This paper summarises recent findings on the causation of stereotypic behaviours and other abnormal repetitive behaviours (ARBs) in captive animals: primarily motivational frustration and/or brain dysfunction, with possible contributory roles also being played by habit-formation and 'coping' effects. We then review the extent to which ARBs occur in zoos and similar, estimating that at least 10 000 captive wild animals are affected worldwide. We argue for 'zero tolerance' of such ARBs, because stress and poor welfare raise ethical issues, while abnormal behavioural phenotypes and possibilities of impaired brain development challenge both the indirect (e.g. educational) and the direct, intrinsic conservation value of affected animals. We then consider five potential means by which ARBs may be tackled: genetic selection; pharmacological treatment; the reinforcement of alternative behaviours; punishment; and environmental enrichment. All except punishment have potentially useful roles to play, but enrichment is the preferred approach: it is most likely to tackle the problems underlying stereotypic behaviours, and thence to improve both welfare and behaviour with few unwanted side-effects. Nevertheless, in zoos, environmental enrichment to date has only had partial success, with no study managing to abolish ARBs in all its subjects-suggesting either that the enrichments currently being used are never quite optimal, or that by the time they are tackled, ARBs have become resistant to change. We suggest some ways in which the effectiveness of enrichments may be enhanced; propose that certain properties of ARBs may usefully help evaluate their likely 'treatability'; and emphasise that if improving welfare is more important than just reducing ARB, then additional measures are needed in order to first, reliably identify those individuals most at risk from poor welfare, and then, to fully evaluate the welfare impact of enrichments. This paper also emphasises, with examples, the enormous potential value of zoo-derived data for helping understand

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In captivity, some species often seem to thrive, while others are often prone to breeding problems, poor health, and repetitive stereotypic behaviour. Within carnivores, for instance, the brown bear, American mink and snow leopard typically adapt well to captivity and show few signs of poor welfare, while the clouded leopard and polar bear are generally hard to breed successfully and/or to prevent from performing abnormal behaviour. Understanding the fundamental source of such differences could enable reproductive success and behavioural normalcy to be improved in zoos and breeding centres, by increasing the appropriateness of the enclosure designs and environmental enrichments offered particular species, and by allowing these to be offered pre-emptively instead of reactively. Here, we demonstrate that a significant proportion of the variation in apparent welfare between captive carnivore species stems from specific aspects of natural behaviour. We tested pre-existing hypotheses that species-typical welfare is predicted by natural hunting behaviour, general activity levels, ranging, or territorial patrolling (all activities that are constrained in captivity), by collating data on median stereotypy levels and infant mortality for multiple captive species, and then regressing these against median values for the relevant aspects of natural behavioural biology (e.g. hunts per day, proportion of flesh in the diet, home-range size, etc.). Our results revealed that instead of relating to foraging (e.g. hunting), as often assumed, carnivore stereotypy levels are significantly predicted by natural ranging behaviour (e.g. home-range size and typical daily travel distances). Furthermore, naturally wide-ranging lifestyles also predicted relatively high captive infant mortality rates. These results suggest that enclosure designs and enrichments focussing on carnivores' ranging tendencies (e.g. providing more space, multiple den sites, greater day-to-day environmental variability/novelty, and/or more control over exposure to aversive or rewarding stimuli) could be particularly effective means of improving welfare; and also, that targeting such enrichment programmes on wide-ranging species, before problems even emerge, might effectively pre-empt their development. Alternatively, species with relatively small ranges could instead be made the focus of future collections and breeding programmes, zoos phasing out wideranging carnivores in favour of those species inherently more suited to current or readily achievable enclosure sizes and enrichment regimes. #

14Keeping elephants in zoos is extremely costly, yet does not yield self-sustaining 15 populations. In Europe, which holds c. half the global zoo elephant population, a long-16 term decline of c.10% per year is expected in both species, if reliant on zoo-bred animals 17 under historically prevailing conditions. Fitness in zoos is compromised in several ways. 29The welfare implications of captivity, and the relative cost and effectiveness of ex versus 30 in situ conservation, determine the value of captive breeding for any given species. 31Several species apparently thrive in zoo conditions (1), and captive breeding has saved 32 some from extinction (2, 3). However, ex situ conservation is typically costlier than in situ 46$53 million per year in maintenance and, in the last decade, over $540 million in facility 47 upgrades worldwide (see Table S1

