A comparison between aquatic birds of lakes and coastal rivers in Florida

Aquatic birds were counted on five Gulf coast Florida rivers to determine if these river systems supported densities, biomass and species richness similar to those found on Florida lakes. Forty-two species were identified and for the species that were found on both Florida streams and lakes similar densities and biomass were encountered. As with Florida lakes, stream bird abundance and species richness were higher in winter months than in summer months, a consequence of migratory bird populations. Total bird abundance, biomass per unit of phosphorus, and species richness per unit of area were similar to data collected on Florida lakes. Thus, Florida rivers are capable of supplying sufficient resources to maintain bird densities, biomass and species richness values similar to lakes of equal size and nutrient concentrations and are therefore important habitats for aquatic bird populations. An examination of individual habitat characteristics indicates that water depth was inversely correlated and submersed aquatic vegetation was positively correlated with bird density, biomass and species richness within the river systems. While both habitat characteristics are important they are also inversely related making it difficult to separate the individual significance of each characteristic.     

Lakes and rivers as islands: species-area relationships in the fish faunas of Ontario

Synopsis Fish species richness in 82 lakes in Ontario, Canada was significantly correlated with surface area. In this region, latitude explained only a small amount of the variation in fish species richness. Thus, our study provides a clear demonstration of the relation between fish species richness and lake area without the confounding effects of latitude and physiography inherent in analyses from broader geographic regions. By comparison with the species-area relationship obtained, we show that acidification clearly depressed the number of fish species in 66 acid-stressed lakes in Ontario. Fish species richness was also significantly correlated with both drainage and surface areas of 21 Ontario rivers. Slopes of species-area regressions of lakes and rivers did not differ significantly, suggesting that species are added to these habitats at similar rates. However, our regression analyses show that rivers support more species of fish per unit surface area of water. Although these results are consistent with some predictions of island biogeography theory, we suggest that fish species richness is more likely to be a simple function of habitat diversity, rather than an equilibrial balance between immigration and extinction.     

An Analysis of Fish Species Richness in Natural Lakes

There is a growing recognition of the need to conserve biodiversity that has been conceptualised in the Convention of Biological Diversity. Maintenance of fish species richness is particularly important, because habitat degradation in inland waters continues to accelerate on a global scale. Here we develop empirical models for predicting fish species richness in natural lakes in various geographical regions of the world. In tropical lakes where fish biodiversity is richer than in temperate lakes, fish species richness can be predicted by a few variables such as lake area and altitude. Low fish species richness in most temperate lakes might be due to the effect of glaciation on colonisation and speciation of fishes. In US, Canadian and northern European lakes, lake acidification is one of the important factors influencing fish species richness. Although limnological characteristics influence fish species richness in temperate lakes, lake area and altitude have greater predictive power. This is in contrast to fish species richness in rivers, which can be reliably predicted by basin area. In the power curves, which describe the relationship between fish species richness and habitat size in lakes and rivers, the exponent is always greater in tropical regions than in temperate regions. Because fish biodiversity is greater in the tropics threats to fish biodiversity through habitat degradation are greater than those in temperate inland waters.     

Importance of landscape context for post‐restoration recovery of riparian vegetation

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Structure of fish assemblages along spatial gradients in a deep subtropical reservoir (Itaipu Reservoir,Brazil-Paraguay border)

Synopsis We assessed the structure of the fish assemblages in the Itaipu Reservoir (Paraná River, Brazil-Paraguay) along a longitudinal or river-dam gradient (composed of riverine, transitional and lacustrine zones) and transversal or upstream–downstream gradients of the tributaries (composed of lotic and lentic stretches of tributaries and reservoir shores). We sampled stations distributed along the environmental gradients quarterly during two years. We caught mostly piscivorous, detritivorous and insectivorous fishes. The structure of the whole fish assemblage both at the regional spatial scale and at the zones of each gradient was significantly non-random according to null models and ordination analysis. When the assemblage was broken into groups, the piscivores and detritivores as well as the predators and prey showed significant non-random structure whereas the insectivores, omnivores, herbivores, benthophages and mud-eaters showed random structure. The first groups are those that had more species and the last mentioned are those with fewer species. The tributaries varied more in composition among themselves than the reservoir shores, showing a high heterogeneity in the transversal gradient. The most widespread species was the introduced piscivore Plagioscion squamosissimus. The loricariids were most abundant in the lotic and lentic stretches of the tributaries. We also observed temporal variability in species composition, mainly in the reservoir shores possibly due to the influence of the upstream floodplain on the migratory fish that periodically use the reservoir as feeding habitat. In the tributaries, the temporal changes were less predictable. A combination of habitat preferences and interactions among species, especially predator–prey relations, might be responsible for the observed structure patterns.     

