Do body size and diet breadth affect partitioning of species diversity? A test with forest Lepidoptera

Two of the major themes resulting from recent macroecological research are the central roles that body size and niche breadth may play as determinants of species geographical distribution. Unanswered questions, however, linger regarding how similarities in body size or niche breadth affect the allocation of α‐ and β‐diversity across spatial scales. Using data on moth diversity in the eastern deciduous forest of North America, we tested the predictions that smaller‐bodied and diet‐restricted species would have lower levels of α‐diversity within forest stands and greater β‐diversity at higher sampling scales compared to larger or more generalist species. Moths were sampled using a nested sampling design consisting of three hierarchical levels: 20 forest stands, 5 sites and 3 ecoregions. Body size for 492 species was estimated as mean forewing length, and diet breadth was assessed from the published literature. Moth species were then classified according to body size (small or large) or diet breadth (generalist or restricted), and partitioning was conducted on each group. Diversity partitions for large‐ and small‐bodied species yielded similar patterns. When observed diversity components differed from those derived from our null model, a consistent pattern was observed: α‐diversity was greater than expected, β‐diversity among forest stands was less than expected, and β‐diversity among sites and ecoregions was higher than expected. In contrast, diet‐restricted moths contributed significantly less to stand‐level α‐diversity than generalist feeders. Furthermore, specialists contributed to a greater proportion of β‐diversity across scales compared to generalist moths. Because absolute measures of β‐diversity among stands were greater for generalists than for restricted feeders, we suggest that regional β‐diversity of forest moths may be influenced by several possible factors: intraspecific aggregation of diet‐restricted species, local fluctuations in population size of eruptive generalists and small geographical distributions of generalist moths than predicted by the geographical extent of putative host plants     

Body size, biomic specialization and range size of African large mammals

Aim The goal of this paper is to examine the relationships between body size, biomic specialization and range size in the African large mammals, which are defined as all the African species corresponding to the orders Primates, Carnivora, Proboscidea, Perissodactyla, Hyracoidea, Tubulidentata, Artiodactyla and Pholidota. Location The study used the large mammal assemblage from Africa. Methods The degree of biomic specialization of African large mammals is investigated using the biomic specialization index (BSI) for each mammal species, based on the number of biomes it inhabits. Range size for each species is measured as the latitudinal extent of the geographical distribution of the species. We have analysed our data using both conventional cross‐species analyses and phylogenetically independent contrasts. Results There is a polygonal relationship between species biomic specialization and body size. While small and large species are biomic specialists, medium‐sized species are distributed along the whole range of biomic specialization. The latitudinal extent–body size relationship is approximately triangular. Small‐bodied species may have either large or small ranges, whereas large‐bodied ones have only large ranges. A positive correlation between latitudinal extent and biomic specialization is evident, although their relationship is better described as triangular. Main conclusions We found a polygonal relationship between species biomic specialization and body size, which agrees with previous arguments that small‐bodied species have more limited dispersal and, therefore, they may come to occupy a lesser proportion of their potential inhabitable biomes. On the other hand, large‐bodied species are constrained to inhabit biomes with a high productivity. A polygonal relationship between species latitudinal extent and body size in African large mammals agrees with previous studies of the relationship between range size and body size in other continents. The independent study of the macroecological pattern in biomic specialization highlights different factors that influence the body size–range size relationship. Although body size is usually implicated as a correlate of both specialization and geographical range size in large mammals, much of the variation in these variables cannot be attributed to size differences but to biome specific factors such as productivity, area, history, etc.     

Evaluating,partitioning, and mapping the spatial autocorrelation component in ecological niche modeling: a new approach based on environmentally equidistant records

