Showing papers in "Journal of Animal Ecology in 2009"

Ecological networks--beyond food webs.

Abstract: Summary 1. A fundamental goal of ecological network research is to understand how the complexity observed in nature can persist and how this affects ecosystem functioning. This is essential for us to be able to predict, and eventually mitigate, the consequences of increasing environmental perturbations such as habitat loss, climate change, and invasions of exotic species. 2. Ecological networks can be subdivided into three broad types: ‘traditional’ food webs, mutualistic networks and host‐parasitoid networks. There is a recent trend towards cross-comparisons among network types and also to take a more mechanistic, as opposed to phenomenological, perspective. For example, analysis of network configurations, such as compartments, allows us to explore the role of co-evolution in structuring mutualistic networks and host‐parasitoid networks, and of body size in food webs. 3. Research into ecological networks has recently undergone a renaissance, leading to the production of a new catalogue of evermore complete, taxonomically resolved, and quantitative data. Novel topological patterns have been unearthed and it is increasingly evident that it is the distribution of interaction strengths and the configuration of complexity, rather than just its magnitude, that governs network stability and structure. 4. Another significant advance is the growing recognition of the importance of individual traits and behaviour: interactions, after all, occur between individuals. The new generation of high-quality networks is now enabling us to move away from describing networks based on species-averaged data and to start exploring patterns based on individuals. Such refinements will enable us to address more general ecological questions relating to foraging theory and the recent metabolic theory of ecology. 5. We conclude by suggesting a number of ‘dead ends’ and ‘fruitful avenues’ for future research into ecological networks.

Cascading top-down effects of changing oceanic predator abundances

Abstract: Summary 1. Top-down control can be an important determinant of ecosystem structure and function, but in oceanic ecosystems, where cascading effects of predator depletions, recoveries, and invasions could be significant, such effects had rarely been demonstrated until recently. 2. Here we synthesize the evidence for oceanic top-down control that has emerged over the last decade, focusing on large, high trophic-level predators inhabiting continental shelves, seas, and the open ocean. 3. In these ecosystems, where controlled manipulations are largely infeasible, ‘pseudo-experimental’ analyses of predator‐prey interactions that treat independent predator populations as ‘replicates’, and temporal or spatial contrasts in predator populations and climate as ‘treatments’, are increasingly employed to help disentangle predator effects from environmental variation and noise. 4. Substantial reductions in marine mammals, sharks, and piscivorous fishes have led to mesopredator and invertebrate predator increases. Conversely, abundant oceanic predators have suppressed prey abundances. Predation has also inhibited recovery of depleted species, sometimes through predator‐prey role reversals. Trophic cascades have been initiated by oceanic predators linking to neritic food webs, but seem inconsistent in the pelagic realm with effects often attenuating at plankton. 5. Top-down control is not uniformly strong in the ocean, and appears contingent on the intensity and nature of perturbations to predator abundances. Predator diversity may dampen cascading effects except where nonselective fisheries deplete entire predator functional groups. In other cases, simultaneous exploitation of predator and prey can inhibit prey responses. Explicit consideration of anthropogenic modifications to oceanic foodwebs should help inform predictions about trophic control. 6. Synthesis and applications . Oceanic top-down control can have important socio-economic, conservation, and management implications as mesopredators and invertebrates assume dominance, and recovery of overexploited predators is impaired. Continued research aimed at integrating across trophic levels is needed to understand and forecast the ecosystem effects of changing oceanic predator abundances, the relative strength of top-down and bottom-up control, and interactions with intensifying anthropogenic stressors such as climate change.

Climate change and unequal phenological changes across four trophic levels: constraints or adaptations?

Abstract: 1. Climate change has been shown to affect the phenology of many organisms, but interestingly these shifts are often unequal across trophic levels, causing a mismatch between the phenology of organisms and their food. 2. We consider two alternative hypotheses: consumers are constrained to adjust sufficiently to the lower trophic level, or prey species react more strongly than their predators to reduce predation. We discuss both hypotheses with our analyses of changes in phenology across four trophic levels: tree budburst, peak biomass of herbivorous caterpillars, breeding phenology of four insectivorous bird species and an avian predator. 3. In our long-term study, we show that between 1988 and 2005, budburst advanced (not significantly) with 0.17 d yr(-1), while between 1985 and 2005 both caterpillars (0.75 d year(-1)) and the hatching date of the passerine species (range for four species: 0.36-0.50 d year(-1)) have advanced, whereas raptor hatching dates showed no trend. 4. The caterpillar peak date was closely correlated with budburst date, as were the passerine hatching dates with the peak caterpillar biomass date. In all these cases, however, the slopes were significantly less than unity, showing that the response of the consumers is weaker than that of their food. This was also true for the avian predator, for which hatching dates were not correlated with the peak availability of fledgling passerines. As a result, the match between food demand and availability deteriorated over time for both the passerines and the avian predators. 5. These results could equally well be explained by consumers' insufficient responses as a consequence of constraints in adapting to climate change, or by them trying to escape predation from a higher trophic level, or both. Selection on phenology could thus be both from matches of phenology with higher and lower levels, and quantifying these can shed new light on why some organisms do adjust their phenology to climate change, while others do not.

The sensitive hare: sublethal effects of predator stress on reproduction in snowshoe hares.

