Showing papers in "Journal of Animal Ecology in 2021"

A 'How-to' Guide for Interpreting Parameters in Habitat-Selection Analyses.

Abstract: Habitat-selection analyses allow researchers to link animals to their environment via habitat-selection or step-selection functions, and are commonly used to address questions related to wildlife management and conservation efforts. Habitat-selection analyses that incorporate movement characteristics, referred to as integrated step-selection analyses, are particularly appealing because they allow modelling of both movement and habitat-selection processes. Despite their popularity, many users struggle with interpreting parameters in habitat-selection and step-selection functions. Integrated step-selection analyses also require several additional steps to translate model parameters into a full-fledged movement model, and the mathematics supporting this approach can be challenging for many to understand. Using simple examples, we demonstrate how weighted distribution theory and the inhomogeneous Poisson point process can facilitate parameter interpretation in habitat-selection analyses. Furthermore, we provide a 'how to' guide illustrating the steps required to implement integrated step-selection analyses using the amt package By providing clear examples with open-source code, we hope to make habitat-selection analyses more understandable and accessible to end users.

The importance of individual-to-society feedbacks in animal ecology and evolution.

Abstract: The social decisions that individuals make-who to interact with and how frequently-gives rise to social structure. The resulting social structure then determines how individuals interact with their surroundings-resources and risks, pathogens and predators, competitors and cooperators. However, despite intensive research on (i) how individuals make social decisions and (ii) how social structure shapes social processes (e.g. cooperation, competition and conflict), there are still few studies linking these two perspectives. These perspectives represent two halves of a feedback loop: individual behaviour scales up to define the social environment, and this environment, in turn, feeds back by shaping the selective agents that drive individual behaviour. We first review well-established research areas that have captured both elements of this feedback loop-host-pathogen dynamics and cultural transmission. We then highlight areas where social structure is well studied but the two perspectives remain largely disconnected. Finally, we synthesise existing research on 14 distinct research topics to identify new prospects where the interplay between social structure and social processes are likely to be important but remain largely unexplored. Our review shows that the inherent links between individuals' traits, their social decisions, social structure, and social evolution, warrant more consideration. By mapping the existing and missing connections among many research areas, our review highlights where explicitly considering social structure and the individual-to-society feedbacks can reveal new dimensions to old questions in ecology and evolution.

Observing the unwatchable: Integrating automated sensing, naturalistic observations and animal social network analysis in the age of big data.

Abstract: In the 4.5 decades since Altmann (1974) published her seminal paper on the methods for the observational study of behaviour, automated detection and analysis of social interaction networks have fundamentally transformed the ways that ecologists study social behaviour. Methodological developments for collecting data remotely on social behaviour involve indirect inference of associations, direct recordings of interactions and machine vision. These recent technological advances are improving the scale and resolution with which we can dissect interactions among animals. They are also revealing new intricacies of animal social interactions at spatial and temporal resolutions as well as in ecological contexts that have been hidden from humans, making the unwatchable seeable. We first outline how these technological applications are permitting researchers to collect exquisitely detailed information with little observer bias. We further recognize new emerging challenges from these new reality-mining approaches. While technological advances in automating data collection and its analysis are moving at an unprecedented rate, we urge ecologists to thoughtfully combine these new tools with classic behavioural and ecological monitoring methods to place our understanding of animal social networks within fundamental biological contexts.

Unifying spatial and social network analysis in disease ecology.

Abstract: Social network analysis has achieved remarkable popularity in disease ecology, and is sometimes carried out without investigating spatial heterogeneity. Many investigations into sociality and disease may nevertheless be subject to cryptic spatial variation, so ignoring spatial processes can limit inference regarding disease dynamics. Disease analyses can gain breadth, power and reliability from incorporating both spatial and social behavioural data. However, the tools for collecting and analysing these data simultaneously can be complex and unintuitive, and it is often unclear when spatial variation must be accounted for. These difficulties contribute to the scarcity of simultaneous spatial-social network analyses in disease ecology thus far. Here, we detail scenarios in disease ecology that benefit from spatial-social analysis. We describe procedures for simultaneous collection of both spatial and social data, and we outline statistical approaches that can control for and estimate spatial-social covariance in disease ecology analyses. We hope disease researchers will expand social network analyses to more often include spatial components and questions. These measures will increase the scope of such analyses, allowing more accurate model estimates, better inference of transmission modes, susceptibility effects and contact scaling patterns, and ultimately more effective disease interventions.

Biologging reveals individual variation in behavioural predictability in the wild.

Abstract: Recent research highlights the ecological importance of individual variation in behavioural predictability. Individuals may not only differ in their average expression of a behavioural trait (their behavioural type) and in their ability to adjust behaviour to changing environmental conditions (individual plasticity), but also in their variability around their average behaviour (predictability). However, quantifying behavioural predictability in the wild has been challenging due to limitations of acquiring sufficient repeated behavioural measures. We here demonstrate how common biologging data can be used to detect individual variation in behavioural predictability in the wild and reveal the coexistence of highly predictable individuals along with unpredictable individuals within the same population. We repeatedly quantified two behaviours-daily movement distance and diurnal activity-in 62 female brown bears Ursus arctos tracked across 187 monitoring years. We calculated behavioural predictability over the short term (50 consecutive monitoring days within 1 year) and long term (across monitoring years) as the residual intra-individual variability (rIIV) of behaviour around the behavioural reaction norm. We tested whether predictability varies systematically across average behavioural types and whether it is correlated across functionally distinct behaviours, that is, daily movement distance and amount of diurnal activity. Brown bears showed individual variation in behavioural predictability from predictable to unpredictable individuals. For example, the standard deviation around the average daily movement distance within one monitoring year varied up to fivefold from 1.1 to 5.5 km across individuals. Individual predictability for both daily movement distance and diurnality was conserved across monitoring years. Individual predictability was correlated with behavioural type where individuals which were on average more diurnal and mobile were also more unpredictable in their behaviour. In contrast, more nocturnal individuals moved less and were more predictable in their behaviour. Finally, individual predictability in daily movement distance and diurnality was positively correlated, suggesting that individual predictability may be a quantitative trait in its own regard that could evolve and is underpinned by genetic variation. Unpredictable individuals may cope better with stochastic events and unpredictability may hence be an adaptive behavioural response to increased predation risk. Coexistence of predictable and unpredictable individuals may therefore ensure adaptable and resilient populations.

