Search

Aim: Addressing global environmental challenges requires access to biodiversity data across wide spatial, temporal and taxonomic scales. Availability of such data has increased exponentially recently with the proliferation of biodiversity databases. However, heterogeneous coverage, protocols, and standards have hampered integration among these databases. To stimulate the next stage of data integration, here we present a synthesis of major databases, and investigate (a) how the coverage of databases varies across taxonomy, space, and record type; (b) what degree of integration is present among databases; (c) how integration of databases can increase biodiversity knowledge; and (d) the barriers to database integration. Location: Global. Time period: Contemporary. Major taxa studied: Plants and vertebrates. Methods: We reviewed 12 established biodiversity databases that mainly focus on geographic distributions and functional traits at global scale. We synthesized information from these databases to assess the status of their integration and major knowledge gaps and barriers to full integration. We estimated how improved integration can increase the data coverage for terrestrial plants and vertebrates. Results: Every database reviewed had a unique focus of data coverage. Exchanges of biodiversity information were common among databases, although not always clearly documented. Functional trait databases were more isolated than those pertaining to species distributions. Variation and potential incompatibility of taxonomic systems used by different databases posed a major barrier to data integration. We found that integration of distribution databases could lead to increased taxonomic coverage that corresponds to 23 years’ advancement in data accumulation, and improvement in taxonomic coverage could be as high as 22.4% for trait databases. Main conclusions: Rapid increases in biodiversity knowledge can be achieved through the integration of databases, providing the data necessary to address critical environmental challenges. Full integration across databases will require tackling the major impediments to data integration: taxonomic incompatibility, lags in data exchange, barriers to effective data synchronization, and isolation of individual initiatives.

Biodiversity contributes to the ecological and climatic stability of the Amazon Basin1,2, but is increasingly threatened by deforestation and fire3,4. Here we quantify these impacts over the past two decades using remote-sensing estimates of fire and deforestation and comprehensive range estimates of 11,514 plant species and 3,079 vertebrate species in the Amazon. Deforestation has led to large amounts of habitat loss, and fires further exacerbate this already substantial impact on Amazonian biodiversity. Since 2001, 103,079–189,755 km2 of Amazon rainforest has been impacted by fires, potentially impacting the ranges of 77.3–85.2% of species that are listed as threatened in this region5. The impacts of fire on the ranges of species in Amazonia could be as high as 64%, and greater impacts are typically associated with species that have restricted ranges. We find close associations between forest policy, fire-impacted forest area and their potential impacts on biodiversity. In Brazil, forest policies that were initiated in the mid-2000s corresponded to reduced rates of burning. However, relaxed enforcement of these policies in 2019 has seemingly begun to reverse this trend: approximately 4,253–10,343 km2 of forest has been impacted by fire, leading to some of the most severe potential impacts on biodiversity since 2009. These results highlight the critical role of policy enforcement in the preservation of biodiversity in the Amazon. Remote-sensing estimates of fires and the estimated geographic ranges of thousands of plant and vertebrate species in the Amazon Basin reveal that fires have impacted the ranges of 77–85% of threatened species over the past two decades.

Massive biological databases of species occurrences, or georeferenced locations where a species has been observed, are essential inputs for modeling present and future species distributions. Location accuracy is often assessed by determining whether the observation geocoordinates fall within the boundaries of the declared political divisions. This otherwise simple validation is complicated by the difficulty of matching political division names to the correct geospatial object. Spelling errors, abbreviations, alternative codes, and synonyms in multiple languages present daunting name disambiguation challenges. The inability to resolve political division names reduces usable data and analysis of erroneous observations can lead to flawed results.Here, we present the Geographic Name Resolution Service (GNRS), an application for the correction, standardization and indexing of world political division names. The GNRS resolves political division names against a reference database that combines names and codes from GeoNames with geospatial object identifiers from the Global Administrative Areas Database (GADM). In a trial resolution of political division names extracted from >270 million species occurrences, only 1.9%, representing just 6% of occurrences, matched exactly to GADM political divisions in their original form. The GNRS was able to resolve, completely or in part, 92% of the remaining 378,568 political division names, or 86% of the full biodiversity occurrence dataset. In an assessment of geocoordinate accuracy for >239 million species occurrences, resolution of political divisions by the GNRS enabled detection of an order of magnitude more errors and an order of magnitude more error-free occurrences. By providing a novel solution to a major data quality impediment, the GNRS liberates a tremendous amount of biodiversity data for quantitative biodiversity research. The GNRS runs as a web service and can be accessed via an API, an R package, and a web-based graphical user interface. Its modular architecture is easily integrated into existing data validation workflows.

