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.     

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.     

Species-energy relationships at the macroecological scale: a review of the mechanisms

Correlations between the amount of energy received by an assemblage and the number of species that it contains are very general, and at the macro-scale such species-energy relationships typically follow a monotonically increasing curve. Whilst the ecological literature contains frequent reports of such relationships, debate on their causal mechanisms is limited and typically focuses on the role of energy availability in controlling the number of individuals in an assemblage. Assemblages from high-energy areas may contain more individuals enabling species to maintain larger, more viable populations, whose lower extinction risk elevates species richness. Other mechanisms have, however, also been suggested. Here we identify and clarify nine principal mechanisms that may generate positive species-energy relationships at the macro-scale. We critically assess their assumptions and applicability over a range of spatial scales, derive predictions for each and assess the evidence that supports or refutes them. Our synthesis demonstrates that all mechanisms share at least one of their predictions with an alternative mechanism. Some previous studies of species-energy relationships appear not to have recognised the extent of shared predictions, and this may detract from their contribution to the debate on causal mechanisms. The combination of predictions and assumptions made by each mechanism is, however, unique, suggesting that, in principle, conclusive tests are possible. Sufficient testing of all mechanisms has yet to be conducted, and no single mechanism currently has unequivocal support. Each may contribute to species-energy relationships in some circumstances, but some mechanisms are unlikely to act simultaneously. Moreover, a limited number appear particularly likely to contribute frequently to species-energy relationships at the macro-scale. The increased population size, niche position and diversification rate mechanisms are particularly noteworthy in this context.     

Abundance, species richness and energy availability in the North American avifauna

Aim To determine how species richness, abundance, biomass, energy use and mean number of individuals per species scale with environmental energy availability in wintering and breeding avian assemblages, and to contrast assemblages of (i) common and rare species and (ii) breeding residents and migrants. To assess whether such patterns are compatible with the ‘more individuals hypothesis’ (MIH) that high‐energy areas are species‐rich because they support larger populations that are buffered against extinction. Location The North American continent (latitudinal range 23.4 °?48.1 °N; longitudinal range 124.2°?68.7° W). Methods Avian species richness, abundance, biomass and energy use were calculated for 295 Resident Bird Count plots. Environmental energy availability was measured using ambient temperature and the Normalized Difference Vegetation Index (NDVI), a close correlate of plant productivity. Analyses took plot area into account, and were conducted (with and without taking habitat type into account) using general linear models and spatial mixed models. Results Positive species–energy relationships were exhibited by both wintering and breeding assemblages, but were stronger in the former. The structure of winter assemblages responded more strongly to temperature than NDVI, while breeding assemblages tended to respond more strongly to NDVI. Breeding residents responded to annual measures of energy availability while breeding migrants and the winter assemblage responded more strongly to seasonal measures. In the winter assemblage, rare and common species exhibited species–energy relationships of a similar strength, but common breeding species exhibited a much stronger relationship than rare breeding species. In both breeding and wintering assemblages, abundance, biomass and energy use increased with energy availability and species richness. Energy availability was a poor predictor of the mean number of individuals per species. Main conclusions The nature of the species–energy relationship varies seasonally and with the manner in which energy availability is measured. Our data suggest that residents are less able to respond to seasonal fluxes in resource availability than long‐distance migrants. Increasing species richness and energy availability is associated with increasing numbers of individuals, biomass and energy use. While these observations are compatible with the MIH our data provide only equivocal support for this hypothesis, as the rarest species do not exhibit the strongest species–energy relationships.     

