Linking species-area and species-energy relationships in Drosophila microcosms

Resource availability is an important constraint on community structure. Some authors have suggested it conceptually links two of the most basic patterns in ecology, the species–area relationship and the latitudinal gradient in species richness. I present the first experimental test of this conjecture, by manipulating both the area and resource concentration of artificial larval drosophilid fly habitats and then allowing colonization from a natural species pool. Both the abundance and species richness of these habitats depended upon the total quantity of resources available, regardless of whether those resources were contained within smaller high-quality habitats or larger poor-quality habitats. While the intercepts of species–area relationships varied with resource concentration, they all collapsed onto the same species–energy curve. These results support the view that energetic constraints are of fundamental importance in structuring ecological communities, and that such constraints may even help explain ecological patterns such as the species–area relationship that do not explicitly address resource availability.     

Diversity-stability relationship varies with latitude in zooplankton

Analyses of temporal patterns of diversity across a wide range of taxa have found that more diverse communities often show smaller compositional changes over time. This generality indicates that high diversity is associated with greater temporal stability in species composition. We examined patterns of diversity and community stability in zooplankton time series data from 36 lakes sampled over a combined 483 years. The species–time relationship was flatter in more species-rich lakes in the temperate zone. However, high-latitude lakes had both low richness and low turnover. These patterns were consistent for turnover both within and among years. Daily, annual and long-term richness were all higher in large lakes while turnover was unaffected by the surface area. Richness on all time scales, as well as turnover within and among years, all declined at high latitude. Species–area relations and latitudinal gradients in richness therefore reflect different temporal components of diversity. Our results suggest that diversity shows strong associations with compositional stability that vary qualitatively across biogeographical provinces. Community stability increases with diversity among lakes in the temperate zone; however, the two are negatively correlated across latitudinal gradients. These patterns indicate that either the direct effects of diversity on stability or their covariance with environmental fluctuations vary with latitude.     

The quest for a null model for macroecological patterns: geometry of species distributions at multiple spatial scales

There have been several attempts to build a unified framework for macroecological patterns. However, these have mostly been based either on questionable assumptions or have had to be parameterized to obtain realistic predictions. Here, we propose a new model explicitly considering patterns of aggregated species distributions on multiple spatial scales, the property which lies behind all spatial macroecological patterns, using the idea we term 'generalized fractals'. Species' spatial distributions were modelled by a random hierarchical process in which the original 'habitat' patches were randomly replaced by sets of smaller patches nested within them, and the statistical properties of modelled species assemblages were compared with macroecological patterns in observed bird data. Without parameterization based on observed patterns, this simple model predicts realistic patterns of species abundance, distribution and diversity, including fractal-like spatial distributions, the frequency distribution of species occupancies/abundances and the species–area relationship. Although observed macroecological patterns may differ in some quantitative properties, our concept of random hierarchical aggregation can be considered as an appropriate null model of fundamental macroecological patterns which can potentially be modified to accommodate ecologically important variables.     

Field theory for biogeography: a spatially explicit model for predicting patterns of biodiversity

Predicting the variation of biodiversity across the surface of the Earth is a fundamental issue in ecology, and in this article we focus on one of the most widely studied spatial biodiversity patterns: the species–area relationship (SAR). The SAR is a central tool in conservation, being used to predict species loss following global climate change, and is striking in its universality throughout different geographical regions and across the tree of life. In this article we draw upon the methods of quantum field theory and the foundation of neutral community ecology to derive the first spatially explicit neutral prediction for the SAR. We find that the SAR has three phases, with a power law increase at intermediate scales, consistent with decades of documented empirical patterns. Our model also provides a building block for incorporating non-neutral biological variation, with the potential to bridge the gap between neutral and niche-based approaches to community assembly.
Ecology Letters (2010) 13: 87–95     

Random sampling, abundance–extinction dynamics and niche-filtering immigration constraints explain the generation of species richness gradients