Effects of hydropower-induced flow perturbations on mayfly (Ephemeroptera) richness and abundance in north Swedish river rapids

Mayfly species richness and abundance were investigated at 52 Swedish river rapids. These were either unregulated, or regulated with or without reduced discharge. Sites impacted by regulation had lower mayfly richness and abundances than unregulated reference sites. The relative importance of reduced discharge, daily fluctuations in flow, flow constancy, and the distance to nearest rapid (as a measure of isolation) was evaluated in multiple regression analyses. These indicated negative effects of diel fluctuations on both abundance and richness, whereas flow constancy was favourable for richness. Neither the distance to nearest rapid nor reduced flow were significantly related to mayfly richness and abundance. In total, 26 mayfly species were recorded. Only Baetis rhodani was found at all sites, but another five species were present at more than 40 sites. The strongest effects were found for species within the Heptageniidae. Nineteen of 20 mayfly species present in both the regulated (with unreduced flow) and unregulated reference rapids were on average more common in the unregulated ones. Mayfly assemblage structure was primarily influenced by regional factors and nutrient status, although daily fluctuations in flow together with rapids dimensions also had a significant influence. No rare species appears to be threatened by hydropower regulation though it is conceivable that depressed abundances in regulated rivers indirectly influence predators and periphyton.     

Effects of anthropogenic habitat disturbance on local pollinator diversity and species turnover across a precipitation gradient

Anthropogenic habitat disturbance can have profound effects on multiple components of forest biotas including pollinator assemblages. We assessed the effect of small-scale disturbance on local richness, abundance, diversity and evenness of insect pollinator fauna; and how habitat disturbance affected species turnover across the landscape and overall diversity along a precipitation gradient in NW Patagonia (Argentina). We evaluated the effect of disturbance on overall pollinator fauna and then separately for bees (i.e. Apoidea) and non-bee pollinators. Locally, disturbed habitats had significantly higher pollinator species richness and abundances than undisturbed habitats for the whole pollinator assemblage, but not for bees or non-bees separately. However, significant differences in species richness between habitats vanished after accounting for differences in abundance between habitat types. At a local scale Shannon–Weaver diversity and evenness did not vary with disturbance. A β diversity index indicated that, across forest types, species turnover was lower between disturbed habitats than between undisturbed habitats. In addition, rarefaction curves showed that disturbed habitats as a whole accumulated fewer species than undisturbed habitats at equivalent sample sizes. We concluded that small patches of disturbed habitat have a negligible effect on local pollinator diversity; however, habitat disturbance reduced β diversity through a homogenization of the pollinator fauna (in particular of bees) across the landscape.     

A preliminary fish assemblage index for a transitional river–reservoir system in southeastern Brazil

Large river–reservoir systems are some of the most difficult aquatic ecosystems to assess because: (1) they typically lack minimally disturbed reference sites; (2) the reservoirs are not natural systems to begin with; and (3) reservoirs with high exchange rates are transitional systems between rivers and lakes. These features are further complicated in Brazil where fish species taxonomy is incomplete (let alone fully described ecologically), where waters naturally have high organic and thermal loadings, and where dams and reservoirs provide most of the nation's electricity and water supplies. As a first step towards generating a biological tool for assessing the effects of reservoirs on rivers, we developed a preliminary River–Reservoir Fish Assemblage Index (RRFAI) in a transitional river–reservoir system in southeastern Brazil. To do so, we gill-netted fish monthly between October 2006 and September 2007 (excluding May and July 2007) immediately upriver of the reservoir, in the upper reservoir, in the lower reservoir, and immediately downriver of the reservoir. In developing our RRFAI we sought fish assemblage metrics to represent ecological characteristics including richness, habitat, trophic, tolerance, and resilience guilds. Despite clear differences in fish assemblage composition between river and reservoir sites, we found 9 metrics common to both systems that were nonredundant and had low sampling variability (number of native species, number of characiform species, number of siluriform species, % omnivorous individuals, % invertivorous individuals, % non-native carnivorous individuals, % intolerant individuals, % tolerant individuals, number of tolerant species). Fish assemblage condition was significantly and consistently lower in the lower reservoir. There was no significant difference between the dry and wet season in RRFAI scores, suggesting that a single season sample should usually suffice. Further research is needed along distinct disturbance gradients in multiple river–reservoir systems in Brazil to confirm the sensitivity of our preliminary RRFAI for assessing the physical and chemical habitat disturbances common to such systems.     

