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.     

Historical isolation,range expansion,and secondary contact of two highly divergent mitochondrial lineages in spotted salamanders (Ambystoma maculatum)

Abstract.— The high species diversity of aquatic and terrestrial faunas in eastern North America has been attributed to range reductions and allopatric diversification resulting from historical climate change. The role these processes may have played in speciation is still a matter of considerable debate; however, their impacts on intraspecific genetic structure have been well documented. We use mitochondrial DNA sequences to reconstruct an intraspecific phylogeny of the widespread North American spotted salamander, Ambystoma maculatum , and test whether phylogenetic patterns conform to regional biogeographical hypotheses about the origins of diversity in eastern North America. Specifically, we address the number and locations of historical refugia, the extent and patterns of postglacial colonization by divergent lineages, and the origin and affinities of populations in the Interior Highland region. Despite apparent morphological uniformity, genetic discontinuities throughout the range of this species suggest that populations were historically fragmented in at least two refugia in the southern Appalachian Mountains. The ranges of these two highly divergent clades expanded northward, resulting in two widely distributed lineages that are sympatric in regions previously proposed as suture zones for other taxa. The evolutionary history of spotted salamander populations underscores the generality of biogeographical processes in eastern North America: despite differences in population size, glacial refugia, and vagility, similar signatures of differentiation are evident among and within widespread taxa.     

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.     

Evolutionary and ecological causes of species richness patterns in North American angiosperm trees

Climate and evolutionary factors (e.g. diversification, time‐for‐speciation, niche conservatism) are both thought to be major drivers of species richness in regional assemblages. However, few studies have simultaneously investigated the relative effects of climate and evolutionary factors on species richness across a broad geographical extent. Here, we assess their relative effects on species richness of angiosperm trees across North America. Species richness of angiosperm trees in 1175 regional assemblages were related to climate and phylogenetic structure using a structural equation modeling (SEM) approach. Climate was quantified based on the mean temperature of the coldest month and mean annual precipitation. Evolutionary factors (time‐for‐speciation vs diversification) were inferred from phylogeny‐based measures of mean root distance, phylogenetic species variability, and net relatedness index. We found that at the continental scale, species richness is correlated with temperature and precipitation with approximately similar strength. In the SEM with net relatedness index and phylogenetic species variability and with all the 1175 quadrats, the total direct effect size of phylogenetic structure on species richness is greater than the total direct effect size of climate on species richness by a factor of 3.7. The specific patterns of phylogenetic structure (i.e. greater phylogenetic distances in more species rich regions) are consistent with the idea that time and niche conservatism drive richness patterns in North American angiosperm trees. We conclude that angiosperm tree species richness in regional assemblages in North America is more strongly related to patterns of phylogenetic relatedness than to climatic variation. The results of the present study support the idea that climatic and evolutionary explanations for richness patterns are not in conflict, and that evolutionary processes explain both the relationship between climate and richness and substantial variation in richness that is independent of climate.     

The region effect on mesoscale plant species richness between eastern Asia and eastern North America

The greater number of plant species in temperate eastern Asia compared to eastern North America has been ascribed to both local environment and regional characteristics, but the relative contributions of each have not been resolved. In this analysis, we related species richness of flowering plants in mesoscale floras (<10⁴ km²) dominated by temperate forest vegetation to area, elevation, latitude, and several climate variables. When analyses were conducted separately within each region, area and, in eastern Asia, elevation, were the primary determinants of species richness. It appears that the number of species in mesic temperate floras within these regions is largely unrelated to the relatively narrow range of local climate factors associated with these floras. Analysis of covariance of the logarithm of species richness with the logarithm of area (b=0.148) and climate measurements as independent variables revealed a region effect, with species richness in eastern Asia exceeding that in eastern North America by 0.294 log₁₀ units, or a factor of 2.0. Similar regional differences in species richness were apparent in floras compiled from larger areas. Understanding differences in plant species richness between regions requires consideration of regional influences, whose effects should be tested in comparative analyses based on floristic surveys of ecologically characterized small areas.     