A Guide to Pre-Processing High-Throughput Animal Tracking Data

Abstract: 1. Modern, high-throughput animal tracking increasingly yields 'big data' at very fine temporal scales. At these scales, location error can exceed the animal's step size, leading to mis-estimation of behaviours inferred from movement. 'Cleaning' the data to reduce location errors is one of the main ways to deal with position uncertainty. Though data cleaning is widely recommended, inclusive, uniform guidance on this crucial step, and on how to organise the cleaning of massive datasets, is relatively scarce. 2. A pipeline for cleaning massive high-throughput datasets must balance ease of use and computationally efficiency, in which location errors are rejected while preserving valid animal movements. Another useful feature of a pre-processing pipeline is efficiently segmenting and clustering location data for statistical methods, while also being scalable to large datasets and robust to imperfect sampling. Manual methods being prohibitively time consuming, and to boost reproducibility, pre-processing pipelines must be automated. 3. We provide guidance on building pipelines for pre-processing high-throughput animal tracking data to prepare it for subsequent analyses. We apply our proposed pipeline to simulated movement data with location errors, and also show how large volumes of cleaned data can be transformed into biologically meaningful 'residence patches', for exploratory inference on animal space use. We use tracking data from the Wadden Sea ATLAS system (WATLAS) to show how pre-processing improves its quality, and to verify the usefulness of the residence patch method. Finally, with tracks from Egyptian fruit bats (Rousettus aegyptiacus), we demonstrate the pre-processing pipeline and residence patch method in a fully worked out example. 4. To help with fast implementation of standardised methods, we developed the R package atlastools, which we also introduce here. Our pre-processing pipeline and atlastools can be used with any high-throughput animal movement data in which the high data-volume combined with knowledge of the tracked individuals' movement capacity can be used to reduce location errors. atlastools is easy to use for beginners, while providing a template for further development. The common use of simple yet robust pre-processing steps promotes standardised methods in the field of movement ecology and leads to better inferences from data.

Proximity to humans affects local social structure in a giraffe metapopulation.

Abstract: 1. Experimental laboratory evidence suggests that animals with disrupted social systems express weakened relationship strengths and have more exclusive social associations, and that these changes have functional consequences. A key question is whether anthropogenic pressures have a similar impact on the social structure of wild animal communities. 2. We addressed this question by constructing a social network from 6 years of systematically collected photographic capture-recapture data spanning 1,139 individual adult female Masai giraffes inhabiting a large, unfenced, heterogeneous landscape in northern Tanzania. We then used the social network to identify distinct social communities, and tested whether social or anthropogenic and other environmental factors predicted differences in social structure among these communities. 3. We reveal that giraffes have a multilevel social structure. Local preferences in associations among individuals scale up to a number of distinct, but spatially overlapping, social communities, that can be viewed as a large interconnected metapopulation. We then find that communities that are closer to traditional compounds of indigenous Masai people express weaker relationship strengths and the giraffes in these communities are more exclusive in their associations. 4. The patterns we characterise in response to proximity to humans reflect the predictions of disrupted social systems. Near bomas, fuelwood cutting can reduce food resources, and groups of giraffes are more likely to encounter livestock and humans on foot, thus disrupting the social associations among group members. Our results suggest that human presence could potentially be playing an important role in determining the conservation future of this megaherbivore.

Detecting and quantifying social transmission using network-based diffusion analysis.

Abstract: Although social learning capabilities are taxonomically widespread, demonstrating that freely interacting animals (whether wild or captive) rely on social learning has proved remarkably challenging. Network-based diffusion analysis (NBDA) offers a means for detecting social learning using observational data on freely interacting groups. Its core assumption is that if a target behaviour is socially transmitted, then its spread should follow the connections in a social network that reflects social learning opportunities. Here, we provide a comprehensive guide for using NBDA. We first introduce its underlying mathematical framework and present the types of questions that NBDA can address. We then guide researchers through the process of selecting an appropriate social network for their research question; determining which NBDA variant should be used; and incorporating other variables that may impact asocial and social learning. Finally, we discuss how to interpret an NBDA model's output and provide practical recommendations for model selection. Throughout, we highlight extensions to the basic NBDA framework, including incorporation of dynamic networks to capture changes in social relationships during a diffusion and using a multi-network NBDA to estimate information flow across multiple types of social relationship. Alongside this information, we provide worked examples and tutorials demonstrating how to perform analyses using the newly developed NBDA package written in the R programming language.

Intra- and interspecific variation in the responses of insect phenology to climate.

Abstract: Phenological change is the most widely documented biological impact of climate change, but shows marked variation in magnitude among populations and species. Thus, quantifying the environmental factors and organismal differences driving this intra- and interspecific variability in phenology is vital to understand and forecast the ecological consequences of climate change. Here, we test intra- and interspecific differences for a set of butterfly species in the organismal sensitivity of flight phenology and its dependence on environmental factors, using as our model system an elevation gradient in a Mediterranean mountain range where temperature and relative humidity vary substantially over space and time. We use field-collected meteorological data, and butterfly counts for 20 univoltine species over 14 years, to test the relative effects on phenology of temperature and relative humidity, the sensitivity of phenology to spatial and temporal variation in temperature and whether ecological traits account for inter-specific variation in sensitivity. For all species, temperature in the months immediately preceding adult emergence had the strongest relationship with phenology. All species appeared earlier in warmer years, with those flying earlier in the season showing the greatest sensitivity to annual (temporal) variation in temperature. However, only a minority of species showed evidence of plastic, space-for-time responses to temperature. Instead, most species showed strong evidence that phenology was more sensitive to temporal than spatial variation in temperature. Our results support the dominant influence of temperature on phenology, even in Mediterranean environments suffering summer drought. They also suggest that accurate forecasts of species' phenological shifts could require the isolation of spatial from temporal components of temperature variation, because the sensitivity of populations and species may differ across these two dimensions. The factors driving synchronisation of phenology over space merit particular research in the context of climate change, given their potential to expose populations simultaneously to environmental extremes.

