Size- and stage-dependence in cause-specific mortality of migratory brown trout.

Evidence-based management of natural populations under strong human influence frequently requires not only estimates of survival but also knowledge about how much mortality is due to anthropogenic vs. natural causes. This is the case particularly when individuals vary in their vulnerability to different causes of mortality due to traits, life history stages, or locations. Here, we estimated harvest and background (other cause) mortality of landlocked migratory salmonids over half a century. In doing so, we quantified among-individual variation in vulnerability to cause-specific mortality resulting from differences in body size and spawning location relative to a hydropower dam. We constructed a multistate mark-recapture model to estimate harvest and background mortality hazard rates as functions of a discrete state (spawning location) and an individual time-varying covariate (body size). We further accounted for among-year variation in mortality and migratory behaviour and fit the model to a unique 50-year time series of mark-recapture-recovery data on brown trout (Salmo trutta) in Norway. Harvest mortality was highest for intermediate-sized trout, and outweighed background mortality for most of the observed size range. Background mortality decreased with body size for trout spawning above the dam and increased for those spawning below. All vital rates varied substantially over time, but a trend was evident only in estimates of fishers' reporting rate, which decreased from over 50% to less than 10% throughout the study period. We highlight the importance of body size for cause-specific mortality and demonstrate how this can be estimated using a novel hazard rate parameterization for mark-recapture models. Our approach allows estimating effects of individual traits and environment on cause-specific mortality without confounding, and provides an intuitive way to estimate temporal patterns within and correlation among different mortality sources.

Individual heterogeneity and early life conditions shape growth in a freshwater top predator.

Body size can have profound impacts on survival, movement, and reproductive schedules shaping individual fitness, making growth a central process in ecological and evolutionary dynamics. Realized growth is the result of a complex interplay between life history schedules, individual variation, and environmental influences. Integrating all of these aspects into growth models is methodologically difficult, depends on the availability of repeated measurements of identifiable individuals, and consequently represents a major challenge in particular for natural populations. Using a unique 30-yr time series of individual length measurements inferred from scale year rings of wild brown trout, we develop a Bayesian hierarchical model to estimate individual growth trajectories in temporally and spatially varying environments. We reveal a gradual decrease in average juvenile growth, which has carried over to adult life and contributed to decreasing sizes observed at the population level. Commonly studied environmental drivers like temperature and water flow did not explain much of this trend and overall persistent and among-year individual variation dwarfed temporal variation in growth patterns. Our model and results are relevant to a wide range of questions in ecology and evolution requiring a detailed understanding of growth patterns, including conservation and management of many size-structured populations.

Seasonal effects of Pacific-based climate on recruitment in a predator-limited large herbivore.

1. Climate is an important factor influencing the population dynamics of large herbivores operating directly on individuals or through its effect on forage characteristics. However, the seasonal effect of climate may differ between forage- and predator-limited populations because of a climatic influence on predation rates. The influence of climate on predator-limited large herbivores is less well known than on forage-limited populations. Further, the effect of Pacific-based climate on large herbivore populations has been rarely assessed. 2. We investigated the effect of the Pacific Decadal Oscillation (PDO), across different seasons, on recruitment in 10 populations (herds) of mountain-dwelling caribou Rangifer tarandus caribou L. in the Yukon Territory, Canada. These low-density populations occur in highly seasonal environments and are considered predator-limited with high neonatal calf mortality. Hence, in most years females do not spend resources through lactational support during the summer and resource intake is devoted to self-maintenance. We predicted that climate affecting environmental conditions at calving would have a strong effect on recruitment via its influence on predation rates. We also predicted that climatic conditions prior to conception could have an effect on recruitment through its influence on female fecundity. We modelled recruitment (n = 165) by seasonal PDO values using generalized linear mixed-effects models with herd-varying coefficients. 3. We found that recruitment variability was best explained by variation in winter climate (beta = 0.110, SE = 0.007) prior to birth (in utero) and May climate (beta = 0.013, SE = 0.006) at calving. There was little support for a pre-conception climate effect influencing female body condition and hence fecundity. These results confirm that recruitment in these populations is limited by predation and that forage-limitation is not a significant factor in their population dynamics. There was considerable variability in herd-specific relationships between the PDO and recruitment. Incorporating herd-specific characteristics, such as variable predator densities or terrain characteristics within a herd range, may shed greater light on the complex relationship between climate and ungulate population dynamics.

