Interspecific patterns of species richness, geographic range size, and body size among New World venomous snakes

Many higher taxa exhibit latitudinal gradients in species richness, geographic range size, and body size. However, these variables are often interdependent, such that examinations of univariate or bivariate patterns alone may be misleading. Therefore, I examined latitudinal gradients in, and relationships between, species richness, geographic range size, and body size among 144 species of New World venomous snakes [families Elapidae (coral snakes) and Viperidae (pitvipers)]. Both lineages are monophyletic, collectively span 99° of latitude, and are extremely variable in body size and geographic range sizes. Coral snakes exhibit highest species richness near the equator, while pitviper species richness peaks in Central America. Species – range size distributions were strongly right-skewed for both families. There was little support for Bergmann's rule or Rapoport's rule for snakes of either family, as neither body size nor range size increased significantly with latitude. However, range area and median range latitude were positively correlated above 15° N, indicating a possible "Rapoport effect" at high northern latitudes. Geographic range size was positively associated with body size. Available continental area strongly influenced range size. Comparative (phylogenetically-based) analyses revealed that shared history is a poor predictor of range size variation within clades. Among vipers, trends in geographic range sizes may have been structured more by historical biogeography than by macroecological biotic factors.     

Elevational patterns of fern species assemblages and richness in central Japan

We conducted field surveys in 807 quadrats to evaluate the elevational belts, boundary and richness patterns of ferns and lycophytes in the temperate region of central Japan. We analysed fern species assemblages at 100 m elevational steps by cluster analysis and tested the number of upper and lower boundaries for elevational intervals against a null model of random distribution of elevational limits. We compared the pattern of fern species richness along the elevational gradients in central Japan with patterns in several locations to evaluate the fern flora in central Japan in relation to the rest of the world. We recorded 261 ferns species in total, which is one-third of the Japanese ferns. We found clear elevational boundaries of fern assemblages at 900 and 1,800 m and three fern elevational zones, which corresponded well to the elevational limits of forest types in central Japan. The pattern of fern species richness in central Japan was an asymmetric hump-shaped pattern that peaked close to the sea level, with the peak of local richness at lower elevations than that of regional richness. We found that the peak of fern species richness along the elevational gradient in Japan was located at lower elevations than that of fern elevational patterns in several locations around the world.     

Elevational patterns of species richness and endemism for some important taxa in the Hengduan Mountains, southwestern China

We describe the elevational patterns of species richness and endemism of some important taxa in the Hengduan Mountains, southwest China. Species richness data came from publications, an online database, herbaria and field work. Species richness was estimated by rarefaction and interpolation. The Hengduan Mountains region was divided into a southern and northern subregion, and all species were assigned to four groups based on their distributional range within this region. The conditional autoregressive model (CAR) was used to relate species richness and explanatory variables. The elevational patterns of total, endemic and non-endemic species richness, at subregion and entire region scales, presented to be unimodal and peaked at similar elevations. Area size was strongly related with species richness, and was more powerful in explaining variation in species richness in the northern subregion than in the southern subregion. A single climatic variable (mean annual rainfall, potential evapotranspiration or moisture index) showed a weak relationship with the elevational pattern of species richness. Area and climatic variables together explained more than 67% of the variation in non-endemic richness, 53% in total richness, and 50% in endemic richness. There were three patterns of endemism at the generic level with increasing elevation: namely endemism increased, decreased, or peaked at middle elevations. All selected taxa have experienced rapid speciation and evolution within this region, which plays an important role in the uniform elevational patterns of total, endemic and non-endemic richness, and in the multiform elevational patterns of endemism. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Latitudinal patterns in European ant assemblages: variation in species richness and body size

Using published distributions of 65 species from the British Isles and northern Europe, we show that ant assemblages change with latitude in two ways. First, as commonly found for many types of organisms, the number of ant species decreased significantly with increasing latitude. For Ireland and Great Britain, species richness also increased significantly with region area. Second, although rarely demonstrated for ectotherms, the body size of ant species, as measured by worker length, increased significantly with increasing latitude. We found that this body-size pattern existed in the subfamily Formicinae and, to a lesser extent, in the Myrmicinae, which together comprised 95% of the ant species in our study area. There was a trend for formicines to increase in size with latitude faster than myrmicines. We also show that the pattern of increasing body size was due primarily to the ranges of ant species shifting to higher latitudes as their body sizes increased, with larger formicines becoming less represented at southerly latitudes and larger myrmicines becoming more represented at northerly latitudes. We conclude by discussing five potential mechanisms for generating the observed body-size patterns: the heat-conservation hypothesis, two hypotheses concerning phylogenetic history, the migration-ability hypothesis, and the starvation-resistance hypothesis.     

