Effects of the 2003 heatwave and climatic warming on mollusc communities of the Saône: a large lowland river and of its two main tributaries (France)

The study of the dynamics of mollusc populations of the Saône and its two main tributaries, the Doubs and Ognon, over several years has provided us with the opportunity of highlighting the consequences of climatic warming and especially of the heatwave of 2003 on these organisms. From 1987 to 2003, the mean temperature of the waters of the Saône upstream of Lyon (Couzon) increased by 1.5°C. In addition, the summer of 2003 was the hottest since 1500 at least. We used correspondence analysis to identify structure change in mollusc data dating from September 1996 to December 2004. The results revealed: (1) during the period from September 1996 to July 2003, a significant progressive change in the mollusc community structure of the Saône upstream of Lyon, probably linked to the increase of temperature; (2) from July to August 2003 during the heatwave, a sudden change in the structure of mollusc communities and a significant decrease of species richness and density of gastropods and bivalves. During 2004, mollusc density and particularly that of Pisidium remained dramatically low. Similar observations were performed at four other sites along the Saône and in the lower reaches of its two main tributaries. This suggests that the resilience of the mollusc populations (i.e., the speed with which they return to a predisturbance state) to the heatwave is low. In this way, as different climatic models have predicted an increase in the frequency of summers as hot as that of 2003 during this century, more than half the mollusc species currently inhabiting the potamic area of the Saône, Doubs and Ognon, and probably other large rivers, are probably directly threatened with extinction.     

The rivers Rhine and Meuse in The Netherlands: present state and signs of ecological recovery

The ecosystems of the rivers Rhine and Meuse have suffered drastic environmental changes, for example because of the regulation of the stream bed and the construction of weirs and dams. Furthermore, discharges of industrial and municipal waste water have caused the water quality of these rivers to deteriorate; this problem became acute in the sixties and seventies. Recently some chemical parameters of water quality have improved in the Rhine, and as a consequence some aquatic communities are showing signs of recovery after decades of severe degradation. This paper describes the present state of the aquatic communities in the Dutch part of the rivers Rhine and Meuse, using published observations on plankton, macrophytes, invertebrates, and fish. The sparse information on the food chain in these rivers is summarized. The main channels of the Rhine and Meuse are characterized by a dense plankton that develops rapidly in the nutrient-rich river waters. The stream beds, now dominated by wave-exposed sand and gravel, have a sparse fauna and flora. The river banks, mostly consolidated by blocks of stone, offer a substratum for numerous benthic organisms, particularly now that the water quality has improved. The floodplain waters and old river channels harbour a flora and fauna rich in species. The degree of water exchange with the river is crucial for the ecological development of the river and its backwaters. Today the freshwater tidal reaches of the rivers occupy a very restricted area, and only remnants of the previously abundant vegetation of rushes are found. Losses in the numbers of animal and plant species, notably those specific to rivers, are evident, but over the last 15 years several species have returned. Allochthonous species (exotics), including crustaceans and molluscs, have also settled in the Rhine and Meuse. Fish species characteristic of these rivers, such as river lamprey, sea trout, barbel, and flounder, have recently been observed in appreciable numbers. The Rhine Action Programme provides a framework for the large-scale rehabilitation of the Rhine. Experiments on re-stocking the Rhine with Atlantic salmon and on the ecological rehabilitation of floodplains are being carried out on the assumption that there will be a further reduction of the pollution load. A similar programme is required for the Meuse.     

Un atoposauridé (Crocodylia,mesosuchia) du portlandien de la Meuse (est de la France)

A small crocodilian skull from the lower Portlandian (Cyrena rugosa beds) of Brauvilliers (Meuse) is referred to an atoposaurid of the genus Alligatorium, apparently closely related to the species A. paintenense from the Tithonian of Franconia. It is the first reportof one of these small continental crocodilians from north-eastern France.     

Impact of a diazinon calamity in 1996 on the aquatic macroinvertebrates in the River Meuse, The Netherlands

A more or less continuous load of pesticides has been noted in the River Meuse in recent years. In April 1996, when high concentrations (up toca. 1 g 1^–1) of the insecticide diazinon were measured in the River Meuse at the Belgian-Dutch border, the maximum concentration for drinking water production was exceeded. This was alerted after activity changes of fish and daphnids in the biological monitoring systems (RIZA). These observations were compared with literature toxicity data of diazinon, in order to determine the ecological impact of this diazinon discharge on the aquatic macroinvertebrates. LC₅₀ values of several aquatic macroinvertebrate species were exceeded. In addition, a high mortality was observed in bioassays with the midgeChironomus riparius and the caddisflyHydropsyche angustipennis. It is inferred that the species composition of the macrofauna community in the River Meuse is likely to be reduced by such an accident. Recolonization of the River Meuse by sensitive macrofauna species may therefore be prevented by incidental pesticide discharges.     

