Social copying drives a tipping point for nonlinear population collapse.

Sudden changes in populations are ubiquitous in ecological systems, especially under perturbations. The agents of global change may increase the frequency and severity of anthropogenic perturbations, but complex populations' responses hamper our understanding of their dynamics and resilience. Furthermore, the long-term environmental and demographic data required to study those sudden changes are rare. Fitting dynamical models with an artificial intelligence algorithm to population fluctuations over 40 y in a social bird reveals that feedback in dispersal after a cumulative perturbation drives a population collapse. The collapse is well described by a nonlinear function mimicking social copying, whereby dispersal made by a few individuals induces others to leave the patch in a behavioral cascade for decision-making to disperse. Once a threshold for deterioration of the quality of the patch is crossed, there is a tipping point for a social response of runaway dispersal corresponding to social copying feedback. Finally, dispersal decreases at low population densities, which is likely due to the unwillingness of the more philopatric individuals to disperse. In providing the evidence of copying for the emergence of feedback in dispersal in a social organism, our results suggest a broader impact of self-organized collective dispersal in complex population dynamics. This has implications for the theoretical study of population and metapopulation nonlinear dynamics, including population extinction, and managing of endangered and harvested populations of social animals subjected to behavioral feedback loops.

Active breeding seabirds prospect alternative breeding colonies.

Compared to other animal movements, prospecting by adult individuals for a future breeding site is commonly overlooked. Prospecting influences the decision of where to breed and has consequences on fitness and lifetime reproductive success. By analysing movements of 31 satellite- and GPS-tracked gull and tern populations belonging to 14 species in Europe and North America, we examined the occurrence and factors explaining prospecting by actively breeding birds. Prospecting in active breeders occurred in 85.7% of studied species, across 61.3% of sampled populations. Prospecting was more common in populations with frequent inter-annual changes of breeding sites and among females. These results contradict theoretical models which predict that prospecting is expected to evolve in relatively predictable and stable environments. More long-term tracking studies are needed to identify factors affecting patterns of prospecting in different environments and understand the consequences of prospecting on fitness at the individual and population level.

Long transient response of vegetation dynamics after four millennia of anthropogenic impacts in an island ecosystem.

Agents of global change commonly have a higher impact on island ecosystem dynamics. In the Mediterranean region, those dynamics have historically been influenced by anthropogenic impacts, for example, the introduction of invasive species and overharvesting of resources. Here, we analysed the spatio-temporal dynamics of vegetation in sa Dragonera island, which experienced a large environmental change ca. 4000 years ago by the arrival of humans. Anthropogenic impacts, such as herbivory by goats and over-logging, ended in the 1970s, while in 2011 the site became the largest Mediterranean island where rats were eradicated. Invasive rats and goats played the ecological role of two endemic species, the cave goat and the giant dormouse, which inhabited the island for more than 5 million years and were rapidly extinct by humans. We used Landsat imagery to explore NDVI as a proxy of vegetation productivity over the years 1984-2021, orthophotos to assess changes in land and vegetation covers and historical plant inventories to study the dynamics in plant diversity. Results showed that those indicators steadily increased both in spring and in summer, while the noise around the trends was partially explained by climate variability. The regime shifts in the temporal dynamics of vegetation productivity suggested a transient from a perturbed to a non-perturbed stable state. Trends in successional dynamics, spatial self-organization and plant diversity also showed the same type of transient dynamics. Historical perturbations related to harvesting (mainly the synergies between goat browsing, burning and forest over-logging) were more important than rat eradication or the influence of climate to explain the vegetation dynamics. Our study shows the transient nature of this small island ecosystem after 4000 years of perturbations and its current path towards vegetation dynamics more controlled by ecological interactions lacking large herbivores and omnivores, drought dynamics and the carrying capacity of the island.

