Latitudinal gradient in species richness of the New World Triatominae (Reduviidae)

Aim To quantify the latitudinal gradient in species richness in the New World Triatominae and to explore the species‐energy and area hypotheses as possible causes. Location The gradient was studied for North and South America, between 43° N and 32° S. Methods A database was constructed containing the geographical distribution of the 118 New World Triatominae species based on data extracted from several published sources. Species richness was recorded as the number of species present within 5° latitudinal bands. We used univariate and multivariate models to analyse the relationship between area within each latitudinal belt, land surface temperature, and potential evapotranspiration as explanatory variables, and species richness. All variables were georeferenced and data were extracted using a GIS. Results Species richness of Triatominae increases significantly from the poles towards the Equator, peaking over the 5°?10 ° S latitudinal band. It increases according to a linear model, both north and south of the Equator, although a quadratic model fits better to southern hemisphere data. Richness correlates with habitable geographical area, when it is analysed through a nonlinear multiple regression factoring out latitude, only in the southern hemisphere. Regarding the species‐energy hypothesis, a multiple regression analysis controlling the effect of latitude shows a significant relationship between temperature and species richness. This effect is more pronounced in the southern hemisphere. Species richness shows a strong longitudinal trend south of the Equator (increasing to the east), but not north of the Equator. This differential pattern is reflected in significant interactions between longitude and both latitude and temperature in models of the species richness of the New World Triatominae. Main conclusions To our knowledge, this is the first time that a latitudinal gradient in species richness has been shown and analysed for obligate haematophagous organisms, and it shows that the species–energy hypothesis can account for this phenomenon. This relationship is stronger in the southern hemisphere.     

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.     

The relationship between geographic area and the latitudinal gradient in species richness in New World birds

One hypothesis for the latitudinal gradient in species richness observed in most animal taxa is that the richness of a region is determined by its geographic area. However, the relationship between geographic area and species richness across regions is generally weak. It has been suggested that this is because species from the tropics spill out of this region of high richness, artificially inflating the richness of other regions. This generates the interesting prediction that the area and richness of extra-tropical regions should be more strongly correlated if tropical species are excluded. We test this prediction using the avifauna of the New World. We find that there is indeed a relationship between the land area and species richness of a region once tropical species are excluded. This relationship is independent of the latitude and productivity of regions. Both latitude and productivity can explain variance in richness unexplained by land area. There is no relationship between land area and species richness if tropical species are not excluded from the analysis, suggesting that tropical species do indeed mask the relationship between richness and area. We conclude that our results generally support the geographic area hypothesis, although tests of its other predictions and on other land masses are required.     

The latitudinal gradient in dispersal constraints: ecological specialisation drives diversification in tropical birds

Physiological and behavioural constraints arising from ecological specialisation are proposed to limit gene flow and promote diversification in tropical lineages. In this study, we use phylogenetic analyses to test this idea in 739 Amazonian bird species. We show that patterns of species and subspecies richness are best predicted by a suite of avian specialisms common in tropical avifaunas but rare in the temperate zone. However, this only applied to niche traits associated with dispersal limitation rather than vagility. These findings are consistent with the view that diversity is promoted by more finely partitioned niches, although not simply by coevolutionary adaptation and niche packing as is often assumed. Instead, they suggest that diversification is driven by dispersal constraints, and that niches characterised by these constraints are biased towards tropical systems. We conclude that specialised tropical niches reduce the likelihood of dispersal across barriers, thereby increasing allopatric diversification and contributing to the latitudinal diversity gradient.     

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.     

The latitudinal gradient of species diversity among North American grasshoppers (Acrididae) within a single habitat: a test of the spatial heterogeneity hypothesis

Abstract. The spatial heterogeneity hypothesis predicts a positive relationship between habitat complexity and species diversity: the greater the heterogeneity of a habitat, the greater the number of species in that habitat. On a regional scale, this hypothesis has been proposed to explain the increases in species diversity from the poles to the tropics: the tropics are more diverse because they contain more habitats. On the local scale, the spatial heterogeneity hypothesis suggests that the tropics are more diverse because they contain more microhabitats. The positive relationship between habitat heterogeneity and species diversity, on the local scale, is well documented. In this paper, we test whether habitat heterogeneity on the local scale can explain the latitudinal gradient of species diversity on the regional scale. We determined the latitudinal gradient of species diversity of 305 species of North American grasshoppers using published distribution maps. We compared the slope of this multihabitat (regional-scale) gradient with the slope of a within-habitat (local-scale) gradient in the prairie grasslands. Our results show no significant difference between the slopes at the two scales. We tested the generality of our results by comparing multi- and within-habitat latitudinal gradients of species diversity for ants, scorpions and mammals using data from the literature. These results are in accordance with those from grasshoppers. We can therefore reject the local-scale spatial heterogeneity hypothesis as a mechanism explaining the regional-scale latitudinal gradient of species diversity. We discuss alternative mechanisms that produce this gradient.     

