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In species with biparental care, the amount of care devoted to offspring is affected by the negotiation rules that the parents adopt. Theoretical models predict that biparental care can be evolutionarily stable if a decrease in parental investment by one parent is only partially compensated by its partner. However, empirical studies have found substantial variability in compensatory behaviour and have mainly used nesting provisioning as a single measure of parental effort. In this study, we investigated parental compensatory behaviour for two parental tasks, offspring provisioning and nest defence. These two tasks are likely characterized by different levels of risk as well as different cost and benefit functions for the parents, which may affect the expected level of compensatory responses. We experimentally widowed (by temporarily removing one parent) male or female Spanish sparrows, Passer hispaniolensis, and measured their compensatory responses to offspring provisioning and nest defence (after predator presentation of green whip snake, Hierophis viridiflavus, models). Parents differed in their compensatory responses in relation to parental task and sex: both sexes partially compensated for offspring provisioning, but females compensated by a larger degree than males. For predator defence, males instead decreased defence behaviour by increasing latency and reducing the number of attacks, while females did not change their behaviour when caring alone. This within-individual comparison indicates that parents adjust their compensatory behaviour according to parental task. We discuss how these differences could arise due to different costs and benefits of extra investments.

Parasites are known to be a key driving force in mate choice and are important for the expression and evolution of ornaments and behavioral traits being used. However, there is little experimental evidence on how the parasite’s burden of the choosing individual is integrated into the mate-choice process and how it affects decision-making, especially in relation to parasite infestation of potential mates. Thus, the aim of our study was to determine whether female house sparrows Passer domesticus adjust their mate preference according to their own as well as the parasite load of prospective partners. To do this, we experimentally manipulated female parasite load and determined their mate preferences prior to and after parasite treatment. We manipulated the chronic coccidian parasite burden of females either by initiating the acute infection phase via re-infecting them with coccidian or by temporally reducing the parasite load of coccidia. We then measured the effect of this manipulation on mate preference by presenting females with a choice of four stimuli: three males with similar ornaments, but unmanipulated, naturally varying chronic coccidiosis levels, and an unmanipulated control female. Additionally, we recorded some males’ behavior in relation to their infection status pointing toward an increased or reduced interest in mating. We found that females preferred highly infested males prior to manipulation, regardless of their own infestation level. However, after manipulation, infested females avoided highly infested males probably in response to the deterioration of their health condition by parasites. Our study suggests that mate-choice decisions are more complex when they are mediated by parasites. The implications of parasites for evolutionary theories of sexual signaling and mate choice are discussed.

Abstract Research on several social fishes has revealed that shoals constituted by familiar individuals behave remarkably differently compared to shoals formed by unfamiliar individuals. However, whether these behavioural changes may arise also in shoals composed by a mixture of familiar and unfamiliar individuals, a situation that may commonly occur in nature, is not clear. Here, we observed the behaviour of Mediterranean killifish (Aphanius fasciatus) shoals that were composed by both familiar and unfamiliar individuals (i.e. individuals were familiar to each other in pairs) and compared it with shoals entirely made by either unfamiliar or familiar individuals. Shoals formed by familiar individuals took longer to emerge from a refuge and swam more cohesively compared to shoals formed by unfamiliar fish. Shoals formed by a mixture of familiar and unfamiliar individuals behaved as shoals formed by unfamiliar individuals. Moreover, mixed shoals did not segregate in pairs according to their familiarity. This study suggests that mixed shoals do not show the behavioural effects of familiarity. Significance statement Laboratory studies have compared the behaviour of shoals formed by familiar fish versus shoals formed by unfamiliar fish, finding notable advantages in the former ones, such as improved antipredator and foraging behaviour. However, comparing these two opposite shoal types may not provide information on the natural situation, because in nature, shoals often change composition. We investigated how shoals formed by a mixture of familiar and unfamiliar fish behaved. We analysed shoals’ preference for open environment versus covers and shoals’ swimming cohesion. Results showed that shoals formed by both familiar and unfamiliar individuals mostly behave like shoals entirely formed by unfamiliar individuals. This suggests that the advantages of social groups formed by familiar fish might be hardly seen in nature for species in which shoal composition changes frequently.

The environment that parents experience can influence their reproductive output and their offspring’s fitness via parental effects. Perceived predation risk can affect both parent and offspring phenotype, but it remains unclear to what extent offspring behavioral traits are affected when the mother is exposed to predation risk. This is particularly unclear in live-bearing species where maternal effects could occur during embryogenesis. Here, using a half-sib design to control for paternal effects, we experimentally exposed females of a live-bearing fish, the guppy (Poecilia reticulata), to visual predator cues and conspecific alarm cues during their gestation. Females exposed to predation risk cues increased their antipredator behaviors throughout the entire treatment. Offspring of mothers exposed to the predation stimuli exhibited more pronounced exploratory behavior, but did not show any significant differences in their schooling behavior, compared to controls. Thus, while maternally perceived risk affected offspring’s exploration during early stages of life, offspring’s schooling behavior could be influenced more by direct environmental experience rather than via maternal cues. Our results suggest a rather limited role in predator-induced maternal effects on the behavior of juvenile guppies.

Background Consistent inter-individual differences in behavioural phenotypes may entail differences in energy efficiency and expenditure, with different fitness payoffs. In colonial-breeding species, inter-individual differences in foraging behaviour may evolve to reduce resource use overlap among conspecifics exploiting shared foraging areas. Furthermore, individual differences in foraging behaviour may covary with individual characteristics, such as sex or physiological conditions. Methods We investigated individual differences in foraging tactics of a colonial raptor, the lesser kestrel (Falco naumanni). We tracked foraging trips of breeding individuals using miniaturized biologgers. We classified behaviours from GPS data and identified tactics at the foraging trip level by cluster analysis. We then estimated energy expenditure associated to each tactic from tri-axial accelerometer data. Results We obtained 489 foraging trips by 36 individuals. Two clusters of trips were identified, one (SF) characterized by more static foraging behaviour and the other (DF) by more dynamic foraging behaviour, with a higher proportion of flying activity and a higher energy expenditure compared to SF. Lesser kestrels showed consistent inter-individual differences in foraging tactics across weather condition gradients, favouring DF trips as solar radiation and crosswind intensity increased. DF trips were more frequent during the nestling-rearing than during the egg incubation stage. Nestlings whose tracked parent was more prone to perform DF trips experienced higher daily mass increase, irrespective of nestling feeding rates. Conclusions Our study provided evidence that breeding lesser kestrels flexibly adopted different foraging tactics according to contingent weather landscapes, with birds showing consistent inter-individual differences in the tendency to adopt a given tactic. The positive correlation between the tendency to perform more energy-demanding DF trips and nestling growth suggests that individual differences in foraging behaviour may play a role in maintaining key life-history trade-offs between reproduction and self-maintenance.