General rightsThis document is made available in accordance with publisher policies. Please cite only the published version using the reference above. Full terms of use are available: http://www.bristol.ac.uk/pure/about/ebr-terms 1 Stereotypic route-tracing in captive Carnivora is predicted by species-typical 1 home range sizes and hunting styles 2 3 Abstract 4In captive conditions (e.g. zoos), some Carnivora species typically show negligible 5 stereotypic behaviour (SB) and reproduce successfully, while others tend to 6 reproduce poorly and be very stereotypic. We used comparative methods to identify 7 species-level risk factors for SB and captive infant mortality (CIM). Candidate 8 predictor variables were natural ranging behaviour, territoriality, aspects of natural 9 foraging, wild activity levels, cranial volume, and IUCN Red List status. Previous 10 research had identified naturally long daily travel distances, and being large-bodied 11 and wide-ranging, as SB risk factors. We nearly doubled the size of this original SB 12 database, and then imposed stricter quality controls (e.g. on minimum sample sizes 13 for inclusion). Analysing the resulting 23-species dataset confirmed naturally large 14 ranges and travel distances as risk factors. It also showed that the range size effect: 15 is independent of body mass (although body mass and range size together predicted 16 SB most strongly); is stronger for stereotypic route-tracing (e.g. pacing) than for all 17 SB forms combined; and explains the apparent daily travel distance effect (which 18 vanished when range size was controlled for). Furthermore, a new finding emerged: 19 that naturally long chase distances during hunts also predicted more severe route-20 tracing. Turning to CIM, previous research had also identified naturally long travel 21 distances and large home ranges as risk factors. We failed to replicate this, or to 22 confidently identify any species-level risk factor (despite CIM significantly varying 23 between related species, at least for Canidae and Ursidae

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We previously reported that African (Loxodonta africana) and Asian (Elephas maximus) female elephants in European zoos have shorter adult lifespans than protected conspecifics in range countries. This effect was the cause of greatest concern in Asian elephants, and risk factors within this species included being zoo-born, transferred between zoos, and possibly removed early from the mother. Here, we investigate these risk factors further; assess fecundity and sustainability in European zoos; and propose testable hypotheses as to the causes of these animals’ problems. Although imported wild-born Asian elephants live longer than zoo-born conspecifics, being imported when juvenile or adult appears no more protective than being imported in infancy, suggesting that the benefits of being wild- rather than zoo-born are conferred early in life. Zoo-born Asian neonates are significantly heavier than those born to working animals in range countries, with a possible tendency to be fatter. In zoos, African elephants have tended to be removed from their mothers at older ages than young Asians, and were also transferred between zoos significantly less often: factors that could possibly underlie this species’ lower calf losses and improving adult survivorship in Europe. Both species have low fecundity in European zoos compared to in situ populations, and are not self-sustaining, declining at approximately 10% per annum if reliant on captive-bred females under historically prevailing conditions. Data from other species suggest that stress and/or obesity are parsimonious explanations for the suite of problems seen. We recommend specific screens for testing these hypotheses, and for potentially identifying vulnerable individuals within the extant zoo populations.

We investigated the survival of 57 rehabilitated juvenile Tawny Owls (Strix aluco) that were 'hard released' (without the provision of a release aviary or support food) by means of radio tracking. The birds were released in the month of August in three consecutive years: 2005, 2006 and 2007, in the counties of Somerset and Hampshire, United Kingdom. Tracking of the owls was successfully carried out for between three and 160 days. Mortality was recorded for 16 birds (28%). The transmitter was shed by 24 (42%) owls, the signal was lost for 12 (21%) and tracking was ceased for five (9%) owls. Survival of the owls was compared with results from previous studies on wild Tawny Owls and also rehabilitated 'soft released' Tawny Owls (released with provision of food and shelter after release) and found to be similar. This study suggests that employing costly and time-consuming soft release techniques may be unnecessary for juvenile Tawny Owls as their survival is not significantly reduced using hard-release methods. Measuring post-release success of rehabilitated birds of prey is discussed in relation to benchmarks used in previous studies.