Environmental enrichments and data-driven welfare indicators for sheltered dogs using telemetric physiological measures and signal processing.

Shelters are stressful environments for domestic dogs which are known to negatively impact their welfare. The introduction of outside stimuli for dogs in this environment can improve their welfare and life conditions. However, our current understanding of the influence of different stimuli on shelter dogs' welfare is limited and the data is still insufficient to draw conclusions. In this study, we collected 28 days (four weeks) of telemetry data from eight male dogs housed in an Italian shelter for a long period of time. During this period, three types of enrichment were introduced into the dogs' pens for one week each: entertaining objects, intraspecific, and interspecific social enrichment, by means of the presence of female conspecifics and the presence of a human. To quantify their impact, we introduce novel metrics as indicators of sheltered dogs' welfare based on telemetry data: the variation of heart rate, muscle activity, and body temperature from an average baseline day, quality of sleep, and the regularity for cyclicity of the aforementioned parameters, based on the day-night cycle. Using these metrics, we show that while all three stimuli statistically improve the dogs' welfare, the variance between individual dogs is large. Moreover, our findings indicate that the presence of female conspecific is the best stimulus among the three explored options which improves both the quality of sleep and the parameters' cyclicity. Our results are consistent with previous research findings while providing novel data-driven welfare indicators that promote objectivity. Thus, this research provides some useful guidelines for managing shelters and improving dogs' welfare.

Genetic inference of the mating system of free-ranging domestic dogs.

Domestication has greatly changed the social and reproductive behavior of dogs relative to that of wild members of the genus Canis, which typically exhibit social monogamy and extended parental care. Unlike a typical gray wolf pack that consists of a single breeding pair and their offspring from multiple seasons, a group of free-ranging dogs (FRDs) can include multiple breeding individuals of both sexes. To understand the consequences of this shift in reproductive behavior, we reconstructed the genetic pedigree of an FRD population and assessed the kinship patterns in social groups, based on genome-wide single-nucleotide polymorphism genotypes. Consistent with behavioral observations, the mating system of the study population was characterized by polygynandry. Instead of the discreet family units observed in wolves, FRDs were linked by a network of kinship relationships that spread across packs. However, we also observed reproduction of the same male-female pairs in multiple seasons, retention of adult offspring in natal packs, and dispersal between neighboring packs-patterns in common with wolves. Although monogamy is the predominant mating system in wolves, polygyny and polyandry are occasionally observed in response to increased food availability. Thus, polygynandry of domestic dogs was likely influenced by the shift in ecological niche from an apex predator to a human commensal.

Neutering Effects on Social Behaviour of Urban Unowned Free-Roaming Domestic Cats.

The "trap, spay/neuter, and release" programs to manage unowned free-roaming cat populations are diffused worldwide and they are largely advised even in countries where the suppression of unowned cats is enforced by law. Despite the massive neutering campaigns in the world, there is little information on the influence of neutering on individual cat behaviour, as well as on the social structure of cat colonies. The aim of this study is to verify such effects. Before neutering, the group consisted of 17 free-roaming domestic cats, who were totally unrestrained. After neutering it consisted of 16 individuals. Data on the outcomes of aggressive, submissive, affiliative, and territorial behaviour were collected, using classic ethological methods ("focal animal", "all occurrences", and "1/0" sampling methods) (735 h of observation before and 537 h after neutering). The dominance hierarchy did not change after neutering. On the other hand, the frequency of aggressive, territorial behaviour as well as proximity among individuals decreased significantly. Affiliative behaviour has been observed among neutered adult males that never performed it before neutering. The results of this study suggest that, after neutering: (i) The group of free-ranging cats was stable over time; (ii) the hierarchical structure of the cat social group did not change; (iii) the general level of activity, decreased; (iv) urine spraying marking behaviour almost disappeared; (v) the level of social proximity among group members decreased, although it tended to increase in some male-male dyads. The most obvious effect of neutering, detected on individual behaviour, was that cats were less active.

Elevated aggression is associated with uncertainty in a network of dog dominance interactions.

Dominance hierarchies are widespread in animal societies and reduce the costs of within-group conflict over resources and reproduction. Variation in stability across a social hierarchy may result in asymmetries in the benefits obtained from hierarchy formation. However, variation in the stability and behavioural costs of dominance interactions with rank remain poorly understood. Previous theoretical models have predicted that the intensity of dominance interactions and aggression should increase with rank, but these models typically assume high reproductive skew, and so their generality remains untested. Here we show in a pack of free-living dogs with a sex-age-graded hierarchy that the central region of the hierarchy was dominated by more unstable social relationships and associated with elevated aggression. Our results reveal unavoidable costs of ascending a dominance hierarchy, run contrary to theoretical predictions for the relationship between aggression and social rank in high-skew societies, and widen our understanding of how heterogeneous benefits of hierarchy formation arise in animal societies.