Ecological limits and diversification rate: alternative paradigms to explain the variation in species richness among clades and regions

Diversification rate is one of the most important metrics in macroecological and macroevolutionary studies. Here I demonstrate that diversification analyses can be misleading when researchers assume that diversity increases unbounded through time, as is typical in molecular phylogenetic studies. If clade diversity is regulated by ecological factors, then species richness may be independent of clade age and it may not be possible to infer the rate at which diversity arose. This has substantial consequences for the interpretation of many studies that have contrasted rates of diversification among clades and regions. Often, it is possible to estimate the total diversification experienced by a clade but not diversification rate itself. I show that the evidence for ecological limits on diversity in higher taxa is widespread. Finally, I explore the implications of ecological limits for a variety of ecological and evolutionary questions that involve inferences about speciation and extinction rates from phylogenetic data.     

A latitudinal gradient of beta diversity for exotic vascular plant species in North America

Determining relationships between the ranges of introduced species and geographical and environmental factors is an important step in understanding the mechanisms and processes of the spread of introduced species. In this study, I examined the beta diversity and latitude relationship for all naturalized exotic species of vascular plants in North America at a continental scale. Beta diversity was calculated as the absolute value of the slope of the relationship between the natural logarithm of the Simpson index of similarity (lnS) and spatial distance between pairs of state‐level exotic floras within four latitudinal zones examined. Relative contributions of spatial distance and environmental difference to species turnover between exotic floras were examined. I found that beta diversity decreased monotonically from low to high latitudes: beta diversity for the southernmost zone was shallower than that for the northernmost zone by a factor of 2.6. Regression models of lnS in relation to spatial distance and environmental (climatic and topographical) difference for each latitudinal zone demonstrated that the explanatory power of these variables diminishes monotonically with latitude: the explained variance in lnS is 70.4%, 62.1%, 53.9%, and 33.9%, respectively, for the four latitudinal zones from south to north. For the southernmost zone, 58.3% of the variance in lnS is explained by climate variables and topography, and spatial distance explains only 2.3% of the variance. In contrast, for the northernmost zone, more than half the amount (22.5%) of the explained variance in lnS is attributable to spatial distance, and the remaining (18.9%) of the explained variance is attributable to climate variables and topography.     

High local species richness of parasitic wasps (Hymenoptera: Ichneumonidae; Pimplinae and Rhyssinae) from the lowland rainforests of Peruvian Amazonia

Abstract.  1. The parasitic wasp family Ichneumonidae (Hymenoptera) is of great interest because it has been claimed that its species richness does not increase with decreasing latitude.
2. No extensive studies of the family have been conducted in South American localities.
3. Arthropods were sampled using 27 Malaise traps in the Allpahuayo–Mishana National Reserve (56 000 ha) in the north-eastern Peruvian Amazonian lowland rainforest. The total duration of the sampling programme was 185 Malaise trap months.
4. Altogether, 88 species were collected. This is one of the highest local pimpline and rhyssine species numbers ever recorded. A comparison with results from Mesoamerica revealed that at equal numbers of individuals sampled, the number of Pimplinae and Rhyssinae species in Peruvian Amazonia is at least twofold compared with lowland locations in Mesoamerica and somewhat higher than in the most species-rich Costa Rican higher altitude localities.
5. Non-parametric methods of estimating species richness were applied. These suggest that additional sampling would yield a considerable number of new Pimplinae and/or Rhyssinae species.     

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.     

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.     

