Environmental conditions experienced upon first breeding modulate costs of early breeding but not age-specific reproductive output in peregrine falcons.

Although once considered uncommon, there is growing evidence of widespread senescence in wildlife populations. However, few studies have examined the traits involved, inter-sexual differences, and environmental correlates of age-specific performance in raptors. We studied age-specific reproductive performance and actuarial senescence (decrease in survival probability with age) in a peregrine falcon population monitored for 21 years. We analysed changes with age in the number of offspring produced and incubation start date. We also inspected variation in lifespan and breeding lifespan (number of breeding occasions in a lifetime). In every case, we assessed associations between variations in traits and age, sex, recruitment age, and environmental conditions (cumulative rainfall during breeding season) experienced upon the first breeding attempt. We found scarce evidence for reproductive senescence. Only the incubation start date in females, which was delayed after approximately 8 cy (calendar years), suggested reproductive senescence in our study population. Regarding actuarial senescence, our data did not support it as we only found evidence of higher juvenile mortality. Furthermore, expected lifespan in peregrines recruited at 2 cy was associated with conditions experienced upon the first breeding attempt. The lifespan and breeding career of individuals recruited as yearlings and experiencing low rainfall upon first breeding did not significantly differ from those recruited as adults. However, those recruited as yearlings and experiencing poor environmental conditions upon the first breeding attempt showed reduced lifespan and breeding lifespan.

Effect of nest composition, experience and nest quality on nest-building behaviour in the Bonelli's Eagle.

In bi-parentally built nests, there is evidence to suggest that nests are extended phenotypic signals that accurately indicate the quality of the building parent/s. Raptors often use a variety of materials to build their nests (natural, such as branches, but also non-natural objects), presumably due to their insulating properties, their suitability to advertise occupancy of the nest, and to decrease pathogen and parasite loads. However, in raptors where both sexes collaborate in nest construction, it is unclear whether nest building (taking the amount of material carried to the nest as the potential predictor) is an indicator of parental quality, and whether the effort expended by both sexes could constitute an honest signal of parental quality to their partners. Between 2011 and 2016, we monitored 16 nests of Bonelli's Eagles (Aquila fasciata), and we examined data on sex, type of material brought to the nest, breeding experience, nest quality, timing, and nest-building investment prior to egg-laying from 32 identifiable Bonelli's Eagles during the pre-laying period to investigate the relative contribution of the sexes to the amount of nest material gathered. Our results indicate that sex is not a determining factor in nest-building effort, and that females did not increase their parental effort in response to the male's contribution, and supply of materials did not increase during the pre-laying period. In contrast, our models showed that: (1) the type of material supplied to the nest by both sexes varied significantly throughout the pre-laying period and (2) nest-building effort was determined by individual experience and nest quality. Therefore, our study suggests that male nest-building behaviour and investment by Bonelli's Eagles cannot be considered as an extended phenotypic signal. The differential use of hard and green material by both sexes in the early and late stages of nest-building period, and the fact that the more experienced individuals contributed a larger amount of material on low quality nests, are discussed in the contexts of signaling nest occupancy to conspecifics and competitors and the decrease of ectoparasite loads during the pre-laying period.

Different increase rate in body mass of two marten species due to climate warming potentially reinforces interspecific competition.

Many species show spatial variation in body size, often associated with climatic patterns. Studying species with contrasting geographical patterns related to climate might help elucidate the role of different drivers. We analysed changes in the body mass of two sympatric medium-sized carnivores-pine marten (Martes martes) and stone marten (Martes foina)-across Europe over 59 years. The body mass of pine marten increased with decreasing latitude, whereas stone marten body mass varied in a more complex pattern across its geographic range. Over time, the average body mass of pine martens increased by 255 g (24%), while stone marten by 86 g (6%). The greatest increase of body mass along both martens' geographic range was observed in central and southern Europe, where both species occur in sympatry. The body mass increase slowed down over time, especially in allopatric regions. The average pine/stone marten body mass ratio increased from 0.87 in 1960 to 0.99 in 2019, potentially strengthening the competition between them. Thus, a differential response in body size to several drivers over time might have led to an adaptive advantage for pine martens. This highlights the importance of considering different responses among interacting species when studying animal adaptation to climate change.

