Phylogenetic structure of mammal assemblages at large geographical scales: linking phylogenetic community ecology with macroecology

Phylogenetic community ecology seeks to explain the processes involved in the formation of species assemblages by analysing their phylogenetic structure, and to date has focused primarily on local-scale communities. Macroecology, on the other hand, is concerned with the structure of assemblages at large geographical scales, but has remained largely non-phylogenetic. Analysing the phylogenetic structure of large-scale assemblages provides a link between these two research programmes. In this paper, I ask whether we should expect large-scale assemblages to show significant phylogenetic structure, by outlining some of the ecological and macroevolutionary processes that may play a role in assemblage formation. As a case study, I then explore the phylogenetic structure of carnivore assemblages within the terrestrial ecoregions of Africa. Many assemblages at these scales are indeed phylogenetically non-random (either clustered or overdispersed). One interpretation of the observed patterns of phylogenetic structure is that many clades underwent rapid biome-filling radiations, followed by diversification slowdown and competitive sorting as niche space became saturated.     

Cross-scale drivers of plant trait distributions in a fragmented forest landscape

During community assembly, plant functional traits are under selective pressure from processes operating at multiple spatial scales. However, in fragmented landscapes, there is little understanding of the relative importance of local-, patch- and landscape-scale processes in shaping trait distributions. Here, we investigate cross-scale influences of landscape change on traits that dictate plant life history strategies in re-assembling plant communities in a fragmented landscape in eastern China. Using forest dynamics plots (FDPs) on 29 land-bridge islands in which all woody plants have been georeferenced and identified to species, we characterized and derived two composite measures of trait variation, representing variation across the leaf economics spectrum and plant size. We then tested for trait shifts in response to local-, patch- and landscape-scale factors, and their potential cross-scale interactions. We found substantial community-wide trait changes along local-scale gradients (i.e. forest edge to interior): more acquisitive leaf economic traits and larger sized species occurred at edges, with a significant increase in trait means and trait range. Moreover, there were significant cross-scale interaction effects of patch and landscape variables on local-scale edge effects. Altered spatial arrangement of habitat in the surrounding landscape (i.e. declining habitat amount and increasing patch density), as well as decreasing area at the patch level, exacerbated edge effects on traits distributions. We suggest that synergistic interactions of landscape- and patch-scale processes, such as dispersal limitation, on local-scale environmental filtering at edges, together shape the spatial distributions of plant life history strategies in fragmented plant communities.     

Does habitat availability determine geographical-scale abundances of coral-dwelling fishes?

The role of local-scale processes in determining large-scale patterns of abundance is a key issue in ecology. To test whether habitat use determines local and large-scale patterns of abundance of obligate coral-dwelling fishes (genus Gobiodon), the author compared habitat availability with the abundance of four species, G. axillaris, G. brochus, G. histrio, and G. quinquestrigatus, among four locations, from the southern Great Barrier Reef to northern Papua New Guinea. Habitat availability, measured at tens of meters, explained 47-65% of the variation in abundance of these species among geographic locations spanning over 2,000 km. Therefore, local-scale patterns of habitat use appear to determine much larger-scale patterns of abundance in these habitat-specialist fish. The abundances of all species, except G. brochus, were also closely associated with particular exposure regimes, independently of the abundance of corals. Broad-scale habitat selection for reef types within locations can most easily explain this pattern. The abundances of all species, except G. brochus, also varied among geographic locations, independently of coral abundances. Therefore, the abundances of these species are influenced by either geographic variation in local-scale processes that was not measured, or additional processes acting at very large spatial scales.     

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.     

