Effects of flyway‐wide weather conditions and breeding habitat on the breeding abundance of migratory boreal waterbirds

Anthropogenic habitat loss and climate change are among the major threats to biodiversity. Bioclimatic zones such as the boreal and arctic regions are undergoing rapid environmental change, which will likely trigger changes in wildlife communities. Disentangling the effects of different drivers of environmental change on species is fundamental to better understand population dynamics under changing conditions. Therefore, in this study we investigate the synergistic effect of winter and summer weather conditions and habitat type on the abundance of 17 migratory boreal waterbird species breeding in Finland using three decades (1986–2015) of count data. We found that above‐average temperatures and precipitations across the western and northern range of the wintering grounds have a positive impact on breeding numbers in the following season, particularly for waterbirds breeding in eutrophic wetlands. Conversely, summer temperatures did not seem to affect waterbird abundance. Moreover, waterbird abundance was higher in eutrophic than in oligotrophic wetlands, but long term trends indicated that populations are decreasing faster in eutrophic than in oligotrophic wetlands. Our results suggest that global warming may apparently benefit waterbirds, e.g. by increased winter survival due to more favourable winter weather conditions. However, the observed population declines, particularly in eutrophic wetlands, may also indicate that the quality of breeding habitat is rapidly deteriorating through increased eutrophication in Finland which override the climatic effects. The findings of this study highlight the importance of embracing a holistic approach, from the level of a single catchment up to the whole flyway, in order to effectively address the threats that waterbirds face on their breeding as well as wintering grounds.     

Spatial heterogeneity in climate change effects decouples the long‐term dynamics of wild reindeer populations in the high Arctic

The ‘Moran effect’ predicts that dynamics of populations of a species are synchronized over similar distances as their environmental drivers. Strong population synchrony reduces species viability, but spatial heterogeneity in density dependence, the environment, or its ecological responses may decouple dynamics in space, preventing extinctions. How such heterogeneity buffers impacts of global change on large‐scale population dynamics is not well studied. Here, we show that spatially autocorrelated fluctuations in annual winter weather synchronize wild reindeer dynamics across high‐Arctic Svalbard, while, paradoxically, spatial variation in winter climate trends contribute to diverging local population trajectories. Warmer summers have improved the carrying capacity and apparently led to increased total reindeer abundance. However, fluctuations in population size seem mainly driven by negative effects of stochastic winter rain‐on‐snow (ROS) events causing icing, with strongest effects at high densities. Count data for 10 reindeer populations 8–324 km apart suggested that density‐dependent ROS effects contributed to synchrony in population dynamics, mainly through spatially autocorrelated mortality. By comparing one coastal and one ‘continental’ reindeer population over four decades, we show that locally contrasting abundance trends can arise from spatial differences in climate change and responses to weather. The coastal population experienced a larger increase in ROS, and a stronger density‐dependent ROS effect on population growth rates, than the continental population. In contrast, the latter experienced stronger summer warming and showed the strongest positive response to summer temperatures. Accordingly, contrasting net effects of a recent climate regime shift—with increased ROS and harsher winters, yet higher summer temperatures and improved carrying capacity—led to negative and positive abundance trends in the coastal and continental population respectively. Thus, synchronized population fluctuations by climatic drivers can be buffered by spatial heterogeneity in the same drivers, as well as in the ecological responses, averaging out climate change effects at larger spatial scales.     

A mean field model for competition: from neutral ecology to the Red Queen

Individual species are distributed inhomogeneously over space and time, yet, within large communities of species, aggregated patterns of biodiversity seem to display nearly universal behaviour. Neutral models assume that an individual's demographic prospects are independent of its species identity. They have successfully predicted certain static, time‐independent patterns. But they have generally failed to predict temporal patterns, such as species ages or population dynamics. We construct a new, multispecies framework incorporating competitive differences between species, and assess the impact of this competition on static and dynamic patterns of biodiversity. We solve this model exactly for the special case of a Red Queen hypothesis, where fitter species are continually arising. The model predicts more realistic species ages than neutral models, without greatly changing predictions for static species abundance distributions. Our modelling approach may allow users to incorporate a broad range of ecological mechanisms.     