A system in which available energy per se controls alpha diversity: marine pelagic birds

An attractive explanation for large-scale gradients of species richness is that trophic energy flux defines living systems. It has yet to be shown that available energy may matter per se, that is, directly and independent of other potential determinants that are usually inescapably correlated (e.g., area, glacial history, or habitat complexity). By using a comprehensive conceptual framework addressing the variation of species richness, we report that in communities of birds regularly foraging in marine pelagic waters during the breeding season, species richness is above all directly linked to the appropriation of metabolic energy. Auxiliary energy supplied by wind and waves is likely to mitigate energetic constraints and thereby codetermine the expansion of niche space, along with an array of other subordinate factors. We emphasize that this system is markedly different from studied communities of terrestrial endotherms or marine exotherms in which habitat complexity and mutagenic solar radiation/temperature, respectively, may be more decisive than the appropriation of trophic energy flux shares as such. While the seabird system suggests that species-energy curves may sometimes directly translate into species-energy relationships, this situation may be rare rather than typical.     

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.     

Abundance matters: a field experiment testing the more individuals hypothesis for richness–productivity relationships

The more individuals hypothesis (MIH) postulates that productivity increases species richness by increasing mean equilibrium population size, thereby reducing the probability of local extinction. We tested the MIH for invertebrates colonizing microcosms that simulated tree holes by manipulating productivity through additions of leaf or animal detritus and subsequently determining the relationships among richness, total abundance, abundance per species, and measures of productivity. We quantified productivity as the rate of microorganism protein synthesis, microorganism metabolic rate, nutrient ion concentration, and type and amount of detritus. Microcosms with animal detritus attracted more species, more individuals per species, and more total individuals than did microcosms with similar amounts of leaf detritus. Relationships between richness or abundance and productivity varied with date. Richness in June increased as a linear function of productivity, whereas the power function predicted by the MIH fit best in July. Abundance in June and July was best described by a power function of productivity, but the linear function predicted by the MIH fit best in September. Abundance per species was best described by a power function of productivity in June and July. Path analysis showed that the indirect effect of productivity through abundance on richness that is predicted by MIH was important in all months, and that direct links between productivity and richness were unnecessary. Our results support many of the predictions of the MIH, but they also suggest that the effects of abundance on richness may be more complex than expected.     

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.     

Species–energy relationships and habitat complexity in bird communities

Species–energy theory is a commonly invoked theory predicting a positive relationship between species richness and available energy. The More Individuals Hypothesis (MIH) attempts to explain this pattern, and assumes that areas with greater food resources support more individuals, and that communities with more individuals include more species. Using a large dataset for North American birds, I tested these predictions of the MIH, and also examined the effect of habitat complexity on community structure. I found qualitative support for the relationships predicted by the MIH, however, the MIH alone was inadequate for fully explaining richness patterns. Communities in more productive sites had more individuals, but they also had more even relative abundance distributions such that a given number of individuals yielded a greater number of species. Richness and evenness were also higher in structurally complex forests compared to structurally more simple grasslands when controlling for available energy.     

Effect of Environmental and Temporal Factors on Patterns of Rarity of Ephemeroptera in Stream of the Brazilian Cerrado

Patterns of species’ abundance and occurrence over time and space allow division of species into (i) common species, which are abundant, but have a low diversity, and (ii) rare species, which are far more diverse and less abundant. Understanding the relationships among these two species groups and how they are affected by environmental conditions is a major challenge for ecologists, especially considering the distinction between local environmental factors and regional factors and variations in abundance over the course of the year. In this study, we focused on the long-term relationship between the abundance of rare and common ephemeropterans and abiotic factors on local and regional scales. Our hypotheses are that common species will be affected primarily by regional environmental variables (i), whereas rare species will be influenced more by temporal variation (ii). Together, both local and regional abiotic variables, plus temporal variation, best explained the abundance of the common species, whereas temporal variation was the best predictor of rare species. Considering the theoretical aspects and the empirical evidence, we discuss the results based on the plasticity of the common species and the life cycle of the rare ones. We believe that our findings reinforce the need for the deconstruction of communities for a deeper understanding of their relationships with abiotic variables and, in particular, the specific aspects of these relationships in the context of the different guilds of the community.     

