The biogeographic history of ruminant faunas determines the phylogenetic structure of their assemblages at different scales

Phylogenetic community structure may help us understand how macroecological and macroevolutionary processes shape assemblages at large geographical scales. In this paper, we test hypotheses linking the formation of large‐scale assemblages, evolutionary processes and macroecology. To provide new insight into ruminant biogeography and evolution, phylogenetic community structure metrics were calculated for faunal assemblages at four hierarchical levels. Phylogenetic relatedness indices (net relatedness index and nearest taxon index) were determined for 59 ruminant assemblages at the landscape scale and scale of their respective climate domains (continuous biome stretches). Species pools at the global and biogeographic realm levels were used to construct null observation models. Significantly, assemblages were selected if they were distributed across biogeographic realms and represented all the world's biomes. Non‐random patterns were also tested for biogeographic realms within the global ruminant species pool. By examining ruminant assemblages at different scales we were able to observe that ruminant faunas show a distribution mainly limited within the boundaries of their biogeographic realms. However, the diversification of some clades was found to be restricted to extremely arid domains in the Sahara and Arabia. The random patterns featured by other extreme climate domains could reflect phylogenetically heterogeneous filling by less biome‐restricted lineages outside Africa.     

Species richness correlates of raw and standardized co‐occurrence metrics

Measuring β‐diversity and changes in species composition across multiple sites and environments is a major research focus in macroecology, and a variety of metrics have been proposed to quantify species co‐occurrence patterns in a species × site occurrence matrix. However, indices of β‐diversity and species co‐occurrence are often statistically dependent on the number of species in an assemblage. We compared the results of several common co‐occurrence metrics with patterns generated by a spatially explicit neutral model simulation. We found that all measures of co‐occurrence and β‐diversity, whether raw, rescaled or standardized by a null model expectation, were highly correlated with the total species richness of the landscape. The one important exception were the effect sizes of the fixed–fixed null model algorithm, which preserves row and column sums of the original matrix during matrix randomization. Our results call for a careful interpretation of meta‐analyses of assemblages that differ widely in species richness. At a minimum, observed species richness should be used as a statistical covariate in regression analyses, and results of the fixed–fixed algorithm should be compared carefully with the results of other randomization tests.     

Landscape‐level tree cover predicts species richness of large‐bodied frugivorous birds in forest fragments

Large‐bodied frugivorous birds play an important role in dispersing large‐sized seeds in Neotropical rain forests, thereby maintaining tree species richness and diversity. Conversion of contiguous forest land to forest fragments is thought to be driving population declines in large‐bodied frugivores, but the mechanistic drivers of this decline remain poorly understood. To assess the importance of fragment‐level versus local landscape attributes in influencing the species richness of large‐bodied (>100 g) frugivorous birds, we surveyed 15 focal species in 22 forest fragments (2.7 to 33.6 ha, avg. = 16.0 ha) in northwest Ecuador in 2014. Fragment habitat variables included density of large trees, canopy openness and height, and fragment size; landscape variables included elevation and the proportion of tree cover within a 1 km radius of each fragment. At both the individual species level, and across the community of 12 species of avian frugivore we detected, there was higher richness and probability of presence in fragments with more tree cover on surrounding land. This tendency was particularly pronounced among some endangered species. These findings corroborate the idea that partially forested land surrounding fragments may effectively increase the suitable habitat for forest‐dwelling frugivorous birds in fragmented landscapes. These results can help guide conservation priorities within fragmented landscapes, with particular reference to retaining trees and reforesting to attain high levels of tree cover in areas between forest patches.     

Putting insects on the map: near‐global variation in sphingid moth richness along spatial and environmental gradients

