A multi-population approach supports common patterns in marine growth and maturation decision in Atlantic salmon (Salmo salar L.) from southern Europe.

This study provides a regional picture of long-term changes in Atlantic salmon growth at the southern edge of their distribution, using a multi-population approach spanning 49 years and five populations. We provide empirical evidence of salmon life history being influenced by a combination of common signals in the marine environment and population-specific signals. We identified an abrupt decline in growth from 1976 and a more recent decline after 2005. As these declines have also been recorded in northern European populations, our study significantly expands a pattern of declining marine growth to include southern European populations, thereby revealing a large-scale synchrony in marine growth patterns for almost five decades. Growth increments during their sea sojourn were characterized by distinct temporal dynamics. At a coarse temporal resolution, growth during the first winter at sea seemed to gradually improve over the study period. However, the analysis of finer seasonal growth patterns revealed ecological bottlenecks of salmon life histories at sea in time and space. Our study reinforces existing evidence of an impact of early marine growth on maturation decision, with small-sized individuals at the end of the first summer at sea being more likely to delay maturation. However, each population was characterized by a specific probabilistic maturation reaction norm, and a local component of growth at sea in which some populations have better growth in some years might further amplify differences in maturation rate. Differences between populations were smaller than those between sexes, suggesting that the sex-specific growth threshold for maturation is a well-conserved evolutionary phenomenon in salmon. Finally, our results illustrate that although most of the gain in length occurs during the first summer at sea, the temporal variability in body length at return is buffered against the decrease in post-smolt growth conditions. The intricate combination of growth over successive seasons, and its interplay with the maturation decision, could be regulating body length by maintaining diversity in early growth trajectories, life histories, and the composition of salmon populations.

Predicting how environmental conditions and smolt body length when entering the marine environment impact individual Atlantic salmon Salmo salar adult return rates.

Populations of Atlantic salmon Salmo salar have experienced precipitous declines in abundance since the 1970s. This decline has been associated with reduced numbers of adult salmon returning to fresh water from their marine migration, i.e., their marine return rates (MRR). Thus, understanding the factors that affect MRR is of crucial conservation importance. The authors used a state-space model with a 13-year time series of individually tagged salmon mark-recapture histories on the River Frome, southern England, to test the effect of smolt body length on their MRR. In addition to smolt length, the model tested for the influence of environmental covariates that were representative of the conditions experienced by the smolts in the early stages of their seaward migration, i.e., from the lower river to the estuary exit. The model indicated that, even when accounting for environmental covariates, smolt body length was an important predictor of MRR. Although larger smolts have a higher probability of returning to their natal river as adults than smaller smolts, and one-sea-winter salmon have a survival rate twice as high as multi-sea-winter salmon, the actual biological mechanisms underpinning this phenomenon remain uncertain. These results have important applications for salmon conservation, as efforts to bolster salmon populations in the freshwater environment should consider methods to improve smolt quality (i.e., body size) as well as smolt quantity.

Spatial synchrony in the response of a long range migratory species (Salmo salar) to climate change in the North Atlantic Ocean.

A major challenge in understanding the response of populations to climate change is to separate the effects of local drivers acting independently on specific populations, from the effects of global drivers that impact multiple populations simultaneously and thereby synchronize their dynamics. We investigated the environmental drivers and the demographic mechanisms of the widespread decline in marine survival rates of Atlantic salmon (Salmo salar) over the last four decades. We developed a hierarchical Bayesian life cycle model to quantify the spatial synchrony in the marine survival of 13 large groups of populations (called stock units, SU) from two continental stock groups (CSG) in North America (NA) and Southern Europe (SE) over the period 1971-2014. We found strong coherence in the temporal variation in postsmolt marine survival among the 13 SU of NA and SE. A common North Atlantic trend explains 37% of the temporal variability of the survivals for the 13 SU and declines by a factor of 1.8 over the 1971-2014 time series. Synchrony in survival trends is stronger between SU within each CSG. The common trends at the scale of NA and SE capture 60% and 42% of the total variance of temporal variations, respectively. Temporal variations of the postsmolt survival are best explained by the temporal variations of sea surface temperature (SST, negative correlation) and net primary production indices (PP, positive correlation) encountered by salmon in common domains during their marine migration. Specifically, in the Labrador Sea/Grand Banks for populations from NA, 26% and 24% of variance is captured by SST and PP, respectively and in the Norwegian Sea for populations from SE, 21% and 12% of variance is captured by SST and PP, respectively. The findings support the hypothesis of a response of salmon populations to large climate-induced changes in the North Atlantic simultaneously impacting populations from distant continental habitats.

Manual fish length measurement accuracy for adult river fish using an acoustic camera (DIDSON).

