Assessing the Welfare of Dairy Cattle

This article attempts to determine the effects of environment (captive or wild) and a simple form of environmental enrichment on the behavior and physiology of a nonhuman animal. Specifically, analyses first compared behavioral budgets and stereotypic behavior of captive coyotes (Canis latrans) in kennels and pens to their counterparts in the wild. Second, experiments examined the effect of a simple form of environmental enrichment for captive coyotes (food-filled bones) on behavioral budgets, stereotypies, and corticosteroid levels. Overall, behavioral budgets of captive coyotes in both kennels and pens were similar to those observed in the wild, but coyotes in captivity exhibited significantly more stereotypic behavior. Intermittently providing a bone generally lowered resting and increased foraging behaviors but did not significantly reduce stereotypic behavior or alter corticosteroid levels. Thus, coyote behavior in captivity can be similar to that exhibited in the wild; in addition, although enrichment can affect proportions of elicited behaviors, abnormal behaviors and corticosteroid levels may require more than a simple form of environmental enrichment for their reduction.     

Environmental Enrichment Effect on Fecal Glucocorticoid Metabolites and Captive Maned Wolf (Chrysocyon brachyurus) Behavior

Environmental enrichment is a technique that may reduce the stress of nonhuman animals in captivity. Stress may interfere with normal behavioral expression and affect cognitive decision making. Noninvasive hormonal studies can provide important information about the stress statuses of animals. This study evaluated the effectiveness of different environmental enrichment treatments in the diminution of fecal glucocorticoid metabolites (stress indicators) of three captive maned wolves (Chrysocyon brachyurus). Correlations of the fecal glucocorticoid metabolite levels with expressed behaviors were also determined. Results showed that environmental enrichment reduced fecal glucocorticoid metabolite levels. Furthermore, interspecific and foraging enrichment items were most effective in reducing stress in two of the three wolves. No definite pattern was found between behavioral and physiological responses to stress. In conclusion, these behavioral and physiological data showed that maned wolves responded positively from an animal well being perspective to the enrichment items presented.     

The effects of foraging demand on social interactions in a laboratory group of Bonnet Macaques

The relationship between foraging demand and social behavior was experimentally studied in a laboratory group of bonnet macaques. Fourteen adult animals were housed in a large outdoor enclosure containing three shallow gravelfilled circular containers that served as the foraging sites. During the experimental foraging sessions raisins were placed in the containers and the social and foraging behaviors of the group were observed for 50 min following the distribution of raisins. Three types of foraging conditions were inter-spersed with one another on different test days: (1) surface load— raisins placed on top of the gravel; (2) buried load— raisins hidden underneath the gravel; and (3) sham load— no raisins placed at the foraging sites. Three basic foraging patterns, defined along a temporal dimension, were seen. One group of animals completed 50% of their total foraging by the end of the first 15 min. A second group foraged more steadily through the session. A third group foraged late, completing 50% of their foraging during the last half of the session. The foraging patterns were similar in the buried and surface condition, although the patterns were more compressed during the surface condition. More aggression and more avoidance of other animals occurred in the buried condition than in the surface condition. Very little foraging occurred during the sham condition. There was no clear relationship between the patterns of interaction during foraging and nonforaging observation sessions. The results suggest the value of manipulative laboratory studies in examining the relationship between ecological variables and social behavior in nonhuman primates.     

Single- or Pair-Housed: Which Is Better for Captive Southern Tamanduas?

The captive environment can limit some important behavioral options for nonhuman animals, which often results in decreased welfare. The companion of a conspecific can be a source of complexity in captivity, but this aspect has received little attention for solitary species. This study investigated the effects of two different housing conditions on the behavior of a solitary species, the Southern tamandua (Tamandua tetradactyla). Six individuals housed singly and four housed in pairs in two Brazilian zoological institutions were observed for 10 days from 09:00h to 17:00h during the rainy season. Each individual was observed for 26:40 hours. Compared with singly housed tamanduas, paired animals were less inactive (single = 74.62 ± 5.2%, paired = 28.51 ± 0.04% of the records), foraged and explored more (single = 11.07 ± 3.0%, paired 31.48 ± 0.01% of the records), and exhibited a diversity of behaviors 102.56% greater than single tamanduas. These results suggest that interactions with conspecifics in captivity, even for solitary animals, can promote the performance of desirable behaviors, which in several species has been correlated with improved welfare.     