Raising a family is not free of conflict for parents, as each parent benefits when its partner provides more care for the offspring. Resolving this conflict requires cooperation between the parents. One way to achieve such cooperation might be to coordinate parental provisioning by synchronizing (i.e. returning to the nest at the same time) or alternating (i.e. taking turns) offspring provisioning at the nest. Empirical studies in birds indicate that pair coordination of the nest visits is common; however, it is unknown whether this behaviour is directly related to different outcomes of sexual conflict, such as brood desertion. We used the rock sparrow, Petronia petronia, a species with high levels of sexual conflict, to explore whether alternation and synchrony of the nest visits were related to later brood desertion. Pairs with no desertion alternated and synchronized their nest visits more than pairs in which one sex deserted. This difference in coordination was not simply a by-product of differences in provisioning by the parents. Synchrony of the visits also increased with offspring age in the pairs with no desertion. We provide evidence, for the first time to our knowledge, that the degree of parental coordination is strongly associated with the ultimate consequence of sexual conflict, brood desertion, supporting the idea that coordination in parental behaviour might promote conflict resolution.

Individuals can behave as either leaders or followers in many taxa of collectively moving animals. Leaders initiate movements and may incur predation risks, while followers are thought to be more risk-averse. As a group encounters different challenges and ecological situations, individuals in the group may change their social role. We investigated leader and follower roles using dyads of captive house sparrow (Passer domesticus) during both exploration of a novel environment and a simulation of predator attack. During the exploration of a novel environment, individuals behaved consistently either as leaders or followers. However, in the simulated attack tests, individuals in the dyads switched their roles, with “followers” leading the escape flights and “leaders” following them. Our study provides evidence of 1) consistent differences between individuals in behavior during social escape and 2) a relationship between social roles across different situations. We suggest that such relationship hinges on individual risk-taking tendencies, which manifest through different social roles across different ecological situations. We further speculate that risk-taking individuals might gain benefits by following risk-averse individuals during an escape flight.

Living in groups is widespread and multiple attributes can affect grouping decisions. In several taxa, social familiarity mediates group formation and maintenance, while phenotypic homogeneity within groups may enhance antipredator responses by exploiting the confusion effect. Although familiarity and homogeneity are crucial attributes for social life, little is known about their relative importance. We used a social fish species, the Mediterranean killifish (Aphanius fasciatus), in which females preferentially shoal with groups characterised by phenotypic homogeneity in the number of vertical bars along the body and explore efficiently a novel environment when shoaling with familiar individuals. We asked if preference for groups that are highly phenotypically homogeneous overrides the preference for familiar groups in a species in which both social familiarity and phenotypic homogeneity are important. We first tested whether fish prefer to associate with a familiar rather than with an unfamiliar shoal while controlling for the phenotypic homogeneity in the number of bars of the stimuli, and we found, as expected, that fish preferentially associated with the familiar group. Secondly, we investigated whether fish prefer to associate with a highly phenotypically homogeneous but unfamiliar group rather than with a phenotypically non-homogeneous but familiar group. Interestingly, we found that fish associated with the group characterised by higher within-shoal homogeneity, irrespective of social familiarity. This finding suggests that homogeneity in body pigmentation may affect shoal assortments more than previously considered and opens further questions concerning the ecological implications, such as predator–prey interactions, of within-shoal homogeneity in body pigmentation. How different group attributes interact to ultimately affect grouping decisions is still an exciting open question. In social fish multiple attributes are known to affect shoaling decisions but data on the relative importance of these attributes is limited. In a social killifish (Aphanius fasciatus) characterized by conspicuous vertical bars along the body we evaluated the relative importance of two crucial group attributes for shoaling decisions: homogeneity in the number of bars and familiarity between individuals. We provide an empirical demonstration that social preference based on familiarity can be completely reversed when phenotypic homogeneity is introduced as a criterion for group choice. A flexible shoaling behaviour provides the opportunity for individuals to maximize the benefits of group membership in response to ecological conditions. Context-dependent shoaling decisions could provide an explanation for the maintenance of grouping preference based on multiple attributes.

Female promiscuity can function to acquire both direct and indirect benefits from their social mate and extra-pair males. In many raptor species, intense mate-feeding significantly contributes to female energy requirements before and during egg laying. Moreover, females may use mate-feeding effort to assess male quality. In this study of the lesser kestrel Falco naumanni, we aimed at experimentally manipulating the female's perception of mate quality by providing females with extra food during egg laying, and evaluated the occurrence of extra-pair paternity in food-supplemented and control broods by parentage analyses. No extra-pair offspring (EPO) was found among 19 food-supplemented broods, whereas EPO occurred in five out of 17 control broods. No significant differences in morphological traits, body condition and reproductive success were found between faithful and unfaithful females. However, clutches containing EPO were laid later in the breeding season. Moreover, un-cuckolded males had longer tarsi than cuckolded ones, indicating larger body size. Hence, extra food provisioning and early breeding reduced the occurrence of EPO in lesser kestrels. In addition, we confirmed the occurrence of intraspecific brood parasitism, as five nestlings were not the offspring of the brooding female. The results of our food-provisioning experiment support the idea that mate-feeding ability is a reliable indicator of male quality, and are in accordance with the hypothesis that male mate-feeding behaviour is a sexually selected trait.