Colonization of Urban Habitats: Tawny Owl Abundance Is Conditioned by Urbanization Structure.

Natural habitats are being altered and destroyed worldwide due to urbanization, leading to a decrease in species abundance and richness. Nevertheless, some species, including tawny owls, have successfully colonized this novel habitat. Consequences at the population level have not been described; thus, our main objective was to describe the effects that urban structure have on the tawny owl population at local and landscape levels. Data were obtained from 527 survey points over 7 months in a large-scale owl survey in the Basque Country (northern Spain) in 2018. At the local scale, the interaction between forest and urban cover affected tawny owl abundance, the optimum being in medium forested areas. The interaction between urban cover and clumpiness index (urban patch distribution) showed a generally negative effect. At the landscape scale, its abundance decreased in complex-shaped urban patches and when distance between them was greater. In conclusion, at the local scale, when a minimal forest structure is present in urbanized areas, the species can exploit it. At the landscape scale, it prefers smaller urban towns to cities. Thinking ahead, the current tendency toward "green capitals" should benefit tawny owl populations.

Phenotypic and environmental correlates of natal dispersal in a long-lived territorial vulture.

Natal dispersal, the movement between the birth and the first breeding site, has been rarely studied in long-lived territorial birds with a long-lasting pre-breeding stage. Here we benefited from the long-term monitoring programs of six populations of Egyptian vultures (Neophron percnopterus) from Spain and France to study how the rearing environment determines dispersal. For 124 vultures, we recorded a median dispersal distance of 48 km (range 0-656 km). Linear models were used to assess the effect of population and individual traits on dispersal distance at two spatial scales. Dispersal distances were inversely related to vulture density in the natal population, suggesting that birds perceive the abundance of conspecifics as a signal of habitat quality. This was particularly true for declining populations, so increasing levels of opportunistic philopatry seemed to arise in high density contexts as a consequence of vacancies created by human-induced adult mortality. Females dispersed further than males, but males were more sensitive to the social environment, indicating different dispersal tactics. Both sexes were affected by different individual attributes simultaneously and interactively with this social context. These results highlight that complex phenotype-by-environment interactions should be considered for advancing our understanding of dispersal dynamics in long-lived organisms.

Winter movement patterns of a globally endangered avian scavenger in south-western Europe.

Partial migration, whereby some individuals migrate and some do not, is relatively common and widespread among animals. Switching between migration tactics (from migratory to resident or vice versa) occurs at individual and population levels. Here, we describe for the first time the movement ecology of the largest wintering population of Egyptian Vultures (Neophron percnopterus) in south-west Europe. We combined field surveys and GPS tracking data from December to February during four wintering seasons (2014-2018). The wintering population consisted on average of 85 individuals (range 58-121; 76% adults and 24% subadults). Individuals were counted at five different roosting sites located near farms, unauthorized carcass deposition sites and authorized carcass deposition sites. Our results show that vultures tend to remain close to the roosting site. Moreover, we observed that females exhibited smaller home range sizes than males, which suggests a possible differential use of food sources. Overall, birds relied more on farms than other available food resources, particularly subadult individuals which exploited more intensively these sites. Our results showed that Egyptian Vultures congregate in significant numbers at specific sites throughout the winter period in south-west Spain and that these roosting and feeding sites should be given some level of legal protection and regular monitoring. Furthermore, predictable food sources might be driving the apparent increase in the non-migratory population of Egyptian Vultures, as observed in other avian species which are also changing their migratory behavior.

Copulatory behaviour in the Bonelli´s Eagle: Assessing the paternity assurance hypothesis.