Aim  The paradigm that species' patterns of distribution, abundance and coexistence are the result of adaptations of the species to their niches has recently been challenged by evidence that similar patterns may be generated by simple random processes. We argue here that a better understanding of macroecological patterns requires an integration of both ecological and neutral stochastic approaches. We demonstrate the utility of such an integrative approach by testing the sampling hypothesis in a species–energy relationship of forest bird species.
Location  A Mediterranean biome in Catalonia, Spain.
Methods  To test the sampling hypothesis we designed a metacommunity model that reproduces the stochastic sampling from a regional pool to predict local species richness variation. Four conceptually different sampling procedures were evaluated.
Results  We showed that stochastic sampling processes predicted a substantial part (over 40%) of the observed variation in species richness, but left considerable variation unexplained. This remaining variation in species richness may be better understood as the result of alternative ecological processes. First, the sampling model explained more variation in species richness when the probability that a species colonises a new locality was assumed to increase with its niche width, suggesting that ecological differences between species matter when it comes to explaining macroecological patterns. Second, extinction risk was significantly lower for species inhabiting high-energy regions, suggesting that abundance–extinction processes play a significant role in shaping species richness patterns.
Main conclusions  We conclude that species–energy relationships may not simply be understood as a result of either ecological or random sampling processes, but more likely as a combination of both.     

The island biogeography of exotic bird species

Aim   A recent upsurge of interest in the island biogeography of exotic species has followed from the argument that they may provide valuable information on the natural processes structuring island biotas. Here, we use data on the occurrence of exotic bird species across oceanic islands worldwide to demonstrate an alternative and previously untested hypothesis that these distributional patterns are a simple consequence of where humans have released such species, and hence of the number of species released.
Location   Islands around the world.
Methods   Statistical analysis of published information on the numbers of exotic bird species introduced to, and established on, islands around the world.
Results   Established exotic birds showed very similar species–area relationships to native species, but different species–isolation relationships. However, in both cases the relationship for established exotics simply mimicked that for the number of exotic bird species introduced. Exotic bird introductions scaled positively with human population size and island isolation, and islands that had seen more native species extinctions had had more exotic species released.
Main conclusion   The island biogeography of exotic birds is primarily a consequence of human, rather than natural, processes.     

Scale dependency of diversity components estimated from primary biodiversity data and distribution maps

Different sources of information about biodiversity may lead to unrealistic or biased estimation of its components, with different patterns according to the scale of analysis. In this study, we analyse patterns of species richness at the local (average alpha) and regional (gamma) scales, and the relationship between them (Whittaker's beta), in central Mexico, using as a source of data for the species' distributions: (1) museum specimen occurrence data for birds, and (2) distribution maps based on ecological niche models developed and refined by experts. We performed analyses at five spatial resolutions (1/32°−1/2°). Scale changes (grain and extent) affected significantly the estimates of average alpha, gamma, and beta. Use of raw occurrence data vs. distribution maps yielded contrasting results, with raw data underestimating alpha and overestimating beta, as functions of area. As regards species–area relationships, our results suggest a natural decomposition of factors into an area-invariant component (related to alpha), and an area dependent factor (related to beta). Most of our results are maintained in a null model that randomizes occurrences without changing observed range-size distributions. From this result we argue that average alpha and Whittaker's beta capture little information about the spatial covariation of species distribution patterns.     

Statistical mechanics unifies different ecological patterns

Recently there has been growing interest in the use of maximum relative entropy (MaxREnt) as a tool for statistical inference in ecology. In contrast, here we propose MaxREnt as a tool for applying statistical mechanics to ecology. We use MaxREnt to explain and predict species abundance patterns in ecological communities in terms of the most probable behaviour under given environmental constraints, in the same way that statistical mechanics explains and predicts the behaviour of thermodynamic systems. We show that MaxREnt unifies a number of different ecological patterns: (i) at relatively local scales a unimodal biodiversity-productivity relationship is predicted in good agreement with published data on grassland communities, (ii) the predicted relative frequency of rare vs. abundant species is very similar to the empirical lognormal distribution, (iii) both neutral and non-neutral species abundance patterns are explained, (iv) on larger scales a monotonic biodiversity-productivity relationship is predicted in agreement with the species-energy law, (v) energetic equivalence and power law self-thinning behaviour are predicted in resource-rich communities. We identify mathematical similarities between these ecological patterns and the behaviour of thermodynamic systems, and conclude that the explanation of ecological patterns is not unique to ecology but rather reflects the generic statistical behaviour of complex systems with many degrees of freedom under very general types of environmental constraints.     

