Disentangling spatiotemporal dynamics in metacommunities through a species-patch network approach.

Colonization and extinction at local and regional scales, and gains and losses of patches are important processes in the spatiotemporal dynamics of metacommunities. However, analytical challenges remain in quantifying such spatiotemporal dynamics when species extinction-colonization and patch gain and loss processes act simultaneously. Recent advances in network analysis show great potential in disentangling the roles of colonization, extinction, and patch dynamics in metacommunities. Here, we developed a species-patch network approach to quantify metacommunity dynamics including (i) temporal changes in network structure, and (ii) temporal beta diversity of species-patch links and its components that reflect species extinction-colonization and patch gain and loss. Application of the methods to simulated datasets demonstrated that the approach was informative about metacommunity assembly processes. Based on three empirical datasets, our species-patch network approach provided additional information about metacommunity dynamics through distinguishing the effects of species colonization and extinction at different scales from patch gains and losses and how specific environmental factors related to species-patch network structure. In conclusion, our species-patch network framework provides effective methods for monitoring and revealing long-term metacommunity dynamics by quantifying gains and losses of both species and patches under local and global environmental change.

Evolutionary and ecological patterns of scatter- and larder-hoarding behaviours in rodents.

Scatter- and larder hoarding are the primary strategies of food-hoarding animals and have important implications for plant-animal interactions and plant recruitment. However, their origins and influencing factors have not been fully investigated across a wide range of taxa. Our systematic literature search amassed data for 183 seed-hoarding rodent species worldwide and tested relationships of seed-hoarding behaviours with phylogenetic signal, functional traits and environmental factors. We found that the evolution of hoarding strategies was not random in phylogeny, and scatter hoarding originated independently multiple times from larder hoarding. Rodents with higher encephalisation quotient (relative brain size), omnivorous diet (related to dependence on seeds) and inhabiting lower latitudes were disproportionately likely to scatter hoard. Despite body mass's potential relationship with competition through food defence, it was associated with food-hoarding strategy only in a few families. Our results show the need to study the community and ecological context of food-hoarding behaviours.

Hilltopping influences spatial dynamics in a patchy population of tiger moths.

Dispersal is a key driver of spatial population dynamics. Dispersal behaviour may be shaped by many factors, such as mate-finding, the spatial distribution of resources, or wind and currents, yet most models of spatial dynamics assume random dispersal. We examined the spatial dynamics of a day-flying moth species (Arctia virginalis) that forms mating aggregations on hilltops (hilltopping) based on long-term adult and larval population censuses. Using time-series models, we compared spatial population dynamics resulting from empirically founded hilltop-based connectivity indices and modelled the interactive effects of temperature, precipitation and density dependence. Model comparisons supported hilltop-based connectivity metrics including hilltop elevation over random connectivity, suggesting an effect of hilltopping behaviour on dynamics. We also found strong interactive effects of temperature and precipitation on dynamics. Simulations based on fitted time-series models showed lower patch occupancy and regional synchrony, and higher colonization and extinction rates when hilltopping was included, with potential implications for the probability of persistence of the patch network. Overall, our results show the potential for dispersal behaviour to have important effects on spatial population dynamics and persistence, and we advocate the inclusion of such non-random dispersal in metapopulation models.

Quantifying phenology and migratory behaviours of hummingbirds using single-site dynamics and mark-detection analyses.

Nuanced understanding of seasonal movements of partially migratory birds is paramount to species and habitat conservation. Using nascent statistical methods, we identified migratory strategies of birds outfitted with radio-frequency identification (RFID) tags detected at RFID feeders in two sites in California, USA. We quantified proportions of migrants and residents and the seasonal phenology for each movement strategy in Allen's and Anna's hummingbirds; we also validated our methodology by fitting our model to obligate migratory black-chinned hummingbirds. Allen's and Anna's hummingbirds exhibited characteristics of facultative migratory behaviour. We also quantified apparent annual survival for each migratory strategy and found that residents had significantly higher probabilities of apparent survival. Low survival estimates for migrants suggest that a high proportion of birds in the migrant group permanently emigrated from our study sites. Considered together, our analyses suggest that hummingbirds in both northern and southern California sites partake in diverse and highly plastic migratory behaviours. Our assessment elucidates the dynamics underlying idiosyncratic migratory behaviours of two species of hummingbirds, in addition to describing a framework for similar assessments of migratory behaviours using the multi-state open robust design with state uncertainty model and single-site dynamics.

