Showing papers in "Journal of Animal Ecology in 2023"

Quantifying the movement, behaviour and environmental context of group-living animals using drones and computer vision.

Abstract: Methods for collecting animal behaviour data in natural environments, such as direct observation and biologging, are typically limited in spatiotemporal resolution, the number of animals that can be observed and information about animals' social and physical environments. Video imagery can capture rich information about animals and their environments, but image-based approaches are often impractical due to the challenges of processing large and complex multi-image datasets and transforming resulting data, such as animals' locations, into geographical coordinates. We demonstrate a new system for studying behaviour in the wild that uses drone-recorded videos and computer vision approaches to automatically track the location and body posture of free-roaming animals in georeferenced coordinates with high spatiotemporal resolution embedded in contemporaneous 3D landscape models of the surrounding area. We provide two worked examples in which we apply this approach to videos of gelada monkeys and multiple species of group-living African ungulates. We demonstrate how to track multiple animals simultaneously, classify individuals by species and age-sex class, estimate individuals' body postures (poses) and extract environmental features, including topography of the landscape and animal trails. By quantifying animal movement and posture while reconstructing a detailed 3D model of the landscape, our approach opens the door to studying the sensory ecology and decision-making of animals within their natural physical and social environments.

Vegetation structure from LiDAR explains the local richness of birds across Denmark.

Abstract: Classic ecological research into the determinants of biodiversity patterns emphasised the important role of three-dimensional (3D) vegetation heterogeneity. Yet, measuring vegetation structure across large areas has historically been difficult. A growing focus on large-scale research questions has caused local vegetation heterogeneity to be overlooked compared with more readily accessible habitat metrics from, for example, land cover maps. Using newly available 3D vegetation data, we investigated the relative importance of habitat and vegetation heterogeneity for explaining patterns of bird species richness and composition across Denmark (42,394 km2 ). We used standardised, repeated point counts of birds conducted by volunteers across Denmark alongside metrics of habitat availability from land-cover maps and vegetation structure from rasterised LiDAR data (10 m resolution). We used random forest models to relate species richness to environmental features and considered trait-specific responses by grouping species by nesting behaviour, habitat preference and primary lifestyle. Finally, we evaluated the role of habitat and vegetation heterogeneity metrics in explaining local bird assemblage composition. Overall, vegetation structure was equally as important as habitat availability for explaining bird richness patterns. However, we did not find a consistent positive relationship between species richness and habitat or vegetation heterogeneity; instead, functional groups displayed individual responses to habitat features. Meanwhile, habitat availability had the strongest correlation with the patterns of bird assemblage composition. Our results show how LiDAR and land cover data complement one another to provide insights into different facets of biodiversity patterns and demonstrate the potential of combining remote sensing and structured citizen science programmes for biodiversity research. With the growing coverage of LiDAR surveys, we are witnessing a revolution of highly detailed 3D data that will allow us to integrate vegetation heterogeneity into studies at large spatial extents and advance our understanding of species' physical niches.

Raising offspring increases ageing: Differences in senescence among three populations of a long-lived seabird, the Atlantic puffin.

Abstract: Actuarial senescence, the decline of survival with age, is well documented in the wild. Rates of senescence vary widely between taxa, to some extent also between sexes, with the fastest life histories showing the highest rates of senescence. Few studies have investigated differences in senescence among populations of the same species, although such variation is expected from population-level differences in environmental conditions, leading to differences in vital rates and thus life histories. We predict that, within species, populations differing in productivity (suggesting different paces of life) should experience different rates of senescence, but with little or no sexual difference in senescence within populations of monogamous, monomorphic species where the sexes share breeding duties. We compared rates of actuarial senescence among three contrasting populations of the Atlantic puffin Fratercula arctica. The data set comprised 31 years (1990-2020) of parallel capture-mark-recapture data from three breeding colonies, Isle of May (North Sea), Røst (Norwegian Sea) and Hornøya (Barents Sea), showing contrasting productivities (i.e. annual breeding success) and population trends. We used time elapsed since first capture (TFC) as a proxy for bird age, and productivity and the winter North Atlantic Oscillation Index (wNAO) as proxies for the environmental conditions experienced by the populations within and outside the breeding season, respectively. In accordance with our predictions, we found that senescence rates differed among the study populations, with no evidence for sexual differences. There was no evidence for an effect of wNAO, but the population with the lowest productivity, Røst, showed the lowest rate of senescence. As a consequence, the negative effect of senescence on the population growth rate (λ) was up to 3-5 times smaller on Røst (Δλ = -0.009) than on the two other colonies. Our findings suggest that environmentally induced differences in senescence rates among populations of a species should be accounted for when predicting effects of climate variation and change on species persistence. There is thus a need for more detailed information on how both actuarial and reproductive senescence influence vital rates of populations of the same species, calling for large-scale comparative studies.

Correction to: Sexual selection and personality: Individual and group-level effects on mating behaviour in red junglefowl.

Abstract: 1Edward Grey Institute, Department of Zoology, University of Oxford, Oxford, UK; 2Department of Biology, University of Florida, Gainesville, FL, USA; 3Behavioural Ecology, Department of Biology, Ludwig Maximilians University of Munich, Munich, Germany; 4Evolution and Ecology Research Centre and School of Biological and Environmental Sciences, University of New South Wales, Sydney, NSW, Australia; 5Department of Ecology, University of Veterinary Medicine Budapest, Budapest, Hungary; 6Department of Physics Chemistry and Biology, IFM Biology, Linköping University, Linköping, Sweden and 7School of Biological Sciences, Monash University, Melbourne, Vic., Australia

Stratification and recovery time jointly shape ant functional reassembly in a Neotropical forest.