Lesser kestrel males display inter-individual variation in melanin-based plumage traits, like ventral plumage colouration and breast/underwing spottiness. We explored whether such plumage variability was associated with single-nucleotide genetic polymorphisms of melanocortin-1-receptor (MC1R) and tyrosinase-related protein 1 (TYRP1), two genes involved in the melanogenesis pathway. No statistically significant associations between single-nucleotide mutations and male plumage traits emerged, though in some cases very rare (

Enhancement of information transfer has been proposed as a key driver of the evolution of coloniality. Transfer of information on location of food resources implies that individuals from the same colony share foraging areas and that each colony can be associated to a specific foraging area. In colonial breeding vertebrates, colony-specific foraging areas are often spatially segregated, mitigating intercolony intraspecific competition. By means of simultaneous GPS tracking of lesser kestrels (Falco naumanni) from neighbouring colonies, we showed a clear segregation of space use between individuals from different colonies. Foraging birds from different neighbouring colonies had home ranges that were significantly more segregated in space than expected by chance. This was the case both between large and between small neighbouring colonies. To our knowledge, the lesser kestrel is the only terrestrial species where evidence of spatial segregation of home ranges between conspecifics from neighbouring colonies has been demonstrated. The observed spatial segregation pattern is consistent with the occurrence of public information transfer about foraging areas and with the avoidance of overexploited areas located between neighbouring colonies. Our findings support the idea that spatial segregation of exploited areas may be widespread among colonial avian taxa, irrespective of colony size.

As a consequence of sexual selection, males and females may exhibit wide behavioural differences, for example, spatial behaviour. In fish, the two sexes often show different exploratory tendencies. This sex difference has been usually studied by testing individual fish. As many fish species live in social groups with different sex composition, the aforementioned approach might not picture the natural variation of the exploratory behaviour expressed by males and females. Here, we observed shoals of four Mediterranean killifish, Aphanius fasciatus, with three different sex compositions (4 females, 4 males, or 2 females and 2 males) during the exploration of a novel environment. Sex composition of the shoals did not predict the latency to emerge from a shelter into the novel environment. However, once emerged, shoals composed by four males displayed reduced exploratory behaviour compared to 4-female and mixed-sex shoals. These results indicated that sex differences in exploration subsist also at group level and highlighted the importance of sex composition in determining the behaviour of the entire shoal.

Many fish species are social, and individuals spend most of their lives in shoals, but sociability can vary greatly between species, populations and individuals. Sociability has been largely studied by measuring the time spent by a focal fish in proximity to one or more conspecifics. To control for the behaviour of the stimulus fish, the conspecifics have often been substituted by a mirror, on the assumption that the subject perceives its mirror image as a conspecific. The validity of the mirror test has recently been questioned, at both the behavioural and the molecular level, because of the discrepancy in fish responses when exposed to a mirror image and to a live conspecific. In this study, we compared the sociability scores of a social fish, the guppy, Poecilia reticulata, obtained using live fish or a mirror as a stimulus, to assess the validity of the mirror test. We found that the sociability score assessed using the standard mirror test was not significantly correlated with the sociability assessed using live stimuli. Nevertheless, we observed a positive correlation between the scores of the two tests when the mirror test was performed in a more naturalistic context, controlling for the minimum distance between the stimulus fish and the mirror. Our findings provide evidence of the reliability of the mirror test as a measure of sociability compared to the test using live conspecifics when certain requirements are satisfied.

Avian eggshell color seems to fulfill multiple functions, some of them being structural and others signaling. In this study, we tested whether or not eggshell coloration may play a role in sexual selection of Tree Sparrows (Passer montanus). According to the “Sexually selected eggshell coloration” hypothesis, eggshell coloration signals female, egg or chick quality and males adjust parental investment according to this signal. Eggs of this species are covered with brown spots and patches, and variation between clutches is high. We found that eggshell coloration correlates with both protoporphyrin and biliverdin, but protoporphyrin concentrations are ten times higher. Eggshell coloration reflects egg and offspring quality, but not female quality. Thus, eggshell coloration may signal female postmating investment in offspring rather than female quality. Furthermore, differential allocation in terms of maternal investment is supported by the fact that females lay more pigmented clutches when mated to males with bigger melanin‐based ornaments relative to their own. Moreover, males invested proportionally more to chicks that hatched from more pigmented clutches. Our correlative results thus seem to support a role of sexual selection in the evolution of eggshell coloration in birds laying brown eggs, pigmented mainly by protoporphyrin.

Aim: Migratory animals regularly move between often distant breeding and non‐breeding ranges. Knowledge about how these ranges are linked by movements of individuals from different populations is crucial for unravelling temporal variability in population spatial structuring and for identifying environmental drivers of population dynamics acting at different spatio‐temporal scales. We performed a large‐scale individual‐based migration tracking study of an Afro‐Palaearctic migratory raptor, to determine the patterns of migratory connectivity of European breeding populations. Location: Europe, Africa. Methods: Migration data were recorded using different devices (geolocators, satellite transmitters, Global Positioning System dataloggers) from 87 individuals breeding in the three core European populations, located in the Iberian, Italian and Balkan peninsulas. We estimated connectivity by the Mantel correlation coefficient (rM), and computed both the degree of separation between the non‐breeding areas of individuals from the same population (i.e. the population spread) and the relative size of the non‐breeding range (i.e. the non‐breeding range spread). Results: European lesser kestrels migrated on a broad front across the Mediterranean Sea and Sahara Desert, with different populations using different routes. Iberian birds migrated to western Sahel (Senegal, Mauritania, western Mali), Balkan birds migrated chiefly to central‐eastern Sahel (Niger, Nigeria, Chad), whereas Italian ones spread from eastern Mali to Nigeria. Spatial differentiation of non‐breeding areas led to a strong migratory connectivity (rM = .58), associated with a relatively high population (637 km) and non‐breeding range (1,149 km) spread. Main conclusions: Our comprehensive analysis of the non‐breeding distribution of European lesser kestrel populations revealed a strong migratory connectivity, a rare occurrence in long‐distance avian migrants. The geographical conformation of the species’ breeding and non‐breeding ranges, together with broad‐front migration across ecological barriers, promoted the differentiation of migratory routes and non‐breeding areas. Strong connectivity could then arise because of both high population spread and broad non‐breeding range. CLH; FCC Energía/Enerstar Villena S.A.; Alcalá de Henares Municipality; EDF Énergies Nouvelles S.A.; EuroNatur; Fundación Iberdrola España, Grant/Award Number: Project Migra; MAVA Foundation; French Ministry of Ecology, Grant/Award Number: Lesser Kestrel National Action Plan; Greek Green Fund; European Commission, Grant/Award Number: LIFE11 NAT/IT/000068, LIFE11 NAT/BG/000360 and NAT/GR/001011; Córdoba Zoo; Seiit‐R‐4; Consejería de Agricultura, Medio Ambiente y Desarrollo Rural de Castilla‐La Mancha; MIUR, Grant/Award Number: PRIN 2010‐2011/20180‐TZKHC.