Mixed-species bird flocks re-assemble interspecific associations across an elevational gradient.

Understanding how non-trophic social systems respond to environmental gradients is still a challenge in animal ecology, particularly in comparing changes in species composition to changes in interspecific interactions. Here, we combined long-term monitoring of mixed-species bird flocks, data on participating species' evolutionary history and traits, to test how elevation affected community assemblages and interspecific interactions in flock social networks. Elevation primarily affected flocks through reassembling interspecific associations rather than modifying community assemblages. Specifically, flock networks at higher elevations (compared to low elevations) had stronger interspecific associations (larger average weighted degree), network connectivity (enhanced network density) and fewer subnetworks. A phylogenetic and functional perspective revealed that associations between similar species weakened, whereas connections between dissimilar and/or random species were unchanged or strengthened with elevation. Likewise, network assortativity for the traits of vertical stratum and breeding period declined with elevation. The overall pattern is a change from modular networks in the lowlands, where species join flocks with other species that have matching traits, to a more open, random system at high elevations. Collectively, this rewiring of interspecific networks across elevational gradients imparts network stability and resiliency and makes mixed-species flocks less sensitive to local extinctions caused by harsh environments.

Variability in plant nutrients reduces insect herbivore performance.

The performance and population dynamics of insect herbivores depend on the nutritive and defensive traits of their host plants. The literature on plant-herbivore interactions focuses on plant trait mean values, but recent studies showing the importance of plant genetic diversity for herbivores suggest that plant trait variance may be equally important. The consequences of plant trait variance for herbivore performance, however, have been largely overlooked. Here we report an extensive assessment of the effects of within-population plant trait variance on herbivore performance using 457 performance datasets from 53 species of insect herbivores. We show that variance in plant nutritive traits substantially reduces mean herbivore performance via non-linear averaging of performance relationships that were overwhelmingly concave down. By contrast, relationships between herbivore performance and plant defence levels were typically linear, with variance in plant defence not affecting herbivore performance via non-linear averaging. Our results demonstrate that plants contribute to the suppression of herbivore populations through variable nutrient levels, not just by having low average quality as is typically thought. We propose that this phenomenon could play a key role in the suppression of herbivore populations in natural systems, and that increased nutrient heterogeneity within agricultural crops could contribute to the sustainable control of insect pests in agroecosystems.

Altered precipitation dynamics lead to a shift in herbivore dynamical regime.

The interaction between endogenous dynamics and exogenous environmental variation is central to population dynamics. Although investigations into the effects of changing mean climate are widespread, changing patterns of variation in environmental forcing also affect dynamics in complex ways. Using wavelet and time series analyses, we identify a regime shift in the dynamics of a moth species in California from shorter to longer period oscillations over a 34-year census, and contemporaneous changes in regional precipitation dynamics. Simulations support the hypothesis that shifting precipitation dynamics drove changes in moth dynamics, possibly due to stochastic resonance with delayed density-dependence. The observed shift in climate dynamics and the interaction with endogenous dynamics mean that predicting future population dynamics will require information on both climatic shifts and their interaction with endogenous density-dependence, a combination that is rarely available. Consequently, models based on historical data may be unable to predict future population dynamics.

The functional roles of species in metacommunities, as revealed by metanetwork analyses of bird-plant frugivory networks.

Understanding how biodiversity and interaction networks change across environmental gradients is a major challenge in ecology. We integrated metacommunity and metanetwork perspectives to test species' functional roles in bird-plant frugivory interactions in a fragmented forest landscape in Southwest China, with consequences for seed dispersal. Availability of fruit resources both on and under trees created vertical feeding stratification for frugivorous birds. Bird-plant interactions involving birds feeding only on-the-tree or both on and under-the-tree (shared) had a higher centrality and contributed more to metanetwork organisation than interactions involving birds feeding only under-the-tree. Moreover, bird-plant interactions associated with large-seeded plants disproportionately contributed to metanetwork organisation and centrality. Consequently, on-the-tree and shared birds contributed more to metanetwork organisation whereas under-the-tree birds were more involved in local processes. We would expect that species' roles in the metanetwork will translate into different conservation values for maintaining functioning of seed-dispersal networks.

