Showing papers in "Journal of Animal Ecology in 2016"

What is the animal doing? Tools for exploring behavioural structure in animal movements

Abstract: Summary Movement data provide a window – often our only window – into the cognitive, social and biological processes that underlie the behavioural ecology of animals in the wild. Robust methods for identifying and interpreting distinct modes of movement behaviour are of great importance, but complicated by the fact that movement data are complex, multivariate and dependent. Many different approaches to exploratory analysis of movement have been developed to answer similar questions, and practitioners are often at a loss for how to choose an appropriate tool for a specific question. We apply and compare four methodological approaches: first passage time (FPT), Bayesian partitioning of Markov models (BPMM), behavioural change point analysis (BCPA) and a fitted multistate random walk (MRW) to three simulated tracks and two animal trajectories – a sea lamprey (Petromyzon marinus) tracked for 12 h and a wolf (Canis lupus) tracked for 1 year. The simulations – in which, respectively, velocity, tortuosity and spatial bias change – highlight the sensitivity of all methods to model misspecification. Methods that do not account for autocorrelation in the movement variables lead to spurious change points, while methods that do not account for spatial bias completely miss changes in orientation. When applied to the animal data, the methods broadly agree on the structure of the movement behaviours. Important discrepancies, however, reflect differences in the assumptions and nature of the outputs. Important trade-offs are between the strength of the a priori assumptions (low in BCPA, high in MRW), complexity of output (high in the BCPA, low in the BPMM and MRW) and explanatory potential (highest in the MRW). The animal track analysis suggests some general principles for the exploratory analysis of movement data, including ways to exploit the strengths of the various methods. We argue for close and detailed exploratory analysis of movement before fitting complex movement models.

COMADRE: a global data base of animal demography

Abstract: 1. The open-data scientific philosophy is being widely adopted and proving to promote considerable progress in ecology and evolution. Open-data global data bases now exist on animal migration, species distribution, conservation status, etc. However, a gap exists for data on population dynamics spanning the rich diversity of the animal kingdom world-wide. This information is fundamental to our understanding of the conditions that have shaped variation in animal life histories and their relationships with the environment, as well as the determinants of invasion and extinction. 2. Matrix population models (MPMs) are among the most widely used demographic tools by animal ecologists. MPMs project population dynamics based on the reproduction, survival and development of individuals in a population over their life cycle. The outputs from MPMs have direct biological interpretations, facilitating comparisons among animal species as different as Caenorhabditis elegans, Loxodonta africana and Homo sapiens. 3. Thousands of animal demographic records exist in the form of MPMs, but they are dispersed throughout the literature, rendering comparative analyses difficult. Here, we introduce the COMADRE Animal Matrix Database, an open-data online repository, which in its version 1.0.0 contains data on 345 species world-wide, from 402 studies with a total of 1625 population projection matrices. COMADRE also contains ancillary information (e.g. ecoregion, taxonomy, biogeography, etc.) that facilitates interpretation of the numerous demographic metrics that can be derived from its MPMs. We provide R code to some of these examples. 4. Synthesis: We introduce the COMADRE Animal Matrix Database, a resource for animal demography. Its open-data nature, together with its ancillary information, will facilitate comparative analysis, as will the growing availability of databases focusing on other aspects of the rich animal diversity, and tools to query and combine them. Through future frequent updates of COMADRE, and its integration with other online resources, we encourage animal ecologists to tackle global ecological and evolutionary questions with unprecedented sample size.

Tackling extremes: challenges for ecological and evolutionary research on extreme climatic events

Abstract: Extreme climatic events (ECEs) are predicted to become more frequent as the climate changes. A rapidly increasing number of studies - though few on animals - suggest that the biological consequences of ECEs can be severe. However, ecological research on the impacts of ECEs has been limited by a lack of cohesiveness and structure. ECEs are often poorly defined and have often been confusingly equated with climatic variability, making comparison between studies difficult. In addition, a focus on short-term studies has provided us with little information on the long-term implications of ECEs, and the descriptive and anecdotal nature of many studies has meant it is still unclear what the key research questions are. Synthesizing the current state of work is essential to identify ways to make progress. We conduct a synthesis of the literature and discuss conceptual and practical challenges faced by research on ECEs. We consider three steps to advance research. First, we discuss the importance of choosing an ECE definition and identify the pros and cons of 'climatological' and 'biological' definitions of ECEs. Secondly, we advocate research beyond short-term descriptive studies to address questions concerning the long-term implications of ECEs, focussing on selective pressures and phenotypically plastic responses and how they might differ from responses to a changing climatic mean. Finally, we encourage a greater focus on multi-event studies that help us understand the implications of changing patterns of ECEs, through the combined use of modelling, experimental and observational field studies. This study aims to open a discussion on the definitions, questions and methods currently used to study ECEs, which will lead to a more cohesive approach to future ECE research.

The challenges of the first migration : movement and behaviour of juvenile vs. adult white storks with insights regarding juvenile mortality

Abstract: Migration conveys an immense challenge, especially for juvenile birds coping with enduring and risky journeys shortly after fledging. Accordingly, juveniles exhibit considerably lower survival rates compared to adults, particularly during migration. Juvenile white storks (Ciconia ciconia), which are known to rely on adults during their first fall migration presumably for navigational purposes, also display much lower annual survival than adults. Using detailed GPS and body acceleration data, we examined the patterns and potential causes of age-related differences in fall migration properties of white storks by comparing first-year juveniles and adults. We compared juvenile and adult parameters of movement, behaviour and energy expenditure (estimated from overall dynamic body acceleration) and placed this in the context of the juveniles' lower survival rate. Juveniles used flapping flight vs. soaring flight 23% more than adults and were estimated to expend 14% more energy during flight. Juveniles did not compensate for their higher flight costs by increased refuelling or resting during migration. When juveniles and adults migrated together in the same flock, the juvenile flew mostly behind the adult and was left behind when they separated. Juveniles showed greater improvement in flight efficiency throughout migration compared to adults which appears crucial because juveniles exhibiting higher flight costs suffered increased mortality. Our findings demonstrate the conflict between the juveniles' inferior flight skills and their urge to keep up with mixed adult-juvenile flocks. We suggest that increased flight costs are an important proximate cause of juvenile mortality in white storks and likely in other soaring migrants and that natural selection is operating on juvenile variation in flight efficiency.

