The effect of climate change on avian offspring production: A global meta-analysis.

Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young.

Fitness benefits of alternated chick provisioning in cooperatively breeding carrion crows.

In most bird species, parents raise offspring cooperatively. In some cases, this cooperation extends to helpers-at-the-nest who assist the breeders with a range of tasks. While cooperative food provisioning might merely arise incidentally, as a result of the efforts of carers that act independently from each other, recent studies suggest that birds may coordinate by taking turns in visiting the nest. However, evidence that such coordination emerges because individuals actively respond to each other's behaviour is controversial, and the potential benefits of carers' alternation remain unknown. We addressed this knowledge gap by analysing a multiyear dataset for cooperatively breeding carrion crows Corvus corone, comprising 8693 nest visits across 50 groups. Our results reveal that turn-taking does occur in this species and that all group members, regardless of their sex and social role (breeder/helper), tend to alternate at the nest with other carers rather than to make repeat visits. Importantly, we found that the body mass of nestlings increased significantly with the degree of carers' alternation, possibly because well-coordinated groups provided food at more regular intervals. Using earlier monitoring data, the observed increase in body mass is predicted to substantially boost postfledging survival rates. Our analyses demonstrate that alternation in nestling provisioning has measurable fitness benefits in this study system. This raises the possibility that cooperatively breeding carrion crows, as well as other bird species with similarly coordinated brood provisioning, exhibit specialized behavioural strategies that enable effective alternation.

En la mayoría de las especies de aves, los padres cooperan entre sí en el cuidado de sus crías. En algunos casos, la cooperación se extiende a individuos ayudantes que asisten a los reproductores en diferentes tareas. El aprovisionamiento cooperativo de las crías puede darse simplemente como resultado del esfuerzo de cuidadores que actúan de forma independiente, pero estudios recientes sugieren que las aves pueden coordinarse, tomando turnos a la hora de visitar el nido. Sin embargo, las evidencias de que dicha coordinación ocurre porque los individuos realmente responden al comportamiento de los demás es controvertida, y los potenciales beneficios de la alternancia al nido son desconocidos. Nosotros abordamos estos aspectos analizando una base de datos plurianual en la corneja negra Corvus corone, que incluye 8693 visitas al nido en 50 grupos. Nuestros resultados revelan que se produce toma de turnos en esta especie y que todos los miembros del grupo, independientemente del sexo y del estatus social (reproductor / ayudante), tienden a alternarse al nido con otros cuidadores, en vez de repetir visitas. Más importante aún, encontramos que la masa corporal de los polluelos aumentaba de forma significativa al aumentar el grado de coordinación de los cuidadores, posiblemente porque los grupos mejor coordinados aprovisionaban las crías a intervalos más regulares. Basándonos en datos anteriores, pudimos calcular que el incremento observado en masa corporal predice un aumento sustancial de la tasa de supervivencia de los volanderos. Nuestros análisis demuestran, por lo tanto, que la alternancia al nido tiene beneficios medibles en eficacia biológica. Esto conlleva la posibilidad de que las cornejas negras cooperativas, al igual que otras especies de aves con aprovisionamiento coordinado de las crías, exhiban estrategias comportamentales especializadas que permitan una eficiente alternancia al nido.

Effects of meteorological conditions on brood care in cooperatively breeding carrion crow and consequences on reproductive success.

Meteorological stressors (e.g., temperature and rain shortage) constrain brood provisioning in some bird species, but the consequences on reproductive success have been rarely quantified. Here we show, in a cooperatively breeding population of carrion crow Corvus corone in Spain, that individual feeding rates decreased significantly with rising air temperatures both in breeders and helpers, while lack of rain was associated with a significant reduction in the effort of the male helpers as compared to the other social categories. Group coordination, measured as the degree of alternation of nest visits by carers, was also negatively affected by rising temperature. Furthermore, we found that the body condition of the nestlings worsened when temperatures were high during the rearing period. Interestingly, the analysis of a long-term data set on crow reproduction showed that nestling body condition steadily deteriorated over the last 26-years. Although many factors may concur in causing population changes, our data suggest a possible causal link between global warming, brood caring behaviour and the decline of carrion crow population in the Mediterranean climatic region of Spain.

Carrion crows learn to discriminate between calls of reliable and unreliable conspecifics.