Abstract: Summary 1. Prey responses to high predation risk can be morphological or behavioural and ultimately come at the cost of survival, growth, body condition, or reproduction. These sub-lethal predator effects have been shown to be mediated by physiological stress. We tested the hypothesis that elevated glucocorticoid concentrations directly cause a decline in reproduction in individual free-ranging female snowshoe hares, Lepus americanus. We measured the cortisol concentration from each dam (using a faecal analysis enzyme immunoassay) and her reproductive output (litter size, offspring birth mass, offspring right hind foot (RHF) length) 30 h after birth. 2. In a natural monitoring study, we monitored hares during the first and second litter from the population peak (2006) to the second year of the decline (2008). We found that faecal cortisol metabolite (FCM) concentration in dams decreased 52% from the first to the second litter. From the first to the second litter, litter size increased 122%, offspring body mass increased 130%, and offspring RHF length increased 112%. Dam FCM concentrations were inversely related to litter size (r 2 ¼ 0AE19), to offspring birth mass (r 2 ¼ 0AE32), and to offspring RHF length (r 2 ¼ 0AE64). 3. In an experimental manipulation, we assigned wild-caught, pregnant hares to a control and a stressed group and held them in pens. Hares in the stressed group were exposed to a dog 1–2 min every other day before parturition to simulate high predation risk. At parturition, unsuccessfulstressed dams (those that failed to give birth to live young) and stressed dams had 837% and 214%, respectively, higher FCM concentrations than control dams. Of those females that gave birth, litter size was similar between control and stressed dams. However, offspring from stressed dams were 37% lighter and 16% smaller than offspring from control dams. Increasing FCM concentration in dams caused the decline of offspring body mass (r 2 ¼ 0AE57) and RHF (r 2 ¼ 0AE52). 4. This is the first study in a free-ranging population of mammals to show that elevated, predatorinduced, glucocorticoid concentrations in individual dams caused a decline in their reproductive output measured both by number and quality of offspring. Thus, we provide evidence that any stressor, not just predation, which increases glucocorticoid concentrations will result in a decrease in reproductive output.

Physical defences wear you down: progressive and irreversible impacts of silica on insect herbivores

Abstract: 1. Silica in the leaves of grasses can act as a defence against both vertebrate and invertebrate herbivores. The mechanisms by which silica affects herbivore performance are not well characterized. Here we expose an insect herbivore Spodoptera exempta to high-silica diets and test two mechanisms by which silica has been proposed to act as a defence. First, that silica reduces the digestibility of leaves and second, that silica causes wear to insect mandibles, both of which could potentially impact on herbivore performance. 2. Silica reduced the efficiency with which S. exempta converted ingested food to body mass and the amount of nitrogen absorbed from their food, leading to reduced insect growth rates. The measure of how efficiently herbivores utilize digested food (ECD) was unaffected by silica. 3. These effects occurred even with short-term exposure to silica-rich diets, but they also increased markedly with the duration of exposure and affected late instars more than early instar larvae. This appears to be due to the progressive impacts of silica with longer exposure times and suggests that herbivores cannot adapt to silica defences, nor do they develop a tolerance for silica with age. 4. Exposure to silica-rich diets caused increased mandible wear in S. exempta. This effect was extremely rapid, occurring within a single instar, further reducing feeding efficiency and growth rates. These effects on insect growth and feeding efficiency are nonreversible, persisting after the herbivore has switched diets. Up to a third of this residual impact can be explained by the degree of mandible wear caused by previous silica-rich diets. 5. The impacts of silica on S. exempta larvae were progressive with exposure time and could not be compensated for, even by switching to a different diet. Thus, herbivores cannot easily adapt to physical defences such as silica, suggesting this defence will have major implications for herbivore fitness.

Undersampling bias: the null hypothesis for singleton species in tropical arthropod surveys

Abstract: Summary 1. Frequency of singletons ‐ species represented by single individuals ‐ is anomalously high in most large tropical arthropod surveys (average, 32%). 2. We sampled 5965 adult spiders of 352 species (29% singletons) from 1 ha of lowland tropical moist forest in Guyana. 3. Four common hypotheses (small body size, male-biased sex ratio, cryptic habits, clumped distributions) failed to explain singleton frequency. Singletons are larger than other species, not gender-biased, share no particular lifestyle, and are not clumped at 0·25‐1 ha scales. 4. Monte Carlo simulation of the best-fit lognormal community shows that the observed data fit a random sample from a community of ~700 species and 1‐2 million individuals, implying approximately 4% true singleton frequency. 5. Undersampling causes systematic negative bias of species richness, and should be the default null hypothesis for singleton frequencies. 6. Drastically greater sampling intensity in tropical arthropod inventory studies is required to yield realistic species richness estimates. 7. The lognormal distribution deserves greater consideration as a richness estimator when undersampling bias is severe.

Classifying movement behaviour in relation to environmental conditions using hidden Markov models.

Abstract: 1. Linking the movement and behaviour of animals to their environment is a central problem in ecology. Through the use of electronic tagging and tracking (ETT), collection of in situ data from free-roaming animals is now commonplace, yet statistical approaches enabling direct relation of movement observations to environmental conditions are still in development. 2. In this study, we examine the hidden Markov model (HMM) for behavioural analysis of tracking data. HMMs allow for prediction of latent behavioural states while directly accounting for the serial dependence prevalent in ETT data. Updating the probability of behavioural switches with tag or remote-sensing data provides a statistical method that links environmental data to behaviour in a direct and integrated manner. 3. It is important to assess the reliability of state categorization over the range of time-series lengths typically collected from field instruments and when movement behaviours are similar between movement states. Simulation with varying lengths of times series data and contrast between average movements within each state was used to test the HMMs ability to estimate movement parameters. 4. To demonstrate the methods in a realistic setting, the HMMs were used to categorize resident and migratory phases and the relationship between movement behaviour and ocean temperature using electronic tagging data from southern bluefin tuna (Thunnus maccoyii). Diagnostic tools to evaluate the suitability of different models and inferential methods for investigating differences in behaviour between individuals are also demonstrated.

