Juan José Sanz

Successful mixed broods in hole-nesting passerines are relatively rare. Increased competition for nesting sites may result from environmental conditions that constrain the breeding period, bringing in contact species that would normally rarely interact. Meteorological conditions preceding the breeding period of 2024 could have delayed the reproduction of Eurasian Nuthatches Sitta europaea relative to Eurasian Blue Tits Cyanistes caeruleus in our study area. Here we report the first case of a successful mixed brood involving Eurasian Nuthatches and Blue Tits. On 4 May, we observed a Blue Tit hatchling in a Nuthatch nest. The Blue Tit, reared in the foster nest, had smaller tarsus and wing length relative to the average measures of all observed Blue Tit nestlings in 2024. However, its mass was in the normal range, and it fledged successfully, along with five Nuthatch ‘siblings’.

Breeding performance and survival are mechanisms through which animal personality affects fitness. Exploration behaviour (EB) is an easily obtainable personality trait that correlates to multiple life-history traits, thus it has been widely used in animal personality studies. Through in situ Novel Environment exploration tests, we assessed the exploration behaviour of blue tits (Cyanistes caeruleus) during two breeding seasons. A principal component analysis revealed two behavioural axes regarding the exploration behaviour: slow-fast and timid-daring. We found that male blue tits are significantly slower explorers than female blue tits, suggesting that evolutionary processes act unequally across sexes in our study area. To investigate these processes, we assessed the relationship between exploration behaviour and two fitness aspects: reproduction and survival. Slower-exploring males bred in nests with significantly shorter incubation periods, and timid males had higher local return ratios. Interactions between male and female EB revealed that disassortative pairs were more successful breeders than assortative ones. Further research should explore potential evolutionary implications of intersexual EB differences.

The rise of temperatures due to global warming is related to a lengthening of the breeding season in many bird species. This allows more pairs to attempt two clutches within the breeding season, thus finishing their breeding activity later in the season and therefore potentially overlapping these with post–breeding moult. We tested whether this occurred in two Spanish great tit Parus major populations. The proportion of pairs laying second clutches increased from 1 % to 32 % over the study period in one of the populations (Sagunto, 1995–2019), while it did not change in the other (Quintos, 2006–2019; mean 5 %). We did not find any temporal trend for moult start date of late–breeding birds in any population. The proportion of individuals of both sexes that overlapped moult and breeding increased in Sagunto. For this latter population, sex and age, but not clutch type, contributed to the variability in the probability of overlapping in late–breeding individuals, this being higher for first–year males and lower for older females.

Territorial behaviour arises as a strategy of ensuring individuals’ access to a variety of potentially limiting resources. While aggressiveness is a well-studied widespread trait across taxa, the mechanisms that allow for a range of aggressive phenotypes to coexist in the wild remains unclear. In this study, we analyse environmental, social and intrinsic variables that can modulate the expression of different strategies of male–male aggressiveness. Furthermore, through network analysis we explore the role of this trait in the establishment of territories during the breeding season as the intensity of different aggressiveness strategies may limit or grant access to resources. Simulating territorial intrusions during the early incubation period, we assessed the aggressiveness of breeding male blue tits ( Cyanistes caeruleus ). We defined three types of conspecific aggressiveness (nonconfrontational intimidating, nonconfrontational cautious and confrontational) and analysed the effect of habitat structure, territory quality, presence of other breeding species and male condition on the type and intensity of the aggressive display. The results obtained suggest that yearling males rely on intimidating behaviour more than older males, that perform more cautious displays. Furthermore, smaller and heavier males opted for confrontational strategies. The density and nature of neighbours, as well as the territory quality and the habitat structure, also conditioned the intensity and type of display. Surprisingly, the network analysis revealed that the intensity of male–male aggressive displays did not condition the establishment of breeding territories. Our results suggest that aggressiveness is a context-specific trait shaped by a complex array of environmental and intrinsic parameters.

Ambient temperature is assumed to be the major cue used by passerines to synchronize their laying and hatching dates to the expected peak of prey availability. While laying eggs, females are still able to fine‐tune their hatching date following increasing or decreasing patterns of ambient temperature, mostly via changes in incubation onset. The onset of incubation behaviour in relation to the laying sequence could have later consequences for the duration of the incubation period and the extent of hatching asynchrony. Clutch size is also known to affect incubation patterns and might therefore condition potential responses to changing temperatures. In this study we assessed the association of ambient temperature and clutch size with the onset of four different incubation behaviours: partial and full nocturnal incubation, and partial and full diurnal incubation. We also evaluated how the onset and duration of each incubation behaviour might predict the duration of diurnal full incubation and the extent of hatching asynchrony. To achieve our aims, we monitored incubation behaviour using temperature data loggers during the egg‐laying period in three Mediterranean Great Tit Parus major populations in three consecutive years. Our results showed that increasing temperatures were related to an advance of diurnal partial incubation, but not its duration or the onset of full incubation behaviour. We did not find any effect of ambient temperatures on nocturnal incubation. However, females lengthened nocturnal partial incubation and delayed the onset of nocturnal full and diurnal partial incubation when laying larger clutches. Longer diurnal incubation before clutch completion was associated with greater hatching asynchrony. Moreover, longer diurnal partial incubation shortened the duration of the full incubation period. In conclusion, increasing ambient temperatures during the egg‐laying period advanced diurnal partial incubation, indirectly shortening the full incubation period and increasing hatching asynchrony.

