Small area and low connectivity constrain the diversity of plant life strategies in temporary ponds

Aim

(i) To determine whether area and connectivity of temporary ponds can predict plant species diversity, and the diversity and abundance of different plant life histories; (ii) To explore whether pond connectivity with the river prior to river regulation predicts better plant diversity patterns than current pond connectivity, suggestive of possible effects of connectivity loss.

Location

Eastern Carpathian Mountains, Romania, Europe.

Methods

We fitted linear and generalized linear models (LM and GLM) to examine whether pond area and current distance from the Olt River predict plant species richness, Shannon diversity and relative cover of different social behaviour types and overall plant species richness and Shannon diversity. Using historical maps, we measured pond distance from the river ca. 60 years before the Olt River was regulated, and we refitted the LM and GLM models using pond area and past distance from the river as independent variables.

Results

Total plant species richness increased with pond area, and it decreased with the distance from the river, but total plant Shannon diversity index was affected, positively, only by pond area. The strength of responses to pond area and connectivity of species richness, Shannon diversity and relative cover varied across the different social behaviour types. Past and current distances between ponds and riverbeds had similar effects on plant diversity, with some evidence for stronger effect of the present connectivity on specialist species Shannon diversity and a weaker effect on disturbance tolerants, generalists and competitors.

Main Conclusions

Pond area and connectivity with the landscape are important predictors of the diversity of plant life history strategies, and therefore, useful tools in pond conservation. Consistent species richness and Shannon diversity responses of wetland specialists to pond area and connectivity make this life history type well suited for monitoring pond condition.     

The origins of species richness in the Hymenoptera: insights from a family-level supertree

Background  

The order Hymenoptera (bees, ants, wasps, sawflies) contains about eight percent of all described species, but no analytical studies have addressed the origins of this richness at family-level or above. To investigate which major subtaxa experienced significant shifts in diversification, we assembled a family-level phylogeny of the Hymenoptera using supertree methods. We used sister-group species-richness comparisons to infer the phylogenetic position of shifts in diversification.     

The complexity of forest borders determines the understorey vegetation

Questions

What are the most important drivers of plant species richness (gamma‐diversity) and species turnover (beta‐diversity) in the field layer of a forest edge? Does the tree and shrub species richness structure and complexity affect the richness of forest and grassland specialist species?

Location

Southeast Sweden.

Methods

We sampled 50 forest edges with different levels of structural complexity in agricultural landscapes. In each border we recorded trees, shrubs and herb layer species in a 50‐m transect parallel with the forest. We investigated species composition and species turnover in relation to the proportions of gaps in the border and the diversity of trees and shrubs.

Results

Total plant species richness in the field layer was mainly explained by the proportion of gaps to areas with full canopy cover and tree diversity. Increasing number of gaps promoted higher diversity of grassland specialist species within the field layer, resulting in open forest borders with the highest overall species richness. Gaps did however have a negative impact on forest species richness. Conversely, increasing forest species richness was positively related to tree diversity, but the number of grassland specialist species was negatively affected by tree diversity.

Conclusions

Managing forest borders, and therefore increasing the area of semi‐open habitats in fragmented agricultural landscapes, provides future opportunities to create a network of suitable habitats for both grassland and deciduous forest specialist species. Such measures therefore have the potential to increase functional connectivity and support dispersal of species in homogeneous forest/agricultural landscapes.     

Patterns of species richness and turnover in endemic amphibians of the Guineo-Congolian rain forest

Aim

The African Guineo-Congolian (GC) region is a global biodiversity hotspot with high species endemism, bioclimatic heterogeneity, complex landscape features, and multiple biogeographic barriers. Bioclimatic and geographic variables influence global patterns of species richness and endemism, but their relative importance varies across taxa and regions and is poorly understood for many faunas. Here, we test the hypothesis that turnover in endemic amphibians of the GC biodiversity hotspot is influenced mainly by the geographic distance between grid cells and secondarily by rainfall- and temperature-related variables.

Location

West and Central Africa.

Major Taxa Studied

Amphibians.