Abstract: 1. Microhabitat differentiation of species communities such as vertical stratification in tropical forests contributes to species coexistence and thus biodiversity. However, little is known about how the extent of stratification changes during forest recovery and influences community reassembly. Environmental filtering determines community reassembly in time (succession) and in space (stratification), hence functional and phylogenetic composition of species communities are highly dynamic. It is poorly understood if and how these two concurrent filters - forest recovery and stratification - interact. 2. In a tropical forest chronosequence in Ecuador spanning 34 years of natural recovery, we investigated the recovery trajectory of ant communities in three overlapping strata (ground, leaf litter, lower tree trunk) by quantifying 13 traits, as well as the functional and phylogenetic diversity of the ants. We expected that functional and phylogenetic diversity would increase with recovery time and that each ant community within each stratum would show a distinct functional reassembly. We predicted that traits related to ant diet would show divergent trajectories reflecting an increase in niche differentiation with recovery time. On the other hand, traits related to the abiotic environment were predicted to show convergent trajectories due to a more similar microclimate across strata with increasing recovery age. 3. Most of the functional traits and the phylogenetic diversity of the ants were clearly stratified, confirming previous findings. However, neither functional nor phylogenetic diversity increased with recovery time. Community-weighted trait means had complex relationships to recovery time and the majority were shaped by a statistical interaction between recovery time and stratum, confirming our expectations. However, most trait trajectories converged among strata with increasing recovery time regardless of whether they were related to ant diet or environmental conditions. 4. We confirm the hypothesized interaction among environmental filters during the functional reassembly in tropical forests. Communities in individual strata respond differently to recovery, and possible filter mechanisms likely arise from both abiotic (e.g., microclimate) and biotic (e.g., diet) conditions. Since vertical stratification is prevalent across animal and plant taxa, our results highlight the importance of stratum-specific analysis in dynamic ecosystems and may generalize beyond ants.

The influence of human activity on predator-prey spatiotemporal overlap.

Abstract: Despite growing evidence of widespread impacts of humans on animal behavior, our understanding of how humans reshape species interactions remains limited. Here, we present a framework that draws on key concepts from behavioral and community ecology to outline four primary pathways by which humans can alter predator-prey spatiotemporal overlap. We suggest that predator-prey dyads can exhibit similar or opposite responses to human activity with distinct outcomes for predator diet, predation rates, population demography, and trophic cascades. We demonstrate how to assess these behavioral response pathways with hypothesis testing, using temporal activity data for 178 predator-prey dyads from published camera trap studies on terrestrial mammals. We found evidence for each of the proposed pathways, revealing multiple patterns of human influence on predator-prey activity and overlap. Our framework and case study highlight current challenges, gaps, and advances in linking human activity to animal behavior change and predator-prey dynamics. By using a hypothesis-driven approach to estimate the potential for altered species interactions, researchers can anticipate the ecological consequences of human activities on whole communities.

Effect of predation risk on parasite transmission from first to second intermediate trematode hosts.

Abstract: 1. Predators can affect parasite-host interactions when directly preying on hosts or their parasites. However, predators may also have non-consumptive indirect effects on parasite-host interactions when hosts adjust their behaviour or physiology in response to predator presence. 2. In this study, we examined how chemical cues from a predatory marine crab affect the transmission of a parasitic trematode from its first (periwinkle) to its second (mussel) intermediate host. 3. Laboratory experiments revealed that chemical cues from crabs lead to a three-fold increase in the release of trematode cercariae from periwinkles as a result of increased periwinkle activity. This positive effect on transmission was contrasted by a tenfold reduction in cercarial infection rates in the second intermediate host when we experimentally exposed mussels to cercariae and predator cues. The low infection rates were caused by a substantial reduction in mussel filtration activity in presence of predator cues, preventing cercariae from entering the mussels. To assess the combined net effect of both processes, we conducted a transmission experiment between infected periwinkles and uninfected mussels. Infection levels of mussels in the treatments with crab cues were 7-fold lower than in mussels without crab chemical cues. This suggests that predation risk effects on mussel susceptibility can counteract the elevated parasite release from first intermediate hosts, with negative net effects on parasite transmission. 4. These experiments highlight that predation risk effects on parasite transmission can have opposing directions at different stages of the parasite's life cycle. Such complex non-consumptive predation risk effects on parasite transmission may constitute an important indirect mechanism affecting prevalence and distribution patterns of parasites in different hosts across their life cycle.

Ecological patterns and processes in the vertical dimension of terrestrial ecosystems.

Abstract: Climatic gradients such as latitude and elevation are considered primary drivers of global biogeography. Yet, alongside these macro-gradients, the vertical space and structure generated by terrestrial plants form comparable climatic gradients but at a fraction of the distance. These vertical gradients provide a spectrum of ecological space for species to occur and co-exist, increasing biodiversity. Further, vertical gradients can serve as pathways for evolutionary adaptation of species traits, leading to a range of ecological specialisations. In this review, we explore the ecological evidence supporting the proposition that the vertical gradient serves as an engine driving the ecology and evolution of species and shaping larger biogeographic patterns in space and time akin to elevation and latitude. Focusing on vertebrate and invertebrate taxa, we synthesised how ecological patterns within the vertical dimension shape species composition, distribution, and biotic interactions. We identify three key ecological mechanisms associated with species traits that facilitate persistence within the vertical environment and draw on empirical examples from the literature to explore these processes. Looking forward, we propose that the vertical dimension provides an excellent study template to explore timely ecological and evolutionary questions. We encourage future research to also consider how the vertical dimension will influence the resilience and response of animal taxa to global change.

Predation cues amplify the effects of parasites on the personality of a keystone grazer.

