Fitness in animals correlates with proximity to discontinuities in body mass distributions

Discontinuous structure in landscapes may cause discontinuous, aggregated species body-mass patterns, reflecting the scales of structure available to animal communities within a landscape. Empirical analyses have shown that the location of species within body mass aggregations, which reflect this scale-specific organization, is non-random with regard to several ecological phenomena, including species extinctions. The propensity of declining species to have body masses proximate to discontinuities suggests that transition zones between scaling regimes ultimately decreases the ecological fitness for some species. We test this proposition using vulnerable and unthreatened fish species in Mediterranean streams with differing levels of human impact. We show that the proximity to discontinuities in body mass aggregations (“distance-to-edge”) of more vs. less fit individuals within vulnerable and unthreatened populations differs. Specifically, regression analysis between the scaled mass index, a proxy of animal fitness, and distance-to-edge reveals negative and positive relationships for vulnerable and unthreatened species, respectively. That is, fitness is higher close to discontinuities in vulnerable populations and toward the center of body mass aggregation groups in unthreatened populations. Our results demonstrate the suitability of the discontinuity framework for scrutinizing non-random patterns of environmental impact in populations. Further exploration of the usefulness of this method across other ecosystems and organism groups is warranted.     

The Use of Discontinuities and Functional Groups to Assess Relative Resilience in Complex Systems

It is evident when the resilience of a system has been exceeded and the system qualitatively changed. However, it is not clear how to measure resilience in a system prior to the demonstration that the capacity for resilient response has been exceeded. We argue that self-organizing human and natural systems are structured by a relatively small set of processes operating across scales in time and space. These structuring processes should generate a discontinuous distribution of structures and frequencies, where discontinuities mark the transition from one scale to another. Resilience is not driven by the identity of elements of a system, but rather by the functions those elements provide, and their distribution within and across scales. A self-organizing system that is resilient should maintain patterns of function within and across scales despite the turnover of specific elements (for example, species, cities). However, the loss of functions, or a decrease in functional representation at certain scales will decrease system resilience. It follows that some distributions of function should be more resilient than others. We propose that the determination of discontinuities, and the quantification of function both within and across scales, produce relative measures of resilience in ecological and other systems. We describe a set of methods to assess the relative resilience of a system based upon the determination of discontinuities and the quantification of the distribution of functions in relation to those discontinuities.     

Functional Richness and Relative Resilience of Bird Communities in Regions with Different Land Use Intensities

Abstract Empirical estimates of the function and resilience of communities under different management regimes can provide valuable information for sustainable natural resource management, but such estimates are scarce to date. We quantified the functional richness and relative resilience of bird communities inhabiting five regions in southeastern Australia that represented different management regimes. First, we show that functional richness and relative resilience were reduced at species-poor sites in all regions. Second, we show that bird communities in agricultural regions had fewer body mass groups and fewer functional groups than expected by chance. This suggests that both the function and the resilience of bird communities in agricultural regions were reduced. The likely mechanisms for the observed loss of function and relative resilience are: (1) the simplification of landscape texture resulting in selective extinction of certain body mass groups; and (2) the selective extinction of certain functional groups that are particularly sensitive to intensive land use. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Spatial Ecological Hierarchies: Coexistence on Heterogeneous Landscapes via Scale Niche Diversification

Spatially heterogeneous environments are generally characterized by nested landscape patterns with resource aggregations on several scales. Empirical studies indicate that such nested landscape patterns impose selection constraints on the perceptive scales of animals, but the underlying selection mechanisms are unclear. We investigated the selection dynamics of perceptive scale within a spatial resource utilization model, where the environment is characterized by its resource distribution and species differ in their perceptive scales and resource preemption capabilities. Using three model landscapes with various resource distributions, we found that the optimal perceptive scale is determined by scale-specific attributes of the landscape pattern and that the number of coexisting species increases with the number of characteristic scales. Based on the results of this model, we argue that resource aggregations on different scales act as distinct resources and that animal species of particular perceptive scales are superior in utilizing resource aggregations of comparable spatial extent. Due to the allometric relationship between body size and perceptive scale, such fitness difference might result in discontinuous body mass distributions.     