Methods

We compiled species-occurrence records via field sampling, online databases, and taxonomic literature. Our study used 1205 unique georeferenced records of 222 amphibian species endemic to the GC region. Patterns of species richness were mapped onto a grid with a spatial resolution of 0.5° × 0.5°. We estimated weighted endemism and tested whether endemism was higher than the expected species richness (randomization test). We quantified species turnover using generalized dissimilarity modelling to evaluate the processes underlying observed patterns of species richness in GC endemic amphibians. We explored bioregionalization using agglomerative hierarchical clustering based on the unweighted pair group method with arithmetic averages.

Results

We identified seven areas within the lower GC region – forests in Cameroon, Gabon, Southern Nigeria, Equatorial Guinea, Republic of Congo, Democratic Republic of Congo, and Cote d'Ivoire – as having high species richness of endemic amphibians. The randomization test returned four major areas of significant weighted endemism: Nigeria-Cameroon mountains, forest regions of the Democratic Republic of Congo, Cote d'Ivoire, and Ghana. Our analysis revealed five bioregions for amphibian endemism, four of which were located within the lower Guineo-Congolian forest. Species turnover was strongly related to the geographic distance between grid cells; contributing bioclimatic variables included precipitation of the warmest quarter, mean temperature of the wettest quarter, and mean diurnal temperature range.

Main Conclusions

Our results indicate that geographic distance between grid cells is the primary determinant of turnover in GC endemic amphibians, with secondary but significant effects of rainfall- and temperature-related variables. Our study identifies key areas of endemic amphibian richness that could be prioritized for conservation actions.     

Fewer chromosomes,more co-occurring species within plant lineages: A likely effect of local survival and colonization

Premise

Plant lineages differ markedly in species richness globally, regionally, and locally. Differences in whole-genome characteristics (WGCs) such as monoploid chromosome number, genome size, and ploidy level may explain differences in global species richness through speciation or global extinction. However, it is unknown whether WGCs drive species richness within lineages also in a recent, postglacial regional flora or in local plant communities through local extinction or colonization and regional species turnover.

Methods

We tested for relationships between WGCs and richness of angiosperm families across the Netherlands/Germany/Czechia as a region, and within 193,449 local vegetation plots.

Results

Families that are species-rich across the region have lower ploidy levels and small monoploid chromosomes numbers or both (interaction terms), but the relationships disappear after accounting for continental and local richness of families. Families that are species-rich within occupied localities have small numbers of polyploidy and monoploid chromosome numbers or both, independent of their own regional richness and the local richness of all other locally co-occurring species in the plots. Relationships between WGCs and family species-richness persisted after accounting for niche characteristics and life histories.

Conclusions

Families that have few chromosomes, either monoploid or holoploid, succeed in maintaining many species in local communities and across a continent and, as indirect consequence of both, across a region. We suggest evolutionary mechanisms to explain how small chromosome numbers and ploidy levels might decrease rates of local extinction and increase rates of colonization. The genome of a macroevolutionary lineage may ultimately control whether its species can ecologically coexist.

     

Bacterial community assembly and turnover within the intestines of developing zebrafish

Background

The majority of animal associated microorganisms are present in digestive tract communities. These intestinal communities arise from selective pressures of the gut habitats as well as host''s genotype are regarded as an extra ‘organ’ regulate functions that have not evolved wholly on the host. They are functionally essential in providing nourishment, regulating epithelial development, and influencing immunity in the vertebrate host. As vertebrates are born free of microorganisms, what is poorly understood is how intestinal bacterial communities assemble and develop in conjunction with the development of the host.

Methodology/Principal Findings

Set within an ecological framework, we investigated the bacterial community assembly and turnover within the intestinal habitats of developing zebrafish (from larvae to adult animals). Spatial and temporal species-richness relationships and Mantel and partial Mantel tests revealed that turnover was low and that richness and composition was best predicted by time and not intestinal volume (habitat size) or changes in food diet. We also observed that bacterial communities within the zebrafish intestines were deterministically assembled (reflected by the observed low turnover) switching to stochastic assembly in the later stages of zebrafish development.

Conclusions/Significance

This study is of importance as it provides a novel insight into how intestinal bacterial communities assemble in tandem with the host''s development (from early to adult stages). It is our hope that by studying intestinal microbiota of this vertebrate model with such or some more refined approaches in the future could well provide ecological insights for clinical benefit. In addition, this study also adds to our still fledgling knowledge of how spatial and temporal species-richness relationships are shaped and provides further mounting evidence that bacterial community assembly and dynamics are shaped by both deterministic and stochastic considerations.     