Movement is the glue connecting home ranges and habitat selection.

Abstract: Summary 1. Animal space use has been studied by focusing either on geographic (e.g. home ranges, species’ distribution) or on environmental (e.g. habitat use and selection) space. However, all patterns of space use emerge from individual movements, which are the primary means by which animals change their environment. 2. Individuals increase their use of a given area by adjusting two key movement components: the duration of their visit and/or the frequency of revisits. Thus, in spatially heterogeneous environments, animals exploit known, high-quality resource areas by increasing their residence time (RT) in and/or decreasing their time to return (TtoR) to these areas. We expected that spatial variation in these two movement properties should lead to observed patterns of space use in both geographic and environmental spaces. We derived a set of nine predictions linking spatial distribution of movement properties to emerging space-use patterns. We predicted that, at a given scale, high variation in RT and TtoR among habitats leads to strong habitat selection and that long RT and short TtoR result in a small home range size. 3. We tested these predictions using moose (Alces alces) GPS tracking data. We first modelled the relationship between landscape characteristics and movement properties. Then, we investigated how the spatial distribution of predicted movement properties (i.e. spatial autocorrelation, mean, and variance of RT and TtoR) influences home range size and hierarchical habitat selection. 4. In landscapes with high spatial autocorrelation of RT and TtoR, a high variation in both RT and TtoR occurred in home ranges. As expected, home range location was highly selective in such landscapes (i.e. second-order habitat selection); RT was higher and TtoR lower within the selected home range than outside, and moose home ranges were small. Within home ranges, a higher variation in both RT and TtoR was associated with higher selectivity among habitat types (i.e. third-order habitat selection). 5. Our findings show how patterns of geographic and environmental space use correspond to the two sides of a coin, linked by movement responses of individuals to environmental heterogeneity. By demonstrating the potential to assess the consequences of altering RT or TtoR (e.g. through human disturbance or climatic changes) on home range size and habitat selection, our work sets the basis for new theoretical and methodological advances in movement ecology.

'You shall not pass!': quantifying barrier permeability and proximity avoidance by animals.

Abstract: Summary 1. Impediments to animal movement are ubiquitous and vary widely in both scale and permeability. It is essential to understand how impediments alter ecological dynamics via their influence on animal behavioural strategies governing space use and, for anthropogenic features such as roads and fences, how to mitigate these effects to effectively manage species and landscapes. 2. Here, we focused primarily on barriers to movement, which we define as features that cannot be circumnavigated but may be crossed. Responses to barriers will be influenced by the movement capabilities of the animal, its proximity to the barriers, and habitat preference. We developed a mechanistic modelling framework for simultaneously quantifying the permeability and proximity effects of barriers on habitat preference and movement. 3. We used simulations based on our model to demonstrate how parameters on movement, habitat preference and barrier permeability can be estimated statistically. We then applied the model to a case study of road effects on wild mountain reindeer summer movements. 4. This framework provided unbiased and precise parameter estimates across a range of strengths of preferences and barrier permeabilities. The quality of permeability estimates, however, was correlated with the number of times the barrier is crossed and the number of locations in proximity to barriers. In the case study we found that reindeer avoided areas near roads and that roads are semi-permeable barriers to movement. There was strong avoidance of roads extending up to c. 1 km for four of five animals, and having to cross roads reduced the probability of movement by 68� 6% (range 3� 5–99� 5%). 5. Human infrastructure has embedded within it the idea of networks: nodes connected by linear features such as roads, rail tracks, pipelines, fences and cables, many of which divide the landscape and limit animal movement. The unintended but potentially profound consequences of infrastructure on animals remain poorly understood. The rigorous framework for simultaneously quantifying movement, habitat preference and barrier permeability developed here begins to address this knowledge gap.

Influences of sampling effort on detected patterns and structuring processes of a Neotropical plant–hummingbird network

Abstract: Virtually all empirical ecological interaction networks to some extent suffer from undersampling. However, how limitations imposed by sampling incompleteness affect our understanding of ecological networks is still poorly explored, which may hinder further advances in the field. Here, we use a plant-hummingbird network with unprecedented sampling effort (2716 h of focal observations) from the Atlantic Rainforest in Brazil, to investigate how sampling effort affects the description of network structure (i.e. widely used network metrics) and the relative importance of distinct processes (i.e. species abundances vs. traits) in determining the frequency of pairwise interactions. By dividing the network into time slices representing a gradient of sampling effort, we show that quantitative metrics, such as interaction evenness, specialization (H2 '), weighted nestedness (wNODF) and modularity (Q; QuanBiMo algorithm) were less biased by sampling incompleteness than binary metrics. Furthermore, the significance of some network metrics changed along the sampling effort gradient. Nevertheless, the higher importance of traits in structuring the network was apparent even with small sampling effort. Our results (i) warn against using very poorly sampled networks as this may bias our understanding of networks, both their patterns and structuring processes, (ii) encourage the use of quantitative metrics little influenced by sampling when performing spatio-temporal comparisons and (iii) indicate that in networks strongly constrained by species traits, such as plant-hummingbird networks, even small sampling is sufficient to detect their relative importance for the frequencies of interactions. Finally, we argue that similar effects of sampling are expected for other highly specialized subnetworks.

