Spatial scale, abundance and the species-energy relationship in British birds

1. The spatial scale of analysis may influence the nature, strength and underlying drivers of macroecological patterns, one of the most frequently discussed of which is the relationship between species richness and environmental energy availability. 2. It has been suggested that species-energy relationships are hump-shaped at fine spatial grains and consistently positive at larger regional grains. The exact nature of this scale dependency is, however, the subject of much debate as relatively few studies have investigated species-energy relationships for the same assemblage across a range of spatial grains. Here, we contrast species-energy relationships for the British breeding avifauna at spatial grains of 1 km x 1 km, 2 km x 2 km and 10 km x 10 km plots, while maintaining a constant spatial extent. 3. Analyses were principally conducted using data on observed species richness. While survey work may fail to detect some species, observed species richness and that estimated using nonparametric techniques were strongly positively correlated with each other, and thus exhibit very similar spatial patterns. Moreover, the forms of species-energy relationships using observed and estimated species richness were statistically indistinguishable from each other. 4. Positive decelerating species-energy relationships arise at all three spatial grains. There is little evidence that the explanatory power of these relationships varies with spatial scale. However, ratios of regional (large-scale) to local (small-scale) species richness decrease with increasing energy availability, indicating that local richness responds to energy with a steeper gradient than does regional richness. Local assemblages thus sample a greater proportion of regional richness at higher energy levels, suggesting that spatial turnover of species richness is lower in high-energy regions. Similarly, a crude measure of temporal turnover, the ratio of cumulative species richness over a 4-year period to species richness in a single year, is lower in high-energy regions. These negative relationships between turnover and energy appear to be causal as both total and mean occupancy per species increases with energy. 5. While total density in 1 km x 1 km plots correlates positively with energy availability, such relationships are very weak for mean density per species. This suggests that the observed association between total abundance and species richness may not be mediated by population extinction rates, as predicted by the more individuals hypothesis. 6. The sampling mechanism suggests that species-energy relationships arise as high-energy areas support a greater number of individuals, and that random allocation of these individuals to local areas from a regional assemblage will generate species-energy relationships. While randomized local species-energy relationships are linear and positive, predicted richness is consistently greater than that observed. The mismatch between the observed and randomized species-energy relationships probably arises as a consequence of the aggregated nature of species distributions. The sampling mechanism, together with species spatial aggregation driven by limited habitat availability, may thus explain the species-energy relationship observed at this spatial scale.     

Dissecting the species-energy relationship

Environmental energy availability can explain much of the spatial variation in species richness. Such species-energy relationships encompass a diverse range of forms, and there is intense debate concerning which of these predominate, and the factors promoting this diversity. Despite this there has been relatively little investigation of whether the form, and relative strength, of species-energy relationships varies with (i) the currency of energy availability that is used, and (ii) the ecological characteristics of the constituent species. Such investigations can, however, shed light on the causal mechanisms underlying species-energy relationships. We illustrate this using the British breeding avifauna. The strength of the species-energy relationship is dependent on the energy metric used, with species richness being more closely correlated with temperature than the Normalized Difference Vegetation Index, which is a strong correlate of net primary productivity. We find little evidence, however, for the thermoregulatory load hypothesis that high temperatures enable individuals to invest in growth and reproduction, rather than thermoregulation, increasing population sizes that buffer species from extinction. High levels of productive energy may also elevate population size, which is related to extinction risk by a negative decelerating function. Therefore, the rarest species should exhibit the strongest species-energy relationship. We find evidence to the contrary, together with little support for suggestions that high-energy availability elevates species richness by increasing the numbers of specialists or predators.     

