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.

Density-dependence and environmental variability have stage-specific influences on European grayling growth.

Fish somatic growth is indeterminate and can be influenced by a range of abiotic and biotic variables. With climate change forecast to increase the frequency of warming and unusual discharge events, it is thus important to understand how these variables currently influence somatic growth and how that might differ for specific age-classes and/ or life stages. Here, we used a 17-year dataset from a chalk stream in southern England to identify the abiotic and biotic influences on the growth of juvenile, sub-adult and adult life stages of European grayling (Thymallus thymallus), a cold-water riverine salmonid. The results revealed that interannual variations in grayling growth were well described by annual- and site-specific abiotic and biotic explanatory variables. We found divergent responses between life stages to increased temperature and unusual discharge during the main growth period with, for example, elevated temperatures related to increased juvenile growth but reduced sub-adult growth, and high discharge events related to increased sub-adult growth yet reduced juvenile growth. Conversely, stage-specific grayling abundance negatively influenced growth at each life stage, though only juvenile growth was impacted by the abundance of a competitor species, brown trout (Salmo trutta). These results emphasise the merits of testing a wide range of environmental and biological explanatory variables on fish growth, and across life stages. They also reveal the importance of maintaining high habitat heterogeneity in rivers to ensure all life stages can reduce their competitive interactions and have access to adequate flow and thermal refugia during periods of elevated environmental stress.

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.

High summer macrophyte cover increases abundance, growth, and feeding of juvenile Atlantic salmon.

Aquatic habitats are severely threatened by human activities. For anadromous species, managing freshwater habitats to maximize production of more, larger juveniles could improve resilience to threats in marine habitats and enhance population viability. In some juvenile salmonid habitats, complexity created by large substrates provides resources and reduces competitive interactions, thereby promoting juvenile production. In lowland rivers, which lack large substrates, aquatic plants might provide similar complexity and enhance fish productivity. To test the influence of aquatic plants on juvenile Atlantic salmon and sympatric brown trout in a lowland river, we directly manipulated the cover of the dominant macrophyte, Ranunculus, in nine sites during summer and autumn for two years. We quantified the abundance, site retention and growth of salmon and trout under high, medium or low Ranunculus cover. To investigate the effects of Ranunculus cover on feeding opportunities and interspecific competition, we quantified available prey biomass and body size, fish diet composition and compared dietary niche overlap. Experimentally increased Ranunculus cover supported higher salmon abundance in summer and autumn, and higher site retention and growth of salmon in summer. Trout abundance and growth were not influenced by Ranunculus cover, but trout site retention doubled in high, relative to low, cover sites. Despite the weak effects of Ranunculus cover on prey availability, salmon and trout inhabiting high cover sites consumed larger prey and a higher biomass of prey. Furthermore, dietary niche overlap was lower in high, relative to low, cover sites, suggesting that abundant Ranunculus reduced interspecific competition. This field experiment shows that high Ranunculus cover can support more and better growing juvenile salmon, and facilitate foraging and co-existence of sympatric salmonid species. Maintaining or enhancing natural macrophyte cover can be achieved through sympathetic in-river and riparian vegetation management and mitigating pressures on them, such as sediment inputs and low flows, or through planting. Further research should test whether macrophyte cover benefits propagate to subsequent life stages, particularly juvenile overwintering associated with high mortality. This knowledge, in combination with our findings, would further clarify whether beneficial juvenile habitat can improve the viability of at-risk salmonid populations. Overall, our findings suggest that the aims of river restoration might be achieved through promotion of in-stream aquatic vegetation.

Warm winters and cool springs negatively influence recruitment of Atlantic salmon (Salmo salar L.) in a southern England chalk stream.