Selective extinction drives taxonomic and functional alpha and beta diversities in island bird assemblages

Abstract: Taxonomic diversity considers all species being equally different from each other and thus disregards species' different ecological functions. Exploring taxonomic and functional aspects of biodiversity simultaneously can better understand the processes of community assembly. We analysed taxonomic and functional alpha and beta diversities of breeding bird assemblages on land-bridge islands in the Thousand Island Lake, China. Given the high dispersal ability of most birds at this spatial scale (several kilometres), we predicted (i) selective extinction driving alpha and beta diversities after the creation of land-bridge islands of varying area and (ii) low taxonomic and functional beta diversities that were not correlated to spatial distance. Breeding birds were surveyed on 37 islands annually from 2007 to 2014. We decomposed beta diversity of breeding birds into spatial turnover and nestedness-resultant components, and related taxonomic and functional diversities to island area and isolation using power regression models (for alpha diversity) and multiple regression models on distance matrices (for beta diversity). We then ran simulations to assess the strength of the correlations between taxonomic and functional diversities. Results revealed that both taxonomic and functional alpha diversities increased with island area. The taxonomic nestedness-resultant and turnover components increased and decreased with difference in area, respectively, but functional counterparts did not. Isolation played a minor role in explaining alpha- and beta-diversity patterns. By partitioning beta diversity, we found low levels of overall taxonomic and functional beta diversities. The functional nestedness-resultant component dominated overall functional beta diversity, whereas taxonomic turnover was the dominant component for taxonomic beta diversity. The simulation showed that functional alpha and beta diversities were significantly correlated with taxonomic diversities, and the observed values of correlations were significantly different from null expectations of random extinction. Our assessment of island bird assemblages validated the predictions of no distance effects and low beta diversity due to pervasive dispersal events among islands and also suggested that selective extinction drives taxonomic and functional alpha and beta diversities. The contrasting turnover and nestedness-resultant components of taxonomic and functional beta diversities demonstrate the importance of considering the multifaceted nature of biodiversity when examining community assembly.

REVIEW: Can habitat selection predict abundance?

Abstract: Habitats have substantial influence on the distribution and abundance of animals. Animals' selective movement yields their habitat use. Animals generally are more abundant in habitats that are selected most strongly. Models of habitat selection can be used to distribute animals on the landscape or their distribution can be modelled based on data of habitat use, occupancy, intensity of use or counts of animals. When the population is at carrying capacity or in an ideal-free distribution, habitat selection and related metrics of habitat use can be used to estimate abundance. If the population is not at equilibrium, models have the flexibility to incorporate density into models of habitat selection; but abundance might be influenced by factors influencing fitness that are not directly related to habitat thereby compromising the use of habitat-based models for predicting population size. Scale and domain of the sampling frame, both in time and space, are crucial considerations limiting application of these models. Ultimately, identifying reliable models for predicting abundance from habitat data requires an understanding of the mechanisms underlying population regulation and limitation.

Predicting the continuum between corridors and barriers to animal movements using Step Selection Functions and Randomized Shortest Paths.

Abstract: The loss, fragmentation and degradation of habitat everywhere on Earth prompts increasing attention to identifying landscape features that support animal movement (corridors) or impedes it (barriers). Most algorithms used to predict corridors assume that animals move through preferred habitat either optimally (e.g. least cost path) or as random walkers (e.g. current models), but neither extreme is realistic. We propose that corridors and barriers are two sides of the same coin and that animals experience landscapes as spatiotemporally dynamic corridor-barrier continua connecting (separating) functional areas where individuals fulfil specific ecological processes. Based on this conceptual framework, we propose a novel methodological approach that uses high-resolution individual-based movement data to predict corridor-barrier continua with increased realism. Our approach consists of two innovations. First, we use step selection functions (SSF) to predict friction maps quantifying corridor-barrier continua for tactical steps between consecutive locations. Secondly, we introduce to movement ecology the randomized shortest path algorithm (RSP) which operates on friction maps to predict the corridor-barrier continuum for strategic movements between functional areas. By modulating the parameter Ѳ, which controls the trade-off between exploration and optimal exploitation of the environment, RSP bridges the gap between algorithms assuming optimal movements (when Ѳ approaches infinity, RSP is equivalent to LCP) or random walk (when Ѳ → 0, RSP → current models). Using this approach, we identify migration corridors for GPS-monitored wild reindeer (Rangifer t. tarandus) in Norway. We demonstrate that reindeer movement is best predicted by an intermediate value of Ѳ, indicative of a movement trade-off between optimization and exploration. Model calibration allows identification of a corridor-barrier continuum that closely fits empirical data and demonstrates that RSP outperforms models that assume either optimality or random walk. The proposed approach models the multiscale cognitive maps by which animals likely navigate real landscapes and generalizes the most common algorithms for identifying corridors. Because suboptimal, but non-random, movement strategies are likely widespread, our approach has the potential to predict more realistic corridor-barrier continua for a wide range of species.

Behavioural flexibility in migratory behaviour in a long-lived large herbivore.

Abstract: Summary Migratory animals are predicted to enhance lifetime fitness by obtaining higher quality forage and/or reducing predation risk compared to non-migratory conspecifics. Despite evidence for behavioural flexibility in other taxa, previous research on large mammals has often assumed that migratory behaviour is a fixed behavioural trait. Migratory behaviour may be plastic for many species, although few studies have tested for individual-level flexibility using long-term monitoring of marked individuals, especially in large mammals such as ungulates. We tested variability in individual migratory behaviour using a 10-year telemetry data set of 223 adult female elk (Cervus elaphus) in the partially migratory Ya Ha Tinda population in Alberta, Canada. We used net squared displacement (NSD) to classify migratory strategy for each individual elk-year. Individuals switched between migrant and resident strategies at a mean rate of 15% per year, and migrants were more likely to switch than residents. We then tested how extrinsic (climate, elk/wolf abundance) and intrinsic (age) factors affected the probability of migrating, and, secondly, the decision to switch between migratory strategies. Over 630 individual elk-years, the probability of an individual elk migrating increased following a severe winter, in years of higher wolf abundance, and with increasing age. At an individual elk level, we observed 148 switching events of 430 possible transitions in elk monitored at least 2 years. We found switching was density-dependent, where migrants switched to a resident strategy at low elk abundance, but residents switched more to a migrant strategy at high elk abundance. Precipitation during the previous summer had a weak carryover effect, with migrants switching slightly more following wetter summers, whereas residents showed the opposite pattern. Older migrant elk rarely switched, whereas resident elk switched more frequently to migrate at older ages. Our results show migratory behaviour in ungulates is an individually variable trait that can respond to intrinsic, environmental and density-dependent forces. Different strategies had opposing responses to density-dependent and intrinsic drivers, providing a stabilizing mechanism for the maintenance of partial migration and demographic fitness in this population.

Reefscapes of fear: predation risk and reef hetero-geneity interact to shape herbivore foraging behaviour.