As temperature increases, predator attack rate is more important to survival than a smaller window of prey vulnerability.

Climate change can have strong effects on species interactions and community structure. Temperature-dependent effects on predator-prey interactions are a major mechanism through which these effects occur. To understand the net effects of predator attack rates and dynamic windows of prey vulnerability, we examined the impacts of temperature on the interaction of a caterpillar (Arctia virginalis) and its ant predator (Formica lasioides). We conducted field experiments to examine attack rates on caterpillars relative to temperature, ant abundance, and body size, and laboratory experiments to determine the effects of temperature on caterpillar growth. We modeled temperature-dependent survival based on the integrated effects of temperature-dependent growth and temperature- and size-dependent predation. Attack rates on caterpillars increased with warming and ant recruitment, but decreased with caterpillar size. Caterpillar growth rates increased with temperature, narrowing the window of vulnerability. The model predicted that net caterpillar survival would decrease with temperature, suggesting that A. virginalis populations could be depressed with future climate warming. Theoretical work suggests that the net outcome of predator-prey interactions with increasing temperature depends on the respective responses of interacting species in terms of velocity across space, whereas the present study suggests the importance of effects of temperature on prey window of vulnerability, or "velocity" across time.

Wet years have more caterpillars: interacting roles of plant litter and predation by ants.

Climate is widely recognized as an important factor that affects temporal and spatial patterns of occurrence and abundance of herbivorous insects, although the ecological mechanisms responsible are poorly understood. We found that precipitation and standing water were positively correlated with locations and years of high abundance of caterpillars of the ranchman's tiger moth, Platyprepia virginalis. We analyzed 30 years of survey data and found that the number of large rainfall events was a better predictor of caterpillar abundance than total annual accumulation. We considered three ecological mechanisms that could drive this relationship and conducted observations and manipulative experiments to evaluate these mechanisms. (1) Rainfall facilitates more plant growth, although we found no evidence that increased food quality or quantity was causing the positive association between precipitation and caterpillar abundance. (2) Large rainfall events cause predatory ground-nesting ants to be less abundant and we found that the number of ants that recruited to local sites was negatively associated with survival and abundance of caterpillars. (3) We found that litter from wet sites provided a refuge from ant predation; litter from wet sites was not beneficial to caterpillars in the absence of ants. Both abiotic factors (precipitation) and biotic factors (predatory ants) affected the temporal and spatial abundance of caterpillars directly and interactively. Climate models predict that rainfall will become more variable, suggesting that populations of this caterpillar may also become more variable in the future.

Seed-predator satiation and Janzen-Connell effects vary with spatial scales for seed-feeding insects.

BACKGROUND AND AIMS: The Janzen-Connell model predicts that common species suffer high seed predation from specialized natural enemies as a function of distance from parent trees, and consequently as a function of conspecific density, whereas the predator satiation hypothesis predicts that seed attack is reduced due to predator satiation at high seed densities. Pre-dispersal predation by insects was studied while seeds are still on parent trees, which represents a frequently overlooked stage in which seed predation occurs. METHODS: Reproductive tree density and seed production were investigated from ten Quercus serrata populations located in south-west China, quantifying density-dependent pre-dispersal seed predation over two years by three insect groups. KEY RESULTS: Acorn infestation was nearly twice as high in the low-seed year as that in the high-seed year, with considerable spatio-temporal variation in the direction and magnitude of density-dependent pre-dispersal seed predation evident. Across whole populations of trees, a high density of reproductive trees caused predator satiation and reduced insect attack in the high-seed year. Within individual trees, and consistent with the Janzen-Connell model, overall insect seed predation was positively correlated with seed production in the low-seed year. In addition, there was variation among insect taxa, with positive density-dependent seed predation by Curculio weevils in the high-seed year and moths in the low-seed year, but apparent density independence by Cyllorhynchites weevils in both years. CONCLUSIONS: The overall trend of negative density-dependent, pre-dispersal seed predation suggests that predator satiation limited the occurrence of Janzen-Connell effects across Q. serrata populations. Such effects may have large impacts on plant population dynamics and tree diversity, depending on the extent to which they are reduced by counteracting positive density-dependent predation for seeds on individual trees and other factors affecting successful recruitment.

