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.     

Competition, sub-lethal mortality and diversity on Southern Ocean coastal rock communities

Overgrowth competition, sub-lethal mortality (some zooids killed but colony survives) and subsequent growth from fragments, and diversity of communities encrusting rocks (pebble through to boulder size) were examined from five Antarctic localities, along a latitudinal gradient. There were distinct gradients in the ecology of both assemblages and individual species with latitude within the Southern Ocean. Compared with warm-water equivalents, the polar assemblages had many fewer species, considerably less variability in species richness, highly transitive interactions and lower incidences of interspecific encounters. There is no gradual transition but a dramatic alteration of assemblage and species level ecology around the margins of the Southern Ocean, which may primarily be due to ice scour. Sub-lethal mortality was common and peaked on different boulder sizes along the Antarctic Peninsula. This is indicative of the disturbance cline along the region.     

How well do we know the Antarctic marine fauna? A preliminary study of macroecological and biogeographical patterns in Southern Ocean gastropod and bivalve molluscs

The aim of this study was to use data for gastropod and bivalve molluscs to determine whether the fauna of the Southern Ocean is sufficiently well known to establish robust biogeographical and macroecological patterns. We chose molluscs for this work because they have been collected by almost every biological expedition to Antarctica, and are relatively well known taxonomically. Sampling of the continental shelf fauna is reasonably full and extensive, although new species are still being described and there are significant gaps in sampling off Wilkes Land and in the Bellingshausen and Amundsen Seas. Species richness was highest in those areas that have been subject to the most intense research activity and this pattern remained even after correction for sampling intensity. The low species richness of the Southern Ocean molluscan fauna compared with many tropical sites is confirmed, and is related principally to the absence of the large number of rare taxa that characterize some tropical assemblages. There is as yet no convincing evidence for a latitudinal cline in molluscan diversity within the Southern Ocean. Multivariate analyses defined biogeographical provinces very similar to those established previously, though they also identified a number of finer-scale sub-provinces including a small area of high diversity off Enderby Land. Most Southern Ocean gastropods and bivalves are rare, with limited distributions; relatively few taxa have circumpolar distributions.     

Global biodiversity and biogeography of mangrove crabs: Temperature,the key driver of latitudinal gradients of species richness

Mangroves are ideal habitat for a variety of marine species especially brachyuran crabs as the dominant macrofauna. However, the global distribution, endemicity, and latitudinal gradients of species richness in mangrove crabs remains poorly understood. Here, we assessed whether species richness of mangrove crabs decreases towards the higher latitudes and tested the importance of environmental factors such as Sea Surface Temperature (SST) in creating the latitudinal gradients in species richness of mangrove crabs. A total of 8262 distribution records of 481 species belonging to six families of mangrove crabs including Camptandriidae, Dotillidae, Macrophthalmidae, Ocypodidae, Sesarmidae, and Oziidae were extracted from open-access databases or collected by the authors, quality controlled, cleaned, and analyzed. Species richness was plotted against 5° latitudinal bands in relation to environmental factors. The R software and ArcGIS 10.6.1 were used to analyze the species latitudinal range and richness as well as to map the distribution of mangrove forest, endemic species, species geographical distribution records, and biogeographic regions. The Indo-West Pacific showed the highest species richness of mangrove crabs where more than 65% of species were found in the Indian Ocean and along the western Pacific Ocean. Our results showed that there are 11 significantly different biogeographic regions of mangrove crabs. The highest endemicity rate was observed in the NW Pacific Ocean (29%). Latitudinal patterns of species richness in Macrophthalmidae, Ocypodidae, and Sesarmidae showed an increasing trend from the poles toward the intermediate latitudes including one dip near the equator. However, latitudinal gradients in Camptandriidae, Dotillidae, and Oziidae were unimodal increasing from the higher latitudes towards the equator. Species richness per 5° latitudinal bands significantly increased following mean SST mean (°C), calcite, euphotic depth (m), and mangrove area (km²) across all latitudes, and tide average within each hemisphere. Species richness significantly decreased with dissolved O₂ (ml l⁻¹) and nitrate (μmol l⁻¹) over all latitudes and in the southern hemisphere. The climax of global latitudinal species richness for some mangrove was observed along latitudes 20° N and 15°–25° S, not at the equator. This can suggest that temperature is probably the key driver of latitudinal gradients of mangrove crabs’ species richness. Species richness and mangrove area were also highly correlated.     