Abstract: Parasites can alter species interactions either by modifying infected host behavior or by influencing behavioral responses in uninfected individuals. Salt marsh ecosystems are characterized by a predator-prey interaction between the keystone grazer, Littoraria irrorata, and its main predator, Callinectes sapidus, both integral players in mediating the productivity of these habitats. Littoraria also acts as the first intermediate host for at least four species of digenetic trematode. Parasite infection has been shown to decrease grazing and climbing in populations of Littoraria, though effects on infected host response to predators has not been investigated. Moreover, how infection might increase or decrease among-individual variation in behavior (i.e., animal personality) is still unknown. Here we ask how trematode infection affects the expression of boldness in the antipredator responses of L. irrorata in both the absence and presence of a predator cue. We find that individual boldness varies substantially, and repeatability tends to increase as the number of stressors increases, with infected individuals exposed to a predator cue showing the strongest expression of behavioral types. Parasitism amplifies this effect, though the parasite itself does not appear to directly induce behavioral changes: infected snails show no evidence of decreased climbing or differences in refuge use as compared to their uninfected counterparts. Infection might therefore drive the expression of condition-dependent personality differences evident only under high-risk conditions. Group infection status strongly influenced behavioral reaction norms: uninfected individuals grouped with an infected snail were more responsive to predation risk, exhibiting increased climbing behavior and spending less time in the water. Here parasites are influencing personality indirectly by inducing avoidance behaviors in healthy individuals, though only in high-risk environments. The potential for exposure to parasites and predators fluctuates greatly across marsh ecosystems. Given the ecological importance of this predator-prey relationship, trematode infection can act as an important, though indirect, determinant of overall salt marsh community structure, health, and function.

Decoupling pioneering traits from latitudinal patterns in a North American bird experiencing a southward range shift.

Abstract: Eco-geographic rules describe spatial patterns in biological trait variation and shed light on the drivers of such variation. In animals, a consensus is emerging that 'pioneering' traits may facilitate range shifts via a set of bold, aggressive, and stress-resilient traits. Many of these same traits are associated with more northern latitudes, and most range shifts in the northern hemisphere indicate northward movement. As a consequence, it is unclear whether pioneering traits are simply corollaries of existing latitudinal variation, or whether they override other well-trodden latitudinal patterning as a unique eco-geographic rule of phenotypic variation. The tree swallow (Tachycineta bicolor) is a songbird undergoing a southward range shift in the eastern United States, in direct opposition of the poleward movement seen in most other native species' range shifts. Because this organic range shift countervails the typical direction of movement, this case study provides for unique ecological insights on organisms and their ability to thrive in our changing world. We sampled female birds across seven populations, quantifying behavioral, physiological, and morphological traits. We also used GIS and field data to quantify a core set of ecological factors with strong ties to these traits as well as female performance. Females at more southern expansion sites displayed higher maternal aggression, higher baseline corticosterone, and more pronounced elevation of corticosterone following a standardized stressor, contrary to otherwise largely conserved latitudinal patterning in these traits. Microhabitat variation explained some quantitative phenotypic variation, but the expansion and historic ranges did not differ in openness, distance to water, or breeding density. This countervailing range shift therefore suggests that pioneering traits are not simply corollaries of existing latitudinal variation, but rather, they may override other well-trodden latitudinal patterning as a unique eco-geographic rule of phenotypic variation.

Population connectivity patterns of genetic diversity, immune responses, and exposure to infectious pneumonia in a metapopulation of desert bighorn sheep.

Abstract: Habitat fragmentation is an important driver of biodiversity loss and can be remediated through management actions aimed at maintenance of natural connectivity in metapopulations. Connectivity may protect populations from infectious diseases by preserving immunogenetic diversity and disease resistance. However, connectivity could exacerbate the risk of infectious disease spread across vulnerable populations. We tracked the spread of a novel strain of Mycoplasma ovipneumoniae in a metapopulation of desert bighorn sheep (Ovis canadensis nelsoni) in the Mojave Desert to investigate how variation in connectivity among populations influenced disease outcomes. M. ovipneumoniae was detected throughout the metapopulation, indicating that the relative isolation of many of these populations did not protect them from pathogen invasion. However, we show that connectivity among bighorn sheep populations was correlated with higher immunogenetic diversity, a protective immune response, and lower disease prevalence. Variation in protective immunity predicted infection risk in individual bighorn sheep and was associated with heterozygosity at genetic loci linked to adaptive and innate immune signaling. Together, these findings may indicate that population connectivity maintains immunogenetic diversity in bighorn sheep populations in this system and has direct effects on immune responses in individual bighorn sheep and their susceptibility to infection by a deadly pathogen. Our study suggests that the genetic benefits of population connectivity could outweigh the risk of infectious disease spread and supports conservation management that maintains natural connectivity in metapopulations.

The importance of a holistic approach to the factors determining population abundances.

Abstract: Research Highlight: Ogilvie, J. E., & CaraDonna, P. J. (2022). The shifting importance of abiotic and biotic factors across the life cycles of wild pollinators. Journal of Animal Ecology, 91, 2412-2423. https://doi.org/10.1111/1365-2656.13825. As global change and its multiple impacts continue to unfold across most of the planet, understanding how populations of wild species respond to changing conditions has become a major focus of ecological studies. Ogilvie and CaraDonna (Ogilvie & CaraDonna, 2022) focus on understanding how biotic and abiotic conditions affect bumblebee abundances. A major advance in their work is that, rather than focusing on a single measure of abundance at a particular life stage for each of the seven bumblebee species they survey (e.g. adult abundance), they focus on understanding the drivers of population abundance across the different stages of the species' life cycles. The authors specifically assess how three factors in particular, climate conditions, floral resource availability and previous life-stage abundances impact these abundances. A main finding in their study is that each of these three factors directly impacted a different life stage, showing that just focusing on a single life-stage would have resulted on a biased and incomplete picture of how abiotic and biotic factors affect bumblebee population dynamics. Studies like this one emphasize the need to focus on understanding the demographic mechanisms that determine population abundances.