Living in a group offers the chance to follow the choices and the behaviours of other individuals. Following a group mate might confer fitness advantages if the group mate knows about resources such as food or shelters. Shoaling fish often follow larger group mates which, in most species, are generally older and therefore more experienced. Yet, the effect of individuals' characteristics other than size on following behaviour remains to be understood. For example, familiar fish and female shoals have been reported as more cohesive, which might be due to a differential tendency to follow in relation to familiarity and sex. Here, we investigated whether size, familiarity, sex, and the interaction between these factors affect fish following behaviour. We observed pairs of differently sized Mediterranean killifish, Aphanius fasciatus, exploring a new environment, and we recorded whether the rear fish followed the front fish when the latter changed swimming direction. In female and male pairs, and in unfamiliar pairs, smaller fish were more likely to follow the directional change of the larger fish than vice versa. In mixed-sex pairs and in familiar pairs, however, size did not affect following behaviour and larger fish followed as much as smaller fish did. Our results revealed that killifish's following decisions are determined by the size of the individuals, their level of familiarity, and their sex. These characteristics may have a notable impact on the behaviour of fish groups in nature.

The performances of different social groups can depend on various characteristics, such as familiarity among their members or the presence of individuals with specific traits. However, it has rarely been investigated how groups perform during an encounter with other conspecifics, even if in the natural environment social groups often run into each other and compete for resources. We investigated whether a certain characteristic of the group (i.e., familiarity) could benefit its members when they are confronted with another group. We designed a novel experimental set-up, creating triads of captive house sparrows (Passer domesticus) and examining whether in a situation of competition for limited resources one triad could gain benefits over the other (consume more mealworms, Tenebrio molitor). While we did not find an effect of previous familiarity among triad members on the triads’ performances, we discovered a group-based difference in the number of mealworms eaten per capita. Group-mates of the very first individual to eat a mealworm (first feeder) ate more mealworms than those in the opposing triad. First feeder individuals also foraged sooner and more than other birds in a subsequent prey consumption assay. Our results suggest that individual performances were influenced by group membership, even when groups were exploiting the same resource simultaneously.

As parental care is costly, it can be expected that there will be a sexual conflict between parents over the individual levels of parental investment because each parent has limited resources to invest in a reproductive event. Theoretical models of parental investment predict that when one parent reduces its parental effort, the other parent should adjust its effort facultatively to compensate for the decrease in the mate’s contribution. Here, we tested for facultative adjustments in care in the rock sparrow, Petronia petronia. In this species, both sexes can desert the brood, creating the potential for strong sexual conflict over parental care to occur. To that end, we examined how rock sparrow parents adjust their level of care in response to an experimental mate removal for a limited time period, mimicking the starting phase of the desertion process. We compared male and female provisioning rates before and after an experimental mate removal. Males behaved according to the prediction of compensatory adjustment, as they allocated more care to the offspring and returned faster to the nest after female removal, whereas females showed no response. Our study shows sexual differences in response to mate removal in a species with biparental care and strong sexual conflict over parental care, and suggests that males may use the female absence to determine their actual parental effort. Significance statement: Strong differences in parental care, with one sex providing more care than the other, are widespread across the animal kingdom. Theoretical models of parental investment predict that when one parent reduces its care, the mate should adjust its care facultatively to compensate. Here, we tested for facultative adjustments in care in the rock sparrow when their mates were experimentally removed for a limited time period. We show that only male behavior was consistent with theoretical predictions as they increased their frequency of provisioning significantly after mate removal, whereas females showed no response. Our study suggests that members of a pair do not make independent decisions, but negotiate to modify their effort in direct response to the prior effort of their mate., Open access funding provided by University of Turku (UTU) including Turku University Central Hospital. This study was financed by project CGL2017-83843-C2-1 to JM from Spanish Ministerio de Economía y Competitividad (MINECO). AC is supported by a postdoctoral fellowship from Fundación Ramón Areces. MP was supported by an FPI grant from MINECO. This work was done while the first author was visiting the University of Padova supported by the Program B José Castillejo (grant CAS18/00139).

