The endangered red panda (Ailurus fulgens) is held in zoos worldwide. The aim of this study was to examine how red pandas are kept and managed in captivity and to compare it with the management guidelines. Sixty-nine zoos, mainly from Europe but also from North America and Australia/New Zealand, responded to our survey. The results revealed that in general zoos follow the management guidelines for most of the investigated issues. The average enclosure is almost four times larger than the minimum size recommended by the management guidelines, although seven zoos have smaller enclosures. About half the zoos do not follow the guidelines concerning visitor access and number of nest boxes. Other issues that may compromise animal welfare include proximity of neighboring carnivore species and placement of nest boxes. Zoo Biol 29: 732-740, 2010. (C) 2010 Wiley-Liss, Inc.

Conservation breeding and reintroduction into the wild can only be an effective management tool if behaviours essential for a life in the wild are maintained in captivity. The aim of this study was to investigate how a protected captive environment influences antipredator behaviour over generations. The red junglefowl (Gallus gallus) was used as a case study. Birds from two different captive populations were followed over four generations. In the last three generations, all birds were hatched and reared in the same indoor settings. Antipredator behaviour was measured in each generation in a standardised test where the birds were exposed to a simulated predator attack. The test was divided into three parts: pre-exposure period, exposure and post-exposure periods. There was an interaction effect between Population and generation (F-3.129 = 4.84, P < 0.01) on behaviour during the pre-exposure period, suggesting that the birds "baseline" agitation level may have been altered differently in the two populations. Population differences were also found during the post-exposure period but the populations tended to become more similar over successive generations in their behaviour after the exposure. Furthermore, there were significant effects of generation (H (d.f. = 1, N = 137) = 10.94, P < 0.05) as well as population (H (d.f. = 1, N = 137) = 5.17, P < 0.05) on the immediate reaction to the simulated predator attack. In conclusion, over four successive generations, the two populations altered their antipredator behaviour and tended to become more similar. This study shows that antipredator behaviour may change over generations in a captive environment. This is likely to be one of the most crucial factors for successful reintroduction into the wild and hence, it is a very important aspect to consider for conservation breeding.

Ett stort antal utrotningshotade djurarter ingår idag i bevarandeprogram världen över. Små populationer hålls då i skyddade miljöer, exempelvis i djurparker, och i många fall är målet att återintroducera djur till naturen. Dessvärre är det vanligt att det uppstår problem när djur återintroduceras vilket kan bero på beteendemässiga anpassningar som uppkommit under tiden i fångenskap. Syftet med den här studien var därför att undersöka beteendemässiga aspekter på bevarandeavel och försöka ta reda på om och hur djur påverkas beteendemässigt av att hållas i skyddade fångenskapsmiljöer. I projektet användes röda djungelhöns (Gallus gallus) som en fallstudie. En granskning av beteendevariation mellan olika populationer av röda djungelhöns i fångenskap konstaterade skillnader i antipredatorbeteende, socialt beteende och födosöksbeteende. Vid en genetisk studie av samma populationer upptäcktes dessutom ett samband mellan genetisk diversitet och beteendevariation som potentiellt kan vara intressant ur ett bevarandeperspektiv. Socialt beteende, födosöksbeteende och olika aspekter av rädsla studerades vidare i populationer med olika bakgrund som fick växa upp tillsammans i en grupp. Resultaten visade att populationerna bara skilde sig åt i rädslebeteenden vilket antyder att denna typ av beteende i större utsträckning påverkas av långvarig avel i en viss fångenskapsmiljö medan socialt beteende och födosöksbeteende istället kan bero på den omedelbara sociala eller fysiska miljön. Antipredatorbeteende studerades också i en longitudinell studie av två populationer över fyra generationer och det visade sig att populationerna blev mer lika varandra ju längre tiden gick då de hölls under likadana miljöförhållanden. Det verkar alltså som om antipredatorbeteende kan förändras av avel i en viss miljö efter bara ett fåtal generationer. Utöver detta studerades även dagliga beteendemönster i olika djurparksmiljöer samt dygnsrytm av galanden hos både vilda populationer och djurparkspopulationer inom artens naturliga utbredningsområde. Resultaten tyder på att sådana beteendemönster inte påverkas nämnvärt av att djur hålls i fångenskap. Fallstudien som presenteras här är ett av de första försöken att, ur ett bevarandeperspektiv, studera hur fångenskapsmiljöer kan påverka djurs beteende och resultaten talar för att dessa aspekter är viktiga att ta hänsyn till vid planering av bevarandeavel.