Human land-uses homogenize stream assemblages and reduce animal biomass production.

Abstract: 1. Human land-use change is a major threat to natural ecosystems worldwide. Nonetheless, the effects of human land-uses on the structure of plant and animal assemblages and their functional characteristics need to be better understood. Furthermore, the pathways by which human land uses affect ecosystem functions, such as biomass production, still need to be clarified. 2. We compiled a unique dataset of fish, arthropod and macrophyte assemblages from 61 stream ecosystems in two Neotropical biomes: Amazonian rainforest and Uruguayan grasslands. We then tested how the cover of agriculture, pasture, urbanization and afforestation affected the taxonomic richness and functional diversity of those three species assemblages, and the consequences of these effects for animal biomass production. Single trait categories and functional diversity were evaluated, combining recruitment and life-history, resource and habitat-use, and body size. 3.The effects of intensive human land-uses on taxonomic and functional diversities were as strong as other drivers known to affect biodiversity, such as local climate and environmental factors. In both biomes, the taxonomic richness and functional diversity of animal and macrophyte assemblages decreased with increasing cover of agriculture, pasture, and urbanization. Human land-uses were associated with functional homogenization of both animal and macrophyte assemblages. Human land-uses reduced animal biomass through direct and indirect pathways mediated by declines in taxonomic and functional diversities. 4. Our findings indicate that converting natural ecosystems to supply human demands results in species loss and trait homogenization across multiple biotic assemblages, ultimately reducing animal biomass production in streams.

Arthropod food webs predicted from body size ratios are improved by incorporating prey defensive properties.

Abstract: Trophic interactions are often deduced from body size differences, assuming that predators prefer prey smaller than themselves because larger prey are more difficult to subdue. This has mainly been confirmed in aquatic ecosystems, but rarely in terrestrial ecosystems, especially in arthropods. Our goal was to validate whether body size ratios can predict trophic interactions in a terrestrial, plant-associated arthropod community and whether predator hunting strategy and prey taxonomy could explain additional variation. We conducted feeding trials with arthropods from marram grass in coastal dunes to test whether two individuals, of the same or different species, would predate each other. From the trial results, we constructed one of the most complete, empirically derived food webs for terrestrial arthropods associated with a single plant species. We contrasted this empirical food web with a theoretical web based on body size ratios, activity period, microhabitat, and expert knowledge. In our feeding trials, predator-prey interactions were indeed largely size-based. Moreover, the theoretical and empirically based food webs converged well for both predator and prey species. However, predator hunting strategy, and especially prey taxonomy improved predictions of predation. Well-defended taxa, such as hard-bodied beetles, were less frequently consumed than expected based on their body size. For instance, a beetle of average size (measuring 4 mm) is 38% less vulnerable than another average arthropod with the same length. Body size ratios predict trophic interactions among plant-associated arthropods fairly well. However, traits such as hunting strategy and anti-predator defences can explain why certain trophic interactions do not adhere to size-based rules. Feeding trials can generate insights into multiple traits underlying real-life trophic interactions among arthropods.

Six decades of North American bird banding records reveal plasticity in migration phenology.

Abstract: The timing of avian migration has evolved to exploit critical seasonal resources, yet plasticity within phenological responses may allow adjustments to interannual resource phenology. The diversity of migratory species and changes in underlying resources in response to climate change make it challenging to generalize these relationships. We use bird banding records during spring and fall migration from across North America to examine macroscale phenological responses to interannual fluctuations in temperature and long-term annual trends in phenology. In total, we examine 19 species of North American wood warblers (family Parulidae), summarizing migration timing from 2,826,588 banded birds from 1961 to 2018 across 46 sites during spring and 124 sites during fall. During spring, warmer spring temperatures at banding locations translated to earlier median passage dates for 16 of 19 species, with an average 0.65-day advancement in median passage for every 1°C increase in temperature, ranging from 0.25 to 1.26 days °C-1 . During the fall, relationships were considerably weaker, with only 3 of 19 species showing a relationship with temperature. In those three cases, later departure dates were associated with warmer fall periods. Projecting these trends forward under climate scenarios of temperature change, we forecast continued spring advancements under shared socioeconomic pathways from 2041-2060 and 2081-2100 and more muted and variable shifts for fall. These results demonstrate the capacity of long-distance migrants to respond to interannual fluctuations in temperatures, at least during the spring, and showcase the potential of North American bird banding data understanding phenological trends across a wide diversity of avian species.

Fencing amplifies individual differences in movement with implications on survival for two migratory ungulates.

Abstract: Fences have recently been recognized as one of the most prominent linear infrastructures on earth. As animals traverse fenced landscapes, they adjust movement behaviors to optimize resource access while minimizing energetic costs of coping with fences. Examining individual responses is key for connecting localized fence effects with population dynamics. We investigated the multi-scale effects of fencing on animal movements, space use, and survival of 61 pronghorn and 96 mule deer on a gradient of fence density in Wyoming, USA. Taking advantage of the recently developed Barrier Behavior Analysis, we classified individual movement responses upon encountering fences (i.e. barrier behaviors). We adopted the reaction norm framework to jointly quantify individual plasticity and behavioral types of barrier behaviors, as well as behavior syndromes between barrier behaviors and animal space use. We also assessed whether barrier behaviors affect individual survival. Our results highlighted a high level individual plasticity encompassing differences in the degree and direction of barrier behaviors for both pronghorn and mule deer. Additionally, these individual differences were greater at higher fence densities. For mule deer, fence density determined the correlation between barrier behaviors and space use, and was negatively associated with individual survival. Yet, these relationships were not statistically significant for pronghorn. By integrating approaches from movement ecology and behavioral ecology with the emerging field of fence ecology, this study provides new evidence that an extraordinarily widespread linear infrastructure uniquely impacts animals at the individual level. Managing landscape for lower fence densities may help prevent irreversible behavioral shifts for wide-ranging animals in fenced landscapes.