Local-scale species-energy relationships in fish assemblages of some forested streams of the Bolivian Amazon

Productivity (trophic energy) is one of the most important factors promoting variation in species richness. A variety of species-energy relationships have been reported, including monotonically positive, monotonically negative, or unimodal (i.e. hump-shaped). The exact form of the relationship seems to depend, among other things, on the spatial scale involved. However, the mechanisms behind these patterns are still largely unresolved, although many hypotheses have been suggested. Here we report a case of local-scale positive species-energy relationship. Using 14 local fish assemblages in tropical forested headwater streams (Bolivia), and after controlling for major local abiotic factors usually acting on assemblage richness and structure, we show that rising energy availability through leaf litter decomposition rates allows trophically specialized species to maintain viable populations and thereby to increase assemblage species richness. By deriving predictions from three popular mechanistic explanations, i.e. the 'increased population size', the 'consumer pressure', and the 'specialization' hypotheses, our data provide only equivocal support for the latter.     

When should species richness be energy limited,and how would we know?

Energetic constraints are fundamental to ecology and evolution, and empirical relationships between species richness and estimates of available energy (i.e. resources) have led some to suggest that richness is energetically constrained. However, the mechanism linking energy with richness is rarely specified and predictions of secondary patterns consistent with energy‐constrained richness are lacking. Here, we lay out the necessary and sufficient assumptions of a causal relationship linking energy gradients to richness gradients. We then describe an eco‐evolutionary simulation model that combines spatially explicit diversification with trait evolution, resource availability and assemblage‐level carrying capacities. Our model identified patterns in richness and phylogenetic structure expected when a spatial gradient in energy availability determines the number of individuals supported in a given area. A comparison to patterns under alternative scenarios, in which fundamental assumptions behind energetic explanations were violated, revealed patterns that are useful for evaluating the importance of energetic constraints in empirical systems. We use a data set on rockfish (genus Sebastes) from the northeastern Pacific to show how empirical data can be coupled with model predictions to evaluate the role of energetic constraints in generating observed richness gradients.     

Relative contribution of abundant and rare species to species-energy relationships

A major goal of ecology is to understand spatial variation in species richness. The latter is markedly influenced by energy availability and appears to be influenced more by common species than rare ones; species-energy relationships should thus be stronger for common species. Species-energy relationships may arise because high-energy areas support more individuals, and these larger populations may buffer species from extinction. As extinction risk is a negative decelerating function of population size, this more-individuals hypothesis (MIH) predicts that rare species should respond more strongly to energy. We investigate these opposing predictions using British breeding bird data and find that, contrary to the MIH, common species contribute more to species-energy relationships than rare ones.     

Species traits and the form of individual species-energy relationships

Environmental energy availability explains much of the spatial variation in species richness at regional scales. While numerous mechanisms that may drive such total species-energy relationships have been identified, knowledge of their relative contributions is scant. Here, we adopt a novel approach to identify these drivers that exploits the composite nature of species richness, i.e. its summation from individual species distributions. We construct individual species-energy relationships (ISERs) for each species in the British breeding avifauna using both solar (temperature) and productive energy metrics (normalized difference vegetation index) as measures of environmental energy availability. We use the slopes of these relationships and the resultant change in deviance, relative to a null model, as measures of their strength and use them as response variables in multiple regressions that use ecological traits as predictors. The commonest species exhibit the strongest ISERs, which is counter to the prediction derived from the more individuals hypothesis. There is no evidence that predatory species have stronger ISERs, which is incompatible with the suggestion that high levels of energy availability increase the length of the food chain allowing larger numbers of predators to exist. We find some evidence that species with narrow niche breadths have stronger ISERs, thus providing one of the few pieces of supportive evidence that high-energy availability promotes species richness by increasing the occurrence of specialist species that use a narrow range of resources.     

Morphological assembly mechanisms in Neotropical bat assemblages and ensembles within a landscape

Empirical studies on bat assemblages have shown that richness is not appreciably influenced by local processes such as ecological interactions. However, most of these studies have been done in large areas that include high heterogeneity, and they analyse all bat species within such areas, and thus they may be not reflecting local but supra-community conditions. We followed an ecomorphological approach to assess how bat assemblages of species from the families Phyllostomidae and Mormoopidae, and ensembles of frugivorous bats, are assembled in local habitats within a single landscape. We measured the volume of the space defined by wing morphology and quantified the average distance between species within such a volume. Then, we related these measures to local richness. Such relationships were contrasted against relationships with random assemblages to test for statistical differences. At the ensemble level of organization, we found that the frugivorous bat morphological assembly mechanism is different from random patterns, and it corresponds to the volume-increasing model. On the other hand, bat assembly mechanisms may be ubiquitous at the assemblage level, because groups of species coexisting in a local habitat and delimited only by phylogeny include more than one ecological group with no potential to interact. Assembling processes are crucial to an understanding of species diversity in local communities, and ecomorphological analyses are very promising tools that may help in their study.     