Continental and regional ranges of North American mammals: Rapoport's rule in real and null worlds

Aim To assess the relationship between species richness and distribution within regions arranged along a latitudinal gradient we use the North American mammalian fauna as a study case for testing theoretical models. Location North America. Methods We propose a conceptual framework based on a fully stochastic mid‐domain model to explore geographical patterns of range size and species richness that emerge when the size and position of species ranges along a one‐dimensional latitudinal gradient are randomly generated. We also analyse patterns for the mammal fauna of North America by comparing empirical results from a biogeographical data base with predictions based on randomization null models. Results We confirmed the validity of Rapoport's rule for the mammals of North America by documenting gradients in the size of the continental ranges of species. Additionally, we demonstrated gradients of mean regional range size that parallel those of continental range. Our data also demonstrated that mean range size, measured both as a continental or a regional variable, is significantly correlated with the geographical pattern in species richness. All these patterns deviated sharply from null models. Main conclusions Rapoport's statement of an areographic relationship between species distribution and richness is highly relevant in modern discussions about ecological patterns at the geographical scale.     

Taxonomic diversity of the terrestrial bird and mammal fauna in temperate and boreal biomes of the northern hemisphere

Abstract. Using comprehensive range information of northern Hemisphere birds and mammals, we assessed the taxonomic diversity of these two groups in four different regions: Europe, east Asia, and western and eastern North America. East Asia is the richest region in the number of bird and mammal species, genera, families and orders, except that mammal species richness is highest in western North America. Eastern North America is taxonomically the poorest region, but when only forest-associated taxa were considered in mammals taxonomic diversity is equally low in Europe and in eastern North America, and in birds, Europe is the least diverse region. Patterns in endemic taxa follow overall taxonomic diversity. The proportion of shared taxa between regions is higher among boreal species and genera than among all taxa. A comparison with tree species diversity underpins the role of east Asia as the most diverse of all northern biota. Largely congruent patterns at different taxonomic levels emphasizes the role of historical processes, such as differential extinction rate in response to paleoenvironmental fluctuations, in producing these patterns, but we stress the need for more research on the coevolution of species diversity and habitat diversity.     

Energy gradients and the geographic distribution of local ant diversity

Geographical diversity gradients, even among local communities, can ultimately arise from geographical differences in speciation and extinction rates. We evaluated three models—energy-speciation, energy-abundance, and area—that predict how geographic trends in net diversification rates generate trends in diversity. We sampled 96 litter ant communities from four provinces: Australia, Madagascar, North America, and South America. The energy-speciation hypothesis best predicted ant species richness by accurately predicting the slope of the temperature diversity curve, and accounting for most of the variation in diversity. The communities showed a strong latitudinal gradient in species richness as well as inter-province differences in diversity. The former vanished in the temperature-diversity residuals, suggesting that the latitudinal gradient arises primarily from higher diversification rates in the tropics. However, inter-province differences in diversity persisted in those residuals—South American communities remained more diverse than those in North America and Australia even after the effects of temperature were removed.     

Tree diversity,tree height and environmental harshness in eastern and western North America

Does variation in environmental harshness explain local and regional species diversity gradients? We hypothesise that for a given life form like trees, greater harshness leads to a smaller range of traits that are viable and thereby also to lower species diversity. On the basis of a strong dependence of maximum tree height on site productivity and other measures of site quality, we propose maximum tree height as an inverse measure of environmental harshness for trees. Our results show that tree species richness is strongly positively correlated with maximum tree height across multiple spatial scales in forests of both eastern and western North America. Maximum tree height co‐varied with species richness along gradients from benign to harsh environmental conditions, which supports the hypothesis that harshness may be a general mechanism limiting local diversity and explaining diversity gradients within a biogeographic region.     

Patterns in the local assembly of Egyptian rodent faunas: Areography and species combinations

We implicate ecological processes in assembly patterns of Egyptian desert rodents using spatial variation of distribution and composition of local assemblages. We also compare our assemblages to prior analyses of North American and Australian small mammal communities in terms of species richness and representation of trophic groups.We studied patterns of occurrence of 29 rodent species among 335 collecting events in 308 sites using museum specimen records resulting from a country-wide survey. The studied taxa vary greatly in their natural histories and ecology. Fully 69% of studied species (20) were localized in 30 or fewer sites (9% of all sites). Site incidence was closely correlated with the geographic range size of species, while local abundance of a species showed little relationship to its geographic range. The species richness of local assemblages ranged from a lone species at 73 sites to 11 species at 3 sites. A total of 214 different combinations were recorded of which 164 (77%) were unique to a single site. G. gerbillus was both the most abundant and most ubiquitous species. Coexistence with other species was positively correlated with incidence and geographic range size. Body mass distribution was remarkably uniform for the fauna as a whole, and influenced the geographic distributions and abundance of individual species.Our sites have low species richness and substantial variability in species composition, which also characterize desert rodent communities elsewhere. Habitat requirements, exclusive distributions of sibling species, low primary productivity, and Egypt's location all influenced species assembly. The Egyptian and Australian deserts supported higher proportions of low richness assemblages compared to North America. As in North America, Egyptian sites were dominated by granivorous species.     