Interactions between fitness components across the life cycle constrain competitor coexistence.

Abstract: Numerous mechanisms can promote competitor coexistence. Yet, these mechanisms are often considered in isolation from one another. Consequently, whether multiple mechanisms shaping coexistence combine to promote or constrain species coexistence remains an open question. Here, we aim to understand how multiple mechanisms interact within and between life stages to determine frequency-dependent population growth, which has a key role stabilizing local competitor coexistence. We conducted field experiments in three lakes manipulating relative frequencies of two Enallagma damselfly species to evaluate demographic contributions of three mechanisms affecting different fitness components across the life cycle: the effect of resource competition on individual growth rate, predation shaping mortality rates, and mating harassment determining fecundity. We then used a demographic model that incorporates carry-over effects between life stages to decompose the relative effect of each fitness component generating frequency-dependent population growth. This decomposition showed that fitness components combined to increase population growth rates for one species when rare, but they combined to decrease population growth rates for the other species when rare, leading to predicted exclusion in most lakes. Because interactions between fitness components within and between life stages vary among populations, these results show that local coexistence is population specific. Moreover, we show that multiple mechanisms do not necessarily increase competitor coexistence, as they can also combine to yield exclusion. Identifying coexistence mechanisms in other systems will require greater focus on determining contributions of different fitness components across the life cycle shaping competitor coexistence in a way that captures the potential for population-level variation.

Host, environment, and anthropogenic factors drive landscape dynamics of an environmentally transmitted pathogen: Sarcoptic mange in the bare-nosed wombat.

Abstract: Understanding the spatial dynamics and drivers of wildlife pathogens is constrained by sampling logistics, with implications for advancing the field of landscape epidemiology and targeted allocation of management resources. However, visually apparent wildlife diseases, when combined with remote-surveillance and distribution modelling technologies, present an opportunity to overcome this landscape-scale problem. Here, we investigated dynamics and drivers of landscape-scale wildlife disease, using clinical signs of sarcoptic mange (caused by Sarcoptes scabiei) in its bare-nosed wombat (BNW; Vombatus ursinus) host. We used 53,089 camera-trap observations from over 3261 locations across the 68,401 km2 area of Tasmania, Australia, combined with landscape data and ensemble species distribution modelling (SDM). We investigated: (1) landscape variables predicted to drive habitat suitability of the host; (2) host and landscape variables associated with clinical signs of disease in the host; and (3) predicted locations and environmental conditions at greatest risk of disease occurrence, including some Bass Strait islands where BNW translocations are proposed. We showed that the Tasmanian landscape, and ecosystems therein, are nearly ubiquitously suited to BNWs. Only high mean annual precipitation reduced habitat suitability for the host. In contrast, clinical signs of sarcoptic mange disease in BNWs were widespread, but heterogeneously distributed across the landscape. Mange (which is environmentally transmitted in BNWs) was most likely to be observed in areas of increased host habitat suitability, lower annual precipitation, near sources of freshwater and where topographic roughness was minimal (e.g. human modified landscapes, such as farmland and intensive land-use areas, shrub and grass lands). Thus, a confluence of host, environmental and anthropogenic variables appear to influence the risk of environmental transmission of S. scabiei. We identified that the Bass Strait Islands are highly suitable for BNWs and predicted a mix of high and low suitability for the pathogen. This study is the largest spatial assessment of sarcoptic mange in any host species, and advances understanding of the landscape epidemiology of environmentally transmitted S. scabiei. This research illustrates how host-pathogen co-suitability can be useful for allocating management resources in the landscape.

Hurricanes affect diversification among individual life courses of a primate population.

Abstract: Extreme climatic events may influence individual-level variability in phenotypes, survival and reproduction, and thereby drive the pace of evolution. Climate models predict increases in the frequency of intense hurricanes, but no study has measured their impact on individual life courses within animal populations. We used 45 years of demographic data of rhesus macaques to quantify the influence of major hurricanes on reproductive life courses using multiple metrics of dynamic heterogeneity accounting for life course variability and life-history trait variances. To reduce intraspecific competition, individuals may explore new reproductive stages during years of major hurricanes, resulting in higher temporal variation in reproductive trajectories. Alternatively, individuals may opt for a single optimal life-history strategy due to trade-offs between survival and reproduction. Our results show that heterogeneity in reproductive life courses increased by 4% during years of major hurricanes, despite a 2% reduction in the asymptotic growth rate due to an average decrease in mean fertility and survival by that is, shortened life courses and reduced reproductive output. In agreement with this, the population is expected to achieve stable population dynamics faster after being perturbed by a hurricane ( ρ = 1.512 $$ \rho =1.512 $$ ; 95% CI: 1.488, 1.538), relative to ordinary years ρ = 1.482 ; 1.475 , 1.490 $$ \left(\rho =1.482;1.475,1.490\right) $$ . Our work suggests that natural disasters force individuals into new demographic roles to potentially reduce competition during unfavourable environments where mean reproduction and survival are compromised. Variance in lifetime reproductive success and longevity are differently affected by hurricanes, and such variability is mostly driven by survival.

Sexual transmission may drive pair similarity of the cloacal microbiome in a polyandrous species.