A major aim of evolutionary biology is to understand why patterns of genomic variation vary among populations and species. Large-scale genomic studies of widespread species are useful for studying how the environment and demographic history shape patterns of genomic divergence, and with the continually decreasing cost of sequencing, such studies are now becoming feasible. Here, we carry out one of the most comprehensive surveys of genomic variation in a wild vertebrate to date; the great tit ( Parus major ) HapMap project. We screened ca 500,000 SNP markers across 647 individuals from 29 populations, spanning almost the entire geographic range of the European great tit subspecies. We found that genome-wide variation was consistent with a recent colonisation across Europe from a single refugium in the Balkans and/or Turkey, with bottlenecks and reduced genetic diversity in island populations. Differentiation across the genome was highly heterogeneous, with clear “islands of differentiation” even among populations which are ostensibly panmictic. These islands of differentiation were consistently found in regions of low recombination, suggesting that background selection can rapidly promote population differentiation among even the most recently colonised populations. We also detected genomic outlier regions that were unique to peripheral great tit populations, most likely as a result of recent directional selection at the range edges of this species. These “unique” outlier regions contained candidate genes for morphology, thermal adaptation and colouration, supporting previous research in this species, and providing avenues for future investigation. Our study suggests that comprehensive screens of genomic variation in wild organisms can provide unique insights into evolution. Author summary Studying patterns of genetic variation is a useful way of determining why populations and species differ in nature. Genetic variation is shaped by natural selection, but also by the present and past size of populations, the amount of migration, and by features of the genome, such as variation in recombination rate, of the organism being studied. Teasing apart the effects of these different processes on genomic diversity is difficult, but one way that this can be achieved is by studying genomic variation across the entire range of a species. We performed a continental-scale analysis of genetic variation in the great tit - a widespread songbird that has been the focus of extensive ecological research. We first used genomic data to reconstruct the historical colonisation of great tits across Europe, and showed that during the last ice age, this species was likely restricted to a single region in Eastern Europe, from which they spread across the continent. We then studied how patterns of variation differ along the genome, and show that recombination rate is a key driver of variation among all populations. Importantly, by comparing many populations we were able to identify genes that have been subject to natural selection in specific geographical regions. We found that natural selection appeared to be strongest in populations on the edges of the great tit’s range acting on traits such as morphology, stress response and colouration. Large-scale genetic analyses such as ours are therefore useful approaches for understanding how evolution operates in the wild.

In the last decades, higher attention has been paid to olfactory perception in birds. As a consequence, a handful of avian species have been discovered to use olfaction in different contexts. Nevertheless, we still have a very limited knowledge about the use of odor cues in avian social life, particularly, in the case of songbirds. Here, we investigate if female house sparrows Passer domesticus show any preference for the odor of kin and nonkin conspecifics and we also test a possible role of familiarity based on male scent in female choice. We performed the experiment with captive birds twice, during the nonbreeding and breeding seasons. Our results show that female house sparrows strongly avoided the odor of unrelated familiar (UF) males, both in the breeding and nonbreeding seasons. Our results suggest recognition for olfactory stimuli related to familiarity and kinship. We suggest that avoidance for UF males is associated with previous experience in this species. Also, we provided further evidence to the use of olfaction in passerine species by using a new experimental setup.

Information about predation risk is of fundamental value in biological communities. Because many prey species have shared predators, eavesdropping on other species' alarms is a widely recognized mechanism underlying the formation of mixed-species groups. However, information transfer may vary both across and within groups because some species provide higher-quality information about predators than others. We tested this phenomenon in Amazonian understory mixed-species flocks of birds in which two sentinel species-the bluish-slate antshrike (Thamnomanes schistogynus) and the dusky-throated antshrike (Thamnomanes ardesiacus)-occupy different habitats and provide alarm calls that are used by eavesdropping flock mates. In a playback experiment, two associate species responded significantly more strongly to alarm calls from the same sentinel species, reflecting the greater reliability of information about predator threats that could affect survival and habitat choice. Our work provides evidence of a repeated asymmetry across space in the available information about threats.

Group-living animals can develop social bonds. Social bonds can be considered a type of social relationship characterized by frequent and consistent affiliative (non-reproductive) interactions. Social bonds with conspecifics bring many advantages, also in terms of direct fitness. A characteristic of social bonds is that they need time to develop. Several studies on humans have emphasized the fact that sharing experiences can affect the strength of social bonds. A similar trend can be spotted in non-human species. For example, a recent experiment showed that if chimpanzees watched a video together with a conspecific, they spent more time in proximity compared to conspecifics with whom they did not actively watch a video. Another experiment on fish showed that individuals who experienced a situation of high predation risk together, showed preference for each other compared to those who did not. As the link between shared experiences and social bonds is not explicitly recognized in non-human animals, the main goal of this work is to propose the exploration of this novel research path. This exploration would contribute to shed light on the evolutionary mechanisms of social bond (or friendship) development and maintenance between individuals in different vertebrate species, from fish to non-human primates.

According to theory, in species in which male variance in reproductive success exceeds that of the females, sons are more costly to produce; females mated with high quality males or those in better condition should produce more sons. In monogamous species, however, the variance in the reproductive success of the two sexes is often similar and mate choice is often mutual, making predictions regarding sex allocation more difficult. In the rock sparrow Petronia petronia, both males and females have a sexually selected yellow patch on the breast, whose size correlates with individual body condition. We investigated whether the brood sex ratio co‐varies with the size of the yellow patch of the father and the mother in a sample of 173 broods (818 chicks) over 8 breeding seasons. While the size of the yellow patch of the mother and the father did not predict per se a deviation from the expected 1:1 sex ratio, brood sex ratios were predicted by the interaction of male and female yellow patch size. This result is surprising, as the ornament is sexually selected by both males and females as an indicator of quality in both sexes and should therefore be inherited by all offspring irrespective of their sex. It indirectly suggests that other sex‐specific traits associated with patch size (e.g. polygyny in males and fecundity in females) may explain the sex allocation bias observed in rock sparrows. Thus, female individual quality alone, as expressed through the size of the yellow patch, was not associated with the biases in sex ratios reported in this study. Our results rather suggest that sex allocation occurs in response to male attractiveness in interaction with female attractiveness. In other words, females tend to preferentially allocate towards the sex of the parent with more developed ornament within the pair., AC was funded by a Juan de la Cierva‐formación postdoctoral grant (FJCI‐2015‐23536; Ministerio de Economía, Industria y Competitividad, MINECO, Spanish Government).