Thermal plasticity in postembryonic life history traits of a widely distributed Collembola: Effects of macroclimate and microhabitat on genotypic differences.

Life history traits in many ectotherms show complex patterns of variation among conspecific populations sampled along wide latitudinal or climatic gradients. However, few studies have assessed whether these patterns can be explained better by thermal reaction norms of multiple life history traits, covering major aspects of the life cycle. In this study, we compared five populations of a Holarctic, numerically dominant soil microarthropod species, Folsomia quadrioculata, sampled from a wide latitudinal gradient (56-81°N), for growth, development, fecundity, and survival across four temperatures (10, 15, 20, and 25°C) in common garden experiments. We evaluated the extent to which macroclimate could explain differences in thermal adaptation and life history strategies among populations. The common garden experiments revealed large genotypic differences among populations in all the traits, which were little explained by latitude and macroclimate. In addition, the life history strategies (traits combined) hardly revealed any systematic difference related to latitude and macroclimate. The overall performance of the northernmost population from the most stochastic microclimate and the southernmost population, which remains active throughout the year, was least sensitive to the temperature treatments. In contrast, performance of the population from the most predictable microclimate peaked within a narrow temperature range (around 15°C). Our findings revealed limited support for macroclimate-based predictions, and indicated that local soil habitat conditions related to predictability and seasonality might have considerable influence on the evolution of life history strategies of F. quadrioculata. This study highlights the need to combine knowledge on microhabitat characteristics, and demography, with findings from common garden experiments, for identifying the key drivers of life history evolution across large spatial scales, and wide climate gradients. We believe that similar approaches may substantially improve the understanding of adaptation in many terrestrial ectotherms with low dispersal ability.

Responses of earthworms to repeated exposure to three biocides applied singly and as a mixture in an agricultural field.

The study aimed at investigating effects of three differently acting biocides; the insecticide esfenvalerate, the fungicide picoxystrobin and the bactericide triclosan, applied individually and as a mixture, on an earthworm community in the field. A concentration-response design was chosen and results were analyzed using univariate and multivariate approaches. Effects on juvenile proportions were less pronounced and more variable than effects on abundance, but effects in general were species- and chemical-specific, and temporal variations distinct. Esfenvalerate and picoxystrobin appeared to elicit stronger effects than triclosan at laboratory-based ECx values, which is in accordance with our previous laboratory study on Eisenia fetida. The mixture affected abundance and juvenile proportions, but the latter only at high mixture concentrations. Esfenvalerate and picoxystrobin appeared to be the main drivers for the mixture's toxicity. Species-specific toxicity patterns question the reliability of mixture toxicity predictions derived on E. fetida for field earthworms. Biocide concentrations equaling EC50s (reproduction) for E. fetida provoked effects on the field earthworms mainly exceeding 50%, indicating effect intensification from the laboratory to field as well as the influence of indirect effects produced by species interactions. The differing results of the present field study and the previous laboratory study imply that lower- and higher-tier studies may not be mutually exclusive, but to be used in complementary.

Optimal body size and energy expenditure during winter: why are voles smaller in declining populations?

Winter is energetically challenging for small herbivores because of greater energy requirements for thermogenesis at a time when little energy is available. We formulated a model predicting optimal wintering body size, accounting for the scaling of both energy expenditure and assimilation to body size, and the trade-off between survival benefits of a large size and avoiding survival costs of foraging. The model predicts that if the energy cost of maintaining a given body mass differs between environments, animals should be smaller in the more demanding environments, and there should be a negative correlation between body mass and daily energy expenditure (DEE) across environments. In contrast, if animals adjust their energy intake according to variation in survival costs of foraging, there should be a positive correlation between body mass and DEE. Decreasing temperature always increases equilibrium DEE, but optimal body mass may either increase or decrease in colder climates depending on the exact effects of temperature on mass-specific survival and energy demands. Measuring DEE with doubly labeled water on wintering Microtus agrestis at four field sites, we found that DEE was highest at the sites where voles were smallest despite a positive correlation between DEE and body mass within sites. This suggests that variation in wintering body mass between sites was due to variation in food quality/availability and not adjustments in foraging activity to varying risks of predation.

