Behavior of polar bear (Ursus maritimus) cubs post-den emergence at the Detroit Zoo.

In the wild, female polar bears (Ursus maritimus) with cubs may spend extended periods of time within the den following initial emergence. As a result, studying behavioral development of cubs at the denning site has been difficult and unreliable. Although care staff at zoological institutions have easier access to animals, every effort is made to minimize intrusions to maternal groups, which presents research limitations. Detroit Zoological Society staff used audio-equipped cameras installed in behind-the-scenes spaces to conduct approximately 9 weeks of monitoring on two female polar bear cubs of the same litter, one of whom (Laerke) was hand-reared while the other (Astra) was mother-reared. Monitoring spanned ages 14-24 weeks and consisted of 12, 5-min focal observations per observation day timed to evenly cover the entire 24-h day. Using generalized linear mixed models, we examined relationships between behavior and time of day, hour, and age. We also conducted descriptive analyses and used these to draw comparisons between the two cubs where appropriate. Despite different rearing environments, both cubs had highly comparable patterns of inactive, locomotory, and independent play behaviors. Astra generally decreased time spent nursing and in proximity to Suka as she aged. Data presented here represent a continuation of previous neonatal observations conducted on Astra, adding to the minimal body of knowledge currently available on first year polar bear cub development. We also report successful mitigation of stereotypic behavior in Laerke. Further studies of captive maternal groups can provide insight into this critical developmental stage, supporting both captive-based breeding efforts and in situ conservation efforts.

The Benefits of Increased Space and Habitat Complexity for the Welfare of Zoo-Housed King Penguins (Aptenodytes patagonicus).

Zoos and aquariums accredited by the Association of Zoos and Aquariums embrace animal welfare as a foundational principle of habitat design. Modern habitats are designed to provide animals with choices and agency over their environment, and to encourage species-appropriate behavior and space use. In 2016, the Detroit Zoological Society opened the Polk Penguin Conservation Center, a 3065.80 m2 facility that features a naturalistic design. The building was designed to optimize animal welfare by incorporating various substrates, nesting sites, and a 1234 kL pool with elements of underwater complexity. The facility houses a mixed-species group of penguins that were previously housed in a smaller habitat that opened in 1968. Between 2015 and 2022, we opportunistically monitored the behavior of ten king penguins (Aptenodytes patagonicus) as they moved back and forth between the two buildings while additional modifications were made to the new habitat. We collected 695 h of behavioral observations and 10,416 h of data from flipper-mounted time-depth recorders. We found that the king penguins spent less time engaged in aggression and more time engaged in swimming and positive social behaviors in the redesigned space. They also spent less time in proximity to other species of penguins and more time alone. These behavioral trends suggest that increased space and environmental complexity had positive welfare benefits for these penguins.

Behavioral development of a captive polar bear (Ursus maritimus) cub in the maternal den.

Two female polar bear (Ursus maritimus) cubs were born to mother Suka at the Detroit Zoo on November 17, 2020. One cub (Laerke) was hand-reared out of medical necessity, while the other cub (Astra) was mother-reared. Using audio-equipped video cameras previously installed in the maternal den, staff were able to conduct uninterrupted continuous monitoring on Suka and Astra for an unprecedented 12 weeks. Data collected during this time were analyzed descriptively. Astra spent a maximum of approximately 20% of any given 24 h period in nursing behaviors. Distribution of time spent nursing was relatively even across the 24 h day. In contrast to the limited data currently available, Astra spent only approximately 11% of the over 2000 h of behavioral data engaged in nursing behaviors. Astra left the nest for the first time in her fourth week of life, and time spent out of the nest gradually increased with age. Suka was observed guiding Astra back to the nest the first few times Astra ventured out. However, maintenance of maternal proximity did not appear to be a motivating factor in patterns of space use. The data presented here represent the longest uninterrupted continuous monitoring of a polar bear cub in the maternal den currently published. Publication of similar data from other institutions would establish more generalizable trends for early infancy development. These data are critical for understanding cub welfare and improving population sustainability and neonate mortality rates.

