Understanding ‘it depends’ in ecology: a guide to hypothesising,visualising and interpreting statistical interactions

Ecologists routinely use statistical models to detect and explain interactions among ecological drivers, with a goal to evaluate whether an effect of interest changes in sign or magnitude in different contexts. Two fundamental properties of interactions are often overlooked during the process of hypothesising, visualising and interpreting interactions between drivers: the measurement scale – whether a response is analysed on an additive or multiplicative scale, such as a ratio or logarithmic scale; and the symmetry – whether dependencies are considered in both directions. Overlooking these properties can lead to one or more of three inferential errors: misinterpretation of (i) the detection and magnitude (Type-D error), and (ii) the sign of effect modification (Type-S error); and (iii) misidentification of the underlying processes (Type-A error). We illustrate each of these errors with a broad range of ecological questions applied to empirical and simulated data sets. We demonstrate how meta-analysis, a widely used approach that seeks explicitly to characterise context dependence, is especially prone to all three errors. Based on these insights, we propose guidelines to improve hypothesis generation, testing, visualisation and interpretation of interactions in ecology.     

Relationships between floristic gradients in a primary succession

Abstract. Correlations between five floristic gradients at small spatial extents (10 - 20 m) and one successional gradient over a larger spatial extent (1.5 km) are analysed. Floristic data (62 taxa) were sampled on five terminal moraines of known age deposited after the ‘Little Ice Age’ (1750–1930). The floristic gradients on the moraine ridges were sampled by two or three transects on each moraine, and the successional gradient by 114 plots on all moraines. The sequential orders of species were determined by constrained and unconstrained Correspondence Analysis. The similarities between species order on the successional gradient and the exposure gradients at the same age were tested against the null hypothesis of no correlation. The null hypothesis was rejected using both Monte Carlo permutation tests and Spearman's rank correlations, except on the oldest moraine. Here a closed canopy has developed, which eliminates the environmental variability associated with ridge morphology. The similarity between the successional gradient and the moraine-ridge gradients is attributed to variation in environmental severity, mainly caused by glacier wind and related factors such as temperature and moisture. Similarity was highest on the moraines in the middle of the glacier foreland, which have many successional stages present and have a relatively exposed relief. The distribution of growth/life-forms along gradients of small spatial extent are comparable to the successional gradient, but lichens, herbs and graminoids differ in their behaviour. The resemblance between species gradients at a small spatial extent and species gradients on a larger spatial extent is interpreted as an ecological self-similar pattern, where young and old substrate are linked to exposed-xeric and protected-mesic habitats, respectively.     

Complementary strengths of spatially-explicit and multi-species distribution models

Species distribution models (SDMs) project the outcome of community assembly processes – dispersal, the abiotic environment and biotic interactions – onto geographic space. Recent advances in SDMs account for these processes by simultaneously modeling the species that comprise a community in a multivariate statistical framework or by incorporating residual spatial autocorrelation in SDMs. However, the effects of combining both multivariate and spatially-explicit model structures on the ecological inferences and the predictive abilities of a model are largely unknown. We used data on eastern hemlock Tsuga canadensis and five additional co-occurring overstory tree species in 35 569 forest stands across Michigan, USA to evaluate how the choice of model structure, including spatial and non-spatial forms of univariate and multivariate models, affects ecological inference about the processes that shape community composition as well as model predictive ability. Incorporating residual spatial autocorrelation via spatial random effects did not improve out-of-sample prediction for the six tree species, although in-sample model fit was higher in the spatial models. Spatial models attributed less variation in occurrence probability to environmental covariates than the non-spatial models for all six tree species, and estimated higher (more positive) residual co-occurrence values for most species pairs. The non-spatial multivariate model was better suited for evaluating habitat suitability and hypotheses about the processes that shape community composition. Environmental correlations and residual correlations among species pairs were positively related, perhaps indicating that residual correlations were due to shared responses to unmeasured environmental covariates. This work highlights the importance of choosing a non-spatial model formulation to address research questions about the species–environment relationship or residual co-occurrence patterns, and a spatial model formulation when within-sample prediction accuracy is the main goal.     