Los agentes del cambio global suelen tener un mayor impacto en la dinámica de los ecosistemas insulares. En la región Mediterránea, esas dinámicas se han visto influenciadas históricamente por impactos antropogénicos, e.g. la introducción de especies invasoras y la sobreexplotación de los recursos. Analizamos aquí la dinámica espacio-temporal de la vegetación en la isla de sa Dragonera, que experimentó un gran cambio ambiental hace unos 4000 años por la llegada de los humanos. Los impactos antropogénicos, como la herbivoría de las cabras y la tala excesiva, terminaron en la década de 1970, mientras que en 2011 se convirtió en la isla mediterránea más grande donde se erradicaron las ratas. Las ratas y cabras invasoras desempeñaron el papel ecológico de dos especies endémicas, el miotrago y el lirón gigante, que habitaron la isla durante más de 5 millones de años y fueron rápidamente extinguidos por los humanos. Usamos imágenes de Landsat para explorar el NDVI (indicador de la productividad de la vegetación) durante los años 1984-2021, ortofotos para evaluar los cambios en la cobertura de la tierra y la vegetación e inventarios históricos de plantas para estudiar la dinámica de su diversidad. Los resultados mostraron que esos indicadores aumentaron constantemente tanto en primavera como en verano, mientras que el ruido en torno a las tendencias se explicaba en parte por la variabilidad climática. Los cambios de régimen en la dinámica temporal de la productividad de la vegetación sugirieron un transitorio de un estado estable perturbado a uno no perturbado. Las tendencias en la dinámica sucesional, la autoorganización espacial y la diversidad de plantas también mostraron el mismo tipo de dinámica transitoria. Las perturbaciones históricas relacionadas con la recolección (principalmente las sinergias entre el pastoreo de cabras, la quema y la tala excesiva de bosques) fueron más importantes que la erradicación de ratas o la influencia del clima para explicar la dinámica de la vegetación. Nuestro estudio muestra la naturaleza transitoria de este ecosistema después de 4000 años de perturbaciones y su trayectoria actual hacia una dinámica de la vegetación más controlada por interacciones ecológicas (que carecen de grandes herbívoros y omnívoros), dinámicas de sequía y la capacidad de carga de la isla.

Multi-species prey dynamics influence local survival in resident and wintering generalist predators.

Stochasticity in food availability influences vital rates such as survival and fertility. Life-history theory predicts that in long-lived organisms, survival should be buffered against environmental stochasticity showing little temporal variability. Furthermore, to optimize survival prospects, many animal species perform migrations to wintering areas where food availability is larger. Species with large latitudinal distribution ranges may show populations that migrate and others that are resident, and they may co-occur in winter. One example of these species is the predatory raptor buzzard Buteo buteo. Here, we test whether temporal variability in the density of five small mammal species of prey inhabiting different habitats (shrubland and forests) influences local annual survival of buzzards in a wintering area depending on their age and residency status (residents versus wintering individuals). We found that prey density explained a considerable amount of annual changes in local survival, which was higher for older and resident birds. This difference in local survival likely corresponded to philopatry to the wintering area, which was larger for residents and increased when prey density was larger. The total density of prey inhabiting open shrublands was the variable explaining more variance in temporal variability of local survival, even though the study area is mostly occupied by woodlands. Temporal population dynamics of the different small mammals inhabiting shrublands were not synchronous, which suggests that buzzards preyed opportunistically on the most abundant prey each winter. Generalist predation may buffer the impact of resource unpredictability for pulsed and asynchronous prey dynamics, typical of small mammals in winter.

High frequency of prospecting for informed dispersal and colonisation in a social species at large spatial scale.

Animals explore and prospect space searching for resources and individuals may disperse, targeting suitable patches to increase fitness. Nevertheless, dispersal is costly because it implies leaving the patch where the individual has gathered information and reduced uncertainty. In social species, information gathered during the prospection process for deciding whether and where to disperse is not only personal but also public, i.e. conspecific density and breeding performance. In empty patches, public information is not available and dispersal for colonisation would be more challenging. Here we study the prospecting in a metapopulation of colonial Audouin's gulls using PTT platform terminal transmitters tagging for up to 4 years and GPS tagging during the incubation period. A large percentage of birds (65%) prospected occupied patches; strikingly, 62% of prospectors also visited empty patches that were colonised in later years. Frequency and intensity of prospecting were higher for failed breeders, who dispersed more than successful breeders. Prospecting and dispersal also occurred mostly to neighbouring patches where population density was higher. GPSs revealed that many breeders (59%) prospected while actively incubating, which suggests that they gathered information before knowing the fate of their reproduction. Prospecting may be enhanced in species adapted to breed in ephemeral habitats, such as Audouin's gulls. Interestingly, none of the tracked individuals colonised an empty patch despite having prospected over a period of up to three consecutive years. Lack of public information in empty patches may drive extended prospecting, long time delays in colonisation and non-linear transient phenomena in metapopulation dynamics and species range expansion.