Impact of alternating overnight housing conditions on welfare measures in a bachelor group of western lowland gorillas (Gorilla gorilla gorilla).

Spatial parameters and design of overnight housing spaces can have significant influences on both nocturnal and diurnal behaviors of western lowland gorillas (Gorilla gorilla gorilla) in captivity. This is especially true for bachelor groups, as adult males have demonstrated a greater tendency to separate from conspecifics overnight. Additionally, previous studies have suggested that the social needs of individuals in bachelor units may shift over time. The bachelor unit at the Detroit Zoo is managed in a unique hybrid strategy in which the group is housed socially for three nights (either indoors or indoors with outdoor access), then separately on the fourth. A 2016 study of this group (n = 3) suggested that individuals had better welfare following social overnight housing as compared to solitary overnight housing. We aimed to reassess how overnight housing conditions affect individual welfare several years later. We revised a previously developed qualitative behavioral assessment tool that included 13 behavioral items spanning both positive and negative indicators of welfare. Animal care staff completed the assessment and collected a fecal sample every day for each gorilla for a period of 3 months. We used Kruskal-Wallis tests to analyze variation in behavioral items and FGMs between conditions. We then used generalized linear mixed models to identify whether overnight housing condition or other potentially confounding variables were driving observed variation in welfare indicators. We found significant variation demonstrating significantly better welfare indicators after being housed separately as compared to being socially housed indoors overnight. Although separate housing appears to be the ideal condition for this group at this point in time, if seasonality permits, social housing with outdoor access may be a feasible alternative to consistent overnight separation for these gorillas.

Behavioral Assessment of Six Reptile Species during a Temporary Zoo Closure and Reopening.

Although reptiles are commonly housed in zoos and aquariums, their welfare is understudied for the diversity of species housed and the taxon's current captive population size. The sensory abilities of reptiles have adapted to the varied ecological niches they inhabit, and these evolutionary adaptations impact how reptiles perceive the stimuli around them-including zoo visitors. This study aimed to assess visitor effects on small groups of six reptile species during a temporary zoo closure due to COVID-19 by measuring behavioral diversity, use of space (measured by a spread of participation index), and select behaviors. The species assessed showed diverse responses. The Catalina Island rattlesnakes (Crotalus catalinensis) demonstrated increased investigation and behavioral diversity after the zoo reopened compared to when the zoo was closed, but the European glass lizards (Pseudopus apodus) showed decreases in the amount of time spent exposed to the observers' view and in their evenness of space use after the zoo was reopened to visitors. The other species, including beaded lizards (Heloderma horridum), Sonoran spiny-tailed iguana (Ctenosaura macrolopha), Arrau turtles (Podocnemis expansa), and dwarf caimans (Paleosuchus palpebrosus), had intermediate changes in their responses to visitor presence.

Survey of current group demographics and management practices of bachelor groups of western lowland gorillas (Gorilla gorilla gorilla) across North America.

Recommendations for the successful management and maintenance of bachelor groups of western lowland gorillas (Gorilla gorilla gorilla) in zoological settings have been an increasingly prevalent focus within the zoological community. Over the past two decades, studies have examined the impact of various environmental factors on the stability of bachelor groups, proposed management strategies for bachelor groups, and compared behavioral trends between bachelor and mixed-sex groups. These studies have clearly demonstrated that bachelor groups are complex social units that require specialized management approaches. In this study, we aimed to assess the extent to which bachelor group management across North American zoos accredited by the Association of Zoos and Aquariums aligns with established recommendations. We distributed a comprehensive survey broadly encompassing habitat and housing, aggression and wounding, group demographics, feeding, and training to the 22 zoos housing bachelor groups as of 2019. We received completed surveys from 19 zoos, representing a total of 21 social units and 59 individual gorillas. We used descriptive statistics to represent the range of current management strategies across the surveyed population and ANOVAs to assess significant variation in key demographic variables. Our results demonstrate that a majority of zoos have adopted the best practices for the formation of social groups established by Stoinski et al. in 2004. However, there is much less standardization across zoos in protocols surrounding training and feeding. Additionally, important variables in the assessment of wounding, such as time of day and location, are often unknown or not observed by animal care professionals. We highlight these two areas as being of particular focus in developing and adhering to consistent protocols across institutions.