The effect of domestication on post-conflict management: wolves reconcile while dogs avoid each other.

Highly cooperative social species are expected to engage in frequent reconciliation following conflicts in order to maintain pack cohesiveness and preserve future cooperation. By contrast, in social species with low reliance on cooperation, reconciliation is expected to be less frequent. Here, we investigate the pattern of reconciliation in four captive wolf packs and four captive dog packs. We provide evidence for reconciliation in captive wolves, which are highly dependent on cooperation between pack members, while domestic dogs, which rely on conspecific cooperation less than wolves, avoided interacting with their partners after conflicts. Occurrence, intensity, latency, duration and initiation of wolf reconciliations appeared to vary as a consequence of a compromise between the costs (e.g. risk of further aggression) and the benefits (e.g. restoring relationship with opponents) of such interactions. Our results are in line with previous findings on various wolf packs living under different social and ecological conditions, suggesting that reconciliation is an important strategy for maintaining functional relationships and pack cohesiveness. However, current results on dogs are in contrast to the only other study showing that reconciliation can occur also in this species. Therefore, the occurrence of reconciliation in dogs may be influenced by social and environmental conditions more than in wolves. Which factors promote and modulate reconciliation in dogs needs to be further investigated.

In wolves, play behaviour reflects the partners' affiliative and dominance relationship.

Puppy packs (consisting of only puppies) and mixed-age packs (composed of puppies and adults) were observed to test whether social play can be used for assessing and establishing social relations in wolves, Canis lupus. Differently from previous studies, we looked at play behaviours in detail, allowing us to categorize play interactions as either competitive or relaxed, and predicted that different types of play would be associated with different relationships between individuals. We found that the more time dyads spent in relaxed play, the more affiliative interactions they exchanged outside of play. In the mixed-age packs, dyads that spent more time in both relaxed and competitive play showed fewer exchanges of aggressive behaviours. Conversely, in puppy packs, the more time dyads spent in competitive play, the more aggressive interactions were exchanged outside of play. Since clear dominance relationships emerged in the mixed-age packs, but not in puppy packs, we suggest that play can help to reduce the frequency of aggressive interactions only when a clear hierarchy exists between pack members. Furthermore, we found that in both puppy and mixed-age packs, dominance relationships were reflected and rarely reversed during play. Finally, dyads with a less clear dominance relationship spent more time playing in a competitive way. Overall, our results support the social assessment hypothesis suggesting that social relationships outside of play are reflected during playful interactions. Moreover, we revealed how different types of play, that is, playing in a competitive or relaxed way, may be related to different social relationships. This distinction between play types has not been acknowledged before but could help researchers better understand the functions of play in different species.

Post-conflict opponent affiliation reduces victim re-aggression in a family group of captive arctic wolves (Canis lupus arctos).

Post-conflict affiliative interactions have been widely investigated in primates but not extensively in other species. Using the Post Conflict-Matched Control (PC-MC) comparison method, this study investigated the patterns of post-conflict opponent affiliation (POA) of a captive family group of 19 arctic wolves (Canis lupus arctos), investigating the correlation with various factors. We found that POAs occurred mainly in the non-feeding context and more often when the victim was dominant and the aggressor subordinate. Furthermore, POAs were more likely to have been initiated by the victim than the aggressor. Victims' stress related behaviours occurred more in PC than MC periods, and more after high vs. low intensity aggressions but they were not more likely to occur after conflicts between wolves with a stronger social bond and POAs did not reduce their rate of occurrence. Our results showed that re-aggression was twice less frequent when a friendly interaction occurred between the aggressor and the victim, and consistent with this, victims engaged in POAs more often than the aggressor. Overall, our results support the hypothesis that POAs in wolves may have been selected for as a mechanism to avoid conflict escalation, which could lead to social disruption and hence jeopardize cooperative activities. The high relatedness among individuals in the pack and the greater dependence of all members on cooperation in breeding and hunting may reduce the importance of 'relationship quality' as a mediating factor of POAs, although dominance relationships, which are directly linked to the risks of further conflicts, do play an important role.

Dominance relationships in a family pack of captive arctic wolves (Canis lupus arctos): the influence of competition for food, age and sex.