Assessing the effect of density on population growth when modeling individual encounter data.

The relative role of density-dependent and density-independent variation in vital rates and population size remains largely unsolved. Despite its importance to the theory and application of population ecology, and to conservation biology, quantifying the role and strength of density dependence is particularly challenging. We present a hierarchical formulation of the temporal symmetry approach, also known as the Pradel model, that permits estimation of the strength of density dependence from capture-mark-reencounter data. A measure of relative population size is built in the model and serves to detect density dependence directly on population growth rate. The model is also extended to account for temporal random variability in demographic rates, allowing estimation of the temporal variance of population growth rate unexplained by density dependence. We thus present a model-based approach that enable to test and quantify the effect of density-dependent and density-independent factors affecting population fluctuations in a single modeling framework. More generally, we use this modeling framework along with simulated and empirical data to show the value of including density dependence when modeling individual encounter data without the need for auxiliary data.

Differential adult survival at close seabird colonies: The importance of spatial foraging segregation and bycatch risk during the breeding season.

Marine megafauna, including seabirds, are critically affected by fisheries bycatch. However, bycatch risk may differ on temporal and spatial scales due to the uneven distribution and effort of fleets operating different fishing gear, and to focal species distribution and foraging behavior. Scopoli's shearwater Calonectris diomedea is a long-lived seabird that experiences high bycatch rates in longline fisheries and strong population-level impacts due to this type of anthropogenic mortality. Analyzing a long-term dataset on individual monitoring, we compared adult survival (by means of multi-event capture-recapture models) among three close predator-free Mediterranean colonies of the species. Unexpectedly for a long-lived organism, adult survival varied among colonies. We explored potential causes of this differential survival by (1) measuring egg volume as a proxy of food availability and parental condition; (2) building a specific longline bycatch risk map for the species; and (3) assessing the distribution patterns of breeding birds from the three study colonies via GPS tracking. Egg volume was very similar between colonies over time, suggesting that environmental variability related to habitat foraging suitability was not the main cause of differential survival. On the other hand, differences in foraging movements among individuals from the three colonies expose them to differential mortality risk, which likely influenced the observed differences in adult survival. The overlap of information obtained by the generation of specific bycatch risk maps, the quantification of population demographic parameters, and the foraging spatial analysis should inform managers about differential sensitivity to the anthropogenic impact at mesoscale level and guide decisions depending on the spatial configuration of local populations. The approach would apply and should be considered in any species where foraging distribution is colony-specific and mortality risk varies spatially.

Complex demographic heterogeneity from anthropogenic impacts in a coastal marine predator.

Environmental drivers, including anthropogenic impacts, affect vital rates of organisms. Nevertheless, the influence of these drivers may depend on the physical features of the habitat and how they affect life history strategies depending on individual covariates such as age and sex. Here, the long-term monitoring (1994-2014) of marked European Shags in eight colonies in two regions with different ecological features, such as foraging habitat, allowed us to test several biological hypotheses about how survival changes by age and sex in each region by means of multi-event capture-recapture modeling. Impacts included fishing practices and bycatch, invasive introduced carnivores and the severe Prestige oil spill. Adult survival was constant but, unexpectedly, it was different between sexes. This difference was opposite in each region. The impact of the oil spill on survival was important only for adults (especially for females) in one region and lasted a single year. Juvenile survival was time dependent but this variability was not synchronized between regions, suggesting a strong signal of regional environmental variability. Mortality due to bycatch was also different between sex, age and region. Interestingly the results showed that the size of the fishing fleet is not necessarily a good proxy for assessing the impact of bycatch mortality, which may be more dependent on the fishing grounds and the fishing gears employed in each season of the year. Anthropogenic impacts affected survival differently by age and sex, which was expected for a long-lived organism with sexual size dimorphism. Strikingly, these differences varied depending on the region, indicating that habitat heterogeneity is demographically important to how environmental variability (including anthropogenic impacts) and resilience influence population dynamics.

Varying demographic impacts of different fisheries on three Mediterranean seabird species.