Vascular plant diversity in eastern Asia and North America: historical and ecological explanations

Taxonomic diversity of vascular plants (ferns, gymnosperms and angiosperms) was compared between eastern Asia and North America. Eastern Asia has significantly higher species richness in all three classes but the difference was greatest in ferns and least in angiosperms. Differences in taxonomic treatments between the two continents are not likely contributors to these patterns. The relationship of regional to global species richness across the three plant classes suggested that diversity patterns were relatively homogeneous at three taxonomic levels. Thus, differences in species richness are established at the family level and are therefore relatively old. The previously noted fact that eastern Asia has a higher proportion of primitive taxa was shown by analyses both among and within plant classes. Diversity patterns across three taxonomic levels (i.e. family, genus and species) of the three classes may reflect the relative historical positions of the two continents (following continental drift) to the centre(s) of their origin, neighbouring land masses, differential speciation/extinction rates, and switches in dominance levels associated with climate change (including glaciation), as well as reproductive/dispersal mechanisms of the three plant classes.     

Beta diversity and latitude in North American mammals: testing the hypothesis of covariation

Several hypotheses attempt to explain the latitudinal gradient of species diversity, but some basic aspects of the pattern remain insufficiently explored, including the effect of scales and the role of beta diversity. To explore such components of the latitudinal gradient, we tested the hypothesis of covariation, which states that the gradient of species diversity should show the same pattern regardless of the scale of analysis. The hypothesis implies that there should be no gradients of beta diversity, of regional range size within regions, and of the slope of the species-area curve. For the fauna of North American mammals, we found contrasting results for bats and non-volant species. We could reject the hypothesis of covariation for non-volant mammals, for which the number of species increases towards lower latitudes, but at different rates depending on the scale. Also, for this group, beta diversity is higher at lower latitudes, the regional range size within regions is smaller at lower latitudes, and z, the slope of the species-area relationship is higher at lower latitudes. Contrarily bats did not show significant deviations from the predictions of the hypothesis of covariation: at two different scales, species richness shows similar trends of increase at lower latitudes, and no gradient can be demonstrated for beta diversity, for regional range size, or for the slopes of the species-area curve. Our results show that the higher diversity of non-volant mammals in tropical areas of North America is a consequence of the increase in beta diversity and not of higher diversity at smaller scales. In contrast, the diversity of bats at both scales is higher at lower latitudes. These contrasting patterns suggest different causes for the latitudinal gradient of species diversity in the two groups that are ultimately determined by differences in the patterns of geographic distribution of the species.     

The effect of the taxon and geographic range size of host eucalypt species on the species richness of gall-forming insects

Abstract This field study was designed to test whether the taxonomic group and geographic range size of a host plant species, usually found to influence insect species richness in other parts of the world, affected the number of gall species on Australian eucalypts. We assessed the local and regional species richness of gall-forming insects on five pairs of closely related eucalypt species. One pair belonged to the subgenus Corymbia, one to Monocalyptus, and three to different sections of Symphyomyrtus. Each eucalypt pair comprised a large and a small geographic range species. Species pairs were from coastal or inland regions of eastern Australia. The total number of gall species on eucalypt species with large geographic ranges was greater than on eucalypt species with small ranges, but only after the strong effect of eucalypt taxonomic grouping was taken into account. There was no relationship between the geographic range size of eucalypt species and the size of local assemblages of gall species, but the variation in insect species composition between local sites was higher on eucalypt species with large ranges than on those with small ranges. Thus the effect of host plant range size on insect species richness was due to greater differentiation between more widespread locations, rather than to greater local species richness. This study confirms the role of the geographic range size of a host plant in the determination of insect species richness and provides evidence for the importance of the taxon of a host plant.     