Metapopulation dynamics on ephemeral patches.

A challenge for conservation management is to understand how population and habitat dynamics interact to affect species persistence. In real landscapes, timing and duration of disturbances can vary, and species' responses to habitat changes will depend on how timing of dispersal and reproduction events relate to the landscape temporal structure. For instance, increasing disturbance frequency may promote extinction of species that are unable to appropriately time their reproduction in an ever-changing habitat and favor species that are able to track habitat changes. We developed a mathematical model to compare the effects of pulsed dispersal, initiated by shifts in habitat quality, with temporally continuous dispersal. We tested the effects of habitat (and population) turnover rates on metapopulation establishment, persistence, and long-term patch occupancy. Pulsed dispersal reduced patch occupancy and metapopulation longevity when habitat patches are relatively permanent. In such cases, demographic extinction was the primary form of local extinction. Conversely, when habitat patches are short-lived and new ones are frequently formed, pulsed dispersal promoted rapid colonization, increased occupancy, and prolonged metapopulation persistence. Our results show that species responsiveness to habitat disturbance is critical to metapopulation persistence, having profound implications for the species likely to persist in landscapes with altered disturbance regimes.

Predation and associational refuge drive ontogenetic niche shifts in an arctiid caterpillar.

Despite the ubiquity of ontogenetic niche shifts, their drivers and consequences are poorly understood. Different nutritional requirements and stage-specific physiological limitations have often been offered as explanations for these life history features, but emerging work has demonstrated that top-down factors may also be important. We studied the roles of predation and associational refuge in ontogenetic niche shifts for a holometabolous insect (Platyprepia virginalis), which shifts habitats and host plants to pupate. We examined the effect of pupation site selection across habitats and host plants by late-instar caterpillars on the rate of predation during the relatively vulnerable pupal stage. Studying the ontogenetic transition from mobile caterpillar to non-feeding, sessile pupa allows isolation of top-down effects from bottom-up, nutritional effects. An observational study supported previous findings that feeding caterpillars preferred marsh habitats, but pupating caterpillars preferred prairie habitats. Experiments demonstrated that caterpillars preferred to pupate within a physically defended plant species. Pupation within this defended plant species resulted in reduced predation (an associational refuge), and removal of the physical defense structures negated the reduced-predation effect. This experiment shows that ontogenetic niche shifts can be driven by predation and can involve facilitation by a host plant that provides a refuge to predation. The co-option of plant chemical defenses by animals is widely established. However, finding a clear example in which an animal exploits a plant's physical defense is rare, especially in the context of ontogenetic niche shifts. This work shows that facilitation mediated by refuge from predation provided by host plants and life-stage-dependent predation risk can interact to shape species' distributions.

Ecological effects of environmental change.

This Special Issue of Ecology Letters presents contributions from an international meeting organised by Centre National de la Recherche Scientifique (CNRS) and Ecology Letters on the broad theme of ecological effects of global environmental change. The objectives of these articles are to synthesise, hypothesise and illustrate the ecological effects of environmental change drivers and their interactions, including habitat loss and fragmentation, pollution, invasive species and climate change. A range of disciplines is represented, including stoichiometry, cell biology, genetics, evolution and biodiversity conservation. The authors emphasise the need to account for several key ecological factors and different spatial and temporal scales in global change research. They also stress the importance of ecosystem complexity through approaches such as functional group and network analyses, and of mechanisms and predictive models with respect to environmental responses to global change across an ecological continuum: population, communities and ecosystems. Lastly, these articles provide important insights and recommendations for environmental conservation and management, as well as highlighting future research priorities.

Neonicotinoid exposure in Tricolored Blackbirds (Agelaius tricolor).