Matching data sets from two different spatial samples

Abstract. Methods for coupling two data sets (species composition and environmental variables for example) are well known and often used in ecology. All these methods require that variables of the two data sets have been recorded at the same sample stations. But if the two data sets arise from different sample schemes, sample locations can be different. In this case, scientists usually transform one data set to conform with the other one that is chosen as a reference. This inevitably leads to some loss of information. We propose a new ordination method, named spatial‐RLQ analysis, for coupling two data sets with different spatial sample techniques. Spatial‐RLQ analysis is an extension of co‐inertia analysis and is based on neighbourhood graph theory and classical RLQ analysis. This analysis finds linear combinations of variables of the two data sets which maximize the spatial cross‐covariance. This provides a co‐ordination of the two data sets according to their spatial relationships. A vegetation study concerning the forest of Chizé (western France) is presented to illustrate the method.     

Geographical and ecological patterns of range size in North American trees

We analysed the degree to which spatial, ecological and phylogenetic factors influenced geographic gradients in range size for all trees native to North America. We compared observed range sizes, measured in either one (latitudinal or longitudinal extent) or two dimensions (range area), with three null models that respected constraints imposed by the geometry of the continent; we tested whether climatic gradients could account for increasing range size with latitude – a Rapoport effect – in North American trees; and whether variation in range size was associated with phylogeny or ecological characteristics of species. We documented an excess of species with small ranges and a lack of widely distributed species compared with null expectations both with and without environmental constraints. We found evidence for a Rapoport effect in North American trees, at different taxonomic levels and for different groups defined by ecological characteristics. This pattern is not an outcome constrained by continental shape and does not depend on the metric for range size, but it is consistent with the climatic variability hypothesis proposed to explain the Rapoport pattern. Finally we showed that trees east of the Rocky Mountains have larger ranges on average than trees to the west or in Mexico and that the proportion of deciduous compared to evergreen species increases with range size. Variation in range size among North American trees is not spatially random, and has a phylogenetic and ecological signal. Consistent with the climatic variability hypothesis, range size of North American trees increases with increasing variability in annual temperature range at higher latitudes.     

Using phylogeographic analyses of gene trees to test species status and processes

A gene tree is an evolutionary reconstruction of the genealogical history of the genetic variation found in a sample of homologous genes or DNA regions that have experienced little or no recombination. Gene trees have the potential of straddling the interface between intra- and interspecific evolution. It is precisely at this interface that the process of speciation occurs, and gene trees can therefore be used as a powerful tool to probe this interface. One application is to infer species status. The cohesion species is defined as an evolutionary lineage or set of lineages with genetic exchangeability and/or ecological interchangeability. This species concept can be phrased in terms of null hypotheses that can be tested rigorously and objectively by using gene trees. First, an overlay of geography upon the gene tree is used to test the null hypothesis that the sample is from a single evolutionary lineage. This phase of testing can indicate that the sampled organisms are indeed from a single lineage and therefore a single cohesion species. In other cases, this null hypothesis is not rejected due to a lack of power or inadequate sampling. Alternatively, this null hypothesis can be rejected because two or more lineages are in the sample. The test can identify lineages even when hybridization and lineage sorting occur. Only when this null hypothesis is rejected is there the potential for more than one cohesion species. Although all cohesion species are evolutionary lineages, not all evolutionary lineages are cohesion species. Therefore, if the first null hypothesis is rejected, a second null hypothesis is tested that all lineages are genetically exchangeable and/or ecologically interchangeable. This second test is accomplished by direct contrasts of previously identified lineages or by overlaying reproductive and/or ecological data upon the gene tree and testing for significant transitions that are concordant with the previously identified lineages. Only when this second null hypothesis is rejected is a lineage elevated to the status of cohesion species. By using gene trees in this manner, species can be identified with objective, a priori criteria with an inference procedure that automatically yields much insight into the process of speciation. When one or more of the null hypotheses cannot be rejected, this procedure also provides specific guidance for future work that will be needed to judge species status.     

Prediction of bird community composition based on point-occurrence data and inferential algorithms: a valuable tool in biodiversity assessments

Local biological communities are made up of species, each of which has its own particular relationship with the environment. To the extent that these autecological niches limit species’ distributions, and by extension community composition, models of species’ ecological niches can predict species composition at particular sites, or at least provide a null hypothesis of potential species composition in the absence of effects of species interactions. We developed distributional predictions (ecological niche models) for 89 species occurring in dry tropical forest in the Balsas Basin of south‐western Mexico using an interpolation technique, and predicted the species likely to occur at 8 sites across the region. Onsite field inventory data were then used to test the community predictions, all of which were statistically significant. These results suggest that inventory efforts can be made more efficient by development beforehand of hypotheses that focus onsite collecting and inventory.     