Total lengths (LT ) of 50 free-swimming fish in a tank, silver carp Hypophthalmichthys molitrix and rainbow trout Oncorhynchus mykiss, were measured using a DIDSON (Dual-frequency IDentification SONar) camera. Using Sound Metrics software, multiple measurements of each fish (LT , side aspect angle and distance from the camera) at different times were analysed by two experienced operators while a subset of data was analysed by two inexperienced operators. The main result showed high variability in intra-fish LT measurements. The number of measurements required to minimise errors and to obtain robust fish measurements (true LT ± 3 cm) was estimated by a bootstrap method. Three to five measurements per fish were recommended for fish surveys in rivers. In this experimental study, aiming to reproduce river conditions, no evidence of fish position (side aspect angle and distance from the camera) effect was detected, but an operator effect (partially explained by training) was observed. General linear mixed models also showed that lengths of the smallest fish (LT < 57 cm) were overestimated and lengths of the largest fish (LT > 57 cm) were underestimated in comparison with their true lengths. In conclusion, we highlight that this technology, like any monitoring methods, returns imperfect observations. We advise DIDSON users to ensure that measurements are carried out correctly in order to draw accurate conclusion from this new technology.

Intra- and interspecific densities shape life-history traits in a salmonid population.

Population dynamics can be regulated through intra- and interspecific density dependence. In species with close ecological requirements, interspecific competition for resources may add to intraspecific density, or even exceed its effect; it may impact single or multiple traits. However, the relative impact of intra- and interspecific densities on demographic parameters has been rarely empirically assessed. We analyzed 18 years of capture-mark-recapture data from brown trout (Salmo trutta) coexisting with Atlantic salmon (Salmo salar) during the juvenile freshwater phase in the Oir River (France) to estimate the relative effects of intra- and interspecific density on trout early life. In trout, a species with optional migration, we estimated the migration probability of young-of-the-year trout out of their natal site, survival probability during the first winter, as well as body size, in relation to both intra- and interspecific density. Trout density correlated negatively with body size and with winter survival in resident trout but not with trout migration. Salmon density correlated positively with trout migration, but no impact was detected on trout body size or survival. Our study highlighted contrasting effects of intra- and interspecific density on trout early life, and the need to account for both factors when studying population dynamics in coexisting species. In particular, by affecting trout migration decision, salmon density may drive trout life history.

Contrasting effects of tropical cyclones on the annual survival of a pelagic seabird in the Indian Ocean.

Tropical cyclones are renowned for their destructive nature and are an important feature of marine and coastal tropical ecosystems. Over the last 40 years, their intensity, frequency and tracks have changed, partly in response to ocean warming, and future predictions indicate that these trends are likely to continue with potential consequences for human populations and coastal ecosystems. However, our understanding of how tropical cyclones currently affect marine biodiversity, and pelagic species in particular, is limited. For seabirds, the impacts of cyclones are known to be detrimental at breeding colonies, but impacts on the annual survival of pelagic adults and juveniles remain largely unexplored and no study has simultaneously explored the direct impacts of cyclones on different life-history stages across the annual life cycle. We used a 20-year data set on tropical cyclones in the Indian Ocean, tracking data from 122 Round Island petrels and long-term capture-mark-recapture data to explore the impacts of tropical cyclones on the survival of adult and juvenile (first year) petrels during both the breeding and migration periods. The tracking data showed that juvenile and adult Round Island petrels utilize the three cyclone regions of the Indian Ocean and were potentially exposed to cyclones for a substantial part of their annual cycle. However, only juvenile petrel survival was affected by cyclone activity; negatively by a strong cyclone in the vicinity of the breeding colony and positively by increasing cyclone activity in the Northern Indian Ocean where they spend the majority of their first year at sea. These contrasting effects raise the intriguing prospect that the projected changes in cyclones under current climate change scenarios may have positive as well as the more commonly perceived negative impacts on marine biodiversity.

The short- and long-term fitness consequences of natal dispersal in a wild bird population.

Dispersal is a key process in population and evolutionary ecology. Individual decisions are affected by fitness consequences of dispersal, but these are difficult to measure in wild populations. A long-term dataset on a geographically closed bird population, the Mauritius kestrel, offers a rare opportunity to explore fitness consequences. Females dispersed further when the availability of local breeding sites was limited, whereas male dispersal correlated with phenotypic traits. Female but not male fitness was lower when they dispersed longer distances compared to settling close to home. These results suggest a cost of dispersal in females. We found evidence of both short- and long-term fitness consequences of natal dispersal in females, including reduced fecundity in early life and more rapid aging in later life. Taken together, our results indicate that dispersal in early life might shape life history strategies in wild populations.

Population regulation of territorial species: both site dependence and interference mechanisms matter.

