Community-based processes behind species richness gradients: contrasting abundance–extinction dynamics and sampling effects in areas of low and high productivity

Aim  To consider the role of local colonization and extinction rates in explaining the generation and maintenance of species richness gradients at the regional scale.
Location  A Mediterranean biome (oak forests, deciduous forests, shrublands, pinewoods, firwoods, alpine heathlands, crops) in Catalonia, Spain.
Methods  We analysed the relative importance of direct and indirect effects of community size in explaining species richness gradients. Direct sampling effects of community size on species richness are predicted by Hubbell's neutral theory of biodiversity and biogeography. The greater the number of individuals in a locality, the greater the number of species expected by random direct sampling effects. Indirect effects are predicted by the abundance–extinction hypothesis, which states that in more productive sites increased population densities and reduced extinction rates may lead to high species richness. The study system was an altitudinal gradient of forest bird species richness.
Results  We found significant support for the existence of both direct and indirect effects of community size in species richness. Thus, both the neutral and the abundance–extinction hypotheses were supported for the altitudinal species richness gradient of forest birds in Catalonia. However, these mechanisms seem to drive variation in species richness only in low-productivity areas; in high-productivity areas, species richness was uncorrelated with community size and productivity measures.
Main conclusions  Our results support the existence of a geographical mosaic of community-based processes behind species richness gradients, with contrasting abundance–extinction dynamics and sampling effects in areas of low and high productivity.     

Plant species and growth form richness along altitudinal gradients in the southwest Ethiopian highlands

Questions: Do growth forms and vascular plant richness follow similar patterns along an altitudinal gradient? What are the driving mechanisms that structure richness patterns at the landscape scale? Location: Southwest Ethiopian highlands. Methods: Floristic and environmental data were collected from 74 plots, each covering 400 m². The plots were distributed along altitudinal gradients. Boosted regression trees were used to derive the patterns of richness distribution along altitudinal gradients. Results: Total vascular plant richness did not show any strong response to altitude. Contrasting patterns of richness were observed for several growth forms. Woody, graminoid and climber species richness showed a unimodal structure. However, each of these morphological groups had a peak of richness at different altitudes: graminoid species attained maximum importance at a lower elevations, followed by climbers and finally woody species at higher elevations. Fern species richness increased monotonically towards higher altitudes, but herbaceous richness had a dented structure at mid‐altitudes. Soil sand fraction, silt, slope and organic matter were found to contribute a considerable amount of the predicted variance of richness for total vascular plants and growth forms. Main Conclusions: Hump‐shaped species richness patterns were observed for several growth forms. A mid‐altitudinal richness peak was the result of a combination of climate‐related water–energy dynamics, species–area relationships and local environmental factors, which have direct effects on plant physiological performance. However, altitude represents the composite gradient of several environmental variables that were interrelated. Thus, considering multiple gradients would provide a better picture of richness and the potential mechanisms responsible for the distribution of biodiversity in high‐mountain regions of the tropics.     

Macroecological patterns of spider species richness across Europe

We analysed the pattern of covariation of European spider species richness with various environmental variables at different scales. Four layers of perception ranging from single investigation sites to the whole European continent were selected. Species richness was determined using published data from all four scales. Correlation analyses and stepwise multiple linear regression were used to relate richness to topographic, climatic and biotic variables. Up to nine environmental variables were included in the analyses (area, latitude, elevation range, mean annual temperature, local variation in mean annual temperature, mean annual precipitation, mean July temperature, local variation in mean July temperature, plant species richness). At the local and at the continental scale, no significant correlations with surface area were found, whereas at the landscape and regional scale, surface area had a significant positive effect on species richness. Factors that were positively correlated with species richness at both broader scales were plant species richness, elevation range, and specific temperature variables (regional scale: local variation in mean annual, and mean July temperature; continental scale: mean July temperature). Latitude was significantly negatively correlated with the species richness at the continental scale. Multiple models for spider species richness data accounted for up to 77% of the total variance in spider species richness data. Furthermore, multiple models explained variation in plant species richness up to 79% through the variables mean July temperature and elevation range. We conclude that these first continental wide analyses grasp the overall pattern in spider species richness of Europe quite well, although some of the observed patterns are not directly causal. Climatic variables are expected to be among the most important direct factors, although other variables (e.g. elevation range, plant species richness) are important (surrogate) correlates of spider species richness.     