Group Composition and Social Structure of Red-Shanked Doucs (Pygathrix nemaeus) at Son Tra Nature Reserve, Vietnam.

Multilevel societies, consisting of multiple one-male multi-female units, are relatively rare among primates, but are more widespread in the odd-nosed colobines than other taxa. Multilevel societies are found particularly in snub-nosed monkeys (Rhinopithecus)and have been debated in studies of proboscis monkeys (Proboscis). While it has been suggested that douc langurs (Pygathrix) may also form multilevel societies, the limited data available make the details of their social organization unclear. We aimed to establish a more comprehensive picture of the social organization of red-shanked doucs (Pygathrix nemaeus) and to address the question of whether this species forms multilevel societies, specifically collections of multiple distinct one-male units hereafter termed "bands." We collected 259 h of behavioral data at Son Tra Nature Reserve in Vietnam from February 2010 to May 2011. The mean band size was approximately 18 individuals. Bands were comprised of approximately 2.7 units, and each unit contained approximately 6.5 individuals. Units had an average sex ratio of 1.0:1.6. We observed fission and fusion behaviors which were not correlated with phenological or weather measures. Activity budget data showed that fission and fusion behaviors between units were positively correlated with activity. Both vocalizations and vigilance increased when units engaged in fission. Based on this evidence, P. nemaeus at Son Tra Nature Reserve appear to engage in daily fission-fusion activity which does not vary between seasons. Additionally, our data suggest that these primates may form multilevel societies made up of distinct units. However, future data including proximity pattern analyses are necessary for confirmation.

Dominance rank, female choice, and reproductive success in semi-free ranging adult male Japanese macaques (Macaca fuscata).

The assumed evolutionary advantage of dominance is increased reproductive success. However, the efficacy of dominance as a mating strategy may be interrupted by any number of variables including female choice, estrous synchrony, and mating by non-troop males. In Japanese macaques (Macaca fuscata), there is evidence both for and against dominance as conferring reproductive success for adult males, with many discussions pointing to the importance of female choice in governing reproductive success in certain populations. In this study, we aimed to evaluate dominance-based versus female choice-based male behavioral strategies and their impact on reproductive success. This study was conducted on a group of Japanese macaques at the Oregon National Primate Research Center. We collected a total of 512 h of behavioral data across two summer study periods in 2018 and 2019. We conducted 15-min focal follows with 1-min instantaneous scans on 17 adult males. Reproductive data were available from genetic records. Using principal components analysis (PCA), we identified males that cluster according to similar behavioral strategies. We then used analysis of variance (ANOVA) and non-parametric ANOVA on ranks to ascertain significant variation in rank and reproductive success between clusters. We found that males that clustered based on high directed aggression held higher rank than less-aggressive male clusters (F = 27.21, df = 4, p < .0001). However, less aggressive male clusters had higher reproductive success (F = 3.50, df = 4, p = .04). There was no variation between affiliative clusters in reproductive success (F = 1.77, df = 3, p = .15). The highly aggressive strategy is effective for attaining high rank, but only resulted in high reproductive success for a single male which likely necessitates alternative strategies. We suggest the operation of female choice within this population, with females preferentially mating with males who are not only affiliative but also less aggressive.

Ranging and territoriality in red-shanked doucs (Pygathrix nemaeus) at Son Tra Nature Reserve, Vietnam.

