Variability in Energy Influences Avian Distribution Patterns Across the USA

Habitat transformations and climate change are among the most important drivers of biodiversity loss. Understanding the factors responsible for the unequal distribution of species richness is a major challenge in ecology. Using data from the North American Breeding Bird Survey to measure species richness and a change metric extracted from the MODerate resolution Imaging Spectroradiometer (MODIS), we examined the influence of energy variability on the geographic distribution of avian richness across the conterminous U.S. and in the different ecoregions, while controlling for energy availability. The analysis compared three groups of birds: all species, Neotropical migrants, and permanent residents. We found that interannual variability in available energy explained more than half of the observed variation in bird richness in some ecoregions. In particular, energy variability is an important factor in explaining the patterns of overall bird richness and of permanent residents, in addition to energy availability. Our results showed a decrease in species richness with increasing energy variability and decreasing energy availability, suggesting that more species are found in more stable and more productive environments. However, not all ecoregions followed this pattern. The exceptions might reflect other biological factors and environmental conditions. With more ecoclimatic variability predicted for the future, this study provides insight into how energy variability influences the geographical patterns of species richness. PR designed study, performed research, analyzed data and wrote paper. CAL, MAL, AMP, VCR, PDC, and EFL designed study and wrote paper.     

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.     

Patterns of species diversity and phylogenetic structure of vascular plants on the Qinghai‐Tibetan Plateau

Large-scale patterns of species richness and the underlying mechanisms regulating these patterns have long been the central issues in biogeography and macroecology. Phylogenetic community structure is a result of combined effects of contemporary ecological interactions, environmental filtering, and evolutionary history, and it links community ecology with biogeography and trait evolution. The Qinghai-Tibetan Plateau provides a good opportunity to test the influence of contemporary climate on shaping species richness because of its unique geological history, cold climate, and high biodiversity. In this study, based on high-resolution distributions of ˜9000 vascular plant species, we explored how species richness and phylogenetic structure of vascular plants correlate with climates on the highest (and species rich) plateau on the Earth. The results showed that most of the vascular plants were distributed on the eastern part of the plateau; there was a strong association between species richness and climate, even after the effects of habitat heterogeneity were controlled. However, the responses of richness to climate remarkably depended on life-forms. Richness of woody plants showed stronger climatic associations than that of herbaceous plants; energy and water availability together regulated richness pattern of woody plants; whereas water availability predominantly regulated richness pattern of herbaceous plants. The phylogenetic structure of vascular species clustered in most areas of the plateau, suggesting that rapid speciation and environment filtering dominated the assembly of communities on the plateau. We further propose that biodiversity conservation in this area should better take into account ecological features for different life-forms and phylogenetic lineages.     

Environmental causes for plant biodiversity gradients

One of the most pervasive patterns observed in biodiversity studies is the tendency for species richness to decline towards the poles. One possible explanation is that high levels of environmental energy promote higher species richness nearer the equator. Energy input may set a limit to the number of species that can coexist in an area or alternatively may influence evolutionary rates. Within flowering plants (angiosperms), families exposed to a high energy load tend to be both more species rich and possess faster evolutionary rates, although there is no evidence that one drives the other. Specific environmental effects are likely to vary among lineages, reflecting the interaction between biological traits and environmental conditions in which they are found. One example of this is demonstrated by the high species richness of the iris family (Iridaceae) in the Cape of South Africa, a likely product of biological traits associated with reproductive isolation and the steep ecological and climatic gradients of the region. Within any set of conditions some lineages will tend to be favoured over others; however, the identity of these lineages will fluctuate with a changing environment, explaining the highly labile nature of diversification rates observed among major lineages of flowering plants.     

Diversity of European habitat types is correlated with geography more than climate and human pressure

