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We used the COVID-19 pandemic closure at the Saint Louis Zoo to examine visitor effects on space use and glucocorticoid levels in banteng, grizzly bears, polar bears, and western lowland gorillas. The study was divided into four six-week phases: closure in spring 2020, reopening in summer 2020, fall 2020, and spring 2021 as a seasonal comparison. Space use data were collected using video, and fecal samples were assayed for glucocorticoids. Generalized linear models were used to examine differences in zone occupancy and glucocorticoids between phases. The banteng spent more time near visitors, and glucocorticoids were only temporarily elevated in two of five animals when visitors returned. The grizzly bears spent more time in their habitat than in the den, and the polar bear spent more time near viewing areas after visitors returned. Glucocorticoids did not differ significantly between the closure and reopening for any bears. The gorillas spent less time close to visitors immediately after reopening but this effect waned by fall; glucocorticoid data were not available. Overall, based on space use and glucocorticoid levels, we suggest visitor effects on the gorillas are neutral, on the grizzly bears are neutral or positive, and are positive on the banteng and polar bear.

Fecal glucocorticoid metabolite (FGM) concentrations, activity, and personality were assessed for 35 fennec foxes (Vulpes zerda) to determine whether animals managed as ambassadors differed from ex...

This study describes patterns of steroid hormone production and activity for banteng (Bos javanicus), an endangered member of the Bovidae family. Using validated assays, concentrations of fecal progestagens, androgens, and glucocorticoids were quantified for four females and one male at the Saint Louis Zoo. A commercial activity monitor was also validated for assessing movement. The devices were then used to characterize activity in relation to season, reproductive status, and fecal steroid concentrations. General linear mixed models assessed differences in activity and steroid concentrations among individuals, in regards to reproductive status and season. Ovulatory cycle patterns, changes in activity around estrus and parturition, and events correlated with increased glucocorticoid production were also documented. Cycle lengths were 24.7 ± 0.4 days, and cycle lengths varied among individuals. Females cycled year-round, but luteal progestagen concentrations, along with glucocorticoids and male androgens, increased during the summer. Activity also increased in the summer. Progestagen concentrations were greater in pregnant females, and the gestation length of one pregnancy was 254 days. Pregnant females were less active overall, but activity increased the day before parturition. Activity was also greater preceding the onset of the luteal phase. The majority of glucocorticoid concentrations were in the range of baseline concentrations. However, a small number of elevated concentrations were correlated with husbandry and veterinary events. This study is the first to validate non-invasive methods for monitoring reproduction, welfare, and activity of banteng. Our results may contribute to the improved management of captive populations.

Modern herpetoculture has seen a rise in welfare-related habitat modifications, although ethologically-informed enclosure design and evidence-based husbandry are lacking. The diversity that exists within snakes complicates standardizing snake welfare assessment tools and evaluation techniques. Utilizing behavioral indicators in conjunction with physiological measures, such as fecal glucocorticoid metabolite concentrations, could aid in the validation of evidence-based metrics for evaluating snake welfare. We increased habitat cleaning, to identify behavioral or physiological indicators that might indicate heightened arousal in snakes as a response to the disturbance. While glucocorticoid metabolite concentrations increased significantly during a period of increased disturbance, this increase was not associated with a significant increase in tongue-flicking, a behavior previously associated with arousal in snakes. Locomotion behavior and the proportion of time spent exposed were also not affected by more frequent habitat cleaning. These results demonstrate the need to further investigate the behavioral and physiological responses of snakes to different aspects of animal care at a species and individual level. They also highlight the need to collect baseline behavioral and physiological data for animals, in order to make meaningful comparisons when evaluating changes in animal care.

Fecal glucocorticoid metabolite (FGM) concentrations, activity, and personality were assessed for 35 fennec foxes (

Special encounters that allow contact between animals and guests are common in zoos and aquariums. Visitors to the Saint Louis Zoo may touch guinea pigs serving as ambassador animals. We evaluated two welfare indicators in ambassador guinea pigs by comparing glucocorticoid levels and body weights between periods when guinea pigs lived in a habitat accessible to the public and while off exhibit. Mean glucocorticoid levels did not differ between sexes or between on- and off-exhibit periods. There was significant individual variation, and females demonstrated greater variation than males. While on exhibit, glucocorticoid levels slightly but significantly increased in males and decreased in females. Moving guinea pigs between habitats only elicited a significant glucocorticoid response when females were moved off exhibit. Temporary closures of the exhibit had no effect on glucocorticoid levels in either sex. Analyses of the impact of handling rates on males found no impact on glucocorticoid levels. Guinea pigs&rsquo, body weights were lower while on exhibit. We conclude that guinea pigs serving as ambassador animals at the Saint Louis Zoo demonstrate comparable physiological profiles while on and off exhibit and, when used in a rotational schedule, are a suitable species for animal encounters involving contact with the public.