Conflict and reconciliation behavior trends of the bottlenose dolphin (Tursiops truncatus)

Wild bottlenose dolphin (Tursiops truncatus) populations display societal structures characterized by numerous and frequent changes in group composition, complex social relationships, and high levels of cooperation, attributes also observed in human and nonhuman primate cultures. Maintaining social relationships under such elemental conditions can frequently create conflict—and the opportunity for reconciliation—among group members. The conflict and reconciliation behavior patterns of the chimpanzee (Pan troglodytes) have been studied extensively; trends are well‐documented in the wild and in captivity. Apparent cultural similarities have prompted several analogous studies on wild and captive bottlenose dolphins. This research attempted to corroborate previous efforts by analyzing the social behavior of seven captive bottlenose dolphins to determine the effects of sex and age on the frequency of conflict and reconciliation, and to investigate the incidence of consolatory behavior within the group. A total of 3,428 interactions involving focal animals, 414 of which were conflict episodes, were documented during 261 hr of videotaped observations. Although the sample size precluded meaningful statistical evaluation of the influence of sex on conflict and reconciliation, participant age was a significant determinant of conflict frequency. Conversely, age did not impact frequency of reconciliation, which only occurred after 18% of all conflict interactions. Little to no definitive evidence of consolation was apparent within the study group. While results partially support the findings of previous dolphin reconciliation research, extensive behavioral studies of wild populations should be conducted before generating broad comparisons between human terms and nonhuman behavioral interactions. Zoo Biol 29:567–585, 2010. © 2009 Wiley‐Liss, Inc.     

Olfactory Enrichment in California Sea Lions (Zalophus californianus): An Effective Tool for Captive Welfare?

In the wild, California sea lions (Zalophus californianus) are exposed to a wide variety of sensory information, which cannot be replicated in captive environments. Therefore, unique procedures are necessary for maintaining physiological and psychological health in nonhuman animals in captivity. The effects of introducing natural scents to captive enclosures have been investigated in a variety of species, yet they have not been examined in marine mammals. This project explored the behavioral effect of scent added to the environment, with the goal of improving the welfare of sea lions in captivity. Two scent types were introduced: (a) natural scents, found in their native environment, and (b) non-natural scents, not found in their native environment. This study examined not only scent enrichment but also the possible evolutionary underpinnings of pinniped olfaction. Scent enrichment was found to significantly impact sea lion behavior as demonstrated by a reduction in pattern swimming, an increase in habitat utilization, and a reduction in stereotypical behavior. However, there were no differences in behavior between natural and non-natural scent conditions.     

Maladaptive behavioural phenotypes in captive‐reared darters (Etheostoma caeruleum,Storer 1845)

The foraging and anti‐predator behaviour of captive‐reared rainbow darters (Etheostoma caeruleum) was compared to their wild‐caught counterparts. Wild‐caught darters responded with appropriate anti‐predator behaviour (reduced foraging activity) when exposed to alarm cues (e.g. stimuli from damaged skin) from both wild‐caught and captive‐reared darters, indicating that the diet in captivity did not inhibit the production of alarm cues. Captive‐reared individuals did not change their level of activity when exposed to alarm cues; however, their significantly lower baseline activity (movement and prey consumption) makes it unclear as to whether they actually failed to recognize risk. Regardless, captive‐reared darters showed little motivation to feed when food became available (i.e. they made few movements to obtain food) and appeared impervious to chemical cues indicating risk. Exposing captive‐reared individuals to both semi‐natural foraging opportunities and predator‐recognition training before their release is recommended.     