Connecting landscape structure and patterns in body size distributions

Understanding the interaction between community structure and landscape structure represents a pressing theoretical challenge of great applied importance considering the increasing structural modification of ecosystems through habitat loss and fragmentation. Dispersal ability and energetic demands coupled to body size determine the landscape structure experienced by an organism, which could essentially be fragmented for small individuals but continuous for large ones. Although discontinuities in species assemblages have been predicted and detected, no explicit association between habitat structure and body size distributions has been demonstrated. In this contribution, we propose that body size structure in local communities should reflect such different perceptions of landscape structure. To this end, we explore this association in a simple metacommunity located in the Atacama Desert, in northern Chile. Using graph theory we found that species of different size and trophic position (carnivores and herbivores) perceive the landscape at contrasting spatial scales. In each community (n = 31) we determined the observed and the expected body size distributions – in a random sample from the metacommunity of 18 727 individuals –, which allowed us to identify the body sizes at which an overrepresentation or underrepresentation of individuals occur. Such aggregations and discontinuities in body sizes were related, for carnivores, to patch location within the landscape, and to the internal banded vegetation pattern within patches for herbivores. Our study shows, for the first time, an empirical connection between the spatial distribution of communities, their local attributes, and the existence and locations of discontinuities and aggregations in body size distributions.     

Ecological Resilience, Biodiversity, and Scale

We describe existing models of the relationship between species diversity and ecological function, and propose a conceptual model that relates species richness, ecological resilience, and scale. We suggest that species interact with scale-dependent sets of ecological structures and processes that determine functional opportunities. We propose that ecological resilience is generated by diverse, but overlapping, function within a scale and by apparently redundant species that operate at different scales, thereby reinforcing function across scales. The distribution of functional diversity within and across scales enables regeneration and renewal to occur following ecological disruption over a wide range of scales. Received 11 April 1997; accepted 9 July 1997.     

Functional Group Change within and across Scales following Invasions and Extinctions in the Everglades Ecosystem

Cross-scale resilience theory predicts that the combination of functional diversity within scales and functional redundancy across scales is an important attribute of ecosystems because it helps these systems resist minor ecological disruptions and regenerate after major disturbances such as hurricanes and fire. Using the vertebrate fauna of south Florida, we quantified how the loss of native species and invasion by nonnatives may alter functional group richness within and across scales. We found that despite large changes in species composition due to potential extinctions and successful invasions by nonnative species, functional group richness will not change significantly within scales, there will not be any significant loss of overall redundancy of ecology function across scales, and overall body mass pattern will not undergo substantial change. However, the types of functions performed will change, and this change may have profound effects on not only the Everglades ecosystem but on the entire landscape of south Florida. Received 14 November 2000; accepted 20 December 2001.     

Habitat structure and body size distributions: cross‐ecosystem comparison for taxa with determinate and indeterminate growth

Habitat structure across multiple spatial and temporal scales has been proposed as a key driver of body size distributions for associated communities. Thus, understanding the relationship between habitat and body size is fundamental to developing predictions regarding the influence of habitat change on animal communities. Much of the work assessing the relationship between habitat structure and body size distributions has focused on terrestrial taxa with determinate growth, and has primarily analysed discontinuities (gaps) in the distribution of species mean sizes (species size relationships or SSRs). The suitability of this approach for taxa with indeterminate growth has yet to be determined. We provide a cross‐ecosystem comparison of bird (determinate growth) and fish (indeterminate growth) body mass distributions using four independent data sets. We evaluate three size distribution indices: SSRs, species size–density relationships (SSDRs) and individual size–density relationships (ISDRs), and two types of analysis: looking for either discontinuities or abundance patterns and multi‐modality in the distributions. To assess the respective suitability of these three indices and two analytical approaches for understanding habitat–size relationships in different ecosystems, we compare their ability to differentiate bird or fish communities found within contrasting habitat conditions. All three indices of body size distribution are useful for examining the relationship between cross‐scale patterns of habitat structure and size for species with determinate growth, such as birds. In contrast, for species with indeterminate growth such as fish, the relationship between habitat structure and body size may be masked when using mean summary metrics, and thus individual‐level data (ISDRs) are more useful. Furthermore, ISDRs, which have traditionally been used to study aquatic systems, present a potentially useful common currency for comparing body size distributions across terrestrial and aquatic ecosystems.     

