Effect of space allowance and earthen flooring on behaviour of farmed blue foxes

This study sought to evaluate the effect of cage space and earthen flooring on the behaviour of individually caged, farmed blue foxes (Alopex lagopus). Three different cage sizes [80 cm long (CL80), 120 cm long (CL120), 240 cm long (CL240); each 105 cm wide×70 cm high] with wire-mesh flooring, and one two-level cage (CL240E) with both wire-mesh (240 cm long×105 cm wide×70 cm high) and earthen flooring (80 cm long×105 cm wide×70 cm high) were employed. Quantitative ethograms were obtained from ten males in each group by videotaping the animals for 144 h monthly from August through November. Altogether 30 different behaviours were described. These were rather similar in all study groups. Examples of behavioural differences included pacing around with a neighbour and the incidence of scratching, which both declined with increasing cage space. Only the foxes in the cage with an earthen flooring (CL240E) exhibited digging behaviour, which averaged 11 min/24 h. The wire-mesh section was distinctly preferred to the earthen-floor section for most behaviours. Foxes in all groups were at their most active from 0800 to 1600 hours. Total activity, including several separate behaviours, declined as winter approached. Locomotor and oral stereotypies were infrequent, and no significant differences were found between the various cage options. For several hours before feeding, the foxes showed increasing levels of stereotypies, but afterwards, stereotypies abruptly declined. Received in revised form: 28 March 2001 Electronic Publication     

Quantification of abnormal repetitive behaviour in captive European starlings (Sturnus vulgaris)

Stereotypies are repetitive, unvarying and goalless behaviour patterns that are often considered indicative of poor welfare in captive animals. Quantifying stereotypies can be difficult, particularly during the early stages of their development when behaviour is still flexible. We compared two methods for objectively quantifying the development of route-tracing stereotypies in caged starlings. We used Markov chains and T-pattern analysis (implemented by the software package, Theme) to identify patterns in the sequence of locations a bird occupied within its cage. Pattern metrics produced by both methods correlated with the frequency of established measures of stereotypic behaviour and abnormal behaviour patterns counted from video recordings, suggesting that both methods could be useful for identifying stereotypic individuals and quantifying stereotypic behaviour. We discuss the relative benefits and disadvantages of the two approaches.     

Stereotypic behavior in Asiatic black and Malayan sun bears

The stereotypies of individually caged Asiatic black bears (Ursus thibetanus) and Malayan sun bears (Helarctos malayanus) were studied in detail. Stereotypies were performed by 27 of the 29 subjects, were primarily locomotory in form (e.g., pacing), and occupied on average 18% (standard error of the mean (SEM)=2.5) of daylight hours. Stereotypy levels during the night were almost negligible and were highly correlated with daytime levels. Total stereotypies peaked prior to food arrival, although oral stereotypies were most frequent after feeding. In general, stereotypies were performed in locations from which food arrival could be viewed, although Asiatic black bears were equally likely to exhibit stereotypy near a neighboring bear. Across individuals, stereotypy frequency was inversely correlated with inactivity and increased with age. Older bears also showed less normal activity and a reduced diversity of normal behavior. Stereotypy levels were unrelated to levels of “compulsive” behavior (e.g., hair plucking) or repetitive self‐sucking–a potential deprivation stereotypy. More frequent stereotypies were performed more invariantly (i.e., were more predictable from one repetition to the next) and in more diverse contexts, namely 1) outside the pre‐feeding period, and 2) during the night. Contrary to observations reported elsewhere, higher frequencies of stereotypy were not associated with reduced behavioral diversity, or with a more elaborate repertoire of stereotypy forms and sequences. Although the two species did not differ in overall frequency, the stereotypies of sun bears appeared to be more food‐motivated than those of Asiatic black bears: the sun bears displayed a higher frequency and diversity of oral stereotypies, and higher levels of pre‐feeding stereotypy, and performed significantly more of their total stereotypies in locations from which they could view food arrival. This study demonstrates how analyzing stereotypies in detail can help identify the motivations that underlie these behaviors, and potentially reveal their degree of establishment–both of which are important factors in stereotypy treatment. Zoo Biol 23:409–430, 2004. © 2004 Wiley‐Liss, Inc.     