Phenology, the study of the timing of cyclical life history events and seasonal changes, is a fundamental aspect of how individual species, communities, and ecosystems will respond to climate change. Both biotic and abiotic phenological patterns are changing rapidly in response to changing seasonal temperatures and other climate-related drivers, and the consequences of these shifts for individual species and entire ecosystems are largely unknown. Landscape-scale simulations can address some of these needs for better predictions by demonstrating how phenology measures can vary with spatial and temporal grain of observations, and how phenological responses can vary with landscape heterogeneity and climate drivers. To explicitly examine the spatial and temporal scale-dependence of multiple phenology measures, we constructed simulated landscapes populated by virtual plant species with realistic phenologies and environmental sensitivities. This approach allowed us to examine phenology measures and environmental sensitivities along a continuum of spatial and temporal grains, while also controlling other aspects of sampling design. By relating measures of phenology calculated at a given spatiotemporal grain to average environmental conditions at that same grain size, we are able to determine observed environmental sensitivities for multiple phenological metrics at that spatial and temporal scale. We demonstrate that different phenological events change distinctly and predictably with spatial and temporal measurement scale, opening the way to incorporating scaling laws into predictions. Using plant flowering as our example, we identify that the timing of the beginnings or ends of an event (e.g., First Flower date, Last Flower date), can be especially sensitive to the spatial and temporal grain (or resolution) of observations. Our work provides an initial assessment of the role of observation scale in landscape phenology, and a general approach for incorporating scale-dependence into predictions of a variety of phenological time series. We thus set the stage for a new generation of empirical research in the field that builds off of multi-scale observations to understand how phenology across Earth9s ecosystems respond to environmental variability and change.

To meet the ambitious objectives of biodiversity and climate conventions, the international community requires clarity on how these objectives can be operationalized spatially and how multiple targets can be pursued concurrently. To support goal setting and the implementation of international strategies and action plans, spatial guidance is needed to identify which land areas have the potential to generate the greatest synergies between conserving biodiversity and nature’s contributions to people. Here we present results from a joint optimization that minimizes the number of threatened species, maximizes carbon retention and water quality regulation, and ranks terrestrial conservation priorities globally. We found that selecting the top-ranked 30% and 50% of terrestrial land area would conserve respectively 60.7% and 85.3% of the estimated total carbon stock and 66% and 89.8% of all clean water, in addition to meeting conservation targets for 57.9% and 79% of all species considered. Our data and prioritization further suggest that adequately conserving all species considered (vertebrates and plants) would require giving conservation attention to ~70% of the terrestrial land surface. If priority was given to biodiversity only, managing 30% of optimally located land area for conservation may be sufficient to meet conservation targets for 81.3% of the terrestrial plant and vertebrate species considered. Our results provide a global assessment of where land could be optimally managed for conservation. We discuss how such a spatial prioritization framework can support the implementation of the biodiversity and climate conventions.