Coral diversity across a disturbance gradient in the Pulau Seribu reef complex off Jakarta, Indonesia

Very few coral reefs are located close enough to metropolitan cities to study the influence of large urban populations on reef communities. Here, we compare the impact of a large-scale disturbance gradient with local-scale disturbance on coral richness, cover, and composition in the Jakarta Bay and Pulau Seribu reef complex off Jakarta, Indonesia. We found no effect of local land-use type of coral reef islands on richness, composition or cover, nor did taxon richness differ among zones at the large-scale. There was, however, a pronounced difference in composition and coral cover among zones. Cover was very low and composition differed markedly in the near-shore zone 1 (Jakarta Bay) where human-induced disturbance is most intense. Cover was highest in the outlying reefs of zone 3. The highly perturbed zone 1 reefs were, furthermore, distinguished by the virtual absence of otherwise abundant coral taxa such as Acropora hyacinthus and Porites rus and the prevalence of taxa such as Oulastrea crispata and Favia maxima. Almost 60% of the spatial variation in composition was related to variation in shelf depth and island size. The importance of shelf depth was related to the prevalence of a strong environmental gradient in reef depth, pollution, and mechanical reef disturbance and salinity from Jakarta Bay to the outlying reefs. Although there was a significant univariate relationship between spatial variation in composition and distance, this did not enter into the multivariate model, except when presence–absence data was used, indicating that environmental processes are the primary structuring forces in determining local coral assemblage composition across the Pulau Seribu complex.     

Local Explanations of Landscape Patterns: Can Analytical Approaches Approximate Simulation Models of Spatial Processes?

Research over the last 100 years has demonstrated the importance of space for ecological processes. Given this importance, it may seem natural to start investigations into broad-scale ecological processes with a comprehensive, broad-scale spatial map. Here we argue that it may sometimes be possible to answer important questions about spatial processes using crude spatial information obtained when a comprehensive map is not available. To present our argument, we first develop a simple simulation model for a perennial plant reproducing and dying on a landscape with different arrangements of suitable and unsuitable sites. We then develop a simple, analytical approximation to predict the fraction of suitable sites that are occupied by the simulated plants. The analytical approximation summarizes the spatial map by using a single parameter that gives the probability that a site adjacent to a suitable site is suitable. Comparing the predictions of both approaches highlights three points: (a) The role of the spatial environment in ecological processes may play out at the local scale. Therefore, studying the local-scale processes may provide insights into landscape patterns. (b) The predictions from the analytical approximation fail noticeably when suitable sites are rare and are distributed randomly (rather than clumped) on the map. In these situations, patches of interconnected suitable sites are very small, and populations within small patches may go extinct via demographic stochasticity. This illustrates how analytical approximations can be used to identify cases when local-scale spatial processes are not sufficient to understand the ecological consequences of space. (c) For many natural systems, constructing the appropriate environmental map needed to study ecological processes is difficult or impossible. However, summary characteristics such as those employed by the analytical approximation may be estimated directly in nature. Therefore, even in the absence of an explicitly spatial broad-scale map, it may be possible to study spatial processes by understanding which local-scale characteristics of space are important. Received 5 May 1997; accepted 31 July 1997.     

Environmental Stress, Bottom-up Effects, and Community Dynamics: Integrating Molecular-Physiological and Ecological Approaches

Environmental stress and nutrient/productivity models predict the responses of community structure along gradients of physical conditions and bottom-up effects. Although both models have succeeded in helping to understand variation in ecological communities, most tests have been qualitative. Until recently, two roadblocks to more quantitative tests in marine environments have been a lack of (1) inexpensive, field-deployable technology for quantifying (e.g.) temperature, light, salinity, chlorophyll, and productivity, and (2) methods of quantifying the sub-organismal mechanisms linking environmental conditions to their ecological expression. The advent of inexpensive remote-sensing technology, adoption of molecular techniques such as quantification of heat-shock proteins and RNA:DNA ratios, and the formation of interdisciplinary alliances between ecologists and physiologists has begun to overcome these roadblocks. An integrated eco-physiological approach focuses on the determinants of: distributional limits among microhabitat patches and along (local-scale) environmental gradients (e.g., zonation); among-site (mesoscale) differences in community pattern; and geographic (macroscale) differences in ecosystem structure. These approaches promise new insights into the physiological mechanisms underlying variation in processes such as species interactions, physical disturbance, survival and growth. Here, we review two classes of models for community dynamics, and present examples of ecological studies of these models in consumer-prey systems. We illustrate the power of new molecular tools to characterize the sub-organismal responses of some of the same consumers and prey to thermal stress and food concentration. Ecological and physiological evidence tends to be consistent with model predictions, supporting our argument that we are poised to make major advances in the mechanistic understanding of community dynamics along key environmental gradients.     

