Patterns of plant species turnover along grazing gradients

Questions: How are plant species distributed along grazing gradients? What is the shape of species richness patterns? How can we test for the existence of potential discontinuities in species turnover pattern? Location: Semi‐deserts in the eastern Caucasus, Azerbaijan, Gobustan district. Methods: We studied the distribution of vascular plant species along transects 900‐m long, perpendicular to five farms, and estimated grazing intensity as current livestock units per distance. We modelled species response curves with Huismann–Olff–Fresco (HOF) models and calculated species turnover by accumulating the first derivatives of all response curves. To test for potential discontinuities in changes of vegetation composition along the grazing gradient, we introduce a new null model based on the individualistic continuum concept that uses permutations of the observed pattern of species responses. Results: Most species show a sigmoidal negative response to grazing intensity, while a few species respond with a unimodal pattern. The monotonic decrease in species richness with increasing grazing intensity marks a process of overgrazing that leads to the complete extirpation of plant species. Although the species turnover pattern shows a clear peak, it does not deviate significantly from the null model of individualistic continuous changes. Conclusions: Our approach offers a method for differentiating between transition zones and continuous shifts in species composition along ecological gradients. It also provides a valuable tool for rangeland management to test state‐and‐transition concepts and gives deeper insights into ecological processes affected by grazing.     

Spatiotemporal changes of beetle communities across a tree diversity gradient

Aim Plant and arthropod diversity are often related, but data on the role of mature tree diversity on canopy insect communities are fragmentary. We compare species richness of canopy beetles across a tree diversity gradient ranging from mono‐dominant beech to mixed stands within a deciduous forest, and analyse community composition changes across space and time. Location Germany’s largest exclusively deciduous forest, the Hainich National Park (Thuringia). Methods We used flight interception traps to assess the beetle fauna of various tree species, and applied additive partitioning to examine spatiotemporal patterns of diversity. Results Species richness of beetle communities increased across the tree diversity gradient from 99 to 181 species per forest stand. Intra‐ and interspecific spatial turnover among trees contributed more than temporal turnover among months to the total γ‐beetle diversity of the sampled stands. However, due to parallel increases in the number of habitat generalists and the number of species in each feeding guild (herbivores, predators and fungivores), no proportional changes in community composition could be observed. If only beech trees were analysed across the gradient, patterns were similar but temporal (monthly) species turnover was higher compared to spatial turnover among trees and not related to tree diversity. Main conclusions The changes in species richness and community composition across the gradient can be explained by habitat heterogeneity, which increased with the mix of tree species. We conclude that understanding temporal and spatial species turnover is the key to understanding biodiversity patterns. Mono‐dominant beech stands are insufficient to conserve fully the regional species richness of the remaining semi‐natural deciduous forest habitats in Central Europe, and analysing beech alone would have resulted in the misleading conclusion that temporal (monthly) turnover contributes more to beetle diversity than spatial turnover among different tree species or tree individuals.     

Turning Up the Heat on a Hotspot: DNA Barcodes Reveal 80% More Species of Geometrid Moths along an Andean Elevational Gradient

We sampled 14,603 geometrid moths along a forested elevational gradient from 1020–3021 m in the southern Ecuadorian Andes, and then employed DNA barcoding to refine decisions on species boundaries initially made by morphology. We compared the results with those from an earlier study on the same but slightly shorter gradient that relied solely on morphological criteria to discriminate species. The present analysis revealed 1857 putative species, an 80% increase in species richness from the earlier study that detected only 1010 species. Measures of species richness and diversity that are less dependent on sample size were more than twice as high as in the earlier study, even when analysis was restricted to an identical elevational range. The estimated total number of geometrid species (new dataset) in the sampled area is 2350. Species richness at single sites was 32–43% higher, and the beta diversity component rose by 43–51%. These impacts of DNA barcoding on measures of richness reflect its capacity to reveal cryptic species that were overlooked in the first study. The overall results confirmed unique diversity patterns reported in the first investigation. Species diversity was uniformly high along the gradient, declining only slightly above 2800 m. Species turnover also showed little variation along the gradient, reinforcing the lack of evidence for discrete faunal zones. By confirming these major biodiversity patterns, the present study establishes that incomplete species delineation does not necessarily conceal trends of biodiversity along ecological gradients, but it impedes determination of the true magnitude of diversity and species turnover.     

