their chances of survival later in life. The objectives of this study were to determine the impact of rearing practices on the behavior of whooping crane chicks. The birds were reared under three commonly used rearing techniques: parent reared (PR), hand reared (HR), and hand reared with exercise (HRE). Fifty-six whooping crane chicks were observed by focal animal sampling from hatch to 20 weeks of age. During these observations, occurrences of comfort behavior, aggression, foraging, nonvigilance, sleep, vigilance, and other types of behavior were collected. Data were analyzed using mixed models repeated measures analysis of variance (ANOVA). Behavior was affected by rearing treatment, age, and time of day. PR birds spent more time being vigilant than HR and HRE birds. An inverse correlation was found between percentage of time foraging and vigilant (r = -0.686, P < 0.0001). However, there were no differences in the behavior of birds reared in HR or HRE programs.

Applied Animal Behaviour Science

The effects of captive rearing on the behavior of newly-released whooping cranes (Grus americana)

Rearing treatments used in captivity to prepare animals for reintroduction to the wild may have a profound effect on behavior and, possibly, affect their survival after reintroduction. This study examined the behaviors of captive-reared whooping cranes (Grus americana) upon their release in Florida to determine if rearing treatments may affect the behavior of the birds and how these affect their chances of survival in the wild. Individually tagged birds were observed at the rearing facility, the U.S. Geological Survey Patuxent Wildlife Research Center in Maryland, from hatch to 20 weeks of age and at the release site in Central Florida for up to 6 weeks post release. The rearing treatments were parent reared (PR), hand reared (HR), and hand reared with exercise (HRE). Observations at the rearing facility are described in a previous paper. At the release site, each bird was observed for 5 min every morning (0700?1000 h) and late afternoon (1500?1800 h) during the 6-week study period. Our results indicated that most of the time, the n = 34 birds were foraging (46.03 ? 1.48%), followed by nonvigilant (20.89 ? 0.73%), vigilant (19.21 ? 0.72%), or performing comfort behaviors (11.61 ? 1.28%). Data were analyzed using mixed models repeated measures ANOVA. There were no significant behavioral differences between HR and HRE birds. PR birds were found in larger groups than HR birds during the first 2 weeks post release and greater than HR and HRE birds afterwards. This may be interpreted as an antipredator strategy for birds that relied on parental guidance during rearing. HR and HRE birds foraged more than PR birds during the first 2 weeks post release and PR birds were more vigilant during the first 2 weeks post release. Across rearing treatments, the percentages of time spent foraging and engaged in vigilant behaviors during rearing were positively correlated with their behavior upon release. If any of these behaviors can be demonstrated to have relevance for the survival of the whooping cranes after release then it may be possible to establish behavioral interventions to increase the frequencies of such behavior, so that they are perpetuated after release.

Applied Animal Behaviour Science

Behavior of feral horses in response to culling and GnRH immunocontraception

Wildlife management actions can alter fundamental behaviors of individuals and groups,which may directly impact their life history parameters in unforeseen ways. This is especially true for highly social animals because changes in one individual&rsquo;s behavior can cascade throughout its social network. When resources to support populations of social animals are limited and populations become locally overabundant, managers are faced with the daunting challenge of decreasing population size without disrupting core behavioral processes. Increasingly, managers are turning to fertility control technologies to supplement culling in efforts to suppress population growth, but little is quantitatively known about how either of these management tools affects behavior. Gonadotropin releasing hormone (GnRH) is a small neuropeptide that performs an obligatory role in mammalian reproduction and has been formulated into the immunocontraceptive GonaCon-BTM. We investigated the influences of this vaccine on behavior of feral horses ( Equus caballus ) at Theodore Roosevelt National Park, North Dakota, USA, for a year preceding and a year following nonlethal culling and GnRH-vaccine treatment. We observed horses during the breeding season and found only minimal differences in time budget behaviors of free-ranging female feral horses treated with GnRH and those treated with saline. The differences observed were consistent with the metabolic demands of pregnancy and lactation. We observed similar social behaviors between treatment groups, reflecting limited reproductive behavior among control females due to high rates of pregnancy and suppressed reproductive behavior among treated females due to GnRH-inhibited ovarian activity. In the treatment year, band stallion age was the only supported factor influencing herding behavior (P < 0.001), harem-tending behavior (P < 0.001), and agonistic behavior (P = 0.02). There was no difference between the mean body condition of control females (4.9 (95% CI = 4.7&ndash;5.1)) and treated females(4.8 (95% CI = 4.7&ndash;4.9)). Band fidelity among all females increased 25.7% in the year fol-lowing vaccination and culling, despite the social perturbation associated with removal of conspecifics. Herding behavior by stallions decreased 50.7% following treatment and culling (P < 0.001), while harem-tending behavior increased 195.0% (P < 0.001). The amount of available forage influenced harem-tending, reproductive, and agonistic behavior in the year following culling and treatment (P < 0.04). These changes reflected the expected nexus between a species with polygynous social structure and strong group fidelity and the large instantaneous change in population density and demography coincident with culling.Behavioral responses to such perturbation may be synergistic in reducing grazing pressure by decreasing energetically expensive competitive behaviors, but further investigation is needed to explicitly test this hypothesis.

Applied Animal Behaviour Science

You can go your own way: No evidence for social behavior based on kinship or familiarity in captive juvenile box turtles

Behavioral interactions between conspecific animals can be influenced by relatedness and familiarity. Compared to other vertebrate taxa, considering such aspects of social behavior when housing captive reptiles has received less attention, despite the implications this could have for informing husbandry practices, enhancing welfare, and influencing outcomes of conservation translocations. In this study, to test how kinship and familiarity influenced social behavior in a reptile, we reared 16 captive-born Eastern Box Turtles ( Terrapene carolina ) under semi-natural conditions in four equally sized groups, where each group comprised pairs of siblings and non-siblings. Using separation distance between pairs of turtles in rearing enclosures as a measure of gregariousness, we found no evidence suggesting siblings more frequently interacted with one another compared to non-relatives over the first five months of life (β = -0.016, 95% CI: -0.117 to 0.084). Average pair separation distance decreased during this time (β = -0.146, 95% CI: -0.228 to -0.063) but may have been due to turtles aggregating around concentrated resources like heat and moist retreat areas as cold winter temperatures approached. When subject were eight months old, we measured repeated separation distances between unique pair combinations in an experimental environment and similarly found no support for gregariousness (associations) being influenced by kinship or familiarity (β = -1.554, 95% CI: -9.956 to 6.848). Additionally, differences in body size between pairs of turtles (β = -22.289, 95% CI: -68.448 to 23.870) nor the five-minute time interval during the 90-minute trial (P ≥ 0.18) had any apparent effect on associations. Agonistic interactions between individuals were never observed. Encouragingly, based on our results, group housing and rearing of juvenile box turtles did not appear to negatively impact their behavioral and physiological well-being. Unlike findings for other taxa, including some reptiles, our results suggest strategically housing groups of juvenile T. carolina to maintain social stability may not be an important husbandry consideration, or even a requirement, when planning releases of captive-reared individuals for conservation purposes.

Applied Animal Behaviour Science