Non-linear feedback processes and a latitudinal gradient in the climatic effects determine green spruce aphid outbreaks in the UK

The role of climatic fluctuations in determining the dynamics of insect populations has been a classical problem in population ecology. Here, we use long-term annual data on green spruce aphid populations at nine localities in the UK for determining the importance of endogenous processes, local weather and large-scale climatic factors. We rely on diagnostic and modelling tools from population dynamic theory to analyse these long-term data and to determine the role of the North Atlantic Oscillation (NAO) and local weather as exogenous factors influencing aphid dynamics. Our modelling suggests that the key elements determining population fluctuations in green spruce aphid populations in the UK are the strong non-linear feedback structure, the high potential for population growth and the effects of winter and spring weather. The results indicate that the main effect of the NAO on green spruce aphid populations is operating through the effect of winter temperatures on the maximum per capita growth rate (R_m). In particular, we can predict quite accurately the occurrence of an outbreak by using a simple logistic model with weather as a perturbation effect. However, model predictions using different climatic variables showed a clear geographical signature. The NAO and winter temperature were best for predicting observed dynamics toward the southern localities, while spring temperature was a much better predictor of aphid dynamics at northern localities. Although aphid species are characterized by complex life-cycles, we emphasize the value of simple and general population dynamic models in predicting their dynamics.     

Signatures of large-scale and local climates on the demography of white-tailed ptarmigan in Rocky Mountain National Park,Colorado, USA

Global climate change may impact wildlife populations by affecting local weather patterns, which, in turn, can impact a variety of ecological processes. However, it is not clear that local variations in ecological processes can be explained by large-scale patterns of climate. The North Atlantic oscillation (NAO) is a large-scale climate phenomenon that has been shown to influence the population dynamics of some animals. Although effects of the NAO on vertebrate population dynamics have been studied, it remains uncertain whether it broadly predicts the impact of weather on species. We examined the ability of local weather data and the NAO to explain the annual variation in population dynamics of white-tailed ptarmigan ( Lagopus leucurus) in Rocky Mountain National Park, USA. We performed canonical correlation analysis on the demographic subspace of ptarmigan and local-climate subspace defined by the empirical orthogonal function (EOF) using data from 1975 to 1999. We found that two subspaces were significantly correlated on the first canonical variable. The Pearson correlation coefficient of the first EOF values of the demographic and local-climate subspaces was significant. The population density and the first EOF of local-climate subspace influenced the ptarmigan population with 1-year lags in the Gompertz model. However, the NAO index was neither related to the first two EOF of local-climate subspace nor to the first EOF of the demographic subspace of ptarmigan. Moreover, the NAO index was not a significant term in the Gompertz model for the ptarmigan population. Therefore, local climate had stronger signature on the demography of ptarmigan than did a large-scale index, i.e., the NAO index. We conclude that local responses of wildlife populations to changing climate may not be adequately explained by models that project large-scale climatic patterns.     

Synchrony and geographical variation of the spruce bark beetle (<Emphasis Type="Italic">Ips typographus</Emphasis>) during a non-epidemic period

Spatio-temporal analyses of non-epidemic bark beetle populations may provide insight in dynamics predisposing for outbreaks. The present article presents a spatio-temporal analysis of the population dynamics of Ips typographus based on pheromone trap data from southeast and mid-Norway in the post-epidemic period 1979–2002. The analyses include regression analyses, hierarchical cluster analysis, and analysis of spatial synchrony of beetle time series and climatic data by means of nonparametric spatial covariance functions. The mean abundance of beetles declined linearly with latitude. In addition, the time series means were higher in areas with high forest productivity and rocky soils predisposed to drought. The time series patterns differed significantly between northern and southern study areas. The regional synchrony of the time series was fairly high (0.38), indicating that some large-scale climatic factor may influence the dynamics. Windfelling was the external variable showing the most parallel pattern of correlation to the beetle dynamics. We thus posit that large windfall events may be a major instigator and synchronizer of beetle outbreaks in areas subjected to regionalized weather systems.     