Species richness, species range size and ecological specialisation among African primates: geographical patterns and conservation implications

The geographical distribution of species richness and species range size of African anthropoid primates (catarrhines) is investigated and related to patterns of habitat and dietary niche breadth. Catarrhine species richness is concentrated in the equatorial regions of central and west Africa; areas that are also characterised by low average species range sizes and increased ecological specificity. Species richness declines with increasing latitude north and south of the equator, while average species range size, habitat and dietary breadth increase. Relationships between species richness, species range size and niche breadth remain once latitudinal and longitudinal effects have been removed. Among areas of lowest species richness, however, there is increased variation in terms of average species range size and niche breadth, and two trends are identified. While most such areas are occupied by a few wide-ranging generalists, others are occupied by range-restricted specialist species. That conservation efforts increasingly focus on regions of high species richness may be appropriate if these regions are also characterised by species that are more restricted in both their range size and their ecological versatility, although special consideration may be required for some areas of low species richness.     

Elevational patterns of frog species richness and endemic richness in the Hengduan Mountains, China: geometric constraints, area and climate effects

We studied frog biodiversity along an elevational gradient in the Hengduan Mountains, China. Endemic and non-endemic elevational diversity patterns were examined individually. Competing hypotheses were also tested for these patterns. Species richness of total frogs, endemics and non-endemics peaked at mid-elevations. The peak in endemic species richness was at higher elevations than the maxima of total species richness. Endemic species richness followed the mid-domain model predictions, and showed a nonlinear relationship with temperature. Water and energy were the most important variables in explaining elevational patterns of non-endemic species richness. A suite of interacting climatic and geometric factors best explained total species richness patterns along the elevational gradient. We suggest that the mid-domain effect was an important factor to explain elevational richness patterns, especially in regions with high endemism.     

Elevational gradients in ant species richness: area, geometry, and Rapoport's rule

Studying the distributions of plants and animals along environmental gradients can illuminate the factors governing and maintaining species diversity. There are two general predictions of how species richness and elevation are related: either species richness decreases monotonically with increasing elevation or richness peaks at mid-elevations. Several processes might contribute to this pattern. In this paper, I examine patterns in ant species richness along elevational gradients in three states in the western US: Colorado, Nevada, and Utah. I test for the effects of available area and the geometric constraints model on species richness patterns. I also test Rapoport's rescue hypothesis, which relates the extent of species' elevational ranges to patterns in species richness. In each state, species richness peaked at mid-elevations. Area explained more variation in species richness than the geometric constraints model in Colorado and Utah, but not in Nevada. Area and geometric constraints together explained 90%, 99%, and 57% of the variation in species richness in Colorado, Nevada, and Utah, respectively. Even though there were peaks at mid-elevations, I still found a strong Rapoport effect. This work suggests that the influences of area and geometric constraints cannot be overlooked when examining patterns in species richness along environmental gradients.     

Elevational patterns of plant richness and their drivers on an Asian mountain

We examined the elevational patterns of plant species along two transects on Mt Seorak, South Korea, and calculated four richness indices from field survey data: total number of species per 100 m elevational band; mean number of species per plot in each elevational band; total estimated number of species per elevational band; and beta diversity of each elevational band. We evaluated the effects of area, mean distance between plots, climatic variables (mean annual temperature and precipitation), and productivity on the richness patterns along the two transects. In total, 235 plant species belonging to 72 families and 161 genera were recorded from 130 plots along the two transects. The analyses revealed different patterns including monotonic decline, and unimodal and multimodal shapes for richness indices of total, woody, and herbaceous plants with the change in elevation along the two transects. The proportion of suitable area (as opposed to rocky areas) was the best predictor for total number of species per elevational band, mean number of species per plot, and total estimated number of species per elevational band of total and herbaceous plants along the two transects. Mean distance between plots was the most important variable for beta diversity of all plant groups. Although regional area, climatic variables, and productivity were important variables for predicting woody plant richness patterns, the effects were not consistent between the two transects. Our study suggests that elevational species richness patterns may differ not only among different plant groups, but also between nearby elevational transects, and that these differences are explained by differences in the underlying mechanisms shaping these patterns.     