Comparative analysis of the floristic richness of alpine communities in the Caucasus and the Central Alps

Abstract. Species richness of selected alpine plant communities in the Teberda Biosphere Reserve (Northwestern Caucasus) and the Davos area (Central Alps) was compared in series of plots from 0.0025 to 100 m². Communities developing under similar ecological conditions and with similar syntaxonomic positions were compared in order to estimate the role of recent environment versus regional historical factors in determining plant community structure and diversity. The floristic richness of the Caucasian and Alpic fens was very similar. The Grasslands and Meadows were quite similar as to floristic richness for plots > 25 cm × 25 cm, but the Caucasian communities had fewer species in smaller plots. The Lichen heath at Teberda was richer than the Caricetum curvulae cetrarietosum for all plot sizes, except the two smallest ones. On the other hand, the plots of the Salix Snowbed community were richer in species than the Caucasian snowbeds for all plot sizes. The Rhododendron Shrubland plots were very similar as to floristic richness in larger plots (4 — 100 m²). Generally, most Alpic communities near Davos were richer in species at small plot sizes than the corresponding communities from Teberda. Caucasian communities were floristically less similar to each other — and thus more discrete — than the Alpic ones. The possible role of different factors controlling floristic richness of the communities is discussed. Our results suggest that recent ecological conditions have a big influence on local floristic diversity and may lead to high similarities between ecologically similar communities from different regions. In addition, the general floristic richness of a regions as well as island effects should be taken into account.     

No evidence of intrinsic spatial processes driving Neotropical savanna vegetation on different substrates

The relationships between floristic patterns and environmental variation in tropical savannas have been the focus of many studies worldwide. However, important aspects of these relationships, such as the role of geographic distance in structuring plant communities, have received little attention. We investigated the individual and combined influences of substrate, climatic, and spatial factors on the floristic‐structural dissimilarity between two savanna physiognomies in the core region of Brazilian savannas: one on plain relief with deep soils and another on steep relief with shallow rocky soils. Ten 1‐ha plots were sampled in each physiognomy. We modeled species abundance using multiple linear models and variance partitioning. Our results indicated that spatial processes that are intrinsically related to species variation have negligible effects on floristic variation. The most important predictors in our models were related to soil characteristics (mainly nutrient availability) and topography (relief and elevation). Consequently, the substrate component exhibited the greatest power (14%) in explaining the floristic‐structural variation in the overall variance partitioning. Our results provide the first demonstration of the individual and combined contributions of substrate, climatic, and spatial factors to the occurrence and abundance of woody species in the most diverse and threatened savanna in the world. We also provide evidence that neutral processes might not be strong predictors of vegetation structure where savanna substrates differ greatly; instead, community structure may be primarily regulated by environmental filters.     

Transit towards floristic homogenization on oceanic islands in the south-eastern Pacific: comparing pre-European and current floras

Aim  To quantify the occurrence of processes of homogenization or differentiation in the vascular flora of six oceanic islands.
Location  Six islands in the south-eastern Pacific drawn from the Desventuradas Archipelago, Easter Island and the Juan Fernández Archipelago.
Methods  Using published floristic studies, we determined the floristic composition of each island at two different stages: (1) pre-European colonization and (2) current flora. We compared changes in the number of shared plants and the floristic similarity among islands for each stage.
Results  The number of plant species doubled from 263 in pre-European flora to 531 species currently. Only three native species became extinct, four natives were translocated among the islands and 271 plant species were introduced from outside. The frequency of plant species shared by two or more islands is higher in the post-European floras than prior to European contact, and the level of floristic similarity between islands increased slightly.
Main conclusions  Considering the low naturalization rate of alien plants, the small number of extinctions and the meagre increase in floristic similarity, these islands are undergoing a slow process of floristic homogenization.     