The quality of a breeding site may have major fitness consequences. A fundamental step to understanding the process of nest-site selection is the identification of the information individuals use to choose high-quality nest sites. For secondary cavity-nesting bird species that do not add nest lining material, organic remains (faeces, pellets) accumulated inside nest cavities during previous breeding events may be a cue for high-quality nest-sites, as they contain information about past successful breeding and may improve thermal insulation of eggs during incubation. However, cavities in which breeding was successful might also contain more nest-dwelling ectoparasites than unoccupied cavities, offering an incentive for prospective parents to avoid them. We exposed breeding cavity-nesting lesser kestrels (Falco naumanni) to nestbox dyads consisting of a dirty (with a thick layer of organic substrate) and a clean nestbox (without organic material). Dirty nestboxes were strongly preferred, being occupied earlier and more frequently than clean ones. Hatching success in dirty nestboxes was significantly higher than in clean ones, suggesting a positive effect of organic nest material on incubation efficiency, while nestbox dirtiness did not significantly affect clutch and brood size. Nestlings from dirty nestboxes had significantly higher ectoparasite load than those from clean nestboxes soon after egg hatching, but this difference was not evident a few days later. Nest substrate did not significantly affect nestling growth. We concluded that nest substrate is a key driver of nest-site choice in lesser kestrels, although the adaptive value of such a strong preference appears elusive and may be context-dependent.

Extended phenotypes as signals are widely distributed among animal taxa. For example, many bird species build eye-catching nests or structures, which can potentially mirror the quality or ability of the builder. Rock sparrow, Petronia petronia, nests are usually overly decorated with feathers belonging to different species. Feather carrying in this and other species seems to play a role beyond their supposed thermoregulatory function, that is, to provide insulation to eggs and developing chicks. In this study, we documented for the first time this intriguing pattern of behaviour in the rock sparrow and experimentally tested its potential role as a sexually selected or status signal by means of a feather supplementation experiment carried out in two distinct populations from Italy and Spain. We found that females were responsible for feather carrying, laid larger clutches and provisioned their young at a lower rate in those nests with experimentally added feathers. Decorated nests sustained fewer intrusions by floater individuals and were defended with greater intensity by both parents than control nests, which supports the role of nest ornamentation as a status signal to conspecifics. Presence of experimental feathers did not significantly increase the frequency with which males provisioned their young but males tended to desert their brood less often and spent more time guarding the brood in experimental nests, indicating that feather presence may also play a role in an intersexual context. Overall, our results allow us to exclude the thermoregulation hypothesis as a likely explanation for the occurrence of these decorations and provide partial evidence for the idea that feather carrying conveys information to the partner and potential competitors. Our study thus supports the notion that nonbodily traits serving a direct (naturally selected) function can also evolve a signalling component.

Background Exploratory behaviour is one of the best-investigated behavioural traits. However, little is known about how differences in familiarity, i.e. in the knowledge and previous experience with a companion can influence the exploration of a novel environment. However, to our knowledge, such a critical feature of the social environment has never been the target of a study relating it to exploratory behaviour in birds. Here we examined if familiarity with a conspecific could affect behavioural responses of individuals confronted with a novel environment. We recorded the latency to land on the ground, latency to feed, time spent feeding and number of sectors visited of 48 female and 48 male house sparrows (Passer domesticus) in an indoor aviary in three contexts: alone (individual context), with an unfamiliar and with a familiar same-sex companion. Results House sparrows landed sooner on the ground when in the familiar context than when in the individual context. Birds in unfamiliar pairs followed each other less than familiar birds, but this difference diminished with time spent exploring. Moreover, males and females differed in their behavioural responses in the unfamiliar context. Females with a familiar companion landed sooner than when they were paired with an unfamiliar conspecific, whereas only the presence of a companion but not familiarity reduced males latency to land on the ground. Finally, when considering the unfamiliar context males had shorter latencies to forage and thus spent more time eating than females. Conclusions The presence or absence of a companion and its familiarity with the focal individual influenced differently the behavioural responses of male and female house sparrows in a novel environment. As house sparrows are strongly sociable, the influence of the social environment is likely to be of paramount importance to understand the selective pressures acting on them, particularly in recently colonized areas with ephemeral food sources. Our results shed light on the complex influence that the social environment has on the behavioural responses of a cosmopolitan bird. Electronic supplementary material The online version of this article (10.1186/s12983-018-0267-8) contains supplementary material, which is available to authorized users.

While the function of ornaments shaped by sexual selection is to attract mates or drive off rivals, these signals may also evolve through social selection, in which the social context affects the fitness of signallers and receivers. Classical ‘mate choice’ experiments often reveal preferences for ornaments, but few studies have considered whether these are strictly sexual or reflect general social preferences. Indeed, an alternative possibility is that ornaments evolve through ‘non-sexual social selection’ (hereafter ‘social selection’). We examined the role of ornamentation (yellow ventral patch) and familiarity (individuals recognize group mates with which they have had previous interactions) on mate choice (opposite-sex stimuli preference) and social choice (same-sex stimuli preference) in both male and female white-eyed bulbuls (Pycnonotus xanthopygos). In the mate choice test, females preferred unfamiliar males with increased yellow. There were no biologically important differences in male preferences based on familiarity or intensity of patch colour. In the social choice test, females preferred to associate with familiar females. Males preferred to associate with familiar males but also preferred to associate with less ornamented males. Our results suggest that ornamentation and familiarity are important features, playing different roles in males and females, in both social and sexual selection processes.

Social foraging is thought to provide the possibility of information transmission between individuals, but this advantage has been proved only in a handful of species and contexts. We investigated how social connections in captive flocks of house sparrows (Passer domesticus) affected the discovery of (i.e. feeding for the first time from) two hidden food patches in the presence of informed flock-mates. At the first-discovered and most-exploited food patch social connections between birds affected the order of discovery and presumably contributed to a greater exploitation of this patch. However, social connections did not affect discovery at the second food patch despite its close spatial proximity. Males discovered the food sources sooner than females, while feeding activity was negatively related to patch discovery. Age had no effect on the order of discovery. Birds that first discovered and fed at the food patches were characterized by higher level of social indifference, i.e. followed others less frequently than other birds in an independent context. Our findings provide experimental evidence for the importance of variable social connections during social foraging in house sparrow flocks, and suggest that social attraction can contribute differently to the exploitation of different patches when multiple food sources are present.