Despite their vast diversity and vital ecological role, insects are notoriously underrepresented in biogeography and conservation, and key broad‐scale ecological hypotheses about them remain untested – largely due to generally incomplete and very coarse spatial distribution knowledge. Integrating records from publications, field work and natural history collections, we used a mixture of species distribution models and expert estimates to provide geographic distributions and emergent richness patterns for all ca 1000 sphingid moth species found outside the Americas in high spatial detail. Total sphingid moth richness, the first for a higher insect group to be documented at this scale, shows distinct maxima in the wet tropics of Africa and the Oriental with notable decay toward Australasia. Using multivariate models controlling for spatial autocorrelation, we found that primary productivity is the dominant environmental variable associated with moth richness, while temperature, contrary to our predictions, is an unexpectedly weak predictor. This is in stark contrast to the importance we identify for temperature as a niche variable of individual species. Despite divergent life histories, both main sub‐groups of moths exhibit these relationships. Tribal‐level deconstruction of richness and climatic niche patterns indicate idiosyncratic effects of biogeographic history for some of the less species‐rich tribes, which in some cases exhibit distinct richness peaks away from the tropics. The study confirms, for a diverse insect group, overall richness associations of remarkable similarity to those documented for vertebrates and highlights the significant within‐taxon structure that underpins emergent macroecological patterns. Results do not, however, meet predictions from vertebrate‐derived hypotheses on how thermoregulation affects the strength of temperature–richness effects. Our study thus broadens the taxonomic focus in this data‐deficient discourse. Our procedures of processing incomplete, scattered distribution data are a template for application to other taxa and regions.     

Species distribution modelling as a macroecological tool: a case study using New World amphibians

Although species distribution modelling (SDM) is widely accepted among the scientific community and is increasingly used in ecology, conservation biology and biogeography, methodological limitations generate potential problems for its application in macroecology. Using amphibian species richness in North and South America, we compare species richness patterns derived from SDM maps and ‘expert’ maps to evaluate if: 1) richness patterns derived from SDM are biased toward climate‐based explanations for diversity when compared to expert maps, since SDM methods are typically based on climatic variables; and 2) SDM is a reliable tool for generating richness maps in hyperrich regions where point occurrence data are limited for many species. We found that although three widely used SDM methods overestimated amphibian species richness in grid cells when compared to expert richness maps in both North and South America due to systematic overestimation of range sizes, diversity gradients were reasonably robust at broad scales. Further, climatic variables statistically explained patterns of richness at similar levels among the different richness sources, although climatic relationships were stronger in the much better known North America than in South America. We conclude that in the face of the high deforestation rates coupled with incomplete data on species distributions, especially in the tropics, SDM represents a useful macroecological tool for investigating broad‐scale richness patterns and the dynamics between species richness and climate.     

rangeMapper: a platform for the study of macroecology of life‐history traits

Aim To introduce rangeMapper, an R package for the study of the macroecological patterns of life‐history traits, and demonstrate its capabilities using three case studies. The first case study addresses an important topic in conservation biology: biodiversity hotspots. Specifically, we investigate the congruence between global hotspots of three parameters that describe avian diversity: species richness, endemic species richness and relative body mass diversity. The second case study investigates a topic of relevance for macroecology: the inter‐specific relationship between range size and body size for avian assemblages, and how it varies geographically. The third case study tackles a methodological problem in macroecology: the influence of map resolution on statistical inference, i.e. the question of whether and how the relationship between species richness and body mass varies with map resolution. Innovation rangeMapper offers a tight integration of spatial and statistical tools for macroecological projects and it relies on a high‐performance database engine which makes it suitable for managing projects using a large number of species. rangeMapper's architecture follows closely the concepts described by Gaston et al. (2008 Journal of Biogeography, 35 , 483–500) and its flexibility allows for both complex data manipulation procedures and easy implementation of new functions. By choosing case studies to cover various technical and conceptual issues we demonstrate rangeMapper's capabilities to address a wide array of questions. Main conclusion rangeMapper ( http://cran.r‐project.org/package=rangeMapper ) is an open source front end software which can be used to address questions in both fundamental ecological research and conservation biology.     