Spatial structure in ecological communities – a quantitative analysis

The spatial structure of communities has recently gained much attention in ecology. Spatial structure comprises an important element in communities, but the literature lacks a thorough investigation about possible among‐organism or among‐ecosystem differences in the degree of spatial structure. Here, I conducted a quantitative review to determine if the degree of spatial structure varied predictably between the major organism types and ecosystems. Spatial structure was quantified as the relative fraction of community variation explained purely by spatial variables (fraction S/E). I integrated data from 322 variation partition analyses both in a generalized linear model (GLM) and using a boosted regression tree method, and showed that a mean of 11.0% of the variation in community composition was explained purely by spatial variables. Across all taxa, a body size–S/E relationship was positive. In GLM, fraction S/E increased highly significantly with study extent, it was highest among terrestrial taxa and higher in ectotherms than in homoiotherms. Spatial structure was also higher in omnivores than in autotrophs. These results suggest that the degree of spatial structure is jointly driven by extrinsic factors such as study extent and ecosystem type, and intrinsic factors such as body size, thermoregulation and interactions between body size and dispersal mode. These results should be important not only for basic research, but also conservation and bioassessment programs would benefit from the information about the magnitude of spatial variation in nature. Synthesis Spatial processes comprise an important element in most ecological communities, but the degree to which spatial structure varies across organisms or ecosystems is poorly known. Here, a quantitative review of 322 variation partition analyses indicated that spatial component varied predictably across ecological communities – it was driven by study extent and ecosystem type as well as by species traits such as body size and thermoregulation. These results give deep insights into the magnitude of spatial variation in nature and should be highly beneficial for conservation and bioassessment programs that are built on the information about how communities vary in space.     

基于无人机影像的草方格生态恢复区植被空间格局演化研究

理解植物群落组成结构的演化对于阐明荒漠化的过程与驱动机制、制定有效的干旱区生态系统恢复措施具有重要价值。研究干旱区植物群落的空间格局的演化过程有助于深入理解荒漠化和生态恢复的过程与机理。目前大量研究关注于植被退化过程中的群落组成结构变化,而对于生态恢复过程中的植物群落空间格局演化的研究尚不多见。干旱区生态系统中植物通常较为稀疏且个体较小,准确提取植物的分布往往需要分辨率极高的遥感数据。近年来,低空无人机遥感技术的快速发展为精细尺度上植被空间格局的研究提供重要技术支持。利用2 cm空间分辨率的低空无人机遥感数据结合地面群落调查,在精细尺度上研究了宁夏沙坡头草方格生态恢复区内植物群落的空间格局变化。研究结果表明,沙坡头地区草方格生态恢复工程实验区域,相对于未实施生态恢复工程的裸露沙丘区域,植物物种多样性和植被盖度显著提高。恢复工程实施4年后,平均植被盖度增加3倍,物种丰富度增加1倍。在植被恢复过程中,随着植被盖度的增加,植被斑块表现出规模上升、破碎化程度下降、形状复杂化、空间自相关减弱等格局特征变化。这些空间格局特征的变化表明大型植被斑块趋于恢复,整体微环境的改善有利于单独生长的植物个体存活,整体上生态系统退化为裸地的风险降低。利用低空无人机遥感手段,对草方格生态恢复工程的植被恢复过程进行了详细、高分辨率的空间格局调查及分析,结合地面群落调查,从多个方面证明了草方格生态恢复措施的有效性。基于无人机的系统格局连续长期监测有助于深入理解干旱区生态恢复机理,对于科学开展荒漠化生态恢复措施也具有重要价值。     

Ecological drift and local exposures drive enteric bacterial community differences within species of Galápagos iguanas

Diet strongly influences the intestinal microbial communities through species sorting. Alternatively, these communicates may differ because of chance variation in local microbial exposures or species losses among allopatric host populations (i.e. ecological drift). We investigated how these forces shape enteric communities of Galápagos marine and land iguanas. Geographically proximate populations shared more similar communities within a host ecotype, suggesting a role for ecological drift during host colonization of the islands. Additionally, evidence of taxa sharing between proximate heterospecific host populations suggests that contemporary local exposures also influence the gut community assembly. While selective forces such as host-bacterial interactions or dietary differences are dominant drivers of intestinal community differences among hosts, historical and contemporary processes of ecological drift may lead to differences in bacterial composition within a host species. Whether such differences in community structure translate into geographic variation in benefits derived from these intimate microbial communities remains to be explored.     

