Human-induced Environmental Change Research Articles

Predators Show Seasonal Predilections for Model Clay Spiders in an Urban Environment

Predator-prey interactions may be altered under human-induced rapid environmental change, such as urbanisation. Extensive clearing in urban areas may leave short-range endemic species, such as mygalomorph spiders, more vulnerable to local extinction through predation in remaining remnants. Predation rates on Australian mygalomorph spiders were assessed using clay models of two size classes (5 cm, 3 cm), during two time periods in 2016 (January–February, July–August). Size and phenology of models resembled the mygalomorph genera Aname and Teyl occurring in these local urban remnants. Local predator guilds were significantly influenced by leaf-litter cover (%) and proportion of surrounding parkland. Preference for spider vs. control models was consistent across all predator types (bird, rodent, lizard and wasp), but specialist spider wasps (Pompilidae) only attacked spider models. Generalist predators (birds, lizards and rodents) were more opportunistic. Lizards and rodents exhibit similar predation behaviour, indicating there may be some inter-specific competition. Invasive generalists (e. g. rodents) or urban adapters (e. g. corvids) are more likely to represent an increased threat to spiders than are co-evolved specialists (e.g. spider wasps).

Survival and reproductive costs of repeated acute glucocorticoid elevations in a captive, wild animal

Organisms are continuously encountering both predictable and unpredictable ecological stressors within their environment. The activation of the hypothalamic-pituitaryadrenal (stress) axis is a fundamental process allowing animals to cope with and respond to such encounters. A main consequence of HPA axis activation is the release of glucocorticoid hormones. Although short-term glucocorticoid elevations lead to changes in physiological and behavioral processes that are often adaptive, our understanding of fitness consequences of repeated acute elevations in glucocorticoid hormones over a longer time period is largely lacking. This is of particular current importance as animals are facing a significant increase in exposure to stressors including those associated with human-induced rapid environmental change. Here, we test fitness-relevant consequences of repeated exposure to glucocorticoids in the absence of natural challenges, by treating wild-caught gravid female eastern fence lizards (Sceloporus undulatus) with a daily transdermal dose of a glucocorticoid hormone until laying. This treatment causes an increase in plasma glucocorticoids that mimics the natural response lizards have when they encounter a stressor in the wild, without confounding effects associated with the encounter itself. This treatment reduced females’ reproductive success (hatching success) and survival. Further, glucocorticoid-induced reductions in reproductive success were greater when females had experienced higher temperatures the previous winter. This demonstrates the potential significant consequences of repeated exposure to acute elevations in glucocorticoid hormones. Additionally, the costs of repeated glucocorticoid elevation may be further exaggerated by an individual’s previous experience, such as the potential compounding effects of winter warming increasing animals’ vulnerability to increased glucocorticoid levels during spring breeding.

Quantifying the effects of climate and anthropogenic change on regional species loss in China.

Human-induced environmental and climate change are widely blamed for causing rapid global biodiversity loss, but direct estimation of the proportion of biodiversity lost at local or regional scales are still infrequent. This prevents us from quantifying the main and interactive effects of anthropogenic environmental and climate change on species loss. Here, we demonstrate that the estimated proportion of species loss of 252 key protected vertebrate species at a county level of China during the past half century was 27.2% for all taxa, 47.7% for mammals, 28.8% for amphibians and reptiles and 19.8% for birds. Both human population increase and species richness showed significant positive correlations with species loss of all taxa combined, mammals, birds, and amphibians and reptiles. Temperature increase was positively correlated with all-taxa and bird species loss. Precipitation increase was negatively correlated with species loss of birds. Human population change and species richness showed more significant interactions with the other correlates of species loss. High species richness regions had higher species loss under the drivers of human environmental and climate change than low-richness regions. Consequently, ongoing human environmental and climate changes are expected to perpetuate more negative effects on the survival of key vertebrate species, particularly in high-biodiversity regions.