Functional diversity in nematode communities across terrestrial ecosystems

Functional diversity can be defined as the distribution of trait values within a community. Hence, functional diversity can be an indicator of habitat filtering and a reliable environmental predictor of ecosystem functioning. However, there is a serious lack of studies that test how functional diversity indices change depending on the environmental conditions. The aim of this study is to provide such evidence by analyzing the distribution and variation of continuous body-mass values (i.e. functional diversity) and related shifts in body length and width in a nematode community.We used a large online dataset on nematode traits to analyze: (i) the distribution of body mass using three functional diversity indices, i.e. functional richness, functional divergence and functional evenness; (ii) the shifts in body-size traits (length and width); and (iii) the body-mass distributions of five trophic groups and of the entire nematode community.Managed grasslands exhibited the widest range of body-mass values while body-mass distribution in arable fields covered the greatest area in comparison to the other ecosystem types. The shift in body size revealed environmental filters that could not have been identified by the study of functional diversity indices per se. We found low values of functional evenness to be associated with high values of functional richness. We provide novel empirical evidence that body-mass distribution within a trophic group mirrors the effects of habitat filtering more than the distribution in the community as a whole. Hence, our trait-based approach, more than functional diversity itself, disclosed soil food-web structure and identified community responses.     

Evolution, Ecology, and Multimodal Distributions of Body Size

The sizes of organisms are determined by their interactions with their environment and related ecological and evolutionary processes. Recent studies of body size distributions across communities show evidence for multimodality. The multiple modes were originally explained as a consequence of textural discontinuities in habitat structure. Because communities consist of species that are drawn from lineages, body size patterns within lineages will affect those that are expressed in communities. We used a cellular automation model to argue that multimodality in body sizes within lineages can arise from a few fundamental evolutionary mechanisms alone. We tested the hypothesis using body size data for 138 fish genera and found strong support for the idea that evolution structures body size distributions. The results suggest, first, that we should expect the distribution of body sizes within lineages to be multimodal and second, that a coherent theory of community body size distributions will need to combine both evolutionary and ecological perspectives. Received 28 January 2002; accepted 21 March 2002     

Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain,Southwest China

Characterizing trait variation across different ecological scales in plant communities has been viewed as a way to gain insights into the mechanisms driving species coexistence. However, little is known about how changes in intraspecific and interspecific traits across sites influence species richness and community assembly, especially in understory herbaceous communities. Here we partitioned the variance of four functional traits (maximum height, leaf thickness, leaf area and specific leaf area) across four nested biological scales: individual, species, plot, and elevation to quantify the scale-dependent distributions of understory herbaceous trait variance. We also integrated the comparison of the trait variance ratios to null models to investigate the effects of different ecological processes on community assembly and functional diversity along a 1200-m elevational gradient in Yulong Mountain. We found interspecific trait variation was the main trait variation component for leaf traits, although intraspecific trait variation ranged from 10% to 28% of total variation. In particular, maximum height exhibited high plasticity, and intraspecific variation accounted for 44% of the total variation. Despite the fact that species composition varied across elevation and species richness decreased dramatically along the elevational gradient, there was little variance at our largest (elevation) scale in leaf traits and functional diversity remained constant along the elevational gradient, indicating that traits responded to smaller scale influences. External filtering was only observed at high elevations. However, strong internal filtering was detected along the entire elevational gradient in understory herbaceous communities, possibly due to competition. Our results provide evidence that species coexistence in understory herbaceous communities might be structured by differential niche-assembled processes. This approach--integrating different biological scales of trait variation--may provide a better understanding of the mechanisms involved in the structure of communities.     

Inferring the Relative Resilience of Alternative States

Ecological systems may occur in alternative states that differ in ecological structures, functions and processes. Resilience is the measure of disturbance an ecological system can absorb before changing states. However, how the intrinsic structures and processes of systems that characterize their states affects their resilience remains unclear. We analyzed time series of phytoplankton communities at three sites in a floodplain in central Spain to assess the dominant frequencies or “temporal scales” in community dynamics and compared the patterns between a wet and a dry alternative state. The identified frequencies and cross-scale structures are expected to arise from positive feedbacks that are thought to reinforce processes in alternative states of ecological systems and regulate emergent phenomena such as resilience. Our analyses show a higher species richness and diversity but lower evenness in the dry state. Time series modeling revealed a decrease in the importance of short-term variability in the communities, suggesting that community dynamics slowed down in the dry relative to the wet state. The number of temporal scales at which community dynamics manifested, and the explanatory power of time series models, was lower in the dry state. The higher diversity, reduced number of temporal scales and the lower explanatory power of time series models suggest that species dynamics tended to be more stochastic in the dry state. From a resilience perspective our results highlight a paradox: increasing species richness may not necessarily enhance resilience. The loss of cross-scale structure (i.e. the lower number of temporal scales) in community dynamics across sites suggests that resilience erodes during drought. Phytoplankton communities in the dry state are therefore likely less resilient than in the wet state. Our case study demonstrates the potential of time series modeling to assess attributes that mediate resilience. The approach is useful for assessing resilience of alternative states across ecological and other complex systems.     