Assessing the accuracy of species distribution models to predict amphibian species richness patterns

Abstract: Summary 1. Evaluating the distribution of species richness where biodiversity is high but has been insufficiently sampled is not an easy task. Species distribution modelling has become a useful approach for predicting their ranges, based on the relationships between species records and environmental variables. Overlapping predictions of individual distributions could be a useful strategy for obtaining estimates of species richness and composition in a region, but these estimates should be evaluated using a proper validation process, which compares the predicted richness values and composition with accurate data from independent sources. 2. In this study, we propose a simple approach to estimate model performance for several distributional predictions generated simultaneously. This approach is particularly suitable when species distribution modelling techniques that require only presence data are used. 3. The individual distributions for the 370 known amphibian species of Mexico were predicted using maxent to model data on their known presence (66 113 presence-only records). Distributions were subsequently overlapped to obtain a prediction of species richness. Accuracy was assessed by comparing the overall species richness values predicted for the region with observed and predicted values from 118 well-surveyed sites, each with an area of c. 100 km 2 , which were identified using species accumulation curves and nonparametric estimators. 4. The derived models revealed a remarkable heterogeneity of species richness across the country, provided information about species composition per site and allowed us to obtain a measure of the spatial distribution of prediction errors. Examining the magnitude and location of model inaccuracies, as well as separately assessing errors of both commission and omission, highlights the inaccuracy of the predictions of species distribution models and the need to provide measures of uncertainty along with the model results. 5. The combination of a species distribution modelling method like maxent and species richness estimators offers a useful tool for identifying when the overall pattern provided by all model predictions might be representing the geographical patterns of species richness and composition, regardless of the particular quality or accuracy of the predictions for each individual species.

Individual variation in reproductive costs of reproduction: high‐quality females always do better

Abstract: 1. Although life-history theory predicts substantial costs of reproduction, individuals often show positive correlations among life-history traits, rather than trade-offs. The apparent absence of reproductive costs may result from heterogeneity in individual quality. 2. Using detailed longitudinal data from three contrasted ungulate populations (mountain goats, Oreamnos americanus; bighorn sheep, Ovis canadensis; and roe deer, Capreolus capreolus), we assessed how individual quality affects the probability of detecting a cost of current reproduction on future reproduction for females. We used a composite measure of individual quality based on variations in longevity (all species), success in the last breeding opportunity before death (goats and sheep), adult mass (all species), and social rank (goats only). 3. In all species, high-quality females consistently had a higher probability of reproduction, irrespective of previous reproductive status. In mountain goats, we detected a cost of reproduction only after accounting for differences in individual quality. Only low-quality female goats were less likely to reproduce following years of breeding than of nonbreeding. Offspring survival was lower in bighorn ewes following years of successful breeding than after years when no lamb was produced, but only for low-quality females, suggesting that a cost of reproduction only occurred for low-quality females. 4. Because costs of reproduction differ among females, studies of life-history evolution must account for heterogeneity in individual quality.

Towards a predictive framework for predator risk effects: the interaction of landscape features and prey escape tactics

Abstract: 1. Risk effects of predators can profoundly affect community dynamics, but the nature of these effects is context dependent. 2. Although context dependence has hindered the development of a general framework for predicting the nature and extent of risk effects, recent studies suggest that such a framework is attainable if the factors that shape anti-predator behaviour, and its effectiveness, in natural communities are well understood. 3. One of these factors, the interaction of prey escape tactics and landscape features, has been largely overlooked. 4. We tested whether this interaction gives rise to interspecific variation in habitat-use patterns of sympatric large marine vertebrates at risk of tiger shark (Galeocerdo cuvier Peron and LeSueur, 1822) predation. Specifically, we tested the a priori hypothesis that pied cormorants (Phalacrocorax varius Gmelin, 1789) would modify their use of shallow seagrass habitats in a manner opposite to that of previously studied dolphins (Tursiops aduncus Ehrenberg, 1833), dugongs (Dugong dugon Muller, 1776), and green turtles (Chelonia mydas Linnaeus, 1758) because, unlike these species, the effectiveness of cormorant escape behaviour does not vary spatially. 5. As predicted, cormorants used interior and edge portions of banks proportional to the abundance of their potential prey when sharks were absent but shifted to interior portions of banks to minimize encounters with tiger sharks as predation risk increased. Other shark prey, however, shift to edge microhabitats when shark densities increase to take advantage of easier escape despite higher encounter rates with sharks. 6. The interaction of landscape features and escape ability likely is important in diverse communities. 7. When escape probabilities are high in habitats with high predator density, risk effects of predators can reverse the direction of commonly assumed indirect effects of top predators. 8. The interaction between landscape features and prey escape tactics can result in a single predator species having differential effects on their sympatric prey that could cascade through ecosystems and should be incorporated into a general framework for context dependence of risk effects.

Can the protein costs of bacterial resistance be offset by altered feeding behaviour

Abstract: 1. Mounting an immune response is likely to be costly in terms of energy and nutrients, and so it is predicted that dietary intake should change in response to infection to offset these costs. The present study focuses on the interactions between a specialist grass-feeding caterpillar species, the African armyworm Spodoptera exempta, and an opportunist bacterium, Bacillus subtilis. 2. The main aims of the study were (i) to establish the macronutrient costs to the insect host of surviving a systemic bacterial infection, (ii) to determine the relative importance of dietary protein and carbohydrate to immune system functions, and (iii) to determine whether there is an adaptive change in the host's normal feeding behaviour in response to bacterial challenge, such that the nutritional costs of resisting infection are offset. 3. We show that the survival of bacterially infected larvae increased with increasing dietary protein-to-carbohydrate (P:C) ratio, suggesting a protein cost associated with bacterial resistance. As dietary protein levels increased, there was an increase in antibacterial activity, phenoloxidase (PO) activity and protein levels in the haemolymph, providing a potential source for this protein cost. However, there was also evidence for a physiological trade-off between antibacterial activity and phenoloxidase activity, as larvae whose antibacterial activity levels were elevated in response to immune activation had reduced PO activity. 4. When given a choice between two diets varying in their P:C ratios, larvae injected with a sub-lethal dose of bacteria increased their protein intake relative to control larvae whilst maintaining similar carbohydrate intake levels. These results are consistent with the notion that S. exempta larvae alter their feeding behaviour in response to bacterial infection in a manner that is likely to enhance the levels of protein available for producing the immune system components and other factors required to resist bacterial infections ('self-medication').

How does abundance scale with body size in coupled size-structured food webs?