Habitat richness, that is, the diversity of ecosystem types, is a complex, spatially explicit aspect of biodiversity, which is affected by bioclimatic, geographic, and anthropogenic variables. The distribution of habitat types is a key component for understanding broad‐scale biodiversity and for developing conservation strategies. We used data on the distribution of European Union (EU) habitats to answer the following questions: (i) how do bioclimatic, geographic, and anthropogenic variables affect habitat richness? (ii) Which of those factors is the most important? (iii) How do interactions among these variables influence habitat richness and which combinations produce the strongest interactions? The distribution maps of 222 terrestrial habitat types as defined by the Natura 2000 network were used to calculate habitat richness for the 10 km × 10 km EU grid map. We then investigated how environmental variables affect habitat richness, using generalized linear models, generalized additive models, and boosted regression trees. The main factors associated with habitat richness were geographic variables, with negative relationships observed for both latitude and longitude, and a positive relationship for terrain ruggedness. Bioclimatic variables played a secondary role, with habitat richness increasing slightly with annual mean temperature and overall annual precipitation. We also found an interaction between anthropogenic variables, with the combination of increased landscape fragmentation and increased population density strongly decreasing habitat richness. This is the first attempt to disentangle spatial patterns of habitat richness at the continental scale, as a key tool for protecting biodiversity. The number of European habitats is related to geography more than climate and human pressure, reflecting a major component of biogeographical patterns similar to the drivers observed at the species level. The interaction between anthropogenic variables highlights the need for coordinated, continental‐scale management plans for biodiversity conservation.     

Geographic variation in the relationship between large-scale environmental determinants and bat species richness

Several studies have already shown the close relationship between geographic gradients of biodiversity and distinct environmental determinants such as energy, environmental heterogeneity and seasonality. Nevertheless, whether and how such relationships vary around the globe remains poorly understood. Here we used spatial models to answer whether the bat species richness-environment relationship on a global scale are constant across geographic space. We also partitioned the contribution of the different environmental determinants on bat species richness at different regions of the globe. We found that the relationship between bat species richness and environment is not constant across geographic space and that the shared contributions of environmental determinants are more important than their unique contributions. We conclude that understanding geographic gradients of biodiversity and its environmental determinants, particularly for bats, is more complex than previously thought because the relationship between species richness and environment varies considerably across geographic space.     

Assessment of the relationships of geographic variation in species richness to climate and landscape variables within and among lineages of North American freshwater fishes

Aim Geographic variation in species richness is a well‐studied phenomenon. However, the unique response of individual lineages to environmental gradients in the context of general patterns of biodiversity across broad spatial scales has received limited attention. The focus of this research is to examine relationships between species richness and climate, topographic heterogeneity and stream channel characteristics within and among families of North American freshwater fishes. Location The United States and Canada. Methods Distribution maps of 828 native species of freshwater fishes were used to generate species richness estimates across the United States and Canada. Variation in species richness was predicted using spatially explicit models incorporating variation in climate, topography and/or stream channel length and stream channel diversity for all 828 species as well as for the seven largest families of freshwater fishes. Results The overall gradient of species richness in North American freshwater fishes is best predicted by a model incorporating variables describing climate and topography. However, the response of species richness to particular climate or landscape variables differed among families, with models possessing the highest predictive ability incorporating data on climate, topography and/or stream channel characteristics within a region. Main conclusions The correlations between species richness and abiotic variables suggest a strong influence of climate and physical habitat on the structuring of regional assemblages of North American freshwater fishes. However, the relationship between these variables and species richness varies among families, suggesting the importance of phylogenetic constraints on the regulation of geographic distributions of species.     

The role of diversification in causing the correlates of dioecy

Dioecy is reported to be correlated with a number of ecological traits, including tropical distribution, woody growth form, plain flowers, and fleshy fruits. Previous analyses have concentrated on determining whether dioecy is more likely to evolve in lineages possessing these traits, rather than considering the speciation and extinction rates of dioecious lineages with certain combinations of traits. To address the association between species richness in dioecious lineages as a function of the ecological traits, we compared the evolutionary success (i.e., relative species richness) of dioecious focal lineages with that of their nondioecious sister groups. This test was repeated for the evolutionary success of randomly chosen nondioecious lineages (control lineages) compared with their nondioecious sister groups. If the possession of certain ecological traits enhances the evolutionary success of dioecious lineages, we predict an association between the presence of these traits and relative species richness in the former, but not latter, set of sister-group comparisons. Dioecious focal lineages with a higher number of these traits experienced higher evolutionary success in sister-group comparisons, whereas no trend was found for the control focal lineages. The increase in evolutionary success was especially true for dioecious focal lineages that had a tropical distribution or fleshy fruit. We discuss how these results provide strong support for differential evolutionary success theories for the correlations between dioecy and the ecological traits considered.     