Multiregional comparison of the ecological and phylogenetic structure of butterfly species richness gradients

Aim To examine butterfly species richness gradients in seven regions/countries and to quantify geographic mean root distance (MRD) patterns. My primary goal is to determine the extent to which an explanation for butterfly richness patterns based on tropical niche conservatism and the evolution of cold tolerance, proposed for the fauna of Canada and the USA, applies to other parts of the world. Location USA/Canada, Mexico, Europe/NW Africa, Transbaikal Siberia, Chile, South Africa and Australia. Methods Digitized range maps for butterfly species in each region were used to map richness patterns in summer (for all areas) and winter (for USA/Canada, Europe/NW Africa and Australia). A phylogeny resolved to subfamily was used to map the geographic MRD patterns. Regression trees and general linear models examined climatic and vegetation correlates of species richness and MRD within and among regions. Results Various combinations of climate and vegetation were strong predictors of species richness gradients within regions, but unresolved ‘regional’ factors contributed to the multiregional pattern. Regionally based differences in phylogenetic structure also exist, but MRD is negatively correlated with temperature both within and across areas. MRD patterns consistent with tropical niche conservatism occur in most areas. With a possible partial exception of Mexico, faunas in cold climates and in mountains are more derived than faunas in lowlands and tropical/subtropical climates. In USA/Canada, Europe and Australia, winter faunas are more derived than summer faunas. Main conclusions The phylogenetic pattern previously found in the USA and Canada is widespread in both the Northern and Southern Hemispheres, and niche conservatism and the evolution of cold tolerance is the likely explanation for the development of the global butterfly species richness gradient over evolutionary time. Contemporary climate also influences species richness patterns but is unlikely to be a complete explanation globally. The importance of climate is also manifested in the seasonal loss of more basal butterfly elements outside the tropics in winter.     

Global patterns in local number of insect galling species

Abstract. We evaluate a three-part hypothesis explaining why gall-inducing insect species richness is so high in scleromorphic vegetation: (1) persistence of low nutrient status scleromorphic leaves facilitates the galling habit in warm temperate latitudes; (2) favourable colonization sites for gallers result from reduced hygrothermal stress, high phenolics in the outer cortex of the gall, and reduced carnivore and fungal attack in the gall; and (3) in more mesic sites, mortality is high due to carnivore attack and invasion of galls by fungi. Over 280 samples of local species of galling herbivorous insects from fourteen countries on all continents except Antarctica revealed a strong pattern of highest richness in warm temperate latitudes, or their altitudinal equivalents. The peak of galling species richness on the latitudinal gradient from the equator into the Arctic was between 25 to 38° N or S. Galling species were particularly diverse in sclerophyllous vegetation, which commonly had greater than twelve species per local sample. In mesic, non-sclerophyllous vegetation types the number of galling species was lower with twelve or fewer species present. Many sites in sclerophyllous vegetation supported between thirteen and forty-six galling species locally, including campina islands in Amazonia, cerrado savanna in central Brazil, the Sonoran Desert in Arizona and Mexico, shrubland in Israel, fynbos in South Africa and coastal scleromorphic vegetation in Australia. At the same latitude, or its elevational equivalent, galling species richness was significantly higher in relatively xeric sites when compared to riparian or otherwise mesic habitats, even when scleromorphic vegetation dominated the mesic sites. The results were consistent with the hypothesis and extend to a more general level the patterns and predictions on the biogeography of gall-inducing insects.     

Latitudinal gradients of parasite species richness in primates

Infectious disease risk is thought to increase in the tropics, but little is known about latitudinal gradients of parasite diversity. We used a comparative data set encompassing 330 parasite species reported from 119 primate hosts to examine latitudinal gradients in the diversity of micro and macroparasites per primate host species. Analyses conducted with and without controlling for host phylogeny showed that parasite species richness increased closer to the equator for protozoan parasites, but not for viruses or helminths. Relative to other major parasite groups, protozoa reported from wild primates were transmitted disproportionately by arthropod vectors. Within the protozoa, our results revealed that vector‐borne parasites showed a highly significant latitudinal gradient in species richness. This higher diversity of vector‐borne protozoa near the tropics could be influenced by a greater abundance or diversity of biting arthropods in the tropics, or by climatic effects on vector behaviour and parasite development. Many vector‐borne diseases, such as leishmaniasis, trypanosomiasis, and malaria pose risks to both humans and wildlife, and nearly one‐third of the protozoan parasites from free‐living primates in our data set have been reported to infect humans. Because the geographical distribution and prevalence of many vector‐borne parasites are expected to increase because of global warming, these results are important for predicting future parasite‐mediated threats to biodiversity and human health.     