Biodiversity and biogeography of southern temperate and polar bryozoans

Aim To describe the distribution of biodiversity and endemism of bryozoans in southern temperate and polar waters. We hypothesized that we would find: (1) no strong latitudinal richness gradient; (2) striking contrasts in richness and endemism between clades and between regions; and (3) that faunal similarity of regions would cluster geographically around each southern continent. Location South Atlantic, Indian and Pacific Oceans and the Southern Ocean. Methods We constructed a data base from known literature, regional data bases and recent finds. We regionalized each southern continent, calculated levels of richness and endemism for each region and continent, and used primer 5 to perform multivariate statistical analysis. Results A third (1681) of global bryozoan species described occur south of 30° S, of which c. 87% were cheilostomes. In richness we found no latitudinal cline and change across longitude was stronger. New Zealand was richest and had the most (60%) endemic species, followed by Antarctica at 57%. There were striking contrasts in regional richness and endemism between clades but the highest levels of between‐region similarity were around Antarctica. The timing of past continent connectivity was reflected. Main conclusions Bryozoans show strong hemispherical asymmetry in richness and, like molluscs and corals, decrease away from Australasia rather than with latitude. Species endemism is much lower in Antarctic bryozoans than previously thought, and as this taxon is not particularly dispersive and is now amongst the best studied regionally, maybe Antarctic endemism in general is lower and Antarctica less cut‐off to species dispersal than previously thought. However, Antarctic generic endemism is double the level previously calculated and regional faunal similarities are much higher than around other continents – both reflecting long‐term isolation. Bryozoans, in contrast to the paradigm of Antarctic fauna, may be fairly robust to predicted climate change. Paradoxically, they may also be one of the best taxa to monitor to sensitively detect marine benthic responses.     

Climatic drivers of hemispheric asymmetry in global patterns of ant species richness

Although many taxa show a latitudinal gradient in richness, the relationship between latitude and species richness is often asymmetrical between the northern and southern hemispheres. Here we examine the latitudinal pattern of species richness across 1003 local ant assemblages. We find latitudinal asymmetry, with southern hemisphere sites being more diverse than northern hemisphere sites. Most of this asymmetry could be explained statistically by differences in contemporary climate. Local ant species richness was positively associated with temperature, but negatively (although weakly) associated with temperature range and precipitation. After contemporary climate was accounted for, a modest difference in diversity between hemispheres persisted, suggesting that factors other than contemporary climate contributed to the hemispherical asymmetry. The most parsimonious explanation for this remaining asymmetry is that greater climate change since the Eocene in the northern than in the southern hemisphere has led to more extinctions in the northern hemisphere with consequent effects on local ant species richness.     

美洲森林群落beta多样性的纬度梯度性

Beta多样性度量不同时空尺度物种组成的变化,是生物多样性的重要组成部分;理解其地理格局和形成机制已成为当前生物多样性研究的热点问题。基于Alwyn H. Gentry在美洲收集的131个森林样方数据,采用倍性和加性分配方法度量群落beta多样性,检验beta多样性随纬度的变化趋势,并分析其形成机制。研究表明:(1) 美洲森林群落beta多样性随纬度增加显著下降,热带和亚热带地区beta多样性高于温带地区;此格局可由物种分布范围的纬度梯度性和不同粒度(grain)下物种丰富度与纬度回归斜率的差异推论得出;(2) 加性分配方法表明beta多样性对各个温度带森林群落gamma多样性的相对贡献率平均为78.2%,并且随纬度升高而降低;(3) 美洲南半球森林群落beta多样性高于其北半球,这可能反映了区域间物种进化和环境变迁历史的差异。此外,还探讨了不同beta多样性计算方法的适用情景,首次证实了森林生态系统群落水平beta多样性的纬度梯度性,这对研究生物多样性的形成机制和生物多样性保护都具有重要的意义。     