Modelling the distribution of a threatened habitat: the California sage scrub

Aim Using predictive species distribution and ecological niche modelling our objectives are: (1) to identify important climatic drivers of distribution at regional scales of a locally complex and dynamic system – California sage scrub; (2) to map suitable sage scrub habitat in California; and (3) to distinguish between bioclimatic niches of floristic groups within sage scrub to assess the conservation significance of analysing such species groups. Location Coastal mediterranean‐type shrublands of southern and central California. Methods Using point localities from georeferenced herbarium records, we modelled the potential distribution and bioclimatic envelopes of 14 characteristic sage scrub species and three floristic groups (south‐coastal, coastal–interior disjunct and broadly distributed species) based upon current climate conditions. Maxent was used to map climatically suitable habitat, while principal components analysis followed by canonical discriminant analysis were used to distinguish between floristic groups and visualize species and group distributions in multivariate ecological space. Results Geographical distribution patterns of individual species were mirrored in the habitat suitability maps of floristic groups, notably the disjunct distribution of the coastal–interior species. Overlap in the distributions of floristic groups was evident in both geographical and multivariate niche space; however, discriminant analysis confirmed the separability of floristic groups based on bioclimatic variables. Higher performance of floristic group models compared with sage scrub as a whole suggests that groups have differing climate requirements for habitat suitability at regional scales and that breaking sage scrub into floristic groups improves the discrimination between climatically suitable and unsuitable habitat. Main conclusions The finding that presence‐only data and climatic variables can produce useful information on habitat suitability of California sage scrub species and floristic groups at a regional scale has important implications for ongoing efforts of habitat restoration for sage scrub. In addition, modelling at a group level provides important information about the differences in climatic niches within California sage scrub. Finally, the high performance of our floristic group models highlights the potential a community‐level modelling approach holds for investigating plant distribution patterns.     

Phytoplankton dynamics in the River Meuse as affected by pollution

The temporal and spatial dynamics of phytoplankton in the River Meuse and a number of physical and chemical parameters were analyzed to gain insight into the pollution level in the river affecting the phytoplankton community. During 1993, samples were taken every two months at eight stations along the River Meuse for physical, chemical and biological characterization of the water. In February, water samples were also taken for laboratory incubations of the natural plankton community. The algae showed a rapid successive development with highest densities (up to 77 mg chlorophylla l^–1) in the middle reach of the river in spring and summer. During winter algal biomass remained very low. A substantial input of nutrients (nitrogen, phosphorus) was observed in the middle reach of the river. Concurrently, an increase in temperature and a reduction in pH was observed. Furthermore, the toxicity of polar organic compounds in the Microtox test showed a strong increase up to the city of Liège and the Belgian/Dutch border; the water quality improved further downstream. The relatively high toxicity at the location Liège was reflected in relatively low growth rates of phytoplankton in the laboratory experiments using water from these locations.     

Environmental factors and community dynamics at the southernmost part of the North American Graminetum – I. On the contribution of climatic factors to temporal variation in species composition

Compositional patterns of vegetation and their relationship to temporal and spatial environmental variation, with emphasis on climatic factors, were investigated in plant communities located in the southernmost portion of the North American Graminetum, in central México. Data from 353 samples, obtained in four ecologically contrasting plant communities during 11 years, were analyzed by partial canonical correspondence analysis. Eight climatic variables and eighteen covariables (seventeen edaphic and one resource management) were included in the ordination. A relationship between floristic change and weather variation, once covariables effects were fitted, was examined. Despite a strong contrast in ecological conditions among study sites, a set of four climatic variables was finally found in which each variable contributed independently and statistically (P < 0.01) to the total variance in the vegetation data. Thus, environmental variables other than climatic could not conceal the important role of weather as a mediator of floristic change through time. Summer precipitation and summer maximum temperature showed the highest correlations with the first two species axes, 0.77 and –0.39, respectively. Contribution of these two climatic variables to variance in the vegetation data explained by environmental variables was approximately 81%. Annual species were abundant during rainy years, while abundance of perennial grasses and shrubs showed no clear relationship to weather variation. This study explicitly probes the important role of rain patterns in shaping structure and composition of semiarid communities and considers how other environmental factors can affect plant communities at the southernmost part of the North American Graminetum.     