Macroecological explanations for differences in species richness gradients: a canonical analysis of South American birds

Aim To evaluate how spatial variation of species richness in different bird orders responds to environmental gradients and determine which order level trait best predicts these relationships. Location South America. Methods A canonical correlation analysis was performed between the species richness in each of 17 bird orders and eight environmental variables in 374, 220 × 220 km cells. Loadings associated with the first two canonical variables were regressed against six order‐level predictors, including diversification level (number of species in each order), body size, median geographical range size and characteristics included in the model to control Type I error rates (the phylogenetic relationship among orders and levels of local‐scale spatial autocorrelation). Results Richness patterns of 14 bird orders were highly correlated with the first canonical axis, indicating that most orders respond similarly to energy‐water gradients (primarily actual evapotranspiration, minimum temperature and potential evapotranspiration). In contrast, species richness within Trochiliformes, Apodiformes and Galliformes were also correlated with the second canonical variable, representing measures of mesoscale climatic variation (range in elevation within cells, minimum temperature, and the interaction term between them) and landcover (habitat diversity). We also found that total diversification within orders was the best predictor of the loadings associated with the first canonical axis, whereas body size of each order best predicted loadings on the second axis. Conclusion Our results broadly support climatic‐related hypotheses as explanations for spatial variation in species richness of different orders. However, both historical (order‐specific variation in speciation rates) and ecological (dispersal of species that evolved by independent processes into areas amenable to birds) processes can explain the relationship between order level traits, such as body size and diversification level, and magnitude of response to current environment, furnishing then guidelines for a further and deeper understanding of broad‐scale diversity gradients.     

Different evolutionary histories underlie congruent species richness gradients of birds and mammals

Aim The global species richness patterns of birds and mammals are strongly congruent. This could reflect similar evolutionary responses to the Earth’s history, shared responses to current climatic conditions, or both. We compare the geographical and phylogenetic structures of both richness gradients to evaluate these possibilities. Location Global. Methods Gridded bird and mammal distribution databases were used to compare their species richness gradients with the current environment. Phylogenetic trees (resolved to family for birds and to species for mammals) were used to examine underlying phylogenetic structures. Our first prediction is that both groups have responded to the same climatic gradients. Our phylogenetic predictions include: (1) that both groups have similar geographical patterns of mean root distance, a measure of the level of the evolutionary development of faunas, and, more directly, (2) that richness patterns of basal and derived clades will differ, with richness peaking in the tropics for basal clades and in the extra‐tropics for derived clades, and that this difference will hold for both birds and mammals. We also explore whether alternative taxonomic treatments for mammals can generate patterns matching those of birds. Results Both richness gradients are associated with the same current environmental gradients. In contrast, neither of our evolutionary predictions is met: the gradients have different phylogenetic structures, and the richness of birds in the lowland tropics is dominated by many basal species from many basal groups, whereas mammal richness is attributable to many species from both few basal groups and many derived groups. Phylogenetic incongruence is robust to taxonomic delineations for mammals. Main conclusions Contemporary climate can force multiple groups into similar diversity patterns even when evolutionary trajectories differ. Thus, as widely appreciated, our understanding of biodiversity must consider responses to both past and present climates, and our results are consistent with predictions that future climate change will cause major, correlated changes in patterns of diversity across multiple groups irrespective of their evolutionary histories.     