Abstract: Predators can exert strong direct and indirect effects on ecological communities by intimidating their prey. The nature of predation risk effects is often context dependent, but in some ecosystems these contingencies are often overlooked. Risk effects are often not uniform across landscapes or among species. Indeed, they can vary widely across gradients of habitat complexity and with different prey escape tactics. These context dependencies may be especially important for ecosystems such as coral reefs that vary widely in habitat complexity and have species-rich predator and prey communities. With field experiments using predator decoys of the black grouper (Mycteroperca bonaci), we investigated how reef complexity interacts with predation risk to affect the foraging behaviour and herbivory rates of large herbivorous fishes (e.g. parrotfishes and surgeonfishes) across four coral reefs in the Florida Keys (USA). In both high and low complexity areas of the reef, we measured how herbivory changed with increasing distance from the predator decoy to examine how herbivorous fishes reconcile the conflicting demands of avoiding predation vs. foraging within a reefscape context. We show that with increasing risk, herbivorous fishes consumed dramatically less food (ca. 90%) but fed at a faster rate when they did feed (ca. 26%). Furthermore, we show that fishes foraging closest to the predator decoy were 40% smaller than those that foraged at further distances. Thus, smaller individuals showed muted response to predation risk compared to their larger counterparts, potentially due to their decreased risk to predation or lower reproductive value (i.e. the asset protection principle). Habitat heterogeneity mediated risk effects differently for different species of herbivores, with predation risk more strongly suppressing herbivore feeding in more complex areas and for individuals at higher risk of predation. Predators appear to create a reefscape of fear that changes the size structure of herbivores towards smaller individuals, increases individual feeding rates, but suppresses overall amounts of primary producers consumed, potentially altering patterns of herbivory, an ecosystem process critical for healthy coral reefs.

How many routes lead to migration? Comparison of methods to assess and characterize migratory movements.

Abstract: Decreasing rate of migration in several species as a consequence of climate change and anthropic pressure, together with increasing evidence of space-use strategies intermediate between residency and complete migration, are very strong motivations to evaluate migration occurrence and features in animal populations. The main goal of this paper was to perform a relative comparison between methods for identifying and characterizing migration at the individual and population level on the basis of animal location data. We classified 104 yearly individual trajectories from five populations of three deer species as migratory or non-migratory, by means of three methods: seasonal home range overlap, spatio-temporal separation of seasonal clusters and the Net Squared Displacement (NSD) method. For migratory cases, we also measured timing and distance of migration and residence time on the summer range. Finally, we compared the classification in migration cases across methods and populations. All methods consistently identified migration at the population level, that is, they coherently distinguished between complete or almost complete migratory populations and partially migratory populations. However, in the latter case, methods coherently classified only about 50% of the single cases, that is they classified differently at the individual-animal level. We therefore infer that the comparison of methods may help point to 'less-stereotyped' cases in the residency-to-migration continuum. For cases consistently classified by all methods, no significant differences were found in migration distance, or residence time on summer ranges. Timing of migration estimated by NSD was earlier than by the other two methods, both for spring and autumn migrations. We suggest three steps to identify improper inferences from migration data and to enhance understanding of intermediate space-use strategies. We recommend (i) classifying migration behaviours using more than one method, (ii) performing sensitivity analysis on method parameters to identify the extent of the differences and (iii) investigating inconsistently classified cases as these may often be ecologically interesting (i.e. less-stereotyped migratory behaviours).

Predator swamping reduces predation risk during nocturnal migration of juvenile salmon in a high-mortality landscape

Abstract: Animal migrations are costly and are often characterized by high predation risk for individuals. Three of the most oft-assumed mechanisms for reducing risk for migrants are swamping predators with high densities, specific timing of migrations and increased body size. Assessing the relative importance of these mechanisms in reducing predation risk particularly for migrants is generally lacking due to the difficulties in tracking the fate of individuals and population-level characteristics simultaneously. We used acoustic telemetry to track migration behaviour and survival of juvenile sockeye salmon (Oncorhynchus nerka) smolts released over a wide range of conspecific outmigration densities in a river associated with poor survival. The landscape was indeed high risk; smolt survival was poor (˜68%) over 13·5 km of river examined even though migration was rapid (generally <48 h). Our results demonstrate that smolts largely employ swamping of predators to reduce predation risk. Increased densities of co-migrant conspecifics dramatically improved survival of smolts. The strong propensity for nocturnal migration resulted in smolts pausing downstream movements until the next nightfall, greatly increasing relative migration durations for smolts that could not traverse the study area in a single night. Smolt size did not appear to impact predation risk, potentially due to unique characteristics of the system or our inability to tag the entire size range of outmigrants. Movement behaviours were important in traversing this high-risk landscape and provide rare evidence for swamping to effectively reduce individual predation risk.

Co-infections and environmental conditions drive the distributions of blood parasites in wild birds

Abstract: Experimental work increasingly suggests that non-random pathogen associations can affect the spread or severity of disease. Yet due to difficulties distinguishing and interpreting co-infections, evidence for the presence and directionality of pathogen co-occurrences in wildlife is rudimentary. We provide empirical evidence for pathogen co-occurrences by analysing infection matrices for avian malaria (Haemoproteus and Plasmodium spp.) and parasitic filarial nematodes (microfilariae) in wild birds (New Caledonian Zosterops spp.). Using visual and genus-specific molecular parasite screening, we identified high levels of co-infections that would have been missed using PCR alone. Avian malaria lineages were assigned to species level using morphological descriptions. We estimated parasite co-occurrence probabilities, while accounting for environmental predictors, in a hierarchical multivariate logistic regression. Co-infections occurred in 36% of infected birds. We identified both positively and negatively correlated parasite co-occurrence probabilities when accounting for host, habitat and island effects. Two of three pairwise avian malaria co-occurrences were strongly negative, despite each malaria parasite occurring across all islands and habitats. Birds with microfilariae had elevated heterophil to lymphocyte ratios and were all co-infected with avian malaria, consistent with evidence that host immune modulation by parasitic nematodes facilitates malaria co-infections. Importantly, co-occurrence patterns with microfilariae varied in direction among avian malaria species; two malaria parasites correlated positively but a third correlated negatively with microfilariae. We show that wildlife co-infections are frequent, possibly affecting infection rates through competition or facilitation. We argue that combining multiple diagnostic screening methods with multivariate logistic regression offers a platform to disentangle impacts of environmental factors and parasite co-occurrences on wildlife disease.