Group housing in row cages: an alternative housing system for juvenile mink

We studied a group housing system as an alternative to the traditional pair housing of juvenile mink. The focus was on both the welfare and production of mink. The pairs were housed in standard mink cages, whereas the groups were in row cage systems consisting of three standard mink cages connected to each other. The welfare of the mink was evaluated by behavioural observations (stereotypies and social contacts), evaluation of the incidence of scars assumed to be caused by biting, and adrenal function (serum cortisol level after adrenocorticotropic hormone (ACTH) administration and adrenal mass). Feed consumption, pelt length, quality and price were used for comparing the two housing systems from the economic point of view. Although the incidence of scars showed that there might have been more aggressive behaviour among the group-housed than among the pair-housed mink, this was not observed unambiguously in behavioural observations, and, at least, aggression did not cause mortality or serious injuries to the animals as has been observed in some earlier studies. In addition, the housing system did not affect pelt size, and, although the quality of the pelts was slightly lower in the group than in pair-housed mink, there was only a tendency for lower pelt prices. The lower pelt prices in the group-housed mink might even be partially compensated for by the group-housed mink eating 10% to 20% less in the late autumn, due to thermoregulatory benefits, than their pair-housed conspecifics. The results on the frequency of stereotypic behaviour (but not adrenal function) suggest that the group-housed animals were possibly less stressed than the pair-housed animals. Group housing of juvenile farmed mink in a row cage system cannot be recommended before the effects on welfare and production are clarified in further studies.     

Behavioural responses to changing landscapes: flock structure and anti-predator strategies of tits wintering in fragmented forests

We analysed the effects of forest fragmentation on the flock structure of insectivorous forest passerines (Parus, Aegithalos, Certhia, Regulus, etc.), and on the anti‐predator behaviour and energy management of blue tits in these flocks. We surveyed flocks in Central Spain during two winters. Flocks in fragments comprised fewer individuals and species than flocks in unfragmented forests. The most abundant species in forest flocks (blue tit, Parus caeruleus, and firecrest, Regulus ignicapillus) were also the most abundant in fragments, while the rarest species in the area never occurred in small woodlots. We investigated how fragmentation and related changes in flock structure affect anti‐predator behaviour of blue tits, a widely distributed species in the area. In fragments but not in forests, blue tits increased scanning rates with decreasing flock size. Vigilance was relaxed when great tits, Parus major, were abundant as flock mates, suggesting that the absence of this dominant species in fragments could intensify anti‐predator behaviour of blue tits. Blue tits enhanced anti‐predator behaviour in the second winter parallel to an increase in the abundance of raptors. This behavioural change was stronger in fragments, where blue tits foraged deeper in the canopy and increased scanning and hopping rates. Under increased predation risk, birds are expected to reduce body mass to improve predator avoidance. On average, blue tits weighed similar in fragments and forests the second winter. However, they accumulated fat along the day in fragments only, and adjusted body mass to body size more closely in that habitat type. This suggests that blue tits perceived fragments as unpredictable habitats where fattening would help avoid starvation, but also as dangerous sites where overweight would further increase the risk of predation. In summary, our results support that fragmentation affects individual behaviour of blue tits, and show the potential of behavioural approaches to unravel how different species face the advancing fragmentation of their habitats.     

Rival imprinting: interspecifically cross-fostered tits defend their territories against heterospecific intruders

Failure to recognize conspecifics in social interactions such as mate choice and aggressive encounters will often result in reduced fitness. Studies on mate choice show that the ability to recognize conspecifics as mates is not universally present at birth, but often needs to be learned. In contrast, little is known about the ontogeny of intrasexual species recognition. To test whether learning influences the recognition of sexual rivals, we compared the aggressive response towards intruders of interspecifically cross-fostered individuals and controls reared by conspecific parents. We simulated territorial intrusion by presenting either a caged individual or playback song near the nest of breeding pairs of great tits, Parus major, and blue tits, P. caeruleus. Great tits reared by blue tit parents responded much more to blue tit stimuli than did great tit controls, and furthermore showed stronger responses to blue tit stimuli than to those of their own species. Blue tits reared by great tits responded much more to great tit stimuli than did blue tit controls. In contrast, blue tits cross-fostered to coal tits, P. ater, did not respond more to coal tits than did blue tit controls. There was a species difference in the response to conspecifics: blue tits cross-fostered to great tits responded more to conspecifics than did cross-fostered great tits. The results were similar for males and females. We conclude that learning influences intrasexual species recognition in these tits. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Effects of early weaning and housing conditions on the development of stereotypies in farmed mink