Fisheries have an enormous economic importance, but reconciling their socio-economic features with the conservation and sustainability of marine ecosystems presents major challenges. Bycatch mortality from fisheries is clearly among the most serious global threats for marine ecosystems, affecting a wide range of top predators. Recent estimates report ca. 200,000 seabirds killed annually by bycatch in European waters. However, there is an urgent need to rigorously estimate actual mortality rates and quantify effects of bycatch on populations. The Mediterranean Sea is one of the most impacted regions. Here, we estimate for the first time both bycatch mortality rates and their population-level effects on three endemic and vulnerable Mediterranean taxa: Scopoli's shearwater, Mediterranean shag, and Audouin's gull, that die in different types of fishing gears: longlines, gillnets and sport trolling, respectively. We use multi-event capture-recapture modelling to estimate crucial demographic parameters, including the probabilities of dying in different fishing gears. We then build stochastic demography models to forecast the viability of the populations under different management scenarios. Longline bycatch was particularly severe for adults of Scopoli's shearwaters and Audouin's gulls (ca. 28% and 23% of total mortality, respectively) and also for immature gulls (ca. 90% of mortality). Gillnets had a lower impact, but were still responsible for ca. 9% of juvenile mortality on shags, whereas sport trolling only slightly influenced total mortality in gulls. Bycatch mortality has high population-level impacts in all three species, with shearwaters having the highest extinction risk under current mortality rates. Different life-history traits and compensatory demographic mechanisms between the three species are probably influencing the different bycatch impact: for shearwaters, urgent conservation actions are required to ensure the viability of their populations. Results will be very useful for guiding future seabird conservation policies and moving towards an ecosystem-based approach to sustainable fisheries management.

Oceanographic drivers and mistiming processes shape breeding success in a seabird.

Understanding the processes driving seabirds' reproductive performance through trophic interactions requires the identification of seasonal pulses in marine productivity. We investigated the sequence of environmental and biological processes driving the reproductive phenology and performance of the storm petrel (Hydrobates pelagicus) in the Western Mediterranean. The enhanced light and nutrient availability at the onset of water stratification (late winter/early spring) resulted in annual consecutive peaks in relative abundance of phytoplankton, zooplankton and ichthyoplankton. The high energy-demanding period of egg production and chick rearing coincided with these successive pulses in food availability, pointing to a phenological adjustment to such seasonal patterns with important fitness consequences. Indeed, delayed reproduction with respect to the onset of water stratification resulted in both hatching and breeding failure. This pattern was observed at the population level, but also when confounding factors such as individuals' age or experience were also accounted for. We provide the first evidence of oceanographic drivers leading to the optimal time-window for reproduction in an inshore seabird at southern European latitudes, along with a suitable framework for assessing the impact of environmentally driven changes in marine productivity patterns in seabird performance.

Climate-driven vital rates do not always mean climate-driven population.

Current climatic changes have increased the need to forecast population responses to climate variability. A common approach to address this question is through models that project current population state using the functional relationship between demographic rates and climatic variables. We argue that this approach can lead to erroneous conclusions when interpopulation dispersal is not considered. We found that immigration can release the population from climate-driven trajectories even when local vital rates are climate dependent. We illustrated this using individual-based data on a trans-equatorial migratory seabird, the Scopoli's shearwater Calonectris diomedea, in which the variation of vital rates has been associated with large-scale climatic indices. We compared the population annual growth rate λi , estimated using local climate-driven parameters with ρi , a population growth rate directly estimated from individual information and that accounts for immigration. While λi varied as a function of climatic variables, reflecting the climate-dependent parameters, ρi did not, indicating that dispersal decouples the relationship between population growth and climate variables from that between climatic variables and vital rates. Our results suggest caution when assessing demographic effects of climatic variability especially in open populations for very mobile organisms such as fish, marine mammals, bats, or birds. When a population model cannot be validated or it is not detailed enough, ignoring immigration might lead to misleading climate-driven projections.

Climate change overruns resilience conferred by temperature-dependent sex determination in sea turtles and threatens their survival.

Temperature-dependent sex determination (TSD) is the predominant form of environmental sex determination (ESD) in reptiles, but the adaptive significance of TSD in this group remains unclear. Additionally, the viability of species with TSD may be compromised as climate gets warmer. We simulated population responses in a turtle with TSD to increasing nest temperatures and compared the results to those of a virtual population with genotypic sex determination (GSD) and fixed sex ratios. Then, we assessed the effectiveness of TSD as a mechanism to maintain populations under climate change scenarios. TSD populations were more resilient to increased nest temperatures and mitigated the negative effects of high temperatures by increasing production of female offspring and therefore, future fecundity. That buffered the negative effect of temperature on the population growth. TSD provides an evolutionary advantage to sea turtles. However, this mechanism was only effective over a range of temperatures and will become inefficient as temperatures rise to levels projected by current climate change models. Projected global warming threatens survival of sea turtles, and the IPCC high gas concentration scenario may result in extirpation of the studied population in 50 years.