Thermal biology,torpor and behaviour in sugar gliders: a laboratory-field comparison

Most studies on animal physiology and behaviour are conducted in captivity without verification that data are representative of free-ranging animals. We provide the first quantitative comparison of daily torpor, thermal biology and activity patterns, conducted on two groups of sugar gliders (Petaurus breviceps, Marsupialia) exposed to similar thermal conditions, one in captivity and the other in the field. Our study shows that activity in captive gliders in an outdoor aviary is restricted to the night and largely unaffected by weather, whereas free-ranging gliders omit foraging on cold/wet nights and may also forage in the afternoon. Torpor occurrence in gliders was significantly lower in captivity (8.4% after food deprivation; 1.1% for all observations) than in the field (25.9%), mean torpor bout duration was shorter in captivity (6.9 h) than in the field (13.1 h), and mean body temperatures during torpor were higher in captivity (25.3°C) than in the field (19.6°C). Moreover, normothermic body temperature as a function of air temperature differed between captive and free-ranging gliders, with a >3°C difference at low air temperatures. Our comparison shows that activity patterns, thermal physiology, use of torpor and patterns of torpor may differ substantially between the laboratory and field, and provides further evidence that functional and behavioural data on captive individuals may not necessarily be representative of those living in the wild.     

Predicting release success of captive‐reared loggerhead shrikes (Lanius ludovicianus) using pre‐release behavior

We assessed whether behavioral markers could be used to evaluate pair compatibility and predict pair bond success of captive‐reared San Clemente loggerhead shrikes (Lanius ludovicianus mearnsi) released into the wild. Potential breeding pairs of shrikes were introduced at the Zoological Society of San Diego's captive‐breeding facility and then moved to release cages located in suitable but unoccupied habitat. Courting pairs were affected negatively by the disturbance of translocation to a new location and generally needed a few days to reinitiate pair bonding in the release enclosures. We separated and returned pairs to captivity when intra‐pair aggression or cessation of all courtship behavior occurred; all other pairs were released into the wild. The rate of nest approaches was the best marker to predict a successful release into the wild (i.e., pairs that remained near the release site and attempted to breed). Additionally, all experienced breeding pairs (i.e., pairs with males and females with prior captive breeding experience) exhibiting nesting behavior were successful, although previous experience alone does not ensure post‐release breeding. Results from this study indicate the importance of assessing behaviors of individuals paired for population augmentation. In using behavioral cues, identifying pairs with a low probability of success is possible, and replacing these pairs with individuals that have a higher likelihood of post‐release success can occur. This strategy will be important to efficiently restore imperiled populations of endangered species while working within temporal and financial constraints. Zoo Biol 0:1–12, 2005. © 2005 Wiley‐Liss, Inc.     

Captive killer whale (Orcinus orca) survival

Killer whales (Orcinus orca) were first placed into captivity in 1961 and are now found in theme parks around the world. Despite successful breeding of captive killer whales since 1985 there is growing concern for their welfare in captivity, which often includes claims of poor survival. We employed Kaplan‐Meier and Cox Proportional hazards models and annual survival rate analyses on 201 captive killer whales to discern how sex, facility (U.S. vs. foreign), captive‐born vs. wild‐captured, pre‐ vs. post‐1 January 1985, and animal age upon entering captivity affect survival. Overall median survival estimate was 6.1 yr, with no difference between male and female survival. Killer whales in U.S. facilities (12.0 yr) demonstrated a significantly higher median survival than those in foreign facilities (4.4 yr), as did whales entering captivity post‐1 January 1985 (11.8 yr) vs. those entering prior to 1 January 1985 (3.9 yr). Median survival for captive‐born (14.1 yr) was significantly higher than wild‐captured killer whales (5.5 yr), though the two failed to differ among the post‐1 January 1985 cohort. Facility location and pre‐ vs. post‐1 January 1985 were predictors of the hazard rate. Survival of captive killer whale cohorts has generally improved through time, although survival to age milestones are poor when compared to wild killer whales.     

Tailored Enrichment Strategies and Stereotypic Behavior in Captive Individually Housed Macaques (Macaca spp.)