Host diversity drives parasite diversity: meta‐analytical insights into patterns and causal mechanisms

Statistical correlations of biodiversity patterns across multiple trophic levels have received considerable attention in various types of interacting assemblages, forging a universal understanding of patterns and processes in free‐living communities. Host–parasite interactions present an ideal model system for studying congruence of species richness among taxa as obligate parasites are strongly dependent upon the availability of their hosts for survival and reproduction while also having a tight coevolutionary relationship with their hosts. The present meta‐analysis examined 38 case studies on the relationship between species richness of hosts and parasites, and is the first attempt to provide insights into the patterns and causal mechanisms of parasite biodiversity at the community level using meta‐regression models. We tested the distinct role of resource (i.e. host) availability and evolutionary co‐variation on the association between biodiversity of hosts and parasites, while spatial scale of studies was expected to influence the extent of this association. Our results demonstrate that species richness of parasites is tightly correlated with that of their hosts with a strong average effect size (r= 0.55) through both host availability and evolutionary co‐variation. However, we found no effect of the spatial scale of studies, nor of any of the other predictor variables considered, on the correlation. Our findings highlight the tight ecological and evolutionary association between host and parasite species richness and reinforce the fact that host–parasite interactions provide an ideal system to explore congruence of biodiversity patterns across multiple trophic levels.     

Global biogeography and ecology of body size in birds

In 1847, Karl Bergmann proposed that temperature gradients are the key to understanding geographic variation in the body sizes of warm-blooded animals. Yet both the geographic patterns of body-size variation and their underlying mechanisms remain controversial. Here, we conduct the first assemblage-level global examination of 'Bergmann's rule' within an entire animal class. We generate global maps of avian body size and demonstrate a general pattern of larger body sizes at high latitudes, conforming to Bergmann's rule. We also show, however, that median body size within assemblages is systematically large on islands and small in species-rich areas. Similarly, while spatial models show that temperature is the single strongest environmental correlate of body size, there are secondary correlations with resource availability and a strong pattern of decreasing body size with increasing species richness. Finally, our results suggest that geographic patterns of body size are caused both by adaptation within lineages, as invoked by Bergmann, and by taxonomic turnover among lineages. Taken together, these results indicate that while Bergmann's prediction based on physiological scaling is remarkably accurate, it is far from the full picture. Global patterns of body size in avian assemblages are driven by interactions between the physiological demands of the environment, resource availability, species richness and taxonomic turnover among lineages.     

More individuals drive the species energy–area relationship in an experimental zooplankton community

The shape of the species–area relationship (SAR) often varies with the amount of available energy; SARs from high‐energy habitats typically have higher intercepts and steeper slopes than SARs from low‐energy habitats. Such patterns are often assumed to result from a shift in the mechanisms of coexistence between high and low energy habitats. However, a plausible but unexplored alternative mechanism emerges from proportional sampling, if there are simply more individuals in larger or more productive habitats, without the need to invoke differing coexistence mechanisms. Here, we examined proportional versus disproportional responses of a diverse assemblage of freshwater zooplankton to manipulations of experimental pond size and energy inputs. We found that higher energy treatments had higher species richness in large, but not small, ponds, leading to a steeper SAR with higher energy input. The total abundances of individuals also increased with energy in large, but not small ponds. By using a sample‐independent rarefaction technique (probability of interspecific encounter), we found that SAR patterns resulted from changes in the total, but not relative, abundance of individuals, and thus proportional, rather than disproportional, responses of species. Overall, our results emphasize the need to consider how both the total and relative abundances of species respond to ecological drivers such as energy and area before inferring the underlying mechanisms that lead to biodiversity patterns. Further, our results may implicate a proportionally smaller influence of energy on patterns of biodiversity when habitats are destroyed.     

Does the colonization of new biogeographic regions influence the diversification and accumulation of clade richness among the Corvides (Aves: Passeriformes)?

Regional variation in clade richness can be vast, reflecting differences in the dynamics of historical dispersal and diversification among lineages. Although it has been proposed that dispersal into new biogeographic regions may facilitate diversification, to date there has been limited assessment of the importance of this process in the generation, and maintenance, of broad‐scale biodiversity gradients. To address this issue, we analytically derive biogeographic regions for a global radiation of passerine birds (the Corvides, c. 790 species) that are highly variable in the geographic and taxonomic distribution of species. Subsequently, we determine rates of historical dispersal between regions, the dynamics of diversification following regional colonization, and spatial variation in the distribution of species that differ in their rates of lineage diversification. The results of these analyses reveal spatiotemporal differences in the build‐up of lineages across regions. The number of regions occupied and the rate of transition between regions both predict family richness well, indicating that the accumulation of high clade richness is associated with repeated expansion into new geographic areas. However, only the largest family (the Corvidae) had significantly heightened rates of both speciation and regional transition, implying that repeated regional colonization is not a general mechanism promoting lineage diversification among the Corvides.     