Limiting climate change to less than 2°C is the focus of international policy under the climate convention (UNFCCC), and is essential to preventing extinctions, a focus of the Convention on Biological Diversity (CBD). The post-2020 biodiversity framework drafted by the CBD proposes conserving 30% of both land and oceans by 2030. However, the combined impact on extinction risk of species from limiting climate change and increasing the extent of protected and conserved areas has not been assessed. Here we create conservation spatial plans to minimize extinction risk in the tropics using data on 289 219 species and modeling two future greenhouse gas concentration pathways (RCP2.6 and 8.5) while varying the extent of terrestrial protected land and conserved areas from

Despite the many studies using trait-based approaches to assess the impact of environmental gradients in forest trait composition, the relative roles of (i) intraspecific variation in community assembly and (ii) microclimatic or fine scale abiotic variation in shaping local trait diversity remain poorly understood. To advance their understanding we tested several assumptions and predictions of trait driver theory (TDT). We quantified the shape of trait distributions related to tree carbon, nutrient economics and stem hydraulics across a small-scale but steep gradient of soil water availability. We utilized a unique and steep environmental gradient in the coastal Brazilian Atlantic forest (restinga) communities that spans a very short distance (207 +/-60 meters). We collected leaf and wood samples of tree species across 42 patches (or plots) of restinga forest. Furthermore, to detect if species directionally shift in niche space, we analyzed species composition in multidimensional hypervolume space. Despite short geographic distances, we observed large shifts in species replacement and intraspecific variation reflected by a directional shift in plant function. Consistent with TDT, we observe (i) trait distributions that are skewed in directions consistent with a forest responding to recent hotter and drier; (ii) peaked trait distributions, indicating strong functional convergence; and (iii) conditions decreasing means and variances of several leaf carbon and nutrient economic traits as well as stem hydraulic traits. Synthesis: Observed species replacements along the water table gradient and interspecific measures of functional diversity (community kurtosis and skewness) are consistent with strong phenotype/environmental matching of plant carbon, nutrient, and hydraulic strategies. We observe environmental filtering in both extremities of the gradient, selecting for acquisitive (wet) to conservative (dry) setup of traits. Similarly, species that span the entire water availability gradient are characterized by directional intraspecific shifts in multi-trait space that mirror interspecific shifts. Strong environmental gradients across short spatial scales provide unique systems to accurately assess assembly processes and address long-held assumptions and timely hypothesis predicted by trait driver theory.

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

The mechanistic origin and shape of body-size distributions within communities are of considerable interest in ecology. A recently proposed light-limitation model provides a good fit to the distribution of tree sizes in a tropical forest plot. The maximum entropy theory of ecology (METE) also predicts size distributions, but without explicit mechanistic assumptions, and thus its predictions should hold in ecosystems generally, regardless of whether they are light limited. A comparison of the form and success of the predictions of the model and the theory can provide insight into the role that mechanisms play in shaping patterns in macroecology. The prediction by the METE of the size distribution of organisms is remarkably similar in form to that of the model: power-law behaviour in the size range where the light-limitation model predicts a power law, and exponential behaviour in the size range where the model predicts an exponential tail. The METE prediction matches data widely, including data in ecosystems where light is not limiting. We show examples for three disparate communities: trees in a tropical forest plot; herbaceous plants in a treeless subalpine meadow; and island arthropods. We conclude that the success of METE's predicted form across systems, including those that are clearly not light limited, enriches our capacity to predict patterns in macroecology without making explicit mechanistic assumptions and provides a unified framework that can capture ubiquitous features of those patterns across diverse ecosystems governed by a variety of mechanisms.

