The species pool hypothesis: the necessity to shift emphasis

The species pool of a biological community is determined as a group of species that inhabit some area and potentially can be included in a given community. The species pool hypothesis, i.e. the assumption that the size of species pool strongly influences species richness of local community can be confirmed if there is positive linear relationship between these two variables. The results of hypothesis testing however are not obvious. For example, correlation between local richness and species pool size can be caused by their dependence on the third variable--capacity of environment. It seems that in case of decreasing area occupied by local community the environmental conditions become more important than species pool size. If that is true, the influence of species pool on local species richness is not significant. However one can estimate the degree of unsaturation of species pool on the basis of relationships between the number of species in small locations occupied by similar local communities and their species pool. We think, that study of local and regional species richness should shift the emphasis--from the analysis of species pool influence on local community richness to the estimation of historical, ecological and anthropogenic factors in variation of species pool size. The local species richness should be considered rather as a tool (allowing to compare the species capacity of biological communities), than as an object of such study.     

The influence of scale and species pool on the relationship between vascular plant species richness and cover in an alpine area in Norway

Recent studies emphasise the potential importance of scale and species pool on the humped-back or unimodal relationship between species richness and productivity. We use a classic phytosociological data-set from Rondane, central south Norway, to evaluate the relative importance of these factors in an alpine area. The effect of species pool is assessed using plot scores from a Correspondence Analysis (CA) of the data. Generalised Additive Models (GAM) are used to relate vascular plant species richness to cover of vascular plants, CA plot scores, and plot area in different combinations. Species richness of vascular plants is unimodally related to total vascular plant cover. Plot scores of the first three CA axes (representing the effect of species pool) have a complex relationship with species richness, but explain a large fraction of the total deviance in richness. A humped relationship between richness and cover remains after accounting for CA plot scores in the model, i.e. the relationship is independent of species pool. The results suggest that the relationship between richness and cover changes from one vegetation type to another, as evaluated statistically through the importance of the interaction between cover and CA scores in explaining variation in richness. Plot area also influences the relationship. A unimodal relationship is only evident when small plot sizes are used, whereas a monotonically increasing relationship is found at large plot sizes. Plot area has the strongest effect on the unimodal relationship between richness and cover, whereas vegetation type has only a minor effect on this relationship. This revised version was published online in June 2006 with corrections to the Cover Date.     

Small-scale plant species richness in calcareous grasslands determined by the species pool, community age and shoot density

Variation in small-scale plant species richness was examined between twenty discrete sites of calcareous alvar grasslands in the western Estonian coastal area. This community type is very species rich at the small-scale. The pattern of diversity has often been explained by variation in ecological conditions which makes it possible to avoid competitive exclusion. Here we test an alternative species pool hypothesis. The species pool was defined as the set of species present in a community, but excluding all those species whose presence may be connected with exceptional environmental conditions for this community. Monte Carlo modelling was used to determine the strengths of the relationships between the non-independent variables: richness vs species pool and average shoot density vs richness. Site age was characterized by its elevation above sea level, since there is a continuous land uplift in the study area. Species richness was positively correlated to the size of the species pool, community age and vegetation density. We conclude that historical processes on both the regional and local levels, which determine the arrival of any particular species to a target community, and not environmental heterogeneity, are responsible for the variations in species richness between communities of this type.     

Size of the local species pool determines invasibility of a C4-dominated grassland

The size of the local species pool (i.e., species surrounding a community capable of dispersal into that community) and other dispersal limitations strongly influence native plant community composition. However, the role that the local species pool plays in determining the invasibility of communities by exotic plants remains to be evaluated. We hypothesized that the richness and abundance of exotic species would be greater in C4‐dominated grassland communities if the local species pool included a larger proportion of exotic species. We also predicted that an increase in the exotic species pool would increase the invasibility of sites thought to be resistant to invasion (annually burned grassland). To test these hypotheses, study plots were established within two long‐term (>20 yr) fire experiments at a tallgrass prairie preserve in NE Kansas (USA). Study plots were surrounded by either a small pool of exotic species (small species pool (SSP) plots; six species) or a larger exotic species pool (large species pool (LSP) plots; 18 species). We found that richness and absolute cover of exotic species was significantly (P<0.001) lower (～70 and 90%, respectively) in annually burned compared to unburned plots, regardless of the size of the exotic species pool. As predicted, exotic species richness was higher (P<0.001) for LSP plots (3.9 per 250 m2) than for SSP plots (0.7 per 250 m2); however, absolute cover was unaffected by the size of the exotic species pool. In the absence of fire, plots with a LSP had four times as many exotic species than SSP plots. An increase in the local exotic species pool also increased the invasibility of annually burned grassland. Indeed, richness of exotic plant species in annually burned LSP plots did not differ from unburned plots with a SSP, indicating that a larger pool of exotic species countered the negative effects of fire. These findings have important implications for predicting how the invasion of plant communities may respond to human‐induced global changes, such as habitat fragmentation. Community characteristics or factors such as frequent fires in grasslands may impart resistance to invasions by exotic species in large, intact ecosystems. However, when a large pool of exotic species is present, frequent fire may not be sufficient to limit the invasions of exotic plants in fragmented landscapes.     

