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.     

Untangling latitudinal richness gradients at higher taxonomic levels: familial perspectives on the diversity of New World bat communities

Aims (i) To describe at the level of local communities latitudinal gradients in the species richness of different families of New World bats and to explore the generality of such gradients. (ii) To characterize the relative effects of changes in the richness of each family to the richness of entire communities. (iii) To determine differences in the rate and direction of latitudinal gradients in species richness within families. (iv) To evaluate how differences among families regarding latitudinal gradients in species richness influence the latitudinal gradient in species richness of entire communities. Location Continental New World ranging from the northern continental United States (Iowa, 42° N) to eastern Paraguay (Canindeyú, 24° S). Methods Data on the species composition of communities came from 32 intensively sampled sites. Analyses focused on species richness of five of nine New World bat families. Multivariate analysis of variance and discriminant function analysis determined and described differences among temperate, subtropical, and tropical climatic zones regarding the species richness of bat families. Simple linear regression described latitudinal gradients in species richness of families. Path analysis was used to describe: (i) the direct effect of latitude on species richness of communities, (ii) the indirect effects of latitude on the species richness of communities through its effect on the species richness of each family, (iii) the relative effects of latitude on the species richness of bat families, and (iv) the relative contribution of each family to variation in the species richness of communities. Results Highly significant differences among climatic zones existed primarily because of a difference between the temperate zone and the tropical and subtropical zones combined. This difference was associated with the high number of vespertilionids in the temperate zone and the high number of phyllostomids in the tropical and subtropical zones. Latitudinal gradients in species richness were contingent on phylogeny. Although only three of the five families exhibited significant gradients, all families except for the Vespertilionidae exhibited indistinguishable increases in species richness with decreases in latitude. The Emballonuridae, Phyllostomidae and Vespertilionidae exhibited significant latitudinal gradients whereby the former two families exhibited the classical increase in species richness with decreasing latitude and the latter family exhibited the opposite pattern. Variation in species richness of all families contributed significantly to variation in the species richness of entire communities. Nonetheless, the Phyllostomidae made a significantly stronger contribution to changes in species richness of communities than did all other families. Much of the latitudinal gradient in species richness of communities could be accounted for by the effects of latitude on the species richness of constituent families. Main conclusions Ecological and evolutionary differences among higher taxonomic units, particularly those differences involving life‐history traits, predispose taxa to exhibit different patterns of diversity along environmental gradients. This may be particularly true along extensive gradients such as latitude. Nonetheless, species rich taxa, by virtue of their greater absolute rates of change, can dominate and therefore define the pattern of diversity at a higher taxonomic level and eclipse differences among less represented taxa in their response to environmental gradients. This is true not only with respect to how bats drive the latitudinal gradient in species richness for all mammals, but also for how the Phyllostomidae drives the latitudinal gradient for all bats in the New World. Better understanding of the mechanistic basis of latitudinal gradients of diversity may come from comparing and contrasting patterns across lower taxonomic levels of a higher taxon and by identifying key ecological and evolutionary traits that are associated with such differences.     

Evolutionary rates do not drive latitudinal diversity gradients

Among the several hypotheses invoked for explaining latitudinal diversity gradients (LDGs), some models classified within the 'evolutionary hypothesis' family assume that LDGs are the direct consequence of latitudinal variations in the speciation and/or extinctions rates. Spatially structured simulations of the biogeographical dispersal of a randomly generated clade refute the central tenet of these explanatory models and indicate that global diversity patterns are combined outcomes of geographic and thermal mid-domain effects under a phylogenetically controlled niche conservatism constraint. The positive correlation observed in several higher taxa between speciation rate and diversity does not involve any causal relationship between these two parameters but is most likely the first order by-product of a positive correlation between temperature and per capita speciation rate.     

A test of Metabolic Theory as the mechanism underlying broad-scale species-richness gradients

Aim To test whether the temperature dependence of individuals’ metabolic rates is the mechanism shaping broad‐scale species‐richness gradients as proposed in the Metabolic Theory of Ecology recently proposed by Allen, Gillooly and Brown. Location North America, north of Mexico. Methods Metabolic Theory predicts that the natural logarithm of species richness will be a linear function of environmental temperature ((kT)⁻¹, where k is Boltzmann's constant and T is temperature in K) with a slope of −0.78. We tested these predictions using the broad‐scale variation in richness of amphibians, reptiles, trees, tiger beetles, butterflies and blister beetles. We tested whether the temperature–richness relationship was linear or curvilinear, and determined the range of temperature values (and geographical area) where the instantaneous slope of the curvilinear temperature–richness relationship was statistically indistinguishable from −0.78, after correcting for spatial autocorrelation. Results We found that for all taxa, temperature–richness relationships were curvilinear. Moreover, for five of six taxa, the slope of this relationship was close to the predicted value for only a narrow range of temperatures. Blister beetles displayed the widest temperature range that is consistent with the Metabolic Theory, covering 45% of the study's geographical area. For the remaining taxa, the geographical range in which the slope is consistent with the predicted value amounts to only 10–20% of North America. Main conclusions For a wide array of taxa in North America, temperature–richness relationships deviate from the pattern predicted by Metabolic Theory. These results demonstrate that the temperature dependence of individuals’ metabolic rates is not the sole cause of broad‐scale diversity gradients. Even in areas where factors other than temperature do not influence productivity, the data do not suggest that richness patterns are determined by the temperature dependence of metabolic rate.     

