The scale and cause of spatial heterogeneity in strength of temporal density dependence

The importance of density dependence in natural communities continues to spark much debate because it is fundamental to population regulation. We used temporal manipulations of density to explore potentially stabilizing density dependence in early survivorship among six local populations of a tropical damselfish (Dascyllus flavicaudus). Specifically, we tested the premise that spatial heterogeneity in the strength of temporal density dependence would reflect variation in density of predators, the agent of mortality. Our field manipulations revealed that mortality among successive cohorts of young fishes was density dependent at each reef, but that its strength varied by approximately 1.5 orders of magnitude. This spatial heterogeneity was well predicted by variation among the six reefs in the density of predatory fishes that consume juvenile damselfishes. Because density dependence arose from competition for enemy-free space within a shelter coral, the mortality consequence of the competition depended on the neighborhood density of predators. Thus, the scale of heterogeneity in the density dependence largely reflected attributes of the environment that shaped the local abundance of predators. These results have important implications for how ecologists explore regulatory processes in nature. Failure to account for spatial variation could frequently yield misleading conclusions regarding density dependence as a stabilizing process, obscure underlying mechanisms influencing its strength, and provide no insight into the spatial scale of the heterogeneity. Further, models of population dynamics will be improved when experimental approaches better estimate the magnitude and causes of variation in strength of stabilizing density dependence.     

Life history traits predict relative abundance in an assemblage of forest caterpillars

Species in a given trophic level occur in vastly unequal abundance, a pattern commonly documented but poorly explained for most taxa. Theoretical predictions of species density such as those arising from the metabolic theory of ecology hold well at large spatial and temporal scales but are not supported in many communities sampled at a relatively small scale. At these scales ecological factors may be more important than the inherent limits to energy use set by allometric scaling of mass. These factors include the amount of resources available, and the ability of individuals to convert these resources successfully into population growth. While previous studies have demonstrated the limits of macroecological theory in explaining local abundance, few studies have tested alternative generalized mechanisms determining abundance at the community scale. Using an assemblage of forest moth species found co-occurring as caterpillars on a single host plant species, we tested whether species abundance on that plant could be explained by mass allometry, intrinsic population growth, diet breadth, or some combination of these traits. We parameterized life history traits of the caterpillars in association with the host plant in both field and laboratory settings, so that the population growth estimate was specific to the plant on which abundance was measured. Using a generalized least-squares regression method incorporating phylogenetic relatedness, we found no relationship between abundance and mass but found that abundance was best explained by both intrinsic population growth rate and diet breadth. Species population growth potential was most affected by survivorship and larval development time on the host plant. Metabolic constraints may determine upper limits to local abundance levels for species, but local community abundance is strongly predicted by the potential for population increase and the resources available to that species in the environment.     

Population dynamics of the barnacle <Emphasis Type="Italic">Chthamalus montagui</Emphasis> at two spatial and temporal scales in northern Spain

This study describes spatial and temporal patterns of variability in population parameters in the barnacle Chthamalus montagui Southward in three localities of northern Spain and evaluates whether density-dependent settlement may regulate population dynamics. The sampling design considered two spatial scales, localities and sites within localities, and two temporal scales, years and six month intervals. Density, amount of free space, mortality, growth rate and magnitude of settlement (both absolute and per unit of free space) were obtained from photographs of permanent quadrats and from direct counts in the field. The number of settlers in scraped and untouched quadrats was used to estimate the importance of the presence of conspecifics in settlement. Significant variation at the two spatial and temporal scales was found for most parameters. Large spatial and temporal variations in adult mortality rate, density, and settlement were observed. Patterns of mortality were not consistent with differences in density among localities. Differences in settlement among localities were maintained through time. We suggest that magnitude of settlement is regulated by persistent features such as topography or local water circulation. We assume that early post-settlement mortality does not differ among localities. In the absence of differential mortality, settlement determines average population density within localities. Within localities, settlement was independent of density and free space. No consistent evidence was found on preferential settlement at the vicinity of conspecifics. The main conclusion is that density-dependent settlement is not relevant for the regulation of the populations of C. montagui in the northern Spain. Regulation might occur by density-dependent processes within the adult fraction of the population and/or the larval phase before settlement.     