Turnover patterns in fish versus macroinvertebrates — implications for conservation planning

Spatial patterns in taxonomic richness and turnover for fish and aquatic macroinvertebrates are compared to assess the relative usefulness of each taxonomic group in mapping biodiversity patterns. Fish and aquatic macroinvertebrate species data for sites down the longitudinal axes of nine rivers in four provinces along the eastern side of South Africa were analysed. Fish and aquatic macroinvertebrate data from previous studies and recent river surveys were used in analyses of species richness and turnover. Fish proved to be not useful for mapping biodiversity patterns, as measured by turnover, whereas aquatic macroinvertebrate species patterns exhibited predictable patterns of turnover with downstream distance. Average turnover rates could be decomposed into turnover of common (‘core’) species, which were accelerated by presence of rare and narrow-range species. Disruptions to the river continuum impacted on the rate of turnover. Consistent with other research on South African rivers, aquatic macroinvertebrate communities could be grouped into upland versus lowland assemblages, and also be defined by longitudinal zones. Fish biodiversity patterns should be viewed at a riverscape scale, whereas macroinvertebrate patterns are more easily discerned at a segment-reach scale, and applied to reflect connectivity and environmental gradients respectively.     

Rodent endemism,turnover and biogeographical transitions on elevation gradients in the northwestern Argentinian Andes

The aim of this research is to relate patterns of endemism and turnover along a local elevation gradient in northwest Argentina with continental biogeographical transitions. Specimen based records constituted the principal source of information to infer rodent distribution along the elevation gradient. I assessed elevational variation of richness, endemism and turnover by means of non-linear regression analysis. Then I identified five distributional patterns based on the overlap of species geographic range. Their frequency along elevation was used to validate biogeographical boundaries inferred by turnover rates. Eleven species out of 37 (30%) are endemic to the study area. Species richness and endemism were hump-shaped. The rate of endemism reached its maximum value at the upper limit of the forest (2500 m). By contrast, species turnover was U-shaped, with a small peak at 1500 m and a maximum at 3500 m. The species’ geographic range patterns were not randomly distributed along elevation but agglomerated at specific elevation. Species turnover and chorological analysis suggest two biogeographical boundaries, a weaker at 1500 m and a stronger at 3500 m. The 1500 m boundary marks the transition from assemblages dominated by Lowland-widespread fauna at lower elevation to Montane (Andean eastern slopes) species at middle elevation. This boundary is characterized by moderate species turnover and high species richness. The strong turnover rate at 3500 and the dominance of highland Andean and Andean-Patagonian species above this elevation suggest the occurrence of the transition between the Neotropical and Andean regions; which is characterised by an almost complete species replacement.     

Seasonal Change of Species Diversity Patterns of Non‐volant Small Mammals along Three Subtropical Elevational Gradients

Our understanding of geographic patterns of species diversity and the underlying mechanisms is increasing rapidly, whereas the temporal variation in these patterns remains poorly understood. We examined the seasonal species richness and species turnover patterns of non‐volant small mammals along three subtropical elevational gradients in southwest China. Small mammal diversity was surveyed in two seasons (early wet season and late wet season) using a standardized sampling protocol. The comparison of species richness patterns between two seasons indicated a temporal component in magnitude and shape, with species richness at high elevations clearly increased during the late wet season. Species richness demonstrated weak correlations with modelled temperature and precipitation. The elevational pattern of species turnover measured by Chao‐Sørenson similarity index also changed seasonally, even though the temporal pattern varied with scale. Species turnover between neighboring elevations at high elevations was slower in the late wet season. Meanwhile, there was an acceleration of species turnover along the whole range of the gradient. The seasonal change in species diversity patterns may be due to population‐level increases in abundance and elevational migration, whereas seasonal variation in factors other than temperature and precipitation may play a greater role in driving seasonal diversity patterns. Our study strongly supports the seasonality in elevational patterns of small mammal diversity in subtropical montane forests. Thus it is recommended that subsequent field surveys consider temporal sampling replicate for elevational diversity studies.     