Predicting small‐mammal responses to climatic warming: autecology,geographic range,and the Holocene fossil record

Forecasting how species will respond to climatic change requires knowledge of past community dynamics. Here we use time‐series data from the small‐mammal fossil records of two caves in the Great Basin of the American West to evaluate how contrasting and variable local paleoclimates have shaped small‐mammal abundance dynamics over the last ～7500 years of climatic change. We then predict how species and communities will respond to future scenarios of increased warming and aridity coupled with continued spread of an invasive annual grass (Bromus tectorum). We find that most community‐level responses to climatic change occur in the mammalian abundance structure at both sites; the dominance of the community by individuals from species with a southern geographic affinity increases with climatic warming. This suggests that responses occurred in situ rather than by the immigration of new taxa over this time interval. Despite predictability at the community‐scale, species‐level relationships between abundance and climate are variable and are not necessarily explained by a species' geographic affinity. Species present at both sites, however, exhibit remarkably similar responses to climate at each site, indicating that species autecology (specifically dietary functional group) is important in determining response to climatic warming. Regression‐tree analyses show remarkable concordance between the two cave faunas and highlight the importance of a granivorous dietary strategy in this desert ecosystem. Under projections of increased temperature and decreased precipitation over the next 50 years, our results indicate that granivores should thrive as communities become more dominated by individuals with a southern geographic affinity. Granivores, however, are negatively impacted by the invasion of cheatgrass. The last century of anthropogenic impacts has thus placed granivores at a greater risk of extinction than predicted under climate‐only scenarios.     

Changes in distribution of waterbirds following prolonged drought reflect habitat availability in coastal and inland regions

Provision of suitable habitat for waterbirds is a major challenge for environmental managers in arid and semiarid regions with high spatial and temporal variability in rainfall. It is understood in broad terms that to survive waterbirds must move according to phases of wet–dry cycles, with coastal habitats providing drought refugia and inland wetlands used during the wet phase. However, both inland and coastal wetlands are subject to major anthropogenic pressures, and the various species of waterbird may have particular habitat requirements and respond individualistically to spatiotemporal variations in resource distribution. A better understanding of the relationships between occurrence of waterbirds and habitat condition under changing climatic conditions and anthropogenic pressures will help clarify patterns of habitat use and the targeting of investments in conservation. We provide the first predictive models of habitat availability between wet and dry phases for six widely distributed waterbird species at a large spatial scale. We first test the broad hypothesis that waterbirds are largely confined to coastal regions during a dry phase. We then examine the contrasting results among the six species, which support other hypotheses erected on the basis of their ecological characteristics. There were large increases in area of suitable habitat in inland regions in the wet year compared with the dry year for all species, ranging from 4.14% for Australian White Ibis to 31.73% for Eurasian Coot. With over half of the suitable habitat for three of the six species was located in coastal zones during drought, our study highlights the need to identify and conserve coastal drought refuges. Monitoring of changes in extent and condition of wetlands, combined with distribution modeling of waterbirds, will help support improvements in the conservation and management of waterbirds into the future.     

Estimating plant responses to climate by direct gradient analysis and geographic distribution analysis

We characterised the climatic behaviour of 53 woody species in terms of the climatic factors that play the main role in controlling species distribution in the study area. Floristic and climatic data were obtained from 150 stands in sites under climatic control (i.e. eu-climatopes). The sampling strategy used allowed a reliable match between floristic and climatic observations. Different methods of frequency analysis and goodness-of-fit tests were used to identify associations between species occurrence and climatic characteristics. The species' responses were summarised by statistics describing ecological preferences and amplitudes, and species were grouped accordingly. A Gaussian response model was fitted to the abundance data along the main climatic gradients for selected species and response surfaces were derived by spatial analysis for a set of indicator species. Frequency analysis methods detected 42 indicator taxa for the Baudiere's Qe drought index, and lower numbers, 34 and 22, respectively, for the mean minimum coldest-month temperature and the daily temperature range in the coldest month. Goodness-of-fit tests revealed a lower number of ecological profiles with statistically significant deviations from equidistribution. We discuss the relative performance of the different methods and suggest that the combined use of statistical tests and frequency analyses may improve estimation of the environmental requirements of species. We also recommend using the species' responses to key environmental factors as reliable criteria in the definition of plant functional types. This revised version was published online in August 2006 with corrections to the Cover Date.     