Ants and people: a test of two mechanisms potentially responsible for the large‐scale human population–biodiversity correlation for Formicidae in Europe

Aim: Recent coarse‐scale studies have shown positive relationships between the biodiversity of plants/vertebrates and the human population. Little is known about the generality of the pattern for invertebrates. Moreover, biodiversity and human population might correlate because they both covary with other factors such as energy availability and habitat heterogeneity. Here we test these two non‐mutually exclusive mechanisms with ant species‐richness data from the Fauna Europaea. Location Forty‐three European countries/regions. Methods We derived mixed models of total, native and exotic ant species richness as a function of human population size/density, controlling for country area, plant species richness (as a proxy for habitat heterogeneity), and mean annual temperature and precipitation (variables related to energy availability). Results Ant species richness increased significantly with increasing human population. This result was confirmed when controlling for variations in country area. Both for human population size/density and for ant species richness, there were positive correlations with temperature but not with precipitation. This finding is in agreement with the energy‐availability hypothesis. However, we observed a negative latitudinal gradient in ant and plant species richness, although not in human population size/density. Plant species richness was positively correlated with ant species richness but not with human population size/density. Thus, there is evidence that this type of habitat heterogeneity can play a role in the observed latitudinal gradient of ant species richness, but not in the positive correlation between ant species richness and human population. The results were confirmed for the 545 native and the 32 exotic ant species reported, and we observed a good correlation between exotic and native ant species richness. Main conclusions Ant species richness in European countries conforms to six macroecological patterns: (1) a negative latitudinal gradient; and a positive (2) species–energy relationship, (3) species–area relationship, (4) correlation with plant species richness, (5) exotic–native species richness correlation, and (6) species–people correlation. There is some evidence for the energy‐availability hypothesis, but little evidence for habitat heterogeneity as an explanation of the large‐scale human population–ant biodiversity correlation. This correlation has implications for the conservation of ant diversity in Europe.     

Human impacts, energy availability and invasion across Southern Ocean Islands

Aim Ongoing biological invasions will enhance the impacts of humans on biodiversity. Nonetheless, the effects of exotic species on diversity are idiosyncratic. Increases in diversity might be a consequence of similar responses by species to available energy, or because of positive relationships between human density, energy and propagule pressure. Here we use data from the Southern Ocean island plants and insects to investigate these issues. Location The Southern Ocean Islands ranging from Tristan da Cunha to Heard Island and South Georgia. Methods Generalized linear models are used to explore the relationships between indigenous and exotic species richness for plants and insects on two different islands. Similar models are used to examine interactions between indigenous and exotic species richness, energy availability and propagule pressure at the regional scale. Results Positive relationships were found between indigenous and exotic species richness at local scales, although for plants, the relationship was partially triangular. Across the Southern Ocean Islands, there was strong positive covariation between indigenous and exotic plant species richness and insect species richness, even taking spatial autocorrelation into account. Both exotic and indigenous plant and insect species richness covaried with energy availability, as did human visitor frequency. When two islands with almost identical numbers of human visits were contrasted, it was clear that energy availability, or perhaps differences in climate‐matching, were responsible for differences in the extent of invasion. Conclusion In plants and insects, there are positive relationships between indigenous and exotic diversity at local and regional scales across the Southern Ocean islands. These relationships are apparently a consequence of similar responses by both groups and by human occupants to available energy. When visitor frequency is held constant, energy availability is the major correlate of exotic species richness, though the exact mechanistic cause of this relationship requires clarification.     

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.     