A comparative hierarchical analysis of bacterioplankton and biofilm metacommunity structure in an interconnected pond system

It is unknown whether bacterioplankton and biofilm communities are structured by the same ecological processes, and whether they influence each other through continuous dispersal (known as mass effects). Using a hierarchical sampling approach we compared the relative importance of ecological processes structuring the dominant fraction (relative abundance ≥0.1%) of bacterioplankton and biofilm communities from three microhabitats (open water, Nuphar and Phragmites sites) at within‐ and among‐pond scale in a set of 14 interconnected shallow ponds. Our results demonstrate that while bacterioplankton and biofilm communities are highly distinct, a similar hierarchy of ecological processes is acting on them. For both community types, most variation in community composition was determined by pond identity and environmental variables, with no effect of space. The highest β‐diversity within each community type was observed among ponds, while microhabitat type (Nuphar, Phragmites, open water) significantly influenced biofilm communities but not bacterioplankton. Mass effects among bacterioplankton and biofilm communities were not detected, as suggested by the absence of within‐site covariation of biofilm and bacterioplankton communities. Both biofilm and plankton communities were thus highly structured by environmental factors (i.e., species sorting), with among‐lake variation being more important than within‐lake variation, whereas dispersal limitation and mass effects were not observed.     

Biomass compensation: Behind the diversity gradients of tidepool fishes

Species richness is unevenly distributed on the Earth, with biodiversity gradients of various spatial scales supposedly being affected by abiotic as well as biotic factors including community traits such as body size spectra and relative abundance patterns. To explore large-scale spatial variation in species diversity and their processes, tidepool fish communities were investigated through an intensive field work conducted on 55 shore sites in south-western Japan. Multiple ecological measures were taken into account to assess changes in local community structures with changes in the number of species. Biomass (total fish wet weight) per unit area showed no systematic change with latitude, while taxa richness and number of individuals tended to increase toward lower latitudes. In addition, median fish body weight scaled positively with latitude, which was more conspicuous in Blenniidae than in Gobiidae. The latitudinal gradient of diversity in tidepool fish assemblages appears to be characterized by partitioning of total biomass that tends to stay constant across latitudes, suggesting the phenomenon of “biomass compensation” whereby body size and abundance/diversity change in opposite directions with latitude. Our study highlights that biomass compensation can be part of processes involved in generating gradients of species richness even without an apparent energy/resource gradient.     

Effects of habitat conditions at hatching time on growth history of offspring European anchovy, <Emphasis Type="Italic">Engraulis encrasicolus</Emphasis>, in the Central Mediterranean Sea

Species distributions are structured by regional and local determinants, which operate at multiple spatial and temporal scales. The purpose of our work was to distinguish the relative roles of local variables, climate, geographical location and post glaciation condition (i.e., delineation between supra- and subaquatic lakes during the post-glacial Ancylus Lake) in explaining variation in macrophyte community composition of all taxa, helophytes and hydrophytes. In addition, we investigated how these four explanatory variable groups affected macrophyte strategy groups based on Grime’s classification. Using partial linear regression and variation partitioning, we found that macrophyte communities are primarily filtered by local determinants together with regional characteristics at the studied spatial scale. We further evidenced that post glaciation condition indirectly influenced on local water quality variables, which in turn directly contributed to the macrophyte communities. We thus suggest that regional determinants interact with local-scale abiotic factors in explaining macrophyte community patterns and examining only regional or local factors is not sufficient for understanding how aquatic macrophyte communities are structured locally and regionally.     