The elevational gradient in Andean bird species richness at the local scale: a foothill peak and a high-elevation plateau

A monotonic decline in species richness with increasing elevation has often been considered a general pattern, but recent evidence suggests that the dominant pattern is hump-shaped with maximum richness occurring at some mid-elevation point. To analyse the relationship between species richness and elevation at a local scale we surveyed birds from lowlands to timberline in the Bolivian Andes. We divided the transect into 12 elevational belts of 250 m and standardized species richness in each belt with both individual- and sample-based rarefaction and richness estimation. The empirical data were then correlated to four explanatory variables: 1) area per elevational belt, 2) elevation (also representing ecosystem productivity), 3) a mid-domain effect (MDE) null model of geometrically constrained empirical range sizes, and 4) a hump-shaped model derived empirically for South American birds representing the regional species pool hypothesis. Local species richness peaked at ca 1000 m elevation, declined sharply to ca 1750 m, and then remained roughly constant. Elevation was the best single predictor, accounting for 78–85% of the variance in the empirical data. A multiple regression model with elevation, area, and MDE explained 85–90% of the variance. Monte Carlo simulations showed that the richness peak at 1000 m is the result of an overlap of two distinct avifaunas (lowland and highland) and that the correlation to MDE in the multiple regression was likely spurious. We recommend complementing correlation analyses involving MDE predictions with an examination of the distribution of range midpoints. The steep decline at mid-elevations was mainly due to a rapid loss of lowland species. The high-elevation plateau is striking and unexpected, but has also been found previously. It cannot be explained at present and exemplifies that despite several decades of research elevational gradients are still not well understood.     

The species pool concept applied to forests in a fragmented landscape: dispersal limitation versus habitat limitation

Abstract. The species pool concept has been used as a theoretical framework for understanding local community richness. A significant problem in putting the concept into practice is the lack of methods for determining the size of the species pool. We tested the hypothesis that species composition of recent forests is primarily determined by the species composition of neighbouring older forests against the null‐hypothesis that species are a random sample of the species occurring in the study area. Forest plant species composition of recently established fragments was significantly correlated with species composition in neighbouring older forests (i.e. the local species pool). When older forest within a neighbourhood of 1000m radius is considered, seed dispersal sources can be found for 91% of the flora in the recent forests. For an individual fragment, dispersal is a much more important determinant of species presence than the environment, with an average of 46% of the total pool excluded from local pools by dispersal limitation and only 8% excluded by environmental limitations. The species richness of recent forests is on average 23% of the local species pool. Several hypotheses are proposed for this low percentage, such as asymmetric competition due to the early successional state or the limited colonization period.     

Predicting distributions of species richness and species size in regional floras: Applying the species pool hypothesis to the habitat templet model

The species pool hypothesis is applied here to the interpretation of ‘hump-shaped’ (unimodal) species richness patterns along gradients of both habitat fertility and disturbance level (the habitat templet). A ‘left-wall’ effect analogous to that proposed for the evolution of organismal complexity predicts a right-skewed unimodal distribution of historical habitat commonness on both gradients. According to the species pool hypothesis, therefore, the distribution of opportunity for net species accumulation (speciation minus extinction) should also have a corresponding unimodal central tendency on both habitat gradients. Two assumptions of this hypothesis are illustrated with particular reference to highly fertile, relatively undisturbed habitats: (i) such habitats have been relatively uncommon in space and time, thus providing relatively little historical opportunity for the origination of species with the traits necessary for effective competitive ability under these habitat conditions; and (ii) species that have evolved adaptation to these habitats are relatively large, thus imposing fundamental ‘packing’ limitations on the number of species that can ‘fit’ within such habitats. Based on these assumptions, the species pool hypothesis defines two associated predictions that are both supported by available data: (a) resident species richness will be relatively low in highly fertile, relatively undisturbed contemporary habitats; and (b) species sizes within regional floras should display as a right-skewed unimodal (log-normal) distribution. The latter is supported here by an analysis of data for 2,715 species in the vascular flora of northeastern North America.     