Drivers of site fidelity in ungulates

Abstract: 1. While the tendency to return to previously visited locations—termed ‘site fidelity’—is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals’ recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity. 2. We compared inter‐year site fidelity in 669 individuals across eight ungulate species fitted with GPS collars and occupying a range of environmental conditions in North America and Africa. We used a distance‐based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size. 3. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren‐ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a ‘win‐stay, lose‐switch’ strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested. 4. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter‐annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species‐specific differences in attraction to known sites, contribute to variation in the expression of this behaviour. 5. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change.

Personality-driven life history trade-offs differ in two subpopulations of free-ranging predators.

Abstract: Consistent individual differences in behaviour (i.e. personality) can be explained in an evolutionary context if they are favoured by life history trade-offs as conceptualized in the pace-of-life syndrome (POLS) hypothesis. Theory predicts that faster-growing individuals suffer higher mortality and that this trade-off is mediated through exploration/risk-taking personality, but empirical support for this remains limited and ambiguous. Equivocal support to the POLS hypothesis suggests that the link between life history and personality may only emerge under certain circumstances. Understanding personality-driven trade-offs would be facilitated by long-term studies in wild populations experiencing different ecological conditions. Here, we tested whether personality measured in semi-captivity was associated with a growth-mortality trade-off via risk-taking in the wild in two subpopulations of juvenile lemon sharks Negaprion brevirostris known to differ in their predator abundance. We expected stronger personality-driven trade-offs in the predator-rich environment as compared to the predator-poor environment. Sharks were captured yearly from 1995 onwards allowing us to obtain long-term data on growth and apparent survival in each subpopulation. We then used a novel open-field assay to test sharks for exploration personality yearly from 2012 to 2017. A subset of the tested sharks was monitored in the field using telemetry to document risk-taking behaviours. We tested (a) if fast explorers in captivity took more risks and grew faster in the wild and (b) if natural selection acted against more explorative, faster-growing sharks. In the subpopulation with fewer predators, more explorative sharks in captivity took more risks in the wild and grew faster. In turn, larger, fast-growing sharks had lower apparent survival. In the predator-rich subpopulation, despite finding selection on fast growth, we found no link between exploration personality and the growth-mortality trade-off. Our study demonstrates that the association between personality and life history is favoured in some ecological contexts but not in others. We identify predator and resource abundance as two main potential drivers of the personality-mediated trade-off and emphasize that future work on the POLS hypothesis would benefit from an approach integrating behaviour and life history across ecological conditions.

The myriad of complex demographic responses of terrestrial mammals to climate change and gaps of knowledge: A global analysis.

Abstract: Approximately 25% of mammals are currently threatened with extinction, a risk that is amplified under climate change. Species persistence under climate change is determined by the combined effects of climatic factors on multiple demographic rates (survival, development and reproduction), and hence, population dynamics. Thus, to quantify which species and regions on Earth are most vulnerable to climate-driven extinction, a global understanding of how different demographic rates respond to climate is urgently needed. Here, we perform a systematic review of literature on demographic responses to climate, focusing on terrestrial mammals, for which extensive demographic data are available. To assess the full spectrum of responses, we synthesize information from studies that quantitatively link climate to multiple demographic rates. We find only 106 such studies, corresponding to 87 mammal species. These 87 species constitute <1% of all terrestrial mammals. Our synthesis reveals a strong mismatch between the locations of demographic studies and the regions and taxa currently recognized as most vulnerable to climate change. Surprisingly, for most mammals and regions sensitive to climate change, holistic demographic responses to climate remain unknown. At the same time, we reveal that filling this knowledge gap is critical as the effects of climate change will operate via complex demographic mechanisms: a vast majority of mammal populations display projected increases in some demographic rates but declines in others, often depending on the specific environmental context, complicating simple projections of population fates. Assessments of population viability under climate change are in critical need to gather data that account for multiple demographic responses, and coordinated actions to assess demography holistically should be prioritized for mammals and other taxa.

Early-life relationships matter: Social position during early life predicts fitness among female spotted hyenas

Abstract: How social development in early-life affects fitness remains poorly understood Though there is growing evidence that early-life relationships can affect fitness, little research has investigated how social positions develop or whether there are particularly important periods for social position development in an animal's life history In long-lived species in particular, understanding the lasting consequences of early-life social environments requires detailed, long-term datasets Here we used a 25-year dataset to test whether social positions held during early development predicted adult fitness Specifically, we quantified social position using three social network metrics: degree, strength and betweenness We determined the social position of each individual in three types of networks during each of three stages of ontogeny to test whether they predict annual reproductive success (ARS) or longevity among adult female spotted hyenas Crocuta crocuta The social positions occupied by juvenile hyenas did predict their fitness, but the effects of social position on fitness measures differed between stages of early development Network metrics when individuals were young adults better predicted ARS, but network metrics for younger animals, particularly when youngsters were confined to the communal den, better predicted longevity than did metrics assessed during other stages of development Our study shows how multiple types of social bonds formed during multiple stages of social development predict lifetime fitness outcomes We suggest that social bonds formed during specific phases of development may be more important than others when considering fitness outcomes

Wild zebra finches that nest synchronously have long-term stable social ties.