The role of evolutionary processes in producing biodiversity patterns, and the interrelationships between taxonomic, functional and phylogenetic biodiversity

Patterns of biodiversity are ultimately the product of speciation and extinction. Speciation serves as the biodiversity pump while extinction serves as the agent that culls global to local levels of biodiversity. Linking these central processes to global and local patterns of biodiversity is a key challenge in both ecology and evolution. This challenge necessarily requires a simultaneous consideration of the species, phylogenetic, and functional diversity across space and the tree of life. In this review, I outline a research framework for biodiversity science that considers the evolutionary and ecological processes that generate and cull levels of biodiversity and that influence the inter-relationships between species, phylogenetic, and functional diversity. I argue that a biodiversity synthesis must begin with a consideration of the inherently ecological process of speciation and end with how global biodiversity is filtered into local-scale plant communities. The review ends with a brief outlook on future challenges for those studying biodiversity, including outstanding hypotheses that need testing and key data limitations.     

The relative of species pools in determining plant species richness: an alternative explanation of species coexistence?

Explanations of the pattern of species have traditionally relied on small-scale, local processes occurring in ecological time. Differences in species richness have associated with different mechanisms avoiding competition, such as spatiotemporal heterogeneity (weaker competitors may find a more favourable place or time) or environmental stress (competition is assumed to be less intensive under difficult conditions). More recently, large-scale process have been taken into account, raising such questions as: which plant species may potentially grow in a certain community? Are evolutionary processes and species dispersal responsible for the differences between communities? The species-pool theory attempts to answer these general questions, and information about species pools is needed for the design of experiments where the number of species in a community is manipulated.     

Ecological uniqueness of macroinvertebrate communities in high-latitude streams is a consequence of deterministic environmental filtering processes

Variation in biological communities is a consequence of stochastic and deterministic factors. Examining the relative importance of these factors helps to understand variation in the whole biodiversity in a region. We examined the roles of stochastic and deterministic factors in structuring macroinvertebrate communities in high-latitude streams across two seasons. We predicted that if communities are the result of deterministic environmental filtering processes, the communities should show strong association with environmental variables, as taxa would be selected according to stream environmental conditions. However, if communities are driven by stochastic factors, they should show strong association with spatial variables, as the distribution of taxa in communities would be driven by spatially related dispersal factors. We studied these predictions by calculating the degree of uniqueness of the streams in terms of their taxonomic and functional community compositions and by modelling the resulting index values using spatial and environmental variables. Our results supported the first prediction where the communities are more influenced by the environmental filtering processes, although indications of the effect of spatial processes in structuring the communities were present especially in autumn. High-latitude stream communities also seem to be sensitive to environmental changes, as even small changes in environment were enough to affect the ecological uniqueness of the streams. These findings highlight the vulnerability of northern streams in the face of the climate change. To maintain biodiversity in high-latitude catchments, it would be important to protect varying habitat conditions, which are the main forces affecting the ecological uniqueness of the streams.     

The structure of helminth communities

The existence of both interactive and isolationist communities of helminths, each of which may be produced by any one of several processes, necessitates a pluralistic view of helminth community structure. A scheme involving three hierarchical scales of communities of helminths is proposed. Interactions between species occur in infracommunities, and should be looked for there. Interactions are often clearer in smaller groups of species, either among core species or within guilds. Infracommunities are samples of helminth communities at two larger scales: the helminth community in the host population provides core species, largely specialists, and that in the host community provides the generalists and the satellite species. The richness of each of the two large-scale communities is affected by various ecological, historical and evolutionary factors. The different concepts which have been applied to parasite communities are based on these factors.     

Biogeography and ecology: towards the integration of two disciplines

Although ecology and biogeography had common origins in the natural history of the nineteenth century, they diverged substantially during the early twentieth century as ecology became increasingly hypothesis-driven and experimental. This mechanistic focus narrowed ecology''s purview to local scales of time and space, and mostly excluded large-scale phenomena and historical explanations. In parallel, biogeography became more analytical with the acceptance of plate tectonics and the development of phylogenetic systematics, and began to pay more attention to ecological factors that influence large-scale distributions. This trend towards unification exposed problems with terms such as ‘community’ and ‘niche,’ in part because ecologists began to view ecological communities as open systems within the contexts of history and geography. The papers in this issue represent biogeographic and ecological perspectives and address the general themes of (i) the niche, (ii) comparative ecology and macroecology, (iii) community assembly, and (iv) diversity. The integration of ecology and biogeography clearly is a natural undertaking that is based on evolutionary biology, has developed its own momentum, and which promises novel, synthetic approaches to investigating ecological systems and their variation over the surface of the Earth. We offer suggestions on future research directions at the intersection of biogeography and ecology.     