Broad-scale geographical patterns in local stream insect genera richness

Comprehensive global studies of stream invertebrate assemblages are rare and have produced contradictory results. To address this shortcoming, we compiled data from 495 published estimates of local genera richness for three orders of stream‐dwelling insects (Ephemeroptera, Plecoptera, Trichoptera) from throughout the world and used these data to describe global geographic patterns in stream insect genera richness and to address two questions: 1) does local stream insect richness vary more with regional historical factors or with local ecological factors?, and 2) to what extent have streams converged in the number of taxa they support?
Maximum genera richness varied sharply across the range of latitude examined from the south to north poles for all three orders of aquatic insects. Ephemeroptera richness showed 3 peaks (～30°S, 10°N, and 40°N) with highest richness near 5–10°N and 40°N latitude. Plecoptera richness was distinctly highest at ～40°N latitude with a similar peak at 40°S latitude. Trichoptera richness showed less latitudinal variation than the other taxa but was slightly higher near the equator and at 40°N and S latitude than at other latitudes. Genera richness generally declined with increasing elevation, except for Plecoptera. Maximum genera richness increased steadily with a measure of regional terrestrial net primary production and declined sharply with a measure of hydrologic disturbance for all orders. Richness varied widely among both biogeographical realms and biomes, although ca 2 times as much variation in richness was associated with biome as biogeographic realm. Richness for each order was highest in different biogeographic realms, but all orders had highest richness in broadleaf forest biomes. These latter results imply that spatial variation in local richness of stream insects is more strongly affected by contemporary ecological factors than by historical biogeography and that maintenance of intact forested landscapes may be critical to the conservation of stream invertebrate faunas.     

A macroecological perspective of diversity patterns in the freshwater realm

1. The aim of this paper is to review literature on species diversity patterns of freshwater organisms and underlying mechanisms at large spatial scales. 2. Some freshwater taxa (e.g. dragonflies, fish and frogs) follow the classical latitudinal decline in regional species richness (RSR), supporting the patterns found for major terrestrial and marine organism groups. However, the mechanisms causing this cline in most freshwater taxa are inadequately understood, although research on fish suggests that energy and history are major factors underlying the patterns in total species and endemic species richness. Recent research also suggests that not all freshwater taxa comply with the decline of species richness with latitude (e.g. stoneflies, caddisflies and salamanders), but many taxa show more complex geographical patterns in across‐regions analyses. These complexities are even more profound when studies of global, continental and regional extents are compared. For example, clear latitudinal gradients may be present in regional studies but absent in global studies (e.g. macrophytes). 3. Latitudinal gradients are often especially weak in the across‐ecosystems analyses, which may be attributed to local factors overriding the effects of large‐scale factors on local communities. Nevertheless, local species richness (LSR) is typically linearly related to RSR (suggesting regional effects on local diversity), although saturating relationships have also been found in some occasions (suggesting strong local effects on diversity). Nestedness has often been found to be significant in freshwater studies, yet this pattern is highly variable and generally weak, suggesting also a strong beta diversity component in freshwater systems. 4. Both geographical location and local environmental factors contribute to variation in alpha diversity, nestedness and beta diversity in the freshwater realm, although the relative importance of these two groups of explanatory variables may be contingent on the spatial extent of the study. The mechanisms associated with spatial and environmental control of community structure have also been inferred in a number of studies, and most support has been found for species sorting (possibly because many freshwater studies have species sorting as their starting point), although also dispersal limitation and mass effects may be contributing to the patterns found. 5. The lack of latitudinal gradients in some freshwater taxa begs for further explanations. Such explanations may not be gained for most freshwater taxa in the near future, however, because we lack species‐level information, floristic and faunistic knowledge, and standardised surveys along extensive latitudinal gradients. A challenge for macroecology is thus to use the best possible species‐level information on well‐understood groups (e.g. fish) or use surrogates for species‐level patterns (e.g. families) and then develop hypotheses for further testing in the freshwater realm. An additional research challenge concerns understanding patterns and mechanisms associated with the relationships between alpha, beta and gamma components of species diversity. 6. Understanding the mechanistic basis of species diversity patterns should preferably be based on a combination of large‐scale macroecological and landscape‐scale metacommunity research. Such a research approach will help in elucidating patterns of species diversity across regional and local scales in the freshwater realm.     

Diversity of Eastern North American Ant Communities along Environmental Gradients

