Post-Eocene climate change, niche conservatism, and the latitudinal diversity gradient of New World birds

Aim The aim of this study was to test a variant of the evolutionary time hypothesis for the bird latitudinal diversity gradient derived from the effects of niche conservatism in the face of global climate change over evolutionary time. Location The Western Hemisphere. Methods We used digitized range maps of breeding birds to estimate the species richness at two grain sizes, 756 and 12,100 km². We then used molecular phylogenies resolved to family to quantify the root distance (RD) of each species as a measure of its level of evolutionary development. Birds were classified as ‘basal’ or ‘derived’ based on the RD of their family, and richness patterns were contrasted for the most basal and most derived 30% of species. We also generated temperature estimates for the Palaeogene across the Western Hemisphere to examine how spatial covariation between past and present climates might make it difficult to distinguish between ecological and evolutionary hypotheses for the current richness gradient. Results The warm, wet tropics support many species from basal bird clades, whereas the northern temperate zone and cool or dry tropics are dominated by species from more recent, evolutionarily derived clades. Furthermore, crucial to evaluating how niche conservatism among birds may drive the hemispherical richness gradient, the spatial structure of the richness gradient for basal groups is statistically indistinguishable from the overall gradient, whereas the richness gradient for derived groups is much shallower than the overall gradient. Finally, modern temperatures and the pattern of climate cooling since the Eocene are indistinguishable as predictors of bird species richness. Main conclusions Differences in the richness gradients of basal vs. derived clades suggest that the hemispherical gradient has been strongly influenced by the differential extirpation of species in older, warm‐adapted clades from parts of the world that have become cooler in the present. We propose that niche conservatism and global‐scale climate change over evolutionary time provide a parsimonious explanation for the contemporary bird latitudinal diversity gradient in the New World, although dispersal limitation of some highly derived clades probably plays a secondary role.     

Different evolutionary histories underlie congruent species richness gradients of birds and mammals

Aim The global species richness patterns of birds and mammals are strongly congruent. This could reflect similar evolutionary responses to the Earth’s history, shared responses to current climatic conditions, or both. We compare the geographical and phylogenetic structures of both richness gradients to evaluate these possibilities. Location Global. Methods Gridded bird and mammal distribution databases were used to compare their species richness gradients with the current environment. Phylogenetic trees (resolved to family for birds and to species for mammals) were used to examine underlying phylogenetic structures. Our first prediction is that both groups have responded to the same climatic gradients. Our phylogenetic predictions include: (1) that both groups have similar geographical patterns of mean root distance, a measure of the level of the evolutionary development of faunas, and, more directly, (2) that richness patterns of basal and derived clades will differ, with richness peaking in the tropics for basal clades and in the extra‐tropics for derived clades, and that this difference will hold for both birds and mammals. We also explore whether alternative taxonomic treatments for mammals can generate patterns matching those of birds. Results Both richness gradients are associated with the same current environmental gradients. In contrast, neither of our evolutionary predictions is met: the gradients have different phylogenetic structures, and the richness of birds in the lowland tropics is dominated by many basal species from many basal groups, whereas mammal richness is attributable to many species from both few basal groups and many derived groups. Phylogenetic incongruence is robust to taxonomic delineations for mammals. Main conclusions Contemporary climate can force multiple groups into similar diversity patterns even when evolutionary trajectories differ. Thus, as widely appreciated, our understanding of biodiversity must consider responses to both past and present climates, and our results are consistent with predictions that future climate change will cause major, correlated changes in patterns of diversity across multiple groups irrespective of their evolutionary histories.     