The home ranges of odd-nosed colobines as reported by a number of studies can vary greatly both within and between taxa. Evaluating Pygathrix, particularly within the context of the odd-nosed colobines, has proven challenging due to the comparable dearth of published data on all three species-Pygathrix nigripes, Pygathrix cinerea, and Pygathrix nemaeus. This dearth has also proven challenging when assessing territoriality within Pygathrix and contextualizing this behavior to odd-nosed colobines. For this study, we set out to assess home range, daily path length, and territoriality in a population of P. nemaeus at Son Tra Nature Reserve in Vietnam. We collected a total of 259 h of behavioral data using 10-min focal follows with 1-min instantaneous scans, including GPS recordings for each follow. We collected phenology and weather data, which were then used with Pearson's correlations to test whether ecological factors influenced variation in ranging or daily path length at both the monthly and seasonal levels. Finally, we used the Mitani-Rodman defendability index to assess territoriality in P. nemaeus. We found that our population of P. nemaeus had a home range of 36.00 ha with a core area of 8.02 ha and an average daily path length of 509.11 ± 244.87 m. Daily path length was correlated to precipitation such that daily path lengths were shorter during increases in rainfall (r(17) = 0.52, p = .03). These measures resulted in a defendability index of 0.75, suggesting that the population of P. nemaus on Son Tra does not travel far enough each day to actively defend territorial borders.

Variation in Adult Male-Juvenile Affiliative Behavior in Japanese Macaques (Macaca fuscata).

Adult males of some primate species are known to positively interact with juveniles. In cases where paternal certainty is high, these behaviors have been largely attributed to the paternal investment hypothesis. Males have also been observed to interact with nonkin juveniles, which has often been explained in terms of mating effort. Here, we examined variation in adult male-juvenile affiliation in semi-free ranging Japanese macaques (Macaca fuscata) at the Oregon National Primate Research Center against possible influencing factors such as age, dominance rank, and female affiliation, and we also tested for fitness benefits. We conducted 154 h of focal observations of 14 adult males from June to September 2018. Males differed significantly in their rate of juvenile-directed affiliation, but not in their fitness in terms of number of offspring. There was a significant positive correlation between rank and age in the group, indicating that, in this group, rank does not conform to the classic inverted-U pattern observed elsewhere in this species. Although there was a significant positive correlation between rank and juvenile-directed affiliation, the highest-ranking male had few offspring and exhibited little juvenile-directed affiliation. These results suggest little to no preliminary support for either the paternal investment or mating effort hypotheses as explanations for juvenile-directed affiliation. This study suggests that there are multiple behavioral strategies for older males that may influence reproductive success.

Behavioral changes following alterations in the composition of a captive bachelor group of western lowland gorillas (Gorilla gorilla gorilla).

Western lowland gorillas (Gorilla gorilla gorilla) in zoos are housed in family or bachelor groups to maximize social opportunities. While wild bachelor groups are transient, all-male groups in zoos may be maintained for many years. Captive bachelor groups need to be carefully monitored, particularly during periods of demographic transition, due to the possibility for escalating aggression. We examined behavioral changes in a bachelor group at the Saint Louis Zoo following two significant alterations in group composition: (1) the introduction of two immature related males in 2011 and (2) the death of the dominant silverback in 2015. Behavioral data were collected on group members using 15 min focal observations with 30-s instantaneous scans, totaling 185.25 hr for the first transition and 115.25 hr for the second transition (with equal effort in baseline and transition periods). We found that the addition of the two subadult males resulted in a significant increase in affiliation in the group often initiated by these new younger individuals, while the rate of abnormal and aggressive behavior did not change significantly. The rate of abnormal, aggressive, or affiliative behavior in the group also did not change significantly following the death of the dominant silverback. Overall, we conclude that this bachelor group and potentially others can endure possibly destabilizing social transitions and remain cohesive social units. Behavioral changes will result from transitions, but the effect on individuals or impact on the group will vary. Therefore, longitudinal monitoring of bachelor groups will be essential for effective management during any major changes.