Abstract: Many animals live and breed in colonies, and yet, with just a few exceptions, the value of the social bonds between colony members has rarely been examined. Social ties are important for group coordination at collective tasks, and social coordination can facilitate synchronized reproduction among colony members. Synchronized reproduction in turn can amplify the benefits of coloniality, such as social foraging and predator avoidance. We conducted a field study to investigate whether synchronized reproduction among individuals in replicated colonies is linked to the strength of their social bond, and whether these strong bonds are maintained beyond the reproductive period. We PIT-tagged wild zebra finches (Taeniopygia guttata), monitoring their reproduction and social foraging over two consecutive years. We then used social network analysis to characterize the strength of social bonds among birds in the population. We show that birds that are more synchronized in their reproductive timing (and breed in the same colony) had significantly stronger social ties both during and after reproduction than expected by chance. Our long-term sampling also revealed that the strong social ties between synchronized breeders were carried over across years. Our study reveals a strong correspondence between synchronized breeding and the social structure of the breeding colony. This suggests that the synchrony between pairs is not a simple process based on opportunity, but a mechanism underpinned by more complex sociality, which could be carried over to other behavioural contexts. The maintenance of cross-contextual social ties across years suggests that social structure could have extensive consequences on the overall life history of individuals in addition to playing a key role for the reproductive dynamics of colonial breeders.

Multiple stressors interact to impair the performance of bumblebee Bombus terrestris colonies

Abstract: 1. Bumblebees are constantly exposed to a wide range of biotic and abiotic stresses which they must defend themselves against to survive. Pathogens and pesticides represent important stressors that influence bumblebee health, both when acting alone or in combination. To better understand bumblebee health, we need to investigate how these factors interact, yet experimental studies to date generally focus on only one or two stressors. 2. The aim of this study is to evaluate how combined effects of four important stressors (the gut parasite Nosema ceranae, the neonicotinoid insecticide thiamethoxam, the pyrethroid insecticide cypermethrin, and the EBI‐fungicide tebuconazole) interact to affect bumblebees at the individual and colony levels. 3. We established seven treatment groups of colonies that we pulse exposed to different combinations of these stressors for two weeks under laboratory conditions. Colonies were subsequently placed in the field for seven weeks to evaluate the effect of treatments on the prevalence of N. ceranae in inoculated bumblebees, expression levels of immunity and detoxification‐related genes, food collection, weight gain, worker and male numbers, and production of worker brood and reproductives. 4. Exposure to pesticide mixtures reduced food collection by bumblebees. All immunity‐related genes were upregulated in the bumblebees inoculated with N. ceranae when they had not been exposed to pesticide mixtures, and bumblebees exposed to the fungicide and the pyrethroid were less likely to have N. ceranae. Combined exposure to the three‐pesticides mixture and N. ceranae reduced bumblebee colony growth, and all treatments had detrimental effects on brood production. The groups exposed to the neonicotinoid insecticide produced 40‐76% fewer queens than control colonies. 5. Our findings show that exposure to combinations of stressors that bumblebees frequently come into contact with have detrimental effects on colony health and performance and could therefore have an impact at the population level. These results also have significant implications for current practices and policies for pesticide risk assessment and use as the combinations tested here are frequently applied simultaneously in the field. Understanding the interactions between different stressors will be crucial for improving our ability to manage bee populations and for ensuring pollination services into the future.

Differential survival throughout the full annual cycle of a migratory bird presents a life-history trade-off.

Abstract: Long-distance migrations are among the most physically demanding feats animals perform. Understanding the potential costs and benefits of such behaviour is a fundamental question in ecology and evolution. A hypothetical cost of migration should be outweighed by higher productivity and/or higher annual survival, but few studies on migratory species have been able to directly quantify patterns of survival throughout the full annual cycle and across the majority of a species' range. Here, we use telemetry data from 220 migratory Egyptian vultures Neophron percnopterus, tracked for 3,186 bird months and across approximately 70% of the species' global distribution, to test for differences in survival throughout the annual cycle. We estimated monthly survival probability relative to migration and latitude using a multi-event capture-recapture model in a Bayesian framework that accounted for age, origin, subpopulation and the uncertainty of classifying fates from tracking data. We found lower survival during migration compared to stationary periods (β = -0.816; 95% credible interval: -1.290 to -0.318) and higher survival on non-breeding grounds at southern latitudes (<25°N; β = 0.664; 0.076-1.319) compared to on breeding grounds. Survival was also higher for individuals originating from Western Europe (β = 0.664; 0.110-1.330) as compared to further east in Europe and Asia, and improved with age (β = 0.030; 0.020-0.042). Anthropogenic mortalities accounted for half of the mortalities with a known cause and occurred mainly in northern latitudes. Many juveniles drowned in the Mediterranean Sea on their first autumn migration while there were few confirmed mortalities in the Sahara Desert, indicating that migration barriers are likely species-specific. Our study advances the understanding of important fitness trade-offs associated with long-distance migration. We conclude that there is lower survival associated with migration, but that this may be offset by higher non-breeding survival at lower latitudes. We found more human-caused mortality farther north, and suggest that increasing anthropogenic mortality could disrupt the delicate migration trade-off balance. Research to investigate further potential benefits of migration (e.g. differential productivity across latitudes) could clarify how migration evolved and how migrants may persist in a rapidly changing world.