Tropical niche conservatism and the species richness gradient of North American butterflies

Aim We explore the potential role of the ‘tropical conservatism hypothesis’ in explaining the butterfly species richness gradient in North America. Its applicability can be derived from the tropical origin of butterflies and the presumed difficulties in evolving the cold tolerance required to permit the colonization and permanent occupation of the temperate zone. Location North America. Methods Digitized range maps for butterfly species north of Mexico were used to map richness for all species, species with distributions north of the Tropic of Capricorn (Extratropicals), and species that also occupy the tropics (Tropicals). A phylogeny resolved to subfamily was used to map the geographical pattern of mean root distance, a metric of the evolutionary development of assemblages. Regression models and general linear models examined environmental correlates of overall richness and for Extratropicals vs. Tropicals, patterns in summer vs. winter, and patterns in northern vs. southern North America. Results Species in more basal subfamilies dominate the south, whereas more derived clades occupy the north. There is also a ‘latitudinal’ richness gradient in Canada/Alaska, whereas in the conterminous USA richness primarily varies longitudinally. Overall richness is associated with broad‐ and mesoscale temperature gradients. The richness of Tropicals is strongly associated with temperature and distance from winter population sources. The richness of Extratropicals in the north is most strongly correlated with the pattern of glacial retreat since the more recent Ice Age, whereas in the south, richness is positively associated with the range of temperatures in mountains and the presence of forests but is negatively correlated with the broad‐scale temperature gradient. Main conclusions The tropical conservatism hypothesis provides a possible explanation for the complex structure of the species richness gradient. The Canada/Alaska fauna comprises temperate, boreal and tundra species that are nevertheless constrained by cold climates and limited vegetation, coupled with possible post‐Pleistocene recolonization lags. In the USA tropical species are constrained by temperature in winter as well as recolonization distances in summer, whereas temperate‐zone groups are richer in cooler climates in mountains and forests, where winter conditions are more suitable for diapause. The evolution of cold tolerance is key to both the evolutionary and ecological patterns.     

Water links the historical and contemporary components of the Australian bird diversity gradient

Aim To document the geographical structure of the historical signal in the continental species richness gradient of birds and evaluate the influences of contemporary and historical climatic conditions on the generation and maintenance of the richness pattern. Location Australia. Methods We used range maps of breeding birds to generate the spatial pattern of species richness at four grain sizes, and two molecular phylogenies to measure the level of evolutionary development of avifaunas at each grain size. We then used simple correlation and path analysis to generate a statistical model of species richness using environmental predictor variables and compared the spatial patterns of richness and mean evolutionary development to identify possible environmental links between richness and net diversification rates across the continent. Results The contemporary richness pattern is well explained statistically by actual evapotranspiration (a measure of water–energy balance), operating both directly and indirectly through plant production, and this is robust to the spatial resolution of the analysis. Further, species richness and the mean level of evolutionary development of faunas show a strong spatial correspondence, such that dry areas support both fewer species and species from more highly derived families, whereas wet areas support more species of both basal and derived families. The evolutionary pattern conforms to a similar pattern known for plants and is probably explained by the increase in aridity in western and central Australia arising in the Miocene. Main conclusion The contemporary bird richness gradient contains a historical signal and reflects the effects of both current levels of water availability as well as changes in rainfall patterns extending over evolutionary time. The historical signal persists even in the absence of obvious hard barriers to dispersal.     

Global patterns of diversification and species richness in amphibians

Geographic patterns of species richness ultimately arise through the processes of speciation, extinction, and dispersal, but relatively few studies consider evolutionary and biogeographic processes in explaining these diversity patterns. One explanation for high tropical species richness is that many species-rich clades originated in tropical regions and spread to temperate regions infrequently and more recently, leaving little time for species richness to accumulate there (assuming similar rates of diversification in temperate and tropical regions). However, the major clades of anurans (frogs) and salamanders may offer a compelling counterexample. Most salamander families are predominately temperate in distribution, but the one primarily tropical clade (Bolitoglossinae) contains nearly half of all salamander species. Similarly, most basal clades of anurans are predominately temperate, but one largely tropical clade (Neobatrachia) contains approximately 96% of anurans. In this article, I examine patterns of diversification in frogs and salamanders and their relationship to large-scale patterns of species richness in amphibians. I find that diversification rates in both frogs and salamanders increase significantly with decreasing latitude. These results may shed light on both the evolutionary causes of the latitudinal diversity gradient and the dramatic but poorly explained disparities in the diversity of living amphibian clades.     