Using an individual-based model to quantify scale transition in demographic rate functions: Deaths in a coral reef fish

Scientifically informed population management requires quantitatively accurate demographic rate functions that apply at the spatial scale at which populations are actually managed, but practical constraints confine most field measurements of such functions to small study plots. This paper employs an individual-based population growth model to extrapolate the death rate function in a well-studied coral reef fish, the bridled goby Coryphopterus glaucofraenum, from the scale of coral reef “cells” at which it was measured to the larger scale of an entire coral reef. Density dependence in the whole-reef function actually proves stronger than in the local function because high goby density occasionally arises in local patches with few refuges from predators, producing very high mortality there. This IBM-based approach extends the reach of scale transition theory by examining considerably more realistic models than standard analytical methods can presently handle.     

Climate, habitat, and species interactions at different scales determine the structure of a Neotropical bat community

Climate, habitat, and species interactions are factors that control community properties (e.g., species richness, abundance) across various spatial scales. Usually, researchers study how a few properties are affected by one factor in isolation and at one scale. Hence, there are few multi-scale studies testing how multiple controlling factors simultaneously affect community properties at different scales. We ask whether climate, habitat structure, or insect resources at each of three spatial scales explains most of the variation in six community properties and which theory best explains the distribution of selected community properties across a rainfall gradient. We studied a Neotropical insectivorous bat ensemble in the Isthmus of Panama with acoustic monitoring techniques. Using climatological data, habitat surveys, and insect captures in a hierarchical sampling design we determined how much variation of the community properties was explained by the three factors employing two approaches for variance partitioning. Our results revealed that most of the variation in species richness, total abundance, and feeding activity occurred at the smallest spatial scale and was explained by habitat structure. In contrast, climate at large scales explained most of the variation in individual species' abundances. Although each species had an idiosyncratic response to the gradient, species richness peaked at intermediate levels of precipitation, whereas total abundance was very similar across sites, suggesting density compensation. All community properties responded in a different manner to the factor and scale under consideration.     

Neighborhood and community interactions determine the spatial pattern of tropical tree seedling survival

Factors affecting survival and recruitment of 3531 individually mapped seedlings of Myristicaceae were examined over three years in a highly diverse neotropical rain forest, at spatial scales of 1-9 m and 25 ha. We found convincing evidence of a community compensatory trend (CCT) in seedling survival (i.e., more abundant species had higher seedling mortality at the 25-ha scale), which suggests that density-dependent mortality may contribute to the spatial dynamics of seedling recruitment. Unlike previous studies, we demonstrate that the CCT was not caused by differences in microhabitat preferences or life history strategy among the study species. In local neighborhood analyses, the spatial autocorrelation of seedling survival was important at small spatial scales (1-5 m) but decayed rapidly with increasing distance. Relative seedling height had the greatest effect on seedling survival. Conspecific seedling density had a more negative effect on survival than heterospecific seedling density and was stronger and extended farther in rare species than in common species. Taken together, the CCT and neighborhood analyses suggest that seedling mortality is coupled more strongly to the landscape-scale abundance of conspecific large trees in common species and the local density of conspecific seedlings in rare species. We conclude that negative density dependence could promote species coexistence in this rain forest community but that the scale dependence of interactions differs between rare and common species.     

Allocation within a generic scaling framework

Barnes and Roderick [Barnes, B., Roderick, M.L., 2004. An ecological framework linking scales across space and time based on self-thinning. Theoret. Popul. Biol. 66, 113–128] developed a generic ecological framework for scaling from individuals to ecosystems. Their approach is general and can be applied to predict above-ground, or total (above- and below-ground), dry mass. In practice, the most common situation is to measure above-ground dry mass, and apply an allometric relationship to estimate the below-ground component. In this paper we develop a general theory for incorporating the dynamics of plant partitioning into the generic framework. We consider the inclusion of allometric relationships between components (such as between roots and shoots), as well as process driven relationships, and illustrate the application of each case. Through this approach, local scale measurements and individual-based dynamic relationships pertaining to plant partitioning can be applied to an understanding of partitioning at the patch (or ecosystem) scale. Moreover, we also demonstrate that the empirically based allometric relationships have, in some circumstances, a physical explanation, providing biological meaning to empirically established allometric constants.     