Population control of an overabundant species achieved through consecutive anthropogenic perturbations.

The control of overabundant vertebrates is often problematic. Much work has focused on population-level responses and overabundance due to anthropogenic subsidies. However, far less work has been directed at investigating responses following the removal of subsidies. We investigate the consequences of two consecutive perturbations, the closure of a landfill and an inadvertent poisoning event, on the trophic ecology (δ13C, δ15N, and δ34S), survival, and population size of an overabundant generalist seabird species, the Yellow-legged Gull (Larus michahellis). We expected that the landfill closure would cause a strong dietary shift and the inadvertent poisoning a decrease in gull population size. As a long-lived species, we also anticipated adult survival to be buffered against the decrease in food availability but not against the inadvertent poisoning event. Stable isotope analysis confirmed the dietary shift towards marine resources after the disappearance of the landfill. Although the survival model was inconclusive, it did suggest that the perturbations had a negative effect on survival, which was followed by a recovery back to average values. Food limitation likely triggered dispersal to other populations, while poisoning may have increased mortality; these two processes were likely responsible for the large fall in population size that occurred after the two consecutive perturbations. Life-history theory suggests that perturbations may encourage species to halt existing breeding investment in order to ensure future survival. However, under strong perturbation pulses the resilience threshold might be surpassed and changes in population density can arise. Consecutive perturbations may effectively manage overabundant species.

From recruitment to senescence: food shapes the age-dependent pattern of breeding performance in a long-lived bird.

We used a long-term data set (26 years) from Audouin's Gull (Larus audouinii), a long-lived seabird, to address the relationship between the age-dependent pattern of reproductive performance and environmental conditions during breeding. Although theoretical models predict that the youngest and oldest breeders (due to inexperience and senescence, respectively) will perform less well than intermediate age classes, few empirical data exist regarding how this expected pattern varies with food availability. To assess the influence of age and food availability (corrected by population size of the main consumers to take into account density dependence) on a number of breeding parameters (laying dates, egg volume, clutch size, and hatching success), we modeled mean and variances of these parameters by incorporating heterogeneity into generalized linear models. All parameters varied with age and to different degrees, depending on food availability. As expected, performance improved with increased food supply, and the observed age pattern was quadratic, with poorer breeding performances occurring in extreme ages. For most parameters (except for laying dates, for which age and food did not interact), the pattern changed with food somewhat unexpectedly; the differences in performance between age classes were higher (i.e., the quadratic pattern was more noticeable) when food was more readily available than when food availability was lower. We suggest that, under poor environmental conditions, only high-quality individuals of the younger and older birds bred and that the differences in breeding performance between age classes were smaller. Although variances for egg volume were constant, variances for laying dates were highest for the youngest breeders and tended to decrease with age, either due to the selection of higher-quality individuals or to a greater frequency of birds skipping breeding with age, especially when food was in low supply. Our results show that mean and variances of breeding parameters changed with age, but that this pattern was different for each parameter and also varied according to food availability. It is likely that, other than food, certain additional factors (e.g., sex, cohort effects, density dependence) also influence changes in breeding performance with age, and this may preclude the finding of a common pattern among traits and among studies on different taxa.

Man-induced activities modify demographic parameters in a long-lived species: effects of poisoning and health policies.

Recent changes in sanitary policies within the European Union (EU) concerning disposal of carcasses of domestic animals and the increase of non-natural mortality factors, such as illegal poisoning, are threatening European vultures. However, the effects of anthropogenic activities on demographic parameters are poorly studied. Using a long-term study (1994-2011) of the threatened Pyrenean Bearded Vulture Gypaetus barbatus population, we assess the variation in the proportion of breeding pairs, egg-laying dates, clutch size, breeding success, and survival following a sharp reduction in food availability in 2005 due to the application of restrictive sanitary policies decreasing livestock carcass availability. We found a delay in laying dates and a regressive trend in clutch size, breeding success, and survival following policy change. The maintenance of specific supplementary feeding stations for Bearded Vultures probably reduced the negative effects of illegal poisoning and food shortages, which mainly affected subadult survival. A drop in food availability may have produced changes in demographic parameters and an increase in mortality due to an increased exposure to contaminated food. As a result, supplementary feeding as a precautionary measure can be a useful tool to reduce illegal poisoning and declines in demographic parameters until previous food availability scenarios are achieved. This study shows how anthropogenic activities through human health regulations that affect habitat quality can suddenly modify demographic parameters in long-lived species, including those, such as survival, with high sensitivity to population growth rate.