Large-scale phylogenetic analyses reveal the causes of high tropical amphibian diversity

Many groups show higher species richness in tropical regions but the underlying causes remain unclear. Despite many competing hypotheses to explain latitudinal diversity gradients, only three processes can directly change species richness across regions: speciation, extinction and dispersal. These processes can be addressed most powerfully using large-scale phylogenetic approaches, but most previous studies have focused on small groups and recent time scales, or did not separate speciation and extinction rates. We investigate the origins of high tropical diversity in amphibians, applying new phylogenetic comparative methods to a tree of 2871 species. Our results show that high tropical diversity is explained by higher speciation in the tropics, higher extinction in temperate regions and limited dispersal out of the tropics compared with colonization of the tropics from temperate regions. These patterns are strongly associated with climate-related variables such as temperature, precipitation and ecosystem energy. Results from models of diversity dependence in speciation rate suggest that temperate clades may have lower carrying capacities and may be more saturated (closer to carrying capacity) than tropical clades. Furthermore, we estimate strikingly low tropical extinction rates over geological time scales, in stark contrast to the dramatic losses of diversity occurring in tropical regions presently.     

Latitudinal Diversity Gradients in New World Bats: Are They a Consequence of Niche Conservatism?

The increase in species diversity from the Poles to the Equator is a major biogeographic pattern, but the mechanisms underlying it remain obscure. Our aim is to contribute to their clarification by describing the latitudinal gradients in species richness and in evolutionary age of species of New World bats, and testing if those patterns may be explained by the niche conservatism hypothesis. Maps of species ranges were used to estimate species richness in a 100 x 100 km grid. Root distances in a molecular phylogeny were used as a proxy for the age of species, and the mean root distance of the species in each cell of the grid was estimated. Generalised additive models were used to relate latitude with both species richness and mean root distance. This was done for each of the three most specious bat families and for all Chiroptera combined. Species richness increases towards the Equator in the whole of the Chiroptera and in the Phyllostomidae and Molossidae, families that radiated in the tropics, but the opposite trend is observed in the Vespertilionidae, which has a presumed temperate origin. In the whole of the Chiroptera, and in the three main families, there were more basal species in the higher latitudes, and more derived species in tropical areas. In general, our results were not consistent with the predictions of niche conservatism. Tropical niche conservatism seems to keep bat clades of tropical origin from colonizing temperate zones, as they lack adaptations to survive cold winters, such as the capacity to hibernate. However, the lower diversity of Vespertilionidae in the Neotropics is better explained by competition with a diverse pre-existing community of bats than by niche conservatism.     

Historical processes enhance patterns of diversity along latitudinal gradients

One of the more vexing issues in ecology is how historical processes affect contemporary patterns of biodiversity. Accordingly, few models have been presented. Two corollary models (centre of origin, time-for-speciation) can be used to make quantitative predictions characterizing the tropical niche conservatism hypothesis and describe diversification as diffusion and subsequent cladogenesis of species away from the place of origin of a higher taxon in the tropics. Predictions derived from such models are: (i) species richness declines toward the periphery of the range of a higher taxon; (ii) taxa are more derived toward the periphery than the centre; (iii) ages of taxa are lower toward the periphery than the centre; and (iv) ages and measures of derivedness are less variable toward the periphery of the range of a higher taxon. I tested these predictions to better understand the formation of one of the most ubiquitous patterns of biodiversity-the latitudinal gradient in species richness. Results indicate well-supported predictions for New World leaf-nosed bats and that diversification has had strong influences on latitudinal gradients of species richness. A better understanding of how evolutionary diversification of taxa contributes to formation of patterns of species richness along environmental gradients is necessary to fully understand spatial variation in biodiversity.     