BACKGROUND: Dominance is one of the most pervasive concepts in the study of wolf social behaviour but recently its validity has been questioned. For some authors, the bonds between members of wolf families are better described as parent-offspring relationships and the concept of dominance should be used just to evaluate the social dynamics of non-familial captive pack members (e.g., Mech & Cluff, 2010). However, there is a dearth of studies investigating dominance relationships and its correlates in wolf family packs. METHODS: Here, we applied a combination of the most commonly used quantitative methods to evaluate the dominance relationships in a captive family pack of 19 Arctic wolves. RESULTS: We found a significant linear and completely transitive hierarchy based on the direction of submissive behaviours and found that dominance relationships were not influenced by the competitive contexts (feeding vs. non-feeding context). A significant linear hierarchy also emerges amongst siblings once the breeding pair (the two top-ranking individuals) is removed from analyses. Furthermore, results suggest that wolves may use greeting behaviour as a formal signal of subordination. Whereas older wolves were mostly dominant over younger ones, no clear effect of sex was found. However, frequency of agonistic (submissive, dominant and aggressive) behaviours was higher between female-female and male-male dyads than female-male dyads and sex-separated linear hierarchies showed a stronger linearity than the mixed one. Furthermore, dominance status was conveyed through different behavioural categories during intra-sexual and inter-sexual interactions. DISCUSSION: Current results highlight the importance of applying a systematic methodology considering the individuals' age and sex when evaluating the hierarchical structure of a social group. Moreover, they confirm the validity of the concept of dominance relationships in describing the social bonds within a family pack of captive wolves.

Play Behavior in Wolves: Using the '50:50' Rule to Test for Egalitarian Play Styles.

Social play is known as a cooperative interaction between individuals involving multiple mechanisms. However, the extent to which the equality of individuals' play styles affects the interaction has not been studied in many species. Dyadic play between wolf puppies, as well as between puppies and adults, was studied to investigate both self-handicapping and offensive behaviors to determine the extent to which wolves engage in play styles where one individual does not dominate the play. Our results did not support the hypothesized '50:50' rule, which suggests that more advantaged individuals should show higher rates of self-handicapping behaviors in order to facilitate play with others. Adult wolves performed significantly less self-handicapping behaviors than their puppy partners, and they performed significantly more offensive behaviors than their puppy partners. While the '50:50' rule was not supported at any time during our study period, dyads consisting of two puppies had significantly more equal play than dyads consisting of one puppy and one adult. These results suggest that wolf puppies are more likely to play on equal terms with similarly-aged play partners, while the dominance status of the partners dictates offensive and self-handicapping behaviors between animals of different ages.

Innate immune responses of young bulls to a novel environment.

Animal welfare during transportation has been investigated in several studies, as opposed to post-transportation phases. In this study, we evaluated the effect of a novel environment after transportation on 26 Friesian bulls, 242 ± 42 day-old, from ten different dairy farms. Animals were shipped to a breeding center in different seasons, and selected parameters of innate immunity (serum bactericidal activity, hemolytic complement, serum albumin, α, ß, and γ-globulins, interleukin-6, TNF-α) were monitored before and after the arrival at days--4/0/4/15/30. Our results showed significant differences of IL-6 and TNF-α protein levels at destination in December (94 ± 1.3 pg/ml) and June (+788 pg/ml), respectively. Moreover, the serum levels of these cytokines increased between days 0 and 15 after the arrival, the modulation of IL-6 being in agreement with established models of physical and/or psychological stress. Concerning the modulation of albumin, alpha and beta-globulins, the highest levels were detected in April, whereas a significant decrease was observed between day 15 and 30 after arrival; on the contrary, γ-globulin levels significantly increased after day 15. The results of this study highlight the occurrence of innate immune responses of young bulls to the combined effects of climate (season) and novel farming conditions.

Social variables affecting mate preferences, copulation and reproductive outcome in a pack of free-ranging dogs.

Mating and reproductive outcome is often determined by the simultaneous operation of different mechanisms like intra-sexual competition, mating preferences and sexual coercion. The present study investigated how social variables affected mating outcome in a pack of free-ranging dogs, a species supposed to have lost most features of the social system of wolves during domestication. We found that, although the pack comprised multiple breeding individuals, both male copulation success and female reproductive success were positively influenced by a linear combination of dominance rank, age and leadership. Our results also suggest that mate preferences affect mating outcome by reinforcing the success of most dominant individuals. In particular, during their oestrous period bitches clearly searched for the proximity of high-ranking males who displayed affiliative behaviour towards them, while they were more likely to reject the males who intimidated them. At the same time, male courting effort and male-male competition for receptive females appeared to be stronger in the presence of higher-ranking females, suggesting a male preference for dominant females. To our knowledge, these results provide the first clear evidence of social regulation of reproductive activities in domestic dogs, and suggest that some common organizing mechanisms may contribute to shape the social organization of both dogs and wolves.