Environmental predictors of global parrot (Aves: Psittaciformes) species richness and phylogenetic diversity

Aim Spatial patterns of phylogenetic diversity (PD) aid our ability to discern diversification rate mechanisms underlying hypotheses for the large‐scale distribution of biodiversity. We develop a predictive framework for the way in which spatial patterns of PD vary with those of species richness, depending on the balance between speciation and extinction rates. Within this framework, diversification processes thought to underlie the productive energy, ambient energy, topographic variability and habitat variety hypotheses predict that gradients of increase in species richness will be associated with: (1) decreasing extinction rates where driven by productive energy, hence increasing relative PD (i.e. PD controlling for species richness, or PD_rel); (2) a similar positive relationship between ambient energy and PD_rel; (3) increasing speciation rates where driven by topographic variability, hence decreasing PD_rel; and (4) no consistent relationship between PD_rel and habitat variety when driven by the latter. We test these predictions using distributional data on parrots. Location Neotropical, Afrotropical, Indo‐Malayan and Australasian realms. Methods Spatial models were used to test the predictions. Results Globally, a positive association between productive energy and PD_rel confirms prediction (1). However, within realms, hump‐shaped relationships suggest the importance of decreasing extinction rates up to a threshold level of productive energy, and the increasing importance of speciation rates thereafter. Ambient energy is positively associated with PD_rel in Australasia, Indo‐Malaya, and globally, supporting prediction (2). However, this is driven by the coincidence of highest PD_rel in areas of high ambient energy and intermediate productive energy (i.e. in seasonal tropical environments), which may be characterized by relatively low speciation and extinction rates. In the Neotropics, increasing topographic variability is associated with decreasing PD_rel and increasing species richness, suggesting an increasing gradient of speciation, supporting prediction (3). Elsewhere, the signal of this mechanism may be obscured by collinearities with energy gradients. The lack of an overall relationship between habitat diversity and PD_rel confirms prediction (4). Main conclusions Spatial patterns of PD_rel in relation to environmental gradients may be sensitive to collinearities among those gradients. Nevertheless, patterns emerge which have implications for the relative importance of speciation and extinction processes in generating latitudinal diversity gradients.     

Latitudinal patterns of mammalian species richness in the New World: the effects of sampling method and faunal group

Abstract. Although the latitudinal gradient of species richness for mammals in North America is well documented, few investigators have quantified the relationship in South America. We examined the pattern in North and South America, at two spatial scales (2.5° and 5°) for each of two sampling methods (quadrats and latitudinal bands). A scale effect was evident for quadrats but not for bands. Significant linear relationships between species richness and latitude were found for three faunal groups: all mammals, nonvolant species, and bats. Effects of area confound the latitudinal relationship. By statistically removing such effects, we found that the latitudinal gradient is not an artifact of the species-area relationship, and that the latitudinal gradients for North and South America were statistically indistinguishable. Our data suggest that both faunal subgroups, nonvolant species and bats, contributed substantially to the overall mammalian pattern. Further, multiple regression analyses showed that only latitude is a necessary variable to explain bat richness; for nonvolant species, in addition to latitude, area and longitude may be important.     

Changes in species richness of vascular plants under the impact of air pollution: a global perspective

Aim To investigate the general pattern of changes in species richness and diversity of vascular plants due to environmental contamination and associated habitat changes imposed by point polluters, and identify the sources of variation in the response of plant communities to industrial pollution. Location Global. Methods We collected species richness and diversity data from 86 studies that were conducted around 60 atmospheric point polluters worldwide and reported in 95 papers (published in 1953–2007). We used meta‐analysis to search for a general effect and to compare between polluter types and plant groups, and linear regression to describe the latitudinal gradient and to quantify relationships between pollution and effect size. Results Although the species richness of vascular plants generally decreased with pollution, the effects were not uniform across the studies. Polluters that cause soil acidification imposed a stronger detrimental effect on plant diversity than industries whose emissions increased soil pH. An overall adverse effect was primarily due to the contribution of non‐ferrous smelters and aluminium plants; the effects of other SO₂‐emitting industries were less detrimental, albeit negative, and the effects of chemical plants, fertilizer factories and cement industries did not differ from zero. Longevity of the pollution impact only made a slight contribution to the detected variation, while adverse effects increased with increase in pollution load. Main conclusions This study is the first demonstration of geographical variation in the responses of plant communities to aerial emissions: adverse effects increased from high to low latitudes, and this pattern was explained primarily by increases in both the diversity of original (undisturbed) communities and mean summer temperatures. The latter result suggests that under a future warmer climate the existing pollution loads may become more harmful. Model calculations indicate that a detectable depauperation of plant communities is unlikely if the polluter emits < 1500 t of SO₂ annually.