Most species data display spatial autocorrelation that can affect ecological niche models (ENMs) accuracy‐statistics, affecting its ability to infer geographic distributions. Here we evaluate whether the spatial autocorrelation underlying species data affects accuracy‐statistics and map the uncertainties due to spatial autocorrelation effects on species range predictions under past and future climate models. As an example, ENMs were fitted to Qualea grandiflora (Vochysiaceae), a widely distributed plant from Brazilian Cerrado. We corrected for spatial autocorrelation in ENMs by selecting sampling sites equidistant in geographical (GEO) and environmental (ENV) spaces. Distributions were modelled using 13 ENMs evaluated by two accuracy‐statistics (TSS and AUC), which were compared with uncorrected ENMs. Null models and the similarity statistics I were used to evaluate the effects of spatial autocorrelation. Moreover, we applied a hierarchical ANOVA to partition and map the uncertainties from the time (across last glacial maximum, pre‐insustrial, and 2080 time periods) and methodological components (ENMs and autocorrelation corrections). The GEO and ENV models had the highest accuracy‐statistics values, although only the ENV model had values higher than expected by chance alone for most of the 13 ENMs. Uncertainties from time component were higher in the core region of the Brazilian Cerrado where Q. grandiflora occurs, whereas methodological components presented higher uncertainties in the extreme northern and southern regions of South America (i.e. outside of Brazilian Cerrado). Our findings show that accounting for autocorrelation in environmental space is more efficient than doing so in geographical space. Methodological uncertainties were concentrated in outside the core region of Q. grandiflora's habitat. Conversely, uncertainty due to time component in the Brazilian Cerrado reveals that ENMs were able to capture climate change effects on Q. grandiflora distributions.     

Challenging Wallacean and Linnean shortfalls: knowledge gradients and conservation planning in a biodiversity hotspot

Knowledge about biodiversity remains inadequate because most species living on Earth were still not formally described (the Linnean shortfall) and because geographical distributions of most species are poorly understood and usually contain many gaps (the Wallacean shortfall). In this paper, we developed models to infer the size and placement of geographical ranges of hypothetical non‐described species, based on the range size frequency distribution of anurans recently described in the Cerrado Biome, on the level of knowledge (number of inventories) and on surrogates for habitat suitability. The rationale for these models is as follow: (1) the range size frequency distribution of these species should be similar to the range‐restricted species, which have been most recently described in the Cerrado Biome; (2) the probability of new discoveries will increase in areas with low biodiversity knowledge, mainly in suitable areas, and (3) assuming range continuity, new species should occupy adjacent cells only if the level of knowledge is low enough to allow the existence of undiscovered species. We ran a model based on the number of inventories only, and two models combining effects of number of inventories and two different estimates of habitat suitability, for a total of 100 replicates each. Finally, we performed a complementary analysis using simulated annealing to solve the set‐covering problem for each simulation (i.e. finding the smallest number of cells so that all species are represented at least once), using extents of occurrence of 160 species (131 real anuran species plus 29 new simulated species). The revised reserve system that included information about unknown or poorly sampled taxa significantly shifted northwards, when compared to a system based on currently known species. This main result can be explained by the paucity of biodiversity data in this part of the biome, associated with its relatively high habitat suitability. As a precautionary measure, weighted by the inferred distribution data, the prioritization of a system of reserves in the north part of the biome appears to be defensible.     

The influence of spatial resolution on macroecological patterns of range size variation: a case study using parrots (Aves: Psittaciformes) of the world

Aim To assess the extent to which the resolution at which geographical range sizes are measured influences macroecological patterns in this variable. Location Global. Methods Data on the geographical ranges of parrot species were digitized, and a Geographic Information System used to produce nine range size estimates for each species using different degrees of spatial resolution. The inter‐correlation of these estimates was then compared, together with their patterns of covariation with population size, body mass and migratory behaviour (across species and controlling for phylogeny), their pattern of phylogenetic correlation, and the frequency distributions of the different measures. Results Strong correlations exist among all nine range size measures across species, albeit that measures of similar spatial resolution are more strongly correlated. All measures show similar patterns of covariation with population size, body mass and migratory behaviour, and similar patterns of phylogenetic correlation. The skewness of frequency distributions increases towards zero as the resolution of the range size measure declines. Main conclusions The results of macroecological analyses are little affected by the resolution with which geographical range sizes are calculated, at least for the parrots of the world. Previously published studies based on crude measures of range size would be unlikely to have produced markedly different conclusions had they used more refined range size metrics.     

Relationships between body size and geographical range size among Australian mammals: has human impact distorted macroecological patterns?