FREQ-Seq: A Rapid,Cost-Effective,Sequencing-Based Method to Determine Allele Frequencies Directly from Mixed Populations

Understanding evolutionary dynamics within microbial populations requires the ability to accurately follow allele frequencies through time. Here we present a rapid, cost-effective method (FREQ-Seq) that leverages Illumina next-generation sequencing for localized, quantitative allele frequency detection. Analogous to RNA-Seq, FREQ-Seq relies upon counts from the >10⁵ reads generated per locus per time-point to determine allele frequencies. Loci of interest are directly amplified from a mixed population via two rounds of PCR using inexpensive, user-designed oligonucleotides and a bar-coded bridging primer system that can be regenerated in-house. The resulting bar-coded PCR products contain the adapters needed for Illumina sequencing, eliminating further library preparation. We demonstrate the utility of FREQ-Seq by determining the order and dynamics of beneficial alleles that arose as a microbial population, founded with an engineered strain of Methylobacterium, evolved to grow on methanol. Quantifying allele frequencies with minimal bias down to 1% abundance allowed effective analysis of SNPs, small in-dels and insertions of transposable elements. Our data reveal large-scale clonal interference during the early stages of adaptation and illustrate the utility of FREQ-Seq as a cost-effective tool for tracking allele frequencies in populations.     

Concordant population dynamics of Lepidoptera herbivores in a forest ecosystem

It is frequently assumed that population fluctuations are largely independent within a community of trophically‐similar species, but this need not be so. If population fluctuations are partly synchronized or concordant, this will produce interannual variability in the community's aggregate abundance and generate temporal variance in ecosystem structure. We studied the community of Lepidoptera inhabiting northern hardwood forests in New Hampshire, USA, to evaluate the hypothesis that fluctuations in consumer communities can arise from concordant dynamics of constituent populations. Interannual comparisons of moth abundances for >75 species sampled at three sites over four years revealed that concordant dynamics contribute strongly to interannual variability in the abundance of consumers. A conspicuous decline in community abundance from 2004 to 2005 was the result of predominantly negative population growth rates of the component species, while an increase in community abundance from 2006 to 2007 was the result of predominantly positive population growth rates. Population dynamics most strongly linked species that feed in the early season (perhaps due to shared responses to climatic effects), but not species that might share natural enemies or host plants. The observed concordant dynamics introduced conspicuous temporal variation in the abundance of primary consumers relative to plants and secondary consumers, thereby altering the forest's trophic structure. Such variance in the aggregate abundance of forest primary consumers could generate time‐lagged fluctuations in abundances of secondary consumers and will generally have important consequences for ecosystem properties and processes that are nonlinear functions of consumer abundance, such as plant community structure and nutrient cycling.     

Reconciling the influence of global climate phenomena on macrofaunal temporal dynamics at a variety of spatial scales

Determining the relative importance of environmental forces on population dynamics is a fundamental question for ecologists. Growing concern over the ecological effects of climate change emphasizes the importance of defining whether broad-scale environmental forces uniformly act upon local populations (hierarchy theory) or cross-scale interactions influence local responses (multiscale theory). This study analyses 13 years of data on species abundances at six sites within a large harbour to determine the effect of the El Niño Southern Oscillation (ENSO). Environmental variables both directly and indirectly related to ENSO were observed to be important predictors of the temporal dynamics of abundance in many species, but the observed effects were not consistent across sites or species. While nearly all species were affected by large temporal and spatial scale variability, smaller temporal scale, location-specific environmental variables (such as wind-generated wave exposure and turbidity) were also generally important, increasing the variability explained by our models by up to 25%. As with many other broad-scale variables, generality of response to ENSO is affected by interactions across time and space with smaller scale heterogeneity. This study therefore suggests that the degree of interaction between broad-scale climatic factors, such as ENSO, with smaller scale variability, will determine the consistency of responses over large spatial scales, and control our ability to predict effects of climate change on coastal and estuarine communities.     

Humpback whale diets respond to variance in ocean climate and ecosystem conditions in the California Current

Large, migratory predators are often cited as sentinel species for ecosystem processes and climate‐related changes, but their utility as indicators is dependent upon an understanding of their response to environmental variability. Documentation of the links between climate variability, ecosystem change and predator dynamics is absent for most top predators. Identifying species that may be useful indicators and elucidating these mechanistic links provides insight into current ecological dynamics and may inform predictions of future ecosystem responses to climatic change. We examine humpback whale response to environmental variability through stable isotope analysis of diet over a dynamic 20‐year period (1993–2012) in the California Current System (CCS). Humpback whale diets captured two major shifts in oceanographic and ecological conditions in the CCS. Isotopic signatures reflect a diet dominated by krill during periods characterized by positive phases of the North Pacific Gyre Oscillation (NPGO), cool sea surface temperature (SST), strong upwelling and high krill biomass. In contrast, humpback whale diets are dominated by schooling fish when the NPGO is negative, SST is warmer, seasonal upwelling is delayed and anchovy and sardine populations display increased biomass and range expansion. These findings demonstrate that humpback whales trophically respond to ecosystem shifts, and as a result, their foraging behavior is a synoptic indicator of oceanographic and ecological conditions across the CCS. Multi‐decadal examination of these sentinel species thus provides insight into biological consequences of interannual climate fluctuations, fundamental to advancing ecosystem predictions related to global climate change.     