Author(s): Newman, Erica A; Wilber, Mark Q; Kopper, Karen E; Moritz, Max A; Falk, Donald A; McKenzie, Don; Harte, John | Abstract: Macroecological studies have established widespread patterns of species diversity and abundance in ecosystems but have generally restricted their scope to relatively steady‐state systems. As a result, how macroecological metrics are expected to scale in ecosystems that experience natural disturbance regimes is unknown. We examine macroecological patterns in a fire‐dependent forest of Bishop pine (Pinus muricata ). We target two different‐aged stands in a stand‐replacing fire regime: a mature stand with a diverse understory and with no history of major disturbance for at least 40 yr, and one disturbed by a stand‐replacing fire 17 yr prior to measurement. We compare properties of these stands with macroecological predictions from the Maximum Entropy Theory of Ecology (METE ), an information entropy‐based theory that has proven highly successful in predicting macroecological metrics in multiple ecosystems and taxa. Ecological patterns in the mature stand more closely match METE predictions than do data from the more recently disturbed, mid‐seral stage stand. This suggests METE 's predictions are more robust in late‐successional, slowly changing, or steady‐state systems than those in rapid flux with respect to species composition, abundances, and organisms’ sizes. Our findings highlight the need for a macroecological theory that incorporates natural disturbance, perturbations, and ecological dynamics into its predictive capabilities, because most natural systems are not in a steady state.

Summary paragraphTo meet the ambitious objectives of biodiversity and climate conventions, countries and the international community require clarity on how these objectives can be operationalized spatially, and multiple targets be pursued concurrently1. To support governments and political conventions, spatial guidance is needed to identify which areas should be managed for conservation to generate the greatest synergies between biodiversity and nature’s contribution to people (NCP). Here we present results from a joint optimization that maximizes improvements in species conservation status, carbon retention and water provisioning and rank terrestrial conservation priorities globally. We found that, selecting the top-ranked 30% (respectively 50%) of areas would conserve 62.4% (86.8%) of the estimated total carbon stock and 67.8% (90.7%) of all clean water provisioning, in addition to improving the conservation status for 69.7% (83.8%) of all species considered. If priority was given to biodiversity only, managing 30% of optimally located land area for conservation may be sufficient to improve the conservation status of 86.3% of plant and vertebrate species on Earth. Our results provide a global baseline on where land could be managed for conservation. We discuss how such a spatial prioritisation framework can support the implementation of the biodiversity and climate conventions.

The severe deforestation of Brazil’s Atlantic Forest and increasing effects of climate change underscore the need to understand how tree species respond to climate and soil drivers. We studied 42 plots of coastal restinga forest, which is highly diverse and spans strong environmental gradients. We determined the forest physiognomy and functional composition, which are physical properties of a community that respond to climate and soil properties, to elucidate which factors drive community-level traits. To identify the most important environmental drivers of coastal Atlantic forest functional composition, we performed a forest inventory of all plants of diameter 5 cm and above. We collected wood samples and leaves from ∼85% of the most abundant plant species and estimated height, aboveground biomass (AGB), and basal area of individual plants, and the community-weighted specific leaf area (SLA). In addition to plant traits, we measured water table depth and 25 physicochemical soil parameters. We then parameterized several models for different hypotheses relating the roles of nutrients and soil to tropical forest diversity and functioning, as represented by plant traits. Hypotheses were formalized via generalized additive models and piecewise structural equation models. Water table depth, soil coarseness, potential acidity, sodium saturation index (SSI) and aluminum concentration were all components of the best models for AGB, height, basal area, and trait composition. Among the 25 environmental parameters measured, those related to water availability (water table depth and coarse sand), followed by potential acidity, SSI, and aluminum consistently emerged as the most important drivers of forest physiognomy and functional composition. Increases in water table depth, coarse sand, and soil concentration of aluminum negatively impacted all the measured functional traits, whereas SSI had a positive effect on AGB and plant height. These results suggest that sodium is not merely tolerated by Atlantic Forest restinga plant communities, but is important to their structure and functioning. Presence of aluminum in the soil had a complex relationship to overall basal area, possibly mediated by soil organic matter.