Sensory System Responses to Human-Induced Environmental Change

Sensory input to the central nervous system is the primary means by which animals respond to variation in their physical and biological environments. It is well established that key threats such as habitat destruction, the introduction of non-native species, and climate change are imposing significant pressures on natural ecosystems, yet surprisingly few studies have examined how these threats impact the senses or determine species’ responses to environmental change. This review focuses on how anthropogenic impacts on aquatic ecosystems can have a detrimental effect on the sensory systems of aquatic organisms and how these modalities can act to influence genetic and non-genetic (e.g. developmental) responses to environmental change, which in turn can cause knock-on effects in a range of other biological systems. Species often exhibit unique sensory specializations that are suited to their behavioral requirements; at present it is unclear whether and how sensory systems have the capacity to respond to environmental change through genetic adaptation and/or sensory plasticity, and on what timescale this might occur. Sensory systems lie at the forefront of how various species respond to environmental perturbation. As such, determining the important role they play in determining fitness is critical for understanding the effects of external processes such as habitat degradation and climate change. Given the current consensus that human impacts and environmental changes are potentially highly detrimental to the delicate balance of the biome, knowing how organisms respond, and to what degree adaptation is physiologically and behaviorally limited, warrants urgent attention.

The timing of energy allocation to reproduction in an important group of marine consumers.

Organisms may energetically finance reproductive effort using energy stored prior to the reproductive period (termed capital breeders) or using energy acquired during the reproductive period (termed income breeders). The specific strategy used has implications for population dynamics as well as for the response to environmental variation. Crabs and other crustaceans have generally been assumed to be capital breeders. Here I demonstrate an experimental procedure used to determine whether crabs are capable of using an income breeding strategy. I then examine data from several published studies from a range of crab species across a broad phylogenetic spectrum that conducted similar experiments to look for evidence of income breeding strategy. I show that income breeding does occur in crabs, but that it appears to be taxon-specific. In particular, I show that income breeding occurs in two species from the family Portunidae, but fail to find evidence for income breeding in other taxa examined. This finding has a range of implications for this ecologically and economically important group of consumers, including implications for their response to human-induced environmental change, their response to fishing pressure, and best practices for aquaculture. The implications of breeding strategy in crabs likely depends on phylogeny (morphology) and ecology, both of which influence the space available for energy storage inside the carapace.

Recovery of coral assemblages despite acute and recurrent disturbances on a South Central Pacific reef

Coral reefs are increasingly threatened by various types of disturbances, and their recovery is challenged by accelerating, human-induced environmental changes. Recurrent disturbances reduce the pool of mature adult colonies of reef-building corals and undermine post-disturbance recovery from newly settled recruits. Using a long-term interannual data set, we show that coral assemblages on the reef slope of Moorea, French Polynesia, have maintained a high capacity to recover despite a unique frequency of large-scale disturbances which, since the 1990s, have caused catastrophic declines in coral cover and abundance. In 2014, only four years after one of the most extreme cases of coral decline documented, abundance of juvenile and adult colonies had regained or exceeded pre-disturbance levels, and no phase-shift to macroalgal dominance was recorded. This rapid recovery has been achieved despite constantly low coral recruitment rates, suggesting a high post-disturbance survivorship of recruits. However, taxonomic differences in coral susceptibility to disturbances and contrasting recovery trajectories have resulted in changes in the relative composition of species. In the present context of global coral reef decline, our study establishes a new benchmark for the capacity of certain benthic reef communities to sustain and recover their coral cover from repeated, intense disturbances.