The welfare of nonhuman animals in captivity is widely dependent on the natural psychological, physical, and behavioral needs of the animals and how adequately these needs are met. Inability to engage in natural behaviors can lead to chronic stress and expression of stereotypic behavior. The majority of research on decreasing stereotypic behavior in captivity addresses problems at the group level and does not account for individual variability in each animal's needs, history, and preferences. This study combined physiological and behavioral measures of well being to comprehensively assess the unique needs of individually housed captive macaques (Macaca spp.) with the aim of developing tailored welfare strategies. Behavioral and hormonal data were collected under 2 conditions: baseline and individualized enrichment. The results showed a significant decrease in stereotypic behavior under the enrichment condition. Additionally, 7 out of 9 individuals showed a decrease in fecal glucocorticoid (stress hormone) levels, indicating a reduction in hypothalamic-pituitary-adrenal activity. Addressing welfare on an individual, rather than group, level allows for a better overall characterization of well being and maximizes the probability of improving the welfare of each animal.     

Rearing environment influences boldness and prey acquisition behavior,and brain and lens development of bull trout

Animals reared in barren captive environments exhibit different developmental trajectories and behaviors than wild counterparts. Hence, the captive phenotypes may influence the success of reintroduction and recovery programs for threatened and endangered species. We collected wild bull trout embryos from the Metolius River Basin, Oregon and reared them in differing environments to better understand how captivity affects the bull trout Salvelinus confluentus phenotype. We compared the boldness and prey acquisition behaviors and development of the brain and eye lens of bull trout reared in conventional barren and more structurally complex captive environments with that of wild fish. Wild fish and captive reared fish from complex habitats exhibited a greater level of boldness and prey acquisition ability, than fish reared in conventional captive environments. In addition, the eye lens of conventionally reared bull trout was larger than complex reared captive fish or same age wild fish. Interestingly, we detected wild fish had a smaller relative cerebellum than either captive reared treatment. Our results suggest that rearing fish in more complex captive environments can create a more wild-like phenotype than conventional rearing practices. A better understanding of the effects of captivity on the development and behavior of bull trout can inform rearing and reintroduction programs though prediction of the performance of released individuals.     

Anointing variation across wild capuchin populations: a review of material preferences, bout frequency and anointing sociality in Cebus and Sapajus

The frequency of anointing bouts and the materials used for self- and social anointing vary across capuchin species in captivity, but there is little published data on capuchin anointing in the wild. Here we present previously unpublished data on anointing behaviors from capuchin monkey populations at ten different field sites and incorporate these data into a review of the anointing literature for captive and wild capuchins. Using a comparative phylogenetic framework, we test four hypotheses derived primarily from captive literature for variation in anointing between wild untufted capuchins (Cebus) and tufted capuchins (Sapajus), including that (1) the frequency of anointing is higher in Cebus, (2) Cebus uses a higher proportion of plant species to insect species for anointing compared with Sapajus, (3) anointing material diversity is higher in Cebus, and (4) social indices of anointing are higher in Cebus. We found that wild Cebus anoints more with plant parts, including fruits, whereas wild Sapajus anoints more with ants and other arthropods. Cebus capucinus in particular uses more plant species per site for anointing compared with other capuchins and may specialize in anointing as an activity independent from foraging, whereas most other capuchin species tend to eat the substances they use for anointing. In agreement with captive studies, we found evidence that wild Cebus anoints at a significantly higher frequency than Sapajus. However, contrary to the captive literature, we found no difference in the range of sociality for anointing between Cebus and Sapajus in the wild. We review anointing in the context of other Neotropical primate rubbing behaviors and consider the evidence for anointing as self-medication; as a mechanism for enhanced sociality; and as a behavioral response to chemical stimuli.     