Seasonal beta-diversity of dry grassland vegetation: Divergent peaks of above-ground biomass and species richness

Question

Temperate grasslands are known for their high plant diversity and distinct seasonality. However, their intra-annual community dynamics are still largely overlooked by ecologists. Therefore, we explored the seasonal alpha- and beta-diversity patterns of vascular plants and their relationships to above-ground biomass in a rocky steppe (Festucion valesiacae).

Location

Pavlov Hills, SE Czech Republic.

Methods

For one year, we monitored the plant community of the rocky steppe at monthly intervals in 42 permanent plots of 0.25 m². We examined seasonal changes in above-ground biomass (estimated from the cover and height of living plant parts) and seasonal beta-diversity, which we partitioned into turnover and nestedness components and their quantitative counterparts: balanced changes and abundance gradients.

Results

We identified a pronounced seasonal pattern of above-ground biomass, species richness and composition. Total above-ground biomass was highest in June (summer), with a peak representing only 60% of total annual production (sum of individual species' maxima). However, the observed peak in species richness occurred in March (early spring), with 80% of the total species number recorded throughout the year. Accordingly, nestedness and abundance gradient patterns differed in the spring months, while seasonal turnover and balanced changes in abundance were generally congruent. Annual, short-lived, and perennial species exhibited different seasonal patterns of species richness and biomass production, although a sharp increase in biomass and a peak in species richness in spring were universal across the community.

Conclusions

Seasonal climatic constraints on plant growth are key determinants of primary production dynamics. Plants adapt to these constraints by adjusting their life cycles in different ways. In dry grasslands, the complexity of plant responses to climatic seasonality can result in seasonal beta-diversity patterns with divergent peaks in biomass and species richness.     

Local Renyi entropic profiles of DNA sequences

Background  

In a recent report the authors presented a new measure of continuous entropy for DNA sequences, which allows the estimation of their randomness level. The definition therein explored was based on the Rényi entropy of probability density estimation (pdf) using the Parzen's window method and applied to Chaos Game Representation/Universal Sequence Maps (CGR/USM). Subsequent work proposed a fractal pdf kernel as a more exact solution for the iterated map representation. This report extends the concepts of continuous entropy by defining DNA sequence entropic profiles using the new pdf estimations to refine the density estimation of motifs.     

Disentangling the historical routes to community assembly in the global epicentre of biodiversity

Aim

The exceptional turnover in biota with elevation and number of species coexisting at any elevation makes tropical mountains hotspots of biodiversity. However, understanding the historical processes through which species arising in geographical isolation (i.e. allopatry) assemble along the same mountain slope (i.e. sympatry) remains a major challenge. Multiple models have been proposed including (1) the sorting of already elevationally divergent species, (2) the displacement of elevation upon secondary contact, potentially followed by convergence, or (3) elevational conservatism, in which ancestral elevational ranges are retained. However, the relative contribution of these processes to generating patterns of elevational overlap and turnover is unknown.

Location

Tropical mountains of Central- and South-America.

Time Period

The last 12 myr.

Major Taxa Studied

Birds.

Methods

We collate a dataset of 165 avian sister pairs containing estimates of phylogenetic age, geographical and regional elevational range overlap. We develop a framework based on continuous-time Markov models to infer the relative frequency of different historical pathways in explaining present-day overlap and turnover of sympatric species along elevational gradients.

Results

We show that turnover of closely related bird species across elevation can predominantly be explained by displacement of elevation ranges upon contact (81%) rather than elevational divergence in allopatry (19%). In contrast, overlap along elevation gradients is primarily (88%) explained by conservatism of elevational ranges rather than displacement followed by elevational expansion (12%).

Main Conclusions

Bird communities across elevation gradients are assembled through a mix of processes, including the sorting, displacement and conservatism of species elevation ranges. The dominant role of conservatism in explaining co-occurrence of species on mountain slopes rejects more complex scenarios requiring displacement followed by expansion. The ability of closely related species to coexist without elevational divergence provides a direct and faster pathway to sympatry and helps explain the exceptional species richness of tropical mountains.     

Intensity-dependent effects of cattle and sheep grazing in sand grasslands — Does livestock type really matter?