Spatial patterns of site occupancy are commonly driven by habitat heterogeneity and are thought to shape population dynamics through a site-dependent regulatory mechanism. When examining this, however, most studies have only focused on a single vital rate (reproduction), and little is known about how space effectively contributes to the regulation of population dynamics. We investigated the underlying mechanisms driving density-dependent processes in vital rates in a Mauritius kestrel population where almost every individual was monitored. Different mechanisms acted on different vital rates, with breeding success regulated by site dependence (differential use of space) and juvenile survival by interference (density-dependent competition for resources). Although territorial species are frequently assumed to be regulated through site dependence, we show that interference was the key regulatory mechanism in this population. Our integrated approach demonstrates that the presence of spatial processes regarding one trait does not mean that they necessarily play an important role in regulating population growth, and demonstrates the complexity of the regulatory process.

Contrasted demographic responses facing future climate change in Southern Ocean seabirds.

1. Recent climate change has affected a wide range of species, but predicting population responses to projected climate change using population dynamics theory and models remains challenging, and very few attempts have been made. The Southern Ocean sea surface temperature and sea ice extent are projected to warm and shrink as concentrations of atmospheric greenhouse gases increase, and several top predator species are affected by fluctuations in these oceanographic variables. 2. We compared and projected the population responses of three seabird species living in sub-tropical, sub-Antarctic and Antarctic biomes to predicted climate change over the next 50 years. Using stochastic population models we combined long-term demographic datasets and projections of sea surface temperature and sea ice extent for three different IPCC emission scenarios (from most to least severe: A1B, A2, B1) from general circulation models of Earth's climate. 3. We found that climate mostly affected the probability to breed successfully, and in one case adult survival. Interestingly, frequent nonlinear relationships in demographic responses to climate were detected. Models forced by future predicted climatic change provided contrasted population responses depending on the species considered. The northernmost distributed species was predicted to be little affected by a future warming of the Southern Ocean, whereas steep declines were projected for the more southerly distributed species due to sea surface temperature warming and decrease in sea ice extent. For the most southerly distributed species, the A1B and B1 emission scenarios were respectively the most and less damaging. For the two other species, population responses were similar for all emission scenarios. 4. This is among the first attempts to study the demographic responses for several populations with contrasted environmental conditions, which illustrates that investigating the effects of climate change on core population dynamics is feasible for different populations using a common methodological framework. Our approach was limited to single populations and have neglected population settlement in new favourable habitats or changes in inter-specific relations as a potential response to future climate change. Predictions may be enhanced by merging demographic population models and climatic envelope models.

Bet-hedging response to environmental variability, an intraspecific comparison.

A major challenge in ecology is to understand the impact of increased environmental variability on populations and ecosystems. To maximize their fitness in a variable environment, life history theory states that individuals should favor a bet-hedging strategy, involving a reduction of annual breeding performance and an increase in adult survival so that reproduction can be attempted over more years. As a result, evolution toward longer life span is expected to reduce the deleterious effects of extra variability on population growth, and consequently on the trait contributing the most to it (e.g., adult survival in long-lived species). To investigate this, we compared the life histories of two Black-browed Albatross (Thalassarche melanophrys) populations breeding at South Georgia (Atlantic Ocean) and Kerguelen (Indian Ocean), the former in an environment nearly three times more variable climatically (e.g., in sea surface temperature) than the latter. As predicted, individuals from South Georgia (in the more variable environment) showed significantly higher annual adult survival (0.959, SE = 0.003) but lower annual reproductive success (0.285 chick per pair, SE = 0.039) than birds from Kerguelen (survival = 0.925, SE = 0.004; breeding success = 0.694, SE = 0.027). In both populations, climatic conditions affected the breeding success and the survival of inexperienced breeders, whereas the survival of experienced breeders was unaffected. The strength of the climatic impact on survival of inexperienced breeders was very similar between the two populations, but the effect on breeding success was positively related to environmental variability. These results provide rare and compelling evidence to support bet-hedging underlying changes in life history traits as an adaptive response to environmental variability.

Long- and short-term influence of environment on recruitment in a species with highly delayed maturity.

Short-term effects of environmental perturbations on various life history traits are reasonably well documented in birds and mammals. But, in the present context of global climate change, there is a need to consider potential long-term effects of natal conditions to better understand and predict the consequences of these changes on population dynamics. The environmental conditions affecting offspring during their early development may determine their lifetime reproductive performance, and therefore the number of recruits produced by a cohort. In this study, we attempted to link recruitment to natal and recent (previous year) conditions in the long-lived black-browed albatross (Thalassarche melanophrys) at Kerguelen Islands. The environmental variability was described using both climatic variables over breeding (sea surface temperature anomaly) and non-breeding grounds (Southern Oscillation index), and variables related to the colony (breeding success and colony size). Immature survival was linked to the breeding success of the colony in the year of birth, which was expected to reflect the average seasonal parental investment. At the cohort level, this initial mortality event may act as a selective filter shaping the number, and presumably the quality (breeding frequency, breeding success probability), of the individuals that recruit into the breeding population. The decision to start breeding was strongly structured by the age of the individuals and adjusted according to recent conditions. An effect of natal conditions was not detected on this parameter, supporting the selection hypothesis. Recruitment, as a whole, was thus influenced by a combination of long- and short-term environmental impacts. Our results highlight the complexity of the influence of environmental factors on such long-lived species, due to the time-lag (associated with a delayed maturity) between the impact of natal conditions on individuals and their repercussion on the breeding population.