Preservation of behavior after fifteen years of isolation: comparisons of wild and captive endangered pupfish in their natural habitat

Endangered species are often maintained in captivity to serve as a safeguard in the event of an extirpation of natural populations. However, wild and captive populations can rapidly diverge in genetic and phenotypic characteristics, including behavior, when maintained in isolation and under different environmental conditions. Here, we compare two populations of the endangered Leon Springs pupfish (Cyprinodon bovinus): a captive population maintained at the Southwestern Native Aquatic Resources and Recovery Center (SNARRC) and the only remaining wild population. Wild fish in the natural Texan Diamond Y desert spring are remnants of a past reintroduction of captive SNARRC fish, which occurred in 1998. The fifteen years of isolation have led to genetic and morphologic differences between the two populations. To assess the behavioral implications of such divergence, we released previously captive C. bovinus into natural habitat and quantified behaviors under natural conditions and in semi-controlled mesocosms and then replicated these conditions with the established natural population. Both populations exhibited similar levels of reproduction, foraging, and agonistic behavior. Despite divergence in genetic and morphometric characteristics, overall behavioral patterns of C. bovinus remained consistent in similar environments. This stability suggests that captive C. bovinus could again be successful establishing in their natural habitat. It is our hope that this investigation will help focus future conservation efforts towards monitoring the persistence of reintroduced C. bovinus in addition to providing a framework for other recovery plans reintroducing captive stock into ancestral habitats.     

Assessing the effects of cognitive experiments on the welfare of captive chimpanzees (Pan troglodytes) by direct comparison of activity budget between wild and captive chimpanzees

We investigated the effects of cognitive experiments by direct comparison of activity budgets between wild and captive chimpanzees. One goal of captive management is to ensure that the activity budgets of captive animals are as similar as possible to those of their wild counterparts. However, such similarity has rarely been achieved. We compared the activity budget among three groups of chimpanzees: wild chimpanzees in Bossou (Guinea, n = 10), and captive chimpanzees who participated in cognitive experiments (experimental chimpanzees, n = 6) or did not participate in the experiments (nonexperimental chimpanzees, n = 6) at the Primate Research Institute (Japan). The experimental chimpanzees voluntarily participated in computer‐controlled cognitive tasks and small pieces of fruits were provided as rewards. The data from captivity were obtained on the experimental days (weekdays) and nonexperimental days (weekends). In both study sites, we followed each chimpanzee from about 7 a.m. until the time when chimpanzees started to rest in the evening. The behaviors were recorded every 1 min. The results showed that on weekdays, feeding time and resting time of the experimental chimpanzees were almost the same as those of wild chimpanzees. However, for the nonexperimental chimpanzees, feeding time was significantly shorter and resting time was longer than those of the wild chimpanzees. In contrast, no difference was found in feeding time or resting time of the two groups of captive chimpanzees on weekends. The results suggested that the cognitive experiments worked as an efficient method for food‐based enrichment. Am. J. Primatol. 73:1231–1238, 2011. © 2011 Wiley Periodicals, Inc.     

Dynamics of genetic adaptation to captivity

Many species require captive breeding to savethem from extinction, with reintroduction intothe wild being the eventual aim of mostprograms. Adaptation to captive environmentstypically results in reduced fitness under wildconditions. Consequently, unintentionaladaptation during captive breeding programs mayseriously compromise the success ofreintroduction programs. However, there islittle experimental evidence on the rate orextent of adaptation for captive populationsmaintained under benign captive conditions forextended periods of time. To investigate thedynamics of genetic adaptation to captivity,large captive populations of Drosophilamelanogaster were assessed for relativefitness under captive conditions for up to 87generations in captivity. Captive fitnessincreased to 3.33 times the initial fitnessover 87 generations. The pattern of adaptationwas curvilinear, with an exponential curveproviding the best fit. Fitness reached 25% ofits maximum within 6 generations, 50% within15 generations, 75% within 31 generations and95% within 67 generations. The model predictedthat the asymptotic level of fitness reachedwould be 3.38 times the initial fitness. Thus,very large genetic adaptations to captivity mayoccur under relatively benign captiveconditions. Captive populations destined forreintroduction need to be managed to minimisegenetic adaptation to captivity.     