Invasions: the perspective of diverse plant communities

Exotic plant invasions represent a threat to natural and managed ecosystems. Understanding of the mechanisms that determine why a given species may invade a given ecosystem, or why some biomes and regions seem more prone to invasions, is limited. One potential reason for this lack of progress may lie in how few studies have addressed invasion mechanisms from the point of view of the invaded community. On the other hand, the renewed debate about the relationship between ecological diversity and ecosystem stability offers the opportunity to revisit existing theory and empirical evidence, and to attempt to investigate which characteristics of plant communities, including their diversity, contribute to their invasibility. Empirical studies have shown both positive and negative relationships between species diversity of resident plant communities and their invasibility by external species. Rather than attempting to build a larger collection of case studies, research now needs to address the mechanisms underlying these relationships. Previous knowledge about the mechanisms favouring invasion needs to be coupled with community theory to form the basis of these new investigations. Modern community theory offers hypotheses and techniques to analyse the invasibility of communities depending on their diversity and other factors, such as species’ life histories and environmental variability. The body of knowledge accumulated in invasion ecology suggests that the role of disturbances, in interaction with fertility, and the importance of interactions with other trophic levels, are specific factors for consideration. In addition, it is essential for future studies to explicitly tease apart the effects of species richness per se from the effects of other components of ecological diversity, such as functional diversity (the number of functional groups) and trophic diversity (the number of interactions among trophic levels).     

Swimming with the giant: coexistence patterns of a new redfin minnow Pseudobarbus skeltoni from a global biodiversity hot spot

Ecological niche theory predicts that coexistence is facilitated by resource partitioning mechanisms that are influenced by abiotic and biotic interactions. Alternative hypotheses suggest that under certain conditions, species may become phenotypically similar and functionally equivalent, which invokes the possibility of other mechanisms, such as habitat filtering processes. To test these hypotheses, we examined the coexistence of the giant redfin Pseudobarbus skeltoni, a newly described freshwater fish, together with its congener Pseudobabus burchelli and an anabantid Sandelia capensis by assessing their scenopoetic and bionomic patterns. We found high habitat and isotope niche overlaps between the two redfins, rendering niche partitioning a less plausible sole mechanism that drives their coexistence. By comparison, environment–trait relationships revealed differences in species–environment relationships, making habitat filtering and functional equivalence less likely alternatives. Based on P. skeltoni's high habitat niche overlap with other species, and its large isotope niche width, we inferred the likelihood of differential resource utilization at trophic level as an alternative mechanism that distinguished it from its congener. In comparison, its congener P. burchelli appeared to have a relatively small trophic niche, suggesting that its trophic niche was more conserved despite being the most abundant species. By contrast, S. capensis was distinguished by occupying a higher trophic position and by having a trophic niche that had a low probability of overlapping onto those of redfins. Therefore, trophic niche partitioning appeared to influence the coexistence between S. capensis and redfins. This study suggests that coexistence of these fishes appears to be promoted by their differences in niche adaptation mechanisms that are probably shaped by historic evolutionary and ecological processes.     