Studies of species diversity patterns across regional environmental gradients seldom consider the impact of habitat type on within-site (alpha) and between-site (beta) diversity. This study is designed to identify the influence of habitat type across geographic and environmental space, on local patterns of species richness and regional turnover patterns of ant diversity in the northeastern United States. Specifically, I aim to 1) compare local species richness in paired open and forested transects and identify the environmental variables that best correlate with richness; and 2) document patterns of beta diversity throughout the region in both open and forested habitat. I systematically sampled ants at 67 sites from May to August 2010, spanning 10 degrees of latitude, and 1000 meters of elevation. Patterns of alpha and beta diversity across the region and along environmental gradients differed between forested and open habitats. Local species richness was higher in the low elevation and warmest sites and was always higher in open habitat than in forest habitat transects. Richness decreased as temperature decreased or elevation increased. Forested transects show strong patterns of decreasing dissimilarity in species composition between sites along the temperature gradient but open habitat transects did not. Maximum temperature of the warmest month better predicted species richness than either latitude or elevation. I find that using environmental variables as key predictors of richness yields more biologically relevant results, and produces simpler macroecological models than commonly used models which use only latitude and elevation as predictors of richness and diversity patterns. This study contributes to the understanding of mechanisms that structure the communities of important terrestrial arthropods which are likely to be influenced by climatic change.     

The biogeography of tropical reef fishes: endemism and provinciality through time

The largest marine biodiversity hotspot straddles the Indian and Pacific Oceans, driven by taxa associated with tropical coral reefs. Centred on the Indo‐Australian Archipelago (IAA), this biodiversity hotspot forms the ‘bullseye’ of a steep gradient in species richness from this centre to the periphery of the vast Indo‐Pacific region. Complex patterns of endemism, wide‐ranging species and assemblage differences have obscured our understanding of the genesis of this biodiversity pattern and its maintenance across two‐thirds of the world's oceans. But time‐calibrated molecular phylogenies coupled with ancestral biogeographic estimates have provided a valuable framework in which to examine the origins of coral reef fish biodiversity across the tropics. Herein, we examine phylogenetic and biogeographic data for coral reef fishes to highlight temporal patterns of marine endemism and tropical provinciality. The ages and distribution of endemic lineages have often been used to identify areas of species creation and demise in the marine tropics and discriminate among multiple hypotheses regarding the origins of biodiversity in the IAA. Despite a general under‐sampling of endemic fishes in phylogenetic studies, the majority of locations today contain a mixture of potential paleo‐ and neo‐endemic fishes, pointing to multiple historical processes involved in the origin and maintenance of the IAA biodiversity hotspot. Increased precision and sampling of geographic ranges for reef fishes has permitted the division of discrete realms, regions and provinces across the tropics. Yet, such metrics are only beginning to integrate phylogenetic relatedness and ancestral biogeography. Here, we integrate phylogenetic diversity with ancestral biogeographic estimation of lineages to show how assemblage structure and tropical provinciality has changed through time.     

宏生态尺度上景观破碎化对物种丰富度的影响

生物多样性的地理格局及其形成机制是宏生态学与生物地理学的研究热点。大量研究表明,景观尺度上的生境破碎化对物种多样性的分布格局具有重要作用,但目前尚不清楚这种作用是否足以在宏生态尺度上对生物多样性地理格局产生显著影响。利用中国大陆鸟类和哺乳动物的物种分布数据,在100 km×100 km网格的基础上生成了这两个类群生物的物种丰富度地理格局,进一步利用普通最小二乘法模型和空间自回归模型研究了物种丰富度与气候、生境异质性、景观破碎化的相关关系。结果表明,景观破碎化因子与鸟类和哺乳动物的物种丰富度都具有显著的关联关系,其方差贡献率可达约30%—50%(非空间模型)和60%—80%(空间模型),略低于或接近于气候和生境异质性因子。方差分解结果显示,景观破碎化因子与气候和生境异质性因子的方差贡献率的重叠部分达20%—40%。相对鸟类而言,景观破碎化对哺乳动物物种丰富度的地理格局具有更高的解释率。     

Neotropical savanna ants show a reversed latitudinal gradient of species richness,with climatic drivers reflecting the forest origin of the fauna

Aim

To evaluate the extent to which ant species richness in Neotropical savannas varies with macrogeographic variables, and to identify the potential climatic drivers of such variation.

Location

The Cerrado savanna biome of central Brazil, in a region spanning ca. 20° of latitude and 18°of longitude.

Methods

Standardized sampling of the arboreal and ground‐dwelling faunas was performed in 29 well‐preserved savanna sites using pitfall traps. Species were classified according to their habitat affinities: open‐savanna specialists, forest‐associated species or habitat generalists. We used generalized linear models to evaluate the importance of geographic (latitude, longitude and elevation) and climatic (mean temperature and three metrics of rainfall) variables as predictors of species richness.