A number of endangered species are currently involved in conservation breeding programs worldwide. Conservation breeding deals with propagation of captive populations, often with the ultimate aim of releasing animals into the wild. However, an alarmingly high proportion of reintroductions have not been successful in establishing viable populations, possibly due to behavioural problems caused by genetic adaptation to captivity. The main aim of this thesis was to study behavioural aspects of conservation breeding and investigate whether, and how, maintenance of small populations in captivity cause behavioural modifications, which could affect the success of reintroductions. Throughout the project, the red junglefowl (Gallus gallus) was used as a case study, representing animals maintained in captive populations. A screening of behavioural variation revealed that captive populations differ in antipredator, social and exploratory behaviours, all of which are central components of life in the wild. A correlation was also found between genetic diversity and behavioural variation. This has not been reported before and may potentially have interesting implications for conservation breeding. When studying the behaviour of populations with different backgrounds being raised together as one group, the results suggested that fear-related behaviours may be more affected by long-term breeding in a certain captive environment than social and exploratory behaviours which seem to be more influenced by the immediate social or physical environment. A longitudinal study of antipredator behaviour in two populations across four generations revealed that the populations became more similar over time when maintained under identical conditions. This demonstrates that effects of a new environment can appear after only a few generations. Furthermore, daily behavioural routines in different captive environments as well as diurnal crowing rhythms in both wild and captive populations were studied in the species’ natural region of distribution and the results suggest that such behavioural patterns are not affected by the captive environments to any notable extent. The present case study is one of the first attempts to, from a conservation perspective, study how captive environments can affect behaviour and the results imply that these aspects are important to take into consideration in conservation breeding programs.

Ex situ conservation of threatened species may lead to behavioural adaptation, which can affect success of reintroduction attempts. In previous studies, we investigated the effects of captivity on the behaviour of red jungle fowl (Gallus gallus) and found that captive populations differed behaviourally as well as genetically. The aim of the present study was to compare the behaviour of two of the previously studied populations, raised under identical conditions. Eggs were collected from birds at Copenhagen zoo (Cop) and Götala research station (Got) and were incubated and hatched together. Twenty-eight birds (16 Got and 12 Cop) were reared together and tested in eight different behavioural tests, measuring different aspects of fear-related behaviours as well as exploratory and social behaviours. The study revealed several differences in fear-related behaviours between the populations but none in exploratory or social behaviours. In general, one of the populations (Cop) showed more intense fear behaviours than the other (Got), which instead were less fearful in their behaviours. This indicates that breeding animals in captivity may lead to behavioural modifications, which can affect the outcome of reintroductions. The results further suggest that fear-related behaviours are dependent on the genetic background of the animals while social behaviours may be more influenced by the social environment. Since fear-related behaviours, such as predator avoidance and fear of humans, are essential for a life in the wild, these aspects are crucial for the breeding of animals in captivity for conservation purposes.

The escalating threats to ecosystems worldwide have lead to a need for efficient methods to breed animals in captivity and to prepare captive-born animals for release back to the wild. However, life in captivity may lead to modifications in the animal’s behaviour mainly by genetic changes, including behavioural adaptations such as reduced predator responses. Such modifications may seriously affect survival after a reintroduction. The present study was a first screening of behavioural and morphological variation between different captive populations in standardized test situations using red junglefowl as a model species. The birds were tested in three different test situations in order to measure anti-predatory behaviour, social behaviour and exploratory behaviour. The results of this study clearly show that there are behavioural differences between the captive populations which potentially can be crucial for the animals in a reintroduction situation. However, the extent to which these differences are due to genetic changes caused by small breeding populations or adaptations to the different captive environments is not yet known, although morphological differences found suggest that genetic variation may cause some of the behavioural differences as well. The differences found imply that life in captivity can affect an animal’s behaviour and even though the red junglefowl is merely used as a model here, this suggests that these aspects may be important to consider also in other species where reintroduction is a more central motive for keeping the animals in captivity.