Female mink pups were weaned at 6, 8 or 10 weeks of age and subjected to two different housing conditions. They were either kept together with a single male sibling in traditional mink cages (30x45x90 cm) or housed socially with all litter-mates in an alternative system consisting of three adjoining traditional cages (90x45x90 cm). All cages were supplied with nest boxes. At 5 months of age, the siblings were removed leaving the females socially isolated in the two different cage systems. Females' stereotypies were quantified by repeated scanning observations under the social housing conditions immediately before removal of the siblings, and again at the age of 7 and 9 months, when the animals had stayed solitary in the two systems for 2 and 4 months. Solitary females showed significantly more stereotypies than females under social housing conditions in both cage systems. Stereotypies were more frequent in the smaller traditional cages. Stereotypies declined from 7 to 9 months of age among solitary animals in traditional cages but not in alternative cages. Early-weaned solitary females in traditional cages showed more stereotypies than later-weaned animals, but only when measured at the age of 7 months. It is suggested that early weaning, individual housing and small cages promote the development of stereotypies in farmed mink. The influence of early weaning on stereotypies seems to decline with age, while effects related to individual housing and small cages appear to be more persistent.     

Blue tits (Cyanistes caeruleus) respond to an experimental change in the aromatic plant odour composition of their nest

Although the use of olfaction by birds is now widely recognised, the olfactory abilities of passerine birds remain poorly explored, for historical reasons. Several studies however suggest that passerines can perceive volatile compounds in several biologically relevant contexts. In Corsica, recent findings suggest that cavity-nesting blue tits may use volatile compounds in the context of nest building and maintenance. Although they build their nests mainly from moss, female blue tits also frequently incorporate fragments of several species of aromatic plants in the nest cup. In field experiments, breeding female blue tits altered their nest maintenance behaviour in response to experimental addition of aromatic plants in their nest. In aviary experiments, captive male blue tits could be trained to detect lavender odour from a distance. Here I report results from a field study aimed to test whether adult blue tits altered their chick-feeding behaviour after an experimental change in nest odour composition. I experimentally added fragments of aromatic plant species that differed from those brought in the nests before the start of the experiment in a set of experimental nests and added moss, the basic nest material, in a set of control nests. Both male and female blue tits hesitated significantly longer entering the nest cavity after addition of new aromatic plant fragments, as compared to moss addition. This response was especially observed during the first visit following the experimental change in nest plant composition. Nest composition treatment had no effect on the time spent in the nest. This study demonstrates that free-ranging blue tits detect changes in nest odour from outside the nest cavity.     

Environmentally Enriched Male Mink Gain More Copulations than Stereotypic,Barren-Reared Competitors

Wild carnivores in zoos, conservation breeding centres, and farms commonly live in relatively small, unstimulating enclosures. Under these captive conditions, in a range of species including giant pandas, black-footed ferrets, and European mink, male reproductive abilities are often poor. Such problems have long been hypothesized to be caused by these animals'' housing conditions. We show for the first time that rearing under welfare-improving (i.e., highly valued and stress-reducing) environmental enrichments enhances male carnivores'' copulatory performance: in mate choice competitions, enriched male American mink (Neovison vison) mated more often than non-enriched males. We screened for several potential mediators of this effect. First was physiological stress and its impact on reproductive physiology; second, stress-mediated changes in morphology and variables related to immunocompetence that could influence male attractiveness; and third, behavioural changes likely to affect social competence, particularly autistic-like excessive routine and repetition (‘perseveration’) as is reflected in the stereotypies common in captive animals. Consistent with physiological stress, excreted steroid metabolites revealed that non-enriched males had higher cortisol levels and lower androgen levels than enriched conspecifics. Their os penises (bacula) also tended to be less developed. Consistent with reduced attractiveness, non-enriched males were lighter, with comparatively small spleens and a trend to greater fluctuating asymmetry. Consistent with impaired social competence, non-enriched males performed more stereotypic behaviour (e.g., pacing) in their home cages. Of all these effects, the only significant predictor of copulation number was stereotypy (a trend suggesting that low bodyweights may also be influential): highly stereotypic males gained the fewest copulations. The neurophysiological changes underlying stereotypy thus handicap males sexually. We hypothesise that such males are abnormally perseverative when interacting with females. Investigating similar problems in other taxa would be worthwhile, since many vertebrates, wild and domestic, live in conditions that cause stereotypic behaviour and/or impair neurological development.     