Mountain summer farms in Røldal,western Norway –; vegetation classification and patterns in species turnover and richness

This paper discusses vegetation types and diversity patterns in relation to environment and land-use at summer farms, a characteristic cultural landscape in the Norwegian mountains. Floristic data (189 taxa) were collected in 130 4-m² sample plots within 12 summer farms in Røldal, western Norway. The study was designed to sample as fully as possible the range of floristic, environmental, and land-use conditions. Vegetation types delimited by two-way indicator species analysis were consistent with results from earlier phytosociological studies. Detrended correspondence analysis and canonical correspondence analysis show that rather than being distinct vegetation types, the major floristic variation is structured along a spatial gradient from summer farm to the surrounding heathland vegetation. Species richness (alpha diversity) was modelled against environmental variables by generalized linear modelling and compositional turnover (beta diversity) by canonical correspondence analysis. Most environmental factors made significant contributions, but the spatial distance-to-farm gradient was the best predictor of both species richness and turnover. While summer farms reduce mean species richness at the plot scale, the compositional heterogeneity of the upland landscapes is increased, thereby creating ‘ecological room’ for additional vegetation types and species. Within an overall similarity across scales, soil variables (pH, base saturation, LOI, phosphate and nitrogen) differed considerably in their explanatory power for richness and turnover. A difference between ‘productivity limiting’ factors and ‘environmental sieves’ is proposed as an explanation. Species turnover with altitude is relatively low in grasslands as compared to heaths.     

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.     

Distribution of Vascular Plant Species Richness Along an Elevational Gradient in the Dongling Mountains, Beijing, China

Quantifying spatial patterns of species richness and determining the processes that give rise to these patterns are core problems In blodlveralty theory. The aim of the present paper was to more accurately detect patterns of vascular species richness at different scales along altitudinal gradients in order to further our understanding of biodlverslty patterns and to facilitate studies on relationships between biodiversity and environmental factors. Species richness patterns of total vascular plants species, including trees, shrubs, and herbs, were measured along an altitudinal gradient on one transect on a shady slope in the Dongling Mountains, near Beijing,China. Direct gradient analysis, regression analysis, and geostatistics were applied to describe the spatial patterns of species richness. We found that total vascular species richness did not exhibit a linear pattern of change with altitude, although species groups with different ecological features showed strong elevational patterns different from total species richness. In addition to total vascular plants, analysis of trees, shrubs, and herbs demonstrated remarkable hierarchical structures of species richness with altitude (i.e. patchy structures at small scales and gradients at large scales). Species richness for trees and shrubs had similar spatial characteristics at different scales, but differed from herbs. These results indicated that species groups with similar ecological features exhibit similar biodlveraity patterns with altitude, and studies of biodiversity based on species groups with similar ecological properties or life forms would advance our understanding of variations in species diversity. Furthermore, the gradients or trends appeared to be due mainly to local variations in species richness means with altitude. We also found that the range of spatial scale dependencies of species richness for total vascular plants, trees, shrubs, and herbs was relatively large. Thus, to detect the relationships betweenspecies richness with environmental factors along altitudinal gradients, it was necessary to quantify the scale dependencies of environmental factors in the sampling design or when establishing non-linear models.     

三江并流地区干旱河谷植物物种多样性海拔梯度格局比较

在滇西北三江并流地区典型干旱河谷段, 在怒江、澜沧江和金沙江的东、西坡共设置了6条海拔梯度样带, 通过标准样地的植物群落调查, 分析各条样带植物的物种丰富度、物种更替率的海拔梯度格局, 并比较了地理和植被变量对分布格局的解释。干旱河谷植被带位于海拔3,000 m以下, 以灌丛和灌草丛为主, 其在各河谷的分布上限自西向东依次升高。植物物种丰富度的分布主要与海拔、流域、经纬度和植被带有关, 沿纬度和海拔梯度升高而显著增加的格局主要表现在草本层和灌木层, 灌木物种丰富度还呈现自西向东显著增加的趋势。怒江的灌木和草本种物种丰富度显著高于金沙江和澜沧江, 三条江的乔木种丰富度差异则不显著。森林带的样方草本物种丰富度显著低于灌草丛带样方, 并且还拥有后者没有的乔木种。不同样带的植物物种更替速率呈现了不一致的海拔梯度格局, 但均在样带海拔下部的灌草丛群落与海拔上部森林群落之间的交错带出现峰值。森林-灌草丛植被交错带在怒江样带处于海拔1,900-2,100 m处, 在澜沧江河谷位于海拔2,300-2,400 m, 在金沙江河谷位于海拔2,700-2,900 m。所有海拔样带的森林段或灌草丛段相对于同一样带不同植被段之间的物种更替程度为最小, 不仅小于同一流域不同样带相同植被段之间物种更替率的均值, 更小于所有样带相同植被段之间的更替率均值。在三条河流6条海拔样带的12个植被带段之间的物种更替变化中, 空间隔离因素可以解释34.2%, 而植被类型差异仅能解释不到0.5%。本研究结果显示了环境差异对不同植被类型物种丰富度的首要影响, 和各河流之间的空间隔离对植物群落构建和物种构成的主要作用。     