Order and disorder in the river continuum: the contribution of continuity and connectivity to floodplain meadow biodiversity

Aim Aspects of connectivity and continuity operating in the River Meuse were analysed for their contribution to the biodiversity of the floodplain. From this analysis of the diversity and composition of the meadow communities, we aimed to derive effective biodiversity conservation strategies. Location The River Meuse is one of the larger rivers in the European Western Plains ecoregion. The alluvial plains of the river have a long history of cultivation, and for these plains the floodplain meadow vegetation is a highly appreciated and valuable nature conservation asset. Method We sampled floodplain meadows from 400 km of the six geomorphic reaches of the middle to lower course of the River Meuse. For each, 50 vascular plant relevés were recorded, representing the spectrum of floodplain meadow communities of that reach. Beta diversity was calculated to quantify similarity in species pools between the reaches. A dissimilarity formula was used to determine the turnover between the reaches, and these dissimilarities were compared with a Mantel test to detect whether species composition of the floodplain meadows exhibited connectivity and continuity between the reaches. Species richness for the floodplain vegetation data of the reaches was compared with data for riparian invertebrate communities. The vegetation data sets were ordinated using detrended correspondence analysis (DCA) to reveal patterns in the floodplain meadow species composition, and the DCA axes were related to plant functional groups and population strategies. The axis scores of the species and plots were linked to river and plant species traits. Results We did not observe an overall continuity trend in similarity, nor one in diversity in a downstream direction. Lateral connectivity was highlighted by the dissimilarity between the reaches and in the influxes of species from adjacent ecoregions. The DCA ordination showed statistically significant separations between reaches and between the plant functional groups. The second DCA axis related to the longitudinal gradient of the river, whereas the first axis showed stronger correlations with river and plant species traits. We termed this axis the ‘disorder axis’. Plant invaders and avoiders are located at the left side of the disorder axis, whereas the true river‐adapted categories of resisters and endurers are at the right extremity. Contributions to the disorder were identified in terms of connectivity with adjacent ecoregions and the physical disturbance regime of natural and anthropogenic perturbations, resulting in community changes between the reaches. Main conclusions We concluded that a single overall strategy for biodiversity conservation of the river or its floodplains is not feasible. Strategies can, however, be derived for separate river reaches based on functional groups in the communities, the disorder characteristics of the reach, and the influence of surrounding ecoregions.     

Long-term changes within the invertebrate and fish communities of the Upper Rhône River: effects of climatic factors

There is increasing evidence that the global climate change is already having measurable biological impacts. However, no study (based on actual data) has assessed the influence of the global warming on communities in rivers. We analyzed long‐term series of fish (1979–1999) and invertebrate (1980–1999) data from the Upper Rhône River at Bugey to test the influence of climatic warming on both communities. Between the periods of 1979–1981 and 1997–1999, the average water temperature of the Upper Rhône River at Bugey has increased by about 1.5°C due to atmospheric warming. In the same period, several dams have been built from 12.5 to 85 km upstream of our study segment and a nuclear power plant has been built on it. Changes in the community structure were summarized using multivariate analysis. The variability of fish abundance was correlated with discharge and temperature during the reproduction period (April–June): low flows and high temperatures coincided with high fish abundance. Beyond abundance patterns, southern, thermophilic fish species (e.g. chub, and barbel) as well as downstream, thermophilic invertebrate taxa (e.g. Athricops, Potamopyrgus) progressively replaced northern, cold‐water fish species (e.g. dace) and upstream, cold‐water invertebrate taxa (e.g. Chloroperla, Protoneumura). These patterns were significantly correlated with thermal variables, suggesting that shifts were the consequences of climatic warming. All analyses were carried out using statistics appropriate for autocorrelated time series. Our results were consistent with previous studies dealing with relationships between fish or invertebrates and water temperature, and with predictions of the impact of climatic change on freshwater communities. The potential confounding factors (i.e. dams and the nuclear power plant) did not seem to influence the observed trends.     