Scale dependence of species-energy relationships: evidence from fishes in thousands of lakes

Variation in the shape of relationships between species richness and different measures of energy may be linked to variation in the spatial scale on which such relationships are measured. We examine scale dependence in the relationship between potential evapotranspiration and the species richness of fishes in 7,885 postglacial lakes. The strength of this relationship is weak across lake communities but strong and positive across groups of lakes or regions. In addition, the strength and slope of this relationship increase significantly as the regional scale of analysis is increased. We interpret the observed patterns in terms of a simple model whereby energy influences the linear character of the species-energy relationship through its influence on spatial turnover in the species composition (beta diversity). Our results suggest that if energy is strongly tied to patterns of site occupancy or abundance, the parameters of species-energy relationships will depend, to a considerable extent, on the scale of measurement. Furthermore, the ability of high-energy regions to accommodate relatively large numbers of rare or infrequent species may underlie any general tendency for the strength or shape of species-energy relationships to change with scale.     

Body size and feeding specificity: macrolepidoptera in Britain

Within a geographic assemblage, large-bodied species of macrolepidopteran moths tend, on average, to be less host-specific than small-bodied. Five possible explanations for this pattern are identified, based respectively on (i) phylogenetic relationships between species, (ii) latitudinal gradients in body size and feeding specificity, (iii) the relationship between range size and body size, (iv) larger body size as a buffer from environmental variation, and (v) the relationship between endophagous host associations and small body size. These mechanisms are tested using data for British macrolepidoptera and also evaluated using evidence from the literature at large. Although some of their assumptions are found to be justified, there is no significant support for any single mechanism. This lack of evidence for previously proposed mechanisms is discussed in the light of a recently proposed alternative explanation which combines theories of host quality and host defence mechanisms.     

Scale‐dependent processes of community assemblyin an African rift lake

1. Ecologists continue to debate whether the assembly of communities of species is more strongly influenced by dispersal limitations or niche‐based factors. Analytical approaches that account for both mechanisms can help to resolve controls of community assembly. 2. We compared littoral snail assemblages in Lake Tanganyika at three different spatial scales (5–25 m, 0.5–10 km and 0.5–27 km) to test whether spatial distance or environmental differences are better predictors of community similarity. 3. At the finest scale (5–25 m), snail assemblages shifted strongly with depth but not across similar lateral distances, indicating a stronger response to environmental gradients than dispersal opportunities. 4. At the two larger scales (0.5–27 km), both environmental similarity and shoreline distance between sites predicted assemblage similarity across sites. Additionally, canonical correspondence analysis revealed that snail abundances were significantly correlated with algal carbon‐to‐nitrogen ratio and wave energy. 5. Our results indicate that the factors governing assemblage structure are scale dependent; niche‐based mechanisms act across all spatial scales, whereas community similarity declines with distance only at larger spatial separations.     

Climate or diet? The importance of biotic interactions in determining species range size

Aim

Species geographical range sizes play a crucial role in determining species vulnerability to extinction. Although several mechanisms affect range sizes, the number of biotic interactions and species climatic tolerance are often thought to play discernible roles, defining two dimensions of the Hutchinsonian niche. Yet, the relative importance of the trophic and the climatic niche for determining species range sizes is largely unknown.

Location

Central and northern Europe.

Time period

Present.

Major taxa studied

Gall-inducing sawflies and their parasitoids.

Methods

We use data documenting the spatial distributions and biotic interactions of 96 herbivore species, and their 125 parasitoids, across Europe and analyse the relationship between species range size and the climatic and trophic dimensions of the niche. We then compare the observed relationships with null expectations based on species occupancy to understand whether the relationships observed are an inevitable consequence of species range size or if they contain information about the importance of each dimension of the niche on species range size.

Results

We find that both niche dimensions are positively correlated with species range size, with larger ranges being associated with wider climatic tolerances and larger numbers of interactions. However, diet breadth appears to more strongly limit species range size. Species with larger ranges have more interactions locally and they are also able to interact with a larger diversity of species across sites (i.e. higher β-diversity), resulting in a larger number of interactions at continental scales.

Main conclusions

We show for the first time how different aspects of species diet niches are related to their range size. Our study offers new insight into the importance of biotic interactions in determining species spatial distributions, which is critical for improving understanding and predictions of species vulnerability to extinction under the current rates of global environmental change.     