Disease dynamics in cyclic populations of field voles (Microtus agrestis): cowpox virus and vole tuberculosis (Mycobacterium microti).

The possible role of pathogens in rodent population cycles has been largely neglected since Elton's 'epidemic hypothesis' of 1931. To revisit this question, 12 adjacent, cyclic but out-of-phase populations of field voles (Microtus agrestis) in North East England were studied and the initial results are presented here. The prevalences of antibodies to cowpox virus and of clinical signs of Mycobacterium microti infection (vole tuberculosis) showed delayed (not direct) density dependence (with a lag of three to six months). This did not result from changes in population structure, even though there were such changes associated with the different phases of the cycle. The prevalences rose as vole numbers rose, and peaked as numbers declined. The apparent lag in the numerical response of infection prevalence to changes in host abundance is consistent with the hypothesis that diseases, singly or in combination, play a hitherto underestimated role in the dynamics of cyclic populations.

Challenging claims in the study of migratory birds and climate change.

Recent shifts in phenology in response to climate change are well established but often poorly understood. Many animals integrate climate change across a spatially and temporally dispersed annual life cycle, and effects are modulated by ecological interactions, evolutionary change and endogenous control mechanisms. Here we assess and discuss key statements emerging from the rapidly developing study of changing spring phenology in migratory birds. These well-studied organisms have been instrumental for understanding climate-change effects, but research is developing rapidly and there is a need to attack the big issues rather than risking affirmative science. Although we agree poorly on the support for most claims, agreement regarding the knowledge basis enables consensus regarding broad patterns and likely causes. Empirical data needed for disentangling mechanisms are still scarce, and consequences at a population level and on community composition remain unclear. With increasing knowledge, the overall support ('consensus view') for a claim increased and between-researcher variability in support ('expert opinions') decreased, indicating the importance of assessing and communicating the knowledge basis. A proper integration across biological disciplines seems essential for the field's transition from affirming patterns to understanding mechanisms and making robust predictions regarding future consequences of shifting phenologies.

Rapid advance of spring arrival dates in long-distance migratory birds.

Several bird species have advanced the timing of their spring migration in response to recent climate change. European short-distance migrants, wintering in temperate areas, have been assumed to be more affected by change in the European climate than long-distance migrants wintering in the tropics. However, we show that long-distance migrants have advanced their spring arrival in Scandinavia more than short-distance migrants. By analyzing a long-term data set from southern Italy, we show that long-distance migrants also pass through the Mediterranean region earlier. We argue that this may reflect a climate-driven evolutionary change in the timing of spring migration.

Mycobacterium microti infection (vole tuberculosis) in wild rodent populations.

Mycobacterium microti (vole tuberculosis) infections in small wild mammals were first described more than 60 years ago in several populations in Great Britain. Few studies of vole tuberculosis have been undertaken since then, and little is known about the relationship between M. microti isolates originating from different populations or at different times or of the prevalence of this infection in wild rodent populations, despite human cases of M. microti infections being increasingly reported. In this study, field voles (Microtus agrestis), bank voles (Clethrionomys glareolus), and wood mice (Apodemus sylvaticus) were found to be infected, with up to 8% having external tuberculous signs, in wild populations in Northumberland and Cheshire, England. Spoligotyping applied directly to the clinical material simultaneously detected and typed M. microti bacteria in skin lesions, lymph glands, and internal abcesses. IS6110 restriction fragment length polymorphism typing of cultured bacteria was used to compare these isolates with previously isolated strains from both animals and humans. This demonstrated that although the current rodent isolates were distinct from those isolated from voles in the 1930s in Great Britain, they had a high degree of similarity to these strains and were distinct from the M. microti isolates from humans, a pig, and a ferret from The Netherlands. Thus, M. microti infection seems to be widespread in wild rodent populations, but more studies are needed to understand how M. microti might be transmitted from animals to humans and to determine better the zoonotic risk posed.