Biodiversity loss is a hallmark of our times, but predicting its consequences is challenging. Ecological interactions form complex networks with multiple direct and indirect paths through which the impacts of an extinction may propagate. Here we show that accounting for these multiple paths connecting species is necessary to predict how extinctions affect the integrity of ecological networks. Using an approach initially developed for the study of information flow, we estimate indirect effects in plant-pollinator networks and find that even those species with several direct interactions may have much of their influence over others through long indirect paths. Next, we perform extinction simulations in those networks and show that although traditional connectivity metrics fail in the prediction of coextinction patterns, accounting for indirect interaction paths allows predicting species' vulnerability to the cascading effects of an extinction event. Embracing the structural complexity of ecological systems contributes towards a more predictive ecology, which is of paramount importance amid the current biodiversity crisis.

The structure of ecological interactions is commonly understood through analyses of interaction networks. However, these analyses may be sensitive to sampling biases in both the interactors (the nodes of the network) and interactions (the links between nodes). These issues may affect the accuracy of empirically constructed ecological networks. We explore the properties of sampled ecological networks by simulating large-scale ecological networks with predetermined topologies, and sampling them with different mathematical procedures. Several types of modular networks were generated, intended to represent a wide variety of communities that vary in size and types of ecological interactions. We sampled these networks with different sampling designs that may be encountered in field experiments. The observed networks generated by each sampling process were analyzed with respect to number and size of components. We show that the sampling effort needed to estimate underlying network properties depends both on the sampling design and on network topology. Networks with random or scale-free modules require more complete sampling compared to networks whose modules are nested or bipartite. Overall, the structure of nested modules was the easiest to detect, regardless of sampling design. Sampling according to species degree was consistently found to be the most accurate strategy to estimate network structure. Conversely, sampling according to module results in an accurate view of certain modules, but fails to provide a global picture of the underlying network. We recommend that these findings are incorporated into the design of projects aiming to characterize large networks of species interactions in the field, to reduce sampling biases. The software scripts developed to construct and sample networks are provided for further explorations of network structure and comparisons to real interaction data., Comment: 35 pages, 4 figures

Although most conservation efforts address the direct, local causes of biodiversity loss, effective long‐term conservation will require complementary efforts to reduce the upstream economic pressures, such as demands for food and forest products, which ultimately drive these downstream losses. Here, we present a wildlife footprint analysis that links global losses of wild birds to consumer purchases across 57 economic sectors in 129 regions. The United States, India, China, and Brazil have the largest regional wildlife footprints, while per‐person footprints are highest in Mongolia, Australia, Botswana, and the United Arab Emirates. A US$100 purchase of bovine meat or rice products occupies approximately 0.1 km2 of wild bird ranges, displacing 1–2 individual birds, for 1 year. Globally significant importer regions, including Japan, the United Kingdom, Germany, Italy, and France, have large footprints that drive wildlife losses elsewhere in the world and represent important targets for consumption‐focused conservation attention.

Aim: Maximum entropy (MaxEnt) models promise a novel approach for understanding community assembly and species abundance patterns. One of these models, the "Maximum Entropy Theory of Ecology" (METE) reproduces many observed species abundance patterns, but is based on an aggregated representation of community structure that does not resolve species identity or explicitly represent species-specific functional traits. In this paper, METE is compared to "Very Entropic Growth" (VEG), a MaxEnt model with a less aggregated representation of community structure that represents species (more correctly, functional types) in terms of their per capita metabolic rates. We examine the contribution of metabolic traits to the patterns of community assembly predicted by VEG and, through aggregation, compare the results with METE predictions in order to gain insight into the biological factors underlying observed patterns of community assembly. Innovation: We formally compare two MaxEnt-based community models, METE and VEG, that differ as to whether or not they represent species-specific functional traits. We empirically test and compare the metabolic predictions of both models, thereby elucidating the role of metabolic traits in patterns of community assembly. Main Conclusions: Our analysis reveals that a key determinant of community metabolic patterns is the "density of species" distribution, defined as the intrinsic number of species with metabolic rates in a given range that are available to a community prior to filtering by environmental constraints. Our analysis suggests that appropriate choice of of the density of species in VEG may lead to more realistic predictions than METE, for which this distribution is not defined, and thus opens up new ways to understanding the link between functional traits and patterns of community assembly.