Species richness at the guild level: effects of species pool and local environmental conditions on stream macroinvertebrate communities

1.?A fundamental question in ecology is which factors determine species richness. Here, we studied the relative importance of regional species pool and local environmental characteristics in determining local species richness (LSR). Typically, this question has been studied using whole communities or a certain taxonomic group, although including species with widely varying biological traits in the same analysis may hinder the detection of ecologically meaningful patterns. 2.?We studied the question above for whole stream macroinvertebrate community and within functional feeding guilds. We defined the local scale as a riffle site and the regional scale (i.e. representing the regional species pool) as a stream. Such intermediate-sized regional scale is rarely studied in this context. 3.?We sampled altogether 100 sites, ten riffles (local scale) in each of ten streams (regional scale). We used the local-regional richness regression plots to study the overall effect of regional species pool on LSR. Variation partitioning was used to determine the relative importance of regional species pool and local environmental conditions for species richness. 4.?The local-regional richness relationship was mainly linear, suggesting strong species pool effects. Only one guild showed some signs of curvilinearity. However, variation partitioning showed that local environmental characteristics accounted for a larger fraction of variance in LSR than regional species pool. Also, the relative importance of the fractions differed between the whole community and guilds, as well as among guilds. 5.?This study indicates that the importance of the local and regional processes may vary depending on feeding guild and trophic level. We conclude that both the size of the regional species pool and local habitat characteristics are important in determining LSR of stream macroinvertebrates. Our results are in agreement with recent large-scale studies conducted in highly different study systems and complement the previous findings by showing that the interplay of regional and local factors is also important at intermediate regional scales.     

Species pools and the "hump-back" model of plant species diversity: an empirical analysis at a relevant spatial scale

We investigated the relative roles of productivity, the species pool, and spatial habitat structure in determining local species richness (alpha diversity) of plant communities within a single, well-defined landscape unit, at spatial and ecological scales where the relationship between community productivity and species diversity often assumes a unimodal or "hump-back" form. At high levels of productivity, the decrease-phase of the unimodal model of the diversity-productivity relationship is typically explained as the dynamic outcome of increased competitive exclusion, but it may also be the passive consequence of a small pool of species possessing attributes necessary to competitively survive in high-fertility environments. We conducted statistical analyses of previously collected data to determine whether variations in local richness in the herbaceous vegetation of a Slovakian mountain valley were best explained by habitat productivity itself (which presumably leads to more intense competition) or by the sizes of the relevant community species pools. We also used measures of spatial habitat structure to investigate the extent to which habitat patchiness influenced patterns of species diversity. In the study system, both community biomass and size of the species pools contributed significantly to local species richness, but the positive effect of the species pools was about twice as important as the negative effect of biomass. The combined area of related associations (alliance area), association perimeter, and habitat patch geometry were all closely related to species pool size.     

Ecosystem consequences of the regional species pool

In this study we analyse with an experiment how the spatial area from which the species pool is sampled, i.e. local vs regional, can have functional consequences for ecosystem processes and structure. Particularly, we analysed how major stocks and processes respond to a change in basal productivity and how this depends on the source of the species pool. The experimental microcosms used in this experiment were developed by inoculating nutrient solution with samples of rockpool sediment containing resting stages. Here we show that communities developed from sediment samples of single origin (local communities) exhibit important differences in ecosystem structure and their response to a productivity gradient compared to systems developed from mixed samples of multiple and spatially dispersed origin (regional communities). The difference between local and regional communities was most pronounced for zooplankton which had much lower abundances in the local communities compared to the regional communities resulting in cascading effects on phytoplankton. We demonstrate that the spatial extend from which the species pool is sampled can affect ecosystem structure and functioning comparable in magnitude to changes in the basal productivity of the system. We discuss with a conceptual model how the regional species pool is important for sustaining diversity in functional groups, and how higher trophic levels are more sensitive to changes in the regional species pool.     