From single fine roots to a black alder forest ecosystem: How system behaviour emerges from single component activities

Based on empirical findings in a natural black alder ecosystem in Northern Germany we developed an individual based model that integrates components of a black alder ecosystem interacting on different levels of organisation. The factors determining seasonal fine root biomass development of forest ecosystems are not yet fully understood.We used an object oriented model approach to investigate this complex matter for black alder trees. Processes like growth, storage, respiration, transport, nutrient mineralisation and uptake as well as interactions among these factors are described on the level of functionally differentiated plant organs (fine roots, coarse roots, stem, branches, leaves) and soil units. The object structure of the model is determined by spatial relations between plant modules as well as between plant modules and their local environment modules.As results of model application we found that (i) on the organ level, spatio-temporal plasticity of (root) growth allocation is related to spatio-temporal variation of resource availability, (ii) on the plant level, balanced root:shoot growth appears in response to variation of available resources light and nutrients, (iii) on the population level, tree stand development (population structure, self-thinning) resulted from coexistence and competition between plant individuals.For the understanding of the root compartment it seems relevant that the model implementation of local scale fine root dynamics is consistent with a self-organised large scale spatial heterogeneity of fine root activity pattern. On the other hand, fine-root dynamics cannot be explained as a result of autonomous dynamics. A reference to above-ground processes is a necessary condition and the overall plant seems to act as an integrator providing boundary conditions for local activity pattern. At the same time fine-root characteristics are of some importance for properties on hierarchically higher levels, e.g. co-existence in a tree population or element cycling in the ecosystem.As a conclusion, modelling of the spatio-temporal dynamics of tree root systems appears as a paradigmatic example of scale and organisation level integrating processes.     

Plant diversity partitioning in Mediterranean croplands: effects of farming intensity, field edge, and landscape context

Farmland biodiversity is affected by factors acting at various spatial scales. However, most studies to date have focused on the field or farm scales that only account for local (alpha) diversity, and these may underestimate the contribution of other diversity components (beta diversity) to total (gamma) farmland diversity. In this work, we aimed to identify the most suitable management options and the scale at which they should be implemented to maximize benefits for diversity. We used a multi-scale additive partitioning approach, with data on plant diversity from 640 plots in 32 cereal crop fields from three agricultural regions of central Spain that differed in landscape configuration. We analyzed the relative contribution to overall plant diversity of different diversity components at various spatial scales and how these diversity components responded to a set of local (application of agri-environment schemes [AES] and position within the field) and landscape (field size and landscape connectivity and composition) factors. Differences in species composition among regions and then among fields within regions contributed most to overall plant diversity. Positive edge effects were found on all diversity components at both the field- and regional scales, whereas application of AES benefited all diversity components only at the field scale. Landscape factors had strong influences on plant diversity, especially length of seminatural boundaries, which increased species richness at both the field and the regional scales. In addition, positive effects of percentage of nonproductive land-uses in the landscape were found on all diversity components at the regional scale. Results showed that components that contributed most to overall plant diversity were not benefited by current AES. We conclude that agri-environmental policies should incorporate and prioritize measures aimed at the maintenance of seminatural boundaries and patches of nonproductive habitats within agricultural landscapes, through landscape planning, cross-compliance, or high nature-value farmland programs. These options will help to conserve overall plant diversity at regional scales, as well as the spillover of plant species from such seminatural elements into crops, especially in Mediterranean areas that still harbor extensive farming and relatively complex landscapes.     