Strong survival selection on seasonal migration versus residence induced by extreme climatic events

Abstract: 1. Elucidating the full eco-evolutionary consequences of climate change requires quantifying the impact of extreme climatic events (ECEs) on selective landscapes of key phenotypic traits that mediate responses to changing environments. Episodes of strong ECE-induced selection could directly alter population composition, and potentially drive micro-evolution. However, to date, few studies have quantified ECE-induced selection on key traits, meaning that immediate and longer-term eco-evolutionary implications cannot yet be considered. 2. One widely-expressed trait that allows individuals to respond to changing seasonal environments, and directly shapes spatio-seasonal population dynamics, is seasonal migration versus residence. Many populations show considerable among-individual phenotypic variation, resulting in 'partial migration'. However, variation in the magnitude of direct survival selection on migration versus residence has not been rigorously quantified, and empirical evidence of whether seasonal ECEs induce, intensify, weaken or reverse such selection is lacking. 3. We designed full-annual-cycle multi-state capture-recapture models that allow estimation of seasonal survival probabilities of migrants and residents from spatio-temporally heterogeneous individual resightings. We fitted these models to nine years of geographically extensive year-round resighting data from partially migratory European shags (Phalacrocorax aristotelis). We thereby quantified seasonal and annual survival selection on migration versus residence across benign and historically extreme non-breeding season (winter) conditions, and tested whether selection differed between females and males. 4. We show that two of four observed ECEs, defined as severe winter storms causing overall low survival, were associated with very strong seasonal survival selection against residence. These episodes dwarfed the weak selection or neutrality evident otherwise, and hence caused selection through overall annual survival. The ECE that caused highest overall mortality and strongest selection also caused sex-biased mortality, but there was little overall evidence of sex-biased selection on migration versus residence. 5. Our results imply that seasonal ECEs and associated mortality can substantially shape the landscape of survival selection on migration versus residence. Such ECE-induced phenotypic selection will directly alter migrant and resident frequencies, and thereby alter immediate spatio-seasonal population dynamics. Given underlying additive genetic variation, such ECEs could potentially cause micro-evolutionary changes in seasonal migration, and thereby cause complex eco-evolutionary population responses to changing seasonal environments.

Disentangling temporal food web dynamics facilitates understanding of ecosystem functioning.

Abstract: Studying how food web structure and function vary through time represents an opportunity to better comprehend and anticipate ecosystem changes. Yet, temporal studies of highly resolved food web structure are scarce. With few exceptions, most temporal food web studies are either too simplified, preventing a detailed assessment of structural properties or binary, missing the temporal dynamics of energy fluxes among species. Using long-term, multi-trophic biomass data coupled with highly resolved information on species feeding relationships, we analysed food web dynamics in the Gulf of Riga (Baltic Sea) over more than three decades (1981-2014). We combined unweighted (topology-based) and weighted (biomass- and flux-based) food web approaches, first, to unravel how distinct descriptors can highlight differences (or similarities) in food web dynamics through time, and second, to compare temporal dynamics of food web structure and function. We find that food web descriptors vary substantially and distinctively through time, likely reflecting different underlying ecosystem processes. While node- and link-weighted metrics reflect changes related to alterations in species dominance and fluxes, unweighted metrics are more sensitive to changes in species and link richness. Comparing unweighted, topology-based metrics and flux-based functions further indicates that temporal changes in functions cannot be predicted using unweighted food web structure. Rather, information on species population dynamics and weighted, flux-based networks should be included to better comprehend temporal food web dynamics. By integrating unweighted, node- and link-weighted metrics, we here demonstrate how different approaches can be used to compare food web structure and function, and identify complementary patterns of change in temporal food web dynamics, which enables a more complete understanding of the ecological processes at play in ecosystems undergoing change.

Multidimensional trophic niche revealed by complementary approaches: Gut content, digestive enzymes, fatty acids and stable isotopes in Collembola

Abstract: Trophic niche differentiation may explain coexistence and shape functional roles of species. In complex natural food webs, however, trophic niche parameters depicted by single and isolated methods may simplify the multidimensional nature of consumer trophic niches, which includes feeding processes such as food choice, ingestion, digestion, assimilation and retention. Here we explore the correlation and complementarity of trophic niche parameters tackled by four complementary methodological approaches, that is, visual gut content, digestive enzyme, fatty acid and stable isotope analyses-each assessing one or few feeding processes, and demonstrate the power of method combination. Focusing on soil ecosystems, where many omnivore species with cryptic feeding habits coexist, we chose Collembola as an example. We compiled 15 key trophic niche parameters for 125 species from 40 studies. We assessed correlations among trophic niche parameters and described variation of these parameters in different Collembola species, families and across life-forms, which represent microhabitat specialisation. Correlation between trophic niche parameters was weak in 45 out of 64 pairwise comparisons, pointing at complementarity of the four methods. Jointly, the results indicated that fungal- and plant-feeding Collembola assimilate storage, rather than structural polysaccharides, and suggested bacterial feeding as a potential alternative feeding strategy. Gut content and fatty acid analyses suggested alignment between ingestion and assimilation/retention processes in fungal- and plant-feeding Collembola. From the 15 trophic niche parameters, six were related to Collembola family identity, suggesting that not all trophic niche dimensions are phylogenetically structured. Only three parameters were related to the life-forms, suggesting that species use various feeding strategies when living in the same microenvironments. Consumers can meet their nutritional needs by varying their food choices, ingestion and digestion strategies, with the connection among different feeding processes being dependent on the consumed resource and consumer adaptations. Multiple methods reveal different dimensions, together drawing a comprehensive picture of the trophic niche. Future studies applying the multidimensional trophic niche approach will allow us to trace trophic complexity and reveal niche partitioning of omnivorous species and their functional roles, especially in cryptic environments such as soils, caves, deep ocean or benthic ecosystems.