Evolutionary and ecological causes of the latitudinal diversity gradient in hylid frogs: treefrog trees unearth the roots of high tropical diversity

Why are there more species in the tropics than in temperate regions? In recent years, this long-standing question has been addressed primarily by seeking environmental correlates of diversity. But to understand the ultimate causes of diversity patterns, we must also examine the evolutionary and biogeographic processes that directly change species numbers (i.e., speciation, extinction, and dispersal). With this perspective, we dissect the latitudinal diversity gradient in hylid frogs. We reconstruct a phylogeny for 124 hylid species, estimate divergence times and diversification rates for major clades, reconstruct biogeographic changes, and use ecological niche modeling to identify climatic variables that potentially limit dispersal. We find that hylids originated in tropical South America and spread to temperate regions only recently (leaving limited time for speciation). There is a strong relationship between the species richness of each region and when that region was colonized but not between the latitudinal positions of clades and their rates of diversification. Temperature seasonality seemingly limits dispersal of many tropical clades into temperate regions and shows significant phylogenetic conservatism. Overall, our study illustrates how two general principles (niche conservatism and the time-for-speciation effect) may help explain the latitudinal diversity gradient as well as many other diversity patterns across taxa and regions.     

The causes of species richness patterns across space, time, and clades and the role of "ecological limits"

A major goal of research in ecology and evolution is to explain why species richness varies across habitats, regions, and clades. Recent reviews have argued that species richness patterns among regions and clades may be explained by "ecological limits" on diversity over time, which are said to offer an alternative explanation to those invoking speciation and extinction (diversification) and time. Further, it has been proposed that this hypothesis is best supported by failure to find a positive relationship between time (e.g., clade age) and species richness. Here, I critically review the evidence for these claims, and propose how we might better study the ecological and evolutionary origins of species richness patterns. In fact, ecological limits can only influence species richness in clades by influencing speciation and extinction, and so this new "alternative paradigm" is simply one facet of the traditional idea that ecology influences diversification. The only direct evidence for strict ecological limits on richness (i.e., constant diversity over time) is from the fossil record, but many studies cited as supporting this pattern do not, and there is evidence for increasing richness over time. Negative evidence for a relationship between clade age and richness among extant clades is not positive evidence for constant diversity over time, and many recent analyses finding no age-diversity relationship were biased to reach this conclusion. More comprehensive analyses strongly support a positive age-richness relationship. There is abundant evidence that both time and ecological influences on diversification rates are important drivers of both large-scale and small-scale species richness patterns. The major challenge for future studies is to understand the ecological and evolutionary mechanisms underpinning the relationships between time, dispersal, diversification, and species richness patterns.     

Determinants of species richness, endemism and turnover in European longhorn beetles

This study assessed the diversity patterns of a large family of beetles, Cerambycidae, in Europe and tested the following hypotheses: 1) richness gradients of this hyperdiverse taxon are driven by water and energy variables; 2) endemism is explained by the same factors, but variation between areas also reflects post‐glacial re‐colonization processes; and 3) faunal composition is determined by the same climatic variables and, therefore, beta diversity (species turnover) is related to richness gradients. Species richness, endemism and beta diversity were modelled using inventories of 37 European territories, built from a database containing the distributions of 609 species. Area, spatial position, and nine topographical and climatic variables were used as predictors in regression and constrained analysis of principal coordinates modelling. Species richness was mostly explained by a temperature gradient, which produced a south‐to‐north decreasing richness gradient. Endemism followed the same pattern, but was also determined by longitudinal variation, peaking in the southwestern and southeastern corners of the continent. Faunal turnover was explained by an important purely spatial pattern and a spatially structured environmental gradient. Thus, contrary to other groups, cerambycid richness was mostly explained by environmental energy, but not by water availability. Endemism was concentrated in the Iberian and Greek peninsulas, but not in Italy. Thus, the latter area may have been the major source of post‐glacial re‐colonization for European longhorn beetles or, otherwise, a poor refuge during glaciations. Turnover patterns were independent of the richness gradient, because northern faunas are nested in southern ones. Turnover, in contrast to richness, was driven by both the independent effects of climate and geographic constraints that might reflect dispersal limitation or stochastic colonization events, suggesting that richness gradients are more environmentally deterministic phenomena than turnover patterns.     