Abstract: 1. Widely observed macro-ecological patterns in log abundance vs. log body mass of organisms can be explained by simple scaling theory based on food (energy) availability across a spectrum of body sizes. The theory predicts that when food availability falls with body size (as in most aquatic food webs where larger predators eat smaller prey), the scaling between log N vs. log m is steeper than when organisms of different sizes compete for a shared unstructured resource (e.g. autotrophs, herbivores and detritivores; hereafter dubbed 'detritivores'). 2. In real communities, the mix of feeding characteristics gives rise to complex food webs. Such complexities make empirical tests of scaling predictions prone to error if: (i) the data are not disaggregated in accordance with the assumptions of the theory being tested, or (ii) the theory does not account for all of the trophic interactions within and across the communities sampled. 3. We disaggregated whole community data collected in the North Sea into predator and detritivore components and report slopes of log abundance vs. log body mass relationships. Observed slopes for fish and epifaunal predator communities (-1.2 to -2.25) were significantly steeper than those for infaunal detritivore communities (-0.56 to -0.87). 4. We present a model describing the dynamics of coupled size spectra, to explain how coupling of predator and detritivore communities affects the scaling of log N vs. log m. The model captures the trophic interactions and recycling of material that occur in many aquatic ecosystems. 5. Our simulations demonstrate that the biological processes underlying growth and mortality in the two distinct size spectra lead to patterns consistent with data. Slopes of log N vs. log m were steeper and growth rates faster for predators compared to detritivores. Size spectra were truncated when primary production was too low for predators and when detritivores experienced predation pressure. 6. The approach also allows us to assess the effects of external sources of mortality (e.g. harvesting). Removal of large predators resulted in steeper predator spectra and increases in their prey (small fish and detritivores). The model predictions are remarkably consistent with observed patterns of exploited ecosystems.

Carry‐over effects in a Pacific seabird: stable isotope evidence that pre‐breeding diet quality influences reproductive success

Abstract: 1. Understanding the interactions between different periods of the annual cycle in migratory animals has been constrained by our inability to track individuals across seasons. In seabirds, virtually nothing is known about how diet quality during the non-breeding period, away from the breeding grounds, might influence subsequent reproductive success. 2. We used stable nitrogen (delta(15)N) and carbon (delta(13)C) isotopes to evaluate the effects of non-breeding diet quality on the timing of breeding and egg size in a population of Cassin's auklets (Ptychoramphus aleuticus) breeding on Triangle Island, British Columbia. Adult feathers are grown during two different periods of the annual cycle, which allowed us to estimate diet quality from the previous fall (October-November) and pre-breeding (February-March) period. 3. We found that the estimated proportion of energetically superior copepods (Neocalanus spp.) in the pre-breeding diet tended to be higher in females that bred earlier and laid larger eggs, whereas energetically poor juvenile rockfish (Sebastes spp.) were dominant in the pre-breeding diets of females that bred later and laid smaller eggs. We detected no effect of fall diet quality on breeding date or egg size, and no effect of pre-breeding diet quality on breeding date in males. 4. Pre-breeding diet quality was not related to body condition measured 1-2 days after laying, which suggests that females may need to attain a threshold condition before they initiate breeding and successfully rear young. 5. Our results suggest that changes in climatic conditions during the pre-breeding period may have severe consequences for reproductive success by influencing breeding date and egg size. Our work emphasizes the importance of determining how events are linked throughout the annual cycle for understanding the fitness and population dynamics of migratory animals.

Taxonomical vs. functional responses of bee communities to fire in two contrasting climatic regions.

Abstract: 1. Valuable insights into mechanisms of community responses to environmental change can be gained by analysing in tandem the variation in functional and taxonomic composition along environmental gradients. 2. We assess the changes in species and functional trait composition (i.e. dominant traits and functional diversity) of diverse bee communities in contrasting fire-driven systems in two climatic regions: Mediterranean (scrub habitats in Israel) and temperate (chestnut forests in southern Switzerland). 3. In both climatic regions, there were shifts in species diversity and composition related to post-fire age. In the temperate region, functional composition responded markedly to fire; however, in the Mediterranean, the taxonomic response to fire was not matched by functional replacement. 4. These results suggest that greater functional stability to fire in the Mediterranean is achieved by replacement of functionally similar species (i.e. functional redundancy) which dominate under different environmental conditions in the heterogeneous landscapes of the region. In contrast, the greater functional response in the temperate region was attributed to a more rapid post-fire vegetation recovery and shorter time-window when favourable habitat was available relative to the Mediterranean. 5. Bee traits can be used to predict the functional responses of bee communities to environmental changes in habitats of conservation importance in different regions with distinct disturbance regimes. However, predictions cannot be generalized from one climatic region to another where distinct habitat configurations occur.

Heterogeneous selection on a heritable temperament trait in a variable environment.

Abstract: 1. Temperament traits increasingly provide a focus for investigating the evolutionary ecology of behavioural variation. Here, we examine the underlying causes and selective consequences of individual variation in the temperament trait 'exploration behaviour in a novel environment' (EB, based on an 8-min assay) in a free-ranging population of a passerine bird, the great tit Parus major. 2. First, we conducted a quantitative genetic analysis on EB using a restricted maximum likelihood-based animal model with a long-term pedigree. Although repeatability was relatively high, EB was only moderately heritable and permanent environment (V(PE)) effects contributed as much to phenotypic variance as additive genetic effects. 3. We then asked whether heterogeneous selection acted on EB at various temporal and spatial scales. Using estimates of lifetime reproductive success, we found evidence of weak negative directional selection acting on EB amongst females which was driven by selection through recruitment, but not fecundity, in one of the four breeding years. There was no evidence of any selection on EB through survival. 4. Heterogeneous selection on EB within seasons was also observed amongst males through fecundity along two fine-scale environmental gradients--local breeding density and habitat quality; we are unaware of any previous equivalent demonstrations. 5. All of these analyses were repeated on a second measure of exploration behaviour (EB(2), measured during a 2-min assay) to facilitate comparison with other studies. EB and EB(2) were strongly correlated to one another at the genetic level, but were only moderately correlated at the phenotypic level and V(PE) was undetected in EB(2). Selection on EB(2) was similar to that on EB; we conclude that both traits are broadly equivalent from an evolutionary perspective. 6. Our analyses suggest that to the extent that the temperament trait 'exploration behaviour' is subject to natural selection in this population, this selection is highly context dependent and most evident along two environmental gradients. Furthermore, the strong V(PE) effect detected suggests that understanding the causes and consequences of variation in this trait will require studies firmly embedded in an environmental context.