Scale-dependent changes in the importance of larval supply and habitat to abundance of a reef fish

While there is great interest in the degree to which local interactions "scale-up" to predict regional patterns of abundance, few studies in marine systems have simultaneously examined patterns of abundance at both the large scale (tens of kilometers) typical of larval movement and the small scale (meters) typical of post-settlement interactions. We addressed this gap by monitoring larval supply, adult survivorship, and giant kelp (Macrocystis pyrifera, a primary habitat-forming species) abundance for 13 populations of kelp bass (Paralabrax clathratus) spread over approximately 200 km in the Santa Barbara Channel, California, USA. At the small, within-site scale, both recruitment and adult survivorship of kelp bass were density-dependent and positively related to kelp abundance. At the larger, among-site scale, the spatial pattern of adult kelp bass abundance was predicted well by the pattern of kelp bass larval supply, but there was a consistent negative spatial relationship between kelp abundance and kelp bass larval supply despite the positive effects of kelp on kelp bass at the smaller spatial scale. This large-scale negative relationship was likely a product of a channel-wide spatial mismatch between oceanographic conditions that favor kelp survival and those that concentrate and distribute fish larvae. These results generally support the recruit-adult hypothesis: kelp bass populations are limited by recruitment at low recruit densities but by density-dependent competition for food resources and/or predator refuges at high recruit densities. At the same time, spatial variation in kelp abundance produced substantial spatiotemporal heterogeneity in kelp bass demographics, which argues for a multispecies, metacommunity approach to predicting kelp bass dynamics.     

Variation at local scales need not impede tests for broader scale patterns

Ecologists are not always mindful of the constraints imposed by their scale of observation and sometimes prematurely attempt broad generalisations or become mesmerised by local details depending on their predilections. We specifically chose a character that is known for its local and unpredictable variation (morphology of kelp) to test the effect of scale on our ability to determine spatial patterns. We compared the morphology of Ecklonia radiata between monospecific and mixed stands of canopy-forming algae across temperate Australia (>5,100 km coastal distance) within a hierarchy of several spatial scales. While E. radiata specimens were generally larger in monospecific than in mixed stands, we failed to observe differences in morphology between stands at many sites and locations. Despite substantial local variation, differences between stands became increasingly clear at broader scales. The frequency of inconsistent differences between stands was greatest at local scales (sites separated by kms), intermediate at intermediate scales (locations separated by 100s of kms) and least at regional scales (regions separated by 1,000s of kms). These observations support the idea that large scale patterns can emerge from apparent stochasticity at small scales, and that unaccountable variation at local scales need not impede tests for similar patterns at broader scales. Most ecologists work at scales where complexity tends to be greatest (i.e. local) and is likely to be explained by special and unique events. It is encouraging, therefore, to observe that patterns can emerge from complexity at local scales to provide new opportunities to answer some of the more interesting questions about the relative importance of processes across the vast parts of the worlds coast.Communicated by M.S. Johnson, Crawley     

Evaluating the Effects of Anthropogenic Stressors on Source‐Sink Dynamics in Pond‐Breeding Amphibians

Although interwetland dispersal is thought to play an important role in regional persistence of pond‐breeding amphibians, few researchers have modeled amphibian metapopulation or source‐sink dynamics. Results of recent modeling studies suggest anthropogenic stressors, such as pollution, can negatively affect density and population viability of amphibians breeding in isolated wetlands. Presumably population declines also result in reduced dispersal to surrounding (often uncontaminated) habitats, potentially affecting dynamics of nearby populations. We used our data on the effects of mercury (Hg) on the American toad ( Bufo americanus) as a case study in modeling the effects of anthropogenic stressors on landscape‐scale amphibian dynamics. We created a structured metapopulation model to investigate regional dynamics of American toads and to evaluate the degree to which detrimental effects of Hg contamination on individual populations can disrupt interpopulation dynamics. Dispersal from typical American toad populations supported nearby populations that would otherwise have been extirpated over long time scales. Through support of such sink populations, dispersal between wetland‐associated subpopulations substantially increased overall productivity of wetland networks, but this effect declined with increasing interwetland distance and decreasing wetland size. Contamination with Hg substantially reduced productivity of wetland‐associated subpopulations and impaired the ability of populations to support nearby sinks within relevant spatial scales. Our results add to the understanding of regional dynamics of pond‐breeding amphibians, the wide‐reaching negative effects of environmental contaminants, and the potential for restoration or remediation of degraded habitats. Evaluación de los Efectos de Estresantes Antropogénicos sobre la Dinámica Fuente‐Vertedero en Anfibios que se Reproducen en Charcas     

