Ecological requirements of Abies alba in the French Alps derived from dendro-ecological analysis

Abstract. We used dendro-ecological techniques to investigate fundamental relationships between climate and growth of Abies alba (silver fir) in eastern France. Seven Abies forests in the Trièves region of the French Alps were chosen to represent a wide range of ecological conditions based on the results of previous forest vegetation surveys. In each forest, four trees were sampled in each of five different stands with two cores per tree. These 280 cores were studied using two separate dendro-ecological methods: the pointer years method (based on extreme growth events), and correlation functions between tree ring-widths and monthly climatic data. Data from 11 meteorological stations were combined to provide a regional analysis of precipitation and minimum and maximum temperatures. The two dendro-ecological methods appear to be complementary, as the first technique emphasizes common and low intensity linear correlations between ring-widths and climatic variations, and the second method emphasizes extreme and unusual climatic events such as exceptionally cold or dry years. Across all sites, drought in the previous year was consistently correlated with a low growth rate; however, other climatic variables varied substantially among sites. For example, drought in the current year reduced growth more in the low elevation sites than in the high elevation sites and severe winter frost reduced growth the most in the high altitude sites and the driest site. Moreover, certain growth responses are better correlated with the age of the stands, the canopy closure and the floristic composition of the community than the abiotic factors, emphasizing the value of dendro-ecological sampling based on phytosociological units.     

Vegetation‐environment relationships in Atlantic European calcareous grasslands

Abstract. The relationship between vegetation and environment was investigated for calcareous grasslands in a region in the west of Spain, France, Britain and Ireland defined by climatic criteria. Vegetation was sampled using objective methods and data collected on soils, land cover, location and management. Climate data were obtained from an available database. Examination of the first axis of vegetation variation as defined by Detrended Correspondence Analysis (DCA) showed a gradient from the Irish and British samples to those from France. The Spanish samples formed a separate group on the second axis. The species composition along the gradients is discussed. Correlations between the vegetation gradients and environmental variables were determined. The strongest correlations with the first DCA axis were for temperature, latitude, soil organic matter, grazing and land cover. The second DCA axis was highly correlated with rainfall, altitude and land cover. The third and fourth DCA axes were more difficult to interpret but appeared to be related to land cover. The results indicate that climate factors are important at this scale, but should not be considered in isolation and that factors relating to land cover and management should also be taken into account.     

Relative importance of abiotic and land use factors in explaining variation in woody vegetation in a French rural landscape

Abstract. In a rural landscape, scale vegetation patterns of woody species are controlled by both abiotic and land use factors. The woody species composition in 126 sample plots was analysed and land use factors and some abiotic parameters were quantified using land register data. The relative importance of land use and abiotic factors was differentiated using a partial Canonical Correspondence Analysis (CCA); the influence of land use in neighbouring areas was explored by changing the scale of land use sampling. The woody species composition appears to be controlled equally by land use and abiotic factors. The fraction of floristic variation in tree species composition explained by land use variables was 33.2 % in 1980 and 30.8 % in 1992, while abiotic variables accounted for 31.2 %. Part of the 17 % of the floristic variance explained is related to the surrounding land use. Thus, when the land use of the sampled plots and the surrounding land use are considered simultaneously, up to 36.9 % of the species variation may be explained. Partial CCA enabled us to quantify the respective proportion of floristic variance which could be explained by land use (36.9 %), abiotic variables alone (20.2 %), shared variance (12.0 %) and unexplained variance (31.2 %). Our results indicated that a delayed effect of variation in land use on plant populations may exist. This delay may result either from population characteristics or from inadequate land use assessment. This study indicates the need for simultaneously examining land use and abiotic patterns in ecological studies, as many Mediterranean-type ecosystems have been shaped by these patterns.     

Potential replacement vegetation: an approach to vegetation mapping of cultural landscapes