Insect species richness tracking plant species richness in a diverse flora: gall-insects in the Cape Floristic Region, South Africa

The Cape Floristic Region (CFR) is one of the most plant-species-rich regions in the world. It is also a warm temperate region and hypothetically should have high gall-insect species richness, making it interesting to investigate the relationship between the insects of the region and the rich flora. The relationship between gall-insect species richness (GSR) and plant richness was investigated for the Fynbos and for representatives of vegetation of the whole CFR. Samples (of up to 600 plants per transect for Fynbos) of woody shrubs were investigated for the presence of galls. The species richness of these insects was quantified, as well as plant species richness for each transect. GSR for Fynbos was compared to global figures for GSR. Fynbos harboured significantly more gall-insect species than other CFR vegetation types. GSR was positively correlated with CFR plant richness. GSR also closely tracked plant richness in Fynbos. GSR was not significantly influenced by other variables (elevation and aspect), suggesting that plant richness per se was an important factor in generating GSR. Fynbos GSR is comparable to other sclerophyllous regions of high GSR globally, corroborating that this vegetation type is conducive to gall-insect diversification. There is likely to be a high percentage of gall-insect endemism in the Fynbos, as might be expected from the high host fidelity of this insect group. Received: 22 September 1997 / Accepted: 16 February 1998     

Plant diversity patterns and their relationships with soil and climatic factors along an altitudinal gradient in the middle Tianshan Mountain area, Xinjiang, China

Patterns of plant diversity along the altitudinal gradient of Tianshan in central Xinjiang, China were examined. Plant and environment characteristics were surveyed from higher, south of Bogeda peak, to lower, north of Guerbantonggute desert. There were a total of 341 vascular plant, 295 herbage, 41 shrub, and seven tree species in the sampled plots. The plant richness of vegetation types generally showed a unimodal pattern along altitude, with a bimodal change of plant species number at 100-m intervals of altitudinal samples. The two belts of higher plant richness were in transient areas between vegetation types, the first in areas from dry grass to forest, and the second from forest to sub-alpine grass and bush. The beta diversity varied with altitudinal changes, with herbaceous species accounting for most species, and thus had similar species turnover patterns to total species. Matching the change of richness of plant species to environmental factors along altitude and correlating these by redundancy analysis revealed that the environmental factors controlling species richness and its pattern were the combined effects of temperature, precipitation, soil water, and nutrition. Water was more important at low altitude, and temperature at high altitude, and soil chemical and physical characters at middle altitudes. This study provides insights into plant diversity conservation of Bogeda Natural Reserve Areas in Tianshan Mountain. Nomenclatures: the scientific name for plants follows Flora of China (Compiling Committee of Flora of China).     

Latitudinal gradients in the species richness of Australian termites (Isoptera)

Abstract Using data on the geographic range of 260 described species in the Atlas of Australian Termites, seven ‘regions’ with more complete data, across a wide range of latitudes were selected for further analysis. For these regions, mean species richness (± SE) was calculated for (i) all species from all families, (ii) Termitidae (197 spp.), (iii) Amitermes spp. (Termitidae, 58 spp.), (iv) all families excluding Amitermes spp. (139 spp.), (v) Termopsidae (5 spp.), (vi) Kalotermitidae (32 spp.) and (vii) Rhinotermitidae (25 spp.). In addition, we compared the Atlas data with species richness for five regions, across a comparable range of latitudes, based on the pooled species richness of described and un-described species given in community studies. No group of termites showed a consistent decline in species richness from tropical to temperate latitudes for either data set. The Atlas data showed similar total species richness from the tropics to the mediterranean southwest, before declining to lowest species richness at the highest latitudes. Species richness of Amitermes spp. and Rhinotermitidae was highest in the southwest. Termopsidae and Kalotermitidae showed no latitudinal pattern in species richness. Community studies showed highest and lowest total species richness in the southwest and at the highest latitudes (south-coastal Western Australia), respectively, and similar species richness from the tropics to arid central Australia. Species richness of. Amitermes spp. was highest in the southwest (31 spp.). Kalotermitidae and Rhinotermitidae showed no clear latitudinal pattern. The latitudinal patterns of species richness for the Australian termites is consistent with that for the Australian vertebrates and ants in that they differ from patterns established for these taxa on other continents.     