Patterns and predictors of β-diversity in the fragmented Brazilian Atlantic forest: a multiscale analysis of forest specialist and generalist birds.

Abstract: Biodiversity maintenance in human-altered landscapes (HALs) depends on the species turnover among localities, but the patterns and determinants of β-diversity in HALs are poorly known. In fact, declines, increases and neutral shifts in β-diversity have all been documented, depending on the landscape, ecological group and spatial scale of analysis. We shed some light on this controversy by assessing the patterns and predictors of bird β-diversity across multiple spatial scales considering forest specialist and habitat generalist bird assemblages. We surveyed birds from 144 point counts in 36 different forest sites across two landscapes with different amount of forest cover in the Brazilian Atlantic forest. We analysed β-diversity among points, among sites and between landscapes with multiplicative diversity partitioning of Hill numbers. We tested whether β-diversity among points was related to within-site variations in vegetation structure, and whether β-diversity among sites was related to site location and/or to differences among sites in vegetation structure and landscape composition (i.e. per cent forest and pasture cover surrounding each site). β-diversity between landscapes was lower than among sites and among points in both bird assemblages. In forest specialist birds, the landscape with less forest cover showed the highest β-diversity among sites (bird differentiation among sites), but generalist birds showed the opposite pattern. At the local scale, however, the less forested landscape showed the lowest β-diversity among points (bird homogenization within sites), independently of the bird assemblage. β-diversity among points was weakly related to vegetation structure, but higher β-diversity values were recorded among sites that were more isolated from each other, and among sites with higher differences in landscape composition, particularly in the less forested landscape. Our findings indicate that patterns of bird β-diversity vary across scales and are strongly related to landscape composition. Bird assemblages are shaped by both environmental filtering and dispersal limitation, particularly in less forested landscapes. Conservation and management strategies should therefore prevent deforestation in this biodiversity hotspot.

Temperature-associated habitat selection in a cold-water marine fish.

Abstract: Habitat selection is a complex process, which involves behavioural decisions guided by the multiple needs and constraints faced by individuals. Climate-induced changes in environmental conditions may alter those trade-offs and resulting habitat use patterns. In this study, we investigated the effect of sea temperature on habitat selection and habitat use of acoustically tagged Atlantic cod (Gadus morhua) at the Norwegian Skagerrak coast. Significant relationships between ocean temperature and habitat selection and use were found. Under favourable sea temperature thresholds (<16 °C), cod selected vegetated habitats, such as eelgrass and macroalgae beds, available in shallow areas. Selection for those habitats was especially high at night, when cod tended to ascend to shallower areas, presumably to feed. Selection and use of those habitats decreased significantly as temperature rose. Under increased sea surface temperature conditions, cod were absent from vegetated shallow habitats, both during the day and night, and selected instead non-vegetated rocky bottoms and sand habitats, available in deeper, colder areas. This study shows the dynamic nature of habitat selection and strongly suggests that cod in this region have to trade off food availability against favourable temperature conditions. Future increases in ocean temperature are expected to further influence the spatial behaviour of marine fish, potentially affecting individual fitness and population dynamics.

Fight-flight or freeze-hide? Personality and metabolic phenotype mediate physiological defence responses in flatfish

Abstract: Survival depends on appropriate behavioural and physiological responses to danger. In addition to active 'fight-flight' defence responses, a passive 'freeze-hide' response is adaptive in some contexts. However, the physiological mechanisms determining which individuals choose a given defence response remain poorly understood. We examined the relationships among personality, metabolic performance and physiological stress responses across an environmental gradient in the olive flounder, Paralichthys olivaceus. We employed four behavioural assays to document the existence of two distinct behavioural types ('bold' and 'shy') in this species. We found consistent metabolic differences between individuals of a given behavioural type across an environmental gradient: shy individuals had overall lower aerobic scope, maximum metabolic rate and standard metabolic rate than bold individuals in both high (25 ppt) and low (3 ppt) salinity. These behavioural and metabolic differences translated into divergent physiological responses during acute stress: shy individuals adopted a passive 'freeze-hide' response by reducing their oxygen consumption rates (akin to shallow breathing) whereas bold individuals adopted an active 'fight-flight' response by increasing their rates of respiration. These distinct defence strategies were repeatable within individuals between salinity treatments. Although it has been suggested theoretically, this is the first empirical evidence that the metabolic response to stressful situations differs between bold and shy individuals. Our results emphasize the importance of incorporating physiological measures to understand the mechanisms driving persistent inter-individual differences in animals.

Beyond neutral and forbidden links: morphological matches and the assembly of mutualistic hawkmoth-plant networks

Abstract: Summary A major challenge in evolutionary ecology is to understand how co-evolutionary processes shape patterns of interactions between species at community level. Pollination of flowers with long corolla tubes by long-tongued hawkmoths has been invoked as a showcase model of co-evolution. Recently, optimal foraging models have predicted that there might be a close association between mouthparts' length and the corolla depth of the visited flowers, thus favouring trait convergence and specialization at community level. Here, we assessed whether hawkmoths more frequently pollinate plants with floral tube lengths similar to their proboscis lengths (morphological match hypothesis) against abundance-based processes (neutral hypothesis) and ecological trait mismatches constraints (forbidden links hypothesis), and how these processes structure hawkmoth–plant mutualistic networks from five communities in four biogeographical regions of South America. We found convergence in morphological traits across the five communities and that the distribution of morphological differences between hawkmoths and plants is consistent with expectations under the morphological match hypothesis in three of the five communities. In the two remaining communities, which are ecotones between two distinct biogeographical areas, interactions are better predicted by the neutral hypothesis. Our findings are consistent with the idea that diffuse co-evolution drives the evolution of extremely long proboscises and flower tubes, and highlight the importance of morphological traits, beyond the forbidden links hypothesis, in structuring interactions between mutualistic partners, revealing that the role of niche-based processes can be much more complex than previously known.