Nonlinear impact of climate on survival in a migratory white stork population.

1. There is growing evidence that ongoing climate change affects populations and species. Physiological limitation and phenotypic plasticity suggest nonlinear response of vital rates to climatic parameters, the intensity of environmental impact might be more pronounced while the frequency of extreme events increases. However, a poor understanding of these patterns presently hampers our predictive capabilities. 2. A recent climatic shift in the Sahel, from droughty to less severe condition, offers a good opportunity to test for an influence of the climatic regime on the response of organisms to their environment. Using a long-term capture-mark-recapture data set on a white stork (Ciconia ciconia) population wintering in Sahel, we investigated potential change in the impact of environmental conditions on survival and recruitment probabilities between 1981 and 2003. 3. We observed a decrease in the strength of the link between survival and Sahel rainfall during the last decade, down to a nondetectable level. Whether Sahel climate was found to affect the survival of storks under droughty conditions, individuals did not seem to respond to climatic variation when precipitation was more abundant. 4. This result gives evidence to a nonlinear response of a migrant bird to wintering environment. Present climate seems to fluctuate within a range of condition providing enough resources to maximize stork's survival. It suggests that whereas inter-annual variability impacted individuals, pluri-annual average condition affected the intensity of this impact. Such pattern may be more widespread than thought, and its modelling will be crucial to predict the impact of future climate change on population dynamics.

Combined spatio-temporal impacts of climate and longline fisheries on the survival of a trans-equatorial marine migrant.

Predicting the impact of human activities and their derivable consequences, such as global warming or direct wildlife mortality, is increasingly relevant in our changing world. Due to their particular life history traits, long-lived migrants are amongst the most endangered and sensitive group of animals to these harming effects. Our ability to identify and quantify such anthropogenic threats in both breeding and wintering grounds is, therefore, of key importance in the field of conservation biology. Using long-term capture-recapture data (34 years, 4557 individuals) and year-round tracking data (4 years, 100 individuals) of a trans-equatorial migrant, the Cory's shearwater (Calonectris diomedea), we investigated the impact of longline fisheries and climatic variables in both breeding and wintering areas on the most important demographic trait of this seabird, i.e. adult survival. Annual adult survival probability was estimated at 0.914±0.022 on average, declining throughout 1978-1999 but recovering during the last decade (2005-2011). Our results suggest that both the incidental bycatch associated with longline fisheries and high sea surface temperatures (indirectly linked to food availability; SST) increased mortality rates during the long breeding season (March-October). Shearwater survival was also negatively affected during the short non-breeding season (December-February) by positive episodes of the Southern Oscillation Index (SOI). Indirect negative effects of climate at both breeding (SST) and wintering grounds (SOI) had a greater impact on survival than longliner activity, and indeed these climatic factors are those which are expected to present more unfavourable trends in the future. Our work underlines the importance of considering both breeding and wintering habitats as well as precise schedules/phenology when assessing the global role of the local impacts on the dynamics of migratory species.

Environmental variation and experience-related differences in the demography of the long-lived black-browed albatross.

1. It has been largely demonstrated that demographic performances of animals increase with age or experience as a result of an improvement of foraging skills, an increasing reproductive effort or a selection process. However, little is known about the age or experience-related response of populations to environmental variations. Theoretical studies consider that age-related variations of the performances are greater under more restricting conditions, but this has rarely been tested. 2. We tested this hypothesis on a long-lived species, black browed albatross Thalassarche melanophrys Temminck, using a long-term capture-mark-recapture data set. We investigated the responses of a population to climate, by studying the effects of climatic factors and breeding experience on survival and breeding success. 3. First-time breeders appear to be poorer performers compared with experienced adults, with lower reproductive success and lower survival. In addition, interannual variations of demographic traits were partly explained by climatic indices, reflecting environmental variations. The survival probability of black-browed albatrosses varied with experience and climate, and differences being greater under harsh conditions. By contrast, the reproductive success of inexperienced individuals was affected by climatic fluctuations in the same way as the experienced ones. 4. First breeding event acts as a strong selective process on the highly heterogeneous class of inexperienced individuals, suggesting the increase in survival and breeding success with experience may mainly reflect a reduction in the heterogeneity among individual qualities.