Local-regional relationships and the geographical distribution of species

Aim Local‐regional (LR) species diversity plots were conceived to assess the contribution of regional and local processes in shaping the patterns of biological diversity, but have been used also to explore the scaling of diversity in terms of its alpha, beta, and gamma components. Here we explore the idea that patterns in the geographical ranges of species over a continent can determine the shape of small region to large region (SRLR) plots, which are equivalent to LR plots when comparing the diversity of sites at two regional scales. Location To test that idea, we analysed the diversity patterns at two regional scales for the mammals of North America, defined as the mainland from Alaska and Canada to Panama. Method We developed a theoretical model relating average range size of species over a large‐scale region with its average regional point species diversity (RPD). Then, we generated a null model of expected SRLR plots based on theoretical predictions. Species diversities at two scales were modelled using linear and saturation functions for Type I and Type II SRLR relationships, respectively. We applied the models to the case of North American mammals by examining the regional diversity and the RPD for 21 large‐scale quadrats (with area equal to 160,000 km²), arranged along a latitudinal gradient. Results Our model showed that continental and large‐scale regional patterns of distribution of species can generate both types of SRLR relationship, and that these patterns can be reflected in LR plots without invoking any kind of local processes. We found that North American nonvolant mammals follow a Type I SRLR relationship, whereas bats follow a Type II pattern. This difference was linked to patterns in which species of the two mammalian groups distribute in geographical space. Conclusion Traditional LR plots and the new SRLR plots are useful tools in exploring the scaling of species diversity and in showing the relationship between distribution and diversity. Their usefulness in comparing the relative role of local and regional processes is, however, very limited.     

The environmental basis of North American species richness patterns among Epicauta (Coleoptera: Meloidae)

Understanding regional variability in species richness is necessary for conservation efforts to succeed in the face of large-scale environmental deterioration. Several analyses of North American vertebrates have shown that climatic energy provides the best explanation of contemporary species richness patterns. The paucity of analyses of insect diversity patterns, however, remains a serious obstacle to a general hypothesis of spatial variation in diversity. We collected species distribution data on a North American beetle genus, Epicauta (Coleoptera: Meloidae) and tested several major diversity hypotheses. These beetles are generally grasshopper egg predators as larvae, and angiosperm herbivores as adults. Epicauta richness is highest in the hot, dry American southwest, and decreases north and east, consistent with the species richness-energy hypothesis. Potential evapotranspiration, which is also the best predictor of richness patterns among North American vertebrates, explains 80.2% of the variability in Epicauta species richness. Net primary productivity and variables measuring climatic heat energy only (such as PET) are not generally comparable, though they are sometimes treated as if they were equivalent. We conclude that the species richness-energy hypothesis currently provides a better overall explanation for Epicauta species richness patterns in North America than other major diversity hypotheses. The observed relationship between climatic energy and regional species richness may provide significant insight into the response of ecological communities to climate change.     

DOES NICHE CONSERVATISM PROMOTE SPECIATION? A CASE STUDY IN NORTH AMERICAN SALAMANDERS

Recent speciation research has generally focused on how lineages that originate in allopatry evolve intrinsic reproductive isolation, or how ecological divergence promotes nonallopatric speciation. However, the ecological basis of allopatric isolation, which underlies the most common geographic mode of speciation, remains poorly understood and largely unstudied. Here, we explore the ecological and evolutionary factors that promote speciation in Desmognathus and Plethodon salamanders from temperate eastern North America. Based on published molecular phylogenetic estimates and the degree of geographic range overlap among extant species, we find strong evidence for a role for geographic isolation in speciation. We then examine the relationship between climatic variation and speciation in 16 sister-taxon pairs using geographic information system maps of climatic variables, new methods for modeling species' potential geographic distributions, and data on geographic patterns of genetic variation. In contrast to recent studies in tropical montane regions, we found no evidence for parapatric speciation along climatic gradients. Instead, many montane sister taxa in the Appalachian Highlands inhabit similar climatic niches and seemingly are allopatric because they are unable to tolerate the climatic conditions in the intervening lowlands. This temporal and spatial-ecological pattern suggests that niche conservatism, rather than niche divergence, plays the primary role in promoting allopatric speciation and montane endemism in this species-rich group of vertebrates. Our results demonstrate that even the relatively subtle climatic differences between montane and lowland habitats in eastern North America may play a key role in the origin of new species.     

Local and regional influences on patterns of parasite species richness of central European fishes

Local, regional and global influences on the patterns of parasite species richness of 39 freshwater fish species from Central Europe were investigated. Host local abundance and host occurrence were considered respectively as local and regional factors, while host geographical range in longitude and latitude was considered as a global factor. Influences of size, ecology and behavior of hosts were also included in a comparative analysis using the independent contrasts method. We considered host habitat, host diet, host shoaling behavior and mobility. We found a positive relationship between local occurrence of fish and global range of their distribution. We confirmed previous findings showing the importance of host behavior and ecology on the variability of parasite species richness. Second, we showed how a global pattern, such as host geographical range, may affect the variability in parasite species richness through its effects on local abundance and distribution of hosts. A negative relationship between endoparasite species richness and host longitudinal range was found. This suggests that fish with eastern distribution live in the boundary of their distribution in Central Europe far from their center of distribution, which should also be characterized by a higher diversity of parasites.     