Beta diversity along environmental gradients: implications of habitat specialization in tropical montane landscapes.

Abstract: 1. Understanding how species in a diverse regional pool are spatially distributed with respect to habitat types is a longstanding problem in ecology. Tropical species are expected to be specialists along environmental gradients, and this should result in rapid compositional change (high beta diversity) across landscapes, particularly when alpha diversity is a small fraction of regional diversity. Corollary challenges are then to identify controlling environmental variables and to ask whether species cluster into discrete community types along a gradient. 2. We investigated patterns of avian species' distributions in the Tilaran mountains of Costa Rica between 1000 m and 1700 m elevation where a strong moisture gradient exists. High beta diversity was found with both auditory counts adjusted for detectability and extensive capture data, revealing nearly complete change in community composition over a few kilometres on the Pacific slope. As predicted, this beta diversity was roughly twice as high as on temperate mountainsides. 3. Partial Mantel analyses and canonical correspondence analysis indicate that change in species composition is highly correlated with change in moisture (and correlated epiphyte cover) at different distances from the continental divide on the Pacific slope. Altitude was not a good predictor of change in species composition, as species composition varies substantially among sites at the same elevation. 4. Detrended correspondence analysis and cluster analysis revealed a zone of rapid transition separating a distinct cloud forest community from rainshadow forest. On the Caribbean slope, where a shallower moisture gradient was predicted to result in lower beta diversity, we found lower rates of compositional change and more continuous species turnover. 5. Results suggest that habitat specialization of birds is likely a strong ecological force generating high beta diversity in montane landscapes. Despite overall rapid rates of species turnover, zones of relatively coherent composition could be identified. 6. Landscapes with such high beta diversity are common in the tropics, although little studied. They offer high benefit/cost opportunities for conservation, particularly as climate change threatens to alter the species composition of communities of habitat specialists.

Linking movement behaviour, dispersal and population processes: is individual variation a key?

Abstract: Summary 1. Movement behaviour has become increasingly important in dispersal ecology and dispersal is central to the development of spatially explicit population ecology. The ways in which the elements have been brought together are reviewed with particular emphasis on dispersal distance distributions and the value of mechanistic models. 2. There is a continuous range of movement behaviours and in some species, dispersal is a clearly delineated event but not in others. The biological complexities restrict conclusions to high-level generalizations but there may be principles that are common to dispersal and other movements. 3. Random walk and diffusion models when appropriately elaborated can provide an understanding of dispersal distance relationships on spatial and temporal scales relevant to dispersal. Leptokurtosis in the relationships may be the result of a combination of factors including population heterogeneity, correlation, landscape features, time integration and density dependence. The inclusion in diffusion models of individual variation appears to be a useful elaboration. The limitations of the negative exponential and other phenomenological models are discussed. 4. The dynamics of metapopulation models are sensitive to what appears to be small differences in the assumptions about dispersal. In order to represent dispersal realistically in population models, it is suggested that phenomenological models should be replaced by those based on movement behaviour incorporating individual variation. 5. The conclusions are presented as a set of candidate principles for evaluation. The main features of the principles are that uncorrelated or correlated random walk, not linear movement, is expected where the directions of habitat patches are unpredictable and more complex behaviour when organisms have the ability to orientate or navigate. Individuals within populations vary in their movement behaviour and dispersal; part of this variation is a product of random elements in movement behaviour and some of it is heritable. Local and metapopulation dynamics are influenced by population heterogeneity in dispersal characteristics and heritable changes in dispersal propensity occur on time-scales short enough to impact population dynamics.

Landscape heterogeneity and marine subsidy generate extensive intrapopulation niche diversity in a large terrestrial vertebrate

Abstract: Summary 1. Inquiries into niche variation within populations typically focus on proximate ecological causes such as competition. Here we examine how landscape heterogeneity and allochthonous (marine) subsidy might ultimately generate intrapopulation niche diversity. 2. Using stable isotope analysis, we detected extensive terrestrial‐marine isotopic niche variation among subpopulations, social groups, and individual grey wolves ( Canis lupus ) that occupy a spatially heterogeneous landscape in coastal British Columbia comprising a mainland area and adjacent archipelago. 3. The inner island subpopulation exhibited the widest isotopic niche in the population, consuming extensive terrestrial and marine resources. Mainland and outer island subpopulations occupied comparatively narrow and primarily terrestrial, and primarily marine, niches respectively. Within these biogeographical subpopulations, social groups also diverged in niche. 4. To support examination at the individual level, we used an isotopic approach to test Van Valen’s (1965) niche variation hypothesis. Consistent with the hypothesis, we observed that among-individual variation increased with subpopulation niche width. 5. Patterns at all levels related to how a spatially heterogeneous coastal landscape structured the competitive environment, which in turn mediated the availability and use of terrestrial and marine resources. Broadly, our results suggest that spatial heterogeneity and allochthonous subsidy ‐ both widespread but commonly subject to contemporary anthropogenic change ‐ might provide novel opportunities for examination and conservation of ecological variation within populations.