Social behavior of captive,group-living Orangutans

The behavior of orang-utans (Pongo pygmaeus)was observed in two captive groups (one adult group, one juvenile group). Activity profiles,animal interactions, and compartmental spatial use for both adult-group and juvenile-group individuals were recorded over a 9-month period. Behavioral repertoires for both groups included large amounts of social activity. Equivalent amounts of social activity were found for each group. The social behavior of juvenile animals involved more active behavior such as play. The social behavior of the adult animals was more subtle, involving social monitoring and allogrooming. These results indicate that orangutans, at least when group-living in captivity, exhibit the potential to display social behavior which is apparently of greater frequency and complexity than that which has been observed in the wild. These findings suggest that the solitary behavior of wild orangutans is not a necessary characteristic of orangutan behavior. Under different environmental conditions orangutans appear to readily adapt socially, and, like other nonhuman primates,they have the capacity to exhibit complex and subtle social behavior. This report is based on part of a senior thesis submitted by Sara D. Edwards     

Do captive waterfowl alter their behaviour patterns during their flightless period of moult?

Many different behavioural changes have been observed in wild waterfowl during the flightless stage of wing moult with birds frequently becoming inactive and reducing time spent foraging. Increased predation risk, elevated energetic demands of feather re-growth and restriction of foraging opportunities are thought to underlie these changes. By studying captive populations of both a dabbling and a diving duck species at the same site, we determined whether captive birds would reflect the behavioural responses of wild waterfowl to moult. The time-budgets of 42 Common Eiders, Somateria mollissima, (a diving duck) and 18 Garganeys, Anas querquedula, (a dabbling duck) were recorded during wing moult (July–August) and non-moult (January) with behaviour recorded under six categories. Despite captivity providing a low predation risk and constant access to food, birds altered their behaviour during the flightless period of wing moult. Time allocated to foraging and locomotion decreased significantly during moult compared to non-moult periods, while resting time increased significantly. Moulting Eiders underwent a greater reduction in time spent foraging and in locomotion compared with Garganeys, which is likely to be in response to a higher energetic cost of foraging in Eiders. It is possible that increased resting in both diving and dabbling ducks reduces their likelihood of detection by predators, while allowing them to remain vigilant. We demonstrate that there is much potential for using captive animals in studies that can augment our knowledge of behaviours of free-living conspecifics, the former being a hitherto under-exploited resource.     

Effects of captivity,diet, and relocation on the gut bacterial communities of white‐footed mice

Microbes can have important impacts on their host's survival. Captive breeding programs for endangered species include periods of captivity that can ultimately have an impact on reintroduction success. No study to date has investigated the impacts of captive diet on the gut microbiota during the relocation process of generalist species. This study simulated a captive breeding program with white‐footed mice (Peromyscus leucopus) to describe the variability in gut microbial community structure and composition during captivity and relocation in their natural habitat, and compared it to wild individuals. Mice born in captivity were fed two different diets, a control with dry standardized pellets and a treatment with nonprocessed components that reflect a version of their wild diet that could be provided in captivity. The mice from the two groups were then relocated to their natural habitat. Relocated mice that had the treatment diet had more phylotypes in common with the wild‐host microbiota than mice under the control diet or mice kept in captivity. These results have broad implications for our understanding of microbial community dynamics and the effects of captivity on reintroduced animals, including the potential impact on the survival of endangered species. This study demonstrates that ex situ conservation actions should consider a more holistic perspective of an animal's biology including its microbes.     

Nutritional and behavioral effects of gorge and fast feeding in captive lions

Nonhuman animals in captivity manifest behaviors and physiological conditions that are not common in the wild. Lions in captivity face problems of obesity, inactivity, and stereotypy. To mediate common problems of captive lions, this study implemented a gorge and fast feeding schedule that better models naturalistic patterns: African lions (Panthera leo) gradually adapted from a conventional feeding program to a random gorge and fast feeding schedule. Digestibility increased significantly and food intake and metabolizable energy intake correspondingly decreased. Lions also showed an increase in appetitive active behaviors, no increase in agonistic behavior, and paced half as frequently on fast days as on feeding days. Thus, switching captive lions to a gorge and fast feeding schedule resulted in improved nutritional status and increased activity.