Arctic plant diversity in the Early Eocene greenhouse

For the majority of the Early Caenozoic, a remarkable expanse of humid, mesothermal to temperate forests spread across Northern Polar regions that now contain specialized plant and animal communities adapted to life in extreme environments. Little is known on the taxonomic diversity of Arctic floras during greenhouse periods of the Caenozoic. We show for the first time that plant richness in the globally warm Early Eocene (approx. 55-52 Myr) in the Canadian High Arctic (76° N) is comparable with that approximately 3500 km further south at mid-latitudes in the US western interior (44-47° N). Arctic Eocene pollen floras are most comparable in richness with today's forests in the southeastern United States, some 5000 km further south of the Arctic. Nearly half of the Eocene, Arctic plant taxa are endemic and the richness of pollen floras implies significant patchiness to the vegetation type and clear regional richness of angiosperms. The reduced latitudinal diversity gradient in Early Eocene North American plant species demonstrates that extreme photoperiod in the Arctic did not limit taxonomic diversity of plants.     

Latitudinal gradients of species richness: a test of the geographic area hypothesis at two ecological scales

The geographic area hypothesis advances area as the primary cause of latitudinal gradients in diversity. The greater area of tropical zones, it suggests, stimulates speciation, inhibits extinction, and leads to increased species richness compared to the situation in smaller temperate and boreal zones. Because bats exhibit exceptionally strong latitudinal gradients of richness at multiple spatial scales in the New World, they are an appropriate system with which to test the geographic area hypothesis. We used range maps for 250 species of New World bats to estimate species richness in biogeographic zones at two hierarchical spatial scales: biome types and provinces. We then conducted a series of regression analyses to evaluate the ability of area to account for latitudinal gradients in species richness. However, spillover (zonal bleeding) of tropical species into extra-tropical zones may mask the species-area relationship and alter perceptions of the latitudinal gradient. To address this issue, we conducted additional analyses excluding tropical species, using a series of increasingly inclusive definitions of tropical ranges. Ecogeographic zones of the New World are not larger at tropical versus extra-tropical latitudes. Moreover, spillover of tropical species into ecogeographic zones within extra-tropical regions generally does not diminish the association between richness and area. Nonetheless, the latitudinal gradient of species richness is strong and significant at both ecogeographic scales. Clearly, area does not drive the latitudinal gradient of bat species richness in the New World. In fact, area represents a source of noise rather than a dominant signal at the focal scale of biome types and provinces in the Western Hemisphere.     

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.     

An evolutionary framework for the polar regions

Abstract: Our knowledge of the nature, generation and maintenance of largescale biodiversity patterns is still far from complete. This is particularly so in the Southern Hemisphere and in the marine realm, where recent taxonomic investigations of Mollusca and other invertebrate groups has cast doubt upon the existence of a simple cline in species richness between the tropics and the pole. Comparatively high regional diversity values for the shelled gastropods and other epifaunal taxa implies a considerable evolutionary legacy; this is supported, at least in part, by available evidence from the fossil record. Certain families within the living gastropod fauna maintain their prominence when traced back 40 m.y., and perhaps even longer; in addition, several Southern Ocean gastropod and bivalve genera can now be traced back to at least the late Eocene. Use of a variety of refugia may have enabled many taxa to survive repeated glacial advances. As we begin to revise our concept of the nature of latitudinal diversity gradients, so we also need to examine regional variations in evolutionary rates. Clearly this is a complex issue. but recourse to a pilot study based on the molluscan fossil record suggests that there may be no significant difference between the rates of radiation of tropical and cold-temperatdpolar taxa. The most diverse clades, which are all tropical, are simply the oldest. What data are available from the fossil record indicate that there is no appreciable latitudinal variation in rates of extinction either. Time, but not necessarily environmental stability, would appear to be crucial to the development of pockets of high taxonomic diversity. Recent improvement in our understanding of the biology of many polar marine invertebrates suggests that life in cold water is not an insuperable evolutionary problem. Of qual importance to any intrinsic properties of organisms which may have governed the differentiation of large-scale biodiversity patterns is the role of extrinsic processes. Foremost among these has almost certainly been repeated range shifts in response to Cenozoic climatic cycles.     