Food web topology and parasites in the pelagic zone of a subarctic lake

Abstract: Summary 1. Parasites permeate trophic webs with their often complex life cycles, but few studies have included parasitism in food web analyses. Here we provide a highly resolved food web from the pelagic zone of a subarctic lake and explore how the incorporation of parasites alters the topology of the web. 2. Parasites used hosts at all trophic levels and increased both food-chain lengths and the total number of trophic levels. Their inclusion in the network analyses more than doubled the number of links and resulted in an increase in important food-web characteristics such as linkage density and connectance. 3. More than half of the parasite taxa were trophically transmitted, exploiting hosts at multiple trophic levels and thus increasing the degree of omnivory in the trophic web. 4. For trophically transmitted parasites, the number of parasite‐host links exhibited a positive correlation with the linkage density of the host species, whereas no such relationship was seen for nontrophically transmitted parasites. Our findings suggest that the linkage density of free-living species affects their exposure to trophically transmitted parasites, which may be more likely to adopt highly connected species as hosts during the evolution of complex life cycles. 5. The study supports a prominent role for parasites in ecological networks and demonstrates that their incorporation may substantially alter considerations of food-web structure and functioning.

Breeding in high-elevation habitat results in shift to slower life-history strategy within a single species.

Abstract: Summary 1 Elevational gradients create environmental variation that is hypothesized to promote variation in life-history strategies. We tested whether differences in life-history strategies were associated with elevation in a songbird, the dark-eyed junco (Junco hyemalis; Aves; A.O.U. 1998). 2 We monitored birds in four replicated sites per elevation, at 2000 m a.s.l. (high elevation) and 1000 m a.s.l. (low elevation), in the Rocky Mountains of Canada. 3 Over 5 years, we measured the following traits and vital rates: egg-laying schedules, morphological indicators of reproductive stage, seasonal reproductive success, indicators of competitive class (age, size, arrival time), and survival rates. 4 We found two main patterns: with an increase in breeding elevation, dark-eyed juncos delayed the development of structures necessary for reproduction (e.g. cloacal protuberance in males) and reduced the duration of their reproductive period to less than half of the time used by low-elevation birds; and 5 Juncos at high-elevation sites had 55–61% lower annual reproductive success and 15 to 20% higher survival rates. While adult juncos at high elevations produced fewer offspring, those offspring were in better condition. Proportions of age and size classes in high- compared to low-elevation populations were similar, suggesting that a life-history trade-off is present, rather than competition forcing inferior competitors to breed in a peripheral habitat. The apparent trade-off between reproduction and survival corresponded to a shorter period of favourable weather and available food in high- compared to low-elevation habitats. 6 Thus, elevation had a strong influence on life-history characteristics of a single species over a short spatial distance, suggesting a shift in life history from a high reproductive strategy at lower elevations to a high survivor strategy at high elevations. 7 This is the first paper to show a shift in avian life-history strategies along an elevational gradient (in both genders, of multiple age classes) when region (latitude) and phylogenetic histories are controlled for.

Microhabitat selection by sea turtles in a dynamic thermal marine environment

Abstract: 1. Reproductive fitness is often compromised at the margins of a species' range due to sub-optimal conditions. 2. Set against this backdrop, the Mediterranean's largest loggerhead sea turtle (Caretta caretta) rookery at Zakynthos (Greece) presents a conundrum, being at a very high latitude for this species, yet hosting a high concentration of nesting. 3. We used visual surveys combined with global positioning system (GPS) tracking to show that at the start of the breeding season, individuals showed microhabitat selection, with females residing in transient patches of warm water. As the sea warmed in the summer, this selection was no longer evident. 4. As loggerhead turtles are ectothermic, this early season warm-water selection presumably speeds up egg maturation rates before oviposition, thereby allowing more clutches to be incubated when sand conditions are optimal during the summer. 5. Active selection of warm waters may allow turtles to initiate nesting at an earlier date.

Effects of individual quality, reproductive success and environmental variability on survival of a long-lived seabird.

Abstract: Summary 1. Heterogeneity in individual quality (i.e. individuals having different performance levels that are consistent throughout life) can drive the demography of iteroparous species, but quality in the context of environmental variability has rarely been evaluated. 2. We investigated the demographic responses of a long-lived seabird, the Adelie penguin ( Pygoscelis adeliae ), to contrasting environmental conditions as a function of reproductive success, breeding quality (BQ) and experience. A continuous index of BQ (BQI) was developed to reflect an individual’s ability, relative to others, to produce viable offspring. 3. First, we assessed the relative importance of costs of reproduction vs. heterogeneity in quality by comparing survival and reproductive probabilities among deferred, successful and unsuccessful breeders under ‘demanding’ conditions using multistate capture‐mark‐recapture modelling. Then, we quantified the influence of BQI on adult survival among experienced breeders vs. the whole study population under both ‘normal’ and ‘demanding’ conditions. 4. Higher survival rates were exhibited by successful (74‐76%) compared to unsuccessful breeders (64%); the former also more frequently reproduced successfully at year t + 1. 5. From 1997 to 2006, adult survival ranged from 64‐79%, with BQI accounting for 91% of variability in the entire study population, but only 17% in experienced breeders. The weakened relationship between BQI and survival in experienced breeders supports the theory that selection during the first reproductive event accounts for a more homogeneous pool of experienced breeders. 6. No significant effect of environmental covariates on survival was evident, suggesting that what appeared to be demanding conditions were within the range that could be buffered by this species. 7. For the first time in seabirds, a quadratic relationship between adult survival and BQI showed that adult survival is shaped by both heterogeneity in quality and reproductive costs. Our study confirms that population demographic patterns are affected by factors at the individuals’ level (e.g., individual quality) that are obscured at population-scale levels.