Thermal forest zone explains regional variations in the diet composition of the Japanese marten (Martes melampus)

Examining biogeographical variations in species’ diet is important for linking key ecological traits between consumers and their environment. While the trophic ecology of the Holarctic martens has been globally synthesised, systematic information on the feeding ecology of martens (Martes spp.) in Asia is still deficient compared to species in Europe and North America. Here we conduct a meta-analysis based on literature describing regional and seasonal variation in diet composition of the Japanese marten (Martes melampus). Across nine study areas in the Japanese archipelago, annual marten diet mainly comprised fruits, invertebrates, and mammals. Cluster analysis divided these into three groups according to the similarity in composition. Dietary niche breadth and the relative frequency of occurrence of the mammalian prey was significantly higher in sub-alpine/cool-temperate than warm-temperate thermal forest zones, whereas the occurrence of fruits was significantly higher in warm-temperate than sub-alpine/cool-temperate zones. Seasonal analysis showed significant effects across all four seasons on the frequency of occurrence of mammals, invertebrates, and fruits. The Japanese marten is a flexible and opportunistic feeder with the potential to maintain its key macronutrient requirements by adapting its feeding strategy to changes in resource variability and environmental conditions. This trophic plasticity may advantage the Japanese marten in coping with human-induced rapid environmental change, as global analysis has projected for martens across other regions.

Increased body size along urbanization gradients at both community and intraspecific level in macro-moths.

Urbanization involves a cocktail of human-induced rapid environmental changes and is forecasted to gain further importance. Urban-heat-island effects result in increased metabolic costs expected to drive shifts towards smaller body sizes. However, urban environments are also characterized by strong habitat fragmentation, often selecting for dispersal phenotypes. Here, we investigate to what extent, and at which spatial scale(s), urbanization drives body size shifts in macro-moths-an insect group characterized by positive size-dispersal links-at both the community and intraspecific level. Using light and bait trapping as part of a replicated, spatially nested sampling design, we show that despite the observed urban warming of their woodland habitat, macro-moth communities display considerable increases in community-weighted mean body size because of stronger filtering against small species along urbanization gradients. Urbanization drives intraspecific shifts towards increased body size too, at least for a third of species analysed. These results indicate that urbanization drives shifts towards larger, and hence, more mobile species and individuals in order to mitigate low connectivity of ecological resources in urban settings. Macro-moths are a key group within terrestrial ecosystems, and since body size is central to species interactions, such urbanization-driven phenotypic change may impact urban ecosystem functioning, especially in terms of nocturnal pollination and food web dynamics. Although we show that urbanization's size-biased filtering happens simultaneously and coherently at both the inter- and intraspecific level, we demonstrate that the impact at the community level is most pronounced at the 800 m radius scale, whereas species-specific size increases happen at local and landscape scales (50-3,200m radius), depending on the species. Hence, measures-such as creating and improving urban green infrastructure-to mitigate the effects of urbanization on body size will have to be implemented at multiple spatial scales in order to be most effective.

Human-induced environmental changes influence habitat use by an ungulate over the long term.

Habitat use and preferences may be subject to spatial and temporal changes. However, long-term studies of species–habitat relationships are the exception. In the present research, long-term trends in habitat use by an alpine ungulate, the Tatra chamois Rupicapra rupicapra tatrica, were analyzed. We examined how environmental changes attributable to climate change, removal of sheep, and habituation to hikers, which took place over the last half-century have changed the spatial distribution of animals. Data on the localities of groups sighted between 1957 and 2013 during autumnal population surveys were used to evaluate habitat associations: these were correlated with year, group size, population size, and climatic conditions. The results indicate that the Tatra chamois is tending, over the long term, to lower its altitude of occurrence, reduce its average distance to hiking trails, and stay less often on slopes with a southerly aspect. These trends are independent of group size, population size, and the weather conditions prevailing during observations, though not for altitude, where increases in air temperature are related to finding chamois at higher elevations. The proportion of alpine meadows and slope in the places used by chamois is correlated with population size, while the proportion of areas with trees and/or shrubs is correlated with group size and air temperature, though long-term changes were not evident for these variables. To the best of our knowledge, this work is the first to document long-term trends in habitat use by ungulates. It shows that a species’ ecology is influenced by human-induced changes: abandonment of pasturage, high-mountain tourism, and climate changes, which constitute the most probable reasons for this aspect of behavioral evolution in the Tatra chamois.