There is increasing awareness of the negative ecological and environmental effects of widespread use of pesticides on the landscape. Spillover or drift of pesticides from agricultural areas has been shown to impact species health, reproduction, and trophic dynamics through both direct and indirect mechanisms. Neonicotinoid insecticides are associated with observed declines of insectivorous and grassland birds, and these environmental pollutants are a significant conservation concern for many species that have experienced past or current population declines. Due to the high efficacy of these modern insecticides in depressing local insect populations, insectivorous birds can be negatively impacted by a pesticide-mediated reduction in food supply. Neonicotinoids may act synergistically with other stressors, such as habitat loss, to exacerbate threats to species or population viability. The Tricolored Blackbird is an insectivorous grassland bird of conservation concern in California, USA. Due to the high association of this species with agricultural habitats, we sought to quantify the amount of neonicotinoid residues in Tricolored Blackbird carcasses as a first step in assessing how this species may be impacted by pesticides. Out of 85 salvaged carcasses sampled (N = 24 adults, N = 3 fledglings, and N = 58 nestlings), only two contained detectable levels of target compounds. These were an adult and one nestling that contained clothianidin residue (40 ppb and 7 ppb, respectively); both of these birds were salvaged from breeding colonies associated with dairy farms in Kern County, California. We suggest that further work is needed to assess neonicotinoid exposure of Tricolored Blackbirds in dairy-associated breeding colonies.

Spatial habitat heterogeneity influences host-pathogen dynamics in a patchy population of Ranchman's tiger moth.

Host-pathogen dynamics are influenced by many factors that vary locally, but models of disease rarely consider dynamics across spatially heterogeneous environments. In addition, theory predicts that dispersal will influence host-pathogen dynamics of populations that are linked, although this has not been examined empirically in natural systems. We examined the spatial dynamics of a patchy population of tiger moths and its baculovirus pathogen, in which habitat type and weather influence dynamics. Theoretical models of host-baculovirus dynamics predict that such variation in dynamics between habitat types could be driven by a range of factors, of which we predict two are likely to be operating in this system: (1) differences in the environmental persistence of pathogens or (2) differences in host intrinsic rates of increase. We used time series models and monitored infection rates of hosts to characterize population and disease dynamics and distinguish between these possibilities. We also examined the role of host dispersal (connectivity) and weather as important contributors to dynamics, using time series models and experiments. We found that the population growth rate was higher, delayed density dependence was weaker, and long-period oscillations had lower amplitudes in high-quality habitat patches. The infection rate was higher on average in high-quality habitat, and this was likely to have been driven by higher mean population densities and no differences in pathogen persistence in different habitats (delayed density dependence). Time series modeling and experiments also showed an interactive effect of temperature and precipitation on moth population growth rates (likely caused by variation in host plant quality and quantity), and an effect of connectivity. Our results showed that spatial heterogeneity, connectivity, climate, and their interactions were important in driving host-baculovirus dynamics. In particular, our study found that connected patches and spatial heterogeneity generated differences in dynamics that only partially aligned with theoretical predictions.

Impacts of top predators and humans on the mammal communities of recovering temperate forest regions.

Top predators are important drivers in shaping ecological community structure via top-down effects. However, the ecological consequences and mechanisms of top predator loss under accelerated human impacts have rarely been quantitatively assessed due to the limited availability of long-term community data. With increases in top predator populations in northern China over the past two decades, forests with varying densities of top predators and humans provide an opportunity to study their ecological effects on mammal communities. We hypothesized a priori of conceptual models and tested these using structural equation models (SEMs) with multi-year camera trap data, aiming to reveal the underlying independent ecological effects of top predators (tigers, bears, and leopards) and humans on mammal communities. We used random forest models and correlations among species pairs to validate results. We found that top predator reduction could be related to augmented populations of large ungulates ("large ungulate release") and mesopredators ("mesopredator release"), consistent with observations of mammal communities in other ecosystems. Additionally, top predator reduction could be related to reduced small mammal abundance. Hierarchical SEMs identified three bottom-up pathways from forest quality to human activities, large ungulates, and some small mammals, and five top-down pathways from human activities and top predators to some small mammals, large ungulates, and mesopredators. Furthermore, our results suggest that humans showed predominant top-down effects on multiple functional groups, partially replacing the role of top predators, rather than be mediated by them; effects of humans and top predators appeared largely independent. Effects of humans on top predators were non-significant. This study provides novel insights into the effects of top predators and humans as super-predators on mammal communities in forest ecosystems and presents cues of bottom-up effects that can be translated into actionable management plans for improving forest quality, thereby supporting top predator recovery and work/life activities of local people.