Fire as a key driver of Earth's biodiversity

Many terrestrial ecosystems are fire prone, such that their composition and structure are largely due to their fire regime. Regions subject to regular fire have exceptionally high levels of species richness and endemism, and fire has been proposed as a major driver of their diversity, within the context of climate, resource availability and environmental heterogeneity. However, current fire‐management practices rarely take into account the ecological and evolutionary roles of fire in maintaining biodiversity. Here, we focus on the mechanisms that enable fire to act as a major ecological and evolutionary force that promotes and maintains biodiversity over numerous spatiotemporal scales. From an ecological perspective, the vegetation, topography and local weather conditions during a fire generate a landscape with spatial and temporal variation in fire‐related patches (pyrodiversity), and these produce the biotic and environmental heterogeneity that drives biodiversity across local and regional scales. There have been few empirical tests of the proposition that ‘pyrodiversity begets biodiversity’ but we show that biodiversity should peak at moderately high levels of pyrodiversity. Overall species richness is greatest immediately after fire and declines monotonically over time, with postfire successional pathways dictated by animal habitat preferences and varying lifespans among resident plants. Theory and data support the ‘intermediate disturbance hypothesis’ when mean patch species diversity is correlated with mean fire intervals. Postfire persistence, recruitment and immigration allow species with different life histories to coexist. From an evolutionary perspective, fire drives population turnover and diversification by promoting a wide range of adaptive responses to particular fire regimes. Among 39 comparisons, the number of species in 26 fire‐prone lineages is much higher than that in their non‐fire‐prone sister lineages. Fire and its byproducts may have direct mutagenic effects, producing novel genotypes that can lead to trait innovation and even speciation. A paradigm shift aimed at restoring biodiversity‐maintaining fire regimes across broad landscapes is required among the fire research and management communities. This will require ecologists and other professionals to spread the burgeoning fire‐science knowledge beyond scientific publications to the broader public, politicians and media.     

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.     

Biodiversity of Jinggangshan Mountain: The Importance of Topography and Geographical Location in Supporting Higher Biodiversity

Diversity is mainly determined by climate and environment. In addition, topography is a complex factor, and the relationship between topography and biodiversity is still poorly understood. To understand the role of topography, i.e., altitude and slope, in biodiversity, we selected Jinggangshan Mountain (JGM), an area with unique topography, as the study area. We surveyed plant and animal species richness of JGM and compared the biodiversity and the main geographic characteristics of JGM with the adjacent 4 mountains. Gleason’s richness index was calculated to assess the diversity of species. In total, 2958 spermatophyte species, 418 bryophyte species, 355 pteridophyte species and 493 species of vertebrate animals were recorded in this survey. In general, the JGM biodiversity was higher than that of the adjacent mountains. Regarding topographic characteristics, 77% of JGM’s area was in the mid-altitude region and approximately 40% of JGM’s area was in the 10°–20° slope range, which may support more vegetation types in JGM area and make it a biodiversity hotspot. It should be noted that although the impact of topography on biodiversity was substantial, climate is still a more general factor driving the formation and maintenance of higher biodiversity. Topographic conditions can create microclimates, and both climatic and topographic conditions contribute to the formation of high biodiversity in JGM.     

Topography, energy and the global distribution of bird species richness

A major goal of ecology is to determine the causes of the latitudinal gradient in global distribution of species richness. Current evidence points to either energy availability or habitat heterogeneity as the most likely environmental drivers in terrestrial systems, but their relative importance is controversial in the absence of analyses of global (rather than continental or regional) extent. Here we use data on the global distribution of extant continental and continental island bird species to test the explanatory power of energy availability and habitat heterogeneity while simultaneously addressing issues of spatial resolution, spatial autocorrelation, geometric constraints upon species' range dynamics, and the impact of human populations and historical glacial ice-cover. At the finest resolution (1 degree), topographical variability and temperature are identified as the most important global predictors of avian species richness in multi-predictor models. Topographical variability is most important in single-predictor models, followed by productive energy. Adjusting for null expectations based on geometric constraints on species richness improves overall model fit but has negligible impact on tests of environmental predictors. Conclusions concerning the relative importance of environmental predictors of species richness cannot be extrapolated from one biogeographic realm to others or the globe. Rather a global perspective confirms the primary importance of mountain ranges in high-energy areas.     