Climate and anthropogenic controls on the carbon cycle of Xingyun Lake, China

The organic and inorganic carbon cycles in lakes are responsive to both natural climate variability and human-induced environmental changes. Here we present an 8600-year sediment record from Xingyun Lake in Yunnan, China that provides insight into carbon cycling and lake primary productivity using stable isotope measurements of organic carbon and nitrogen, carbon to nitrogen ratio, inorganic carbon isotopes, and mass accumulation rates of carbon, nitrogen, and calcite. The early to middle Holocene is characterized by relative stability in most proxies although some variation in organic carbon isotopes reflects fluctuating terrestrial organic matter inputs. The middle to late Holocene, from 5300 to 3300 years BP, is marked by amplified variability in nearly all proxies and suggests either increased C4 vegetation or increased primary productivity. These changes are coincident with declining lake levels and increased aridity throughout the Indian Summer Monsoon region. After 3300 years BP, the greatest variability is in nitrogen isotopes and coincides closely with human settlement in the region. These changes are accompanied by further indications of increased primary productivity, suggesting eutrophic lake conditions. While we cannot definitively link human activity to these changes, the balance of evidence suggests that other mechanisms are unlikely. This study supports previous work that anthropogenic activities began measurably impacting the landscape in southwestern China within the last few millennia.

Introgression among three rockfish species (Sebastes spp.) in the Salish Sea, northeast Pacific Ocean.

Interspecific hybridization is often seen as a major conservation issue, potentially threatening endangered species and decreasing biodiversity. In natural populations, the conservation implications of hybridization depends on both on anthropogenic factors and the evolutionary processes maintaining the hybrid zone. However, the timeline and patterns of hybridization in the hybrid zone are often not known. Therefore, species conservation becomes a concern when recent anthropogenic changes influence hybridization and not if hybridization is part of a long-term process. Here, we use sequence data from one mitochondrial gene, three nuclear introns and one nuclear exon to estimate the direction, geographic extent, frequency and possible timeline of hybridization between three rockfish species (Sebastes auriculatus, S. caurinus, S. maliger) in the Salish Sea, Washington, USA. We show that (i) introgression occurred much more frequently in the Salish Sea than on the outer coast, (ii) introgression was highly asymmetrical from S. maliger into the other two species, (iii) almost 40% of individuals in the Salish Sea were hybrids, with frequency of hybrids increasing with isolation from the coast, and (iv) all hybrids were later generation backcrosses rather than F1 hybrids. Our results suggest long-standing low-level hybridization rather than recent onset of interbreeding because of human induced environmental change, possibly facilitated by specific environmental conditions in the sub-basins of the Salish Sea, and by differences in population sizes during recolonization of the area after the last glaciation. This rockfish hybrid system, with asymmetrical introgression and the maintenance of parental species, may prove useful to study both mechanisms that maintain species boundaries and that facilitate speciation in the presence of rapid environmental change.

Urban blackbirds have shorter telomeres.

Urbanization, one of the most extreme human-induced environmental changes, represents a major challenge for many organisms. Anthropogenic habitats can have opposing effects on different fitness components, for example, by decreasing starvation risk but also health status. Assessment of the net fitness effect of anthropogenic habitats is therefore difficult. Telomere length is associated with phenotypic quality and mortality rate in many species, and the rate of telomere shortening is considered an integrative measure of the 'life stress' experienced by an individual. This makes telomere length a promising candidate for examining the effects of urbanization on the health status of individuals. We investigated whether telomere length differed between urban and forest-dwelling common blackbirds (Turdus merula). Using the terminal restriction fragment assay, we analysed telomere length in yearlings and older adults from five population dyads (urban versus forest) across Europe. In both age classes, urban blackbirds had significantly shorter telomeres (547 bp) than blackbirds in natural habitats, indicating lower health status in urban blackbirds. We propose several potential hypotheses to explain our results. Our findings show that even successful city dwellers such as blackbirds pay a price for living in these anthropogenic habitats.