Extinction and artificial reduction in the size of geographical ranges of many species have occurred extensively across the globe because of human activities. In particular, Australian mammals have suffered heavily in the last two hundred years, with the highest number of reported cases of mammal extinctions anywhere. In the present study, we investigated the extent to which human impact has affected contemporary macroecological patterns in Australian terrestrial mammals. After examining patterns relating to body size and range size among the contemporary mammal fauna, we removed the effects of the last two hundred years of human impact by exploring patterns in the pre‐European assemblage. This permitted us to determine whether contemporary macroecological patterns are distortions of pre‐European patterns. In contrast to the expected pattern of a significant positive relationship between body size and range size, our results showed no significant association for the complete fauna in both cross‐species and phylogenetic analyses, even when data were corrected for species extinctions and range reductions. Analyses within families and among species with the same dietary strategy revealed three significant positive relationships (Macropodidae, Peramelidae, and herbivores) and one significant negative relationship (insectivores) within the contemporary assemblage that disappeared when the pre‐European assemblage was analysed. A positive relationship also emerged in the pre‐European assemblage for the Vombatidae that was not apparent in the contemporary fauna. Thus, correcting for human impact revealed important distortions among contemporary macroecological relationships that have been brought about by human‐induced range reduction and extinction. These findings not only provide further evidence that the Australian continent presents a unique and valuable opportunity with which to test the generality of macroecological patterns, but they also have important ramifications for the analysis and interpretation of contemporary macroecological datasets.     

Sampling effort and species richness assessment: a case study on Brazilian spiders

The knowledge on the geographical distribution of species is essential for building biogeographical and macroecological hypotheses. However, information on this regard is not distributed uniformly in space and usually come from biased sampling. The aim of this study is to quantify the influence of spatial distribution of sampling effort on the assessment of spider species richness in Brazil. We used a database of spider distribution records in Brazil, based on the taxonomic and biodiversity survey literature. The results show that the Atlantic Forest was better sampled and had the highest spider species richness among the Brazilian biomes. The Amazon, though having large collecting gaps and high concentration of records around major cities and rivers, showed the second highest number of species. The Pampa had a large number of records, but these are concentrated near a major city in the transition zone with the Atlantic Forest. The Cerrado, Caatinga and Pantanal had shown to be poorly sampled and, consequently, were among the lesser known biomes regarding the spider fauna. A linear regression analysis showed that the spider species richness in Brazil is strongly correlated to the number of records. However, we have identified areas potentially richest in species, which strongly deviate from the predicted by our analyses. Our results show that it is possible to access the spatial variation in species richness, as long as the variation in sampling effort is taken into account.     

Local richness and distribution of the lizard fauna in natural habitat mosaics of the Brazilian Cerrado

We investigate local lizard richness and distribution in central Brazilian Cerrado, harbouring one of the least studied herpetofaunas in the Neotropical region. Our results are based on standardized samplings at 10 localities, involving 2917 captures of 57 lizard species in 10 families. Local richness values exceeded most presented in earlier studies and varied from 13 to 28 species, with modal values between 19 and 28 species. Most of the Cerrado lizard fauna is composed of habitat‐specialists with patchy distributions in the mosaic of grasslands, savannas and forests, resulting in habitat‐structured lizard assemblages. Faunal overlap between open and forested habitats is limited, and forested and open areas may act as mutual barriers to lizard distribution. Habitat use is influenced by niche conservatism in deep lineages, with iguanians and gekkotans showing higher use of forested habitats, whereas autarchoglossans are richer and more abundant in open habitats. Contrary to trends observed in Cerrado birds and large mammals, lizard richness is significantly higher in open, interfluvial habitats that dominate the Cerrado landscape. Between‐localities variation in lizard richness seems tied to geographical distance, landscape history and phylogenetic constraints, factors operating in other well‐studied lizard faunas in open environments. Higher richness in dominant, open interfluvial habitats may be recurrent in Squamata and other small‐bodied vertebrates, posing a threat to conservation as these habitats are most vulnerable to the fast, widespread and ongoing process of habitat destruction in central Brazil.     