Aims

By analysing cattle- and sheep-grazed sand grasslands, we tested the following hypotheses: (i) livestock type has a stronger effect on the vegetation characteristics than grazing intensity; (ii) sheep grazing results in lower biomass and species and functional diversity than cattle grazing, regardless of intensity; and (iii) increased grazing intensity causes a shift of the trait composition in grasslands.

Location

Sand grasslands in the Nyírség region, East Hungary.

Methods

We selected 26 sand grassland sites grazed by cattle or sheep and classified them into four intensity levels. Vegetation composition was surveyed in 2 m × 2 m plots. We harvested the above-ground biomass from 20 cm × 20 cm plots; then dried and sorted it to live biomass, litter, moss, and lichen. We compared Rao dissimilarity index, species richness, Shannon diversity, evenness, and the community-weighted means of nine vegetative and generative traits along a grazing intensity gradient. We calculated functional richness, evenness, and divergence for comparison.

Results

We found that some diversity metrics and community-weighted means of most studied traits were significantly affected by grazing intensity. Several characteristics were also affected by the interaction of grazing intensity and livestock type, but none of the studied characteristics was affected by livestock type in itself. Increasing Rao dissimilarity index peaking at the fourth grazing intensity level was detected, but for other multitrait indices, no such changes were proven, except for functional divergence, which was the lowest at the first intensity level. Graminoid, forb, and litter biomass were significantly affected by intensity, but none of the biomass fractions was affected by livestock type.

Conclusions

We suggest that for the management of sand grasslands, grazing intensity should be carefully adjusted, considering not only livestock units per hectare. For practical recommendations, well-defined, long-term experiments studying different livestock and habitat types along an intensity gradient would be essential.     

Different Assembly Processes Drive Shifts in Species and Functional Composition in Experimental Grasslands Varying in Sown Diversity and Community History

Background

The prevalence of different biotic processes (limiting similarity, weaker competitor exclusion) and historical contingency due to priority effects are in the focus of ongoing discussions about community assembly and non-random functional trait distributions.

Methodology/Principal Findings

We experimentally manipulated assembly history in a grassland biodiversity experiment (Jena Experiment) by applying two factorially crossed split-plot treatments to all communities: (i) duration of weeding (never weeded since sowing or cessation of weeding after 3 or 6 years); (ii) seed addition (control vs. seed addition 4 years after sowing). Spontaneous colonization of new species in the control treatment without seed addition increased realized species richness and functional richness (FRic), indicating continuously denser packing of niches. Seed addition resulted in forced colonization and increased realized species richness, FRic, functional evenness (FEve) and functional divergence (FDiv), i.e. higher abundances of species with extreme trait values. Furthermore, the colonization of new species led to a decline in FEve through time, suggesting that weaker competitors were reduced in abundance or excluded. Communities with higher initial species richness or with longer time since cessation of weeding were more restricted in the entry of new species and showed smaller increases in FRic after seed addition than other communities. The two assembly-history treatments caused a divergence of species compositions within communities originally established with the same species. Communities originally established with different species converged in species richness and functional trait composition over time, but remained more distinct in species composition.

Conclusions/Significance

Contrasting biotic processes (limiting similarity, weaker competitor exclusion) increase functional convergence between communities initially established with different species. Historical contingency with regard to realized species compositions could not be eradicated by cessation of weeding or forced colonization and was still detectable 5 years after application of these treatments, providing evidence for the role of priority effects in community assembly.     

Comparing alpha-diversity between plants and birds in a remnant wetland: evidence for a threshold and implication for management

In a Mediterranean patchy wetland of central Italy, we analyzed the relationship between the number of bird species, expressed in terms of bird alpha diversity, and plant alpha diversity, expressed as Hill number. This number (the exponential of the Shannon entropy) is considered one of the most strong and reliable indexes of alpha-diversity, synthesizing the information on evenness, richness and diversity in one single metric. We observed a progressive increase of the mean values of bird alpha diversity when plant alpha diversity increases along Hill number. Bird alpha diversity shows an abrupt increase between the first and the second of four categories of plant alpha diversity (0–1, >1–2, >2–3, >3), indicating a threshold response in all the groups considered (breeding, wintering and total bird assemblages). This marked decline of bird species richness at around 1 in the Hill index should represent an alarm for managers: wetland sites at or below this level of plant alpha diversity are likely to be experiencing a drastic decrease in bird species richness, both in spring (breeding birds) and in winter (wintering birds).     