Are zebra mussels restricted by toxicant levels in the River Meuse? a review

This paper questions if chemical barriers prevent the occurrence of benthic invertebrates in the river Meuse. To this purpose an ecotoxicological analysis is presented, using published observations on the zebra musselDreissena polymorpha, a tolerant species that maintains populations in the river. Zebra mussels collected or exposed at the Belgian-Dutch border contained high levels of several groups of toxicants, and mixture toxicity is likely to occur. A recently developed bio-assay, using the filtration rate of the zebra mussel, demonstrated strong inhibitory effects of water from the river Meuse. To determine which (combination of) toxicants cause such effects, laboratory experiments with toxicant mixtures were carried out. It was demonstrated that in a mixture of five metals (Cu, Zn, Ni, Cd, Pb) the metals contributed to the toxicity of the mixture below the No Observed Effect Concentrations (NOEC) for these metals when tested individually. The average concentration of Cu, Zn and possibly Pb in Meuse water exceed the NOEC values for filtration rate. Thus, it seems likely that joint effects of different (groups of) toxicants in the river Meuse cause the overall toxicity of the water for the zebra mussel, explaining the marginal populations at the test site, while other river species are absent there. It is suggested that toxicants in the river Meuse restrict the recolonisation of the river by invertebrate species more sensitive than the zebra mussel. This will be studied using invertebrates of different sensitivities (molluscs, arthropods), enabling the assessment of toxicity at different stages of water quality improvement of the river Meuse.     

Multivariate approach to the classification and ordination of the forest ecosystem of Nandiar valley western Himalayas

QuestionsDoes the vegetation composition of the forests of Nandiar Valley correlate with climatic, topographic and edaphic variables? Is it possible to identify plant communities through indicator species in relation to environmental gradients? Can this approach of classification and ordination will be helpful for conservation planning?LocationForests of Nandiar Valley, Moist temperate Western Himalayas Pakistan.MethodsEighty stands were selected for quantitative and qualitative characteristic of vegetation between an elevations of 525–3817 m. Species composition was recorded by using 400 m long transects. GPS, climatic, edaphic and topographic data were recorded for each sampling site. The relationship between habitat types, species composition and distribution along with climatic, edaphic and topographic variables were analyzed using TWINSPAN, Cluster analysis and DCA ordination.ResultsSum 325 vascular plants species belonging to 97 families were recorded. Diversity index and species richness was maximum in the moist temperate zone. Classification and ordination showed that the variance in species data was 7.07. Two-ways indicator species analysis classified the vegetation into eight plant communities. Indicator species analysis revealed that slope aspect, wind speed, temperature, dew point, wet bulb, pH, organic matter and phosphorous were the strongest parameters (p ≤ 0.05) determining plant community composition and indicator species in each habitat. The results also show the strength of the environment – species relationship using Monte Carlo procedures. DCA ordination grouped different species having similar habitat and habitats having common species.ConclusionsThe multivariate analysis of the vegetation along with environmental variables of Nandiar valley confirmed the indicators of each sort of vegetation communities/microclimatic zones which could further be used in conservation planning and management not only in studied area but also in the adjacent regions as well as in the areas exhibit similar sort of climatic, edaphic and topographic conditions.     

Regional differences in floristic change in South Swedish oak forests as related to soil chemistry and land use

Abstract. Changes of the ground layer vegetation between 1983 and 1993 were studied on permanent oak forest plots in Skåne and eastern Småland, two regions of South Sweden rich in oak but with different climatic conditions. 79 of the 123 more common species differed considerably in frequency between Skåne and Småland. The oak forests in Skåne contained more typical forest species, whereas the number of light-demanding grassland species was higher in Småland. Almost half of the more common species showed significant changes in frequency between 1983 and 1993. Several species of clearings and some light-demanding grassland species increased their frequency in Skåne. They were probably favoured by canopy thinning which occurred in 65 out of 95 permanent plots. Only two species decreased in frequency in Skåne, but 14 species showed a decrease in Småland. All were typical grassland species and their decline may be regarded as a long-term response to ceased grazing and tree canopy closure as almost no tree cutting occurred during the study period in Småland. Total species richness increased with management intensity in Skåne. This was due to decreasing species losses and increasing establishment of new species at intense management. A small general increase in species numbers was recorded in Småland. Species richness was more closely related to soil pH in Skåne than in Småland. The floristic variability between the oak forest plots was closely related to soil pH in both Skåne and Småland. We suggest that changes of light conditions as a result of changes in forest management are important for current secondary succession in South Swedish oak forests, whereas the general floristic pattern is to a large extent controlled by soil acidity.     