Human impacts on the species-area relationship in reef fish assemblages

The relationship between species richness and area is one of the oldest, most recognized patterns in ecology. Here we provide empirical evidence for strong impacts of fisheries exploitation on the slope of the species–area relationship (SAR). Using comparative field surveys of fish on protected and exploited reefs in three oceans and the Mediterranean Sea, we show that exploitation consistently depresses the slope of the SAR for both power-law and exponential models. The magnitude of change appears to be proportional to fishing intensity. Results are independent of taxonomic resolution and robust across coral and rocky reefs, sampling protocols and statistical methods. Changes in species richness, relative abundance and patch occupancy all appear to contribute to this pattern. We conclude that exploitation pressure impacts the fundamental scaling of biodiversity as well as the species richness and spatial distribution patterns of reef fish. We propose that species–area curves can be sensitive indicators of community-level changes in biodiversity, and may be useful in quantifying the human imprint on reef biodiversity, and potentially elsewhere.     

Abundance and distribution within a guild of benthic stream fishes: local processes and regional patterns

1. Spatial patterns at regional and local scales were examined for evidence that species interactions can influence distribution and abundance within a guild of benthic fishes in upland streams of Oklahoma, U.S.A. Three groups of community patterns were examined: the species–area relationship, species–habitat associations, and interspecific associations.
2. The species–area relationship for riffle habitats was compared to a null species–area model based on random placement. The observed species–area curve was steeper resulting in less species per unit area in small streams than predicted by the null model.
3. Small, species-poor streams had summed fish densities at least as high as larger, species-rich streams, suggesting density compensation.
4. Several significant patterns of negative covariation were found among species at the regional scale, before and after statistically accounting for effects due to measured habitat variables.
5. For two of these negatively covarying taxa ( Cottus carolinae and Etheostoma spectabile ), the influence of each species on the distribution of the other was evaluated experimentally in field enclosures varying in depth and current velocity. The sculpin C . carolinae caused a shift in habitat use by the darter E . spectabile , but no reciprocal shift was found.
6. These results indicate an agreement between local and regional patterns of distribution for C . carolinae and E . spectabile and suggest that biotic interactions can influence regional patterns of distribution for species within this guild.     

Inferring species abundance distribution across spatial scales

A long-standing problem in ecology is to understand how the species–abundance distribution (SAD) varies with sampling scale. The problem was first characterized by Preston as the veil line problem. Although theoretical and empirical studies have now shown the nonexistence of the veil line, this problem has generated much interest in scaling biodiversity patterns. However, research on scaling SAD has so far exclusively focused on the relationship between the SAD in a smaller sampling area and a known SAD assumed for a larger area. An unsolved challenge is how one may predict species–abundance distribution in a large area from that of a smaller area. Although upscaling biodiversity patterns (e.g. species–area curve) is a major focus of macroecological research, upscaling of SAD across scale is, with few exceptions, ignored in the literature. Methods that directly predict SAD in a larger area from that of a smaller area have just started being developed. Here we propose a Bayesian method that directly answers this question. Examples using empirical data from tropical forests of Malaysia and Panama are employed to demonstrate the use of the method and to examine its performance with increasing sampling area. The results indicate that only 10-15% of the total census area is needed to adequately predict species abundance distribution of a region. In addition to species abundance distributions, the method also predicts well the regional species richness.     

Abundance and distribution within a guild of benthic stream fishes: local processes and regional patterns

1. Spatial patterns at regional and local scales were examined for evidence that species interactions can influence distribution and abundance within a guild of benthic fishes in upland streams of Oklahoma, U.S.A. Three groups of community patterns were examined: the species–area relationship, species–habitat associations, and interspecific associations.
2. The species–area relationship for riffle habitats was compared to a null species–area model based on random placement. The observed species–area curve was steeper resulting in less species per unit area in small streams than predicted by the null model.
3. Small, species-poor streams had summed fish densities at least as high as larger, species-rich streams, suggesting density compensation.
4. Several significant patterns of negative covariation were found among species at the regional scale, before and after statistically accounting for effects due to measured habitat variables.
5. For two of these negatively covarying taxa ( Cottus carolinae and Etheostoma spectabile ), the influence of each species on the distribution of the other was evaluated experimentally in field enclosures varying in depth and current velocity. The sculpin C . carolinae caused a shift in habitat use by the darter E . spectabile , but no reciprocal shift was found.
6. These results indicate an agreement between local and regional patterns of distribution for C . carolinae and E . spectabile and suggest that biotic interactions can influence regional patterns of distribution for species within this guild.     