Abstract: All animals host a microbial community within and on their reproductive organs, known as the reproductive microbiome. In free-living birds, studies on the sexual transmission of bacteria have typically focused on a few pathogens instead of the bacterial community as a whole, despite a potential link to reproductive function. Theory predicts higher sexual transmission of the reproductive microbiome in females via the males' ejaculates and higher rates of transmission in promiscuous systems. We studied the cloacal microbiome of breeding individuals of a socially polyandrous, sex-role-reversed shorebird, the red phalarope (Phalaropus fulicarius). We expected (i) higher microbial diversity in females compared to males; (ii) low compositional differentiation between sexes; (iii) lower variation in composition between individuals (i.e. microbiome dispersion) in females than in males; (iv) convergence in composition as the breeding season progresses as a consequence of sexual transmission and/or shared habitat use; and (v) higher similarity in microbial composition between social pair members than between two random opposite-sex individuals. We found no or small between-sex differences in cloacal microbiome diversity/richness and composition. Dispersion of predicted functional pathways was lower in females than in males. As predicted, microbiome dispersion decreased with sampling date relative to clutch initiation of the social pair. Microbiome composition was significantly more similar among social pair members than among two random opposite-sex individuals. Pair membership explained 21.5% of the variation in taxonomic composition and 10.1% of functional profiles, whereas temporal and sex effects explained only 0.6%-1.6%. Consistent with evidence of functional convergence of reproductive microbiomes within pairs, some select taxa and predicted functional pathways were less variable between social pair members than between random opposite-sex individuals. As predicted if sexual transmission of the reproductive microbiome is high, sex differences in microbiome composition were weak in a socially polyandrous system with frequent copulations. Moreover, high within-pair similarity in microbiome composition, particularly for a few taxa spanning the spectrum of the beneficial-pathogenic axis, demonstrates the link between mating behaviour and the reproductive microbiome. Our study is consistent with the hypothesis that sexual transmission plays an important role in driving reproductive microbiome ecology and evolution.

Reproductive senescence and mating tactic interact and conflict to drive reproductive success in a passerine.

Abstract: An understanding of the drivers of individual fitness is a fundamental component of evolutionary ecology and life-history theory. Reproductive senescence, mate and mating tactic choice, and latent heterogeneity in individual quality interact to affect individual fitness. We sought to disentangle the effects of these fitness drivers, where longitudinal data are required to understand their respective impacts. We used reproductive allocation and success data from a long-term (1989-2018) study of white-throated dippers (Cinclus cinclus) in Switzerland to simultaneously examine the effects of female and male age, mating tactic, nest initiation date, and individual heterogeneity on reproductive performance. We modeled quadratic and categorical effects of age on reproductive parameters. The probability of polygyny increased with age in both sexes before declining in older age classes. Similarly, hatching probability in monogamous pairs and the number of nestlings hatched in both monogamous and polygynous pairs increased with female age before declining later in life. As predicted, offspring survival in monogamous pairs increased with male age before declining in older age classes, but male age had no effect on offspring survival in polygynous nesting attempts. Our results demonstrate that parental age, mating tactic, and individual heterogeneity all affect reproductive success, and that the impacts of senescent decline are expressed across different demographic components as a function of sex-specific senescent decline and mating tactic.

Resource competition drives an invasion‐replacement event among shrew species on an island

Abstract: Invasive mammals are responsible for the majority of native species extinctions on islands. While most of these extinction events will be due to novel interactions between species (e.g. exotic predators and naive prey), it is more unusual to find incidences where a newly invasive species causes the decline/extinction of a native species on an island when they normally coexist elsewhere in their overlapping mainland ranges. We investigated if resource competition between two insectivorous small mammals was playing a significant role in the rapid replacement of the native pygmy shrew Sorex minutus in the presence of the recently invading greater white-toothed shrew Crocidura russula on the island of Ireland. We used DNA metabarcoding of gut contents from >300 individuals of both species to determine each species' diet and measured the body size (weight and length) during different stages of the invasion in Ireland (before, during and after the species come into contact with one another) and on a French island where both species have long coexisted (acting as a natural 'control' site). Dietary composition, niche width and overlap and body size were compared in these different stages. The body size of the invasive C. russula and composition of its diet changes between when it first invades an area and after it becomes established. During the initial stages of the invasion, individual shrews are larger and consume larger sized invertebrate prey species. During later stages of the invasion, C. russula switches to consuming smaller prey taxa that are more essential for the native species. As a result, the level of interspecific dietary overlap increases from between 11% and 14% when they first come into contact with each other to between 39% and 46% after the invasion. Here we show that an invasive species can quickly alter its dietary niche in a new environment, ultimately causing the replacement of a native species. In addition, the invasive shrew could also be potentially exhausting local resources of larger invertebrate species. These subsequent changes in terrestrial invertebrate communities could have severe impacts further downstream on ecosystem functioning and services.

Lab cognition going wild: Implementing a new portable touchscreen system in vervet monkeys.

Abstract: Touchscreen technology has provided researchers with opportunities to conduct well-controlled cognitive tests with captive animals, allowing researchers to isolate individuals, select participants based on specific traits, and control aspects of the environment. In this study, we aimed to investigate the potential utility of touchscreen technology for the study of cognition in wild vervet monkeys. We assessed the viability of touchscreen testing by comparing rates of participation between wild and sanctuary-housed vervets. Additionally, we compared performance on a simple associative learning task in order to verify that wild participants are able to engage meaningfully with a touchscreen task presented in their natural environment. We presented eight groups of vervet monkeys (four wild and four sanctuary groups, totalling 240 individuals) with a portable touchscreen device. The touchscreen displayed tasks in which food rewards could be gained by touching a stimulus displayed on the screen. We assessed individuals' likelihood of interacting with the touchscreen, their frequency of participation, and their performance on a simple associative learning task. We found that sanctuary-housed monkeys were more likely to interact with the touchscreen. Participation in wild vervet monkeys was influenced by sex and age. However, monkeys in the two contexts (sanctuary vs. wild) did not differ in their performance on a simple associative learning task. This study demonstrates that touchscreen technology can be successfully deployed in a population of wild primates. This gives us a starting point to test animal cognition under natural conditions that include varying group composition, environmental challenges and ongoing activities such as foraging, which are challenging to recreate in captivity. While rates of participation were lower than those found in captivity, reasonable sample sizes can be achieved, and wild primates can successfully learn touchscreen tasks in a manner comparable to their captive counterparts.