Risk-taking behavior, urbanization and the pace of life in birds

Despite growing appreciation of the importance of considering a pace-of-life syndrome (POLS) perspective to understand how animals interact with their environment, studies relating behavior to life history under altered environmental conditions are still rare. By means of a comparative analysis of flight initiation distances (i.e., the distance at which an animal takes flight when a human being is approaching) across > 300 bird species distributed worldwide, we document here the existence of a POLS predicted by theory where slow-lived species tend to be more risk-averse than fast-lived species. This syndrome largely emerges from the influence of body mass, and is highly dependent on the environmental context. Accordingly, the POLS structure vanishes in urbanized environments due to slow-lived species adjusting their flight distances based on the perception of risk. While it is unclear whether changes in POLS reflect plastic and/or evolutionary adjustments, our findings highlight the need to integrate behavior into life history theory to fully understand how animals tolerate human-induced environmental changes. Animals can often respond to changing environmental conditions by adjusting their behavior. However, the degree to which different species can modify their behavior depends on their life history strategy and on the environmental context. Species-specific perception of risk is a conspicuous example of adjustable behavior tightly associated with life history strategy. While there is a general tendency of higher risk aversion in rural than city-dwelling birds, it is dependent on the species’ life history strategy. Slow-lived species are more prone to adjust their flight initiation distances based on the perception of risk, allowing humans to approach closer in urban than rural environments. Behavior must therefore be taken into account together with life history to reliably assess species’ vulnerability at the face of ongoing environmental change.

Flood-related risks in Ho Chi Minh City and ways of mitigation

With an ever-growing population of around 10 million inhabitants (officially 7.9 in 2013), Ho Chi Minh City (HCMC) is set to become one of the largest cities in South East Asia and already occupies a major economic role in the area. To accommodate the increasing population, the megacity now stretches out in an urban continuum covering more than 800 square kilometers and is currently growing at a rate of 3.2% per year. If the neighboring provinces around HCMC are included, the total population reaches nearly 18 million people.This paper attempts to describe the interplay between HCMC and flood-related risks and offer some guidelines to deal with inundations. The potential risks of flooding by rain, tsunami and/or dam failure upstream of the city are evaluated and contextualized within the perspective of climate and human-induced environmental changes. The region is highly vulnerable to the combined effects of subsidence and rising sea levels and has already led to serious flooding that may extend spatially before the end of the century. We propose possible preventative solutions to urban flooding using a multi-pronged approach to issues regarding urban development and suggest a redevelopment strategy for major infrastructure projects.

Is reduced freshwater flow in Tigris-Euphrates rivers driving fish recruitment changes in the Northwestern Arabian Gulf?

Like fishing, natural regime shifts and human-induced environmental changes are often equally important factors in driving fish stock declines. In the Northwestern Arabian Gulf, many fish stocks are declining, raising questions about the reduction in the flow of Tigris-Euphrates rivers. Here we investigate the relationship between Tigris-Euphrates river flow and the estimated recruitment patterns from assessment models. We found a positive correlation between the estimated finfish recruitment trends and the flow of Tigris-Euphrates rivers. Additionally, the assessment model showed remarkably weak compensation ratio, likely indicating a reduction in the productivity of nursery area of two finfish stocks but not in that of the crustacean stock. Our investigation would be very critical in providing guidelines to the government agencies in the Northwestern Arabian Gulf as well as countries of Tigris-Euphrates basins: to consider the impacts associated with reductions in Tigris-Euphrates river flows on the ecosystem services of the region.

Foraging plasticity in seabirds: A non-invasive study of the diet of greater crested terns breeding in the Benguela region.