Body size and latitudinal gradients in regional diversity of New World birds

Are latitudinal gradients in regional diversity random or biased with respect to body size? Using data for the New World avifauna, I show that the slope of the increase in regional species richness from the Arctic to the equator is not independent of body size. The increase is steepest among small and medium‐sized species, and shallowest among the largest species. This is reflected in latitudinal variation in the shape of frequency distributions of body sizes in regional subsets of the New World avifauna. Because species are added disproportionately in small and medium size classes towards low latitudes, distributions become less widely spread along the body size axis than expected from the number of species. These patterns suggest an interaction between the effects of latitude and body size on species richness, implying that mechanisms which vary with both latitude and body size may be important determinants of high tropical diversity in New World birds.     

A latitudinal gradient in large-scale beta diversity for vascular plants in North America

Species turnover, or beta diversity, has been predicted to decrease with increasing latitude, but few studies have tested this relationship. Here, we examined the beta diversity–latitude relationship for vascular plants at a continental scale, based on complete species lists of native vascular plants for entire states or provinces in North America (north of Mexico). We calculated beta diversity as the slope of the relationship between the natural logarithm of the Jaccard index (ln J ) for families, genera or species, and both geographic distance and climate difference within five latitude zones. We found that beta diversity decreased from south to north; within latitude zones, it decreased from species to genera and families. Geographic and climatic distance explained about the same proportion of the variance in ln J in zones south of c. 50°N. North of this latitude, nearly all the explained variance in ln J was attributable to geographic distance. Therefore, decreasing beta diversity from south to north reflects decreasing climate differentiation within more northerly latitude zones, and primarily post-glacial dispersal limitation north of 50°N.     

Invasion by alligator weed,Alternanthera philoxeroides,is associated with decreased species diversity across the latitudinal gradient in China

Aims Invasive species occurrence and their effects on biodiversity may vary along latitudes. We examined the occurrence (species cover) and relative dominance (importance value) of invasive alligator weed, Alternanthera philoxeroides, in its terrestrial habitat in China through a large-scale latitudinal field investigation.Methods We established 59 plots along the latitudinal transect from 21°N to 37°N. We recorded species name, abundance, height and individual species coverage of plants in every quadrat. We then measured α-species diversity variations associated with the A. philoxeroides community across the latitudinal range. We also analyzed the effect of latitude on plant species' distributions in this community by using canonical correspondence analysis (CCA).Important findings We found that species cover and importance value of A. philoxeroides increased in areas <35°N, but decreased at higher latitudes. Lower latitudes supported greater species diversity than higher latitudes. Small-scale invasion of A. philoxeroides was associated with higher species diversity, but community diversity was lower when A. philoxeroides species cover exceeded 36%. Community plant species changed from mesophyte to hygrophyte gradually from low to high latitude. Our research suggests that latitude had significant influences on community diversity which interacted with the biotic resistance of a community and impact of invasion. Consequently, A. philoxeroides may become more invasive and have greater negative impacts on community species diversity in higher latitudes as global climate changes.     

The latitudinal position of peak marine diversity in living and fossil biotas

Aim Peak marine taxonomic diversity has only rarely occurred at or near the equator during the Phanerozoic Eon, in contrast to the present‐day pattern. This fundamental difference is difficult to reconcile because the latitude at which peak diversity occurs for living marine taxa has not yet been explicitly determined at a broad taxonomic and spatial scale. Here, we attempt to determine this value in order to compare the contemporary and fossil patterns directly. Location Our data are global in coverage. Methods We used a literature compilation of 149 present‐day marine latitudinal diversity gradients. We summed the number of marine taxa that exhibited peak diversity within 10° latitudinal bins. In addition, we recorded locality data, general habitat (benthic/pelagic), and the taxonomic level of the study organisms. Results We found that peak diversity for most sampled marine taxa currently occurs between 10° and 20° N, even after correcting for a Northern Hemisphere sampling bias. Moreover, this peak position is a global phenomenon: it is found across habitats and higher taxa, within all sampled ocean basins, and on both sides of the Atlantic and Pacific oceans. Benthic taxa, which dominate our data, exhibit one peak at 10°–20° N, while pelagic taxa exhibit a peak at 10°–20° N and an additional peak at 10°–20° S, producing a distinct trough at the equator. Main conclusions Our data indicate that peak marine diversity for many taxa is currently within 10°–20° N rather than at the equator, and that this is not likely to result from either undersampling at lower latitudes or the pattern being dominated by a particular taxon. Possible explanations may include a coincidence with the intertropical convergence zone, a mid‐domain effect, abundant shallow marine habitat, or high ocean temperatures at latitudes nearest the equator. Regardless of its exact cause, the position of peak diversity should be considered a fundamental feature of the latitudinal diversity gradient that must be accounted for within attempts to explain the latter’s existence.     

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.     

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.