An allometric model of home range formation explains the structuring of animal communities exploiting heterogeneous resources

Understanding and predicting the composition and spatial structure of communities is a central challenge in ecology. An important structural property of animal communities is the distribution of individual home ranges. Home range formation is controlled by resource heterogeneity, the physiology and behaviour of individual animals, and their intra‐ and interspecific interactions. However, a quantitative mechanistic understanding of how home range formation influences community composition is still lacking. To explore the link between home range formation and community composition in heterogeneous landscapes we combine allometric relationships for physiological properties with an algorithm that selects optimal home ranges given locomotion costs, resource depletion and competition in a spatially‐explicit individual‐based modelling framework. From a spatial distribution of resources and an input distribution of animal body mass, our model predicts the size and location of individual home ranges as well as the individual size distribution (ISD) in an animal community. For a broad range of body mass input distributions, including empirical body mass distributions of North American and Australian mammals, our model predictions agree with independent data on the body mass scaling of home range size and individual abundance in terrestrial mammals. Model predictions are also robust against variation in habitat productivity and landscape heterogeneity. The combination of allometric relationships for locomotion costs and resource needs with resource competition in an optimal foraging framework enables us to scale from individual properties to the structure of animal communities in heterogeneous landscapes. The proposed spatially‐explicit modelling concept not only allows for detailed investigation of landscape effects on animal communities, but also provides novel insights into the mechanisms by which resource competition in space shapes animal communities.     

Hierarchical and scale‐dependent effects of fishing pressure and environment on the structure and size distribution of parrotfish communities

Parrotfishes are considered to have a major influence on coral reef ecosystems through grazing the benthic biota and are also primary fishery targets in the Indo‐Pacific. Consequently, the impact of human exploitation on parrotfish communities is of prime interest. As anthropogenic and environmental factors interact across spatial scales, sampling programs designed to disentangle these are required by both ecologists and resource managers. We present a multi‐scale examination of patterns in parrotfish assemblage structure, size distribution and diversity across eight oceanic islands of Micronesia. Results indicate that correlates of assemblage structure are scale‐dependent; biogeographic distributions of species and island geomorphology hierarchically influenced community patterns across islands whereas biophysical features and anthropogenic pressure influenced community assemblage structure at the within‐island scale. Species richness and phylogenetic diversity increased with greater broad‐scale habitat diversity associated with different island geomorphologies. However, within‐island patterns of abundance and biomass varied in response to biophysical factors and levels of human influence unique to particular islands. While the effect of fishing activities on community composition and phylogenetic diversity was obscured across island types, fishing pressure was the primary correlate of mean parrotfish length at all spatial scales. Despite widespread fishery‐induced pressure on Pacific coral reefs, the structuring of parrotfish communities at broad spatial scales remains a story largely dependent on habitat. Thus, we propose better incorporation of scale‐dependent habitat effects in future assessments of overexploitation on reef fish assemblages. However, strong community‐level responses within islands necessitate an improved understanding of the phylogenetic and functional consequences of altering community structure.     

放牧干扰梯度下川西亚高山植物群落的组合机理

为了阐明放牧干扰对川西亚高山区域植物群落的组合过程以及群落结构的影响, 研究了放牧干扰梯度下的功能群均匀度和群落谱系结构的变化趋势。结果显示: 在干扰较轻的阔叶林与针叶林样地, 部分样方的功能群均匀度显著高于无效模型, 随着干扰梯度的增强, 功能群均匀度呈线性下降, 样方平均值从0.930降至0.840, 其高于无效模型的次数也逐渐降低, 干扰程度较大的草甸中出现部分样方的功能群均匀度显著低于无效模型。随着干扰程度的增强, 群落的谱系结构指数也呈逐渐上升趋势, 净关联指数平均值由-0.634逐渐增加至2.360, 邻近类群指数由-0.158上升至2.179。草甸与低矮灌丛受干扰较为严重, 其大部分样方的谱系结构指数显著高于随机群落, 表明干扰群落的谱系结构呈聚集分布。功能群均匀度与谱系结构的变化趋势一致, 表明生境筛滤效应与种间竞争作用的平衡决定着群落的组合过程。干扰降低了竞争作用, 促进了少数耐干扰功能群的优势地位, 造成功能群均匀度下降, 同时通过生境筛滤作用, 使群落的谱系结构呈现出聚集分布; 而未干扰的群落中由于竞争作用的效应, 功能群均匀度较高, 谱系结构也更加分散。研究区域植物群落的功能群均匀度与物种丰富度呈负相关, 表明物种间特别是相似物种间的竞争限制了群落的物种多样性。研究结果说明, 生态位分化和物种间的相互竞争在物种共存与群落组合中具有重要作用。     