Abstract. The concept of mapping potential replacement vegetation (PRV) is proposed as a parallel to potential natural vegetation (PNV). Potential replacement vegetation (PRV) is an abstract and hypothetical vegetation which is in balance with climatic and soil factors currently affecting a given habitat, with environmental factors influencing the habitat from outside such as air pollution, and with an abstract anthropogenic influence (management) of given type, frequency and intensity. For every habitat, there is a series of possible PRV-types corresponding to the different anthropogenic influences, e.g. grazing, mowing, trampling or growing cereals. The PRV-concept is especially useful in large-scale mapping (scales > 1 : 25 000) of small areas where replacement vegetation is the focus of attention for managers and land-use planners, for example in nature reserves where the aim is conservation of replacement vegetation managed in a traditional way, or in restoration ecology where the concept may be used for defining restoration goals and evaluating the success of restoration efforts. At smaller scales, PRV-mapping may be useful for revealing the biogeographical patterns of larger areas which may be different from the corresponding PNV patterns, because replacement vegetation and natural vegetation may respond to environmental gradients at different scales. An example of medium-scale PRV-mapping through the coincidence of diagnostic species of vegetation types, based on species distribution grid data, is presented. In cultural landscapes, the advantage of using the PRV-concept instead of PNV is its direct relationship to the replacement vegetation. In the habitat mapping with respect to the replacement vegetation, the PRV concept yields more valuable results than the mapping of actual vegetation, as the latter is strongly affected by spatially variable anthropogenic influences which may be largely independent from climatic and soil factors.     

Invasibility of a coastal strip in NE Spain by alien plants

Abstract. The alien plant species and the ecological factors that facilitate their invasion to a coastal strip in the Baix Camp region (Tarragona, NE Spain) were studied. A detailed inventory of the area showed that 20% of the plant species, most of them from the American Continent, were aliens, many of which were strongly invasive. At the habitat level, the relationships between the invasive behaviour and a number of autecological, ecological and habitat variables were analysed by means of logistic analyses. In the best model five variables were related to alien plant invasions: climatic affinity, disturbance of the invaded habitat, route of introduction, soil moisture during summer, and life form. To elucidate the environmental factors that could be responsible for alien plant invasions at a regional scale, the number of alien taxa and the environmental features at 13 localities of the western Mediterranean area were analysed. The warm lowlands, subjected to low annual rainfall and high population density, exhibited the greatest reception capacity for alien plants. It is concluded that various specific environmental features – high mean temperatures, the abundance of riparian systems, the existence of biotopes with favourable water balance which harbour a great richness of alien species and the high human pressure to which the enclave is subjected, combined with the ecological requirements of the alien species and certain species attributes (biotype) can interact at different scales and have caused the over‐representation of alien plants.     

Effects of age and distance on the composition of mixed deciduous forest fragments in an agricultural landscape

Abstract. Forest patches in central Belgium were inventoried twice for the presence or absence of forest plant species to study the effects of age and distance on species composition. All forests in the study area were subdivided based on their land use history. To avoid effects of autocorrelated environmental characteristics on species composition, habitat homogeneity was indirectly investigated using a TWINSPAN classification of the vegetation data. Two major habitats (alluvial and non‐alluvial forests) were distinguished and analysed separately. Patterns of species composition were investigated at the landscape level using Mantel tests. Species composition similarity measures were calculated between all pairs of fragments based on the floristic data. A highly significant correlation was found between species composition similarity and inter‐patch distance. Difference in age, which we used as a measure for habitat quality, was less important in explaining species composition patterns. The effects of spatial configuration became significant when difference in age was accounted for, and a partial correlation was calculated between inter‐patch distance and species composition similarity. Different results were found for alluvial and non‐alluvial forest types. Alluvial forests were more influenced by the spatial configuration than the non‐alluvial. For the non‐alluvial forest type effects measured with the difference in age between forest fragments obscured the effects of inter‐patch distance. Based on our findings we suggest that species composition is not only internally generated, but external processes such as differential colonization caused by varying degrees of isolation may be of overriding importance.     

Effects of soil erosion on the floristic composition of plant communities on marl in northeast Spain

Abstract. This study explored the validity of three responses of vegetation to increased soil erosion: reduction of vegetation cover, number of species and reduced substitution of species. 201 relevés, including edaphic and geomorphological data, were surveyed in the intensely eroded Eocene marls of the Prepyrenees (NE Spain). Changes in plant species’ presence in relevés from different degradation stages were compared. The level of vegetation degradation was defined as the total phanerogam cover which, in the studied area, was correlated to the degree of soil erosion. The considered trends were validated. Reduction of phanerogam cover and species number were gradual from low to high‐eroded areas. Vegetation degradation explained 48% of the species number variance. In the later stages of degradation a significant substitution of species was not observed, only a lower frequency of occurrence of several species that appeared in the whole set of relevés. Through the process of degradation, 47% of species displayed significantly reduced frequencies as degradation increased, none showed a significant increase in frequency. It is concluded that there are no characteristic species in these plant communities that survive in the severely eroded marls. Among the few species that had increased in frequency, most only colonised favourable micro‐environments.     

