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

This paper reviews a way of investigating health and welfare problems in captive wild animals (e.g., those in zoos, aviaries, aquaria, or aquaculture systems) that has great potential, but to date has been little used: systematically comparing species with few or no health and welfare issues to those more prone to problems. Doing so empirically pinpoints species-typical welfare risk and protective factors (such as aspects of their natural behavioral biology): information which can then be used to help prevent or remedy problems by suggesting new ways to improve housing and husbandry, and by identifying species intrinsically best suited to captivity. We provide a detailed, step-by-step "how to" guide for researchers interested in using these techniques, including guidance on how to statistically control for the inherent similarities shared by related species: an important concern because simple, cross-species comparisons that do not do this may well fail to meet statistical assumptions of non-independence. The few relevant studies that have investigated captive wild animals' welfare problems using this method are described. Overall, such approaches reap value from the great number and diversity of species held in captivity (e.g., the many thousands of species held in zoos); can yield new insights from existing data and published results; render previously intractable welfare questions (such as "do birds need to fly?" or "do Carnivora need to hunt?") amenable to study; and generate evidence-based principles for integrating animal welfare into collection planning.

Understanding why some species thrive in captivity, while others struggle to adjust, can suggest new ways to improve animal care. Approximately half of all Psittaciformes, a highly threatened order, live in zoos, breeding centres and private homes. Here, some species are prone to behavioural and reproductive problems that raise conservation and ethical concerns. To identify risk factors, we analysed data on hatching rates in breeding centres (115 species, 10 255 pairs) and stereotypic behaviour (SB) in private homes (50 species, 1378 individuals), using phylogenetic comparative methods (PCMs). Small captive population sizes predicted low hatch rates, potentially due to genetic bottlenecks, inbreeding and low availability of compatible mates. Species naturally reliant on diets requiring substantial handling were most prone to feather-damaging behaviours (e.g. self-plucking), indicating inadequacies in the composition or presentation of feed (often highly processed). Parrot species with relatively large brains were most prone to oral and whole-body SB: the first empirical evidence that intelligence can confer poor captive welfare. Together, results suggest that more naturalistic diets would improve welfare, and that intelligent psittacines need increased cognitive stimulation. These findings should help improve captive parrot care and inspire further PCM research to understand species differences in responses to captivity.

Abnormal repetitive behaviours (ARBs) are associated with past, or present, welfare problems and are common elements of the behavioural repertoire of some captive animals, including birds. Millions of birds from hundreds of species are held in various captive settings, yet most avian ARB research to-date focuses on just a handful of these. Therefore, our knowledge of ARBs and, by implication, welfare, of a taxonomically diverse range of avian species is poorly understood. The purpose of this review is to begin to address this by providing a useful overview of ARBs across captive avian species. Taking advantage of the research effort on well-studied species, we pool current findings relating to avian ARBs into a coherent framework, highlight gaps in understanding, and use this to give a reference point for future research in both these and other species. We adopt Tinbergen's 'Four Questions' approach to comprehensively consider ARBs from each of his four perspectives. We begin with presenting studies on ARB development, describe how physiological predispositions and early-life housing and experiences impact ARB risk in later life. Next we outline internal causal triggers for ARBs, such as the effects of neurotransmitters, hormones, and dietary deficiencies, and discuss external, environmental triggers for ARBs. In the evolution section, we detail the influence of species' evolutionary history on ARB, and use findings from early molecular studies on laying hens to discuss heritability and genes associated with ARB. The benefits of using cross-species studies to determine underlying evolutionary drivers of ARBs are also illustrated with an example from Psittaciformes. In discussing ARB 'function', we make two tentative suggestions for potential examples of ARB performance allowing a bird to cope, and also consider situations where ARB may be functionless. We then summarise, and discuss, these four interacting perspectives on avian ARBs. To finish, the benefits of Tinbergen's approach are shown in a worked example of an ARB in one species, demonstrating how this valuable framework leads to the most complete understanding of ARB. Thus, by utilising Tinbergen's Four Questions, our review provides a platform for future