Personality‐related differences in response to habitat in Mediterranean blue tits

Habitat‐specific selection pressures have been widely recognized, but whether selection favours different personality types in different habitats has rarely been evaluated. This study aimed to test whether personality‐related differences in annual reproductive success differed between two populations of blue tits (Cyanistes caeruleus) occupying different Mediterranean habitats (oakwood and pinewood). We measured exploration and parental provisioning behaviours and used a path analysis to ask how the interplay between these two behavioural traits affected reproductive success in each of these two habitats. We found that blue tits breeding in the pinewood were slow‐exploring compared to blue tits breeding in the oakwood, suggesting the occurrence of personality‐related differences in settlement, or behavioural plasticity in response to habitat. Exploration behaviour and feeding rates were positively associated, suggesting that they affect each other or that there is an environmental factor affecting both traits simultaneously. Finally, fast explorers were favoured in the pinewood, while there was no selection acting on exploration behaviour in the oak habitat. These findings emphasize the importance of integrating habitat selection, plasticity and personality in the study of behavioural evolution.     

The effect of preweaning and postweaning housing on the behaviour of the laboratory mouse (Mus musculus)

Standard housing for laboratory mice severely restricts natural behaviour and the control that the animal has over its environment. Providing the cage with objects is a method that has been used to both increase environmental complexity, promote the performance of natural behaviour and provide greater controllability for the animal. This method of furnishing cages has mostly been studied in adult animals, and little is known about the influence that the preweaning environment has on the behaviour of mice as adults. This study aimed to investigate the effects on mice behaviour of preweaning and postweaning housing environment. In this experiment, 64 pairs of animals of the strain C57BL/6J were used. Half of the animals were born and reared until weaning in standard cages and the other half in cages twice the size of the standard and furnished with nesting material, a cardboard tube, a PVC nest box and a wooden chewblock. After weaning, half the animals in each group were changed to the other type of cage, whereas the other half remained in the same environment; in both cases they were kept in single-sex pairs of littermates. Behaviour during the dark, active period was studied through video recordings. We found no main effects of preweaning environment on behaviour; however, mice moved from furnished to standard cages at weaning showed a decrease in inactive behaviour at four weeks of age. Mice housed after weaning in standard cages spent less time inactive, and more time engaging in activities like feeding and drinking, self-grooming and allogrooming. A sex difference was also found, in that females showed a greater performance of exploratory behaviour as well as a greater prevalence of stereotypies. The use of different objects and locations within the furnished cage was also analysed at both ages. Results show that at eight weeks of age mice spent more time at the top of the cage, and that the use of the nest box (although not for resting) increased between four and eight weeks. Mice were found to use the nest box as a nesting site/sleeping place only at age four weeks, whereas they always used the nesting material for sleeping.     