Zoological institutions often use immersive, naturalistic exhibits to create an inclusive atmosphere that is inviting for visitors while providing for the welfare of animals in their collections. In this study, we investigated physiological changes in salivary cortisol and blood pressure, as well as psychological changes among visitors before and after a walk through the River's Edge, an immersive, naturalistic exhibit at the Saint Louis Zoo. Study participants had a significant reduction in salivary cortisol and blood pressure after walking through the exhibit. Psychological assessments of mood found that most visitors felt happier, more energized, and less tense after the visit. Additionally, participants who spent more time in River's Edge, had visited River's Edge prior to the study, and had seen more exhibits at the Zoo prior to entering River's Edge experienced greater psychological and/or physiological benefits. We conclude that immersive, naturalistic exhibits in zoos can elicit positive changes in physiological and psychological measures of health and well-being and argue for a greater scientific focus on the role of zoos and other green spaces in human health.

Somali wild ass (Equus africanus somaliensis) are the most endangered species of wild equid, and little is known about their reproductive biology. Here, we describe social interactions and glucocorticoid production of five pairs of Somali wild ass females and their foals, which were born within a two-month period at the Saint Louis Zoo. Behavioral data were recorded from 0 to 3 months (Infant), and again from 6 to 12 months of age (Older Foal). Fecal samples were collected three times a week for 12 months, starting when foals were born, and analyzed for glucocorticoids using a validated assay. Generalized linear mixed models assessed differences in behavior and glucocorticoid concentrations between foals and mothers, in regard to foal sex, foal age, and season. Nursing bout frequency and total nursing duration, but not bout duration, decreased with age. Low nursing rates were still observed in 11 to 12-month-old foals, suggesting that weaning occurs after one year. Rates of social behaviors were similar between mothers and their foals, and rates of agonistic behavior were similar to rates of affiliative behavior. Male foals initiated and received agonistic behaviors more than female foals. Patterns of interactions varied with foal age. Infants interacted primarily with their mothers, whereas interactions among foals were common during the Older Foal period. Glucocorticoid concentrations of foals and mothers varied in relation to foal age; concentrations for both were highest in the first months of life. Seasonal differences in glucocorticoid concentrations were also observed, but no evidence of elevated glucocorticoid production was found except around the time of parturition. This suggests that neither mothers nor foals experienced chronic stress. Our study presents new information on the biology of Somali wild asses and it is one of the few to document behavior and physiology in a non-domestic equid. This information may contribute to improved management of ex situ populations by enhancing individual welfare and increasing reproductive success.

Fruit doves (Ptilinopus) constitute a genus of small to medium-sized, brightly colored arboreal birds, whose diets consist entirely of fruit. Little is known about the behavior of fruit doves because most species inhabit dense forests and are difficult to observe in the wild. This study describes the parental behavior of Jambu fruit dove pairs (Ptilinopus jambu) in a captive breeding program at the Saint Louis Zoo. Continuous video recordings were made of three pairs which raised a total of eight squabs over 2 years; daily rates of parental and squab behaviors were quantified. Overall, females were present at the nest, brooded their squabs, pecked, and attempted feedings more often than males. Parents also cared for their squabs at different times throughout the day. Males fed and brooded squabs during the middle of the day, while females fed throughout the day and brooded in the morning, evening, and overnight. Feeding rates were lower than those described for seed-eating doves (Columbinae), with hours between consecutive feedings, and squabs rarely begged before feeding events. Most squab behaviors involved initiating or terminating brooding and self-preening. These behaviors increased as squabs approached fledging, and coincided with a shift from full to partial brooding, and a decrease in parental allopreening. Older squabs also initiated feeding less frequently. Together, these data provide the first description of parental behavior in a Ptilinopus fruit dove. The results of this study may help improve captive breeding efforts, which are likely to become increasingly important for future conservation and reintroduction programs.