Previous work suggests that juvenile salmon recruitment in rain-fed rivers is negatively influenced by warm and wet winters and cool springs. We tested whether this is generally applicable to a southern England chalk stream characterized by comparatively stable discharges and temperatures. We found that warm spawning and cool emergence temperatures negatively influenced juvenile recruitment between 2015 and 2020. Together these findings suggest an ability to predict juvenile productivity from water temperature records around spawning and fry emergence, thereby allowing time for management interventions in years of unfavourable temperatures.

Under what circumstances does the capture and tagging of wild Atlantic salmon Salmo salar smolts affect probability of return as adults?

Adult return rates for wild Atlantic salmon Salmo salar smolts captured in a rotary screw trap and tagged with coded wire (CW) tags were compared with a control group, using detections from passive integrated transponder (PIT) antennae systems over 7 years in a small chalk stream in southern England, U.K. Compared with control smolts, capture and CW-tagging of experimental smolts affected detected return rates only under certain conditions, with a decreased return probability for smolts caught and tagged following mild winter river temperature anomalies and during the night. Similarly, analysis of the experimental smolts revealed that capture and CW-tagging following mild winters decreased their probability of return as adults. There were also marginal positive effects of length at PIT-tagging as parr and length at CW-tagging as smolts, on individual probability of return as adult. The results support the hypothesis that the effect of procedures involving the capture and tagging of migrating wild S. salar smolts will vary with the circumstances under which they are performed. The implications of the findings are considered in the context of ongoing investigations to derive and report marine return rates for S. salar in support of national and international stock assessments and in developing best practice.

Review: Allee effects in social species.

Allee effects have important implications for many aspects of basic and applied ecology. The benefits of aggregation of conspecific individuals are central to Allee effects, which have led to the widely held assumption that social species are more prone to Allee effects. Robust evidence for this assumption, however, remains rare. Furthermore, previous research on Allee effects has failed to adequately address the consequences of the different levels of organisation within social species' populations. Here, we review available evidence of Allee effects and model the role of demographic and behavioural factors that may combine to dampen or strengthen Allee effects in social species. We use examples across various species with contrasting social structure, including carnivores, bats, primates and eusocial insects. Building on this, we provide a conceptual framework that allows for the integration of different Allee effects in social species. Social species are characterised by nested levels of organisation. The benefits of cooperation, measured by mean individual fitness, can be observed at both the population and group levels, giving rise to "population level" and "group level" Allee effects respectively. We also speculate on the possibility of a third level, reporting per capita benefits for different individuals within a group (e.g. castes in social insects). We show that group size heterogeneity and intergroup interactions affect the strength of population-level demographic Allee effects. Populations with higher group size heterogeneity and in which individual social groups cooperate demonstrate the weakest Allee effects and may thus provide an explanation for why extinctions due to Allee effects are rare in social species. More adequately accounting for Allee effects in social species will improve our understanding of the ecological and evolutionary implications of cooperation in social species.

Long-term field data and climate-habitat models show that orangutan persistence depends on effective forest management and greenhouse gas mitigation.

BACKGROUND: Southeast Asian deforestation rates are among the world's highest and threaten to drive many forest-dependent species to extinction. Climate change is expected to interact with deforestation to amplify this risk. Here we examine whether regional incentives for sustainable forest management will be effective in improving threatened mammal conservation, in isolation and when combined with global climate change mitigation. METHODOLOGY/PRINCIPAL FINDINGS: Using a long time-series of orangutan nest counts for Sabah (2000-10), Malaysian Borneo, we evaluated the effect of sustainable forest management and climate change scenarios, and their interaction, on orangutan spatial abundance patterns. By linking dynamic land-cover and downscaled global climate model projections, we determine the relative influence of these factors on orangutan spatial abundance and use the resulting statistical models to identify habitat crucial for their long-term conservation. We show that land-cover change the degradation of primary forest had the greatest influence on orangutan population size. Anticipated climate change was predicted to cause reductions in abundance in currently occupied populations due to decreased habitat suitability, but also to promote population growth in western Sabah by increasing the suitability of presently unoccupied regions. CONCLUSIONS/SIGNIFICANCE: We find strong quantitative support for the Sabah government's proposal to implement sustainable forest management in all its forest reserves during the current decade; failure to do so could result in a 40 to 80 per cent regional decline in orangutan abundance by 2100. The Sabah orangutan is just one (albeit iconic) example of a forest-dependent species that stands to benefit from sustainable forest management, which promotes conservation of existing forests.