Can behaviour and physiology mitigate effects of warming on ectotherms? A test in urban ants.

Abstract: Global climate change is expected to have pervasive effects on the diversity and distribution of species, particularly ectotherms whose body temperatures depend on environmental temperatures. However, these impacts remain difficult to predict, in part because ectotherms may adapt or acclimate to novel conditions or may use behavioural thermoregulation to reduce their exposure to stressful microclimates. Here we examine the potential for physiological and behavioural changes to mitigate effects of environmental warming on five species of ants in a temperate forest habitat subject to urban warming. We worked in eight urban and eight non-urban forest sites in North Carolina, USA; sites experienced a 1.1°C range of mean summer air temperatures. At each site, we documented species-specific microclimates (ant operative temperatures, Te ) and ant activity on a transect of 14 bait stations at three times of day. In the laboratory, we measured upper thermal tolerance (CTmax ) and thermal preference (Tpref ) for each focal species. We then asked whether thermal traits shifted at hotter sites, and whether ants avoided non-preferred microclimates in the field. CTmax and Tpref did not increase at warmer sites, indicating that these populations did not adapt or acclimate to urban warming. Consistent with behavioural thermoregulation, four of the five species were less likely to occupy baits where Te departed from Tpref . Apparent thermoregulation resulted from fixed diel activity patterns that helped ants avoid the most inappropriate temperatures but did not compensate for daily or spatial temperature variation: Hotter sites had hotter ants. This study uses a novel approach to detect behavioural thermoregulation and sublethal warming in foraging insects. The results suggest that adaptation and behaviour may not protect common temperate forest ants from a warming climate, and highlight the need to evaluate effects of chronic, sublethal warming on small ectotherms.

Plant–hummingbird pollination networks exhibit limited rewiring after experimental removal of a locally abundant plant species

Abstract: Mutualistic relationships, such as those between plants and pollinators, may be vulnerable to the local extinctions predicted under global environmental change. However, network theory predicts that plant–pollinator networks can withstand species loss if pollinators switch to alternative floral resources (rewiring). Whether rewiring occurs following species loss in natural communities is poorly known because replicated species exclusions are difficult to implement at appropriate spatial scales. We experimentally removed a hummingbird-pollinated plant, Heliconia tortuosa, from within tropical forest fragments to investigate how hummingbirds respond to temporary loss of an abundant resource. Under the rewiring hypothesis, we expected that behavioural flexibility would allow hummingbirds to use alternative resources, leading to decreased ecological specialization and reorganization of the network structure (i.e. pairwise interactions). Alternatively, morphological or behavioural constraints—such as trait-matching or interspecific competition—might limit the extent to which hummingbirds alter their foraging behaviour. We employed a replicated Before-After-Control-Impact experimental design and quantified plant–hummingbird interactions using two parallel sampling methods: pollen collected from individual hummingbirds (‘pollen networks’, created from >300 pollen samples) and observations of hummingbirds visiting focal plants (‘camera networks’, created from >19,000 observation hours). To assess the extent of rewiring, we quantified ecological specialization at the individual, species and network levels and examined interaction turnover (i.e. gain/loss of pairwise interactions). H. tortuosa removal caused some reorganization of pairwise interactions but did not prompt large changes in specialization, despite the large magnitude of our manipulation (on average, >100 inflorescences removed in exclusion areas of >1 ha). Although some individual hummingbirds sampled through time showed modest increases in niche breadth following Heliconia removal (relative to birds that did not experience resource loss), these changes were not reflected in species- and network-level specialization metrics. Our results suggest that, at least over short time-scales, animals may not necessarily shift to alternative resources after losing an abundant food resource—even in species thought to be highly opportunistic foragers, such as hummingbirds. Given that rewiring contributes to theoretical predictions of network stability, future studies should investigate why pollinators might not expand their diets after a local resource extinction.

A life-history spectrum of population responses to simultaneous change in climate and land use.

Abstract: 1. Climate and land-use change are two of the primary threats to global biodiversity; however, each species within a community may respond differently to these facets of global change. Although it is typically assumed that species use the habitat that is advantageous for survival and reproduction, anthropogenic changes to the environment can create ecological traps, making it critical to assess both habitat selection (e.g., where species congregate on the landscape) and the influence of selected habitats on the demographic processes that govern population dynamics. 2. We used a long-term (1958-2011), large-scale, multi-species data set for waterfowl that spans the United States and Canada to estimate species-specific responses to climate and land use variables in a landscape that has undergone significant environmental change across space and time. We first estimated the effects of change in climate and land use variables on habitat selection and population dynamics for nine species. We then hypothesized that species-specific responses to environmental change would scale with life-history traits, specifically: longevity, nesting phenology, and female breeding site fidelity. 3. We observed species-level heterogeneity in the demographic and habitat selection responses to climate and land use change, which would complicate community-level habitat management. Our work highlights the importance of multi-species monitoring and community-level analysis, even among closely related species. 4. We detected several relationships between life-history traits, particularly nesting phenology, and species' responses to environmental change. One species, the early-nesting northern pintail (Anas acuta), was consistently at the extreme end of responses to land use and climate predictors and has been a species of conservation concern since their population began to decline in the 1980s. They, and the blue-winged teal, also demonstrated a positive habitat selection response to the proportion of cropland on the landscape that simultaneously reduced abundance the following year, indicative of susceptibility to ecological traps. 5. By distilling the diversity of species' responses to environmental change within a community, our methodological approach and findings will help improve predictions of community responses to global change and can inform multi-species management and conservation plans in dynamic landscapes that are based on simple tenets of life-history theory.