Partner choice on the basis of an individual's reliability is expected to stabilize social interactions. In this experiment, we tested whether carrion crows (Corvus corone corone) learn to differentiate between calls of reliable or unreliable individuals. Crows were kept in an aviary that comprised four visually but not acoustically isolated compartments, separated by a central room. In an association phase, a dead crow placed in the central compartment was visible only to one of the four crow groups, whilst alert calls of a conspecific were played back. Therefore, these calls were reliable for that group, but unreliable for the three other groups. The procedure was repeated, using a different reliable caller for each group. In two test sessions, 1 month apart, reliable and unreliable model individuals were played back, but no dead crow was presented. We quantified birds' attention behaviour and the number of vocalisations emitted. In the association phase, crows were more attentive towards the reliable compared with the unreliable stimuli and called more in response to reliable compared to unreliable individuals. In the test and repeat phase, attention behaviour did not differ between reliability conditions, but the pattern of vocal behaviour reversed, with crows calling less frequent when listening to reliable compared with unreliable calls. Vocal responses of crows suggest that they can discriminate between reliable and unreliable callers.

Chicks of the great spotted cuckoo may turn brood parasitism into mutualism by producing a foul-smelling secretion that repels predators.

The great spotted cuckoo (Clamator glandarius) is an important brood parasite of carrion crows (Corvus corone corone) in northern Spain. We recently found that, unlike what is commonly known for cuckoo-host interactions, the great spotted cuckoo has no negative impact on average crow fitness in this region. The explanation for this surprising effect is a repulsive secretion that the cuckoo chicks produce when they are harassed and that may protect the brood against predation. Here, we provide details on the chemical composition of the cuckoo secretion, as well as conclusive evidence that the dominating volatile chemicals in the secretion are highly repellent to model species representative of common predators of the crows. These results support the notion that, in this particular system, the production of a repulsive secretion by the cuckoo chicks has turned a normally parasitic interaction into a mutualistic one.

Lazy group members are substitute helpers in carrion crows.

In many cooperatively breeding societies, helping effort varies greatly among group members, raising the question of why dominant individuals tolerate lazy subordinates. In groups of carrion crows Corvus corone corone, helpers at the nest increase breeders' reproductive success, but chick provisioning is unevenly distributed among non-breeders, with a gradient that ranges from individuals that work as much as the breeders to others that completely refrain from visiting the nest. Here we show that lazy non-breeders represent an insurance workforce that fully compensates for a reduction in the provisioning effort of another group member, avoiding a decrease in reproductive success. When we temporarily impaired a carer, decreasing its nest attendance, the laziest non-breeders increased their provisioning rate and individuals that initially refrained from visiting the nest started helping. Breeders, in contrast, did not increase chick provisioning. This shows that lazy non-breeders can buffer a sudden unfavourable circumstance and suggests that group stability relies on the potential contribution of group members in addition to their current effort.

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 family's 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.

Behavioural responses to olfactory cues in carrion crows.

Until recently, the use of olfactory signals in birds has been largely ignored, despite the fact that birds do possess a fully functioning olfactory system and have been shown to use odours in social and foraging tasks, predator detection and orientation. The present study investigates whether carrion crows (Corvus corone corone), a bird species living in complex social societies, respond behaviourally to olfactory cues of conspecifics. During our experiment, carrion crows were observed less often close to the conspecific scent compared to a control side. Because conspecific scent was extracted during handling, a stressful procedure for birds, we interpreted the general avoidance of the 'scent' side as disfavour against a stressed conspecific. However, males, unlike females, showed less avoidance towards the scent of a familiar individual compared to an unfamiliar one, which might reflect a stronger interest in the information conveyed and/or willingness to provide social support.

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.

From parasitism to mutualism: unexpected interactions between a cuckoo and its host.

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.

Kin selection in cooperative alliances of carrion crows.

In most cooperative vertebrates, delayed natal dispersal is the mechanism that leads to the formation of kin societies. Under this condition, the possibility that kin-based cooperative breeding is an unselected consequence of dispersal patterns can never be ruled out because helpers can only help their relatives. Here we show that a population of carrion crows (Corvus corone corone) fully fits the central prediction of kin selection theory that cooperative breeding should arise among relatives. On their territory, resident breeders are aided not only by nonbreeding retained offspring but also by immigrants (mainly males), with whom they share matings. Philopatry cannot account, however, for the high degree of genetic relatedness found between breeders and immigrants of the same sex that cooperate at a nest, indicating that crows actively choose to breed cooperatively with their relatives.