Vegetation and ecological gradients in abandoned salt pans in western France

Abstract. Salt marshes along the Atlantic coast of France have been converted into solar salt pans since the 7th century. Salt production declined strongly from the 18th century onward, leading to the abandonment of many of these salt pans. High soil salinity is a residue from the original salt production and varies among salt pans according to time since abandonment, the current flooding regime and the effect of drainage measures. The relationships between the plant communities and seven environmental variables were investigated by Detrended Correspondence Analyses (DCA). Duration of flooding, electric conductivity and sodium saturation were most strongly related to variation in vegetation. The Heleo‐chareto‐Hippuridetum vulgaris and the Caricetum ripariae occur in lagoons with slightly saline soil that are flooded for the longest time period; the Scirpetum maritimi compacti occurs in salt pans with saline soil flooded for a long period; the Alopecuro‐Juncetum gerardii occurs on saline soil where flooding is of short duration whereas the Carici‐Lolietum perennis is never flooded and occurs on only slightly saline soil. Soil salinity and duration of flooding provide a satisfactory explanation of the variation in species composition in abandoned salt pans but land‐use practices, especially grazing, have to be taken into account to fully understand their floristic composition.     

Changes in landscape structure in the northwestern Alps over the last 7000 years: lessons from soil charcoal

Abstract. Current land‐use a bandonment and the current rise in temperature in the Alps both suggest that tree limits may change. When it is assumed that the climate of the early mid‐Holocene between 8000 and 5000 yr before present is analogous to that of the predicted climate of the late 21st century, palaeo‐ecological studies of the early Holocene may provide data for the prediction of the vegetation pattern in a century from now. It appears that mid‐Holocene charcoal assemblages can be used to reconstruct the spatial patterns of the vegetation before, or during, the practice of slash‐and‐burn. Correspondence analysis (CA) of charcoal assemblages shows that an important ecological gradient is determined by elevation. However CA also shows that charcoal assemblages in profiles between 1700 and 2100 m a.s.l. are roughly stratified: the more recent assemblages from the topmost centimetres of soil are intermediate between the lowermost assemblages and assemblages from higher elevations. This suggests that the woody communities at the highest elevation were located at lower elevations at a later date. The taxonomic diversity of the soil charcoal assemblages has been compared to that of present‐day phytosociological relevés after transformation to charcoal‐equivalent data. This comparison revealed that the vegetation pattern along the altitudinal gradient in the mid‐Holocene was different from that at present. The assemblages indicate that some communities disappeared, that Picea is a late‐Holocene invading species, and that there is no strict modern analogue for the vegetation structure prior to that of 3000 yr ago. The past structure of the woody vegetation was also different from that of today. Although past vegetation is not a good analogue for predicting future vegetation patterns, it still has potential as an indicator for the potential presence of tree species where there is none today. If we assume a temperature rise, and take into account current trends of landscape use abandonment, then we can expect strong vegetation dynamics at the upper tree line in the future: Abies alba may expand to occupy elevations of ca. 1800–2000 m in mixed communities with Picea abies, Pinus sylvestris and hardwood species, and Pinus cembra may expand up to 2500–2700 m a.s.l.     

Linear relationships between aboveground biomass and plant cover in low open herbaceous vegetation

Abstract. Four herbaceous plant species of a sand dune area and several herb species of an open early‐successional patch were investigated for the occurrence of a simple relationship between aboveground biomass and plant cover. Without exception linear regressions of aboveground biomass on plant cover were found with slope factors depending on the growth form of the species. These results suggest that (early) growth of herbaceous plants in low and (temporarily) open vegetation is not affected by possible constraints caused by a decreasing ratio of plant cover to aboveground biomass. The obtained linear relationships could be used for rapid non‐destructive determination of aboveground biomass by image‐analysis of cover data.     

