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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.
FULL pack 109 fuck little bitches 10
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In this study, we investigated the social variables affecting both male and female mate preferences in a pack of free-ranging dogs (Canis familiaris), i.e. those domestic dogs whose movements, activities and reproduction are not constrained by human beings and that, according to recent studies, may actually represent the most numerous category of domestic dogs in the world [58]. In areas where they have access to abundant food resources directly or indirectly provided by human beings, free-ranging dogs can live in stable packs formed by multiple breeding individuals of both sexes [59], [60], [61], [62], [63], [64], [65], [66], [67], [68], [69], [70]. There are several claims in the scientific literature that, due to the effects of domestication on their behaviour, free-ranging dogs are unable to form structured social groups and retain very little of the social organization of wolves (Canis lupus), that are their closest living relatives [71], [72], [73], [74]. In particular, the presence of multiple breeding individuals in dog groups seems in sharp contrast with the structure of wolf family groups, usually comprising a single dominant breeding pair and a number of subordinate non-breeding helpers [75], [76], [77], and has led researchers to conclude that domestic dogs lack any social regulation of reproductive activities [72], [74]. However, recent studies [65], [68], [70] have demonstrated that free-ranging dogs show a complex social organization characterized by age-graded dominance hierarchies in which males tend to be dominant over females of similar age, although females often are dominant over younger males. Furthermore, dominance relationships are expressed both in agonistic interactions and in affiliative greeting ceremonies [65] and, as in wolves [78], older dominant individuals usually lead the collective movements of the pack [68], [70]. Nevertheless, it remains to be demonstrated whether dominance relationships in dogs result in some kind of social control of reproduction within the group, as has been observed in wolves (see references above).
In our dog pack, the dominance rank of males achieved through competition outside the oestrous period [65] affected the direct competition among males during courtship: subordinate males had lower copulation success partly because they were intimidated by dominant males who often did not need to attack to keep them away from females. Nevertheless, our results indicate that male intra-sexual competition also affected female preferences. Our data on male-female affiliative interactions during oestrus suggest that bitches prefer to mate with high-ranking males. In particular, oestrus bitches approached high-ranking males at a significantly higher rate than they approached low-ranking males, thus showing a preference for maintaining a close proximity with dominant males. Moreover, high-ranking males were also those who showed more frequently affiliative behaviour towards oestrus females, and this strategy apparently was successful because they achieved a higher frequency of copulation than low-ranking dogs (who showed lower levels of affiliation to oestrus females). Conversely, male aggressiveness towards females during oestrous did not seem to function effectively to constrain female mate preference. Although, unlike other studies [80], [81], [86], we did not observe male dogs forcing females into mating, some males used aggression to try to intimidate females that refused to mate with them. Moreover, sometimes males showed aggressive behaviour towards females who were attempting to mate with other males. Females typically refused to mate with these aggressive males by actively avoiding, attacking or chasing them. In particular, our results show that old high-ranking females were more aggressive than young subordinate females in refusing male mount attempts.
Our results also suggest that female intra-sexual competition affected male mate preferences. Using male mating effort as a proxy for male mate choice, we found that male dogs did not distribute this effort evenly among oestrus females. In fact, male courting effort and male-male competition for receptive females (measured as rate of mount attempts and proportion of interrupted mounts) appeared to be stronger in the presence of higher-ranking females, suggesting that males prefer to mate with dominant females. It is possible that male preferences for dominant females are functional because high-ranking females, based on our study, seem to have higher reproductive success than low-ranking ones. Rank-related reproductive asymmetries in bitches may result from competition for food resources in which high-ranking females usually prevail over subordinates [65], [70]. Another speculation is that the differential reproductive success may be a consequence of infanticide by dominant females. Although we never observed infanticide in our population of dogs, it has been documented in several canid species (e.g. Canis lupus [108], Canis latrans [109], Canis aureus [110]), and also in captive dingoes [111] that descend from domestic dogs [112]. Avoidance of intra-sexual competition may explain why, during our study and in others [60], some pregnant females dispersed from the pack before giving birth. Although the number of dispersing females in our study was too low to allow a statistical comparison, it is worth to note that they obtained a moderate reproductive success, whereas among resident females only the highest in rank had a considerable reproductive success (see Table 1). However, we cannot rule out that, in our study, some kind of undetected paternal investment by high-ranking males might have increased the reproductive success of their preferred females. So, an alternative interpretation of our results is that high-ranking females had higher reproductive success than low-ranking ones because high-ranking males provided them with paternal investment in exchange for being preferred as mates.
The presence of multiple breeding individuals in dog packs might be explained functionally as an adaptive consequence of the domestication process. As suggested by several authors [58], [120], [121], unrestricted dog populations have adapted to scavenge from human refuses that are abundant and do not follow marked seasonal fluctuations. This continuous availability of food may have favored the loss of seasonal reproductive behavior, and may have allowed dogs to reproduce in their natal pack during the first year of their life, once they reach full body weight [58]. Furthermore, the abundance of food resources experienced by free-ranging dogs may have led to a decrease in the level of within group competition for food and in the degree of reproductive suppression of subordinates [70]. Conversely, unlike dogs, wolves are seasonal breeders and they rarely reach sufficient body size to reproduce until their second breeding season at the age of about 22 months, when they usually disperse from their natal pack [58], [122], [123], [124], [125]. However, wolf packs with multiple breeders can be found where food resources are unusually abundant and some individuals delay dispersal [76], [77], [112], [126], [127], [128], [129], [130], [131]. On the other hand, in a pack of feral dogs studied in an area with harsh weather and limited food availability, only one female gave birth during a two years period, and pup rearing apparently was shared by several group members [132]. Although detailed data about the social relationships among the members of this group are lacking, this example suggests that, under some extreme ecological conditions, dogs can form packs whose structure may be even more similar to that of wolf packs, and highlights the considerable social flexibility of this species.
The adaptation of dogs to exploit human refuse as a food resource may also account for the apparent reduction in allofeeding of lactating mothers observed in this species relative to wolves [58], [70]. Lactating female wolves spend a considerable portion of their time with pups at the denning site, and they are provisioned with food by the other pack members who perform most of the hunting [119], [133], [134], [135], [136]. Unlike wolves, most free-ranging dogs do not need to hunt to feed, and they can often rely on food sources that are presumably more predictable in terms of location and time than wolves' prey [70]. This usually allows lactating bitches to place their dens in the vicinity of human refuse, and thus to join their pack during feeding, while reducing the time during which pups are left alone [70]. So, the ecological conditions found in a domestic environment may have driven the evolution of an increased independency of bitches in raising their pups relative to female wolves. 041b061a72