Great hornbills (Buceros bicornis) are a long-lived, monogamous species that forms strong pair-bonds, and mate compatibility is thought to be important for successful reproduction. Within AZA, great hornbills are listed as a red SSP. The population consists of a limited number of individuals that do not breed reliably, and improving reproduction is a top priority for the Coraciiformes TAG. To better understand mating behavior and evaluate mate compatibility, this study documented the behavior of pairs of great hornbills during and immediately after courtship. Using live observations, the study followed one female, an experienced and successful breeder, as she was paired with four successive males over 11 breeding seasons. Initially, males frequently vocalized, investigated the nest, and approached the female. As the female spent more time in the nest, these behaviors were replaced by regurgitation and food offering. The female was most often observed plastering and vocalizing. Behavioral differences between successful and unsuccessful pairs, possibly indicative of pair compatibility, included rates of approaching, billing, and biting. Numerous behaviors occurred more frequently during years that a chick hatched, including pseudoregurgitation, regurgitation, offering food items, and nest investigation. Males also spent more time in proximity to both the female and the nest during years that a chick hatched. Together, these results suggest that the amount of time pairs spend in proximity, the amount of time a male spends near the nest, and the frequency of certain behaviors may help evaluate compatibility and the likelihood of successful reproduction for pairs of great hornbills.

Although reproduction in the domestic horse has been well described, less is known about reproduction in wild equids. This study describes endocrine patterns associated with estrous cycles and pregnancy for Somali wild asses (Equus africanus somaliensis), an endangered African equid. Fecal samples were collected three times per week for more than 2 years from five female Somali wild asses at the Saint Louis Zoo; progestagen and estrogen metabolites were quantified using commercially available immunoassays. Progestagen analysis indicated that cycle lengths were 27.2 ± 1.2 days and females cycled throughout the year. Progestagen levels during early pregnancy were low and not sustained above baseline until approximately 40 weeks prior to partition. Concentrations increased markedly around 16 weeks prior to delivery and peaked 2-3 weeks before birth. Fecal estrogen levels also increased significantly starting 40-45 weeks before parturition and reached their maximal value approximately 20 weeks prior to birth. Neither foal heat nor lactational suppression of estrus was observed, and females cycled within 45 days after delivery. These data are the first to describe the reproductive physiology of Somali wild asses. As the species faces increasing threats in the wild, this information may support conservation efforts by assisting with ex situ breeding programs.

Birds in multi-nestling broods often experience reduced growth and elevated nutritional and social stress as a result of competition for parental resources. While responses are often species-specific, experimentally increasing brood size has been shown to decrease growth and increase production of both testosterone and corticosterone in nestling passerines. To investigate the relationship between brood size, growth, and steroid hormone production in eastern bluebirds, we cross-fostered nestlings to small, medium, and large broods. Body mass, skeletal size, serum testosterone, and baseline serum corticosterone concentration were measured prior to fledging. Bluebird nestlings raised in large broods weighed less than nestlings in small and medium-sized broods and secreted elevated concentrations of testosterone. Nestling corticosterone concentrations did not vary in response to brood size or body condition, but concentrations were higher in males compared to females. Our results suggest that nestling bluebirds experiencing nutritional and social stress increase testosterone production. Elevated concentrations may enhance begging and competitive abilities in smaller young. Higher concentrations of corticosterone in males may be related to sex-biased provisioning. Further experimental work is needed to ascertain the effects of elevated testosterone concentrations in nestling bluebirds, as well as the causes and consequences of elevated corticosterone for male nestlings.

We studied androgen production during development in nestling Eastern Screech-Owls (Megascops asio) and hypothesized that gender and hatch order might influence serum levels of testosterone and androstenedione. Testosterone levels were highest immediately after hatching and declined significantly in the 4 weeks leading to fledging. The average level of testosterone for 1–7 day-old owls was 3.99 ± 0.68 ng/ml. At 22–28 days of age, the average testosterone level for nestling owls was 0.83 ± 0.18 ng/ml. Testosterone levels did not differ between males or females. The average testosterone level for male nestlings was 2.23 ± 0.29 ng/ml and 2.39 ± 0.56 ng/ml for female nestlings. The average level of androstenedione for nestling owls was 1.92 ± 0.11 ng/ml and levels remained constant throughout development. Levels were significantly higher in males than females. The average androstenedione level was 1.77 ± 0.16 ng/ml for male nestlings and 1.05 ± 0.24 ng/ml for female nestlings. Hatching order did not ...