Trait-based filtering from the regional species pool into local grassland communities

Aims For plants to establish in a local community from a pool of possible colonizers from the region, it must pass through a series of filters. Which of the filters is most important in this process has been much debated. In this study, we explored how species are filtered from the regional species pool into local communities. The aim was to determine if differences in species abundance and functional traits could explain which species from the regional species pool establish at the local scale and if the filtering differed between grassland communities.Methods This study took place in a cultivated landscape in southeastern Sweden. We estimated plant species abundance in 12 ex-arable field sites and 8 adjacent seminatural grassland sites and in a 100-m radius around the center of each site. We used Monte Carlo simulations to examine if species abundance and functional traits (height, seed mass, clonal abilities, specific leaf area and dispersal method) controlled the filtering of species from the regional pool into local communities.Important findings On average, only 28% of species found in the regional pool established in the ex-arable field sites and 45% in the seminatural grassland sites, indicating that the size of the regional species pool was not limiting local richness. For both grassland types, species abundance in the regional pool was positively correlated with species occurrence at the local scale. We found evidence for both species interaction filtering and dispersal limitation influencing the local assembly. Both local and regional processes were thus influencing the filtering of species from the regional species pool into local communities. In addition, the age of the communities influenced species filtering, indicating that community assembly and the importance of different filters in that process change over succession.     

Dissecting NDVI–species richness relationships in Hawaiian dry forests

Aim A growing body of research has used the normalized difference vegetation index (NDVI) as a proxy for productivity to predict species richness. Yet the mechanisms that produce the relationship between NDVI and species richness remain unclear because of correlated biotic and abiotic factors that influence NDVI. In this study we investigated different biotic and abiotic effects that potentially drive plant species richness–productivity relationships. Location Hawaiian Islands, USA. Methods We quantified woody plant species richness, structure (density, basal area and canopy height), and species composition along a precipitation gradient of 14 Hawaiian dry forest plots. We then used structural equation models combined with 10 years of satellite data to disentangle the effects of precipitation, structure and NDVI‐estimated productivity on species richness. Results Underlying the simple correlation between NDVI and species richness was the indirect effect of precipitation and direct effect of forest structure. The best‐fit model showed there was no direct effect of NDVI on species richness. Main conclusions Our results demonstrate that complex relationships drive simple correlations between species richness and productivity. Considering the mechanisms and underlying factors driving NDVI–species richness relationships could improve predictions of species diversity as satellite measures of productivity have an increasingly important role in habitat mapping, species distribution modelling and predictions for global change.     

Effects of area,environmental status and environmental variation on species richness per unit area in Mediterranean wetlands

Abstract. We propose a mechanistic model to relate α- and γ-diversity to area per se, moisture status and environmental variation (local and total), and explored the effects these abiotic variables have on species richness per unit area (α-diversity) for plant communities in a network of wetland habitats located in a Mediterranean mountainous region of central Spain. In this study, environmental status is measured as actual evapotranspiration (as an expression of energy), slope and soil wetness, and environmental variation refers to slope variation and soil wetness variation. Species richness per unit area was related to soil wetness, soil wetness variation, ground slope and ground slope variation. There were also positive correlations among moisture status and environmental variation variables. There is a joint effect of slope and soil wetness variation in explaining species richness per unit area of these wetland habitats, but area effects and energy are relatively unimportant. We conclude that species richness per unit area of wetland vegetation can be explained by moisture status and local environmental variation, and that habitat area may not have an important effect. Area could affect γ-diversity directly through random sampling and/or indirectly through increasing β-diversity, and energy may be important in areas with larger energy ranges. Complete surveys of environmental status, local and total environmental variation, and their associated species assemblages are needed to explain the processes that give rise to the rule that larger areas have larger species richness.     

The relationship between nutrient availability,shoot biomass and species richness in grassland and wetland communities

The relationship was studied between shoot biomass, nutrient concentration in the soil and number of species per unit area. The study was carried out in two different parts of the Netherlands, the Gelderse Vallei (east of Amersfoort) and the Westbroekse Zodden (northwest of Utrecht). Four series of vegetation and soil samples were taken: one series in grassland and wetland communities, one series in grassland communities, one series in fen communities and one series in only one wetland community. The two series in grassland communities show a negative correlation between shoot biomass and species number and a positive correlation between shoot biomass and nutrient concentration in the soil. The opposite was found in the series in the fen communities: there was a positive correlation between species number and shoot biomass and a negative correlation between shoot biomass and nutrient concentrations. The series of samples that had been taken in only one wetland community showed an optimum curve for the relation between shoot biomass and number of species. It is concluded that in the plant communities studied the species richness per unit area increases with increasing productivity at low production levels (< 400–500 g/m²) and decreases with increasing productivity at higher production levels (> 400–500 g/m²).     