Marine predators, such as seabirds, are useful indicators of marine ecosystem functioning. In particular, seabird diet may reflect variability in food-web composition due to natural or human-induced environmental change. Diet monitoring programmes, which sample diet non-invasively, are valuable aids to conservation and management decision-making. We investigated the diet of an increasing population of greater crested terns Thalasseus bergii in the Western Cape, South Africa, during three successive breeding seasons (2013 to 2015), when populations of other seabirds feeding on small pelagic schooling fish in the region were decreasing. Breeding greater crested terns carry prey in their bills, so we used an intensive photo-sampling method to record their diet with little disturbance. We identified 24,607 prey items from at least 47 different families, with 34 new prey species recorded. Fish dominated the diet, constituting 94% of prey by number, followed by cephalopods (3%), crustaceans (2%) and insects (1%). The terns mainly targeted surface-schooling Clupeiformes, with anchovy Engraulis encrasicolus the most abundant prey in all three breeding seasons (65% overall). Prey composition differed significantly between breeding stages and years, with anchovy most abundant at the start of the breeding season, becoming less frequent as the season progressed. The proportion of anchovy in the diet also was influenced by environmental factors; anchovy occurred more frequently with increasing wind speeds and was scarce on foggy days, presumably because terns rely in part on social facilitation to locate anchovy schools. The application of this intensive and non-invasive photo-sampling method revealed an important degree of foraging plasticity for this seabird within a context of locally reduced food availability, suggesting that, unlike species that specialise on a few high-quality prey, opportunistic seabirds may be better able to cope with reductions in the abundance of their preferred prey.

Multi-scale habitat selection by two declining East Asian waterfowl species at their core spring stopover area

Animals respond to their environment at multiple spatial scales that each require different conservation measures. Waterbirds are key bio-indicators for globally threatened wetland ecosystems but their multi-scale habitat selection mechanisms have rarely been studied. Using satellite tracking data and Maximum entropy modeling, we studied habitat selection of two declining waterfowl species, the Greater White-fronted Goose (Anser Albifrons) and the Tundra Bean Goose (A. serrirostris), at three spatial scales: landscape (30, 40, 50 km), foraging (10, 15, 20 km) and roosting (1, 3, 5 km). We hypothesized that the landscape-scale habitat selection was mainly based on relatively coarse landscape metrics, while more detailed landscape features were taken into account for the foraging- and roosting- scale habitat selection. We found that both waterfowl species preferred areas with a larger percentage of wetland and waterbodies at the landscape scale, aggregated waterbodies surrounded by scattered croplands at the foraging scale, and well-connected wetlands and well-connected middle-sized waterbodies at the roosting scale. The main difference in habitat selection for the two species occurred at the landscape and foraging scale; factors at the roosting scale were similar. We suggest that conservation activities should focus on enhancing the aggregation and connectivity of waterbodies and wetlands, and developing less aggregated cropland in the surroundings. Our approach could guide waterbird conservation practices and wetland management by providing effective measures to improve habitat quality in the face of human-induced environmental change.

Contemporary evolution of a Lepidopteran species, Heliothis virescens, in response to modern agricultural practices.

Adaptation to human-induced environmental change has the potential to profoundly influence the genomic architecture of affected species. This is particularly true in agricultural ecosystems, where anthropogenic selection pressure is strong. Heliothis virescens primarily feeds on cotton in its larval stages, and US populations have been declining since the widespread planting of transgenic cotton, which endogenously expresses proteins derived from Bacillus thuringiensis (Bt). No physiological adaptation to Bt toxin has been found in the field, so adaptation in this altered environment could involve (i) shifts in host plant selection mechanisms to avoid cotton, (ii) changes in detoxification mechanisms required for cotton-feeding vs. feeding on other hosts or (iii) loss of resistance to previously used management practices including insecticides. Here, we begin to address whether such changes occurred in H.virescens populations between 1997 and 2012, as Bt-cotton cultivation spread through the agricultural landscape. For our study, we produced an H.virescens genome assembly and used this in concert with a ddRAD-seq-enabled genome scan to identify loci with significant allele frequency changes over the 15-year period. Genetic changes at a previously described H.virescens insecticide target of selection were detectable in our genome scan and increased our confidence in this methodology. Additional loci were also detected as being under selection, and we quantified the selection strength required to elicit observed allele frequency changes at each locus. Potential contributions of genes near loci under selection to adaptive phenotypes in the H.virescens cotton system are discussed.