THE ROLE OF CLIMATIC TOLERANCES AND SEED TRAITS IN REDUCED EXTINCTION RATES OF TEMPERATE POLYGONACEAE

The latitudinal diversity gradient (LDG) is one of the most striking and consistent biodiversity patterns across taxonomic groups. We investigate the species richness gradient in the buckwheat family, Polygonaceae, which exhibits a reverse LDG and is, thus, decoupled from dominant gradients of energy and environmental stability that increase toward the tropics and confound mechanistic interpretations. We test competing age and evolutionary diversification hypotheses, which may explain the diversification of this plant family over the past 70 million years. Our analyses show that the age hypothesis, which posits that clade richness is positively correlated with the ecological and evolutionary time since clade origin, fails to explain the richness gradient observed in Polygonaceae. However, an evolutionary diversification hypothesis is highly supported, with diversification rates being 3.5 times higher in temperate clades compared to tropical clades. We demonstrate that differences in rates of speciation, migration, and molecular evolution insufficiently explain the observed patterns of differential diversification rates. We suggest that reduced extinction rates in temperate clades may be associated with adaptive responses to selection, through which seed morphology and climatic tolerances potentially act to minimize risk in temporally variable environments. Further study is needed to understand causal pathways among these traits and factors correlated with latitude.     

Variation in the composition and structure of tropical savannas as a function of rainfall and soil texture along a large-scale climatic gradient in the Northern Territory, Australia

Abstract. Variation in structural and compositional attributes of tropical savannas are described in relation to variation in annual rainfall and soil texture along a subcontinental-scale gradient of rainfall in the wet-dry tropics of the Northern Territory, Australia. Rainfall varies along the gradient from over 1500 mm p.a. in the Darwin region ( c . 12° S) to less than 500 mm in the Tennant Creek region ( c . 18° S). Soils are patchy, and sands, loams and clays may occur in all major districts within the region. We utilized a large data set (1657 quadrats ° 291 woody species; with numerous measured and derived sample variables) covering an area of 0.5 million km². Correlations between floristic composition of woody species and environmental variables were assessed using DCA ordination and vector fitting of environmental variables. Vectors of annual rainfall and soil texture were highly correlated with variation in species composition. Multiple regression analyses incorporating linear and quadratic components of mean annual rainfall and topsoil clay content were performed on three structural attributes (tree height, tree cover, tree basal area) and two compositional attributes (woody species richness, deciduous tree species richness). Tree height declined with decreasing rainfall; cover, basal area, woody species richness and deciduous species richness all declined with decreasing rainfall and increasing soil clay content. Regression models accounted for between 17% and 45% of the variation in the data sets. Variation in other factors such as soil depth, landscape position and recent land-use practices (for which there were no data on an individual quadrat basis) are likely to have contributed to the large residual variation in the data set.     

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.     

A PHYLOGENETIC PERSPECTIVE ON ELEVATIONAL SPECIES RICHNESS PATTERNS IN MIDDLE AMERICAN TREEFROGS: WHY SO FEW SPECIES IN LOWLAND TROPICAL RAINFORESTS?

Differences in species richness at different elevations are widespread and important for conservation, but the causes of these patterns remain poorly understood. Here, we use a phylogenetic perspective to address the evolutionary and biogeographic processes that underlie elevational diversity patterns within a region. We focus on a diverse but well-studied fauna of tropical amphibians, the hylid frogs of Middle America. Middle American treefrogs show a "hump-shaped" pattern of species richness (common in many organisms and regions), with the highest regional diversity at intermediate elevations. We reconstructed phylogenetic relationships among 138 species by combining new and published sequence data from 10 genes and then used this phylogeny to infer evolutionary rates and patterns. The high species richness of intermediate elevations seems to result from two factors. First, a tendency for montane clades to have higher rates of diversification. Second, the early colonization of montane regions, leaving less time for speciation to build up species richness in lowland regions (including tropical rainforests) that have been colonized more recently. This "time-for-speciation" effect may explain many diversity patterns and has important implications for conservation. The results also imply that local-scale environmental factors alone may be insufficient to explain the high species richness of lowland tropical rainforests, and that diversification rates are lower in earth's most species-rich biome.     