Modeled global invasive potential of Asian gypsy moths, Lymantria dispar

Asian populations of gypsy moths, Lymantria dispar (L.) (Lepidoptera: Lymantriidae), remain poorly characterized – indeed, they are not presently accorded any formal taxonomic status within the broader species. Their ecology is similarly largely uncharacterized in the literature, except by assumption that it will resemble that of European populations. We developed ecological niche models specific to Asian populations of the species, which can in turn be used to identify a potential geographic distributional area for the species. We demonstrated statistically significant predictivity of distributional patterns within the East Asian range of these populations; projecting the Asian ecological niche model to Europe, correspondence with European distributions was generally good, although some differences may exist; projecting the ecological niche model globally, we characterized a likely potential invasive distribution of this set of populations across the temperate zone of both Northern and Southern Hemisphere.     

A framework for assessing the biodiversity and fishery aspects of marine reserves

1.  Resource management agencies are often charged with managing natural resources for economic and social goals, while also protecting and conserving biodiversity and ecosystem function. However, this may not always be possible. Ecosystem-based management is frequently suggested as a way to achieve multiple objectives in resource management and requires that trade-offs among conflicting objectives be identified and an effective means to utilize these trade-offs developed.
2.  We examine the relationship between area and species richness in a diverse assemblage of fishes along the US West Coast and then use parameters from this relationship as input for a model that considers trade-offs between fisheries yield and the number of species protected by different management strategies.
3.  The species–area relationship ( S  =  cA ^z ) for fishes along the US Pacific coast is well described by the relationship S  =   16·18 A ^0·226.
4.  There are nearly linear trade-offs between diversity and yield when fishing effort is low. However, the trade-offs become nonlinear as fishing effort increases and imposing MPAs increases both the conservation and fisheries value of the system when the system is overfished.
5.   Synthesis and applications . Solving conflicts between fisheries and conservation requires attention as to how conservation benefits accrue as fishing effort is reduced. However, scientists often lack quantitative information about the trade-offs inherent in human activities such as fisheries. The approach we develop here can begin to help frame the questions to be posed and evaluate the likely consequences of different management options.     

The power of time: spatiotemporal scaling of species diversity

The species–area relationship (SAR) provides the foundation for much of theoretical ecology and conservation practice. However, by ignoring time the SAR offers an incomplete model for biodiversity dynamics. We used long‐term data from permanent plots in Kansas grasslands, USA, to show that the increase in the number of species found with increasing periods of observation takes the same power‐law form as the SAR. A statistical model including time, area, and their interaction explains 98% of variation in mean species number and demonstrates that while the effect of time depends on area, and vice versa, time has strong effects on species number even at relatively broad spatial scales. Our results suggest equivalence of underlying processes in space and time and raise questions about the diversity estimates currently used by basic researchers and conservation practitioners.     

A null model of exotic plant diversity tested with exotic and native species-area relationships

At large spatial scales, exotic and native plant diversity exhibit a strong positive relationship. This may occur because exotic and native species respond similarly to processes that influence diversity over large geographical areas. To test this hypothesis, we compared exotic and native species–area relationships within six North American ecoregions. We predicted and found that within ecoregions the ratio of exotic to native species richness remains constant with increasing area. Furthermore, we predicted that areas with more native species than predicted by the species–area relationship would have proportionally more exotics as well. We did find that these exotic and native deviations were highly correlated, but areas that were good (or bad) for native plants were even better (or worse) for exotics. Similar processes appear to influence exotic and native plant diversity but the degree of this influence may differ with site quality.     

The terminology and use of species–area relationships: a response to Dengler (2009)

Dengler ( Journal of Biogeography , 2009, 36 , 728–744) addresses issues regarding species–area relationships (SARs), but fails to settles those issues. He states that only certain types of sampling schemes should be used to construct SARs, but is not consistent in the criteria that he uses to include some sampling schemes but not others. He argues that a sampling scheme of contiguous plots will be more accurate in extrapolating beyond the sampled area, but logic tells us that a dispersed sampling scheme is likely to be more accurate. Finally, he concludes that the 'true' SAR is a power function, but this conclusion is inconsistent with his results and with the results of others. Rather than defining a narrow framework for SARs, we need to recognize that the relationship between area and species richness is scale- and system-dependent. Different sampling schemes serve different purposes, and a variety of functional relationships are likely to hold. Further theoretical and empirical work is needed to resolve these issues fully.     