Testing the species pool hypothesis for mire vegetation: exploring the influence of pH specialists and habitat history

The Evolutionary species pool hypothesis (ESPH) predicts that historically more common habitats will be richer in species because they have had greater opportunity for the evolution of suitably adapted species. We explored the relationship between mire species richness and pH, an important environmental variable in mires, in two regions that differ in habitat pH distribution: the West Carpathians and Bulgaria. Mire habitats in both the West Carpathians and Bulgaria demonstrate support for the ESPH prediction that habitats with more common pH values host more species. We also explored the influence of habitat history by examining the distribution of generalists and specialists along gradients of habitat pH, using extensive community-level vegetation data from European mires in these two regions. We found a striking pattern with the distribution of pH-specialists having three distinct peaks in both regions, whereas the total species pool peaked in near neutral pH habitats in both regions. Because the peaks in specialist richness do not correspond to regional pH distribution patterns, we hypothesize that historical explanations may be important, and that habitats currently rich in pH-specialists may have historically acted as pleniglacial refugia for many mire species. Our findings support the general predictions of the ESPH, but further suggest that historical processes such as patterns of glacial refugia, may significantly influence contemporary species distributions and the diversity of plant species in mire habitats.     

Small-scale species richness in forest canopy gaps: the role of niche limitation versus the size of the species pool

Abstract. The form of the relationship between local species richness and the number of species in the surrounding region can be used as a test between competing theories of community structure. For 32 canopy gaps in New Zealand Nothofagus forest, we examined the relationship between the number of vascular plant species in 0.2-m² quadrats within the gap and the species richness of the whole gap. We found no evidence that competition for a limited number of niches placed an upper limit on the number of locally co-occurring species. Rather, the mean number of species in quadrats within canopy gaps increased in direct proportion to gap species richness. This relationship held after we controlled for potentially confounding factors, including variation in forest floor substrate, and gap size, age, shape and orientation. Our results suggest that even over relatively small spatial scales, local species richness can be constrained by the size of the species pool in the immediately surrounding region.     

Island biogeography of bats in Baja California, Mexico: patterns of bat species richness in a near-shore archipelago

Aim We studied the relationship between the size and isolation of islands and bat species richness in a near‐shore archipelago to determine whether communities of vagile mammals conform to predictions of island biogeography theory. We compared patterns of species richness in two subarchipelagos to determine whether area per se or differences in habitat diversity explain variations in bat species richness. Location Islands in the Gulf of California and adjacent coastal habitats on the Baja California peninsula in northwest Mexico. Methods Presence–absence surveys for bats were conducted on 32 islands in the Gulf of California using acoustic and mist‐net surveys. We sampled for bats in coastal habitats of four regions of the Baja peninsula to characterize the source pool of potential colonizing species. We fitted a semi‐log model of species richness and multiple linear regression and used Akaike information criterion model selection to assess the possible influence of log₁₀ area, isolation, and island group (two subarchipelagos) on the species richness of bats. We compared the species richness of bats on islands with greater vegetation densities in the southern gulf (n = 20) with that on drier islands with less vegetation in the northern gulf (n = 12) to investigate the relationship between habitat diversity and the species richness of bats. Results Twelve species of bats were detected on islands in the Gulf of California, and 15 species were detected in coastal habitats on the Baja peninsula. Bat species richness was related to both area and isolation of islands, and was higher in the southern subarchipelago, which has denser vegetation. Log₁₀ area was positively related to bat species richness, which increased by one species for every 5.4‐fold increase in island area. On average, richness declined by one species per 6.25 km increase in isolation from the Baja peninsula. Main conclusions Our results demonstrate that patterns of bat species richness in a near‐shore archipelago are consistent with patterns predicted by the equilibrium theory of island biogeography. Despite their vagility, bats may be more sensitive to moderate levels of isolation than previously expected in near‐shore archipelagos. Differences in vegetation and habitat xericity appear to be associated with richness of bat communities in this desert ecosystem. Although observed patterns of species richness were consistent with those predicted by the equilibrium theory, similar relationships between species richness and size and isolation of islands may arise from patch‐use decision making by individuals (optimal foraging strategies).     

The species richness–biomass relationship in herbaceous plant communities: what difference does the incorporation of root biomass data make?