Environmental harshness,latitude and incipient speciation

Are rates of evolution and speciation fastest where diversity is greatest – the tropics? A commonly accepted theory links the latitudinal diversity gradient to a speciation pump model whereby the tropics produce species at a faster rate than extra‐tropical regions. In this issue of Molecular Ecology, Botero et al. ( 2014 ) test the speciation pump model using subspecies richness patterns for more than 9000 species of birds and mammals as a proxy for incipient speciation opportunity. Rather than using latitudinal centroids, the authors investigate the role of various environmental correlates of latitude as drivers of subspecies richness. Their key finding points to environmental harshness as a positive predictor of subspecies richness. The authors link high subspecies richness in environmental harsh areas to increased opportunities for geographic range fragmentation and/or faster rates of trait evolution as drivers of incipient speciation. Because environmental harshness generally increases with latitude, these results suggest that opportunity for incipient speciation is lowest where species richness is highest. The authors interpret this finding as incompatible with the view of the tropics as a cradle of diversity. Their results are consistent with a growing body of evidence that reproductive isolation and speciation occur fastest at high latitudes.     

Multiregional comparison of the ecological and phylogenetic structure of butterfly species richness gradients

Aim To examine butterfly species richness gradients in seven regions/countries and to quantify geographic mean root distance (MRD) patterns. My primary goal is to determine the extent to which an explanation for butterfly richness patterns based on tropical niche conservatism and the evolution of cold tolerance, proposed for the fauna of Canada and the USA, applies to other parts of the world. Location USA/Canada, Mexico, Europe/NW Africa, Transbaikal Siberia, Chile, South Africa and Australia. Methods Digitized range maps for butterfly species in each region were used to map richness patterns in summer (for all areas) and winter (for USA/Canada, Europe/NW Africa and Australia). A phylogeny resolved to subfamily was used to map the geographic MRD patterns. Regression trees and general linear models examined climatic and vegetation correlates of species richness and MRD within and among regions. Results Various combinations of climate and vegetation were strong predictors of species richness gradients within regions, but unresolved ‘regional’ factors contributed to the multiregional pattern. Regionally based differences in phylogenetic structure also exist, but MRD is negatively correlated with temperature both within and across areas. MRD patterns consistent with tropical niche conservatism occur in most areas. With a possible partial exception of Mexico, faunas in cold climates and in mountains are more derived than faunas in lowlands and tropical/subtropical climates. In USA/Canada, Europe and Australia, winter faunas are more derived than summer faunas. Main conclusions The phylogenetic pattern previously found in the USA and Canada is widespread in both the Northern and Southern Hemispheres, and niche conservatism and the evolution of cold tolerance is the likely explanation for the development of the global butterfly species richness gradient over evolutionary time. Contemporary climate also influences species richness patterns but is unlikely to be a complete explanation globally. The importance of climate is also manifested in the seasonal loss of more basal butterfly elements outside the tropics in winter.     