A simultaneous test of synchrony causal factors in muskrat and mink fur returns at different scales across Canada

BACKGROUND: Synchrony among populations has been attributed to three major hypotheses: dispersal, the Moran effect, and trophic-level interactions. Unfortunately, simultaneous testing of these hypotheses demands complete and detailed data, which are scarce for ecological systems. METHODOLOGY/PRINCIPAL FINDINGS: Hudson's Bay Company data on mink and muskrat fur returns in Canada represent an excellent opportunity to test these hypotheses because of the detailed spatial and temporal data from this predator-prey system. Using structural equation modelling, support for each hypothesis was evaluated at two spatial scales: across Canada and dividing the country into three regions longitudinally. Our results showed that at both scales mink synchrony is a major factor determining muskrat synchrony, supporting the hypothesis of trophic-level interactions, but the influence of winter precipitation synchrony is also important in eastern Canada. Moreover, mink synchrony is influenced principally by winter precipitation synchrony at the level of all Canada (Moran effect), but by distance at regional level, which might suggest some influence of dispersal at this level. DISCUSSION/SIGNIFICANCE: Our result is one of the few reports of synchrony mediated by trophic-level interactions, highlighting the importance of evaluation of scale effects in population synchrony studies.     

The species pool concept applied to forests in a fragmented landscape: dispersal limitation versus habitat limitation

Abstract. The species pool concept has been used as a theoretical framework for understanding local community richness. A significant problem in putting the concept into practice is the lack of methods for determining the size of the species pool. We tested the hypothesis that species composition of recent forests is primarily determined by the species composition of neighbouring older forests against the null‐hypothesis that species are a random sample of the species occurring in the study area. Forest plant species composition of recently established fragments was significantly correlated with species composition in neighbouring older forests (i.e. the local species pool). When older forest within a neighbourhood of 1000m radius is considered, seed dispersal sources can be found for 91% of the flora in the recent forests. For an individual fragment, dispersal is a much more important determinant of species presence than the environment, with an average of 46% of the total pool excluded from local pools by dispersal limitation and only 8% excluded by environmental limitations. The species richness of recent forests is on average 23% of the local species pool. Several hypotheses are proposed for this low percentage, such as asymmetric competition due to the early successional state or the limited colonization period.     

A practical approach to the study of spatial structure in simple cases of heterogeneous vegetation

Abstract. Spatial heterogeneity is a characteristic of most natural ecosystems which is difficult to handle analytically, particularly in the absence of knowledge about the exogenous factors responsible for this heterogeneity. While classical methods for analysis of spatial point patterns usually require the hypothesis of homogeneity, we present a practical approach for partitioning heterogeneous vegetation plots into homogeneous subplots in simple cases of heterogeneity without drastically reducing the data. It is based on the detection of endogenous variations of the pattern using local density and second‐order local neighbour density functions that allow delineation of irregularly shaped subplots that could be considered as internally homogeneous. Spatial statistics, such as Ripley's K‐function adapted to analyse plots of irregular shape, can then be computed for each of the homogeneous subplots. Two applications to forest ecological field data demonstrate that the method, addressed to ecologists, can avoid misinterpretations of the spatial structure of heterogeneous vegetation stands.     

Diversity patterns of plant functional types in relation to fire regime and previous land use in Mediterranean woodlands