The magnitude and selectivity of natural and multiple anthropogenic mortality causes in hunted brown bears

Abstract: Summary 1. The population dynamic and evolutionary effects of harvesting are receiving growing attention among biologists. Cause-specific estimates of mortality are necessary to determine and compare the magnitude and selectivity of hunting and other types of mortalities. In addition to the logistic and financial constraints on longitudinal studies, they are complicated by the fact that nonhunting mortality in managed populations usually consists of a mix of natural and human-caused factors. 2. We used multistate capture‐recapture (MCR) models to estimate cause-specific survival of brown bears ( Ursus arctos ) in two subpopulations in Sweden over a 23-year period. In our analysis, we distinguished between legal hunting and other sources of mortality, such as intraspecific predation, accidents, poaching, and damage control removals. We also tested whether a strong increase in harvest quotas after 1997 in one of the subpopulations affected vulnerability to legal hunting. 3. Although only a fraction of mortalities other than legal hunting could be considered natural, this group of causes showed a general pattern of demographic selectivity expected from natural mortality regimes in populations of long-lived species, namely greater vulnerability of young animals. On the other hand, demographic effects on hunting vulnerability were weak and inconsistent. Our findings support the assumption that hunting and other mortalities were additive. 4. As expected, an increase in hunting pressure coincided with a correspondingly large increase in vulnerability to hunting in the affected subpopulation. Because even unbiased harvest can lead to selective pressures on life-history traits, such as size at primiparity, increasing harvest quotas may not only affect population growth directly, but could also alter optimal life-history strategies in brown bears and other carnivores. 5. Legal hunting is the most conveniently assessed and the most easily managed cause of mortality in many wild populations of large mammals. Although legal hunting is the single-most important cause of mortality for brown bears in Sweden, the combined mortality from other causes is of considerable magnitude and additionally shows greater selectivity in terms of sex and age than legal hunting. Therefore, its role in population dynamics and evolution should not be underestimated.

A slow life in hell or a fast life in heaven: demographic analyses of contrasting roe deer populations.

Abstract: 1. Environmental conditions shape population growth through their impact on demographic parameters. While knowledge has accumulated concerning the effects of population density and climatic conditions, a topical question now concerns how predation and harvest influence demographic parameters and population growth (lambda). 2. We performed a comparative demographic analysis based on projection matrix models for female roe deer. Population-specific matrices were parameterized based on longitudinal data from five intensively monitored populations in Norway and France, spanning a large variability in environmental characteristics such as densities of large predators, hunter harvest and seasonality. 3. As expected for a large iteroparous vertebrate, temporal variation was invariably higher in recruitment than in adult survival, and the elasticity of adult survival was consistently higher than that of recruitment. However, the relative difference in elasticity of lambda to recruitment and adult survival varied strongly across populations, and was closely correlated with adult survival. 4. Different traits accounted for most of the variance in lambda in different ecological settings. Adult survival generally contributed more in populations with low mean adult survival and low mean growth rate during the study period. Hunters and predators (Eurasian lynx and red foxes) occurred in two of our study populations and contributed substantially to the variance in lambda, accounting for a total of 35% and 70% in the two populations respectively. 5. Across populations, we did not find any evidence that roe deer increased their reproductive output when faced with harsh conditions, resulting in some populations having negative growth rates. 6. Generation time, a measure of the speed of the life-history cycle, increased from less than 4 years in the most productive population ('roe deer heaven') to more than 6 years in declining populations facing predation from lynx, red fox and hunters ('roe deer hell'), and was tightly and inversely correlated with lambda. Such a deceleration of the life cycle in declining populations might be a general feature in large herbivores. 7. Our results shows that the plethora of environmental conditions faced by populations of large herbivores also induce high intraspecific variation in their ranking along the 'fast-slow' continuum of life-history tactics.

Timing is everything: flexible phenology and shifting selection in a colonial seabird

Abstract: 1. In order to reproduce successfully in a temporally varying environment, iteroparous animals must exhibit considerable behavioural flexibility across their lifetimes. By adjusting timing of breeding each year, parents can ensure optimal overlap between the energy intensive period of offspring production and the seasonal peak in favourable environmental conditions, thereby increasing their chances of successfully rearing young. 2. Few studies investigate variation among individuals in how they respond to fluctuating conditions, or how selection acts on these individual differences, but this information is essential for understanding how populations will cope with rapid environmental change. 3. We explored inter-annual trends in breeding time and individual responses to environmental variability in common guillemots Uria aalge, an important marine top predator in the highly variable California Current System. Complex, nonlinear relationships between phenology and oceanic and climate variables were found at the population level. Using a novel application of a statistical technique called random regression, we showed that individual females responded in a nonlinear fashion to environmental variability, and that reaction norm shape differed among females. 4. The pattern and strength of selection varied substantially over a 34-year period, but in general, earlier laying was favoured. Females deviating significantly from the population mean laying date each year also suffered reduced breeding success, with the strength of nonlinear selection varying in relation to environmental conditions. 5. We discuss our results in the wider context of an emerging literature on the evolutionary ecology of individual-level plasticity in the wild. Better understanding of how species-specific factors and local habitat features affect the timing and success of breeding will improve our ability to predict how populations will respond to climate change.

Comparison of social networks derived from ecological data: implications for inferring infectious disease dynamics.

Abstract: Social network analyses tend to focus on human interactions. However, there is a burgeoning interest in applying graph theory to ecological data from animal populations. Here we show how radio-tracking and capture–mark–recapture data collated from wild rodent populations can be used to generate contact networks. Both radio-tracking and capture–mark–recapture were undertaken simultaneously. Contact networks were derived and the following statistics estimated: mean-contact rate, edge distribution, connectance and centrality. Capture–mark–recapture networks produced more informative and complete networks when the rodent density was high and radio-tracking produced more informative networks when the density was low. Different data collection methods provide more data when certain ecological characteristics of the population prevail. Both sets of data produced networks with comparable edge (contact) distributions that were best described by a negative binomial distribution. Connectance and closeness were statistically different between the two data sets. Only betweenness was comparable. The differences between the networks have important consequences for the transmission of infectious diseases. Care should be taken when extrapolating social networks to transmission networks for inferring disease dynamics.