The projected degradation of subtropical coral assemblages by recurrent thermal stress

Abstract: Subtropical coral assemblages are threatened by similar extreme thermal stress events to their tropical counterparts. Yet, the mid- and long-term thermal stress responses of corals in subtropical environments remain largely unquantified, limiting our capacity to predict their future viability. The annual survival, growth and recruitment of 311 individual corals within the Solitary Islands Marine Park (Australia) was recorded over a 3-year period (2016-2018), including the 2015/2016 thermal stress event. These data were used to parameterise integral projection models quantifying the effect of thermal stress within a subtropical coral assemblage. Stochastic simulations were also applied to evaluate the implications of recurrent thermal stress scenarios predicted by four different Representative Concentration Pathways. We report differential shifts in population growth rates (λ) among coral populations during both stress and non-stress periods, confirming contrasting bleaching responses among taxa. However, even during non-stress periods, the observed dynamics for all taxa were unable to maintain current community composition, highlighting the need for external recruitment sources to support the community structure. Across all coral taxa, projected stochastic growth rates (λs ) were found to be lowest under higher emissions scenarios. Correspondingly, predicted increases in recurrent thermal stress regimes may accelerate the loss of coral coverage, species diversity and structural complexity within subtropical regions. We suggest that these trends are primarily due to the susceptibility of subtropical specialists and endemic species, such as Pocillopora aliciae, to thermal stress. Similarly, the viability of many tropical coral populations at higher latitudes is highly dependent on the persistence of up-current tropical systems. As such, the inherent dynamics of subtropical coral populations appear unable to support their future persistence under unprecedented thermal disturbance scenarios.

A citizen science approach reveals long-term social network structure in an urban parrot, Cacatua galerita

Abstract: Parrots are often referenced in discussions of social and cognitive complexity, yet relatively little is known of their social organisation in the wild. In particular, the presence of long-lasting social ties has been highlighted as a hallmark of social complexity, however the presence of such ties can be masked in fission-fusion systems like that exhibited by most parrot species. Social network analysis has the potential to elucidate such multi-level dynamics. While most parrot species are tropical canopy dwellers, a subset have successfully colonised urban habitats, where they are often the focus of much public interest. Our study takes advantage of this to use citizen-science to collect observations of wing-tagged sulphur-crested cockatoos in central Sydney and record their social associations over multiple years. Using a specifically designed mobile phone application "Wingtags", we collected over >27,000 citizen-science reports of wing-tagged cockatoos, and built social networks from spatial-temporal co-occurrences in observations for 130 tagged birds. To validate this novel methodology, we GPS-tagged a subset of wing-tagged birds and compared networks built from both data collection methods. We then examined correlates of social network structure before exploring the temporal dynamics of network structure and social associations. Social networks constructed from GPS data and citizen science data were highly correlated, suggesting that this novel methodology is robust. Network structure exhibited little seasonal variability and was largely driven by roost site choice; however, individuals also showed a surprising degree of mixing between roosts in their foraging associations. Finally, within this larger fission-fusion system, individuals tended to maintain specific social ties for long periods of time. There was an effect of age on these temporal dynamics, with aging individuals increasing both social stability and longevity of associations. Our findings highlight the utility of citizen-science to measure social networks in urban species, and add to the evidence that long-lasting social associations can persist in fission-fusion social systems such as those observed in wild sulphur-crested cockatoos.

Effects of social structure and management on risk of disease establishment in wild pigs.

Abstract: Contact heterogeneity among hosts determines invasion and spreading dynamics of infectious disease, thus its characterization is essential for identifying effective disease control strategies. Yet, little is known about the factors shaping contact networks in many wildlife species and how wildlife management actions might affect contact networks. Wild pigs in North America are an invasive, socially structured species that pose a health concern for domestic swine given their ability to transmit numerous devastating diseases such as African swine fever (ASF). Using proximity loggers and GPS data from 48 wild pigs in Florida and South Carolina, USA, we employed a probabilistic framework to estimate weighted contact networks. We determined the effects of sex, social group and spatial distribution (monthly home-range overlap and distance) on wild pig contact. We also estimated the impacts of management-induced perturbations on contact and inferred their effects on ASF establishment in wild pigs with simulation. Social group membership was the primary factor influencing contacts. Between-group contacts depended primarily on space use characteristics, with fewer contacts among groups separated by >2 km and no contacts among groups >4 km apart within a month. Modelling ASF dynamics on the contact network demonstrated that indirect contacts resulting from baiting (a typical method of attracting wild pigs or game species to a site to enhance recreational hunting) increased the risk of disease establishment by ~33% relative to direct contact. Low-intensity population reduction (<5.9% of the population) had no detectable impact on contact structure but reduced predicted ASF establishment risk relative to no population reduction. We demonstrate an approach for understanding the relative role of spatial, social and individual-level characteristics in shaping contact networks and predicting their effects on disease establishment risk, thus providing insight for optimizing disease control in spatially and socially structured wildlife species.

Sea temperature effects on depth use and habitat selection in a marine fish community

Abstract: Understanding the responses of aquatic animals to temperature variability is essential to predict impacts of future climate change and to inform conservation and management. Most ectotherms such as fish are expected to adjust their behaviour to avoid extreme temperatures and minimize acute changes in body temperature. In coastal Skagerrak, Norway, sea surface temperature (SST) ranges seasonally from 0 to over 20°C, representing a challenge to the fish community which includes cold-, cool- and warm-water affinity species. By acoustically tracking 111 individuals of Atlantic cod Gadus morhua, pollack Pollachius pollachius and ballan wrasse Labrus bergylta in 2015-2018, we examined how coexisting species within a fish community adjusted their behaviour (i.e. vertical distribution in the water column and habitat selection) to cope with the thermal variation. Mixed-effect models showed that thermal preference was a main driver of behaviour and habitat use of the fish community in a southern Norwegian fjord. Cod used colder waters, compared with pollack and ballan wrasse. Increases in SST during summer were associated with the use of deeper, colder waters by cod, especially by larger individuals, and conversely with the occupancy of shallower areas by pollack and ballan wrasse. During winter, when SST dropped and the thermal stratification reversed, pollack and ballan wrasse moved to deeper, relatively warmer areas, while cod selected shallower, colder habitats. Although habitat selection was affected by temperature, species-specific habitat selection was observed even when temperature was similar throughout habitats. This study shows how cohabiting fish species respond to thermal heterogeneity, suggesting that (a) temperature regulates the access to the different depths and habitats and (b) behavioural plasticity may be an important factor for coping with temperature variability and potentially for adaptation to climate change.