Genetic diversity and structure related to expansion history and habitat isolation: stone marten populating rural\u2013urban habitats

BackgroundPopulation genetic diversity and structure are determined by past and current evolutionary processes, among which spatially limited dispersal, genetic drift, and shifts in species distribution boundaries have major effects. In most wildlife species, environmental modifications by humans often lead to contraction of species’ ranges and/or limit their dispersal by acting as environmental barriers. However, in species well adapted to anthropogenic habitat or open landscapes, human induced environmental changes may facilitate dispersal and range expansions. In this study, we analysed whether isolation by distance and deforestation, among other environmental features, promotes or restricts dispersal and expansion in stone marten (Martes foina) populations.ResultsWe genotyped 298 martens from eight sites at twenty-two microsatellite loci to characterize the genetic variability, population structure and demographic history of stone martens in Poland. At the landscape scale, limited genetic differentiation between sites in a mosaic of urban, rural and forest habitats was mostly influenced by isolation by distance. Statistical clustering and multivariate analyses showed weak genetic structuring with two to four clusters and a high rate of gene flow between them. Stronger genetic differentiation was detected for one stone marten population (NE1) located inside a large forest complex. Genetic differentiation between this site and all others was 20% higher than between other sites separated by similar distances. The genetic uniqueness index of NE1 was also twofold higher than in other sites. Past demographic history analyses showed recent expansion of this species in north-eastern Poland. A decrease in genetic diversity from south to north, and MIGRAINE analyses indicated the direction of expansion of stone marten.ConclusionsOur results showed that two processes, changes in species distribution boundaries and limited dispersal associated with landscape barriers, affect genetic diversity and structure in stone marten. Analysis of local barriers that reduced dispersal and large scale analyses of genetic structure and demographic history highlight the importance of isolation by distance and forest cover for the past colonization of central Europe by stone marten. This confirmed the hypothesis that human-landscape changes (deforestation) accelerated stone marten expansion, to which climate warming probably has also been contributing over the last few decades.

An evolutionary framework outlining the integration of individual social and spatial ecology.

Behaviour is the interface between an organism and its environment, and behavioural plasticity is important for organisms to cope with environmental change. Social behaviour is particularly important because sociality is a dynamic process, where environmental variation influences group dynamics and social plasticity can mediate resource acquisition. Heterogeneity in the ecological environment can therefore influence the social environment. The combination of the ecological and social environments may be interpreted collectively as the "socioecological environment," which could explain variation in fitness. Our objective was to outline a framework through which individual social and spatial phenotypes can be integrated and interpreted as phenotypes that covary as a function of changes in the socioecological environment. We propose the socioecological environment is composed of individual behavioural traits, including sociality and habitat selection, both of which are repeatable, potentially heritable and may reflect animal personality traits. We also highlight how ecological and social niche theory can be applied to the socioecological environment framework, where individuals occupy different socioecological niches. Individual sociality and habitat selection are also density-dependent, and theory predicts that density-dependent traits should affect reproduction, survival, and therefore fitness and population dynamics. We then illustrate the proximate links between sociality, habitat selection and fitness as well as the ultimate, and possibly adaptive, consequences associated with changes in population density. The ecological, evolutionary and applied implications of our proposed socioecological environment framework are broad and changes in density could influence individual fitness and population dynamics. For instance, human-induced environmental changes can influence population density, which can affect the distribution of social and spatial phenotypes within a population. In summary, we outline a conceptual framework that incorporates individual social and spatial behavioural traits with fitness and we highlight a range of ecological and evolutionary processes that are likely associated with the socioecological environment.