Personality and urbanization: behavioural traits and <Emphasis Type="Italic">DRD4</Emphasis> SNP830 polymorphisms in great tits in Barcelona city

Most examples of adaptation to the urban environment relate to plasticity processes rather than to natural selection. Personality, however, defined as consistent individual differences in behaviour related to exploration, caution, and neophobia, is a good behavioural candidate character to study natural selection in relation to the urban habitat due to its heritable variation. The aim of this paper was to analyse variation in personality by comparing urban and forest great tits Parus major using standard tests of exploratory behaviour and boldness. We studied personality in 130 wild great tits captured in Barcelona city and nearby forests and found that urban birds were more explorative and bolder towards a novel object than forest birds. Genotype frequencies of the DRD4 SNP830 polymorphism, a gene region often associated with personality variation, varied significantly between forest and urban birds. Behavioural scores, however, were not correlated with this polymorphism in our population. Exploration scores correlated to boldness for forest birds but not for urban birds. Our findings suggest that the novel selection pressures of the urban environment favour the decoupling of behavioural traits that commonly form behavioural syndromes in the wild.     

Test multipler Hypothesen zum Auftreten von stereotypen Verhaltensweisen bei Großkatzen im Zoo Leipzig

We tested multiple hypotheses on the occurrence of stereotypic behaviour in leopards (Panthera pardus), snow leopards (Uncia uncia) and tigers (Panthera tigris) at Leipzig Zoo. Stereotypies in all three species occurred exclusively as pacing. Stereotypic pacing was observed in 79% of the 14 animals and occupied on average 19% of the time spent active. Stereotypies were unaffected by species, age, sex and social housing, respectively. Animals showed significantly more stereotypic pacing before feeding. Stereotypic pacing was performed significantly more often in locations from which food arrival could be first viewed. Older animals tended to show more stereotypic pacing after feeding. Animals that generally exhibited more stereotypic behaviour paced significantly more often after feeding. Our results suggest that stereotypies may represent an exaggerated response of food reward. Furthermore, stereotypies may become emancipated from their original motivation over the course of an individual's development. Stereotypies thus could possibly be remedied by increasing the feeding (i.e. reward) frequency, if applied from an animal's early age.     

Sexual imprinting and the origin of obligate brood parasitism in birds

We discuss two pathways along which obligate brood parasitism (OBP) may evolve and examine some of the critical steps that must be passed by letting great tits Parus major be reared by blue tits Parus caeruleus in a field experiment. The cross-fostered chicks survived well in blue tit nests, but their local recruitment and reproductive success was much lower than that of controls. The effect was strongest when great tits grew up with siblings of the host species rather than with conspecific siblings in blue tit nests. The low success seemed to be caused by misimprinting because the cross-fostered birds behaved like blue tits in several aspects (species association, alarm calls, and aggressive response by resident females to caged intruders). Some birds of both sexes were apparently so strongly imprinted that they did not attract or accept a social mate of their own species. We conclude that imprinting may be necessary for OBP to evolve in birds because the parasite must be attracted to the nests of the host species to add eggs and thereby continue the parasitic life cycle. However, strong imprinting may also prevent OBP from occurring if parasitic offspring seek a mate from the host species.     

Does Rearing Laying Hens in Aviaries Adversely Affect Long-Term Welfare following Transfer to Furnished Cages?

This study tests the hypothesis that hens that are reared in aviaries but produce in furnished cages experience poorer welfare in production than hens reared in caged systems. This hypothesis is based on the suggestion that the spatial restriction associated with the transfer from aviaries to cages results in frustration or stress for the aviary reared birds. To assess the difference in welfare between aviary and cage reared hens in production, non-beak trimmed white leghorn birds from both rearing backgrounds were filmed at a commercial farm that used furnished cage housing. The videos were taken at 19 and 21 weeks of age, following the birds'' transition to the production environment at 16 weeks. Videos were analysed in terms of the performance of aversion-related behaviour in undisturbed birds, comfort behaviour in undisturbed birds, and alert behaviour directed to a novel object in the home cage. A decrease in the performance of the former behaviour and increase in the performance of the latter two behaviours indicates improved welfare. The results showed that aviary reared birds performed more alert behaviour near to the object than did cage reared birds at 19 but not at 21 weeks of age (P = 0.03). Blood glucose concentrations did not differ between the treatments (P>0.10). There was a significant difference in mortality between treatments (P = 0.000), with more death in aviary reared birds (5.52%) compared to cage birds (2.48%). The higher mortality of aviary-reared birds indicates a negative effect of aviary rearing on bird welfare, whereas the higher duration of alert behavior suggests a positive effect of aviary rearing.     