Conservation of species is to a large extent carried out by zoos where animals are kept in small populations under protected conditions. The ultimate goal of such ex situ conservation programs is to eventually reintroduce the animals into natural habitats to provide support for the survival of the species. However, reintroduction has encountered considerable difficulty in the past, often due to behavioural deficiencies. Maintaining animals in captivity may lead to behavioural modifications as a result of adaptation to the captive environment, altered selection pressures and loss of genetic variation. The overall aim of this thesis was to study behavioural and genetic aspects of ex situ conservation and investigate whether maintenance of smal1 populations in captivity causes modifications which can affect the survival and reproduction capacity of the animals in a hypothetical reintroduction situation. Throughout the project the red junglefowl (Callus gallus) is used as a model for ex situ conservation populations. In Paper I, the behavioural variation between four captive populations of red junglefowl was studied. The birds were tested in different test situations; anti-predatory behaviour test, social behaviour test and exploratory and sociality test. The results showed clear behavioural differences between the populations, some of which are relevant from a conservation perspective. In Paper 11, the genetic relationships between the populations were examined by microsatellite analysis and possible correlations with the results of Paper I were investigated. The results showed that the populations were genetically differentiated and that all populations had lost a considerable amount of their assumed original genetic variation. Furthermore, the genetic variation of each population fo11owed the same pattern as behavioural variation ranks. The populations which had the highest genetic variation were also the ones showing the most behavioural variation in most behavioural variables. The results of Papers I and II imply that maintenance in captivity can affect an animal behaviourally as well as genetically. Even though the red junglefowl is merely used as a model here, the studies suggest that these issues are important to consider also in species where reintroduction is a more central motive for keeping the animals in captivity.

Bevarande av utrotningshotade djur sker idag i stor utsträckning i samarbete med djurparker. Målet med sådana s.k. ex situ-bevarandeprogram är att så småningom återintroducera djur till naturen för att skydda arten från utrotning. Återintroduktioner har dock visat sig vara problematiska i många fall. Att hålla djur i fångenskap kan leda till beteendemässiga förändringar till följd av anpassning till den skyddade miljön, förändrade selektionstryck och förlust av genetisk variation. Detta skulle kunna förklara en del av de misslyckade återintroduktionerna och för att bevarandeprogrammen ska bli effektivare krävs mer kunskap om vad som sker när djur placeras fångenskap. Det övergripande syftet med den här studien var därför att undersöka beteendemässiga och genetiska aspekter på ex situ-bevarande samt att utreda om hållandet av små djurpopulationer i fångenskap kan leda till förändringar som kan påverka framgången av en återintroducering. Röda djungelhöns (Gallus gallus) användes som modell för att kunna undersöka detta. I artikel I studerades beteendeskillnader mellan fyra populationer av röda djungelhöns. Hönsen studerades i olika testsituationer för att mäta antipredatorbeteende, socialt beteende och utforskningsbeteende. Resultaten visade klara skillnader i beteende mellan de olika populationerna och flera av dessa är relevanta utifrån ett bevarandeperspektiv. I artikel II analyserades det genetiska förhållandet mellan populationerna med hjälp av mikrosatellitmarkörer och möjliga samband med resultaten från artikel I undersöktes. Resultaten visade att populationerna var genetiskt differentierade och att de inom varje population hade förlorat en stor del av sin ursprungliga genetiska diversitet. När resultaten från de två artiklarna jämfördes visade det sig dessutom att den genetiska diversiteten följde samma mönster som rankingen av beteendemässig variation. De populationer med mest genetisk variation var också de som visade mest beteendemässig variation. Resultaten från artikel I och II antyder att hållande av djur i exempelvis djurparker kan påverka djuren både beteendemässigt och genetiskt. Röda djungelhöns användes här som en modell men resultaten tyder på att det här är viktigt att tänka på även hos arter där återintroducering är ett mer centralt motiv för att hålla djuren i fångenskap.

1. Although social behaviour is a major factor affecting the coping of poultry in production environments little is known about how it has been affected by intensive selection processes in fowl. We attempted to clarify selection effects on overall repertoire and occurrence of different social behaviours as well as on aggressive responses to re-grouping with unfamiliar birds by comparing high-producing White Leghorn layers to wild type Red Junglefowl.

2. In the first experiment we observed 8 stable mixed sex groups/breed each consisting of four 24-week-old birds previously familiar to each other. During 9 consecutive days, a wide range of social signals, sexual and aggressive interactions as well as spacing behaviour and activity were recorded over a 12-h photoperiod.

3. In the second experiment, starting at 19 weeks of age, 16 single sex groups of three birds from each breed were formed by mixing unfamiliar individuals. Aggressive behaviours were recorded 0, 5, 24 and 48 h after re-grouping.

4. Results from the stable groups indicated that the repertoire of social behaviours has been preserved during selection with few changes in frequencies and intensities. However, Leghorns showed a more cohesive spacing pattern than junglefowl.

5. In the second experiment, aggressive activity was higher immediately and after 24 h following re-grouping in Leghorns, but there was a drop in the aggressiveness at 5 h to the same level as junglefowl. We suggest that this may indicate poorer social learning capacity with a weaker ability to cope with group disruptions compared to the ancestral breed.