Experimental evolution of dispersal: Unifying theory, experiments and natural systems.

Abstract: Dispersal is a central life history trait that affects the ecological and evolutionary dynamics of populations and communities. The recent use of experimental evolution for the study of dispersal is a promising avenue for demonstrating valuable proofs of concept, bringing insight into alternative dispersal strategies and trade-offs, and testing the repeatability of evolutionary outcomes. Practical constraints restrict experimental evolution studies of dispersal to a set of typically small, short-lived organisms reared in artificial laboratory conditions. Here, we argue that despite these restrictions, inferences from these studies can reinforce links between theoretical predictions and empirical observations and advance our understanding of the eco-evolutionary consequences of dispersal. We illustrate how applying an integrative framework of theory, experimental evolution and natural systems can improve our understanding of dispersal evolution under more complex and realistic biological scenarios, such as the role of biotic interactions and complex dispersal syndromes.

DeepWild: Application of the pose estimation tool DeepLabCut for behaviour tracking in wild chimpanzees and bonobos.

Abstract: Studying animal behaviour allows us to understand how different species and individuals navigate their physical and social worlds. Video coding of behaviour is considered a gold standard: allowing researchers to extract rich nuanced behavioural datasets, validate their reliability, and for research to be replicated. However, in practice, videos are only useful if data can be efficiently extracted. Manually locating relevant footage in 10,000 s of hours is extremely time-consuming, as is the manual coding of animal behaviour, which requires extensive training to achieve reliability. Machine learning approaches are used to automate the recognition of patterns within data, considerably reducing the time taken to extract data and improving reliability. However, tracking visual information to recognise nuanced behaviour is a challenging problem and, to date, the tracking and pose-estimation tools used to detect behaviour are typically applied where the visual environment is highly controlled. Animal behaviour researchers are interested in applying these tools to the study of wild animals, but it is not clear to what extent doing so is currently possible, or which tools are most suited to particular problems. To address this gap in knowledge, we describe the new tools available in this rapidly evolving landscape, suggest guidance for tool selection, provide a worked demonstration of the use of machine learning to track movement in video data of wild apes, and make our base models available for use. We use a pose-estimation tool, DeepLabCut, to demonstrate successful training of two pilot models of an extremely challenging pose estimate and tracking problem: multi-animal wild forest-living chimpanzees and bonobos across behavioural contexts from hand-held video footage. With DeepWild we show that, without requiring specific expertise in machine learning, pose estimation and movement tracking of free-living wild primates in visually complex environments is an attainable goal for behavioural researchers.

Patrolling the border: Billfish exploit the hypoxic boundary created by the world's largest oxygen minimum zone.

Abstract: Pelagic predators must contend with low prey densities that are irregularly distributed and dynamic in space and time. Based on satellite imagery and telemetry data, many pelagic predators will concentrate horizontal movements on ephemeral surface fronts-gradients between water masses-because of enhanced local productivity and increased forage fish densities. Vertical fronts (e.g. thermoclines, oxyclines) can be spatially and temporally persistent, and aggregate lower trophic level and diel vertically migrating organisms due to sharp changes in temperature, water density or available oxygen. Thus, vertical fronts represent a stable and potentially energy rich habitat feature for diving pelagic predators but remain little explored in their capacity to enhance foraging opportunities. Here, we use a novel suite of high-resolution biologging data, including in situ derived oxygen saturation and video, to document how two top predators in the pelagic ecosystem exploit the vertical fronts created by the oxygen minimum zone of the eastern tropical Pacific. Prey search behaviour was dependent on dive shape, and significantly increased near the thermocline and hypoxic boundary for blue marlin Makaira nigricans and sailfish Istiophorus platypterus, respectively. Further, we identify a behaviour not yet reported for pelagic predators, whereby the predator repeatedly dives below the thermocline and hypoxic boundary (and by extension, below the prey). We hypothesize this behaviour is used to ambush prey concentrated at the boundaries from below. We describe how habitat fronts created by low oxygen environments can influence pelagic ecosystems, which will become increasingly important to understand in the context of global change and expanding oxygen minimum zones. We anticipate that our findings are shared among many pelagic predators where strong vertical fronts occur, and additional high-resolution tagging is warranted to confirm this.

Species-level drivers of avian centrality within seed-dispersal networks across different levels of organisation.

Abstract: Bird-plant seed-dispersal networks are structural components of ecosystems. The role of bird species in seed-dispersal networks (from less [peripheral] to more connected [central]), determines the interaction patterns and their ecosystem services. These roles may be driven by morphological and functional traits as well as evolutionary, geographical and environmental properties acting at different spatial extents. It is still unknown if such drivers are equally important in determining species centrality at different network levels, from individual local networks to the global meta-network representing interactions across all local networks. Using 308 networks covering five continents and 11 biogeographical regions, we show that at the global meta-network level species' range size was the most important driver of species centrality, with more central species having larger range sizes, which would facilitate the interaction with a higher number of plants and thus the maintenance of seed-dispersal interactions. At the local network level, body mass was the only driver with a significant effect, implying that local factors related to resource availability are more important at this level of network organisation than those related to broad spatial factors such as range sizes. This could also be related to the mismatch between species-level traits, which do not consider intraspecific variation, and the local networks that can depend on such variation. Taken together, our results show that the drivers determining species centrality are relative to the levels of network organisation, suggesting that prediction of species functional roles in seed-dispersal interactions requires combined local and global approaches.