Elevational patterns of species richness and density of rattan palms (Arecaceae: Calamoideae) in Central Sulawesi, Indonesia

We studied species richness and density of rattan palms in 50 plots of 10 × 100 m² each between 250 m and 2420 m in eight sites in Lore Lindu National Park (LLNP), Central Sulawesi. Rattans were observed in all sample sites, representing three genera and 34 species. The elevational patterns for species richness and density were humped-shaped with maxima around 1000 m. Polynomial models of second order explained 59 and 32% of species richness and density with the factor elevation, respectively. A majority of rattan species (65%) overlapped between 1000 and 1100 m elevation, while a pronounced change in the rattan flora occurred above 1100 m. Commercially important rattan species (Calamus zollingeri, C. ornatus var. celebicus, Daemonorops macroptera) were not observed above 1250 m. The change of species assemblage was significantly related to elevation (56%), followed by geographical distance (47%) and precipitation (40%). Less than 10% of LLNP is lowland forests, much of which is threatened by agricultural intensification. In contrast, montane forests are well represented in the park and high elevation forests are not subject to agricultural conversion or intensive harvesting of rattan and other forest products.     

Multiple environmental determinants of regional species richness and effects of geographic range size

Understanding patterns of species richness at broad geographic extents remains one of the most challenging yet necessary scientific goals of our time. Many hypotheses have been proposed to account for spatial variation in species richness; among them, environmental determinants have played a central role. In this study, we use data on regional bat species richness in the New World to study redundancy and complementarity of three environmental hypotheses: energy, heterogeneity and seasonality. We accomplish this by partitioning variation in species richness among components associated with unique and combined effects of variables from each hypotheses, and by partitioning the overall richness gradient into gradients of species with varying breadths of geographic distribution. These three environmental hypotheses explain most variation in the species richness gradient of all bats, but do not account for all positive spatial autocorrelation at short distances. Although environmental predictors are highly redundant, energy and seasonality explain different and complementary fractions of variation in species richness of all bats. On the other hand, heterogeneity variables contribute little to explain this gradient. However, results change dramatically when richness is estimated for groups of species with different sizes of geographic distribution. First, the amount of variation explained by environment decreases with a decrease in range size; this suggests that richness gradients of small‐ranged species can not be explained as easily as those of broadly distributed species, as has been implied by analyses that do not consider differences in range size among species. Second, the relative contribution of environmental predictors to explained variation also changes with change in range size. Seasonality and energy are good predictors of species with broad distributions, but they loose almost all explanatory power for richness of species with small ranges. In contrast, heterogeneity, which is a relatively poor predictor of richness of species with large ranges, becomes the main predictor of richness gradients of species with restricted distributions. This suggests that range size is a different dimension on which heterogeneity and other environmental characteristics are complementary to each other. Our results suggest that determinants of species richness gradients might be complex, or at least more complex than many studies have previously suggested.     

Country-based patterns of total species richness,endemicity, and threatened species richness in African rodents and insectivores

Country-based patterns of total species richness, endemicity, and threatened species richness in African rodents and insectivores are studied in this paper. We found several patterns which were similar between insectivores and rodents. Indeed, in both groups we observed: (i) a significantly uneven distribution of species richness across countries and geographic regions with highest species richness peaks being in Middle Africa and lowest peaks in Northern Africa, (ii) species richness increasing with rainfall but being independent on a country’s surface area, (iii) in each country, the insectivore total species richness and endemic species richness increases were positively correlated with rodent total species richness and endemic species richness increases. However, number of endemics peaked in South Africa and D.R. Congo in both groups, but also in Tanzania for Insectivores and in Ethiopia for rodents. In addition, the highest numbers of threatened species occurred in D.R. Congo, Rwanda and Uganda for rodents and in South Africa, Tanzania and Cameroon for insectivores. The conservation implications of these results were discussed.     