Integrated population modelling reveals a perceived source to be a cryptic sink

Abstract: Demographic links among fragmented populations are commonly studied as source-sink dynamics, whereby source populations exhibit net recruitment and net emigration, while sinks suffer net mortality but enjoy net immigration. It is commonly assumed that large, persistent aggregations of individuals must be sources, but this ignores the possibility that they are sinks instead, buoyed demographically by immigration. We tested this assumption using Bayesian integrated population modelling of Greenland white-fronted geese (Anser albifrons flavirostris) at their largest wintering site (Wexford, Ireland), combining capture-mark-recapture, census and recruitment data collected from 1982 to 2010. Management for this subspecies occurs largely on wintering areas; thus, study of source-sink dynamics of discrete regular wintering units provides unprecedented insights into population regulation and enables identification of likely processes influencing population dynamics at Wexford and among 70 other Greenland white-fronted goose wintering subpopulations. Using results from integrated population modelling, we parameterized an age-structured population projection matrix to determine the contribution of movement rates (emigration and immigration), recruitment and mortality to the dynamics of the Wexford subpopulation. Survival estimates for juvenile and adult birds at Wexford and adult birds elsewhere fluctuated over the 29-year study period, but were not identifiably different. However, per capita recruitment rates at Wexford in later years (post-1995) were identifiably lower than in earlier years (pre-1995). The observed persistence of the Wexford subpopulation was only possible with high rates of immigration, which exceeded emigration in each year. Thus, despite its apparent stability, Wexford has functioned as a sink over the entire study period. These results demonstrate that even large subpopulations can potentially be sinks, and that movement dynamics (e.g. immigration) among winters can dramatically obscure key processes driving subpopulation size. Further, novel population models which integrate capture-mark-recapture, census and recruitment data are essential to correctly ascribing source-sink status and accurately informing development of site-safeguard networks.

Integrating the pace-of-life syndrome across species, sexes and individuals: covariation of life history and personality under pesticide exposure.

Abstract: The pace-of-life syndrome (POLS) hypothesis integrates covariation of life-history traits along a fast-slow continuum and covariation of behavioural traits along a proactive-reactive personality continuum. Few studies have investigated these predicted life-history/personality associations among species and between sexes. Furthermore, whether and how contaminants interfere with POLS patterns remains unexplored. We tested for covariation patterns in life history and in behaviour, and for life-history/personality covariation among species, among individuals within species and between sexes. Moreover, we investigated whether pesticide exposure affects covariation between life history and behaviour and whether species and sexes with a faster POLS strategy have a higher sensitivity to pesticides. We reared larvae of four species of Ischnura damselflies in a common garden experiment with an insecticide treatment (chlorpyrifos absent/present) in the final instar. We measured four life-history traits (larval growth rate during the pesticide treatment, larval development time, adult mass and life span) and two behavioural traits (larval feeding activity and boldness, each before and after the pesticide treatment). At the individual level, life-history traits and behavioural traits aligned along a fast-slow and a proactive-reactive continuum, respectively. Species-specific differences in life history, with fast-lived species having a faster larval growth and development, a lower mass at emergence and a shorter life span, suggested that time constraints in the larval stage were predictably driving life-history evolution both in the larval stage and across metamorphosis in the adult stage. Across species, females were consistently more slow-lived than males, reflecting that a large body size and a long life span are generally more important for females. In contrast to the POLS hypothesis, there was only little evidence for the expected positive coupling between life-history pace and proactivity. Pesticide exposure decreased larval growth rate and affected life-history/personality covariation in the most fast-lived species. Our study supports the existence of life-history and behavioural continua with limited support for life-history/personality covariation. Variation in digestive physiology may explain this decoupling of life history and behaviour and provide valuable mechanistic insights to understand and predict the occurrence of life-history/personality covariation patterns.

Adding constraints to predation through allometric relation of scats to consumption.

Abstract: A thorough understanding of mechanisms of prey consumption by carnivores and the constraints on predation help us in evaluating the role of carnivores in an ecosystem. This is crucial in developing appropriate management strategies for their conservation and mitigating human-carnivore conflict. Current models on optimal foraging suggest that mammalian carnivores would profit most from killing the largest prey that they can subdue with minimal risk of injury to themselves. Wild carnivore diets are primarily estimated through analysis of their scats. Using extensive feeding experiments (n = 68) on a wide size range (4·5-130 kg) of obligate carnivores - lion, leopard, jungle cat and domestic cat, we parameterize biomass models that best relate consumption to scat production. We evaluate additional constraints of gut fill, prey digestibility and carcass utilization on carnivory that were hereto not considered in optimal foraging studies. Our results show that patterns of consumption to scat production against prey size are similar and asymptotic, contrary to established linear models, across these carnivores after accounting for the effect of carnivore size. This asymptotic, allometric relationship allowed us to develop a generalized model: biomass consumed per collectable scat/predator weight = 0·033-0·025exp(-4·284(prey weight/predator weight)) , which is applicable to all obligate carnivores to compute prey biomass consumed from scats. Our results also depict a relationship for prey digestibility which saturates at about 90% for prey larger than predator size. Carcass utilization declines exponentially with prey size. These mechanisms result in digestible biomass saturating at prey weights approximately equal to predator weight. Published literature on consumption by tropical carnivores that has relied on linear biomass models is substantially biased. We demonstrate the nature of these biases by correcting diets of tiger, lion and leopard in recent publications. Our analysis suggests that consumption of medium-sized prey was significantly underestimated, while large prey consumption was grossly overestimated in large carnivore diets to date. We highlight that additional constraints of prey digestibility and utilization combined with escalating handling time and risks of killing large prey make prey larger than the predator size unprofitable for obligate carnivores.