The influence of host diversity and composition on epidemiological patterns at multiple spatial scales

Spatial patterns of pathogen prevalence are determined by ecological processes acting across multiple spatial scales. Host-pathogen interactions are influenced by community composition, landscape structure, and environmental factors. Explaining prevalence patterns requires an understanding of how local determinants of infection, such as community composition, are mediated by landscape characteristics and regional-scale environmental drivers. Here we investigate the role of local community interactions and the effects of landscape structure on the dynamics of barley and cereal yellow dwarf viruses (B/CYDV) in the open meadows of the Cascade Mountains of Oregon. B/CYDV is an aphid-transmitted, generalist pathogen of over 100 wild and cultivated grass species. We used variance components analysis and model selection techniques to partition the sources of variation in B/CYDV prevalence and to determine which abiotic and biotic factors influence host-pathogen interactions in a Cascades meadowsystem. B/CYDV prevalence in Cascades meadows varied by host species identity, with a significantly higher proportion of infected Festuca idahoensis individuals than Elymus glaucus or Bromus carinatus. Although there was significant variation in prevalence among host species and among meadows in the same meadow complex, there was no evidence of any significant variation in prevalence among different meadow complexes at a larger spatial scale. Variation in prevalence among meadows was primarily associated with the local community context (host identity, the relative abundance of different host species, and host species richness) and the physical landscape attributes of the meadow. These results highlight the importance of local host community composition, mediated by landscape characteristics such as meadow aspect, as a determinant of the spatial pattern of infection of a multi-host pathogen.     

Linking patch-use behavior, resource density, and growth expectations in fish

Optimality theory rests on the assumptions that short-term foraging decisions are driven by variation in environmental quality, and that these decisions have important implications for long-term fitness. These assumptions, however, are rarely tested in a field setting. We linked behavioral foraging decisions in food patches with measures of environmental quality covering larger spatial (resource density) or temporal (growth parameters) scales. In 10 lakes, we measured the food density at which benthic fish give up foraging in experimental food patches (giving-up density, GUD), quantified the biomass of benthic invertebrates, and calculated the maximum individual size (L(infinity)) of bream (Abramis brama L.), a typical benthivore in these lakes. We found positive relationships between resource density and both GUD and L(infinity), and a positive relationship between L(infinity) and GUD. Prey characterized as vulnerable to predation contributed most to the relationships between resource density and either GUD or L(infinity). A path analysis showed that resource density and L(infinity) directly explained 54% and 28%, respectively, of the variation in GUD, whereas 86% of the variation in L(infinity) was explained by resource density, with mostly indirect contribution from GUD. We conclude that the short-term foraging behavior of benthivores matched our expectations based on optimality theory by being positively linked to variables on environmental quality operating at both a larger spatial scale and a longer temporal scale.     