Species with more volatile population dynamics are differentially impacted by weather

Climatic variation has been invoked as an explanation of population dynamics for a variety of taxa. Much work investigating the link between climatic forcings and population fluctuation uses single-taxon case studies. Here, we conduct comparative analyses of a multi-decadal dataset describing population dynamics of 50 co-occurring butterfly species at 10 sites in Northern California. Specifically, we explore the potential commonality of response to weather among species that encompass a gradient of population dynamics via a hierarchical Bayesian modelling framework. Results of this analysis demonstrate that certain weather conditions impact volatile, or irruptive, species differently as compared with relatively stable species. Notably, precipitation-related variables, including indices of the El Niño Southern Oscillation, have a more pronounced impact on the most volatile species. We hypothesize that these variables influence vegetation resource availability, and thus indirectly influence population dynamics of volatile taxa. As one of the first studies to show a common influence of weather among taxa with similar population dynamics, the results presented here suggest new lines of research in the field of biotic–abiotic interactions.     

Bacterial colonization and extinction on marine aggregates: stochastic model of species presence and abundance

Organic aggregates provide a favorable habitat for aquatic microbes, are efficiently filtered by shellfish, and may play a major role in the dynamics of aquatic pathogens. Quantifying this role requires understanding how pathogen abundance in the water and aggregate size interact to determine the presence and abundance of pathogen cells on individual aggregates. We build upon current understanding of the dynamics of bacteria and bacterial grazers on aggregates to develop a model for the dynamics of a bacterial pathogen species. The model accounts for the importance of stochasticity and the balance between colonization and extinction. Simulation results suggest that while colonization increases linearly with background density and aggregate size, extinction rates are expected to be nonlinear on small aggregates in a low background density of the pathogen. Under these conditions, we predict lower probabilities of pathogen presence and reduced abundance on aggregates compared with predictions based solely on colonization. These results suggest that the importance of aggregates to the dynamics of aquatic bacterial pathogens may be dependent on the interaction between aggregate size and background pathogen density, and that these interactions are strongly influenced by ecological interactions and pathogen traits. The model provides testable predictions and can be a useful tool for exploring how species‐specific differences in pathogen traits may alter the effect of aggregates on disease transmission.     

崇明东滩冬季水鸟生态位分析

依据2003年冬季对崇明东滩自然保护区越冬水鸟的种类、数量、生境类型分布的最新调查数据,以及上海农林局10年来积累的越冬水鸟食性与形态的数据,采取聚类分析方法对崇明东滩冬季鸟类的群落生态进行研究,从鸟类的取食空间生态位、食性生态位以及形态生态位三个维度确认其生态资源分配状况,并由此确认了占据优势种群地位的鸟类在不同生态位维度上的分离是群落结构处于稳定状态的主要原因。     

Little evidence for climate effects on local-scale structure and dynamics of California kelp forest communities

Climatic variables such as temperature and precipitation play an important role in controlling local and regional scale differences in population dynamics and species distributions, and large-scale climatic events such as El Niño southern oscillation (ENSO) have been shown to affect population dynamics of key species in many ecosystems, particularly in kelp forests. Few studies have been able to evaluate the consequences of climate variables on the structure and dynamics of biological communities, in large part because the lack of data at appropriate spatial and temporal scales has made it difficult to adequately address local-scale responses of species and communities to such events over relevant time scales. Here, we combined an unprecedented dataset of kelp forest species' abundances from the Channel Islands, California with data for several local, regional, and global scale climatic variables to evaluate the temporal and spatial scale at which one can detect community-wide effects of climate variables, in particular ENSO events. We found large and significant local-scale differences in community structure, but these differences were not related to differences in climatic variables. Moreover, giant kelp abundance, which has been shown to be highly sensitive to water temperature and storm disturbance, was a poor predictor of community differences, and all communities tended to decline in abundance over the 20-year sampling period, suggesting a press perturbation to the system such as PDO cycles or sustained fishing pressure. Although ENSO events can have dramatic impacts on the abundance and distribution of giant kelp itself across the range of the species, such events appear to have little effect on local-scale kelp forest community structure or dynamics.     