Wildfire severity reduces richness and alters composition of soil fungal communities in boreal forests of western Canada

Wildfire is the dominant disturbance in boreal forests and fire activity is increasing in these regions. Soil fungal communities are important for plant growth and nutrient cycling postfire but there is little understanding of how fires impact fungal communities across landscapes, fire severity gradients, and stand types in boreal forests. Understanding relationships between fungal community composition, particularly mycorrhizas, and understory plant composition is therefore important in predicting how future fire regimes may affect vegetation. We used an extreme wildfire event in boreal forests of Canada's Northwest Territories to test drivers of fungal communities and assess relationships with plant communities. We sampled soils from 39 plots 1 year after fire and 8 unburned plots. High‐throughput sequencing (MiSeq, ITS) revealed 2,034 fungal operational taxonomic units. We found soil pH and fire severity (proportion soil organic layer combusted), and interactions between these drivers were important for fungal community structure (composition, richness, diversity, functional groups). Where fire severity was low, samples with low pH had higher total fungal, mycorrhizal, and saprotroph richness compared to where severity was high. Increased fire severity caused declines in richness of total fungi, mycorrhizas, and saprotrophs, and declines in diversity of total fungi and mycorrhizas. The importance of stand age (a surrogate for fire return interval) for fungal composition suggests we could detect long‐term successional patterns even after fire. Mycorrhizal and plant community composition, richness, and diversity were weakly but significantly correlated. These weak relationships and the distribution of fungi across plots suggest that the underlying driver of fungal community structure is pH, which is modified by fire severity. This study shows the importance of edaphic factors in determining fungal community structure at large scales, but suggests these patterns are mediated by interactions between fire and forest stand composition.     

Identifying resilience mechanisms to recurrent ecosystem perturbations

The complex nature of ecological systems limits the unambiguous determination of mechanisms that drive resilience to natural disturbance or anthropogenic stress. Using eight-year time series data from boreal lakes with and without bloom formation of an invasive alga (Gonyostomum semen, Raphidophyceae), we studied resilience of phytoplankton communities in relation to recurring bloom impacts. We first characterized phytoplankton community dynamics in both lake types using univariate metrics of community structure (evenness, species richness, biovolume and Simpson diversity). All metrics, except species richness, were substantially altered and showed an inherent stronger variability in bloom lakes relative to reference lakes. We assessed resilience mechanisms using a multivariate time series modelling technique. The models captured clear successional dynamics of the phytoplankton communities in all lakes, whereby different groups of species were substituted sequentially over the ice-free period. The models also identified that G. semen impacts in bloom lakes were only manifested within a single species group, not across species groups, highlighting the rapid renewal of the phytoplankton communities upon bloom collapse. These results provide empirical support of the cross-scale resilience model. Cross-scale resilience could provide an explanation for the paradox that similar species richnesses are seen in bloom-forming lakes and reference lakes despite the clear difference between the community features of the two different sets of lakes investigated.     

Functional richness, functional evenness, and use of niche space in parasite communities

Most efforts aimed at elucidating the factors responsible for the variation in species richness among different parasite communities have focused on host characteristics such as body mass or diet. Independently of host features, however, the way in which parasites use resources within the host may also affect the species richness of the community. The distribution of parasite individuals or biomass in niche space determines whether host resources are used evenly, or whether there are gaps in resource use, with some parts of the total niche being underutilized. Here, the concepts of functional richness and functional evenness are applied to parasite communities for the first time, using simple indices. Measurements of the distribution of species in niche space within communities, such as mapping the distribution of helminths along the length of the host's intestine, is standard practice in parasitology. In such cases, functional richness is simply the proportion of the total number of intestine sections available that are used by at least 1 worm, whereas functional evenness measures the evenness in the distribution of worm numbers or biomass across all niche sections that are occupied. Data on cestode communities of elasmobranchs are used to illustrate the use of these indices, and to show how important they can be in tests of ecological hypotheses. The indices presented here capture essential features of resource use in parasite communities, and can be useful tools for comparative analyses.