Results

The total number of species recorded at each site varied more than twofold (from 59 to 144), and latitude was the best geographic correlate of overall species richness. However, contrary to the expected pattern, more species were found at higher than lower latitudes. This reversed latitudinal pattern of diversity occurred for both the arboreal and ground‐dwelling faunas, and for the habitat generalists and forest specialists. The savanna specialists showed a mid‐latitudinal peak in diversity. Overall, there was a significant positive association between rainfall and species richness, but the strength of this relationship varied with ant habitat affinity.

Main conclusions

The Cerrado ant fauna shows a reverse latitudinal gradient in species diversity, and this can be explained by increasing rainfall during the warmest months of the year (and therefore in plant productivity) with increasing latitude. The sensitivity of Cerrado ant diversity to declining rainfall contrasts with the high resilience to aridity of the Australian savanna ant fauna, and this reflects the contrasting evolutionary histories of these faunas. Our findings highlight the importance of historical processes as drivers of intercontinental contrasts in macroecological patterns.     

Towards a panbiogeography of the seas

A contrast is drawn between the concept of speciation favoured in the Darwin–Wallace biogeographic paradigm (founder dispersal from a centre of origin) and in panbiogeography (vicariance or allopatry). Ordinary ecological dispersal is distinguished from founder dispersal. A survey of recent literature indicates that ideas on many aspects of marine biology are converging on a panbiogeographic view. Panbiogeographic conclusions supported in recent work include the following observations: fossils give minimum ages for groups and most taxa are considerably older than their earliest known fossil; Pacific/Atlantic divergence calibrations based on the rise of the Isthmus of Panama at 3 Ma are flawed; for these two reasons most molecular clock calibrations for marine groups are also flawed; the means of dispersal of taxa do not correlate with their actual distributions; populations of marine species may be closed systems because of self‐recruitment; most marine taxa show at least some degree of vicariant differentiation and vicariance is surprisingly common among what were previously assumed to be uniform, widespread taxa; mangrove and seagrass biogeography and migration patterns in marine taxa are best explained by vicariance; the Indian Ocean and the Pacific Ocean represent major biogeographic regions and diversity in the Indo‐Australian Archipelago is related to Indian Ocean/Pacific Ocean vicariance; distribution in the Pacific is not the result of founder dispersal; distribution in the south‐west Pacific is accounted for by accretion tectonics which bring about distribution by accumulation and juxtaposition of communities; tectonic uplift and subsidence can directly affect vertical distribution of marine communities; substantial parallels exist between the biogeography of terrestrial and marine taxa; biogeographically and geologically composite areas are tractable using panbiogeographic analysis; metapopulation models are more realistic than the mainland/island dispersal models used in the equilibrium theory of island biogeography; and regional biogeography is a major determinant of local community composition. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84 , 675–723.     

Multiscale Patterns of Riparian Plant Diversity and Implications for Restoration

Planning riparian restoration to resemble historic reference conditions requires an understanding of both local and regional patterns of plant species diversity. Thus, understanding species distributions at multiple spatial scales is essential to improve restoration planting success, to enhance long‐term ecosystem functioning, and to match restoration planting designs with historic biogeographic distributions. To inform restoration planning, we examined the biogeographic patterns of riparian plant diversity at local and regional scales within a major western U.S.A. drainage, California's Sacramento—San Joaquin Valley. We analyzed patterns of species richness and complementarity (β‐diversity) across two scales: the watershed scale and the floodplain scale. At the watershed scale, spatial patterns of native riparian richness were driven by herbaceous species, whereas woody species were largely cosmopolitan across the nearly 38,000 km² study area. At the floodplain scale, riparian floras reflected species richness and dissimilarity patterns related to hydrological and disturbance‐driven successional sequences. These findings reinforce the importance of concurrently evaluating both local and regional processes that promote species diversity and distribution of native riparian flora. Furthermore, as restoration activities become more prevalent across the landscape, strategies for restoration outcomes should emulate the patterns of species diversity and biogeographic distributions found at regional scales.     