Abstract. We explore patterns of diversity of plant functional types (PFTs) in Mediterranean communities subjected to landscape‐scale fire disturbances in a mosaic of uncultivated and old fields stands. We use regenerative and growth form attributes to establish two sets of PFTs of perennial species living in shrublands and pine forests of NE Spain. We test the following hypotheses: 1. Fire frequency decreases regenerative PFTs diversity by negatively selecting attributes with low regenerative efficiency. 2. Fire history has more influence on regenerative than on growth form PFTs. 3. The lowest diversity of growth form PFTs will be in old fields without recent fires. We surveyed stands of different combinations of fire and land use histories. Fire history included areas without fires in the last twenty years (unburned), sites burned in 1982 (1‐burned), and sites burned in 1982 and 1994 (2‐burned). Land use histories considered terraced old fields, and uncultivated stands on stony soils. We analysed patterns of PFT abundance and diversity at the stand level, and across the landscape (among stands absolute deviations from sample medians of the relative cover of PFTs). At the stand level, fire had more influence on the diversity patterns of regenerative PFTs than on growth form PFTs. Fire decreased the diversity of regenerative PFTs, due to the elimination of the species without effective mechanisms to post‐fire regeneration. This effect was not observed across the landscape, but seeders showed more variation in stands with longer history without fire. Land use contributed to explain the diversity patterns of growth form PFT (i.e. the number of growth form PFTs was lower in uncultivated, unburned sites), but it did not influence regenerative PFTs diversity. Patterns of PFTs diversity reflect the response to ecological processes operating at the landscape level. Overall, regenerative and growth form PFTs appear to be more sensitive to the fire history than to the past land use.     

Stand and neighbourhood parameters as determinants of plant species richness in a managed forest

Abstract. Patterns of species richness in a managed forest were related to ecological factors at two spatial scales. Local variables within a plot (5 m × 5 m) included 13 soil classes and nine stand classes. Neighbourhood variables were defined within a 25 m radius from the focal plot and were measured using a Neighbourhood Diversity Index (NDI - total diversity of different stand classes) and nine Neighbourhood Scores (NS - relative dominance of a given stand class). 224 species were registered in the survey of which 138 were true forest species. Local parameters (stand and soil class) were weak predictors of total and forest species richness. High total and forest species richness were significantly related to high values of NDI (i.e. heterogeneous neighbourhoods). Regression trees showed that total species richness was best predicted by the amount of roadside habitat in the neighbourhood and NDI. Forest species richness was positively related to NDI and negatively related to the amount of coniferous stands in the neighbourhood.     

Analysing ecological networks of species interactions

Network approaches to ecological questions have been increasingly used, particularly in recent decades. The abstraction of ecological systems – such as communities – through networks of interactions between their components indeed provides a way to summarize this information with single objects. The methodological framework derived from graph theory also provides numerous approaches and measures to analyze these objects and can offer new perspectives on established ecological theories as well as tools to address new challenges. However, prior to using these methods to test ecological hypotheses, it is necessary that we understand, adapt, and use them in ways that both allow us to deliver their full potential and account for their limitations. Here, we attempt to increase the accessibility of network approaches by providing a review of the tools that have been developed so far, with – what we believe to be – their appropriate uses and potential limitations. This is not an exhaustive review of all methods and metrics, but rather, an overview of tools that are robust, informative, and ecologically sound. After providing a brief presentation of species interaction networks and how to build them in order to summarize ecological information of different types, we then classify methods and metrics by the types of ecological questions that they can be used to answer from global to local scales, including methods for hypothesis testing and future perspectives. Specifically, we show how the organization of species interactions in a community yields different network structures (e.g., more or less dense, modular or nested), how different measures can be used to describe and quantify these emerging structures, and how to compare communities based on these differences in structures. Within networks, we illustrate metrics that can be used to describe and compare the functional and dynamic roles of species based on their position in the network and the organization of their interactions as well as associated new methods to test the significance of these results. Lastly, we describe potential fruitful avenues for new methodological developments to address novel ecological questions.     

Natural disturbances and the physiognomy of pine savannas: A phenomenological model