In many social species, when an individual is associated with familiar conspecifics, it displays an array of behaviours that may confer benefits (e.g., increased boldness and faster habituation to novel environments). In fish, these effects of familiarity have been studied using individuals of only one sex or juveniles. Since shoals often vary regarding sex composition and males and females show different social behaviours, we hypothesised that social familiarity's effects vary with group sex composition. We tested this hypothesis by comparing the exploratory behaviour of groups of two females, two males, and one male and one female Mediterranean killifish, Aphanius fasciatus, which were either familiar or unfamiliar. Pairs of familiar females were bolder than pairs of unfamiliar females, whereas males showed the opposite trend. Pairs of familiar females also showed faster habituation to the novel environment and, at the beginning of the test, were more cohesive compared to pairs of unfamiliar females. Pairs of familiar mixed-sex fish habituated faster to the novel environment than unfamiliar pairs. Pairs of familiar males did not show any beneficial effect of familiarity relative to pairs of unfamiliar males. Hence, the effects of social familiarity on exploratory behaviour, and likely the associated benefits, appear to depend on the sex composition of the pair in the Mediterranean killifish.

The White-spectacled Bulbul, Pycnonotus xanthopygos, is an abundant and possibly invasive species in Turkey, where it has gradually expanded its distribution and breeding range in both western and southeastern directions. This study focused on its breeding biology, which is still poorly known. The breeding activity extends from February until September. The preferred nesting areas are mainly gardens and maquis groves, where 24 different nesting tree species have been identified. The clutch size is 3.3 ± 0.8 eggs per pair, nesting success 68%, hatching success 94%, fledgling success 95%, and overall breeding success 89%. While nesting success differs significantly between the years, we found no significant differences in hatching, fledging, and overall breeding success between the years studied. Despite favourable climatic conditions in the Mediterranean region, the species makes only one brood per year in a relatively extended breeding season extending over seven months, and has a relatively a hig...

Increasing intensification in vineyards has detrimental effects on biodiversity. Although several studies addressed this topic, the reproductive outcomes of model organisms in vineyards received little attention. We carrie d out the first study on birds nesting in natural nests on vines, focusing on nest density, breeding performance and nest-site selection in organic and conventional systems and in two contrasting trellising systems, pergola (taller, with more spaced rows and a denser canopy) vs. spalliera . We surveyed 228 nests of six species and analysed nest densities and final fates as a function of vineyard management and trellising system by means of GLM(M)s. 51% of nests were abandoned before egg-laying and the probability of early abandonment was positively related to the amount of access farmers had for management activities. The number of nests was four times higher in pergola than in spalliera vineyards, likely due to pergola 's complex and tree-like structure. Organic or conventional management did not affect nest density, probably due to reduced differences in terms of management practices between them. Breeding success was low and marginally affected by the interaction of the management and trellising systems, being higher in conventional pergola . Nests were preferentially built on larger vines and were placed at an increasing height on the vine as the season progressed. Nests built at greater heights had greater success. We provide some management recommendations for bird conservation in vineyards: promoting pergola and maintaining older vineyards, limiting grass mowing in April and May, and in compatibility with agricultural management, optimising the amount of farmers' access to vineyards.

Experimental evidence suggests that reproductive interference between heterospecifics can seriously affect individual fitness; support from field studies for such an effect has, however, remained scarce. We studied reproductive interference in 25 natural breeding ponds in an area where two ranid frogs, Rana dalmatina and Rana temporaria, co-occur. The breeding seasons of the two species usually overlap and males of both species are often found in amplexus with heterospecific females, even though matings between heterospecifics produce no viable offspring. We estimated species abundance ratios based on the number of clutches laid and evaluated fertilization success. In ponds with low spatial complexity and a species abundance ratio biased towards R. temporaria, the average fertilization success of R. dalmatina eggs decreased, while this relationship was not detectable in spatially more complex ponds. Fertilization success of R. temporaria did not decrease with increasing relative numbers of heterospecifics. This asymmetry in fitness effects of reproductive interference may be attributed to R. temporaria males being more competitive in scramble competition for females than R. dalmatina males. Our study is among the first to demonstrate that in natural breeding populations of vertebrates interference among heterospecifics has the potential to substantially lower reproductive success at the population level, which may in turn affect population dynamics.

To explain the evolution of egg colouration in open cup nesting species, a number of functions have been suggested. Recent studies focus on the role of eggshell colour as a postmating sexually selected trait of females which manipulates male parental investment. A basic prediction of this hypothesis is that egg pigmentation reflects female quality. In this study we examine whether there is a relationship between eggshell colouration and either female quality or egg quality in reed warblers. This open cup nesting species has eggs that are heavily spotted with brownish marks on a bluish-green background. We used several parameters describing female and egg quality, and measured eggshell colouration at the blunt pole and the egg centre, deriving four colour variables from colour spectrometry. To determine egg quality parameters, the third egg of each clutch was sampled and analysed. To determine female quality, females were trapped shortly after egg laying, and several morphological and a single conditional ...

Invasive alien predators (IAP) are spreading on a global scale-often with devastating ecological effects. One reason for their success may be that prey species fail to recognize them due to a lack of co-evolutionary history. We performed a comprehensive test of this "prey naivete" hypothesis using a novel approach: we tested whether predator-naive tadpoles of the agile frog (Rana dalmatina) display antipredator behavior upon encountering chemical cues produced by native, invasive (established or recent) or allopatric fishes (four perciforms, four siluriforms, and two cypriniforms). We studied the influence of population origin on predator-detection ability by presenting chemical cues to predator-naive tadpoles that originated from fishless hill-ponds or fish-infested floodplain populations. Before trials, we fed fishes with tadpoles or an alternative food to test whether direct chemical cues from the predator's diet influences the tadpoles' recognition of potential predators. Tadpoles reduced their activity upon exposure to cues from native and long-established invasive perciforms, but not in response to recent invaders, allopatric predators, or to any siluriforms. Also, predators that were previously fed with tadpoles did not universally induce behavioral defensedefenses upon first encounter. Finally, tadpoles originating from isolated hill-ponds exhibited higher baseline activity and responded in weaker fashion than their conspecifics from floodplain populations, which co-exist with predatory fishes. Our results indicate that tadpoles may be vulnerable to invading predatory fishes due to their inability to recognize them as dangerous, though their ability to recognize invasive IAP may evolve rapidly, in fewer than 30 generations.