Macroecological studies have generally restricted their scope to relatively steady-state systems, and as a result, how biodiversity and abundance metrics are expected to scale in disturbance-dependent ecosystems is unknown. We examine macroecological patterns in a fire-dependent forest of Bishop pine (Pinus muricata). We target two different-aged stands in a stand-replacing fire regime, one a characteristically mature stand with a diverse understory, and one more recently disturbed by a stand-replacing fire (17 years prior to measurement). We compare the stands using macroecological metrics of species richness, abundance and spatial distributions that are predicted by the Maximum Entropy Theory of Ecology (METE), an information-entropy based theory that has proven highly successful in predicting macroecological metrics across a wide variety of systems and taxa. Ecological patterns in the mature stand more closely match METE predictions than do data from the recently disturbed stand. This suggests METE’s predictions are more robust in late-successional, slowly changing, or steady-state systems than those in rapid flux with respect to species composition, abundances, and organisms’ sizes. Our findings highlight the need for a macroecological theory that incorporates natural disturbance and other ecological perturbations into its predictive capabilities, because most natural systems are not in a steady state.

Anthropogenic (or human-caused) wildfire is an increasingly important driver of ecological change on Pacific islands including southeastern Polynesia, but fire ecology studies are almost completely absent for this region. Where observations do exist, they mostly represent descriptions of fire effects on plant communities before the introduction of invasive species in the modern era. Understanding the effects of wildfire in southeastern Polynesian island vegetation communities can elucidate which species may become problematic invasives with continued wildfire activity. We investigate the effects of wildfire on vegetation in three low-elevation sites (45-379 m) on the island of Mo’orea in the Society Islands, French Polynesia, which are already heavily impacted by past human land use and invasive exotic plants, but retain some native flora. In six study areas (3 burned and 3 unburned comparisons), we placed 30 transects across sites and collected species and abundance information at 390 points. We analyzed each local community of plants in three categories: natives, those introduced by Polynesians before European contact (1767 C.E.), and those introduced since European contact. Burned areas had the same or lower mean species richness than paired comparison sites. Although wildfire did not affect the proportions of native and introduced species, it may increase the abundance of introduced species on some sites. Non-metric multidimensional scaling indicates that (not recently modified) comparison plant communities are more distinct from one another than are those on burned sites. We discuss conservation concerns for particular native plants absent from burned sites, as well as invasive species (including Lantana camara and Paraserianthes falcataria) that may be promoted by fire in the Pacific.

Networks provide one of the best representations for ecological communities, composed of many species with sometimes complex connections between them. Yet the methodological literature allowing one to analyze and extract meaning from ecological networks is dense, fragmented, and unwelcoming. We provide a general overview to the field of using networks in community ecology, outlining both the intent of the different measures, their assumptions, and the contexts in which they can be used. When methodologically justified, we suggest good practices to use in the analysis of ecological networks. We anchor this synopsis with examples from empirical studies, and conclude by highlighting what identified as needed future developments in the field.

Many fundamental properties of ecological systems and interactions are tied to body size and a related metric, the metabolic rate distribution, both within and across species. A previously proposed maximum entropy theory of ecology (METE) predicts numerous interrelated macroecological patterns, including spatial distributions of individuals within species, abundance distributions across species, species area relationships, and distributions of metabolic rates of all individuals within a community. Extensive tests of METE's macroecological predictions generally support the theory, but two related predictions have not been evaluated against full community census data: the distribution of metabolic rates of individuals within species as a function of the abundance of the species and the distribution of average individual metabolic rates across species. We test the metabolic predictions of METE for herbaceous plants in a subalpine meadow and show that while this theory realistically predicts the distribution of individual metabolic rates across the entire community, the within- and across-species predictions generally fail. We also test the energy-equivalence type prediction that arises as a consequence of the prediction for the distribution of average individual metabolic rates across species. We suggest several possible explanations for the empirical deviations from theory, and distinguish between the expected deviations caused by ecological disturbance and those deviations that might be corrected within the theory.