Richness patterns, species distributions and the principle of extreme deconstruction

Aim To analyse the global patterns in species richness of Viperidae snakes through the deconstruction of richness into sets of species according to their distribution models, range size, body size and phylogenetic structure, and to test if environmental drivers explaining the geographical ranges of species are similar to those explaining richness patterns, something we called the extreme deconstruction principle. Location Global. Methods We generated a global dataset of 228 terrestrial viperid snakes, which included geographical ranges (mapped at 1° resolution, for a grid with 7331 cells world‐wide), body sizes and phylogenetic relationships among species. We used logistic regression (generalized linear model; GLM) to model species geographical ranges with five environmental predictors. Sets of species richness were also generated for large and small‐bodied species, for basal and derived species and for four classes of geographical range sizes. Richness patterns were also modelled against the five environmental variables through standard ordinary least squares (OLS) multiple regressions. These subsets are replications to test if environmental factors driving species geographical ranges can be directly associated with those explaining richness patterns. Results Around 48% of the total variance in viperid richness was explained by the environmental model, but richness sets revealed different patterns across the world. The similarity between OLS coefficients and the primacy of variables across species geographical range GLMs was equal to 0.645 when analysing all viperid snakes. Thus, in general, when an environmental predictor it is important to model species geographical ranges, this predictor is also important when modelling richness, so that the extreme deconstruction principle holds. However, replicating this correlation using subsets of species within different categories in body size, range size and phylogenetic structure gave more variable results, with correlations between GLM and OLS coefficients varying from –0.46 up to 0.83. Despite this, there is a relatively high correspondence (r = 0.73) between the similarity of GLM‐OLS coefficients and R² values of richness models, indicating that when richness is well explained by the environment, the relative importance of environmental drivers is similar in the richness OLS and its corresponding set of GLMs. Main conclusions The deconstruction of species richness based on macroecological traits revealed that, at least for range size and phylogenetic level, the causes underlying patterns in viperid richness differ for the various sets of species. On the other hand, our analyses of extreme deconstruction using GLM for species geographical range support the idea that, if environmental drivers determine the geographical distribution of species by establishing niche boundaries, it is expected, at least in theory, that the overlap among ranges (i.e. richness) will reveal similar effects of these environmental drivers. Richness patterns may be indeed viewed as macroecological consequences of population‐level processes acting on species geographical ranges.     

Do New World pitvipers “scale‐down” at high elevations? Macroecological patterns of scale characters and body size

Bergmann's rule describes the macroecological pattern of increasing body size in response to higher latitudes and elevations. This pattern is extensively documented in endothermic vertebrates, within and among species; however, studies involving ectotherms are less common and suggest no consistent pattern for amphibians and reptiles. Moreover, adaptive traits, such as epidermal features like scales, have not been widely examined in conjunction with Bergmann's rule, even though these traits affect physiological processes, such as thermoregulation, which are hypothesized as underlying mechanisms for the pattern. Here, we investigate how scale characters correlate with elevation among 122 New World pitviper species, representing 15 genera. We found a contra‐Bergmann's pattern, where body size is smaller at higher elevations. This pattern was mainly driven by the presence of small‐bodied clades at high elevations and large‐bodied clades at low elevations, emphasizing the importance of taxonomic scope in studying macroecological patterns. Within a subset of speciose clades, we found that only Crotalus demonstrated a significant negative relationship between body size and elevation, perhaps because of its wide elevational range. In addition, we found a positive correlation between scale counts and body size but no independent effect of elevation on scale numbers. Our study increases our knowledge of Bergmann's rule in reptiles by specifically examining characters of squamation and suggests a need to reexamine macroecological patterns for this group.     

Macroecological analyses reveal historical factors influencing seed dispersal strategies in Brazilian Araucaria forests