Pulsed resources and climate-induced variation in the reproductive traits of wild boar under high hunting pressure

Abstract: 1. Identifying which factors influence age and size at maturity is crucial for a better understanding of the evolution of life-history strategies. In particular, populations intensively harvested, hunted or fished by humans often respond by displaying earlier age and decreased size at first reproduction. 2. Among ungulates wild boar (Sus scrofa scrofa L.) exhibit uncommon life-history traits, such as high fertility and early reproduction, which might increase the demographic impact of varying age at first reproduction. We analysed variation in female reproductive output from a 22-year long study of an intensively hunted population. We assessed how the breeding probability and the onset of oestrus responded to changes of female body mass at different ages under varying conditions of climate and food availability. 3. Wild boar females had to reach a threshold body mass (27-33 kg) before breeding for the first time. This threshold mass was relatively low (33-41% of adult body mass) compared to that reported in most other ungulates (about 80%). 4. Proportions of females breeding peaked when rainfall and temperature were low in spring and high in summer. Climatic conditions might act through the nutritional condition of females. The onset of oestrus varied a lot in relation to resources available at both current and previous years. Between none and up to 90% of females were in oestrus in November depending on the year. 5. Past and current resources accounted for equivalent amount of observed variations in proportions of females breeding. Thus, wild boar rank at an intermediate position along the capital-income continuum rather than close to the capital end where similar-sized ungulates rank. 6. Juvenile females made a major contribution to the yearly reproductive output. Comparisons among wild boar populations facing contrasted hunting pressures indicate that a high demographic contribution of juveniles is a likely consequence of a high hunting pressure rather than a species-specific life-history pattern characterizing wild boar.

Temperature and life history: experimental heating leads female tree swallows to modulate egg temperature and incubation behaviour

Abstract: Summary 1. Life-history decisions are strongly affected by environmental conditions. In birds, incubation is energetically expensive and affected significantly by ambient temperature. We reduced energetic constraints for female tree swallows ( Tachycineta bicolor ) by experimentally heating nests during incubation by an average of 6·9 ° C to test for changes in incubation behaviour. 2. Females in heated boxes (hereafter ‘heated females’) increased time spent incubating and maintained higher on-bout and off-bout egg temperatures. This indicates that female energetic constraints, not maximizing developmental conditions of offspring, determine incubation investment. Furthermore, this result suggests that embryonic developmental conditions in unmanipulated nests are suboptimal. 3. We found individual variation in how females responded to experimental heating. Early-laying (i.e. higher phenotypic quality) females with heated nests increased egg temperatures and maintained incubation constancy, while later-laying (lower quality) heated females increased incubation constancy. Changes in egg temperature were due to changes in female behaviour and not due directly to increases in internal nest-box temperatures. 4. Behaviour during the incubation period affected hatching asynchrony. Decreased variation in egg temperature led to lower levels of hatching asynchrony, which was also generally lower in heated nests. 5. Our study finds strong support for the prediction that intermittent incubators set their incubation investment at levels dictated by energetic constraints. Furthermore, females incubating in heated boxes allocated conserved energy primarily to increased egg temperature and increased incubation attentiveness. These results indicate that studies investigating the role of energetics in driving reproductive investment in intermittent incubators should consider egg temperature and individual variation more explicitly.

Reproductive interference determines persistence and exclusion in species interactions

Abstract: Summary 1. Reproductive interference is a negative interspecific sexual interaction that adversely affects the fitness of males and females during reproductive process. Theoretical studies suggest that because reproductive interference is characterized by positive frequency dependence it is far more likely to cause species exclusion than the density dependence of resource competition. However, the respective contributions of resource competition and reproductive interference to species exclusion, which have been frequently observed in many competition studies, remain unclear. 2. We show that reproductive interference is a far more critical cause of species exclusion than resource competition in the competition between Callosobruchus bean weevil species. In competition experiments over several generations, we manipulated the initial relative abundance of the adzuki bean beetle, Callosobruchus chinensis, and the southern cowpea beetle, Callosobruchus maculatus. When the initial adult ratio of C. chinensis : C. maculatus were 6 : 2 and 4 : 4, C. chinensis excluded C. maculatus. However, when C. maculatus was four times more abundant than C. chinensis at the start, we observed the opposite outcome. 3. A behavioural experiment using adults of the two species revealed asymmetric reproductive interference. The fecundity and longevity of C. maculatus females, but not those of C. chinensis females, decreased when the females were kept with heterospecific males. Fecundities of females of both species decreased as the number of heterospecific males increased. In contrast, resource competition at the larval stage resulted in higher survival of C. maculatus than of C. chinensis. 4. These results suggest that the positive frequency-dependent effect of reproductive interference resulted in species exclusion, depending on the initial population ratio of the two species, and the asymmetry of the interference resulted in C. chinensis being dominant in this study, as in previous studies. Classical competition studies should be reviewed in light of this evidence for reproductive interference.

Host density and human activities mediate increased parasite prevalence and richness in primates threatened by habitat loss and fragmentation.

Abstract: 1. Habitat loss and fragmentation are the principal causes of the loss of biological diversity. In addition, parasitic diseases are an emerging threat to many animals. Nevertheless, relatively few studies have tested how habitat loss and fragmentation influence the prevalence and richness of parasites in animals. 2. Several studies of nonhuman primates have shown that measures of human activity and forest fragmentation correlate with parasitism in primates. However, these studies have not tested for the ecological mechanism(s) by which human activities or forest fragmentation influence the prevalence and richness of parasites. 3. We tested the hypothesis that increased host density due to forest fragmentation and loss mediates increases in the prevalence and richness of gastrointestinal parasites in two forest primates, the Tana River red colobus (Procolobus rufomitratus, Peters 1879) and mangabey (Cercocebus galeritus galeritus, Peters 1879). We focused on population density because epidemiological theory states that host density is a key determinant of the prevalence and richness of directly transmitted parasites in animals. 4. The Tana River red colobus and mangabey are endemic to a highly fragmented forest ecosystem in eastern Kenya where habitat changes are caused by a growing human population increasingly dependent on forest resources and on clearing forest for cultivation. 5. We found that the prevalence of parasites in the two monkeys was very high compared to primates elsewhere. Density of monkeys was positively associated with forest area and disturbance in forests. In turn, the prevalence and richness of parasites was significantly associated with monkey density, and attributes indicative of human disturbance in forests. 6. We also found significant differences in the patterns of parasitism between the colobus and the mangabey possibly attributable to differences in their behavioural ecology. Colobus are arboreal folivores while mangabeys are terrestrial habitat generalists.