Parasite specialization in a unique habitat: hummingbirds as reservoirs of generalist blood parasites of Andean birds

Abstract: Understanding how parasites fill their ecological niches requires information on the processes involved in the colonization and exploitation of unique host species. Switching to hosts with atypical attributes may favour generalists broadening their niches or may promote specialization and parasite diversification as the consequence. We analysed which blood parasites have successfully colonized hummingbirds, and how they have evolved to exploit such a unique habitat. We specifically asked (i) whether the assemblage of Haemoproteus parasites of hummingbirds is the result of single or multiple colonization events, (ii) to what extent these parasites are specialized in hummingbirds or shared with other birds and (iii) how hummingbirds contribute to sustain the populations of these parasites, in terms of both prevalence and infection intensity. We sampled 169 hummingbirds of 19 species along an elevation gradient in Southern Ecuador to analyse the host specificity, diversity and infection intensity of Haemoproteus by molecular and microscopy techniques. In addition, 736 birds of 112 species were analysed to explore whether hummingbird parasites are shared with other birds. Hummingbirds hosted a phylogenetically diverse assemblage of generalist Haemoproteus lineages shared with other host orders. Among these parasites, Haemoproteus witti stood out as the most generalized. Interestingly, we found that infection intensities of this parasite were extremely low in passerines (with no detectable gametocytes), but very high in hummingbirds, with many gametocytes seen. Moreover, infection intensities of H. witti were positively correlated with the prevalence across host species. Our results show that hummingbirds have been colonized by generalist Haemoproteus lineages on multiple occasions. However, one of these generalist parasites (H. witti) seems to be highly dependent on hummingbirds, which arise as the most relevant reservoirs in terms of both prevalence and gametocytaemia. From this perspective, this generalist parasite may be viewed as a hummingbird specialist. This challenges the current paradigm of how to measure host specialization in these parasites, which has important implications to understand disease ecology.

Nematode parasite diversity in birds: the role of host ecology, life history and migration.

Abstract: Previous studies have found that migratory birds generally have a more diverse array of pathogens such as parasites, as well as higher intensities of infection. However, it is not clear whether this is driven by the metabolic and physiological demands of migration, differential selection on host life-history traits or basic ecological differences between migratory and non-migratory species. Parasitic helminths can cause significant pathology in their hosts, and many are trophically transmitted such that host diet and habitat use play key roles in the acquisition of infections. Given the concurrent changes in avian habitats and migratory behaviour, it is critical to understand the degree to which host ecology influences their parasite communities. We examined nematode parasite diversity in 153 species of Anseriformes (water birds) and Accipitriformes (predatory birds) in relation to their migratory behaviour, diet, habitat use, geographic distribution and life history using previously published data. Overall, migrators, host species with wide geographic distributions and those utilizing multiple aquatic habitats had greater nematode richness (number of species), and birds with large clutches harboured more diverse nematode fauna with respect to number of superfamilies. Separate analyses for each host order found similar results related to distribution, habitat use and migration; however, herbivorous water birds played host to a less diverse nematode community compared to those that consume some animals. Birds using multiple aquatic habitats have a more diverse nematode fauna relative to primarily terrestrial species, likely because there is greater opportunity for contact with parasite infectious stages and/or consumption of infected hosts. As such, omnivorous and carnivorous birds using aquatic habitats may be more affected by environmental changes that alter their diet and range. Even though there were no overall differences in their ecology and life history compared with non-migrators, migratory bird species still harboured a more diverse array of nematodes, suggesting that this behaviour places unique demands on these hosts and warrants further study.

Street lighting: sex‐independent impacts on moth movement

Abstract: Artificial lights have become an integral and welcome part of our urban and peri-urban environments. However, recent research has highlighted the potentially negative ecological consequences of ubiquitous artificial light. In particular, insects, especially moths, are expected to be negatively impacted by the presence of artificial lights. Previous research with light traps has shown a male-biased attraction to light in moths. In this study, we sought to determine whether street lights could limit moth dispersal and whether there was any sex bias in attraction to light. More specifically, we aimed to determine sex-specific attraction radii for moths to street lights. We tested these hypotheses by collecting moths for 2 years at an experimental set-up. To estimate the attraction radii, we developed a Markov model and related it to the acquired data. Utilizing multinomial statistics, we found that attraction rates to lights in the middle of the matrix were substantially lower than predicted by the null hypothesis of equal attraction level (0·44 times). With the Markov model, we estimated that a corner light was 2·77 times more attractive than a wing light with an equivalentre attraction radius of c. 23 m around each light. We found neither sexual differences in the attraction rate nor in the attraction radius of males and females. Since we captured three times more males than females, we conclude that sex ratios are representative of operational sex ratios or of different flight activities. These results provide evidence for street lights to limit moth dispersal, and that they seem to act equally on male and female moths. Consequently, public lighting might divide a suitable landscape into many small habitats. Therefore, it is reasonable to assume (i) that public lighting near hedges and bushes or field margins reduces the quality of these important habitat structures and (ii) that public lighting may affect moth movement between patches.

Experimental manipulation of floral scent bouquets restructures flower-visitor interactions in the field.

Abstract: A common structural feature of natural communities is the non-random distribution of pairwise interactions between organisms of different trophic levels. For plant-animal interactions, it is predicted that both stochastic processes and functional plant traits that facilitate or prevent interactions are responsible for these patterns. However, unbiased manipulative field experiments that rigorously test the effects of individual traits on community structure are lacking. We address this gap by manipulating floral scent bouquets in the field. Manipulation of floral scent bouquets led to quantitative as well as qualitative restructuring of flower-visitor networks, making them more generalized. Olfactometer trials confirmed both positive and negative responses to scent bouquets. Our results clearly show that the distribution of insect visitors to the two abundant study plant species reflects the insects' species-specific preferences for floral scents, rather than for visual or morphological floral traits. Thus, floral scents may be of major importance in partitioning flower-visitor interactions. Integrating experimental manipulations of plant traits with field observations of interaction patterns thus represents a promising approach for revealing the processes that structure species assemblages in natural communities.