Measuring biodiversity to explain community assembly: a unified approach

One of the oldest challenges in ecology is to understand the processes that underpin the composition of communities. Historically, an obvious way in which to describe community compositions has been diversity in terms of the number and abundances of species. However, the failure to reject contradictory models has led to communities now being characterized by trait and phylogenetic diversities. Our objective here is to demonstrate how species, trait and phylogenetic diversity can be combined together from large to local spatial scales to reveal the historical, deterministic and stochastic processes that impact the compositions of local communities. Research in this area has recently been advanced by the development of mathematical measures that incorporate trait dissimilarities and phylogenetic relatedness between species. However, measures of trait diversity have been developed independently of phylogenetic measures and conversely most of the phylogenetic diversity measures have been developed independently of trait diversity measures. This has led to semantic confusions particularly when classical ecological and evolutionary approaches are integrated so closely together. Consequently, we propose a unified semantic framework and demonstrate the importance of the links among species, phylogenetic and trait diversity indices. Furthermore, species, trait and phylogenetic diversity indices differ in the ways they can be used across different spatial scales. The connections between large‐scale, regional and local processes allow the consideration of historical factors in addition to local ecological deterministic or stochastic processes. Phylogenetic and trait diversity have been used in large‐scale analyses to determine how historical and/or environmental factors affect both the formation of species assemblages and patterns in species richness across latitude or elevation gradients. Both phylogenetic and trait diversity have been used at different spatial scales to identify the relative impacts of ecological deterministic processes such as environmental filtering and limiting similarity from alternative processes such as random speciation and extinction, random dispersal and ecological drift. Measures of phylogenetic diversity combine phenotypic and genetic diversity and have the potential to reveal both the ecological and historical factors that impact local communities. Consequently, we demonstrate that, when used in a comparative way, species, trait and phylogenetic structures have the potential to reveal essential details that might act simultaneously in the assembly of species communities. We highlight potential directions for future research. These might include how variation in trait and phylogenetic diversity alters with spatial distances, the role of trait and phylogenetic diversity in global‐scale gradients, the connections between traits and phylogeny, the importance of trait rarity and independent evolutionary history in community assembly, the loss of trait and phylogenetic diversity due to human impacts, and the mathematical developments of biodiversity indices including within‐species variations.     

Population-scale drivers of individual arrival times in migratory birds

1. In migratory species, early arrival on the breeding grounds can often enhance breeding success. Timing of spring migration is therefore a key process that is likely to be influenced both by factors specific to individuals, such as the quality of winter and breeding locations and the distance between them, and by annual variation in weather conditions before and during migration. 2. The Icelandic black-tailed godwit Limosa limosa islandica population is currently increasing and, throughout Iceland, is expanding into poorer quality breeding areas. Using a unique data set of arrival times in Iceland in different years for individuals of known breeding and wintering locations, we show that individuals breeding in lower quality, recently occupied and colder areas arrive later than those from traditionally occupied areas. The population is also expanding into new wintering areas, and males from traditionally occupied winter sites also arrive earlier than those occupying novel sites. 3. Annual variation in timing of migration of individuals is influenced by large-scale weather systems (the North Atlantic Oscillation), but between-individual variation is a stronger predictor of arrival time than the NAO. Distance between winter and breeding sites does not influence arrival times. 4. Annual variation in timing of migration is therefore influenced by climatic factors, but the pattern of individual arrival is primarily related to breeding and winter habitat quality. These habitat effects on arrival patterns are likely to operate through variation in individual condition and local-scale density-dependent processes. Timing of migration thus appears to be a key component of the intricate relationship between wintering and breeding grounds in this migratory system.