The Southern African orchid flora: composition, sources and endemism

Aim The Southern African orchid flora is taxonomically well known, but the biogeographical and diversity patterns have not yet been analysed. In particular, we want to establish whether (a) it is, like the Southern African flora in general, more diverse than would be expected from its latitude and area; (b) it is an African flora, or whether it contains palaeoendemic relicts of a Gondwanan orchid flora; (c) the diversity and endemism in the orchid flora is concentrated in particular biomes and habitat types; and (d) the patterns of endemism in the flora can be accounted for by current environmental parameters, or whether we need to invoke historical explanations. Location Southern Africa. Methods We used the recent floristic account of the Southern African orchids, in conjunction with a data base of over 14,642 herbarium records, to assign the species and subspecies of Southern African orchids to biomes, habitats, and clades. We explored the relationship between the number and endemism of entities (species, subspecies and varieties) and the biomes and habitats. We compared the richness of this flora with that of 31 other regions from all continents and latitudes, to establish whether the Southern African orchid flora is richer or poorer than expected. We assigned the Southern African orchid species to 16 monophyletic clades and mapped the global distribution of these clades to establish the continental affinities of the flora. Main conclusions The Southern African orchid flora is not any more diverse than could be expected from its latitude or area, while the two tropical African floras included were less diverse than expected. Latitude is an excellent predictor of regional orchid species richness; this might indicate that available habitat is more important for orchid diversity than gross area available, since latitude is probably correlated with the extent of suitable habitat. The Southern African orchid flora is clearly an African flora, since all clades are also found in tropical Africa, while many of them are absent from the Americas or Asia. Conversely, while most African orchid clades are also found in Southern Africa, both the Americas and Asia contain many clades absent from Africa. The distribution of orchid entities among the biomes in Southern Africa is very uneven, with two of the seven biomes totally devoid of orchids. Habitats and biomes that have no equivalent in tropical Africa are high in endemism, and habitats and biomes which are also well developed in tropical Africa are low in endemism. Endemism appears largely explained in terms of modern habitats. However, two patterns (the high endemism in the Succulent Karoo and the lack of endemism in the southern Cape among epiphytic orchids) may also be explained in terms of Quaternary climatic changes.     

Older species: a rejuvenation on coral reefs?

Aim To discuss the theory that the present high species diversity and apomorphic character of the coral reef ecosystem is because of the historic accumulation of basal species from marginal habitats. Location The Indo‐West Pacific Ocean. Methods The examination of biogeographical patterns from the standpoint of paleontology, phylogeny, genetics, and empirical data. Results Fossil patterns from several clades indicate a gradient of increasing average generic age that extends outward from the high diversity reefs. Successful species that give rise to new species, genera, and families apparently originate from high diversity locations. The tropics have been a major source of evolutionary novelty, not simply a refuge that has accumulated diversity. Many plesiomorphic clades, that once dominated the shallow tropics, are now limited to the deep sea and other safe places. Recent research on several tropical fish families indicates that more apomorphic species inhabit the high diversity reefs. Genetic studies suggest that a decrease in genetic variation extends from the diversity centre toward the outer reaches of the Indo‐West Pacific. Empirical data show that it is extremely difficult for species from low diversity areas to invade places of higher diversity. Main conclusions There is no convincing evidence to indicate that basal species from marginal habitats have been able to accumulate on the coral reefs. Once such species have been displaced from a high diversity environment, there is apparently no return. The evolutionary innovations that contribute to the origination of new phyletic lines take place under conditions of high diversity and maximum competition.     

Extinction and time help drive the marine‐terrestrial biodiversity gradient: is the ocean a deathtrap?

The marine‐terrestrial richness gradient is among Earth's most dramatic biodiversity patterns, but its causes remain poorly understood. Here, we analyse detailed phylogenies of amniote clades, paleontological data and simulations to reveal the mechanisms underlying low marine richness, emphasising speciation, extinction and colonisation. We show that differences in diversification rates (speciation minus extinction) between habitats are often weak and inconsistent with observed richness patterns. Instead, the richness gradient is explained by limited time for speciation in marine habitats, since all extant marine clades are relatively young. Paleontological data show that older marine invasions have consistently ended in extinction. Simulations show that marine extinctions help drive the pattern of young, depauperate marine clades. This role for extinction is not discernible from molecular phylogenies alone, and not predicted by most previously hypothesised explanations for this gradient. Our results have important implications for the marine‐terrestrial biodiversity gradient, and studies of biodiversity gradients in general.