Determinants of extinction-colonization dynamics in Mediterranean butterflies: the role of landscape, climate and local habitat features.

Many species are found today in the form of fragmented populations occupying patches of remnant habitat in human-altered landscapes. The persistence of these population networks requires a balance between extinction and colonization events assumed to be primarily related to patch area and isolation, but the contribution of factors such as the characteristics of patch and matrix habitats, the species' traits (habitat specialization and dispersal capabilities) and variation in climatic conditions have seldom been evaluated simultaneously. The identification of environmental variables associated with patch occupancy and turnover may be especially useful to enhance the persistence of multiple species under current global change. However, for robust inference on occupancy and related parameters, we must account for detection errors, a commonly overlooked problem that leads to biased estimates and misleading conclusions about population dynamics. Here, we provide direct empirical evidence of the effects of different environmental variables on the extinction and colonization rates of a rich butterfly community in the western Mediterranean. The analysis was based on a 17-year data set containing detection/nondetection data on 73 butterfly species for 26 sites in north-eastern Spain. Using multiseason occupancy models, which take into account species' detectability, we were able to obtain robust estimates of local extinction and colonization probabilities for each species and test the potential effects of site covariates such as the area of suitable habitat, topographic variability, landscape permeability around the site and climatic variability in aridity conditions. Results revealed a general pattern across species with local habitat composition and landscape features as stronger predictors of occupancy dynamics compared with topography and local aridity. Increasing area of suitable habitat in a site strongly decreased local extinction risks and, for a number of species, both higher amounts of suitable habitat and more permeable landscapes increased colonization rates. Nevertheless, increased topographic variability decreased the extinction risk of bad dispersers, a group of species with significantly lower colonization rates. Our models predicted higher sensitivity of the butterfly assemblages to deterministic changes in habitat features rather than to stochastic weather patterns, with some relationships being clearly dependent on the species' traits.

Ecological and evolutionary implications of food subsidies from humans.

Human activities are the main current driver of global change. From hunter-gatherers through to Neolithic societies-and particularly in contemporary industrialised countries-humans have (voluntarily or involuntarily) provided other animals with food, often with a high spatio-temporal predictability. Nowadays, as much as 30-40% of all food produced in Earth is wasted. We argue here that predictable anthropogenic food subsidies (PAFS) provided historically by humans to animals has shaped many communities and ecosystems as we see them nowadays. PAFS improve individual fitness triggering population increases of opportunistic species, which may affect communities, food webs and ecosystems by altering processes such as competition, predator-prey interactions and nutrient transfer between biotopes and ecosystems. We also show that PAFS decrease temporal population variability, increase resilience of opportunistic species and reduce community diversity. Recent environmental policies, such as the regulation of dumps or the ban of fishing discards, constitute natural experiments that should improve our understanding of the role of food supply in a range of ecological and evolutionary processes at the ecosystem level. Comparison of subsidised and non-subsidised ecosystems can help predict changes in diversity and the related ecosystem services that have suffered the impact of other global change agents.

Contrasting effects of climatic variability on the demography of a trans-equatorial migratory seabird.