Abstract. Question: The decline of the Pinus palustris ecosystems has resulted from anthropogenic influences, such as conversion to pine plantation forestry, agriculture and land development, all of which are closely related to increases in human populations. Other effects, however, have arisen from alterations in disturbance regimes that maintain the structure and function of these ecosystems. How have alterations of the disturbance regime altered the physiognomy of ‘old‐growth’ stands, and what are the implications for ecosystem conservation and restoration? Methods: In contrast to models that emphasize close interactions among the vertically complex strata, we develop a conceptual phenomenological model for the physiognomic structure of Pinus palustris stands. We relate two natural disturbances (tropical storms and fire) that affect different stages of the life cycle to different aspects of the physiognomic structure. We then compare overstorey stand structure and ground cover composition of two old‐growth longleaf stands near the extremes of different composite disturbance regimes: the Wade Tract (frequent hurricanes and fire) and the Boyd Tract (infrequent hurricanes and long‐term fire exclusion). Results: We predict that tropical storms and fires have different effects on stand physiognomy. Tropical storms are periodic, and sometimes intense, whereas fires are more frequent and less intense. Hurricanes directly influence the overstorey via wind‐caused damage and mortality, and indirectly influence the herb layer by altering the spatial distribution of shading and litter accumulation. Fire exerts direct effects on juvenile stages and indirect effects on the herb layer via fine fuel consumption and selective mortality of potential competitors of P. palustris juveniles. These differences in effects of disturbances can result in widely different physiognomies for P. palustris stands. Finally, some global climate change scenarios have suggested that changes may occur in tropical storm and fire regimes, altering frequency and severity. Such changes may greatly affect pine stands, and ultimately entire pine savanna ecosystems. Conclusions: Our phenomenological model of disturbance regimes in Pinus palustris old‐growth produces very different physiognomies for different disturbances regimes that reflect natural process and human management actions. This model can be used to derive restoration strategies for pine savannas that are linked to reinstitution of important ecological processes rather than specific physiognomic states.     

Masting,mixtures and modes: are two models better than one?

A key hypothesis in population ecology is that synchronous and intermittent seed production, known as mast seeding, is driven by the alternating allocation of carbohydrates and mineral nutrients between growth and reproduction in different years, i.e. ‘resource switching’. Such behaviour may ultimately generate bimodal distributions of long‐term flower and seed production, and evidence of these patterns has been taken to support the resource switching hypothesis. Here, we show how a widely‐used statistical test of bimodality applied by many studies in different ecological contexts may fail to reject the null hypothesis that focal probability distributions are unimodal. Using data from five tussock grass species in South Island, New Zealand, we find clear evidence of bimodality only when flowering patterns are analyzed with probabilistic mixture models. Mixture models provide a theory oriented framework for testing hypotheses of mast seeding patterns, enabling the different responses underlying medium‐ and high‐ versus non‐ and low‐flowering years to be modelled more realistically by associating these with distinct probability distributions. Coupling theoretical expectations with more rigorous statistical approaches will empower ecologists to reject null hypotheses more often.     

Carbon balances of bioenergy systems using biomass from forests managed with long rotations: bridging the gap between stand and landscape assessments

Studies report different findings concerning the climate benefits of bioenergy, in part due to varying scope and use of different approaches to define spatial and temporal system boundaries. We quantify carbon balances for bioenergy systems that use biomass from forests managed with long rotations, employing different approaches and boundary conditions. Two approaches to represent landscapes and quantify their carbon balances – expanding vs. constant spatial boundaries – are compared. We show that for a conceptual forest landscape, constructed by combining a series of time‐shifted forest stands, the two approaches sometimes yield different results. We argue that the approach that uses constant spatial boundaries is preferable because it captures all carbon flows in the landscape throughout the accounting period. The approach that uses expanding system boundaries fails to accurately describe the carbon fluxes in the landscape due to incomplete coverage of carbon flows and influence of the stand‐level dynamics, which in turn arise from the way temporal system boundaries are defined on the stand level. Modelling of profit‐driven forest management using location‐specific forest data shows that the implications for carbon balance of management changes across the landscape (which are partly neglected when expanding system boundaries are used) depend on many factors such as forest structure and forest owners’ expectations of market development for bioenergy and other wood products. Assessments should not consider forest‐based bioenergy in isolation but should ideally consider all forest products and how forest management planning as a whole is affected by bioenergy incentives – and how this in turn affects carbon balances in forest landscapes and forest product pools. Due to uncertainties, we modelled several alternative scenarios for forest products markets. We recommend that future work consider alternative scenarios for other critical factors, such as policy options and energy technology pathways.     

Global patterns of body size evolution are driven by precipitation in legless amphibians