The principle of Maximum Entropy (MaxEnt) promises a novel approach for understanding community assembly. Despite reproducing a variety of observed species abundance patterns, MaxEnt models in ecology have been hampered by disparate model assumptions and interpretations. A recurring challenge is that MaxEnt predictions are highly sensitive to the level of detail used to describe the community being modeled, and there seems to be no reason to prefer one level of detail over another. Here we present of formal unification of two previously developed MaxEnt models which differ in their level of detail, but which are otherwise mathematically similar. The less detailed model, “Maximum Entropy Theory of Ecology” (METE), does not resolve species identity or explicitly represent species-specific traits. The more detailed model, “Very Entropic Growth” (VEG), defines each separate species by its per capita metabolic rate e and assumes a “density of species” function ρ ( e ) representing the distribution of e in the metacommunity. A formal comparison of METE and VEG then highlights ρ ( e ) as a key determinant of local community assembly. In particular, appropriate choice of ρ ( e ) in VEG can produce more realistic predictions for the metabolic-rank distribution of local communities than METE, which does not explicitly account for metacommunity structure. This opens new avenues of inquiry about what determines metacommunity structure in nature and suggests possible ways to improve METE.

Although Borrelia burgdorferi sensu lato (s.l.) are found in a great diversity of vertebrates, most studies in North America have focused on the role of mammals as spirochete reservoir hosts. We investigated the roles of birds as hosts for subadult Ixodes pacificus ticks and potential reservoirs of the Lyme disease spirochete B. burgdorferi sensu stricto (s.s.) in northwestern California. Overall, 623 birds representing 53 species yielded 284 I. pacificus larvae and nymphs. We used generalized linear models and zero-inflated negative binomial models to determine associations of bird behaviors, taxonomic relationships and infestation by I. pacificus with borrelial infection in the birds. Infection status in birds was best explained by taxonomic order, number of infesting nymphs, sampling year, and log-transformed average body weight. Presence and counts of larvae and nymphs could be predicted by ground- or bark-foraging behavior and contact with dense oak woodland. Molecular analysis yielded the first reported detection of Borrelia bissettii in birds. Moreover, our data suggest that the Golden-crowned Sparrow (Zonotrichia atricapilla), a non-resident species, could be an important reservoir for B. burgdorferi s.s. Of 12 individual birds (9 species) that carried B. burgdorferi s.l.-infected larvae, no birds carried the same genospecies of B. burgdorferi s.l. in their blood as were present in the infected larvae removed from them. Possible reasons for this discrepancy are discussed. Our study is the first to explicitly incorporate both taxonomic relationships and behaviors as predictor variables to identify putative avian reservoirs of B. burgdorferi s.l. Our findings underscore the importance of bird behavior to explain local tick infestation and Borrelia infection in these animals, and suggest the potential for bird-mediated geographic spread of vector ticks and spirochetes in the far-western United States.

A theory of macroecology based on the maximum information entropy (MaxEnt) inference procedure predicts that the log-log slope of the species-area relationship (SAR) at any spatial scale is a specified function of the ratio of abundance, N(A), to species richness, S(A), at that scale. The theory thus predicts, in generally good agreement with observation, that all SARs collapse onto a specified universal curve when local slope, z(A), is plotted against N(A)/S(A). A recent publication, however, argues that if it is assumed that patterns in macroecology are independent of the taxonomic choices that define assemblages of species, then this principle of "taxon invariance" precludes the MaxEnt-predicted universality of the SAR. By distinguishing two dimensions of the notion of taxon invariance, we show that while the MaxEnt-based theory predicts universality regardless of the taxonomic choices that define an assemblage of species, the biological characteristics of assemblages should under MaxEnt, and do in reality, influence the realism of the predictions.