Patterns of composition and species richness of crustaceans and aquatic insects along environmental gradients in Mediterranean water bodies

Differences in the dynamics of ecological processes between Mediterranean and colder temperate aquatic systems could imply different patterns in faunal communities in terms of composition and biodiversity (i.e. species richness and rarity). In order to identify some of these patterns the crustacean and aquatic insect composition and biodiversity of four water body types, classified according to their salinity and water permanence, were compared. Moreover, the relationships between species richness and water, pond and landscape variables were analysed. A total number of 91 water bodies located throughout Catalunya (NE Iberian Peninsula) were sampled. Three species assemblages were observed: one for permanent freshwaters, another for temporary freshwaters, and a third one for saline waters (SW), since permanent and temporary saline water bodies had similar composition. Differences in salinity were associated with proportion of crustaceans versus insects and with singularity. Thus, saline ponds had a higher proportion of crustaceans, and lower values of singularity. Conductivity was significantly related to total (crustaceans plus insects) richness, and also related to insect richness. The main difference between the models obtained for crustacean species richness and insect species richness is the significance of landscape variables in the latter, and this fact could be related to the different dispersion types of these two faunal groups: active for insects versus passive for crustaceans. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: R. Céréghino, J. Biggs, B. Oertli & S. Declerck The ecology of European ponds: defining the characteristics of a neglected freshwater habitat     

Breeding bird species richness in Taiwan: distribution on gradients of elevation, primary productivity and urbanization

Aim To examine the richness of breeding bird species in relation to elevation, primary productivity and urbanization. Location The island of Taiwan (120°–122° E, 22°–25° N). Methods We arranged bird species richness (BSR) data from 288 bird censuses undertaken in Taiwan into a 2 × 2 km quadrat system and calculated average values of elevation, primary productivity [surrogated by normalized difference vegetation index (NDVI)], and urbanization (surrogated by road density and percentage of built area) for each 2 × 2 km quadrat. Results Bird species richness showed a hump‐shaped relationship with elevation. It increased with elevation from sea level (10–64 species per 2 × 2 km quadrat), peaked around 2000 m (43–76 species), and then decreased with elevation towards its minimum at the highest elevation. Road density and percentage of built area decreased with elevation, and NDVI showed a hump‐shaped relationship with elevation and inverse relationships with road density and percentage of built area. BSR increased with NDVI and decreased with road density and percentage of built area. Linear and cubic terms of elevation together explained 31.3% of the variance in BSR, and road density explained additional 3.4%. The explanatory power of NDVI on BSR was insignificant after the effects of elevation and road density had been justified. Main conclusions We argue that urbanization plays an important role in the BSR of Taiwan. Urbanization might indirectly decrease BSR through decreasing primary productivity and therefore change the hypothetical inverse relationship between BSR and elevation into a hump‐shaped relationship. We also propose a time hypothesis that the biotic communities in the mid‐elevation zone of Taiwan had relatively longer periods of existence during the Pleistocene glacial cycles, which might be one underlying process of the observed hump‐shaped relationship between species diversity and elevation.     

An empirical test of the mass effect determinant of species richness

The mass effect determinant of species richness proposed by Shmida & Wilson (1985) was empirically tested at the interface of undisturbed, native vegetation with recovering vegetation on a reclaimed surface mine. The observed decrease in native species richness with distance away from the source area was consistent with the predicted pattern based on Shmida & Wilson's (1985) model.Taxonomic nomenclature follows Hitchcock & Cronquist (1983) except genus Artemisia, which follows Beetle & Johnson (1982).This research was supported by Grant No. DEB81-01827 from the National Science Foundation. Neil West and Charles Romesburg aided in the experimental design and analysis. Patricia Johnson assisted in computer programming.     

Statutory protected areas and avian species richness in Britain

An effective portfolio of protected areas should, all else being equal, give rise to positive relationships between the amount of protected land in a region and the numbers of species present. Tests of this prediction are, however, extremely scarce, and most do not control for the potentially confounding effects of environmental factors that influence broad geographic trends in biodiversity. Here, we document the form of the relationship between species richness and coverage by protected areas using the British avifauna as a case study. We contrast relationships that arise for breeding and wintering assemblages, considering both all species collectively and threatened species only. We use spatially explicit multiple regression analyses that take into account environmental factors previously shown to exert a marked influence on avian species richness in Britain (temperature and altitude). Avian species richness and the amount of protected land are consistently positively correlated with each other, and the slopes of these relationships do not differ between assemblages (breeding/wintering and all species/threatened species). Explanatory power is, however, very weak which may be indicative of the ability of conservation measures in the wider landscape to maintain avian species richness, reducing any distinctive influence of protected areas. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.