The expression of the gene Slc7a11 promotes the antioxidant capacity of cells by providing them with cysteine that can be used for the synthesis of glutathione (GSH), the most important intracellular antioxidant. In melanocytes, intracellular cysteine can also enter melanosomes and get incorporated in the pheomelanin pigment synthesis pathway, thus decreasing cysteine availability for GSH synthesis and potentially creating chronic oxidative stress. We thus hypothesized that a mechanism limiting the use of intramelanocytic cysteine for pheomelanin synthesis in environmental conditions generating oxidative stress may be physiologically advantageous and favored by natural selection. Here we searched for evidence of such a mechanism by comparing the influence of melanocytic Slc7a11 expression on pheomelanin‐based pigmentation in developing Eurasian nuthatch Sitta europaea nestlings from two populations differing in predation risk, a natural source of oxidative stress. Pheomelanin synthesis and pigmentation tended to increase with Slc7a11 expression in the low‐risk population as expected from the activity of this gene, but decreased with Slc7a11 expression in the high‐risk population. The same was not observed in the expression of five other genes influencing pheomelanin synthesis without affecting cysteine availability in melanocytes. The influence of body condition on the intensity of pheomelanin‐based pigmentation also differed between populations, being positive in the low‐risk population and negative in the high‐risk population. The resulting pigmentation of birds was more intense in the high‐risk population. These findings suggest that birds perceiving high predation risk may limit the use of cysteine for pheomelanin synthesis, which becomes independent of Slc7a11 expression. Some birds may have thus evolved the ability to adjust their pigmentation phenotype to environmental stress.

Certain populations of long‐distance migratory birds are suffering declines, which may be attributed to effects of climate change. In this article, we have analysed a long‐term (1991–2015) data set on a pied flycatcher Ficedula hypoleuca population breeding in nest‐boxes in a Mediterranean montane oak forest, exploring the trends in population size due to changes in nestling recruitment, female survival and female immigration. We have related these changes in population parameters to local climate, winter NAO index and to breeding density. During the last 25 yr the population has declined by half, mainly in association with a decrease in nestling mass and structural size which had repercussions on the probability of nestling recruitment to the population. Lower local nestling recruitment in certain years was linked to lower female immigration rate in the same years. On the other hand, the local survival of females remained stable throughout the study period. Laying date and breeding success were negatively affected by local temperatures while breeding, recruitment rate likewise by minimum temperature prior to breeding in April. As minimum April temperatures have increased across the study period, this may have affected recruitment and immigration rates negatively. On the other hand, tarsus length and body mass of nestlings were positively associated with winter NAO index, pointing to more global climatic links. Moreover, there was also a negative temporal trend in body mass of adults, implying increasingly difficult conditions for breeding. Declining recruit production in the study area could be attributed to a mismatch between the timing of arrival and breeding in the population, and the peak of food availability in this area.

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.

Capsule: The nestling diet of Blue Tits Cyanistes caeruleus is influenced by parental effort and habitat type, and consequently has an impact on breeding success. Aims: In a three-year study, we compared the nestling diet of Blue Tits in two Mediterranean forests (pinewood and oakwood) and tested its implications for breeding success. Methods: Adults were captured at the nest to obtain morphological measurements, and provisioning behaviour was recorded when chicks were 11 days old. Nestling tarsus length and body mass were measured on day 13 after hatching. Results: Caterpillars constituted the largest proportion of nestling diet in both habitats, however, higher numbers and biomass of noctuid, as well as higher numbers of tortricid larvae, were provided to nestlings in the pinewood. Furthermore, females provided tortricids more often than males, whereas males supplied more geometrid larvae and spiders. We found a more generalist diet for nestlings raised in the pinewood. Also, a greater number of young fledged when their diet included more tortricids and was more generalist, and Blue Tit nestlings raised on a diet with a higher number of spiders were in better body condition. Conclusion: Differences in nestling diet between habitat types contribute to explain variance in breeding performance and therefore demonstrate diverse foraging behaviour strategies among populations.