Small mammal species richness and turnover along elevational gradient in Yulong Mountain,Yunnan, Southwest China

Understanding the species diversity patterns along elevational gradients is critical for biodiversity conservation in mountainous regions. We examined the elevational patterns of species richness and turnover, and evaluated the effects of spatial and environmental factors on nonvolant small mammals (hereafter “small mammal”) predicted a priori by alternative hypotheses (mid‐domain effect [MDE], species–area relationship [SAR], energy, environmental stability, and habitat complexity]) proposed to explain the variation of diversity. We designed a standardized sampling scheme to trap small mammals at ten elevational bands across the entire elevational gradient on Yulong Mountain, southwest China. A total of 1,808 small mammals representing 23 species were trapped. We observed the hump‐shaped distribution pattern of the overall species richness along elevational gradient. Insectivores, rodents, large‐ranged species, and endemic species richness showed the general hump‐shaped pattern but peaked at different elevations, whereas the small‐ranged species and endemic species favored the decreasing richness pattern. The MDE and the energy hypothesis were supported, whereas little support was found for the SAR, the environmental stability hypothesis, and the habitat complexity. However, the primary driver(s) for richness patterns differed among the partitioning groups, with NDVI (the normalized difference vegetation index) and MDE being the most important variables for the total richness pattern. Species turnover for all small mammal groups increased with elevation, and it supported a decrease in community similarity with elevational distance. Our results emphasized for increased conservation efforts in the higher elevation regions of the Yulong Mountain.     

Structure of the species--energy relationship

The relationship between energy availability and species richness (the species-energy relationship) is one of the best documented macroecological phenomena. However, the structure of species distribution along the gradient, the proximate driver of the relationship, is poorly known. Here, using data on the distribution of birds in southern Africa, for which species richness increases linearly with energy availability, we provide an explicit determination of this structure. We show that most species exhibit increasing occupancy towards more productive regions (occurring in more grid cells within a productivity class). However, average reporting rates per species within occupied grid cells, a correlate of local density, do not show a similar increase. The mean range of used energy levels and the mean geographical range size of species in southern Africa decreases along the energy gradient, as most species are present at high productivity levels but only some can extend their ranges towards lower levels. Species turnover among grid cells consequently decreases towards high energy levels. In summary, these patterns support the hypothesis that higher productivity leads to more species by increasing the probability of occurrence of resources that enable the persistence of viable populations, without necessarily affecting local population densities.     

Elevational patterns of functional diversity and trait of Delphinium (Ranunculaceae) in Hengduan Mountains,China

Elevational patterns of trait occurrence and functional diversity provide an important perspective for understanding biodiversity. However, previous studies have mostly examined functional diversity at the community scale. Here, we examined large-scale patterns of trait occurrence and functional diversity in Delphinium along an elevational gradient from 1000 to 5700 m in the Hengduan Mountains, SW China. Elevational distribution and trait data of 102 Delphinium species were compiled to evaluate the patterns of interspecific traits, species richness, and functional diversity. We found that the distribution of species richness showed a unimodal curve that peaked between 3500 and 4000 m; functional diversity and traits showed different patterns along an elevational gradient. The functional diversity increased at a lower rate along an elevation gradient, whereas species richness continued to increase. Species with large ranges and non-endemic species were most affected by geometric constraints. Richness of species endemic to the Hengduan Mountains peaked at higher elevations, likely due to increased speciation and restricted dispersion under alpine conditions. We conclude that the middle elevation region is not only the functionally richest but also the most functionally stable region for Delphinium, which could be insurance against environmental change. Extreme conditions and strong environmental filters in an alpine environment may cause the convergence of species traits, which could relate to reducing nutrient trait investment and increasing reproductive trait investment. We conclude that large-scale studies are consistent with previous studies at the community scale. This may indicate that the relationship between functional diversity and species richness across different scales is the same.     