When a female bird begins incubation before clutch completion, the nestlings hatch sequentially, and a size hierarchy forms within the brood. This size hierarchy may be minimized or exacerbated through differential allocation of resources to eggs across the laying order. In this study, we characterize intra-clutch variation in cockatiel clutches by measuring egg mass, yolk mass, and concentrations of yolk testosterone, androstenedione, and corticosterone. Cockatiels are a long-lived member of the Psittaciformes. Because asynchronous hatching may reduce sibling competition and allow for extended development periods in long-lived birds, we predicted that female cockatiels would allocate maternal resources in a way that would reinforce the brood size hierarchy. Significant within-clutch differences in egg size and steroid concentrations were observed. Eggs at the end of the laying sequence were smaller and had significantly smaller yolks than eggs early in the laying order. Fifth-laid eggs, as well as first-laid eggs, contained significantly lower concentrations of testosterone than eggs in other positions of the laying sequence. No differences in yolk androstenedione concentration were observed. Yolk corticosterone concentrations increased linearly with laying order. Together, these patterns might reinforce the brood size hierarchy created by asynchronous hatching.

Fruit doves (Ptilinopus) constitute a genus of small to medium-sized, brightly colored arboreal birds, whose diets consist entirely of fruit. Little is known about the behavior of fruit doves because most species inhabit dense forests and are difficult to observe in the wild. This study describes the parental behavior of Jambu fruit dove pairs (Ptilinopus jambu) in a captive breeding program at the Saint Louis Zoo. Continuous video recordings were made of three pairs which raised a total of eight squabs over 2 years; daily rates of parental and squab behaviors were quantified. Overall, females were present at the nest, brooded their squabs, pecked, and attempted feedings more often than males. Parents also cared for their squabs at different times throughout the day. Males fed and brooded squabs during the middle of the day, while females fed throughout the day and brooded in the morning, evening, and overnight. Feeding rates were lower than those described for seed-eating doves (Columbinae), with hours between consecutive feedings, and squabs rarely begged before feeding events. Most squab behaviors involved initiating or terminating brooding and self-preening. These behaviors increased as squabs approached fledging, and coincided with a shift from full to partial brooding, and a decrease in parental allopreening. Older squabs also initiated feeding less frequently. Together, these data provide the first description of parental behavior in a Ptilinopus fruit dove. The results of this study may help improve captive breeding efforts, which are likely to become increasingly important for future conservation and reintroduction programs.

Great hornbills (Buceros bicornis) are a long-lived, monogamous species that forms strong pair-bonds, and mate compatibility is thought to be important for successful reproduction. Within AZA, great hornbills are listed as a red SSP. The population consists of a limited number of individuals that do not breed reliably, and improving reproduction is a top priority for the Coraciiformes TAG. To better understand mating behavior and evaluate mate compatibility, this study documented the behavior of pairs of great hornbills during and immediately after courtship. Using live observations, the study followed one female, an experienced and successful breeder, as she was paired with four successive males over 11 breeding seasons. Initially, males frequently vocalized, investigated the nest, and approached the female. As the female spent more time in the nest, these behaviors were replaced by regurgitation and food offering. The female was most often observed plastering and vocalizing. Behavioral differences between successful and unsuccessful pairs, possibly indicative of pair compatibility, included rates of approaching, billing, and biting. Numerous behaviors occurred more frequently during years that a chick hatched, including pseudoregurgitation, regurgitation, offering food items, and nest investigation. Males also spent more time in proximity to both the female and the nest during years that a chick hatched. Together, these results suggest that the amount of time pairs spend in proximity, the amount of time a male spends near the nest, and the frequency of certain behaviors may help evaluate compatibility and the likelihood of successful reproduction for pairs of great hornbills.

Aquatic organisms may use different mechanisms to locate an odour source, including odour-gated rheotaxis and chemotaxis. However, the effects of spatial and temporal information contained within odour signals in guiding orientation are under debate. In nature, turbulence and plume meandering give rise to signal intermittency, which organisms must overcome to locate an odour source successfully. In this study, crayfish were presented with odour plumes formed by a continuously released jet or a pulsed jet ranging from 0.5 to 3 Hz. Crayfish oriented significantly differently in pulsed odour plumes differed significantly from than in a continuously released jet plume. In a continuous odour plumes, crayfish walked faster, had more accurate heading and turning angles, and stopped less often. The results suggest that crayfish are able to orient in a pulsed odour plumes, but that the spatial and temporal complexity of a continuous released odour plume allowed them to be more successful to find the odour source.