Patch dynamics and local succession in a sandstone area with frequent disturbance

Abstract. A system of sand talus cones in a small forested rocky sandstone area was investigated to determine the importance of disturbance‐related dynamics and mesoclimate to vegetation differentiation. These cones (usually 3–12 m long) are formed by the accumulation of sand at the foot of sandstone rocks and are subject to frequent disturbance by the transport of sand. Vegetation was recorded both at the whole‐cone level and at the within‐cone level; the latter was approached by means of a transect of small squares along each cone. Soil profiles were recorded at the upper, centre and bottom parts of the cones. To express mesoclimatic differentiation among the cones within the rocky area, the potential insolation was calculated from the horizon angular height and the likelihood of thermal inversions was estimated by the height of the cones above the valley bottom. The major environmental factor correlated with variation in cone vegetation is the disturbance dynamics as determined by the structure of the soil profile; active cones (with sand at the surface) are colonized by different species as compared with non‐active cones (with surface covered by humus sediments). There is a clear primary succession gradient from plants able to cope with continuous sand transport to stands of tall bryophytes, small shrubs and herbs. Quite often the same gradient is found within cones, with late successional vegetation confined to the lower, stabilized parts of the cones, while the upper part is still being affected by sand transport. The differentiation along the gradient of disturbance is much stronger than the differentiation due to climatic or other gradients. Indirect data indicate that the long‐term average rate of sand accumulation is ca. 1 mm/yr. If sand transport ceases, Sphagnum peat accumulates on some of the cones; the depth of accumulated Sphagnum remains may reach 50 cm. The development of the peat layer is but weakly correlated with the measured variables suggesting that random processes at the beginning of Sphagnum establishment may be a driving force. By measuring Sphagnum growth and decomposition, and the peat density, we estimated the time needed for their development to be several hundreds of years.     

The role of spatio‐temporal heterogeneity in the establishment and maintenance of Suaeda maritima in salt marshes

Abstract. The effects of disturbance and microtopography on the organization and dynamics of plant communities were studied in a European salt marsh located in the Bay of Mont St. Michel, France. The existence of seed trapping mechanisms was also tested. The study took place in the lower and middle marsh plant communities dominated by the perennials Puccinellia maritima and Halimione portulacoides, respectively and associated with the annual Suaeda maritima. Three treatments were used in series of plots placed in each community: (1) vegetation removal and root destruction to a depth of 10 cm and refilling, (2) non‐remnant herbicide treatment without vegetation removal and (3) creation of depressions (20 cm deep). These treatments were compared with adjacent control plots. The first year of the experiment showed that the perennials facilitated the establishment of Suaeda by trapping its seeds. Estimation of cover, density and biomass over 5 yr following the disturbances showed that in the first 2 yr Suaeda dominated the disturbed plots. Thereafter Suaeda was gradually eliminated by competitive exclusion after ca. 3 yr in the zone originally dominated by Puccinellia maritima and after 4 yr in the zone occupied by Halimione portulacoides. Depressions constituted refuge habitats for Suaeda by limiting competition with the perennials but also led to a high risk of mortality with temporal fluctuations in density. Despite a period of investigation limited to 5 yr, our study demonstrated that natural disturbances of various types occurred and influenced the dynamics of Suaeda, Halimione and Puccinellia. We deduced that natural disturbances and microtopography are responsible for the maintenance of the habitat in a state of non‐equilibrium by favouring the establishment of both spatial and temporal environmental heterogeneity. These conditions appear to be particularly favourable for the maintenance of annual species such as Suaeda maritima.     

Broad‐scale vegetation‐environment relationships in Eurasian high‐latitude areas

Question: How is tundra vegetation related to climatic, soil chemical, geological variables and grazing across a very large section of the Eurasian arctic area? We were particularly interested in broad‐scale vegetation‐environment relationships and how well do the patterns conform to climate‐vegetation schemes. Material and Methods: We sampled vegetation in 1132 plots from 16 sites from different parts of the Eurasian tundra. Clustering and ordination techniques were used for analysing compositional patterns. Vegetation‐environment relationships were analysed by fitting of environmental vectors and smooth surfaces onto non‐metric multidimensional scaling scattergrams. Results: Dominant vegetation differentiation was associated with a complex set of environmental variables. A general trend differentiated cold and continental areas from relatively warm and weakly continental areas, and several soil chemical and physical variables were associated with this broad‐scaled differentiation. Especially soil chemical variables related to soil acidity (pH, Ca) showed linear relationships with the dominant vegetation gradient. This was closely related to increasing cryoperturbation, decreasing precipitation and cooler conditions. Remarkable differences among relatively adjacent sites suggest that local factors such as geological properties and lemming grazing may strongly drive vegetation differentiation. Conclusions: Vegetation differentiation in tundra areas conforms to a major ecocline underlain by a complex set of environmental gradients, where precipitation, thermal conditions and soil chemical and physical processes are coupled. However, local factors such as bedrock conditions and lemming grazing may cause marked deviations from the general climate‐vegetation models. Overall, soil chemical factors (pH, Ca) turned out to have linear relationship with the broad‐scale differentiation of arctic vegetation.     