Determinants of species richness, endemism and turnover in European longhorn beetles

This study assessed the diversity patterns of a large family of beetles, Cerambycidae, in Europe and tested the following hypotheses: 1) richness gradients of this hyperdiverse taxon are driven by water and energy variables; 2) endemism is explained by the same factors, but variation between areas also reflects post‐glacial re‐colonization processes; and 3) faunal composition is determined by the same climatic variables and, therefore, beta diversity (species turnover) is related to richness gradients. Species richness, endemism and beta diversity were modelled using inventories of 37 European territories, built from a database containing the distributions of 609 species. Area, spatial position, and nine topographical and climatic variables were used as predictors in regression and constrained analysis of principal coordinates modelling. Species richness was mostly explained by a temperature gradient, which produced a south‐to‐north decreasing richness gradient. Endemism followed the same pattern, but was also determined by longitudinal variation, peaking in the southwestern and southeastern corners of the continent. Faunal turnover was explained by an important purely spatial pattern and a spatially structured environmental gradient. Thus, contrary to other groups, cerambycid richness was mostly explained by environmental energy, but not by water availability. Endemism was concentrated in the Iberian and Greek peninsulas, but not in Italy. Thus, the latter area may have been the major source of post‐glacial re‐colonization for European longhorn beetles or, otherwise, a poor refuge during glaciations. Turnover patterns were independent of the richness gradient, because northern faunas are nested in southern ones. Turnover, in contrast to richness, was driven by both the independent effects of climate and geographic constraints that might reflect dispersal limitation or stochastic colonization events, suggesting that richness gradients are more environmentally deterministic phenomena than turnover patterns.     

物种丰富度垂直分布格局及影响机制

物种丰富度分布格局是一定地域内物种丰富度沿三维空间的立体分布,包括物种丰富度在经度、纬度和垂直梯度(海拔高度和海水深度)三个维度上的空间分异。近年来物种多样性的垂直分布格局与机制研究得到了生物地理学家和生态学家的重视。物种丰富度的垂直分布格局存在多种类型,但随海拔增加而物种数减少的单调递减模型和中海拔物种丰富度最高的单峰模型较为常见。目前在机制研究中验证较多的是气候稳定性、生物因子(种间相互作用)、能量、生境异质性、干扰、进化时间、物种分化速率、面积、中域效应(mid-domain effect)、生态位保守性(niche conservatism)等假说和机制。物种丰富度的分布格局是多方面因素综合作用的结果;由于地理、地形、气候、地质演化历史、物种库和进化历史、物种分化速率、干扰等差异,在不同地区存在着特别的物种丰富度空间分布格局和机制;处于同一地区的不同类群的物种也因进化扩散历史和生态适应能力不同而呈现多样化的分布格局。因此,对不同地区和类群的物种丰富度格局和机制进行研究应具体分析后才能得到可信结论。     

Forest biodiversity gradients and the human impact in Annapurna Conservation Area, Nepal

Patterns of biodiversity, environment and human impact were studied in 57 sample plots in an 1,178 ha forest area in a rural mountain area of Nepal that is administrated by the Annapurna Conservation Area Project. Alpha-, beta- and gamma-diversity was measured or estimated for six groups of organisms: trees, shrubs, climbers, herbs, polypores and mycorrhizal fungi, and the recorded patterns were correlated with a set of environmental variables. Human impact in terms of fuelwood collection, selective cutting and grazing was found to influence species diversity patterns in all organism groups. Species richness of trees, climbers and polypores at plot level (alpha-diversity) generally responded negatively to human impact, whereas species richness of herbs and shrubs showed a positive relation. Species turnover (beta-diversity), measured as length of the DCA first axis, was significantly correlated to distance to village for all species groups. This indicates that the human impact is very important for the biodiversity patterns in the study area, and that biodiversity connected to undisturbed forest habitats are potentially threatened in the area. The results are discussed in the context of practical conservation. A proposal for future management zones addressing protection of biodiversity without limiting the local use of the forest resource is put forward.     

Spatial patterns along an elevation gradient in high altitude grasslands,Brazil

The Brazilian high altitude grasslands are important biodiversity centers that contribute to human well‐being far outside their boundaries. They host many vulnerable species that may go locally extinct as a result of small changes in the environment. Performing studies on local species distribution is important to guide biodiversity conservation in these areas. This study aimed to characterize a high altitude grassland plant community along an elevation gradient in southeastern Brazil. Simple logistic regression, chi‐squared tests and indicator species analysis were performed to investigate respectively whether: 1) the occurrence of botanical families was related to the altitude; 2) the richness and abundance of life forms varied along the gradient and 3) there were populations associated with specific altitudinal belts. Rubiaceae and Polygalaceae had higher occurrence probability at higher altitudes. Hemicryptophytes and chamaephytes seem to be the most favored life forms under local extreme conditions such as low temperatures and drought. The vegetation spectra varied significantly along the gradient, highlighting the major role of the turnover of habitats on a local scale for the underlying patterns of species distribution. Upper elevations encompassed a high number of indicator species, and further analyses of indicator species of high altitude grasslands will provide clues about adaptability of local species, important to consider in ecosystem management and conservation.     