Sleeping with the ‘enemy’: hybridization of an endangered tree weta

While hybridization is an important part of the evolutionary process, for rare species mating with more common species hybridization can increase the risks of extinction. By mating with heterospecifics rare species waste valuable reproductive resources and as a result population sizes may decline. If introgression occurs, the rare species can become genetically swamped by alleles from the more common species, rendering it effectively extinct. As a consequence of these risks, hybridization with the more common species Hemideina femorata (Canterbury tree weta) may lead to the extinction of the range restricted species H. ricta (Banks Peninsula tree weta) on Banks Peninsula. The current study uses spatial interpolation to model the distribution of each species and the potential sympatric zone to guide sampling efforts. Polymorphic microsatellite markers and mitochondrial sequence data were used to determine the extent of hybridization between H. ricta and H. femorata. The results confirm that hybridization is possible between these species. However, hybrids and introgression appear to be very rare, suggesting that reproductive isolating barriers are present but incomplete. The threat of extinction to H. ricta via hybridization with H. femorata is low but extreme loss of habitat may cause changes to population densities that could increase the risks of hybridization. Therefore, landowners should be encouraged to conserve native bush.     

The more‐individuals hypothesis revisited: the role of community abundance in species richness regulation and the productivity–diversity relationship

Species richness increases with energy availability, yet there is little consensus as to the exact processes driving this species–energy relationship. The most straightforward explanation is the more‐individuals hypothesis (MIH). It states that higher energy availability promotes a higher total number of individuals in a community, which consequently increases species richness by allowing for a greater number of species with viable populations. Empirical support for the MIH is mixed, partially due to the lack of proper formalisation of the MIH and consequent confusion as to its exact predictions. Here, we review the evidence of the MIH and evaluate the reliability of various predictions that have been tested. There is only limited evidence that spatial variation in species richness is driven by variation in the total number of individuals. There are also problems with measures of energy availability, with scale‐dependence, and with the direction of causality, as the total number of individuals may sometimes itself be driven by the number of species. However, even in such a case the total number of individuals may be involved in diversity regulation. We propose a formal theory that encompasses these processes, clarifying how the different factors affecting diversity dynamics can be disentangled.     

Amphibian Species Contribute Similarly to Taxonomic,but not Functional and Phylogenetic Diversity: Inferences from Amphibian Biodiversity on Emei Mountain

Understanding the relationships between species,communities,and biodiversity are important challenges in conservation ecology.Current biodiversity conservation activities usually focus on species that are rare,endemic,distinctive,or at risk of extinction.However,empirical studies of whether such species contribute more to aspects of biodiversity than common species are still relatively rare.The aim of the present study was to assess the contribution of individual amphibian species to different facets of biodiversity,and to test whether species of conservation interest contribute more to taxonomic,functional,and phylogenetic diversity than do species without special conservation status.To answer these questions,19 000 simulated random communities with a gradient of species richness were created by shuffling the regional pool of species inhabiting Emei Mountain.Differences of diversity values were then computed before and after removing individual species in these random communities.Our results indicated that although individual species contributed similarly to taxonomic diversity,their contribution to functional and phylogenetic diversity was more idiosyncratic.This was primarily driven by the diverse functional attributes of species and the differences in phylogenetic relationships among species.Additionally,species of conservation interest did not show a significantly higher contribution to any facet of biodiversity.Our results support the claims that the usefulness of metrics based only on species richness is limited.Instead,assemblages that include species with functional and phylogenetic diversity should be protected to maintain biodiversity.     

Species diversity preserved in different numbers of nature reserves of the same total area

The expected number of species occurring in different numbers of reserves of the same total area is examined on different assumptions of the spatial distribution and the probability of extinction. The advantage of one large reserve or several smaller ones of equal total area depends on the spatial distributions of species and the stage after the establishement of reserves. In general, several smaller reserves maintain more species immediately after the establishments unless the spatial distribution are uniform or random, whereas one large reserve excels several smaller ones after some rare species have gone extinct unless the spatial distributions are strongly contagious. Since the extinction of rare species must be facilitated as the size of each reserve reduces, the area of a reserve should be larger than the critical area that ensures the persistence of the species. Hence it is concluded that one or a few large reserves are a better strategy in order to maintain the species diversity.     