Cryptic diversity in a fig wasp community—morphologically differentiated species are sympatric but cryptic species are parapatric

A key debate in ecology centres on the relative importance of niche and neutral processes in determining patterns of community assembly with particular focus on whether ecologically similar species with similar functional traits are able to coexist. Meanwhile, molecular studies are increasingly revealing morphologically indistinguishable cryptic species with presumably similar ecological roles. Determining the geographic distribution of such cryptic species provides opportunities to contrast predictions of niche vs. neutral models. Discovery of sympatric cryptic species increases alpha diversity and supports neutral models, while documentation of allopatric/parapatric cryptic species increases beta diversity and supports niche models. We tested these predictions using morphological and molecular data, coupled with environmental niche modelling analyses, of a fig wasp community along its 2700‐km latitudinal range. Molecular methods increased previous species diversity estimates from eight to eleven species, revealing morphologically cryptic species in each of the four wasp genera studied. Congeneric species pairs that were differentiated by a key morphological functional trait (ovipositor length) coexisted sympatrically over large areas. In contrast, morphologically similar species, with similar ovipositor lengths, typically showed parapatric ranges with very little overlap. Despite parapatric ranges, environmental niche models of cryptic congeneric pairs indicate large regions of potential sympatry, suggesting that competitive processes are important in determining the distributions of ecologically similar species. Niche processes appear to structure this insect community, and cryptic diversity may typically contribute mostly to beta rather than alpha diversity.     

Biological invasions and the neutral theory

Aim  The invasion of natural communities by alien species represents a serious threat, but creates opportunities to learn about community functions. Neutral theory proposes that the niche concept may not be needed to explain the assemblage and diversity of natural communities, challenging the classical view of community ecology and generating a lasting debate. Biological invasions, when considered as natural experiments, can be used to contrast some of the predictions of neutral and classic niche theories.
Location  Global.
Methods  We use data from biological invasions as natural experiments to contrast some of the fundamental predictions of neutral theory.
Results  Some emerging patterns did not differ from neutral model expectations (e.g. the relationship between native and exotic species richness, invasibility of resource-rich habitats, and the relationship between propagule release and invasion success). Nevertheless, other patterns (e.g. experimental evidence of the relationship between diversity and susceptibility to invasion, the invasion of communities with a low resource availability, invasiveness related to species traits) contrasted with the predictions that can be inferred from neutral theory.
Main conclusions  Neutral theory correctly highlights the need to include randomness in models of community structure. Biological invasion patterns show that neutral forces are important in structuring natural communities, but the patterns differ from those inferred from a complete neutral model. For biodiversity-conservation purposes, the implications of accepting or not accepting neutral theory as a process-based theory are very important.     

Assessing species saturation: conceptual and methodological challenges

Is there a maximum number of species that can coexist? Intuitively, we assume an upper limit to the number of species in a given assemblage, or that a lineage can produce, but defining and testing this limit has proven problematic. Herein, we first outline seven general challenges of studies on species saturation, most of which are independent of the actual method used to assess saturation. Among these are the challenge of defining saturation conceptually and operationally, the importance of setting an appropriate referential system, and the need to discriminate among patterns, processes and mechanisms. Second, we list and discuss the methodological approaches that have been used to study species saturation. These approaches vary in time and spatial scales, and in the variables and assumptions needed to assess saturation. We argue that assessing species saturation is possible, but that many studies conducted to date have conceptual and methodological flaws that prevent us from currently attaining a good idea of the occurrence of species saturation.     

Concurrent niche and neutral processes in the competition-colonization model of species coexistence

The importance of neutral dynamics is contentiously debated in the ecological literature. This debate focuses on neutral theory's assumption of fitness equivalency among individuals, which conflicts with stabilizing fitness that promotes coexistence through niche differentiation. I take advantage of competition-colonization trade-offs between species of aquatic micro-organisms (protozoans and rotifers) to show that equalizing and stabilizing mechanisms can operate simultaneously. Competition trials between species with similar colonization abilities were less likely to result in competitive exclusion than for species further apart. While the stabilizing mechanism (colonization differences) facilitates coexistence at large spatial scales, species with similar colonization abilities also exhibited local coexistence probably due to fitness similarities allowing weak stabilizing mechanisms to operate. These results suggest that neutral- and niche-based mechanisms of coexistence can simultaneously operate at differing temporal and spatial scales, and such a spatially explicit view of coexistence may be one way to reconcile niche and neutral dynamics.     