青藏高原高寒草地植物物种丰富度及其与环境因子和生物量的关系

在青藏高原进行了大范围的群落调查 ,研究高原的两种主要草地群落类型———高寒草甸和高寒草原的植物物种丰富度及其变化。结果表明 :(1)在 5 0个样地 2 5 0个 1m× 1m的样方中 ,共出现 2 6 7种植物 ,其中高寒草甸179种 ,高寒草原 135种。在高寒草甸 ,1m2 样方内物种数最多为 32种 ,最少的仅为 3种 ;在高寒草原 ,物种数最多为 18种 /m2 ,最少的仅为 2种 /m2 。 (2 )物种丰富度随经度和纬度的增加呈增加趋势 ;随海拔的上升呈减少趋势。对物种丰富度与环境因子之间进行逐步回归 ,发现物种丰富度与生长季降水和温暖指数呈显著正相关。 (3)物种丰富度与地上生物量呈显著正相关。     

The potential contribution of vegetation ecology to biodiversity research

The contribution of vegetation ecology to the study of biodiversity depends on better communication between the different research paradigms in ecology. Recent developments in vegetation theory and associated statistical modelling techniques are reviewed for their relevance to biodiversity. Species composition and collective properties such as species richness vary as a continuum in a multi-dimensional environmental space; a concept which needs to be incorporated into biodiversity studies. Different kinds of environmental gradients can be recognised and species responses to them vary. Species response curves of eucalypts to an environmental gradient of mean annual temperature have been shown to exhibit a particular pattern of skewed response curves. Generalised linear modelling (GLM) and generalised additive modelling (GAM) techniques are important tools for biodiversity studies. They have successfully distinguished the contribution of environmental (climatic) and spatial (history and species dispersal ability) variables in determining forest tree composition in New Zealand. Species richness studies are examined at global, regional and local scales. At all scales, direct and resource environmental gradients need to be incorporated into the analysis rather than indirect gradients e.g. latitude which have no direct physiological influence on biota. Evidence indicates that species richness at the regional scale is sensitive to environment, confounding current studies on local/regional species richness relationships. Plant community experiments require designs based on environmental gradients rather than dependent biological properties such as productivity or species richness to avoid confounding the biotic components. Neglect of climatic and other environmental gradients and the concentration on the collective properties of species assemblages has limited recent biodiversity studies. Conservation evaluation could benefit from greater use of the continuum concepts and statistical modelling techniques of vegetation ecology. The future development of ecology will depend on testing the different assumptions of competing research paradigms and a more inclusive synthesis of ecological theory.     

Floristic diversity of an eastern Mediterranean dwarf shrubland: the importance of soil pH

Questions: Does plant species richness and composition of eastern Mediterranean dwarf shrubland (phrygana) correlate with soil pH? How important is the effect of pH on species diversity in relation to other environmental factors in this ecosystem? What is the evolutionary background of the diversity–pH relationship? Location: Western Crete, Greece. Methods: Species composition of vascular plants, soil and other environmental variables were sampled in 100‐m² plots on acidic and basic bedrock in phrygana vegetation. The relationships between species composition and environmental variables (including climate) were tested using canonical correspondence analysis, and relationships between species richness and environment using correlation and regression analyses. Data were analysed separately for different plant functional types based on life form and life span. Results: Although soil pH varied across a narrow range (5.9‐8.1), species composition changed significantly along the pH gradient within all plant functional types. For most functional types, the effect of soil pH on species composition was stronger than that of other environmental variables. Species richness of annuals, geophytes and suffruticose chamaephytes increased with soil pH, while richness of hemicryptophytes and shrubs was not correlated with pH. Conclusions: The results are consistent with the evolutionary species pool hypothesis. High numbers of calcicole annuals, geophytes and suffruticose chamaephytes may be a result of the evolution of these groups on base‐rich dry soils in the Mediterranean climate. In contrast, hemicryptophytes, a life form typical of the temperate zone, evolved on both acidic and basic soils and therefore their species numbers do not respond to soil pH across the narrow range studied. The lack of a relationship between shrub species richness and pH is difficult to explain: it may reflect the more diverse or older origin of Mediterranean woody species and their conservative niches.