Spatial variability in woody species richness along altitudinal gradient in a lowland-dryland site, Lokapel Turkana, Kenya

The woody species richness patterns in three 2–4 km long transects, approximately 1–3 km apart in a lowland (600–700 m) dryland around Lokapel in Turkana northern Kenya was analyzed in 2003 at 200–500 m intervals using the Point-Centred Quarter (PCQ) method involving 51 observation points. Transect 1 and 2 were set along ephemeral runoff channels locally known as lagga with wet season flow westwards from the Lokapel Hills to the Turkwell River. Transect 3 was a cross-cutting profile dissecting the area initially downhill from the Lokapel Hills and later gently uphill eastwards towards Lokichar. The altitude at each of the 51 observation sites was recorded using a GPS and the woody species identified through local knowledge and taxonomic aids. The results showed that the overall integrated altitudinal gradient for the three transects was approximately 100 m. A total of 43 species of trees and shrubs were identified. The Shannon index showed that Transect 2 had the highest diversity of woody species followed by Transect 1 and Transect 3 while the Sorensen’s index indicated qualitative dissimilarity between all the transects. The results of regression analysis indicated that woody species richness increased linearly with elevation in only one transect but regression analysis of height of woody plants and altitude indicated that only about 20% of the variation in the height of woody plants was accountable by altitude. The spatial analysis of woody species-richness and altitudinal gradient showed a dual peak pattern with the main richness peak in low lying areas below 700 m which was mainly within or close to the riparian floodplain environment of the Turkwell River. A minor richness peak was also identified in higher lying areas around the Lokapel hills. The species richness pattern was similar to the hump-shaped altitudinal species-richness pattern which has been recorded widely around the world but mainly in large-scale studies.     

Turnover and nestedness in subtropical dung beetle assemblages along an elevational gradient

Aim

We investigated changes in dung beetle β‐diversity components along a subtropical elevational gradient, to test whether turnover or nestedness‐related processes drive the dissimilarity of assemblages at spatial and temporal scales.

Location

An elevational gradient (200–1,600 m a.s.l.) of the Atlantic Forest in southern Brazil.

Methods

We investigated the extent to which β‐diversity varied along the elevational gradient (six elevations) at both spatial (among sites at different elevations) and temporal (different months at the same site) scales. We compared both the turnover and nestedness‐related dissimilarity of species and genera using multiple‐site or multiple‐month measures and tested whether these measurements were different from random expectations.

Results

A mid‐elevation peak in species richness along the elevational gradient was observed, and the lowest richness occurred at the highest elevations. We found two different groups of species, lowland and highland species, with a mixing of groups at intermediate elevations. The turnover component of β‐diversity was significantly higher for both spatial (i.e. elevational) and temporal changes in species composition. However, when the data for genera by site were considered, the elevational turnover value decreased in relative importance. Nestedness‐related processes are more important for temporal dissimilarity patterns at higher elevation sites.

Main conclusions

Spatial and temporal turnover of dung beetle species is the most important component of β‐diversity along the elevational gradient. High‐elevation assemblages are not subsets of assemblages that inhabit lower elevations, but this relationship ceases when β‐diversity is measured at the generic level. Environmental changes across elevations may be the cause of the differential establishment of distinctive species, but these species typically belong to the same higher taxonomic rank. Conservation strategies should consider elevational gradients in case‐specific scenarios as they may contain distinct species assemblages in lowlands vs. highlands.

     

Bioindicators of edge effects within Atlantic Forest remnants: Conservation implications in a threatened biodiversity hotspot

Aim

Deforestation of the Atlantic Forest of eastern Paraguay has been recent but extensive, resulting in a fragmented landscape highly influenced by forest edges. We examined edge effects on multiple dimensions of small mammalian diversity.

Location

Forest fragments of eastern Paraguayan Atlantic Forest.

Methods

We trapped small mammal species at different distances from the forest edge (DTE) in reserves and estimated multiple dimensions of diversity per site. Similarity analysis identified species clusters that best described the patterns of diversity across reserves. Multivariate ordination and linear mixed models were used to determine the influence of DTE on various dimensions of small mammal diversity.