Sheltered from the storm? Population viability analysis of a rare endemic under periodic catastrophe regimes

Rare species are important targets for biodiversity conservation efforts because rarity often equates to small populations and increased endangerment. Rare species are prone to stochastic extinction events and may be particularly susceptible to catastrophes. Therefore, understanding how rare species respond to disturbances is critical for evaluating extinction risk and guiding conservation managers. Population viability analyses (PVAs) are essential for assessing rare species' status yet they seldom consider catastrophic events. Accordingly, we present a PVA of a rare tropical epiphyte, Lepanthes caritensis (Orchidaceae), under simulated disturbance regimes to evaluate its demographics and extinction risk. We aimed to test how demographic models incorporating catastrophes affect population viability estimates. Our goal was to better guide management of these orchids and other rare plants. Results revealed L. caritensis numbers have declined recently, but projected growth rates indicated that most subpopulations should increase in size if undisturbed. Still, projection models show that moderate catastrophes reduce growth rates, increase stochasticity in subpopulation sizes, and elevate extinction risk. Severe catastrophes had a more pronounced effect in simulations; growth rates fell below replacement level, there was greater variation in projected population sizes, and extinction risk was significantly higher. PVAs incorporating periodic catastrophes indicate that rare species may have greater extinction probabilities than standard models suggest. Thus, precautionary conservation measures should be taken in disturbance prone settings and we encourage careful monitoring after environmental catastrophes. Future rare plant PVAs should incorporate catastrophes and aim to determine if rescue and reintroduction efforts are necessary after disturbances to insure long-term population viability.     

The species-area-energy relationship

Area and available energy are major determinants of species richness. Although scale dependency of the relationship between energy availability and species richness (the species-energy relationship) has been documented, the exact relationship between the species-area and the species-energy relationship has not been studied explicitly. Here we show, using two extensive data sets on avian distributions in different biogeographic regions, that there is a negative interaction between energy availability and area in their effect on species richness. The slope of the species-area relationship is lower in areas with higher levels of available energy, and the slope of the species-energy relationship is lower for larger areas. This three-dimensional species-area-energy relationship can be understood in terms of probabilistic processes affecting the proportions of sites occupied by individual species. According to this theory, high environmental energy elevates species' occupancies, which depress the slope of the species-area curve.     

Patterns and correlates of plant diversity differ between common and rare species in a neotropical dry forest

Determining which factors affect species richness is important for conservation theory and practice. However, richness of common and rare species may be affected by different factors. We use an extensive inventory of woody plants from a tropical dry forest landscape in Yucatan, Mexico to assess the unique effects of environmental variables, spatial dependence of sampling sites, forest stand age and the combined effect of all groups of variables on species richness of woody plants with different levels of rarity (common, intermediate, rare, very rare)—according to their abundance, habitat specificity and spatial distribution range in the landscape. Analyzing separately common species and those with different levels of rarity uncovered contrasting patterns and correlates of species richness that were not apparent when focusing on all woody plants. In particular, richness of common and intermediate species was influenced mainly by environmental factors, whereas richness of very rare species was affected mostly by the unique effect of spatial dependence of sampling sites, suggesting a main role of environmental filtering and dispersal limitation, respectively. However, common and very rare species also responded inversely to some landscape metrics, revealing contrasting environmental preferences of these groups of species. These contrasting results suggest different underlying mechanisms and the need for very different conservation strategies. Therefore, basic and applied research on tropical forest biodiversity should consider separately species with different levels of rarity, focusing on which factors control variation in each level, and paying special attention to very rare species, generally the most specious and vulnerable to local extinction.     