Limited evidence for the demographic Allee effect from numerous species across taxa.

Extensive theoretical work on demographic Allee effects has led to the latent assumption that they are ubiquitous in natural populations, yet current empirical support for this phenomenon is sparse. We extended previous single-taxon analyses to evaluate the empirical support for demographic Allee effects in the per capita population growth rate of 1198 natural populations spanning all major taxa. For each population, we quantified the empirical support for five population growth models: no growth (random walk); exponential growth, with and without an Allee effect; and logistic growth, with and without an Allee effect. We used two metrics to quantify empirical support, information-theoretic and Bayesian strength of evidence, and observed top-rank frequency. The Ricker logistic model was both the most supported and most frequently top-ranked model, followed by random walk. Allee models had a combined relative support of 12.0% but were top-ranked in only 1.1% of the time series. Accounting for local climate variation and measurement error caused the loss of top-ranked Allee models, although the latter also increased their relative support. The 13 time series exhibiting Allee models were shorter and less variable than other time series, although only three were non-trending. Time series containing observations at low abundance were not more likely and did not show higher support for Allee effect models. We conclude that there is relatively high potential for demographic Allee effects in these 1198 time series but comparatively few observed cases, perhaps due to the influences of climate and measurement error.

Safety in numbers: extinction arising from predator-driven Allee effects.

Experimental evidence of extinction via an Allee effect (AE) is a priority as more species become threatened by human activity. Kramer & Drake (2010) begin the International Year of Biodiversity with the important--but double-edged--demonstration that predators can induce an AE in their prey. The good news is that their experiments help bridge the knowledge gap between theoretical and empirical AEs. The bad news is that this predator-driven AE precipitates the prey extinction via a demographic AE. Although their findings will be sensitive to departures from their experimental protocol, this link between predation and population extinction could have important consequences for many prey species.

Space invaders? A search for patterns underlying the coexistence of alien black rats and Galápagos rice rats.

The introduction and spread of the black rat Rattus rattus is believed to have caused the worst decline of any vertebrate taxon in Galápagos. However, the "extinct" Santiago rice rat Nesoryzomys swarthi has recently been rediscovered in sympatry with R. rattus providing the first exception to this general pattern of displacement. We carried out an exploratory investigation of this novel system with the aim of identifying patterns that may facilitate the apparent coexistence of the two species. We carried out an extensive survey of Santiago Island to map the current distribution of the endemic rice rat and to explore broad scale distribution-habitat associations. We then used live-trapping, radio-tracking, and spool-and-line tracking to quantify abundance-habitat correlations and to test for evidence of interspecific spatial segregation, alteration of N. swarthi activity patterns (spatial and temporal), and microhabitat partitioning. We found that N. swarthi has disappeared from part of its historical range and appears to be restricted to a 14 km stretch of the north-central coast, characterised by high density of the cactus Opuntia galapageia. In contrast, the generalist R. rattus was found at all survey sites. We found no evidence of spatial segregation, and home range size, temporal activity and density of N. swarthi did not vary with local density of R. rattus. However, pre-dawn and post-dusk N. swarthi activity levels increased with R. rattus density perhaps reflecting an increase in foraging effort necessary to compensate for the costs of interspecific exploitation or interference competition. The distribution, microhabitat selection, and abundance-habitat relations of N. swarthi suggest that the endemic cactus O. galapageia may facilitate interspecific coexistence. Further research should include a comparison of inter-seasonal resource preference and foraging activity of the two species coupled with replicated field experiments to confirm and quantify competition and to elucidate the mechanism of competitive coexistence.