Body size shapes ecological interactions across and within species, ultimately influencing the evolution of large‐scale biodiversity patterns. Therefore, macroecological studies of body size provide a link between spatial variation in selection regimes and the evolution of animal assemblages through space. Multiple hypotheses have been formulated to explain the evolution of spatial gradients of animal body size, predominantly driven by thermal (Bergmann's rule), humidity (‘water conservation hypothesis’) and resource constraints (‘resource rule’, ‘seasonality rule’) on physiological homeostasis. However, while integrative tests of all four hypotheses combined are needed, the focus of such empirical efforts needs to move beyond the traditional endotherm–ectotherm dichotomy, to instead interrogate the role that variation in lifestyles within major lineages (e.g. classes) play in creating neglected scenarios of selection via analyses of largely overlooked environment–body size interactions. Here, we test all four rules above using a global database spanning 99% of modern species of an entire Order of legless, predominantly underground‐dwelling amphibians (Gymnophiona, or caecilians). We found a consistent effect of increasing precipitation (and resource abundance) on body size reductions (supporting the water conservation hypothesis), while Bergmann's, the seasonality and resource rules are rejected. We argue that subterranean lifestyles minimize the effects of aboveground selection agents, making humidity a dominant selection pressure – aridity promotes larger body sizes that reduce risk of evaporative dehydration, while smaller sizes occur in wetter environments where dehydration constraints are relaxed. We discuss the links between these principles with the physiological constraints that may have influenced the tropically‐restricted global radiation of caecilians.     

A test of ecological succession hypotheses using 55‐year time‐series data for 361 boreal forest stands

Aim There has been much work on succession over many decades, but succession fundamentals are still debated because of the reliance on chronosequences and dendrochronological reconstruction, both of which are problematic approaches. Here we use time‐series data to test four hypotheses that lie at the heart of successional theory: (1) the neighbourhood effect hypothesis – tree species abundance is time dependent; (2) the density‐dependence hypothesis – a rare species is more favoured over time; (3) the resource ratio hypothesis – species that can grow at the lowest resource level tend to dominate resource limited sites through succession; and (4) the intermediate disturbance hypothesis – intermediate disturbances increase the abundance of rare species. Location Central boreal forest in Canada (47°50′–50°10′ N; 80°10′–85°50′ W). Methods We used repeated measurements from sequential aerial photography and ground surveys for 361 fire‐origin stands that were measured over a c. 55‐year period. Results Shade‐intolerant Pinus banksiana decreased, tolerant Thuja occidentalis increased, intolerant Populus spp. and Betula papyrifera displayed a U‐shaped trend, and intermediate‐tolerant Picea spp. and tolerant Abies balsamea did not change with time since fire, showing evidence of negative, positive, or neutral neighbourhood effects. Species either persisted for longer or increased more in non‐conspecific stands, and had higher increases in abundance when associated with species of contrasting shade tolerance, supporting the density‐dependence hypothesis and indicating shade‐tolerance complementarity as a mechanism for coexistence. Resource‐poor soils favoured those species capable of tolerating limited resources, whereas rich soils permitted invasion and promoted mixtures, supporting the resource ratio hypothesis. Intermediate disturbances increased the invasibility of rare species. Main conclusions Contrary to previous studies where time since a stand‐replacing disturbance is the sole predictor for succession, our study shows that time alone is either an insufficient predictor or is irrelevant to species dynamics in the boreal forest. Rather, density dependence, site resource and intermediate disturbances are key mechanisms in species dynamics and coexistence over time.     

基于不同零模型的点格局分析

在种群空间格局研究中,定量分析格局及其形成过程已成为生态学家的主要目标。在量化分析的众多方法中,点格局分析是最常用的方法,而在选择零模型时,完全空间随机模型以外的复杂零模型很少使用,实际上,这些零模型可能有助于认识格局的内在特征。为此,我们在研究实例中,选择完全空间随机模型(complete spatial randomness)、泊松聚块模型(Poisson cluster process)和嵌套双聚块模型(nested double-cluster process)对典型草原处于不同恢复演替阶段的羊草(Leymus chinensis)种群空间格局进行了分析。结果发现:完全空间随机模型仅能检测种群在不同尺度下的格局类型;而通过泊松聚块模型和嵌套双聚块模型检验表明,在恢复演替的初期阶段,羊草种群在小尺度范围内偏离泊松聚块模型,而在整个取样范围内完全符合嵌套双聚块模型;随着恢复演替时间的推移,在恢复演替的后期,在整个取样尺度上,羊草种群与泊松聚块模型相吻合。这是很有意义的生态学现象。这一实例表明在应用点格局分析种群空间格局时,仅通过完全空间随机模型的检验来分析格局特征,或许很难论证复杂的生态过程,而选择一些完全空间随机模型以外的较复杂的零模型,可能发现一些有价值的生态学现象,对揭示格局掩盖下的内在机制有所裨益。