Predictors of mammal species richness in KwaZulu-Natal,South Africa

The management of multi-functional landscapes warrants better knowledge of environment-richness associations at varying disturbance levels and habitat gradients. Intensive land-use patterns for agricultural purposes lead to fragmentation of natural habitat resulting in biodiversity loss that can be measured using landscape metrics to assess mammalian richness. Since carnivores and herbivores are likely to show different responses to disturbance, we calculated carnivore, non-carnivore, and total mammal species richness from camera surveys using a first order Jackknife Estimator. Richness was compared along a habitat gradient comprising coastal forest, Acacia thicket, and highland in KwaZulu-Natal, South Africa. We used standardized OLS regression models to identify climatic and disturbance variables, and landscape metrics as predictors of species richness. The estimated total and non-carnivore species richness were highest in coastal forest, while carnivore species richness was highest in highland followed by coastal forest and Acacia thicket. Average monthly maximum temperature was a significant predictor of all richness groups, and precipitation of the wettest month and isothermality determined total and non-carnivore species richness, respectively. These climatic variables possibly limit species distribution because of physiological tolerance of the species. Total mammal richness was determined by mean shape (+) and habitat division (−) while diversity (+) and patch richness (−) explained carnivore species richness. Mean shape index (+) influenced non-carnivore richness. However, habitat division and patch richness negatively influenced total mammal richness. Though habitat patch size and contiguity had a weak positive prediction, these metrics demonstrated the importance of habitat connectivity for maintaining mammal richness. The identification of these climatic and landscape patterns is important to facilitate future landscape management for mammal conservation in forest-mosaics.     

What's on the horizon for macroecology?

Over the last two decades, macroecology – the analysis of large‐scale, multi‐species ecological patterns and processes – has established itself as a major line of biological research. Analyses of statistical links between environmental variables and biotic responses have long and successfully been employed as a main approach, but new developments are due to be utilized. Scanning the horizon of macroecology, we identified four challenges that will probably play a major role in the future. We support our claims by examples and bibliographic analyses. 1) Integrating the past into macroecological analyses, e.g. by using paleontological or phylogenetic information or by applying methods from historical biogeography, will sharpen our understanding of the underlying reasons for contemporary patterns. 2) Explicit consideration of the local processes that lead to the observed larger‐scale patterns is necessary to understand the fine‐grain variability found in nature, and will enable better prediction of future patterns (e.g. under environmental change conditions). 3) Macroecology is dependent on large‐scale, high quality data from a broad spectrum of taxa and regions. More available data sources need to be tapped and new, small‐grain large‐extent data need to be collected. 4) Although macroecology already lead to mainstreaming cutting‐edge statistical analysis techniques, we find that more sophisticated methods are needed to account for the biases inherent to sampling at large scale. Bayesian methods may be particularly suitable to address these challenges. To continue the vigorous development of the macroecological research agenda, it is time to address these challenges and to avoid becoming too complacent with current achievements.     

High species diversity and turnover in granite inselberg floras highlight the need for a conservation strategy protecting many outcrops

Determining patterns of plant diversity on granite inselbergs is an important task for conservation biogeography due to mounting threats. However, beyond the tropics there are relatively few quantitative studies of floristic diversity, or consideration of these patterns and their environmental, biogeographic, and historical correlates for conservation. We sought to contribute broader understanding of global patterns of species diversity on granite inselbergs and inform biodiversity conservation in the globally significant Southwest Australian Floristic Region (SWAFR). We surveyed floristics from 16 inselbergs (478 plots) across the climate gradient of the SWAFR stratified into three major habitats on each outcrop. We recorded 1,060 species from 92 families. At the plot level, local soil and topographic variables affecting aridity were correlated with species richness in herbaceous (HO) and woody vegetation (WO) of soil‐filled depressions, but not in woody vegetation on deeper soils at the base of outcrops (WOB). At the outcrop level, bioclimatic variables affecting aridity were correlated with species richness in two habitats (WO and WOB) but, contrary to predictions from island biogeography, were not correlated with inselberg area and isolation in any of the three habitats. Species turnover in each of the three habitats was also influenced by aridity, being correlated with bioclimatic variables and with interplot geographic distance, and for HO and WO habitats with local site variables. At the outcrop level, species replacement was the dominant component of species turnover in each of the three habitats, consistent with expectations for long‐term stable landscapes. Our results therefore highlight high species diversity and turnover associated with granite outcrop flora. Hence, effective conservation strategies will need to focus on protecting multiple inselbergs across the entire climate gradient of the region.