Body size determines the strength of the latitudinal diversity gradient

In most groups of organisms, the species richness decreases from the tropics to the poles. The mechanisms causing this latitudinal diversity gradient are still controversial. We present data from a comprehensive weighted meta-analysis on the strength of the latitudinal gradient in relation to body size. We sampled literature data on the correlation between species richness and latitude for a variety of organisms, ranging from trees to protozoa. In addition, own data on the presence of large-scale diversity patterns for diatoms were included, both for local and regional species richness. The strength of the latitudinal gradient was positively correlated to the size of the organisms. Strongest decreases of species richness to the poles was found for large organisms like trees and vertebrates, whereas meiofauna, protozoa and diatoms showed weak or no correlations between species richness and latitude. These results imply that latitudinal gradients are shaped by non-equilibrium (regional) processes and are persistent under conditions of dispersal limitation.     

Latitudinal gradient of species diversity: multi-scale variability in rocky intertidal sessile assemblages along the Northwestern Pacific coast

This study examined the latitudinal gradient of species diversity of rocky intertidal sessile assemblages on the slopes of rocks along the Northwestern Pacific coast of Japan, located between 31°N and 43°N, by explicitly incorporating an hierarchical spatial scale into the monitoring design. The specific questions were to examine: (1) whether there is a latitudinal gradient of regional diversity, (2) how spatial components of the regional diversity (local diversity and turnover diversity) vary with latitude depending on spatial scale, and (3) whether the latitudinal gradient differs between different measures of species diversity, i.e. species richness and Simpsons diversity index. We measured coverage and the presence or absence of all sessile organisms in a total of 150 census plots established at five shores in each of six regions. The results showed that there were clear latitudinal gradients in regional species richness and in species turnover among shores. However, these patterns were not reflected in smaller-scale local species richness. For Simpsons diversity index, there was no evidence of latitudinal clines either in regional diversity or in spatial components. These results suggest that relative abundance of common species does not vary along latitude, while the number of rare species increases with decreasing latitude.

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An erratum to this article can be found at     

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.     

Patterns of functional diversity across an extensive environmental gradient: vertebrate consumers, hidden treatments and latitudinal trends

Over the last two decades, although much has been learned regarding the multifaceted nature of biodiversity, relatively little is known regarding spatial variation in constituents other than species richness. This is particularly true along extensive environmental gradients such as latitude. Herein, we describe latitudinal gradients in the functional diversity of New World bat communities. Bat species from each of 32 communities were assigned to one of seven functional groups. Latitudinal gradients existed for the richness, diversity and scaled‐dominance of functional groups. No significant patterns were observed for evenness of functional groups. Measures of functional diversity were different in magnitude and increased towards the equator at a faster rate than expected given the underlying spatial variation in species richness. Thus, latitudinal gradient in species richness alone do not cause the latitudinal gradient in functional diversity. When variation in species composition of the regional fauna of each community was incorporated into analyses, many differences between observed and simulated patterns of functional diversity were not significant. This suggests that those processes that determine the composition of regional faunas strongly influence the latitudinal gradient in functional diversity at the local level. Nonetheless, functional diversity was lower than expected across observed sites. Community‐wide responses to variation in the quantity and quality of resources at the local level probably contribute to differences in functional diversity at local and regional scales and enhance beta diversity.     

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.     

Weak links: 'Rapoport's rule' and large-scale species richness patterns

Many hypotheses have been proposed to explain regional species richness patterns. Among these, ‘Rapoport's rule’ has sparked considerable controversy by stating that the latitudinal gradient in species richness can be explained indirectly as a function of narrower geographic ranges for species at low latitudes. Annual climatic variability, or deviation from mean climatic conditions, has been hypothesized to moderate this phenomenon. Furthermore, taxa that avoid much of this seasonality, such as temperate zone insects that enter diapause or species that migrate, were predicted to show reduced latitudinal gradients in richness. I test the suggested link between ‘Rapoport's rule’ and species richness for two higher level insect taxa as well as for the class Mammalia. Although these taxa exhibit the well-known latitudinal gradient in species richness, simple annual climatic variability and deviation from mean annual climatic conditions provide very poor predictions of species richness in each of them. Potential evapotranspiration, a measurement of ambient climatic energy, explains most of the observed variance in regional species richness patterns for all three taxa, consistent with the species richness-energy hypothesis. I find no support for an indirect link between ‘Rapoport's rule’ and terrestrial species richness patterns in North America.