Results

There was an increase in richness and abundance along a DTE gradient, and remnants with higher edge:area ratios showed higher richness and abundance, independent of remnant size. Species at edges were generalists, open-habitat species or exotic species (spillover effect). We found higher phylogenetic diversity and functional richness and divergence towards forest edges. Spillover of non-forest and invasive species best explained richness, generalist forest species best explained total abundance, abundance of Hylaeamys megacephalus best explained diversity and evenness metrics and the presence of Marmosa paraguayana best explained various phylogenetic diversity models. None of the models that included megafauna or social factors were shown to be important in explaining patterns as a function of DTE.

Main Conclusions

We found strong support for a spillover effect and mixed support for complementary resource use and enhanced habitat resources associated with ecotones. Generalists characterized edge assemblages but not all generalists were equivalent. Edges showed more phylogenetically and functionally distinct assemblages than the interior of remnants. There was a conservation of functional diversity; however, open-habitat species, habitat generalists and exotic species boosted diversity near forest edges. Mechanisms governing diversity along forest edges are complex; disentangling those mechanisms necessitates the use of multiple dimensions of diversity.     

Ancient origin of high taxonomic richness among insects

Insects are a hyper-diverse group, comprising nearly three-quarters of all named animal species on the Earth, but the environmental drivers of their richness and the roles of ecological interactions and evolutionary innovations remain unclear. Previous studies have argued that family-level insect richness increased continuously over the evolutionary history of the group, but inclusion of extant family records artificially inflated the relative richness of younger time intervals. Here we apply sampling-standardization methods to a species-level database of fossil insect occurrences, removing biases present in previous richness curves. We show that insect family-richness peaked 125 Ma and that Recent values are only 1.5–3 times as high as the Late Palaeozoic. Rarefied species-richness data also tentatively suggest little or no net increase in richness over the past 125 Myr. The Cretaceous peak in family richness was coincident with major radiations within extant groups but occurred prior to extinctions within more basal groups. Those extinctions may in part be linked to mid-Cretaceous floral turnover following the evolution of flowering plants. Negligible net richness change over the past 125 Myr implies that major radiations within extant groups were offset by reduced richness within groups that are now relict or extinct.     

It's a long way to the top: Plant species diversity in the transition from managed to old‐growth forests

Questions

Do vascular plant species richness and beta‐diversity differ between managed and structurally complex unmanaged stands? To what extent do species richness and beta‐diversity relate to forest structural attributes and heterogeneity?

Location

Five national parks in central and southern Italy.

Methods

We sampled vascular plant species composition and forest structural attributes in eight unmanaged temperate mesic forest stands dominated or co‐dominated by beech, and in eight comparison stands managed as high forests with similar environmental features. We compared plant species richness, composition and beta‐diversity across pairs of stands (unmanaged vs managed) using GLMM s. Beta‐diversity was quantified both at the scale of each pair of stands using plot‐to‐plot dissimilarity matrices (species turnover), and across the whole data set, considering the distance in the multivariate species space of individual plots from their centroid within the same stand (compositional heterogeneity). We modelled the relationship between species diversity (richness and beta‐diversity) and forest structural heterogeneity and individual structural variables using GLMM s and multiple regression on distance matrices.

Results

Species composition differed significantly between managed and unmanaged stands, but not richness and beta‐diversity. We found weak evidence that plant species richness increased with increasing levels of structural heterogeneity and canopy diversification. At the scale of individual stands, species turnover was explained by different variables in distinct stands, with variables related to deadwood quantity and quality being selected most often. We did not find support for the hypothesis that compositional heterogeneity varies as a function of forest structural characteristics at the scale of the whole data set.

Conclusions

Structurally complex unmanaged stands have a distinct herb layer species composition from that of mature stands in similar environmental conditions. Nevertheless, we did not find significantly higher levels of vascular plant species richness and beta‐diversity in unmanaged stands. Beta‐diversity was related to patterns of deadwood accumulation, while for species richness the evidence that it increases with increasing levels of canopy diversification was weak. These results suggest that emulating natural disturbance, and favouring deadwood accumulation and canopy diversification may benefit some, but not all, facets of plant species diversity in Apennine beech forests.

     

How gradients of climate and soil determine Mediterranean annual-rich dryland vegetation

Questions

Dryland annual plant communities constitute the most species-rich small-scale vegetation in the Mediterranean. Nevertheless, the composition and diversity of these units and the factors controlling their variation are still insufficiently understood. Therefore, we investigated species composition and richness patterns in relation to important environmental gradients provided by climate and soil.