So far, in all studies on the much-discussed hump-backed relationship between plant community productivity and species richness, productivity has been assessed through plant shoot biomass, i.e. it has been ignored that frequently most of the biomass is produced below ground. We revisited the 27 grassland and forest field-layer communities, studied earlier by Zobel and Liira, to sample root biomass, plant total biomass and root/shoot allocation, and learn how the incorporation of below-ground biomass data would affect the shape of the hump-backed relationship. In order to avoid scaling artefacts we estimated richness as the average count of species per 500 plant ramets (absolute richness). We also included relative richness measures. Relative richness was defined as richness per 500 ramets/size of the actual species pool (the set of species present in the community), relative pool size was defined as size of the actual species pool/size of the regional species pool (the set of species available in the region and capable of growing in the given community).
The biomass-absolute richness relationship was humped, irrespective of the biomass measure used, the hump being most obvious when plant total biomass was used as the independent variable. Evidently, the unimodal richness–productivity curve is not a sampling artefact, as suspected by Oksanen. However, relative richness was not related to community biomass (above-ground, below-ground or total). The hump-backed curve is shaped by the sizes of actual species pools in communities, implying that processes which are responsible for small-scale diversity pattern mainly operate on the community level.
Neither absolute nor relative richness were significantly related to root/shoot allocation. The presumably stronger (asymmetric) shoot competition at greater allocation to shoots appears not to suppress small-scale richness. However, there is a significant relationship between relative pool size and root/shoot allocation. Relatively more species from regional species pools are able to enter and persist in communities with more biomass allocated into roots.     

Fish species richness and incidence patterns in isolated and connected stream pools: effects of pool volume and spatial position

I tested the effects of pool size and spatial position (upstream or downstream) on fish assemblage attributes in isolated and connected pools in an upland Oklahoma stream, United States. I hypothesized that there would be fundamental differences between assemblages in these two pool types due to the presence or absence of colonization opportunities. Analyses were carried out at three ecological scales: (1) the species richness of pool assemblages, (2) the species composition of pool assemblages, and (3) the responses of individual species. There were significant species-volume relationships for isolated and connected pools. However, the relationship was weaker and there were fewer species, on average, in isolated pools. For both pool types, species incidences were significantly nested such that species-poor pools tended to be subsets of species-rich pools, a common pattern that ultimately results from species-specific differences in colonization ability and/or extinction susceptibility. To examine the potential importance of these two processes in nestedness patterns in both pool types, I made the following two assumptions: (1) probability of extinction should decline with increasing pool size, and (2) probability of immigration should decline in an upstream direction (increasing isolation). When ordered by pool volume, only isolated pools were significantly nested suggesting that these assemblages were extinction-driven. When ordered by spatial position, only connected pools were significantly nested (more species downstream) suggesting that differences in species-specific dispersal abilities were important in structuring these assemblages. At the individual-species level, volume was a significant predictor of occurrence for three species in isolated pools. In connected pools, two species showed significant position effects, one species showed a pool volume effect, and one species showed pool volume and position effects. These results demonstrate that pool size and position within a watershed are important determinants of fish species assemblage structure, but their importance varies with the colonization potential of the pools. Isolated pool assemblages are similar to the presumed relaxed faunas of montane forest fragments and land bridge islands, but at much smaller space and time scales. Received: 6 December 1996 / Accepted: 10 December 1996     

The role of historical processes in determining tree species richness in the forests of Western Caucasus

To estimate the role of history in variation of tree species richness in the forests of the Western Caucasus we analyzed correlation between their local richness (S--the mean number of species per 300 m2) and size of actual species pool (N--the number of species per 1 ha). If compared communities are differently distant from the point of evolutionary equilibrium one should expect a significant variation in correlation between S and N (determined with the greater sensitivity of N than S in respect of historical factors). The lower value of N/S corresponds to less saturated level of historically determined species richness. A mean N/S ratio in Japana temperate broadleaved forests (Masaki et al., 1999) provided the basis for analysis. The present tree species richness of the forest communities in the 1 ha plots seem essentially determined by the historical processes. The mountain forest communities of Western Caucasus are characterized on the average with lower saturation level of the actual tree species pool in comparison with the Japan temperate broad-leaved forest communities. On the Western Caucasus the middle mountain beech and coniferous-broadleaved communities (400-1600 m a.s.l.) are characterized with the higher saturation level of the actual species pool in comparison with communities located lower and higher. These results confirm published historical reconstructions, according to which the middle mountain forest communities in the Western Caucasus are older than forests located higher or lower. Present low mountain forests of the southern (to Black Sea) and the northern macroslopes of the Western Caucasus are characterized with similar saturation level of the actual species pool. These data agree with the assumption of Dolukhanov (1980) that low mountain zone of the southern macroslope was not a refuge for tree species in Pleistocene.