The value of georeferenced collection records for predicting patterns of mosquito species richness and endemism in the Neotropics

Abstract.  1. Determining large-scale distribution patterns for mosquitoes could advance knowledge of global mosquito biogeography and inform decisions about where mosquito inventory needs are greatest.
2. Over 43 000 georeferenced records are presented of identified and vouchered mosquitoes from collections undertaken between 1899 and 1982, from 1853 locations in 42 countries throughout the Neotropics. Of 492 species in the data set, 23% were only recorded from one location, and Anopheles albimanus Wiedemann is the most common species.
3. A linear log–log species–area relationship was found for mosquito species number and country area. Chile had the lowest relative density of species and Trinidad-Tobago the highest, followed by Panama and French Guiana.
4. The potential distribution of species was predicted using an Ecological Niche Modelling (ENM) approach. Anopheles species had the largest predicted species ranges, whereas species of Deinocerites and Wyeomyia had the smallest.
5. Species richness was estimated for 1° grids and by summing predicted presence of species from ENM. These methods both showed areas of high species richness in French Guiana, Panama, Trinidad-Tobago, and Colombia. Potential hotspots in endemicity included unsampled areas in Panama, French Guiana, Colombia, Belize, Venezuela, and Brazil.
6. Argentina, The Bahamas, Bermuda, Bolivia, Cuba, and Peru were the most under-represented countries in the database compared with known country species occurrence data. Analysis of species accumulation curves suggested patchiness in the distribution of data points, which may affect estimates of species richness.
7. The data set is a first step towards the development of a global-scale repository of georeferenced mosquito collection records.     

Species-area relationships from a spatially explicit neutral model in an infinite landscape

We use recently developed technical methods to study species–area relationships from a spatially explicit extension of Hubbell's neutral model on an infinite landscape. Our model includes variable dispersal distances and exhibits qualitatively different behaviour from the cases of nearest-neighbour dispersal and finite periodic landscapes that have previously been studied. We show that different dispersal distances and even different dispersal kernels produce identical species–area curves up to rescaling of the two axes. This scaling property provides a straightforward method for fitting the model to empirical data. The species–area curves display all three phases observed empirically and enable the exponent describing the power law relationship for species–area curves to be identified as the gradient at the central phase. This exponent can take all values between 0 and 1 and is given by a simple function of the speciation rate, independent of all other model variables.     

Global patterns of diversity among the specialist birds of riverine landscapes

1. Despite the growing view that biodiversity provides a unifying theme in river ecology, global perspectives on richness in riverine landscapes are limited. As a result, there is little theory or quantitative data on features that might have influenced global patterns in riverine richness, nor are there clear indications of which riverine landscapes are important to conservation at the global scale. As conspicuous elements of the vertebrate fauna of riverine landscapes, we mapped the global distributions of all of the world's specialist riverine birds and assessed their richness in relation to latitude, altitude, primary productivity and geomorphological complexity (surface configuration). 2. Specialist riverine birds, typical of high‐energy riverine landscapes and dependent wholly or partly on production from river ecosystems, occur in 16 families. They are represented by an estimated 60 species divided equally between the passerines and non‐passerines. Major radiation has occurred among different families on different continents, indicating that birds have evolved several times into the niches provided by riverine landscapes. 3. Continental richness varies from four species in Europe to 28 in Asia, with richness on the latter continent disproportionately larger than would be expected from a random distribution with respect to land area. Richness is greatest in mountainous regions at latitudes of 20–40°N in the riverine landscapes of the Himalayan mountains, where 13 species overlap in range. 4. Family, genus and species richness in specialist riverine birds all increase significantly with productivity and surface configuration (i.e. relief). However, family richness was the best single predictor of the numbers of species or genera. In keeping with the effect of surface configuration, river‐bird richness peaks globally at 1300–1400 m altitude, and most species occur typically on small, fast rivers where they feed predominantly on invertebrates. Increased lengths of such streams in areas of high relief and rainfall might have been responsible for species–area effects. 5. We propose the hypothesis that the diversity in channel forms and habitats in riverine landscapes, in addition to high temperature and primary productivity, have been prerequisites to the development of global patterns in the richness of specialist riverine organisms. We advocate tests of this hypothesis in other taxonomic groups. We draw attention, however, to the challenges of categorically defining riverine organisms in such tests because (i) rivers grade into many other habitat types across several different ecotones and (ii) `terrestrialisation' processes in riverine landscapes means that they offer habitat for organisms whose evolutionary origins are not exclusively riverine.