Phenological synchronization of seasonal bird migration with vegetation greenness across dietary guilds.

Abstract: The seasonal movement of animals has been linked to seasonal variation in ecological productivity, and it has been hypothesized that primary consumers synchronize migration with vegetation phenology. Within temperate regions of the Northern Hemisphere, herbivorous bird species often track the phenology of vegetation greenness during spring migration. Phenological synchronization with vegetation greenness by migratory birds in other dietary guilds, across the full extent of their annual distributions during both spring and autumn migration, has not been explored. Here, we document population-level associations with a remotely sensed measure of vegetation greenness for 230 North American migratory bird species in seven dietary guilds across the full annual cycle using eBird occurrence information for the combined period 2006-2018. Evidence of phenological synchronization was strongest for omnivores, herbivores, herbivore-granivores and granivores during spring and autumn migration, except for omnivores in the west during spring migration. Strong evidence of synchronization was also observed for insectivores during spring migration and carnivores during spring and autumn migration that migrated across the entire breadth of the continent. The level of evidence declined for insectivores in the west and east during spring migration, and for nectarivores in the west during spring and autumn migration. Limited evidence was also found for insectivores in the east during autumn migration, insectivores in the west and the centre of the continent during spring and autumn migration, and carnivores in the west during spring migration. Carnivores in the west during autumn migration showed the weakest evidence of synchronization. We found broad support across an array of dietary guilds for phenological coupling between vegetation greenness and seasonal bird migration within North America. Our results highlight the potential for many migratory bird species to encounter phenological mismatches as vegetation phenology responds to climate change. Our findings emphasize the need to better understand the environmental cues that regulate migratory behaviour across dietary guilds, consumer levels and migration tactics.

Increased acclimation ability accompanies a thermal niche shift of a recent invasion

Abstract: Globalization is removing dispersal barriers for the establishment of invasive species and enabling their spread to novel climates. New thermal environments in the invaded range will be particularly challenging for ectotherms, as their metabolism directly depends on environmental temperature. However, we know little about the role climatic niche shifts play in the invasion process, and the underlining physiological mechanisms. We tested if a thermal niche shift accompanies an invasion, and if native and introduced populations differ in their ability to acclimate thermal limits. We used an alien ant species-Tapinoma magnum-which recently started to spread across Europe. Using occurrence data and accompanying climatic variables, we measured the amount of overlap between thermal niches in the native and invaded range. We then experimentally tested the acclimation ability in native and introduced populations by incubating T. magnum at 18, 25 and 30°C. We measured upper and lower critical thermal limits after 7 and 21 days. We found that T. magnum occupies a distinct thermal niche in its introduced range, which is on average 3.5°C colder than its native range. Critical thermal minimum did not differ between populations from the two ranges when colonies were maintained at 25 or 30°C, but did differ after colony acclimation at a lower temperature. We found twofold greater acclimation ability of introduced populations to lower temperatures, after prolonged incubation at 18°C. Increased acclimation ability of lower thermal limits could explain the expansion of the realized thermal niche in the invaded range, and likely contributed to the spread of this species to cooler climates. Such thermal plasticity could be an important, yet so far understudied, factor underlying the expansion of invasive insects into novel climates.

Reproductive senescence in a short-lived fish.

Abstract: Reproductive senescence is an age-associated decline in reproductive performance, which often arises as a trade-off between current and future reproduction. Given that mortality is inevitable, increased allocation into current reproduction is favoured despite costs paid later in life. This assumption is violated in organisms with post-maturity growth whose reproductive output increases long after maturity. While reproductive senescence is frequently studied in animals with determinate growth at maturity, such as insects or mammals, we have very limited understanding of reproductive senescence in organisms with an extensive post-maturity growth period. The fact that many post-maturity growers experience strong adult mortality leads to conflicting expectations for reproductive senescence. The aim of this study was to investigate how co-occurrence of rapid life history and post-maturity growth mould reproductive senescence in a short-lived killifish, Nothobranchius furzeri, using longitudinal data on laboratory and wild-type populations. We followed the individual fecundity, fertility and fertilization of 132 singly housed fish from the perspectives of chronological and biological age. At the onset of senescence, the sex-specific contribution to decrease in fertilization capacity was investigated. Allocation trade-offs were estimated through the association between reproductive parameters and life span, and between early-life and late-life fecundity. We demonstrate that female fecundity increased steadily after maturity and reproductive senescence occurred long after the growth asymptote. The prime age for fecundity coincided with 50% female survival and consequent decline in fecundity implies an association with somatic deterioration. Reproductive senescence in fertilization rate was stronger in females than in males. Females with high early fecundity experienced a long life span and high late-life fecundity, discounting the role of allocation trade-offs in reproductive senescence. The present study reports a clear case of reproductive senescence in a fish with a long post-maturation growth period, unusually rapid development and short life span. The onset of reproductive senescence was postponed compared to animals that cease growing at sexual maturity. Fish and other animals with post-maturity growth have long been considered insusceptible to ageing but this conclusion may be related to the previous lack of longitudinal data rather than to the absence of reproductive senescence in such organisms.