We examined copulatory behaviour in the Bonelli´s Eagle (Aquila fasciata) at nesting sites in the eastern zone of the Baetic Cordillera, southern Spain, between 2010 and 2012. We observed the copulatory behaviour of 15 pairs during the pre-laying period. Bonelli´s Eagles commenced sexual activity ca. 69 days before egg-laying. Ninety-six percent of mounting attempts were successful. Bonelli´s Eagle pairs averaged 99.8 copulation attempts per clutch, with an average copulation frequency of 0.86 copulation attempts per day. Pairs displayed a daily bimodal pattern of copulation activity, with copulations occurring most frequently in the evening. We used our data to test three predictions with regard to the paternity assurance hypothesis. Prediction 1, that within-pair copulations increase with local breeding density, was rejected because our models showed no evidence for it. Prediction 2, that within-pair copulations increase during the female fertile period, was marginally supported. Finally, Prediction 3, that mate attendance increases during the female fertile period, was also rejected because mate-guarding did not increase as the fertile period approached. However, mate-guarding was positively correlated with within-pair copulation frequency. Moderate copulation rates compared to other raptors and the absence of mate-guarding suggest that, in the study area, Bonelli´s Eagles exhibit only partially adaptive behaviour to assure their paternity. A possible explanation could be related to the low number of extra-pair encounters observed (opportunities for which appear to be rare), although the gradual increase in within-pair copulations during the female fertile period is consistent with the sperm competition hypothesis. The results are discussed based on the signalling hypothesis, which proposes that raptors signal territory ownership to conspecifics, and possibly to other raptor species, by copulating frequently and conspicuously in the defended nesting area.

Ungulate vehicle collisions in a peri-urban environment: consequences of transportation infrastructures planned assuming the absence of ungulates.

Ungulate vehicle collisions (UVC) provoke serious damage, including human casualties, and a large number of measures have been developed around the world to avoid collisions. We analyse the main factors involved in UVC in a road network built in the absence of ungulates, where mitigation structures to avoid UVC were not adequately considered. Ungulate population greatly increased during the last two decades and now Roe Deer and Wild Boars are widely distributed over the study area, but even after this increase, the road network was not adapted to avoid UVC. A total of 235 Roe Deer (RDVC) and 153 Wild Boar vehicle collisions (WBVC) were recorded between January 2008 and December 2011. We randomly selected 289 sample points (87 RDVC, 60 WBVC and 142 controls) separated by at least 500 metres from the next closest point and measured 19 variables that could potentially influence the vehicle collisions. We detected variations in the frequency of RDVC on a monthly basis, and WBVC was higher at weekends but no significant differences were detected on a monthly basis. UVC were more likely to occur at locations where sinuosity of the road, velocity, surface of shrub and deciduous forest area were greater, the presence of fences entered with positive relationship and distance to the nearest building was less. RDVC were more likely to occur at locations where timber forest area increased and distance to the nearest building decreased and WBVC was related to open fields cover and also to the presence of fences. Sinuosity and velocity entered in both cases as significant factors. Major roads, in which the traffic volume is greater and faster, caused more accidents with ungulates than secondary roads. Nowadays, the high frequency of ungulate road-kills deserves a new strategy in order to adapt infrastructure and adopt mitigation measures.

Large-scale patterns of genetic variation in a female-biased dispersing passerine: the importance of sex-based analyses.

Dispersal affects the distribution, dynamics and genetic structure of natural populations, and can be significantly different between sexes. However, literature records dealing with the dispersal of migratory birds are scarce, as migratory behaviour can notably complicate the study of dispersal. We used the barn swallow Hirundo rustica as model taxon to investigate patterns of genetic variability in males and in females of a migratory species showing sex-biased dispersal. We collected blood samples (n = 186) over the period 2006 to 2011 from adults (H. r. rustica subspecies) nesting in the same breeding site at either high (Ireland, Germany and Russia) or low (Spain, Italy and Cyprus) latitude across Europe. We amplified the Chromo Helicase DNA gene in all birds in order to warrant a sex-balanced sample size (92 males, 94 females). We investigated both uniparental (mitochondrial ND2 gene) and biparental (microsatellite DNA: 10 loci) genetic systems. The mtDNA provided evidence for demographic expansion yet no significant partition of the genetic variability was disclosed. Nevertheless, a comparatively distant Russian population investigated in another study, whose sequences were included in the present dataset, significantly diverged from all other ones. Different to previous studies, microsatellites highlighted remarkable genetic structure among the studied populations, and pointed to the occurrence of differences between male and female barn swallows. We produced evidence for non-random patterns of gene flow among barn swallow populations probably mediated by female natal dispersal, and we found significant variability in the philopatry of males of different populations. Our data emphasize the importance of taking into account the sex of sampled individuals in order to obtain reliable inferences on species characterized by different patterns of dispersal between males and females.