The altitudinal gradient of vascular plant richness in Aurland, western Norway

Differences in vascular plant species richness; along the altitudinal gradient in the Aurland area of western Norway have been investigated. Based on field surveys, as complete lists as possible of all vascular plants have been compiled for each 100 m altitudinal band, from sea level to the highest mountain (1764 m). For each of the 18 altitudinal bands, climatic data have been estimated. A total of 444 vascular plant species were recorded. Highest species richness (263 species) occurred in the 600–700 in band, whereas the uppermost band had only 10 species. There are minor differences in species number between the altitudinal bands < 1000 m. Partial least squares regression shows that species richness for the overall altitudinal gradient is well predicted by mean July and January temperatures and mean annual precipitation. Species turnover is highest in the 100–200 m. 600–700 m. and 1400–1500 m altitudinal bands. In terms of the gradient in summer temperature, the study supports the generally assumed linear relationship between July temperature and the number of vascular plant species between 700 and 1500 m corresponding with a mean July temperature range of 7–11°C. The study shows a decrease of ca 30 vascular plant species with a 1°C decrease in mean July temperature, and that the “climatic vascular plant limit” is here estimated to occur at a mean July temperature of 2.4°C. Above 1500 and below 700 m. species number is lower than expected based on summer iemperature conditions alone. The 700–800 m band represents the highest floristic difference compared to the other bands. Ordination and classification analyses of the floristic compositional data of all the bands highlight the 600–800 and 1500–1600 m altitudinal bands as the major biotic boundaries along the gradient. No major discontinuity in species richness, composition, or turnover was consistently found, however, at the 1100–1200 m band representing the forest-limit ecotone in Aurland.     

云南干热河谷地区种子植物资源与开发利用初探

云南干热河谷植被均为耐旱的旱生植被,主要有以下三种类型：1）半稀树草原（Semi-savanna）型植被;2）肉质多刺灌丛;3）在金沙江上游滇西北的亚热带性的干热河谷地区分布着类似马基(Maquis-like）型的多刺小叶矮生灌丛。按其产物和用途可分为6类：药用植物、纤维植物、树脂和树胶植物、芳香油植物以及淀粉植物等。资源破坏性开发和环境的变化,使现存资源的保护和开发利用成为目前的主要问题。     

Floristic homogenization by native ruderal and alien plants in north-east Spain: the effect of environmental differences on a regional scale

Aim To evaluate the relative potential contribution of native ruderals and aliens to plant homogenization at a regional scale, after taking into account the effect of diverse environmental distances. Location Catalonia (north‐east Spain) Methods We have used the flora module of the BDBC project (Catalonian Database of Biodiversity), which provides information on plant species distribution per 10 × 10 km Universal Transverse Mercator (UTM) cell. Pairwise floristic similarities of: (1) total, (2) native non‐ruderal, (3) native ruderal, and (4) alien vascular plant species have been calculated for a particularly well‐sampled subset of UTM cells, using a modified version of the Simpson index. These similarities have been compared per UTM pair using Mantel tests, before and after considering their relative association with geographical, climatic and landscape distances from linear regression models. The floristic similarity of both total and native non‐ruderal species was also correlated with the proportion of alien and native ruderal species after discounting the effects of environmental distances. Results The proportion of variance explained by environmental correlates was highest for the floristic similarity of native non‐ruderal plants and lowest for that of aliens. In all plant groups, climatic distance was the main significant variable of species similarity. Geographical distance was only significant for total and native non‐ruderal species and was of secondary importance in both cases. Landscape distance was not significant in any case. Similarities among both aliens and native ruderals were significantly higher than among native non‐ruderals, but these differences disappeared after removing the effect of environmental distances. Main conclusions Species similarity between sites may depend on differences in environmental factors other than geographical distance. This has to be taken into account when exploring the implications for biotic homogenization. In the case of Catalonian flora, the potentially homogenizing effect of native ruderal and alien species seems to be associated with their lower dependence on geographical distance and climatic factors compared with those of native, non‐ruderal species.     

Floristic and structural variation in the Tamaulipan thornscrub,northeastern Mexico

Abstract. A generalized research strategy is presented for identifying the ecological effects of the physical environment and management in a poorly known region of subtropical, semiarid thornscrub in northeastern Mexico. Vegetation samples were stratified across a small number of climatic subregions, substrate types and topographic situations. Classification analysis and PCo A of the species x site matrix of incidence data after application of the Information Statistic were used. The analyses suggested that the regional variation in climate, substrates and topography was responsible for the major floristic differences in the vegetation. The distributions of most plant species were related to the variation in the physical environment. PCo A of the species x site cover data after application of the Bray-Curtis dissimilarity metric revealed evidence of vegetation change due to overgrazing in each major floristic group, but not to selective cutting for timber and firewood.