Meerkat helpers buffer the detrimental effects of adverse environmental conditions on fecundity, growth and survival

Abstract: Recent comparative studies show that cooperative breeding is positively correlated with harsh and unpredictable environments and it is suggested that this association occurs because helpers buffer the negative effects of adverse ecological conditions on fitness. In the Kalahari, rainfall varies widely between- and within years, affecting primary production and the availability of the principal prey of cooperatively breeding Kalahari meerkats, Suricata suricatta. Our study aimed to establish whether the presence and number of helpers buffer the negative effects of variation in rainfall on the fecundity and body mass of breeding females, and the survival and growth of pups. We investigate the relationship between group size and variation in rainfall on dominant female fecundity, body mass, and offspring survival and growth using an additive modelling approach on 21 years of individual-based records of the life histories of individual meerkats. We show that breeding female fecundity is reduced during periods of low rainfall but that the effects of low rainfall are mitigated by increases in group size and body mass because heavier females and those in larger groups have increased fecundity and reduced interbirth intervals. Pup growth and survival are also reduced during periods of low rainfall, but only in smaller groups. Our results support the suggestion that cooperative breeding mitigates the detrimental effects of adverse environmental conditions and may enhance the capacity of species to occupy environments where food availability is low and unpredictable.

Effects of climate-change-driven gradual and acute temperature changes on shark and ray species

Abstract: Climate change is altering distributions and abundances of marine species through both gradual and acute changes in temperature and productivity. Due to their high mobility and metabolic rates, elasmobranchs (sharks and rays) are likely to redistribute across latitudes and depths as they thermoregulate, but little is known about their responses to these climatic changes, which could vary widely across this diverse group of species. Here, we assessed how species with differing mobility and ecology responded to gradual changes in daily sea surface temperature (SST) and acute temperature anomalies, caused by the El Nino-Southern Oscillation (ENSO), at Cocos Island, Costa Rica, the site of multiple marine heatwaves. We used generalized linear mixed models to analyse 34,342 records of relative abundance or frequency of occurrence for seven shark and ray species collected in 27 years (1993-2019) by a dive company. We compared effect sizes for SST and the Oceanic Nino Index across the different species, which vary widely in body size and mobility. Large, mobile species responded strongly but inconsistently to temperature. For scalloped hammerhead sharks Sphyrna lewini, a 1℃ rise in SST reduced counts by over 14%, and dropped the occurrence of their large schools by almost one-fifth (19.4%). Mobula ray occurrence also declined substantially with a few degrees rise in SST, whereas tiger shark Galeocerdo cuvier occurrence sharply increased. These species also had divergent responses to the ENSO: S. lewini and G. cuvier were sighted with greater frequency during La Nina events, and their abundance dropped considerably during El Nino events-over a twofold decline between a strong La Nina and strong El Nino for S. lewini. In contrast, Mobula rays showed little response to ENSO. The smaller and sedentary Triaenodon obesus exhibited the weakest response of all species to both SST and the ENSO, reflecting its lower metabolic rates and mobility. Climate change will continue to impact elasmobranchs, even for smaller and more localized species, with the potential to impact the effectiveness of marine protected areas (MPAs). Our results compel further work on the diversity of elasmobranch responses to environmental change.

Sharks surf the slope: Current updrafts reduce energy expenditure for aggregating marine predators.

Abstract: An animal's energy landscape considers the power requirements associated with residing in or moving through habitats. Within marine environments, these landscapes can be dynamic as water currents will influence animal power requirements and can change rapidly over diel and tidal cycles. In channels and along slopes with strong currents, updraft zones may reduce energy expenditure of negatively buoyant fishes that are also obligate swimmers. Despite marine predators often residing within high-current area, no study has investigated the potential role of the energetic landscape in driving such habitat selectivity. Over 500 grey reef sharks Carcharhinus amblyrhynchos reside in the southern channel of Fakarava Atoll, French Polynesia. We used diver observations, acoustic telemetry and biologging to show that sharks use regions of predicted updrafts and switch their core area of space use based on tidal state (incoming versus outgoing). During incoming tides, sharks form tight groups and display shuttling behaviour (moving to the front of the group and letting the current move them to the back) to maintain themselves in these potential updraft zones. During outgoing tides, group dispersion increases, swimming depths decrease and shuttling behaviours cease. These changes are likely due to shifts in the nature and location of the updraft zones, as well as turbulence during outgoing tides. Using a biomechanical model, we estimate that routine metabolic rates for sharks may be reduced by 10%-15% when in updraft zones. Grey reef sharks save energy using predicted updraft zones in channels and 'surfing the slope'. Analogous to birds using wind-driven updraft zones, negatively buoyant marine animals may use current-induced updraft zones to reduce energy expenditure. Updrafts should be incorporated into dynamic energy landscapes and may partially explain the distribution, behaviour and potentially abundance of marine predators.

Local prey shortages drive foraging costs and breeding success in a declining seabird, the Atlantic puffin

Abstract: As more and more species face anthropogenic threats, understanding the causes of population declines in vulnerable taxa is essential. However, long-term datasets, ideal to identify lasting or indirect effects on fitness measures such as those caused by environmental factors, are not always available. Here we use a single year but multi-population approach on populations with contrasting demographic trends to identify possible drivers and mechanisms of seabird population changes in the north-east Atlantic, using the Atlantic puffin, a declining species, as a model system. We combine miniature GPS trackers with camera traps and DNA metabarcoding techniques on four populations across the puffins' main breeding range to provide the most comprehensive study of the species' foraging ecology to date. We find that puffins use a dual foraging tactic combining short and long foraging trips in all four populations, but declining populations in southern Iceland and north-west Norway have much greater foraging ranges, which require more (costly) flight, as well as lower chick-provisioning frequencies, and a more diverse but likely less energy-dense diet, than stable populations in northern Iceland and Wales. Together, our findings suggest that the poor productivity of declining puffin populations in the north-east Atlantic is driven by breeding adults being forced to forage far from the colony, presumably because of low prey availability near colonies, possibly amplified by intraspecific competition. Our results provide valuable information for the conservation of this and other important North-Atlantic species and highlight the potential of multi-population approaches to answer important questions about the ecological drivers of population trends.