Daniela Canestrari

Effect of parentage and relatedness on the individual contribution to cooperative chick care in carrion crows Corvus corone corone

Carrion crows Corvus corone corone in northern Spain form complex cooperative groups that contain non-dispersing offspring and immigrants. Reproduction is often shared among group members, with polyandry prevalent over joint egg laying. However, due to incest avoidance or sexual immaturity, reproduction is potentially available only to a fraction of group members (“potential breeders”), while others do not breed (“non-breeders”). We combined molecular data with video-recorded observations at nests in order to investigate whether potential breeders and non-breeders adjust their individual effort in nestling feeding respectively to their level of parentage and the relatedness with the chicks. Overall, parents of at least one chick in the brood showed the highest feeding rates. Among potential breeders, genetic mothers and fathers fed chicks at significantly higher rates than individuals with no parentage, but they did not adjust their effort according to the proportion of offspring generated. Current direct fitness benefits are therefore important in determining a high provisioning effort, but crows may lack a mechanism to finely assess their share of parentage. Among non-breeders, males contributed more than females to chick feeding, but we found no significant correlation between feeding rate and relatedness to the nestling. We discuss how the latter result can be reconciled with the fact that kin selection has been shown to be important in shaping the crow cooperative society.

Experimentally increased food resources in the natal territory promote offspring philopatry and helping in cooperatively breeding carrion crows.

Kin-based societies, where families represent the basic social unit, occur in a relatively small number of vertebrate species. In the majority of avian kin societies, families form when offspring prolong their association with the parents on the natal territory. Therefore, the key to understanding the evolution of families in birds is to understand natal philopatry (i.e. the tendency to remain on the natal territory). It has been shown that, within populations, the strength of the association between parents and offspring (i.e. family stability) increases when offspring dispersal is constrained by external environmental factors, but it is unclear whether and how family wealth influences juvenile dispersal decisions. Here, we show that young carrion crows (Corvus corone corone) from territories that were food-supplemented year-round were more philopatric and more likely to help at their familys nest than the unfed ones. The results suggest that offspring philopatry and helping behaviour are influenced by the quality of ‘home’ and that the availability of food resources positively affects the cohesion of the family.

COOPERATIVELY BREEDING GROUPS OF CARRION CROW(CORVUS CORONE CORONE) IN NORTHERN SPAIN

Abstract The Carrion Crow (Corvus corone) is almost invariably reported as a species breeding as unassisted pairs, but a population of Carrion Crows (Corvus c. corone) from northern Spain that we have been studying since 1995 regularly exhibits cooperative breeding. The Spanish population thus demonstrates that cooperative breeding can vary greatly across populations. Most of the breeding territories (73.3%) in Spain were held by cohesive groups, which consisted of up to nine birds (mode = 3 birds). The proportion of fledglings that delayed dispersal for one year varied between 12.2 and 47.5% for different cohorts. The corresponding values for juveniles postponing dispersal for two years were 4.9 and 19.2%. Philopatry was skewed toward males. Some social groups contained immigrants. Therefore, delayed juvenile dispersal was not the only route towards sociality. Up to three helpers provided food to the nestling and at least one helper was found in all the groups surveyed. However, some group members refrained from visiting nests and thus the contribution to nestling care was highly variable among individuals.

Reproductive success increases with group size in cooperative carrion crows, Corvus corone corone

The effect of group size and the number of helpers on reproductive success is crucial to understand the evolution and maintenance of cooperative breeding. In northern Spain, carrion crows form kin-groups (three to nine individuals) where up to five individuals contribute to rearing the young. Using data from 99 different territories, followed over 10 years, we showed that group size was positively correlated with the annual number of fledglings produced, after controlling for the potential confounding effect of territory quality. This occurred through: (1) an increased probability for larger groups of renesting after early nest failure; (2) a higher probability of nest success; and (3) a higher number of fledglings produced in successful attempts. Video-recorded observations at the nests showed that chicks received more food in larger groups during the first 10 days of life, when the risk of starvation is highest, suggesting a role of allofeeding by additional carers in augmenting the reproductive success of a group. In crows, indirect and direct benefits through increased production of young are therefore available to nonbreeding group members.

From Parasitism to Mutualism: Unexpected Interactions Between a Cuckoo and Its Host

Predation Favors Parasitism Parasitism in birds often results in ejection or starvation of the hosts nestlings. Consequently, many host bird species have evolved protective behavior such as mobbing and parasite egg rejection. Curiously, some host species show no parasite avoidance behaviors; for example, the crow Corvus corone corone tolerates cuckoo chicks among its own brood. In a long-term study, Canestrari et al. (p. 1350) found that crow nests containing a cuckoo chick had lower rates of predation because the parasites chicks secrete a noxious repellent substance. Overall, in years of high predation pressure, the presence of cuckoos improves the crows breeding success, but when there are fewer predators around, parasitism reduces crow fitness. The carrion crow Corvus corone corone can benefit from parasitism by the great spotted cuckoo Clamator glandarius. Avian brood parasites lay eggs in the nests of other birds, which raise the unrelated chicks and typically suffer partial or complete loss of their own brood. However, carrion crows Corvus corone corone can benefit from parasitism by the great spotted cuckoo Clamator glandarius. Parasitized nests have lower rates of predation-induced failure due to production of a repellent secretion by cuckoo chicks, but among nests that are successful, those with cuckoo chicks fledge fewer crows. The outcome of these counterbalancing effects fluctuates between parasitism and mutualism each season, depending on the intensity of predation pressure.