Location

Central Crete, Greece.

Methods

The study is based on 82 plots of 4 m² sampled at altitudes between 11 and 1400 m a.s.l. We conducted vegetation relevés and soil analyses. We used generalised additive models to model species richness and community characteristics along the studied gradients. We then performed distance-based redundancy analysis to determine the main environmental factors influencing species composition. To determine species of diagnostic value for bedrock types, we applied an indicator species analysis. Correlation tests were used to test the performance of the South Aegean Plant Indicator Values on our dataset.

Results

We recorded 347 taxa (species and subspecies) of 43 plant families, and mean species numbers of 47.2 ± 12.5 per plot. While overall species richness varied only slightly along the analysed environmental gradients, significant changes were observed for relative proportions of species from different life forms and families. Soil pH and elevation had the highest influence on the variation in species composition (23.3% explained). We found 22 species indicative of calcareous rock and 24 species indicative of lime-deficient rock types. The South Aegean Plant Indicator Values were relatively strongly correlated with environmental variables.

Conclusions

Results indicate considerable species turnover both along climatic (elevation) and soil gradients, highlighting the special importance of soil pH. The data provided by our study are expected to supply relevant ecological background information for a pending classification of East Mediterranean annual-rich vegetation.     

Biodiversity assessment in incomplete inventories: leaf litter ant communities in several types of Bornean rain forest

Biodiversity assessment of tropical taxa is hampered by their tremendous richness, which leads to large numbers of singletons and incomplete inventories in survey studies. Species estimators can be used for assessment of alpha diversity, but calculation of beta diversity is hampered by pseudo-turnover of species in undersampled plots. To assess the impact of unseen species, we investigated different methods, including an unbiased estimator of Shannon beta diversity that was compared to biased calculations. We studied alpha and beta diversity of a diverse ground ant assemblage from the Southeast Asian island of Borneo in different types of tropical forest: diperocarp forest, alluvial forest, limestone forest and heath forests. Forests varied in plant composition, geology, flooding regimes and other environmental parameters. We tested whether forest types differed in species composition and if species turnover was a function of the distance between plots at different spatial scales. As pseudo-turnover may bias beta diversity we hypothesized a large effect of unseen species reducing beta diversity. We sampled 206 ant species (25% singletons) from ten subfamilies and 55 genera. Diversity partitioning among the four forest types revealed that whereas alpha species richness and alpha Shannon diversity were significantly smaller than expected, beta-diversity for both measurements was significantly higher than expected by chance. This result was confirmed when we used the unbiased estimation of Shannon diversity: while alpha diversity was much higher, beta diversity differed only slightly from biased calculations. Beta diversity as measured with the Chao-Sørensen or Morisita-Horn Index correlated with distance between transects and between sample points, indicating a distance decay of similarity between communities. We conclude that habitat heterogeneity has a high influence on ant diversity and species turnover in tropical sites and that unseen species may have only little impact on calculation of Shannon beta diversity when sampling effort has been high.     

Biodiversity patterns of vascular plant species in mountain vegetation in the Faroe Islands

Biodiversity patterns of vascular plant species were studied along altitudinal gradients in the Faroe Islands. Plants were sampled from five different mountains (150–856 m a.s.l.) at 50 m altitudinal intervals. Included in the study were 107 vascular plant species. In order to compare only altitudes with the same number of plots, three different analyses were carried out. One analysis included five mountains from 250 to 750 m a.s.l., one had three mountains from 150 to 750 m a.s.l., and the last one had two mountains from 750 to 850 m a.s.l. The patterns of biodiversity were evaluated on the basis of species richness as the total number of species at each altitudinal interval, as species turnover between altitudes and in relation to the Shannon‐Wiener index. Similar patterns were found for species richness in the three analyses, although richness was higher along the whole transect when five mountains were included. For the Shannon‐Wiener index, only small differences were found among the three analyses. A maximum was seen at 250 m a.s.l. and again at 500 m a.s.l. both in richness and in the Shannon‐Wiener index. Maximum species turnover was found at mid‐altitudes. Total vegetation cover followed the same pattern as richness. In addition to climate, the altitudinal variation of biodiversity may be affected by grazing.