Individual quality explains variation in reproductive success better than territory quality in a long-lived territorial raptor.

Evolution by natural selection depends on the relationship between individual traits and fitness. Variation in individual fitness can result from habitat (territory) quality and individual variation. Individual quality and specialization can have a deep impact on fitness, yet in most studies on territorial species the quality of territory and individuals are confused. We aimed to determine if variation in breeding success is better explained by territories, individual quality or a combination of both. We analysed the number of fledglings and the breeding quality index (the difference between the number of fledglings of an individual/breeding pair and the average number of fledglings of the monitored territories in the same year) as part of a long term (16 years) peregrine falcon (Falco peregrinus) monitoring program with identification of individuals. Using individual and territory identities as correlates of quality, we built Generalised Linear Models with Mixed effects, in which random factors depicted different hypotheses for sources of variation (territory/individual quality) in the reproductive success of unique breeding pairs, males and females, and assessed their performance. Most evidence supported the hypothesis that variation in breeding success is explained by individual identity, particularly male identity, rather than territory. There is also some evidence for inter year variations in the breeding success of females and a territory effect in the case of males. We argue that, in territorial species, individual quality is a major source of variation in breeding success, often masked by territory. Future ecological and conservation studies on habitat use should consider and include the effect of individuals, in order to avoid misleading results.

Do long lived seabirds reduce the negative effects of acute pollution on adult survival by skipping breeding? A study with European storm petrels (Hydrobates pelagicus) during the "Prestige" oil-spill.

We estimated the survival probability of breeding European storm petrels before, during and after a severe oil-spill. We hypothesized that petrels might have deserted the breeding colony to maximize their own survival probability and we expected no major change on adult survival probabilities as a consequence of the spill. We used an information-theoretical approach and multi-model inference to assess the strength of the evidence in favour of different hypotheses. Evidence contained in the data clearly supported the non-effect of the spill on adult survival hypothesis while punctual impact of the spill on survival and expanded (3 years) impact alternatives received less support. The effect size of the spill on averaged survival estimates was negligible in every case. We suggest that petrels minimized the impact of acute pollution by not investing in reproduction. We suggest that short-medium term management actions after oil-spills and similar catastrophes should focus on ecosystem restoration.

Short-term effects of the prestige oil spill on the peregrine falcon (Falco peregrinus).

We have monitored the distribution, population status, breeding success, turnover rate and diet of a Peregrine Falcon population in Bizkaia (North of Spain) since 1997. On the 13th November 2002, the tanker Prestige sunk off La Coruña (NW Spain) causing an oil spill that affected the whole of the Cantabrian Coast and the Southwest of France. The total number of birds affected by the Prestige oil spill was expected to be between 115,000 and 230,000, some of them raptors. The loss of clutches during the incubation period increased significantly and was correlated with the loss of females. Moreover, the turnover rate of the population increased from 21% to 30%. The polycyclic aromatic hydrocarbon concentrations in the eggs, collected from five nests after they were deserted, ranged from 21.20 ng/g to 461.08 ng/g, values which are high enough to cause the death of the embryos and poisoning of adult birds. The effects of pollution reached inland since some inland-breeding falcons prey on shorebirds that use rivers during their migratory flights. As the Prestige oil spill has clearly resulted in increased rates of adult mortality and reduced fertility, we suggest that the environmental authorities urgently undertake measures aimed at protecting the Peregrine Falcon in Bizkaia.