A new approach to functional interpretation of vegetation data

Abstract. In this paper we present a new approach to the simultaneous analysis of a species composition data set, an environmental gradient data set and a functional attribute data set. We demonstrate its advantages in terms of statistical modelling including model development and assessment as well as subsequent prediction. Our method is applied to a set of case data deriving from experimental wetland microcosms including 20 species, 12 treatment combinations and a classification of species into functional groups. Acknowledging that lack of independence between samples and over‐interpretation of data may lead to overly optimistic assessment of model performance, we use cross‐validation with different subsets of data to obtain realistic model performance measures. It is shown that although the outcome of the wetland experiment is predictable in terms of experimental treatments and taxo‐nomic species, the functional groups cannot be used to explain the variation in species frequencies in the experiment. We compare the method with recently published approaches to the functional analysis of vegetation data, and discuss its applied perspectives.     

Spatial variation in a grassland on soil rich in heavy metals

Abstract. The vegetation in a grassland area in the prefecture of Kilkis (N. Greece), known for its surface Cu-mineralization, was studied. 43 quadrats were established along a transect through an area where the vegetation formed patches of different size. Cover-abundance and frequency estimates for all species were made. Normal Association Analysis revealed five quadrat groups characterized by Trifolium scabrum, Linaria pelisseriana, Anthoxanthum ovatum, Gypsophila muralis and Minuartia hirsuta ssp. falcata. 43 soil samples were analysed for pH, organic C, CaCO₃, total content of Fe, Zn, Pb, Cu, Mn, Ca, Mg, K, Na and soil texture. From a discriminant analysis performed on soil data five soil groups resulted, which are highly related to the five vegetation groups. Thymus sibthorpii, Minuartia hirsuta ssp. falcata and Rumex acetosella are the most important taxa with respect to physiognomy and patchi-ness of the vegetation. The number of species in each group of quadrats is affected by the relative favourableness or severity of the soil conditions. Metal contents, mainly Zn, Cu, Mg and Na, and soil texture are considered to be among the main factors controlling the structure and physiognomy of the vegetation.     

Dynamics of understory vegetation in an old-growth boreal coniferous forest, 1988–1993

Abstract. Understorey vegetation changes in a South Norwegian old-growth coniferous forest were studied between 1988 and 1993 in 200 1-m² vegetation plots. Our aims were to quantify the amount of between-year compositional change, and to elaborate the environmental basis for long-term vegetation change, including the previously identified gradient structure with a major gradient related to topography (and soil nutrient status and soil depth) and a minor gradient reflecting paludification and canopy coverage. Species richness (yearly mean and cumulative species number) and change in species richness differed between vascular plants and cryptogams, and between forest types. The number of vascular plant species decreased in pine forest in dry years; bryophyte species number increased in spruce forest. Statistically significant vegetation change, as tested by constrained ordination (CCA) with time as the constraining variable, is demonstrated for most one-year periods and for the five-year period in most forest types. Vegetation change along identified gradients, measured as plot displacement along DCA ordination axes, also occurred. The magnitude of year-to-year vegetation change was related neither to forest type nor to one-year period; different responses to climatic and environmental change were observed in each forest type. The largest average displacement observed, from medium-rich spruce forest towards poor spruce forest, was interpreted as a long-term trend. Humus-layer pH decreased by ca. 0.25 units from 1988 to 1993, most strongly in medium-rich spruce forest where exchangeable Ca decreased and Al and Mn increased strongly. Our study supports the hypothesis that vascular plants show a long-term and broad-scale response to soil acidification. Change in bryophyte composition is linked to some very long growing-seasons. Detailed analysis of short-term vegetation dynamics enhances the interpretation of long-term changes and stresses the complementarity of univariate and multivariate methods in the analysis of vegetation change.     

Spatial variation in vegetation and abiotic factors related to the occurrence of a ring‐forming sedge

Abstract. The clonal sedge Carex humilis forms rings of densely aggregated ramets in a dry grassland community in Central Europe. We describe the small‐scale spatial variation, both in abiotic factors and vegetation, in relation to these rings. Compared to the surrounding vegetation the cover of plants, other than C. humilis, was significantly lower both in the central area of rings and within the rings themselves. The vegetation structure was also different. The soil was more fertile in the central area and within the ring than in the surroundings, measured both directly and by the abiotic response values of the vascular plants. We conclude that neither resource depletion nor competition from other plants were likely to be responsible for the low ramet density in the central area of C. humilis rings. Instead, we suggest that the ring form is caused either by the deposition of growth inhibiting substances or by intrinsic morphological rules.     