Interactions among shrub cover and the soil microclimate may determine future Arctic carbon budgets.

Arctic and Boreal terrestrial ecosystems are important components of the climate system because they contain vast amounts of soil carbon (C). Evidence suggests that deciduous shrubs are increasing in abundance, but the implications for ecosystem C budgets remain uncertain. Using midsummer CO(2) flux data from 21 sites spanning 16° of latitude in the Arctic and Boreal biomes, we show that air temperature explains c. one-half of the variation in ecosystem respiration (ER) and that ER drives the pattern in net ecosystem CO(2) exchange across ecosystems. Woody sites were slightly stronger C sinks compared with herbaceous communities. However, woody sites with warm soils (> 10 °C) were net sources of CO(2) , whereas woody sites with cold soils (< 10 °C) were strong sinks. Our results indicate that transition to a shrub-dominated Arctic will increase the rate of C cycling, and may lead to net C loss if soil temperatures rise.

Spatial clustering of habitat structure effects patterns of community composition and diversity.

Natural ecosystems often show highly productive habitats that are clustered in space. Environmental disturbances are also often nonrandomly distributed in space and are either intrinsically linked to habitat quality or independent in occurrence. Theoretical studies predict that configuration and aggregation of habitat patch quality and disturbances can affect metacommunity composition and diversity, but experimental evidence is largely lacking. In a metacommunity experiment, we tested the effects of spatially autocorrelated disturbance and spatial aggregation of patch quality on regional and local richness, among-community dissimilarity, and community composition. We found that spatial aggregation of patch quality generally increased among-community dissimilarity (based on two measures of beta diversity) of communities containing protozoa and rotifers in microcosms. There were significant interacting effects of landscape structure and location of disturbances on beta diversity, which depended in part on the specific beta diversity measures used. Effects of disturbance on composition and richness in aggregated landscapes were generally dependent on distance and connectivity among habitat patches of different types. Our results also show that effects of disturbances in single patches cannot directly be extrapolated to the landscape scale: the predictions may be correct when only species richness is considered, but important changes in beta diversity may be overlooked. There is a need for biodiversity and conservation studies to consider the spatial aggregation of habitat quality and disturbance, as well as connectivity among spatial aggregations.

The importance of host plant limitation for caterpillars of an arctiid moth (Platyprepia virginalis) varies spatially.

Spatial dynamic theories such as source-sink models frequently describe habitat-specific demographies, yet there are surprisingly few field studies that have examined how and why interacting species vary in their dynamics across multiple habitat types. We studied the spatial pattern of interaction between a chewing herbivore and its primary larval host plant in two habitat types. We found that the interaction between an arctiid caterpillar (Platyprepia virginalis) and its host (Lupinus arboreus) differed in wet vs. upland dry habitats, as did yearly population dynamics for the caterpillar. In upland sites, there was a strong positive relationship between lupine cover and the abundance of caterpillars although this relationship was not apparent in wet sites. Additionally, in wet sites, caterpillar populations were larger and less variable across years. Caterpillars appeared to exhibit source-sink dynamics, with the time-averaged finite growth rate lamda > 1 in wet sites (sources), lamda < 1 in upland dry sites (sinks), and predominant source-to-sink movement of late-instar caterpillars. Populations in upland dry sites also went locally extinct in years of low regional abundance. Emigration from wet sites could potentially explain the lack of coupling of herbivore and host plant dynamics in these sites. These results indicate that movement and other factors affecting demography are habitat-specific and have important implications for trophic control. Acknowledging such complexity makes simple models of trophic control seem overly general but may allow us to formulate more broadly applicable ecological models.