How to capture fish in a school? Effect of successive predator attacks on seabird feeding success

Abstract: Prey aggregations, such as fish schools, attract numerous predators. This typically leads to the formation of multispecific groups of predators. These aggregations can be seen both as a place of increased competition and as a place of possible facilitation between predators. Consequently, the functional role of such predator-prey aggregation is uncertain, and its effect on individual feeding success is virtually unknown. Using underwater film footage of different predators feeding on fish schools during the sardine run in South Africa, we directly measured the in situ feeding success of individual Cape gannets Morus capensis in different foraging situations. We determined the types of Cape gannet attacks (direct plunge dive or plunge dive followed by underwater pursuit) and we measured the occurrences and timing of attacks from the different species (mostly Cape gannets and long-beaked common dolphins Delphinus capensis). We also estimated the size of the targeted fish schools. These observations were complemented with a simulation model to evaluate the cumulative effect of successive predator attacks on the prey aggregation structure. The probability to capture a fish in one feeding attempt by Cape gannets averaged 0·28. It was lower when gannets engaged in underwater prey pursuit after the plunge compared to direct plunge (0·13 vs. 0·36). We found no effect of the number of prey on gannets' feeding success. However, the timing and frequency of attacks influenced strongly and positively the feeding success of individuals. The probability to capture a fish was the lowest (0·16) when no attack occurred in the few seconds (1-15 s) prior to a dive and the highest (˜0·4, i.e. more than twice) when one or two attacks occurred during this time window. The simulation model showed that a prey aggregation disorganized just after an attack and that the maximum of disturbance was obtained a few seconds after the initiation of the successive attacks. Our study suggests that, in multispecies predator assemblages, the cumulative effect (through disorganization of school cohesiveness) of the multiple species attacking a prey aggregation may increase the feeding success of each individual. Therefore, facilitation between predators is likely to overcome competition in these multispecific assemblages.

Adaptive strategies in nocturnally migrating insects and songbirds: contrasting responses to wind

Abstract: Animals that use flight as their mode of transportation must cope with the fact that their migration and orientation performance is strongly affected by the flow of the medium they are moving in, that is by the winds. Different strategies can be used to mitigate the negative effects and benefit from the positive effects of a moving flow. The strategies an animal can use will be constrained by the relationship between the speed of the flow and the speed of the animal's own propulsion in relation to the surrounding air. Here we analyse entomological and ornithological radar data from north-western Europe to investigate how two different nocturnal migrant taxa, the noctuid moth Autographa gamma and songbirds, deal with wind by analysing variation in resulting flight directions in relation to the wind-dependent angle between the animal's heading and track direction. Our results, from fixed locations along the migratory journey, reveal different global strategies used by moths and songbirds during their migratory journeys. As expected, nocturnally migrating moths experienced a greater degree of wind drift than nocturnally migrating songbirds, but both groups were more affected by wind in autumn than in spring. The songbirds' strategies involve elements of both drift and compensation, providing some benefits from wind in combination with destination and time control. In contrast, moths expose themselves to a significantly higher degree of drift in order to obtain strong wind assistance, surpassing the songbirds in mean ground speed, at the cost of a comparatively lower spatiotemporal migratory precision. Moths and songbirds show contrasting but adaptive responses to migrating through a moving flow, which are fine-tuned to the respective flight capabilities of each group in relation to the wind currents they travel within.

The contribution of developmental experience vs. condition to life history, trait variation and individual differences.

Abstract: 1. Developmental experience, for example food abundance during juvenile stages, is known to affect life history and behaviour. However, the life history and behavioural consequences of developmental experience have rarely been studied in concert. As a result, it is still unclear whether developmental experience affects behaviour through changes in life history, or independently of it. 2. The effect of developmental experience on life history and behaviour may also be masked or affected by individual condition during adulthood. Thus, it is critical to tease apart the effects of developmental experience and current individual condition on life history and behaviour. 3. In this study, we manipulated food abundance during development in the western black widow spider, Latrodectus hesperus, by rearing spiders on either a restricted or ad lib diet. We separated developmental from condition-dependent effects by assaying adult foraging behaviour (tendency to attack prey and to stay on out of the refuge following an attack) and web structure multiple times under different levels of satiation following different developmental treatments. 4. Spiders reared under food restriction matured slower and at a smaller size than spiders reared in ad lib conditions. Spiders reared on a restricted diet were more aggressive towards prey and built webs structured for prey capture, while spiders reared on an ad lib diet were less aggressive and built safer webs. Developmental treatment affected which traits were plastic as adults: restricted spiders built safer webs when their adult condition increased, while ad lib spiders reduced their aggression when their adult condition increased. The amount of individual variation in behaviour and web structure varied with developmental treatment. Spiders reared on a restricted diet exhibited consistent variation in all aspects of foraging behaviour and web structure, while spiders reared on an ad lib diet exhibited consistent individual variation in aggression and web weight only. 5. Developmental experience affected the average life history, behaviour and web structure of spiders, but also shaped the amount of phenotypic variation observed among individuals. Surprisingly, developmental experience also determined the particular way in which individuals plastically adjusted their behaviour and web structure to changes in adult condition.

The evolution of labile traits in sex- and age-structured populations.

Abstract: Many quantitative traits are labile (e.g. somatic growth rate, reproductive timing and investment), varying over the life cycle as a result of behavioural adaptation, developmental processes and plastic responses to the environment. At the population level, selection can alter the distribution of such traits across age classes and among generations. Despite a growing body of theoretical research exploring the evolutionary dynamics of labile traits, a data-driven framework for incorporating such traits into demographic models has not yet been developed. Integral projection models (IPMs) are increasingly being used to understand the interplay between changes in labile characters, life histories and population dynamics. One limitation of the IPM approach is that it relies on phenotypic associations between parents and offspring traits to capture inheritance. However, it is well-established that many different processes may drive these associations, and currently, no clear consensus has emerged on how to model micro-evolutionary dynamics in an IPM framework. We show how to embed quantitative genetic models of inheritance of labile traits into age-structured, two-sex models that resemble standard IPMs. Commonly used statistical tools such as GLMs and their mixed model counterparts can then be used for model parameterization. We illustrate the methodology through development of a simple model of egg-laying date evolution, parameterized using data from a population of Great tits (Parus major). We demonstrate how our framework can be used to project the joint dynamics of species' traits and population density. We then develop a simple extension of the age-structured Price equation (ASPE) for two-sex populations, and apply this to examine the age-specific contributions of different processes to change in the mean phenotype and breeding value. The data-driven framework we outline here has the potential to facilitate greater insight into the nature of selection and its consequences in settings where focal traits vary over the lifetime through ontogeny, behavioural adaptation and phenotypic plasticity, as well as providing a potential bridge between theoretical and empirical studies of labile trait variation.