Aim Our objective was to evaluate the influence of distinct macroecological factors (space, floristics and environment) on the variation in seed dispersal strategies of shrubs and trees in the Araucaria forest biome in the southern and south‐eastern Brazilian highlands. We hypothesize that history‐related factors (space, floristics) are major determinants of the proportion of endo‐ and syn‐zoochorous species in Araucaria forests, despite the current gradients in environmental conditions. Location Araucaria forest is the main forest biome in southern Brazil, at altitudes above 500 m a.s.l. (latitude ≤ 30° S). Their northern limit in Brazil is at latitude 20° S, where forests occur at elevations above 1000 m a.s.l. Methods We compiled information from 27 floristic checklists comprising shrub and tree species distributed along the geographical range of the Brazilian Araucaria forest biome. We classified species as zoochorous and non‐zoochorous, based on morphological attributes of their diaspores. Sites were described by geographical coordinates (latitude, longitude) and five environmental variables. We evaluated separately the influence of floristic, spatial and environmental variables on zoochory using correspondence analysis and linear regressions. Further, we evaluated causal connections between these variable groups using Mantel tests and path analysis. Results Zoochory increased with both latitude and longitude. Regression analysis showed that rainfall seasonality was the only environmental variable explaining the variation in the proportion of zoochorous species. All pairwise Mantel correlations between space, rainfall seasonality, floristics and zoochory were significant. Path analysis showed that rainfall seasonality was strongly determined by spatial distances between sites, and floristics was directly determined by rainfall seasonality. Further zoochory was mostly determined by floristics. Main conclusions Taking into account historical factors in the interpretation of macroecological patterns improves our understanding of biodiversity gradients. Hypotheses based on long‐term dynamics of distinct floristic groups provide some useful insights into patterns shown by studies elsewhere. Here we offer an analytical solution to incorporate history‐related factors into macroecological analyses. While history‐based hypotheses do not replace any other ideas concerning macroecological patterns, they are likely to improve our understanding on factors determining present‐day ecological patterns.     

Conservation biogeography of anurans in Brazilian Cerrado

The increasing rates of declines in anuran populations worldwide are creating demands for urgent strategies to maximize conservation efforts. This may be critical in regions for which few detailed data on diversity, abundance and distribution are available, such as in the Cerrado of Central Brazil. In this paper, we used a macroecological approach based on the extent of occurrence of 131 species of Anura (Amphibia) in the Cerrado region to design a regional network of potential areas that preserves all anuran species. The final network, obtained using a simulation annealing algorithm based on complementarity, has a total of 17 cells, widely distributed throughout the biome. Minimum costs solutions were obtained in respect to total human population size, soybean production and bovine density, because these are the factors associated with human occupation that historically are more likely to cause broad scale habitat losses. The macro-scale approach used here can provide overall guidelines for conservation and define the focus for more local and effective conservation efforts.     

Anuran species richness,complementarity and conservation conflicts in Brazilian Cerrado

Broad-scale correlations between species richness and human population suggest that processes driving species richness, mainly related to high ecological productivity, may also drive human populations. However, it is still under debate if this coincidence implies conflicts between biodiversity conservation and human development. In this paper, we analyzed the relationships among human population size, species richness and irreplaceability in Brazilian Cerrado. We analyzed a dataset with 131 species of anurans distributed in 181 cells with 1° of spatial resolution covering the biome. We found a positive correlation between human population size and anuran species richness (r = 0.46; P = 0.033 with 19.5 geographically effective degrees of freedom, v*), but the irreplaceability of each cell was poorly correlated with human population size (r = 0.075; P = 0.323; v* = 173.9). The 17 cells in the 97 optimal reserve networks contained a total human population ranging from 2942,195 to 4319,845 people, representing on average 11.8% of the human population in the entire Cerrado grid. The comparison of these observed values with 10,000 values from randomly generated networks suggests a relatively high flexibility in optimal complementarity sets for reserve selection. Our results indicated that correlation between richness and human population does not necessarily result in conflicts, given the opportunities for conciliating conservation and development. However, the analyses performed here are initial explorations within the framework of conservation biogeography, so more detailed studies are necessary to establish conservation planning at regional and local scales.     

Metacommunity‐scale biodiversity regulation and the self‐organised emergence of macroecological patterns

There exist a number of key macroecological patterns whose ubiquity suggests that the spatio‐temporal structure of ecological communities is governed by some universal mechanisms. The nature of these mechanisms, however, remains poorly understood. Here, we probe spatio‐temporal patterns in species richness and community composition using a simple metacommunity assembly model. Despite making no a priori assumptions regarding biotic spatial structure or the distribution of biomass across species, model metacommunities self‐organise to reproduce well‐documented patterns including characteristic species abundance distributions, range size distributions and species area relations. Also in agreement with observations, species richness in our model attains an equilibrium despite continuous species turnover. Crucially, it is in the neighbourhood of the equilibrium that we observe the emergence of these key macroecological patterns. Biodiversity equilibria in models occur due to the onset of ecological structural instability, a population‐dynamical mechanism. This strongly suggests a causal link between local community processes and macroecological phenomena.     