Biodiversity Increases the Productivity and Stability of Phytoplankton Communities

Global biodiversity losses provide an immediate impetus to elucidate the relationships between biodiversity, productivity and stability. In this study, we quantified the effects of species richness and species combination on the productivity and stability of phytoplankton communities subject to predation by a single rotifer species. We also tested one mechanism of the insurance hypothesis: whether large, slow-growing, potentially-defended cells would compensate for the loss of small, fast-growing, poorly-defended cells after predation. There were significant effects of species richness and species combination on the productivity, relative yield, and stability of phytoplankton cultures, but the relative importance of species richness and combination varied with the response variables. Species combination drove patterns of productivity, whereas species richness was more important for stability. Polycultures containing the most productive single species, Dunaliella, were consistently the most productive. Yet, the most species rich cultures were the most stable, having low temporal variability in measures of biomass. Polycultures recovered from short-term negative grazing effects, but this recovery was not due to the compensation of large, slow-growing cells for the loss of small, fast-growing cells. Instead, polyculture recovery was the result of reduced rotifer grazing rates and persisting small species within the polycultures. Therefore, although an insurance effect in polycultures was found, this effect was indirect and unrelated to grazing tolerance. We hypothesize that diverse phytoplankton assemblages interfered with efficient rotifer grazing and that this “interference effect” facilitated the recovery of the most productive species, Dunaliella. In summary, we demonstrate that both species composition and species richness are important in driving patterns of productivity and stability, respectively, and that stability in biodiverse communities can result from an alteration in consumer functioning. Our findings underscore the importance of predator-prey dynamics in determining the relationships between biodiversity, productivity and stability in producer communities.     

Disentangling the relative effects of ambient energy,water availability,and energy–water balance on pteridophyte species richness at a landscape scale in China

Understanding what factors generate geographic variation in species richness is a fundamental goal of ecology and biogeography. Water and energy are considered as the major environmental factors influencing large-scale patterns of species richness, but their roles vary among taxa and regions. Pteridophytes are an ideal group of organisms for examining the relationship between species richness and their environment because the distribution of pteridophytes is usually in equilibrium with contemporary climate to a greater degree than those of seed plants and most terrestrial vertebrates partly due to the lightness of their spores, which is highly capable of long-distance dispersal by wind, and partly due to their single-spore reproduction strategy. Using correlation and regression analyses and structural equation modeling technique, we examine the relationship of pteridophyte species richness in 151 localities from across China with environmental factors representing energy, water, and energy–water balance. We found that pteridophyte species richness is correlated to water availability more strongly than to ambient energy. Furthermore, we found that of all environmental variables considered, energy–water balance has played the most important role in regulating pteridophyte species richness gradients in China.     

How Many Parasites Species a Frog Might Have? Determinants of Parasite Diversity in South American Anurans

There is an increasing interest in unveiling the dynamics of parasite infection. Understanding the interaction patterns, and determinants of host-parasite association contributes to filling knowledge gaps in both community and disease ecology. Despite being targeted as a relevant group for conservation efforts, determinants of the association of amphibians and their parasites in broad scales are poorly understood. Here we describe parasite biodiversity in South American amphibians, testing the influence of host body size and geographic range in helminth parasites species richness (PSR). We also test whether parasite diversity is related to hosts’ phylogenetic diversity. Results showed that nematodes are the most common anuran parasites. Host-parasite network has a nested pattern, with specialist helminth taxa generally associated with hosts that harbour the richest parasite faunas. Host size is positively correlated with helminth fauna richness, but we found no support for the association of host geographic range and PSR. These results remained consistent after correcting for uneven study effort and hosts’ phylogenic correlation. However, we found no association between host and parasite diversity, indicating that more diversified anuran clades not necessarily support higher parasite diversity. Overall, considering both the structure and the determinants of PRS in anurans, we conclude that specialist parasites are more likely to be associated with large anurans, which are the ones harbouring higher PSR, and that the lack of association of PSR with hosts’ clade diversification suggests it is strongly influenced by ecological and contemporary constrains.