Elevational changes in vascular plants richness,diversity, and distribution pattern in Abune Yosef mountain range,Northern Ethiopia

The aim of this research is to investigate the patterns of vascular plant species richness,diversity,and distribution along an elevation gradient in the Abune Yosef mountain range,Ethiopia.Preferential systematic sampling was employed to collect vegetation and environmental data along the elevation gradient.We found that plant species richness declines monotonically from low to high elevations.Specifically,vascular plant species richness and diversity were lower in the Afroalpine grassland(high elevation)than in the Dry evergreen Afromontane forest and Ericaceous forest(low elevations).In contrast,endemic vascular plant richness was significantly higher in the Afroalpine grassland than in the Dry evergreen Afromontane forest and Ericaceous forest.Elevation showed a significant impact on the richness,diversity,and endemism of vascular plants.According to Sorensen's coefficient,the similarity between Dry evergreen Afromontane forest and Ericaceous forest vegetation types is higher(32%)than the similarity between Ericaceous forest and Afroalpine grassland(18%).Only 5%similarity was recorded between the Dry evergreen Afromontane forest and Afroalpine grassland.Growth forms showed different elevationai richness patterns.Trees and liana increased monotonically up to 3300 m.Shrub and herb richness patterns followed a hump-shaped and inverted hump-shaped pattern along the elevation gradient.The elevation patterns of vascular plant species richness,diversity,and growth form in the present study may be attributed to differences in management intensity,spatial heterogeneity,microclimatic variations,and anthropogenic disturbances.     

Contrasting patterns of body size for Daphnia species that segregate by habitat

Summary We investigate how body size of two coexisting Daphnia species varies among 7 lakes that represent a gradient of predation risk. The two species segregate vertically in stratified lakes; D. galeata mendotae is typically smaller and more eplimnetic than D. pulicaria. The extent of vertical habitat partitioning, however, varies seasonally within and among lakes in apparent response to predation intensity by epilimnetic planktivorous fishes. Daphnia pulicaria uses the epilimnion at low levels of fish predation but is restricted to the hypolimnion under high fish predation, whereas D. galaeta mendotae always utilizes the epilimnion. The species display contrasting patterns of genetic variation in neonate size and size at maturity. D. pulicaria is larger in lakes with higher fish and Chaoborus densities whereas D. galeata mendotae is smaller. This contrast in body size in lakes with high predation is associated with greater habitat segregation in those lakes. In lakes with low predation risk, the two species are similar in body size at birth and maturity.Authorship order alphabetical     

Density, body mass and parasite species richness of terrestrial mammals

We investigated the relationships between helminth species richness and body mass and density of terrestrial mammals. Cross-species analysis and the phylogenetically independent contrast method produced different results. A non-phylogenetic approach (cross-species comparisons) led to the conclusion that parasite richness is linked to host body size. However, an analysis using phylogenetically independent contrasts showed no relationship between host body size and parasite richness. Conversely, a non-phylogenetic approach generated a negative relationship between parasite richness and host density, whereas the independent contrast method showed the opposite trend – that is, parasite richness is positively correlated with host density. From an evolutionary perspective, our results suggest that opportunities for parasite colonization depend more closely on how many hosts are available in a given area than on how large the hosts are. From an epidemiological point of view, our results confirm theoretical models which assume that host density is linked to the opportunity of a parasite to invade a population of hosts. Our findings also suggest that parasitism may be a cost associated with host density. Finally, we provide some support for the non-linear allometry between density and mammal body mass (Silva and Downing, 1995), and explain why host density and host body mass do not relate equally to parasite species richness.     

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.     

Generic species richness and body mass in North American mammals: Support for the inverse relationship of body size and speciation rate

Generic species richness, the number of species per genus, is examined as a function of mean generic body mass for extant North American mammals. Species richness decreases as an inverse power function with increased mass, and the Spearman rank correlation coefficient of the logio transformed data is significant (r_s= ‐0.37). When the data are partitioned by trophic level, the relationship is not statistically significant for carnivores but strengthens for herbivores (r_s= ‐0.46). This interesting but incidental effect is due to the negligible number of diminutive and excessively large carnivores, which is in turn determined by foraging strategies. Alternate hypotheses for the “right‐skewed”; size distribution of modern North American mammals, such as disproportionate extinction of large species, differential species longevity, and a geographical scaling function, are rejected in favor of the proposition that elevated levels of speciation are restricted to animals of small body mass, as originally proposed by Gould and Eldredge (1977). This phenomenon is explained as a function of habitat restriction and particularly in herbivores, limited home range size. Aquatic mammals, regardless of body size, speciate rarely. Cope's Rule, the tendency of many animal groups to evolve towards large size, is understood as a probabilistic statement reflecting the phylogenetic tendencies of a disproportionately high number of small species alive at any given point in time.