Cryptic lineages and Pleistocene population expansion in a Brazilian Cerrado frog

Diversification of South American species endemic to open habitats has been attributed to both Tertiary events and Pleistocene climatic fluctuations. Nonetheless, phylogeographical studies of taxa in these regions are few, precluding generalizations about the timing and processes leading to differentiation and speciation. We inferred population structure of Hypsiboas albopunctatus, a frog widely distributed in the Brazilian Cerrado. Three geographically distinct lineages were recovered in our phylogeny. The Chapada dos Guimarães (CG) clade was the first to diverge from other populations and contains multiple haplotypes from a single population in western Cerrado, probably representing a cryptic species. The southeast clade (SE) includes populations along the southeastern limit of the range within the historical distribution of the Brazilian Atlantic forest. Finally, the Central Cerrado (CC) group includes haplotypes from the interior of Brazil that are paraphyletic relative to the SE clade. Analyses of historical demography indicate significant population expansion in the CC and SE populations, likely associated with colonization of newly formed open habitats. The divergence of populations in the CG clade occurred in the late Miocene, concordant with the uplift of the central Brazilian plateau. Divergence of the SE clade from the CC occurred during the mid‐Pleistocene. Thus, both Tertiary geological events and Pleistocene climatic fluctuations promoted divergences among lineages. Our study reveals a complex history of diversification in the Cerrado, a morphoclimatic domain highly threatened because of anthropogenic habitat alteration. We identified surprisingly deep divergences in a widely distributed frog, indicating that the Cerrado is not a barrier‐free habitat and that its diversity is likely underestimated.     

Biogeography of body size in Pacific island birds

Many insular vertebrates have undergone rapid and dramatic changes in body size compared to their mainland counterparts. Here we explore the relationship between two well known patterns of island body size – the tendency for large‐bodied species to dwarf and small‐bodied species to get larger on islands, known as the “island rule”, and the scaling of maximum and minimum body size of island assemblages with island area. Drawing on both fossil and modern data, we examined the relationship between body size and island area in Pacific island birds, both within clades and at the island assemblage level. We found that the size of the smallest bird on each island decreased with island area while the maximum body size increased with island area. Similarly, within clades the body size of small‐bodied groups decreased and large‐bodied groups increased from small to large islands, consistent with the island rule. However, the magnitude of size change within clades was not sufficient to explain the overall scaling of maximum size with island area. Instead, the pattern was driven primarily by the evolution of very large, flightless birds on large islands. Human‐mediated extinctions on islands over the past few millennia severely impacted large, flightless birds, to the effect that this macroecological pattern has been virtually erased. After controlling for effects of biogeographic region and island area, we found island productivity to be the best predictor of maximum size in flightless birds. This result, and the striking similarities in maximum body size between flightless birds and island mammals, suggests a common energetic mechanism linking body size and landmass area in both the island rule and the scaling of island body size extremes.     

Abundance–occupancy relationships in deep sea wood fall communities

The generally positive relationship between the number of sites a species occupies and its average abundance within those sites provides an important link between population processes occurring at different spatial scales. Although such abundance–occupancy relationships (AORs) have been documented across a very wide range of taxa and in many different environments, little is known of such patterns in Earth's largest ecosystem, the deep sea. Wood falls – derived from natural or anthropogenic inputs of wood into the oceans – constitute an important deep‐sea habitat, habouring their own unique communities ultimately entirely dependent on the wood for chemical energy. In this study we take advantage of the unique features of an experimental wood fall deployment to examine AORs for the first time in deep‐sea invertebrates. The study design combines advantages of both experimental (tractability, control of key environmental parameters) and observational (natural colonisation by taxonomically diverse communities) studies. We show that the interspecific AOR is strongly positive across the 48 species occurring over 32 wood fall communities. The precise form of the AOR is mediated by both species‐level life history (body size) and by the colonisation stage at which communities were harvested, but not by environmental energy (wood fall size). Temporal dynamics within species are also generally consistent with positive intraspecific AORs. This support for positive AORs in the deep sea is an important extension of a macroecological generality into a new environment offering considerable potential for further testing and developing mechanistic macroecological theories.