Applying ecological site concepts and state‐and‐transition models to a grazed riparian rangeland

Ecological sites and state‐and‐transition models are useful tools for generating and testing hypotheses about drivers of vegetation composition in rangeland systems. These models have been widely implemented in upland rangelands, but comparatively, little attention has been given to developing ecological site concepts for rangeland riparian areas, and additional environmental criteria may be necessary to classify riparian ecological sites. Between 2013 and 2016, fifteen study reaches on five creeks were studied at Tejon Ranch in southern California. Data were collected to describe the relationship between riparian vegetation composition, environmental variables, and livestock management; and to explore the utility of ecological sites and state‐and‐transition models for describing riparian vegetation communities and for creating hypotheses about drivers of vegetation change. Hierarchical cluster analysis was used to classify the environmental and vegetation data (15 stream reaches × 4 years) into two ecological sites and eight community phases that comprised three vegetation states. Classification and regression tree (CART) analysis was used to determine the influence of abiotic site variables, annual precipitation, and cattle activity on vegetation clusters. Channel slope explained the greatest amount of variation in vegetation clusters; however, soil texture, geology, watershed size, and elevation were also selected as important predictors of vegetation composition. The classification tree built with this limited set of abiotic predictor variables explained 90% of the observed vegetation clusters. Cattle grazing and annual precipitation were not linked to qualitative differences in vegetation. Abiotic variables explained almost all of the observed riparian vegetation dynamics—and the divisions in the CART analysis corresponded roughly to the ecological sites—suggesting that ecological sites are well‐suited for understanding and predicting change in this highly variable system. These findings support continued development of riparian ecological site concepts and state‐and‐transition models to aid decision making for conservation and management of rangeland riparian areas.     

The shrubland vegetation in western Shewa,Ethiopia and its possible recovery

Shrubland vegetation and environmental data in western Shewa, Ethiopia have been analysed. Vegetation data include cover-abundance values of vascular plant species; environmental data comprise physical and chemical properties of the soil, altitude, slope, grazing and browsing pressure. The vegetation data were subj ected to hierarchical and non-hierarchical classification and ordination with correspondence analysis. The classification resulted in seven different vegetation types, ranging from grassland with scattered shrubs to degenerated forest. Ordination of the data and biplot analysis showed that the vegetation is influenced by anthropogenic factors and altitudinal variation. Sand content is related to a low level of anthropogenic influence whereas silt content is related to a high level. This is explained by historical events rather than by the present situation. Total nitrogen, organic carbon, altitude and slope are positively correlated and these variables are negatively related to anthropogenic influences. The shrubland vegetation may have expanded from lower altitudes and drier sites as forests gradually disappeared. The recovery of an economically more rewarding vegetation type may be achieved through pro viding alternative sources of fuel and construction and through prohibiting cultivation and grazing in the shrublands on the hillsides. Regeneration can be accelerated by actively introducing seedlings of tree species that do not need a heavy canopy cover for establishment and growth.     

The vegetation of roadside verges with respect to environmental gradients in southern New Zealand

Abstract. A survey of the vegetation of roadside verges was made across the southern part of the South Island of New Zealand. Samples were taken at 10-km intervals along selected roads providing a climatic range from the suboceanic conditions of the east coast into semi-arid Central Otago, and from Central Otago through the Southern Alps to the hyper-oceanic areas of high rainfall on the west Coast. The variation in the floristic composition is associated mainly with variation in rainfall, continentality, altitude, soil acidity, soil organic matter, and presence of forest. Sites in the arable and pastoral regions on the eastern side of the Southern Alps support a herbaceous vegetation consisting mostly of exotic species of European origin, with a few native grasses scattered through the drier and less fertile sites. A greater proportion of native species is found at higher altitudes. Roadside vegetation in the area of high rainfall to the west is characterized by indigenous ferns and woody species, although vegetation adjacent to cleared areas is more similar to that on roadsides adjacent to farmland on the east coast. The pattern of distribution of both native and exotic species is strongly related to altitudinal and climatic gradients, and the environmental responses of the exotic species are similar to those recorded in Europe. This suggests a colonization of all available sites by the exotic species, despite the relatively short time since their introduction to New Zealand, rather than an incomplete invasion.