The effects of feeding and housing on the behaviour of the laboratory rabbit

The effects of housing, feeding time and diet composition on the behaviour of the laboratory rabbit were examined. The animals were caged individually in single or double metal cages with perforated metal floors, metal walls, and bars in the front, or kept as a group in floor pens. The light/dark cycle was 12/12 h with light from 04:00 to 16:00 h and 30 min twilight. One experiment compared feeding equal energy levels of a high energy diet (10.1 MJ/kg) and with a low energy diet (7.0 MJ/kg) at 08:00 h. The second experiment compared feeding the high energy diet at 08:00 h and at 14:00 h. In both studies the behaviour of the rabbits was recorded between 08:00 and 14:00 h and between 16:00 and 22:00 h. Feeding the animals at 14:00 h reduced abnormal behaviour during the dark period compared to feeding at 08:00 h, whereas no difference in behaviour could be detected between feeding a high-energy and a low-energy diet at 08:00 h. Animals in floor pens generally showed less abnormal behaviour than caged animals. The results indicate that the welfare for caged rabbits can be improved by feeding the animals in the afternoon rather than in the morning.     

Aromatic herbs in Corsican blue tit nests: The ‘Potpourri’ hypothesis

This study reports that Corsican blue tit (Parus caeruleus ogliastrae) nests contain between one to five aromatic herb species between the onset of egg laying till the chicks’ finished growth 13 d after hatching. An herb removal experiment during the chick stage shows that blue tits bring fresh aromatic material 1–5 d after herb removal. Nests with a series of distinct odour classes easily perceived by humans have never been reported in birds. A new ‘Potpourri’ hypothesis is proposed that may explain the functional significance of this behaviour.     

Parental provisioning behaviour plays a key role in linking personality with reproductive success

Repeatable behavioural traits (‘personality’) have been shown to covary with fitness, but it remains poorly understood how such behaviour–fitness relationships come about. We applied a multivariate approach to reveal the mechanistic pathways by which variation in exploratory and aggressive behaviour is translated into variation in reproductive success in a natural population of blue tits, Cyanistes caeruleus. Using path analysis, we demonstrate a key role for provisioning behaviour in mediating the link between personality and reproductive success (number of fledged offspring). Aggressive males fed their nestlings at lower rates than less aggressive individuals. At the same time, their low parental investment was associated with increased female effort, thereby positively affecting fledgling production. Whereas male exploratory behaviour was unrelated to provisioning behaviour and reproductive success, fast-exploring females fed their offspring at higher rates and initiated breeding earlier, thus increasing reproductive success. Our findings provide strong support for specific mechanistic pathways linking components of behavioural syndromes to reproductive success. Importantly, relationships between behavioural phenotypes and reproductive success were obscured when considering simple bivariate relationships, underlining the importance of adopting multivariate views and statistical tools as path analysis to the study of behavioural evolution.     

Realized heritability and repeatability of risk-taking behaviour in relation to avian personalities

Personalities are general properties of humans and other animals. Different personality traits are phenotypically correlated, and heritabilities of personality traits have been reported in humans and various animals. In great tits, consistent heritable differences have been found in relation to exploration, which is correlated with various other personality traits. In this paper, we investigate whether or not risk-taking behaviour is part of these avian personalities. We found that (i) risk-taking behaviour is repeatable and correlated with exploratory behaviour in wild-caught hand-reared birds, (ii) in a bi-directional selection experiment on 'fast' and 'slow' early exploratory behaviour, bird lines tend to differ in risk-taking behaviour, and (iii) within-nest variation of risk-taking behaviour is smaller than between-nest variation. To show that risk-taking behaviour has a genetic component in a natural bird population, we bred great tits in the laboratory and artificially selected 'high' and 'low' risk-taking behaviour for two generations. Here, we report a realized heritability of 19.3 +/- 3.3% (s.e.m.) for risk-taking behaviour. With these results we show in several ways that risk-taking behaviour is linked to exploratory behaviour, and we therefore have evidence for the existence of avian personalities. Moreover, we prove that there is heritable variation in more than one correlated personality trait in a natural population, which demonstrates the potential for correlated evolution.