Prediction of plant species distributions across six millennia

The usefulness of species distribution models (SDMs) in predicting impacts of climate change on biodiversity is difficult to assess because changes in species ranges may take decades or centuries to occur. One alternative way to evaluate the predictive ability of SDMs across time is to compare their predictions with data on past species distributions. We use data on plant distributions, fossil pollen and current and mid-Holocene climate to test the ability of SDMs to predict past climate-change impacts. We find that species showing little change in the estimated position of their realized niche, with resulting good model performance, tend to be dominant competitors for light. Different mechanisms appear to be responsible for among-species differences in model performance. Confidence in predictions of the impacts of climate change could be improved by selecting species with characteristics that suggest little change is expected in the relationships between species occurrence and climate patterns.     

Biotic attrition from tropical forests correcting for truncated temperature niches

Species migration in response to warming temperatures is expected to lead to ‘biotic attrition,’ or loss of local diversity, in areas where the number of species emigrating or going locally extinct exceeds the number immigrating. Biotic attrition is predicted be especially severe in the low‐lying hot tropics since elevated temperatures may surpass the observed tolerances of most extant species. It is possible, however, that the estimated temperature niches of many species are inaccurate and truncated with respect to their true tolerances due to the absence of hotter areas under current global climate. If so, these species will be capable of persisting in some areas where future temperatures exceed current temperatures, reducing rates of biotic attrition. Here, we use natural history collections data to estimate the realized thermal niches of > 2000 plant species from the tropical forests of South America. In accord with the truncation hypothesis, we find that the thermal niches of species from hot lowland areas are several degrees narrower than the thermal niches of species from cooler areas. We estimate rates of biotic attrition for South American tropical forests due to temperature increases ranging from 1 to 5 °C, and under two niche assumptions. The first is that the observed thermal niches truly reflect the plant's tolerances and that the reduction in niche breadth is due to increased specialization. The second is that lowland species have the same mean thermal niche breadth as nonlowland and nonequatorial species. The differences between these two models are dramatic. For example, using observed thermal niches we predict an almost complete loss of plant diversity in most South American tropical forests due to a 5 °C temperature increase, but correcting for possible niche truncation we estimate that most forests will retain > 50–70% of their current species richness. The different predictions highlight the importance of using fundamental vs. realized niches in predicting the responses of species to global climate change.     

Competition and specialization in an African forest carnivore community

Globally, human activities have led to the impoverishment of species assemblages and the disruption of ecosystem function. Determining whether this poses a threat to future ecosystem stability necessitates a thorough understanding of mechanisms underpinning community assembly and niche selection. Here, we tested for niche segregation within an African small carnivore community in Kibale National Park, Uganda. We used occupancy modeling based on systematic camera trap surveys and fine‐scale habitat measures, to identify opposing preferences between closely related species (cats, genets, and mongooses). We modeled diel activity patterns using kernel density functions and calculated the overlap of activity periods between related species. We also used co‐occupancy modeling and activity overlap analyses to test whether African golden cats Caracal aurata influenced the smaller carnivores along the spatial and/or temporal axes. There was some evidence that related species segregated habitat and activity patterns. Specialization was particularly strong among forest species. The cats and genets partitioned habitat, while the mongooses partitioned both habitat and activity period. We found little evidence for interference competition between African golden cats and other small carnivores, although weak interference competition was suggested by lower detection probabilities of some species at stations where African golden cats were present. This suggests that community assembly and coexistence in this ecosystem are primarily driven by more complex processes. The studied carnivore community contains several forest specialists, which are typically more prone to localized extinction. Preserving the observed community assemblage will therefore require the maintenance of a large variety of habitats, with a particular focus on those required by the more specialized carnivores.