While it is well established that females prefer to mate with well‐ornamented males, the influence of perceptive and cognitive processes on the expression of female mate choice is still poorly known. It has been suggested that the female perception of a male's attractiveness is not absolute, but depends on the other males with which he is compared that have been previously encountered (comparative evaluation). We investigated whether mate preference in bearded reedlings (Panurus biarmicus) is dependent on or independent of social context in relation to two different traits: beard and tail lengths. Each female had a choice between two to three males with different modifications of beard and tail. For each female, three different experiments were conducted (one binary and two trinary tests). We found that when females are presented with options that vary antagonistically with respect to two ornaments (binary test), some individuals prefer one trait while others the other trait. This indicates that in our bearded reedlings population exists a mate preference polymorphisms. Moreover, we found that the presence of a third stimulus, irrespective of the initial preference, reduced the strength of the initial preference – what we can call a “preference dilution effect.” Our results suggest that the female's choice may be constrained by her cognitive abilities when she is simultaneously presented with several options varying for two uncorrelated traits.

Several studies have investigated how individuals’ social status or behavioural and physiological traits affect social organisation in group-living species, but the potential role of ornamental traits has been rarely studied. Here, we analysed initiator–follower interactions in relation to experimentally manipulated sexually selected ornaments in captive flocks of male bearded reedlings (Panurus biarmicus). We manipulated beard length or tail length on the most and least frequently followed individuals in unisex flocks and examined how trait manipulation affected social prestige (i.e. frequency of being followed) and centrality (i.e. frequency of following others) of these individuals. We found that prestige/centrality after manipulation was significantly positively associated with their before-manipulation measure, whereas body condition did not affect social position within flocks. Similarly, the manipulation of the ornamental traits had no significant effect on the social positions of the manipulated i...

Colour variation in birds is often used to signal functional differences between individuals and sexes, though white plumage has usually been disregarded because white feathers were thought to be cheap to produce and hence unreliable signals. Here, we provide evidence for sex-specific morphological and colour differences in the strikingly patterned but apparently monomorphic Dia- mond Firetail (Stagonopleura guttata). We found that males had longer and darker tails, wider lores and a darker bill, while females had significantly more white flank spots. These gender-specific trait differences could be used to signal individual quality in sexual or social interactions, and hence we examined the correlation between these traits and the cell-mediated immune response (phytohaemag- glutinin, PHA test). During both the breeding and non- breeding seasons, we found a positive correlation between PHA response and white flank spot number as well as their ultra-violet reflectance in females, and a negative correla- tion with white chroma and hue. In males, there was a positive correlation between PHA response and red rump reflectance (hue, red-chroma). There was no association between PHA response and either tail length or lore depth in either sex. These results add to a growing body of evidence that female spot number and white plumage reflectance signal quality.

In seasonally fluctuating environments, timing of reproduction is a crucial determinant of fitness. Studies of birds show that late breeding attempts generally result in offspring of lower reproductive value, with lower recruitment and long-term survival prospects. Several proximate mechanisms, including a seasonal decline of immune system functioning, may lead to a seasonal decline of offspring fitness. We investigated seasonal variation in offspring quality by subjecting first- and second-brood chicks of a sexually size dimorphic species, the European starling Sturnus vulgaris, to an immune challenge with a bacterial endotoxin (LPS), and evaluated their growth and physiological response in terms of total plasma antioxidant capacity (TAC), concentration of reactive oxygen metabolites and hematocrit. LPS challenge did not affect chick growth or oxidative status. However, hematocrit of second-brood chicks was higher in LPS chicks compared to controls. Body mass halfway through the rearing period (days 8–9 post-hatching), TAC and hematocrit were lower among second- vs. first-brood chicks. Interestingly, sexual dimorphism in body mass at days 8–9 post-hatching markedly differed between broods, first-brood males being 4.7% and second-brood males 22.7% heavier than their sisters, respectively. Pre-fledging mortality occurred among second-brood chicks only and was strongly female-biased. Our findings suggest that starling chicks, even if in poor conditions, are little affected by a bacterial challenge, at least in the short-term. Moreover, our study indicates that sex differences in body size, possibly mediated by sex-specific maternal investment in egg size, may heavily impact on pre-fledging survival in a different way in the course of the breeding season, resulting in sex-specific seasonal decline of offspring fitness. Finally, we suggest that levels of circulating antioxidants should be regarded among the proximate causes of the association between timing of fledging and long-term survival in avian species.

Plumage colour can be used as an honest signal to convey health and status, which has traditionally been examined in the sexual selection context of choosy females and elaborate males. We use a model avian system to study the role of plumage coloration in a social context such as inter- and intrasexual competition over food resources. The diamond firetail (Stagonopleura guttata) is an endemic Australian finch: females have more white flank spots than males, and white spot number was correlated with cell-mediated immune response in females. We use two experimental designs to test the role of white flank spots for feeding dominance and dominance discrimination in a group-living bird. The results from two-choice trials and from single-arena trials showed that female ornamentation was consistently important in social food contests, and males consistently responded to female spot number. Females with higher spot number fed first, in trials with males and ⁄or females. Also, females preferred to feed next to test birds with low spot number, but males showed no preference for feeding next to birds with few or many spots. Finally, latency to feed was predicted by spot number: both males and females had longer latency to feed if test birds had more spots than the focal birds. We conclude that female, but not male, ornamentation was important for inter- and intrasexual food competition. This is one of the very few studies to show that the same plumage ornament can have a different function between the sexes as a signal of social status. Moreover, this study shows that white plumage can be a signal of dominance.