Reconciling empirical ecology with neutral community models

Neutral community models embody the idea that individuals are ecologically equivalent, having equal fitness over all environmental conditions, and describe how the spatial dynamics and speciation of such individuals can produce a wide range of patterns of distribution, diversity, and abundance. Neutral models have been controversial, provoking a rush of tests and comments. The debate has been spurred by the suggestion that we should test mechanisms. However, the mechanisms and the spatial scales of interest have never clearly been described, and consequently, the tests have often been only peripherally relevant. At least two mechanisms are present in spatially structured neutral models. Dispersal limitation causes clumping of a species, which increases the strength of intraspecific competition and reduces the strength of interspecific competition. This may prolong coexistence and enhance local and regional diversity. Speciation is present in some neutral models and gives a donor-controlled input of new species, many of which remain rare or are short lived, but which directly add to species diversity. Spatial scale is an important consideration in neutral models. Ecological equivalence and equal fitness have implicit spatial scales because dispersal limitation and its emergent effects operate at population levels, and populations and communities are defined at a chosen spatial scale in recent neutral models; equality is measured relative to a metacommunity, and this necessitates defining the spatial scale of that metacommunity. Furthermore, dispersal has its own scales. Thorough empirical tests of neutral models will require both tests of mechanisms and pattern-producing ability, and will involve coupling theoretical models and experiments.     

A general model for analyzing Taylor's spatial scaling laws

Taylor's spatial scaling law concerns the relation between the variance and the mean population counts within areas of a given size. For a range of area sizes, the log of the variance often is an approximately linear function of the mean with a slope between 1 and 2, depending on the range of areas considered. In this paper, we investigate this relationship theoretically for random quadrat samples within a large area. The model makes a distinction between the local point process determining the position of each individual and the population density described by a spatial covariance function. The local point process and the spatial covariance of population density both contribute to the general relationship between the mean and the variance in which the slope may begin at 1, increase to 2, and decrease to 1 again. It is demonstrated by an example that the slope theoretically may exceed 2 by a small amount for very regular patterns that generate spatial covariance functions that increase in certain intervals. We also show how properties of population dynamics in space and time determine this relationship.     

Scale dependence of reproductive failure in fragmented Echinacea populations

I investigated reproduction in a three-year study of Echinacea angustifolia, purple coneflower, growing in a fragmented prairie landscape. I quantified the local abundance of flowering conspecifics at individual-based spatial scales and at a population-based spatial scale. Regression analyses revealed that pollen limitation increased while seed set and fecundity decreased with isolation of individual plants. Isolation, defined as the distance to the k(th) nearest flowering conspecific, was a good predictor of pollen limitation, for all nearest neighbors considered (k = 1-33), but the strength of the relationship, as quantified by R2, peaked at intermediate scales (k = 2-18). The relationship of isolation to seed set and fecundity was similarly strongest at intermediate scales (k = 3-4). The scale dependence of individual density effects on reproduction (density of flowering plants within x meters) resembled that of isolation. Analyses at a population-based scale showed that pollen limitation declined significantly with population size. Seed set and fecundity also declined with population size, but significantly so only in 1998. Whether quantifying local abundance with population- or individual-based measures, reproductive failure due to pollen limitation is a consistent consequence of Echinacea scarcity. However, individual-based measures of local abundance predicted pollen limitation from a wider sample of plants with a simpler model than did population size. Specifically, the largest site, a nature preserve, is composed of plants with intermediate individual isolation and, as predicted, intermediate pollen limitation, but its large population size poorly predicted population mean pollen limitation.     

Regional variation in the hierarchical partitioning of diversity in coral-dwelling fishes

The size of the regional species pool may influence local patterns of diversity. However, it is unclear whether certain spatial scales are less sensitive to regional influences than others. Additive partitioning was used to separate coral-dwelling fish diversity to its alpha and beta components, at multiple scales, in several regions across the Indo-Pacific. We then examined how the relative contribution of these components changes with increased regional diversity. By employing specific random-placement null models, we overcome methodological problems with local-regional regressions. We show that, although alpha and beta diversities within each region are consistently different from random-placement null models, the increase in beta diversities among regions was similar to that predicted once heterogeneity in coral habitat was accounted for. In contrast, alpha diversity within single coral heads was limited and increased less than predicted by the null models. This was correlated with increased intraspecific aggregation in more diverse regions and is consistent with ecological limitations on the number of coexisting species at the local scale. These results suggest that, apart from very small spatial scales, variation in the partitioning of fish diversity along regional species richness gradients is driven overwhelmingly by the corresponding gradients in coral assemblage structure.