Abstract A variety of factors influences the formation of hierarchical structures, and can include an altered aggressive state, an ability to physically dominate, and previous agonistic experience. Using male Orconectes rusticus, we tested the duration of the winner effect by varying the time between a winning encounter and a subsequent encounter by a 20, 40 or 60-minute interval. Varying the time between the two fights significantly altered the probabilities of initiating fight behaviour and of winning a fight. A crayfish with a 20-minute delay between its winning experience and its subsequent fight was significantly less likely to initiate fight behaviour and significantly more likely to win its next fight than was an animal whose next fight was delayed for 40 or 60 minutes. We then investigated whether the dynamics of this winner effect were influenced by perception of odour signals during agonistic interactions by blocking the chemo- and mechanoreceptors on the antennae and antennules to prevent reception of relevant cues communicating social status. Individuals fighting an opponent with this loss of sensory information were significantly more likely to initiate a fight, but then escalated at a slower rate to a higher fight intensity level. In addition, individuals had a decreased chance of winning an agonistic bout against an opponent deprived of sensory input from the antennae and antennules.

of the world after each molt. That is, ommatidia viewing the most anterior region of the animal's visual space now sample a more lateral region of visual space. As a consequence, the retinotopic map in the brain must undergo comparable rearrangements to accommodate inputs from new ommatidia sampling visual space in front of the animal. The retinotopic map has been determined for adult animals at the first two synaptic layers in the brain (6), but that of the juvenile remains to be studied. The retinotopic map must be plastic to accommodate the changing retinal mosaic as the eye grows. Supported by grants from the National Science Foundation, National Eye Institute, and the National Institutes of Mental Health. C. McGuiness and S. Meadors received REU Fellowships from the National Science Foundation. We thank the f the world after each molt. That is, ommatidia viewing the most terior egion f the animal's visual space now sample a more ateral region f visual space. As a consequence, the retinotopic ap in the brain must undergo comparable rear ng ments to ccommodate inputs from new ommatidia sampling visual space front of the animal. The retinotopic map has been determined for dult animals at the first two synaptic layers n the b ain (6), but at of the juvenil remains to be studied. The retinotopic map ust be plastic o accommodate the changing retinal mosaic as the Monomoy National Wildlife Refuge, Morris Island, Chatham, Massachusetts.

Leach's storm-petrels accumulate large amounts of body mass throughout the nestling period. Approximately 4days before fledging, nestlings weigh 50-100% more than adults. In order to shed this excess mass, nestlings engage in behavioral anorexia, and leave the burrow when they are light enough to fly. During this prefledging period, we measured several plasma hormones (corticosterone, thyroxine, and testosterone) to determine whether the behavioral changes associated with fledging are correlated with endocrine changes. In several species, including petrels, corticosterone levels are known to increase near fledging. Reduced food consumption has been shown to elevate corticosterone levels and decrease thyroxine levels in nestling birds. In nestling storm-petrels, levels of both corticosterone and thyroxine increased. Storm-petrels were found to secrete measurable levels of testosterone, but levels did not change during the prefledging period. Increased corticosterone levels might be part of an endocrine signal that initiates changes in feeding behavior, or may result from reduced food intake. Elevated thyroxine levels may be related to metabolic changes involved in mass loss. Future experimental work is needed to ascertain whether the described endocrine changes are responsible for, or result from, prefledging changes in petrel feeding behavior.

Female birds deposit significant amounts of steroid hormones into the yolks of their eggs. Studies have demonstrated that these hormones, particularly androgens, affect nestling growth and development. In order to measure androgen concentrations in avian egg yolks, most authors follow the extraction methods outlined by Schwabl (1993. Proc. Nat. Acad. Sci. USA 90:11446–11450). We describe a simplified method for extracting androgens from avian egg yolks. Our method, which has been validated through recovery and linearity experiments, consists of a single ethanol precipitation that produces substantially higher recoveries than those reported by Schwabl (1993. Proc. Nat. Acad. Sci. USA 90:11446–11450). Zoo Biol 28:137–143, 2009. © 2008 Wiley-Liss, Inc.