Determinants of lepidopteran community composition and species diversity in eastern deciduous forests: roles of season, eco-region and patch size

Ecologists have long recognized that factors operating at both local and regional scales influence whether a given species occurs in an ecological community. The relative roles of variables manifested at local and regional scales on community structure, however, remain an unexplored issue for many faunas. To address this question, we compared the community composition and species diversity of forest Lepidoptera between (i) large forest tracts in historically glaciated and unglaciated regions of the eastern deciduous forest in North America, and (ii) large and small forest patches within a highly fragmented forest landscape. Specifically, we used seasonally stratified sampling to test whether regional and local differences in moth communities were related to variation in stand structure and floristic composition. At the local scale, we tested three alternative hypotheses describing the effects of patch size on moth species richness: species impoverishment, species replacement, or species supplementation. Cluster analysis revealed significant compositional differences in moth communities sampled between (i) early and late seasons, (ii) glaciated and unglaciated forest eco‐regions, and (iii) large and small forest patches. Canonical correspondence analysis suggested that floristic variation at regional scales had a greater role in determining moth community composition than local vegetation or patch‐size effects. Species richness was higher in the glaciated North Central Tillplain, and was attributable to a more diverse herbaceous feeding moth assemblage. Finally, we found evidence that both species impoverishment and species replacement processes structure the moth fauna of small woodlots; the richness of moths with larvae that feed on woody plants decreased with patch area, but herbaceous feeding species increased in diversity in smaller patches. Thus, our results suggest that local and regional differences in moth community structure are mediated by differences in host‐plant resources attributable to regional biogeographic history and local differences in patch size. Because community composition appeared to be more sensitive to environmental variation than species richness, we suggest that monitoring lepidopteran species diversity in forests will not detect significant changes in species composition due to environmental change.     

Species richness in plant genera disjunct between temperate eastern Asia and North America

The species richness of 109 amphi-Pacific disjunct genera was examined in eastern Asia and North America. Although the entire flora of eastern Asia contains approximately one-third more species than that of North America, the difference in species richness among disjunct taxa is less. When woody and herbaceous genera are considered separately, the former exhibit a strong diversity bias favouring eastern Asia whereas there is no significant difference in diversity between continents among herbaceous genera. This result is not due to habitat differences between woody and herbaceous genera, because the disjunct herbs inhabit primarily moist forests and woodlands. This result is also not related to relative phylogenetic advancement, even though older major lineages of plants tend to have a predominance of woody taxa. Woody genera are distributed in lower latitudes than herbaceous genera on both continents, and both woody and herbaceous genera are distributed in lower latitudes in eastern Asia than in North America. The North American temperate flora is primarily a relict of a flora form 7 more widespread throughout the Northern Hemisphere. Contemporary patterns of diversity suggest that the effects of climate changes in the late Tertiary were less severe in eastern Asia and promoted diversification, but were more severe in North America and may have caused widespread extinction. The difference in the effect of climate change on diversity in herbaceous and woody lineages reflects the different ecological relationships of species having these contrasting life forms. Clearly, the contemporary floras of eastern Asia and North America bear the imprint of history and emphasize the important interface between ecological relationships and evolutionary responses.     

Climate and history explain the species richness peak at mid-elevation for Schizothorax fishes (Cypriniformes: Cyprinidae) distributed in the Tibetan Plateau and its adjacent regions

Aim  We studied elevational species richness patterns of Schizothorax fishes and identified the roles of ecological and evolutionary factors in shaping the patterns of elevational diversity.
Location  The Tibetan Plateau and its adjacent regions.
Methods  We assembled distribution and altitude data for all Schizothorax species using the literature. We merged ecological and evolutionary approaches to test the relationships between species richness and ecological factors (climate, area, the mid-domain effect) or evolutionary factors (diversification rates and time of colonization).
Results  We found that species richness of Schizothorax fishes peaked at mid-elevations. Rainfall, area, the mid-domain effect and diversification rate were weak predictors of the richness pattern. Temperature showed a nonlinear relationship with species richness. Temperature and time of colonization were the most important variables in explaining the elevational diversity pattern.
Main conclusion  Our findings indicate that the time-for-speciation effect and niche conservatism play important roles in variation of species richness.