Large-scale seasonal climatic indices, such as the North Atlantic Oscillation (NAO) index or the Southern Oscillation Index (SOI), account for major variations in weather and climate around the world and may influence population dynamics in many organisms. However, assessing the extent of climate impacts on species and their life-history traits requires reliable quantitative statistical approaches. We used a new analytical tool in mark-recapture, the multi-event modelling, to simultaneously assess the influence of climatic variation on multiple demographic parameters (i.e. adult survival, transient probability, reproductive skipping and nest dispersal) at two Mediterranean colonies of the Cory's shearwater Calonectris diomedea, a trans-equatorial migratory long-lived seabird. We also analysed the impact of climate in the breeding success at the two colonies. We found a clear temporal variation of survival for Cory's shearwaters, strongly associated to the large-scale SOI especially in one of the colonies (up to 66% of variance explained). Atlantic hurricane season is modulated by the SOI and coincides with shearwater migration to their wintering areas, directly affecting survival probabilities. However, the SOI was a better predictor of survival probabilities than the frequency of hurricanes; thus, we cannot discard an indirect additive effect of SOI via food availability. Accordingly, the proportion of transients was also correlated with SOI values, indicating higher costs of first reproduction (resulting in either mortality or permanent dispersal) when bad environmental conditions occurred during winter before reproduction. Breeding success was also affected by climatic factors, the NAO explaining c. 41% of variance, probably as a result of its effect in the timing of peak abundance of squid and small pelagics, the main prey for shearwaters. No climatic effect was found either on reproductive skipping or on nest dispersal. Contrarily to what we expect for a long-lived organism, large-scale climatic indexes had a more pronounced effect on survival and transient probabilities than on less sensitive fitness parameters such reproductive skipping or nest dispersal probabilities. The potential increase in hurricane frequency because of global warming may interact with other global change agents (such as incidental bycatch and predation by alien species) nowadays impacting shearwaters, affecting future viability of populations.

Drivers of resource allocation for breeding under variable environments in a bet hedger.

The evolutionary theory of life histories predicts that there is a trade-off between survival and reproduction: since adult survival in long-lived organisms is high, then breeding investment is more variable and more dependent on conditions (e.g. food availability and individual experience). Clutch features influence fitness prospects, but how a bet hedger builds its clutch in temporally varying environments is quite unknown. Using 27-year data on 2847 clutches of known-age breeders, we analyse how Audouin's gulls (Larus audouinii), a species showing a combination of conservative and adaptive bet-hedging breeding strategies, can allocate energy by laying clutches and eggs of different sizes. Results show that both food availability and age influenced clutch size and total egg volume in a clutch. Interestingly, we found an interaction between food and age on egg parameters: total volume in two-egg clutches, laid mostly by younger breeders, did not significantly change with food availability and the quadratic pattern in clutch size over the range of ages was less marked as long as food conditions became harsher. With increased food, females invested more by building larger first eggs, whereas they were more conservative on second and third eggs. Furthermore, asymmetries in egg volume within three-egg clutches increased with food availability for old females. Egg size profiles of two-egg clutches suggest that gulls should exhibit progressive reduction of the size of the third egg before shifting to a two-egg clutch size. Food availability influenced all parameters studied, whereas age affected the amount of energy allocated for producing eggs (their size and number) but not the way of allocating those energies (i.e. asymmetries within the clutch). Despite the range of factors affecting the clutch, results suggest that females can allocate the amount of resources in a clutch optimally to increase their fitness under variable environments via bet-hedging.

Increased grey wolf diurnality in southern Europe under human-restricted conditions.

Wolves have been the archetype of wildlife persecution by humans for centuries all over the world, and still are heavily persecuted in some regions. Facultative diurnal/nocturnal wild mammals are known to become more nocturnal when persecuted. Conversely, little is known regarding the possibility of wolves becoming more diurnal if not persecuted. We took advantage of a 9-year natural experiment of restricted human access to a restored coal mine debris dump to study the daily activity patterns of wolves under conditions of infrequent human presence. Results were compared with a paired control site with frequent human use. Circadian wolf activity was monitored using camera traps (3 years in human-restricted site; 2 years in control). Additionally, data from two GPS-GSM-collared wolves monitored in a second control site were also analyzed. In our control sites, wolves were nearly inactive during daylight hours. In contrast, in the human-restricted site wolves extended their activity toward noon, with a daily activity peak between 10:00 and 12:00, and showed some activity throughout the entire circadian 2-h interval cycle considered. Wolves clearly had higher diurnality in the human-restricted area with 78% greater incidence of capture with remote cameras during the day than in the control site. We suggest that the shift toward increased diurnality was related to the loss of fear of humans. Evidence in support of this hypothesis comes from flight initiation distance (FID) data. Wolves showed relatively short FIDs when faced with a human observer (range 70-183 m) in broad daylight at the human-restricted site, but were so afraid of humans in the control site that we were unable to conduct FID trials there. Based on these results, we suggest that wolves may increase their diurnality in those European countries with currently increasing movement of human populations from rural to urban areas and that do not conduct lethal control of wolves. This would represent a historical landmark for a species that has been persecuted for many centuries. However, such behavioral shifts could bring new human-wolf conflicts that would require new policies.