History, environment and social behaviour: experimentally induced cooperative breeding in the carrion crow

Kin–based cooperative breeding, where grown offspring delay natal dispersal and help their parents to rear new young, has a long history in some avian lineages. Family formation and helping behaviour in extant populations may therefore simply represent the retention of ancestral features, tolerated under current conditions, rather than a current adaptive process driven by environmental factors. Separating these two possibilities challenges evolutionary biologists because of the tight coupling that normally exists between phylogeny and the environmental distribution of species and populations. The carrion crow Corvus corone corone, which exhibits extreme interpopulational variation in the extent of cooperative breeding, with populations showing no delayed dispersal and helping at all, provides a unique opportunity for an experimental approach. Here we show that offspring of non–cooperative carrion crows from Switzerland will remain on the natal territory and express helping behaviour when raised in a cooperative population in Spain. When we transferred carrion crow eggs from Switzerland to Spain, five out of six transplanted juveniles delayed dispersal, and two of those became helpers in the following breeding season. Our results provide compelling experimental evidence of the causal relationship between current environmental conditions and expression of cooperative behaviour.

Costs of chick provisioning in cooperatively breeding crows : an experimental study

The costs of providing care to offspring and how these costs influence individual allocation of resources to current and future reproduction are likely to be important in the evolution of cooperative breeding. We reduced the costs of offspring care experimentally in 12 groups of cooperatively breeding carrion crows, Corvus corone corone, by providing supplementary food throughout the breeding season. Neither nonbreeders nor breeders from supplemented territories significantly increased their levels of provisioning effort. However, unfed crows lost more weight than fed crows and, in contrast to fed crows, lost weight in relation to provisioning effort. Furthermore, breeders decreased their provisioning rates in groups with more than three caregivers, supporting both the idea that provisioning chicks is costly and the conclusion that crows invest in self-maintenance rather than in the current brood when costs are reduced.

False feedings at the nests of Carrion crows Corvus corone corone

False feedings, when individuals visit the nest but refrain from feeding the chicks, occur in some cooperative species and have been interpreted in the white-winged chough (Corcorax melanorhamphos) as active deception by helpers towards the rest of the group. In a cooperatively breeding population of carrion crows (Corvus corone corone) 81.5% of the individuals that provided nestling care showed various kinds of false feedings: arriving at the nest with no food, consuming part or all the food brought to the nest, or taking back from a chick’s gape the food that had just been delivered. False feedings occurred on average during 16.3% of nest visits, with some individuals performing them at very high rates (up to 64% of nest visits). False feedings occurred at similar rates in unassisted pairs and groups with helpers, and breeding females showed false feeding at significantly higher rates than other group members. Furthermore, individuals showed false feedings regardless of whether they were alone on the nest or in the presence of other group members, and false feedings did not provoke aggression by the rest of the group. False feedings are not likely to represent deceptive help in the carrion crow. We suggest that crows evaluate the chicks’ condition during nest visits and that false feedings occur as result of a trade-off between their own hunger and the chicks’ needs.

High begging intensity of great spotted cuckoo nestlings favours larger-size crow nest mates

In nests of birds parasitized by a larger non-evicting brood parasite, host chicks typically are at disadvantage in competing for food and often starve. However, when host chicks are larger, they may benefit from the presence of the parasite, which contributes to the net brood begging signal but cannot monopolize the food brought to the nest. Here, we show that, despite a higher begging intensity, great spotted cuckoos (Clamator glandarius) did not outcompete larger size carrion crow (Corvus corone corone) nestlings. Furthermore, cuckoos’ exaggerated begging allowed crow nest mates to decrease their begging intensity without negative consequences on food intake. Assuming an energetic cost to chicks of begging intensely, our results suggest that crow chicks sharing the nest with a cuckoo may obtain an advantage that should be weighed against the loss of indirect fitness due to parasitism.

Spatiotemporal variation of host use in a brood parasite: the role of the environment

Avian brood parasites should target the most profitable host species, but current conditions might locally influence their choice, producing geographic mosaics of coevolution. Throughout Europe, the magpie Pica pica has been invariably reported as the primary host of the great spotted cuckoo Clamator glandarius, whereas the carrion crow Corvus corone is the secondary one. However, we found that this pattern reversed in northern Spain, where up to 70% of carrion crow nests were parasitized versus 20% of magpie nests. In southern Spain, conversely, parasitism increased proportionally in both hosts (up to approximately 90% of available nests) throughout the 3 years of study. Surprisingly, magpies provided the best reproductive output for cuckoos in both areas, in contrast with cuckoo’s preference for the crow host in the north. Genetic data ruled out the presence of different host-specific races in this brood parasite, dismissing the hypothesis that a prevalence of different gentes at the 2 sites explained the observed variability in host choice. Instead, we found that magpie nests in the south were easier to reach and more scattered than in the north, where cuckoos preferentially targeted nests that were less concealed and more isolated. We suggest that the habitat constraints parasitism on magpies in the north, driving cuckoo host choice toward the crows. The coevolutionary scenario therefore includes a 3-way interaction, where the pressure that the parasite puts on a host species in a given place critically depends on the environmentally mediated interaction between the same parasite and a different host.