Searching for differences in the behavioural response of piglet groups subjected to novel situations.

The Backtest (BT), the Open Field (OF) and the Novel Object (NO) tests have been used to identify individual reaction patterns in piglets and to measure parameters that previous studies have shown to be correlated to the coping strategies of animals. The BT allows for the classification of piglets into two different "coping styles": high-resisting (HR) and low-resisting (LR), which respectively correspond to a (pro-)active and passive (or reactive) behavioural response. During previous research, the subjects were tested singularly, so the aim of this study was to investigate if differences between HR and LR animals could be detected when piglets are tested in a group using the OP and NO tests. A total of 132 piglets were subjected to the BT and then were housed in groups consisting of four individuals each so as to obtain HR pens, LR pens and mixed pens. We found differences in the pigs' behaviour during the OF and the NO tests and according to the type of group. Individual differences in the behavioural response of piglets to the Backtest were not predictive of the behavioural response of the animals subjected to the OF or to NO tests. Our results show that there are no relevant differences between HR and LR piglets when they are subjected in a group to novel situations.

Free-ranging dogs assess the quantity of opponents in intergroup conflicts.

In conflicts between social groups, the decision of competitors whether to attack/retreat should be based on the assessment of the quantity of individuals in their own and the opposing group. Experimental studies on numerical cognition in animals suggest that they may represent both large and small numbers as noisy mental magnitudes subject to scalar variability, and small numbers (≤4) also as discrete object-files. Consequently, discriminating between large quantities, but not between smaller ones, should become easier as the asymmetry between quantities increases. Here, we tested these hypotheses by recording naturally occurring conflicts in a population of free-ranging dogs, Canis lupus familiaris, living in a suburban environment. The overall probability of at least one pack member approaching opponents aggressively increased with a decreasing ratio of the number of rivals to that of companions. Moreover, the probability that more than half of the pack members withdrew from a conflict increased when this ratio increased. The skill of dogs in correctly assessing relative group size appeared to improve with increasing the asymmetry in size when at least one pack comprised more than four individuals, and appeared affected to a lesser extent by group size asymmetries when dogs had to compare only small numbers. These results provide the first indications that a representation of quantity based on noisy mental magnitudes may be involved in the assessment of opponents in intergroup conflicts and leave open the possibility that an additional, more precise mechanism may operate with small numbers.

Management of feral domestic cats in the urban environment of Rome (Italy).

In Italy, which is rabies-free, the national Law No. 281 [Legge Nazionale 14 agosto 1991. No. 281: Legge Quadro in materia di animali di affezione e prevenzione del randagismo. Gazz. Uff. Rep. Ital. no 203 del 30 agosto 1991: p. 3] on the management of pets and on the control of feral cats has introduced the no-kill policy for this species. Thus, "trap-neuter-release" (TNR) programs have been carried out for >10 years. In this paper we present data on registered colonies and censused cats in Rome from 1991 to 2000; the results of the neutering campaign from 1991 to 2000; and a survey, on 103 cat colonies, on the effects of demographic control of urban feral-cat colonies in the city of Rome, carried out by the local Veterinary Public Services (VPS) in collaboration with the associations of cat care-takers. In 10 years almost 8000 were neutered and reintroduced in their original colony. The spay/neuter campaigns brought about a general decrease in cat number but the percentage of cat immigration (due to abandonment and spontaneous arrival) is around 21%. This suggests that all these efforts without an effective education of people to control the reproduction of house cats (as a prevention for abandonment) are a waste of money, time and energy.

Bold attitude makes male urban feral domestic cats more vulnerable to Feline Immunodeficiency Virus.

Individual differences in behaviour are a phenomenon that is more and more attracting the attention of scientists. Among the other reasons, behavioural individuality occurs because selection favours the adoption of different tactics by individuals. It is now widely recognized that within many vertebrate species, individuals vary along an axis the extremes of which are represented by individuals 'bold' and 'shy', sometimes called 'proactive' and 'reactive'. Here we present the case of feral domestic cats (Felis catus L.) living in group in the urban environment where showing bold attitudes is linked to the benefit of a high annual reproductive success but, on the other hand, to a high probability to be infected by the Feline Immunodeficiency Virus (FIV), a lethal disease caused by a retrovirus. In this species, natural selection has probably favoured proactive temperament in spite of the cost represented by getting the disease. In fact, proactive individuals, even if FIV positive, reproduce more than reactive individuals before the last stage of FIV-infection (AIDS) characterized by a loss of immunological defences and subsequent opportunistic infections. Evolutionary implications are discussed.