Correlates of extinction risk of birds from two Indonesian islands

Size of distributional range, position in the range, body size and diet are some of the ecological traits that may correlate with local abundance. Evolutionary phenomena such as taxon cycles, acting over much greater time periods, may also influence abundance and promote species extinction. This paper assesses which of a wide range of ecological and historic traits best predict the variation in abundance of tropical forest birds on Sumba and Buru islands in Wallacea (Indonesia). In addition we seek to determine which traits predict species' ability to adapt to secondary or logged forest. The most important correlates of both abundance and ability to transfer were those related to the evolutionary history of the species within the Wallacean Archipelago and not the traits that were more directly related to species ecology. These relationships are maintained when allowance is made for phylogenetic relationships. Our interpretation of the results is that recent colonists to an island are initially rare in the indigenous forest habitat but concomitant with an adaptation to local conditions they gradually become more abundant and taxonomically distinct from other populations of the same species. These results apparently contradict the taxon cycle hypothesis but this may be a result of our focus on indigenous forest habitats rather than on a wider range dominated by anthropogenic ones.     

Severe extinction and rapid recovery of mammals across the Cretaceous–Palaeogene boundary,and the effects of rarity on patterns of extinction and recovery

The end‐Cretaceous mass extinction ranks among the most severe extinctions of all time; however, patterns of extinction and recovery remain incompletely understood. In particular, it is unclear how severe the extinction was, how rapid the recovery was and how sampling biases might affect our understanding of these processes. To better understand terrestrial extinction and recovery and how sampling influences these patterns, we collected data on the occurrence and abundance of fossil mammals to examine mammalian diversity across the K‐Pg boundary in North America. Our data show that the extinction was more severe and the recovery more rapid than previously thought. Extinction rates are markedly higher than previously estimated: of 59 species, four survived (93% species extinction, 86% of genera). Survival is correlated with geographic range size and abundance, with widespread, common species tending to survive. This creates a sampling artefact in which rare species are both more vulnerable to extinction and less likely to be recovered, such that the fossil record is inherently biased towards the survivors. The recovery was remarkably rapid. Within 300 000 years, local diversity recovered and regional diversity rose to twice Cretaceous levels, driven by increased endemicity; morphological disparity increased above levels observed in the Cretaceous. The speed of the recovery tends to be obscured by sampling effects; faunas show increased endemicity, such that a rapid, regional increase in diversity and disparity is not seen in geographically restricted studies. Sampling biases that operate against rare taxa appear to obscure the severity of extinction and the pace of recovery across the K‐Pg boundary, and similar biases may operate during other extinction events.     

Life in a Double-Hotspot: The Transformation of Hawaiian Passerine Bird Diversity following Invasion and Extinction

Although recent research has shown that non-indigenous species often increase local-scale species richness, few have documented how such increases translate into compositional changes across biological scales. In particular, transformations of biodiversity patterns may be acute within regions that are simultaneously extinction and invasion hotspots (i.e. double-hotspots), such as the Hawaiian Islands. Nevertheless, modification of diversity relationships in such places are rarely quantified. Here, I utilize passerine non-indigenous species introductions and native species extinctions on Hawaii to quantitatively explore the changing relationship between within- (alpha), between- (beta), and across-island (gamma) diversity. My results indicate that, even after incorporating the enrichment effects of non-indigenous species invasions, across-island passerine diversity has dropped substantially. Nevertheless, within-island diversity has remained largely unchanged, or in some cases increased. Perhaps the more profound changes in diversity have come from the loss of between-island diversity. Because nearly all native Hawaiian passerines are extinct or near extinction, the current diversity relationships are inordinately influenced by patterns in the transportation and establishment of non-indigenous birds. These human-induced ‘dispersal’ patterns are markedly different from natural ones. In addition, these dispersal patterns may be unique to vagile species such as birds, thus indicating that transformations of diversity within other groups (e.g. plants or freshwater fishes) currently inhabiting Hawaii may differ. These results suggest the need to explore how alteration of diversity relationships translate into the loss of ecosystem services, or other valued components of biodiversity.