Long-term variation in trematode (Trematoda, Digenea) component communities associated with intertidal gastropods is linked to abundance of final hosts

Trematodes are abundant parasites essential for maintaining stability of marine intertidal ecosystems. Despite the great ecological significance of trematodes, long-term dynamics of their communities remains practically unstudied. This study, based on 12-year-long seasonal monitoring of infection of Littorina and Hydrobia snails with 16 trematode species in the White Sea, aimed to reveal factors determining long-term variation in infection of intertidal snails by trematode parthenitae and larvae. Using the state-of-the-art method of singular spectrum analysis, we revealed trends in this variation and assessed their significance. Interestingly, these trends were not associated with oceanic and climatic parameters but were mostly determined by changes in abundance of the trematode final hosts. Moreover, the prevalence trends turned out to be connected with both large-scale events and local factors at the scale of the intertidal site. The main factors determining the long-term dynamics of the trematode component communities in the study area were the decreasing abundance of birds due to growing anthropogenic disturbance and the increasing abundance of the three-spined stickleback. The analysis of long-term trends of trematode prevalence in intertidal snails may be a sensitive indicator of the abundance dynamics of final hosts in the coastal areas of temperate and northern seas.     

Analysis of trophic interactions reveals highly plastic response to climate change in a tri-trophic High-Arctic ecosystem

As a response to current climate changes, individual species have changed various biological traits, illustrating an inherent phenotypic plasticity. However, as species are embedded in an ecological network characterised by multiple consumer–resource interactions, ecological mismatches are likely to arise when interacting species do not respond homogeneously. The approach of biological networks analysis calls for the use of structural equation modelling (SEM), a multidimensional analytical setup that has proven particularly useful for analysing multiple interactions across trophic levels. Here we apply SEM to a long-term dataset from a High-Arctic ecosystem to analyse how phenological responses across three trophic levels are coupled to snowmelt patterns and how changes may cascade through consumer–resource interactions. Specifically, the model included the effect of snowmelt on a High-Arctic tri-trophic system of flowers, insects and waders (Charadriiformes), with latent factors representing phenology (timing of life history events) and performance (abundance or reproduction success) for each trophic level. The effects derived from the model demonstrated that the time of snowmelt directly affected plant and arthropod phenology as well as the performance of all included trophic levels. Additionally, timing of snowmelt appeared to indirectly influence wader phenology as well as plant, arthropod and wader performance through effects on adjacent trophic levels and lagged effects. The results from the tri-trophic community presented here emphasise that effects of climate on species in consumer–resource systems may propagate through trophic levels.     

Multiple environmental changes interact to modify species dynamics and invasion rates

Multiple aspects of the environment often change at the same time, influencing populations directly by modifying their physiology, but also indirectly by influencing other interacting species. The impacts of each environmental change upon population dynamics are usually assumed be independent of the state of other aspects of the environment, despite evidence at the individual level indicating that the combined impacts are often non‐additive. The importance of indirect effects mediated through community interactions also has high uncertainty. We used experimental microcosms to determine whether environmental factors interact to affect species dynamics and the relative importance of direct and indirect effects on species dynamics. We factorially manipulated three aspects of the environment (temperature, food availability and salinity) and examined reciprocal invasions of competing protist species under each environment. Experimental observations were used to parameterize a dynamic model of the system. Using this model and a novel variance decomposition method, we examined the mechanisms by which environmental changes altered species invasion rates. The three environmental factors interacted when modifying species growth rates, intra‐ and interspecific competition, causing the impact of each environmental change on species dynamics to depend crucially on the state of other aspects of the environment. Indirect changes in the abundance of the resident competitor and its interspecific competitive ability were the main cause of environmental driven variation in invasion rates, whilst direct effects on species intrinsic growth rates were relatively unimportant. This indicates that, to understand and ultimately predict species and community responses to multiple environmental changes, we should consider their joint impacts and the mechanisms by which they interact to modify key ecological processes such as competition.