Spatial and temporal patterns of species richness in a riparian landscape

Aim To test for control of vascular plant species richness in the riparian corridor by exploring three contrasting (although not mutually exclusive) hypotheses: (1) longitudinal patterns in riparian plant species richness are governed by local, river‐related processes independent of the regional species richness, (2) riparian plant species richness is controlled by dispersal along the river (longitudinal control), and (3) the variation in riparian plant species richness mirrors variation in regional richness (lateral control). Location The riparian zones of the free‐flowing Vindel River and its surrounding river valley, northern Sweden. Methods We used data from three surveys, undertaken at 10‐year intervals, of riparian reaches (200‐m stretches of riverbank) spanning the entire river. In addition, we surveyed species richness of vascular plants in the uplands adjacent to the river in 3.75‐km² large plots along the same regional gradient. We explored the relationship between riparian and upland flora, and various environmental variables. We also evaluated temporal variation in downstream patterns of the riparian flora. Results Our results suggest that local species richness in boreal rivers is mainly a result of local, river‐related processes and dispersal along the corridor. The strongest correlation between species richness and the environment was a negative one between species number and soil pH, but pH varied within a narrow range. We did not find evidence for a correlation between species richness on regional and local scales. We found that the local patterns of species richness for naturally occurring vascular plants were temporally variable, probably in response to large‐scale disturbance caused by extreme floods. Most previous studies have found a unimodal pattern of species richness with peaks in the middle reaches of a river. In contrast, on two of three occasions corresponding to major flooding events, we found that the distribution of species richness of naturally occurring vascular plants resembled that of regional diversity: a monotonic decrease from headwater to coast. We also found high floristic similarity between the riparian corridor and the surrounding landscape. Main conclusions These results suggest that local processes control patterns of riparian species richness, but that species composition is also highly dependent on the regional species pool. We argue that inter‐annual variation in flood disturbance is probably the most important factor producing temporal variability of longitudinal species richness patterns.     

Declines in littoral species richness across both spatial and temporal nutrient gradients: a palaeolimnological study of two taxonomic groups

1. Using a palaeolimnological approach in shallow lakes, we quantified the species richness responses of diatoms and Cladocera to phosphorus enrichment. We also examined differences in species richness responses between littoral and pelagic assemblages of our focal communities. To address both spatial and temporal relationships, our study includes an analysis of both surface sediments from 40 lakes and of a lake sediment record spanning c. 120 years. The objective of our study was to determine whether similar species richness patterns occurred across trophic levels, as well as along spatial and temporal gradients. 2. We found that both diatom and Cladocera species richness estimates significantly declined with increasing phosphorus across space and through time. When the assemblages were subdivided according to known habitat preferences, littoral biodiversity maintained a negative trend, whereas pelagic species richness tended to show no relationship with phosphorus. 3. Negative productivity–diversity patterns have been observed across almost all palaeolimnological studies that span large productivity gradients. This congruence in patterns is most likely due to the similarity in data collection methods and in focal communities studied. The contrasting responses between littoral and pelagic assemblages may be explained by the differences in physical habitat and the pool of taxa in each of these environments. Consistent with the literature, we found statistical support for the idea that littoral diversity declines could be explained by an interaction between macrophytes and nutrients along strong trophic gradients. The general lack of a diversity response in our pelagic assemblages could be attributable to the limited pool of subfossil taxa. The response of the pelagic diatom could also be related to their broad range of nutrient tolerances. 4. The observed negative response of species richness to phosphorus enrichment, particularly in the littoral assemblages, has implications for ecosystems functioning because communities with reduced biodiversity often are less resilient to anthropogenic change.     

Composition and dispersal of riverine and lake phytoplankton communities in connected systems with different water retention times

1. Lake phytoplankton community structure may be influenced by both internal factors (predation, competition, resource constraints) and external ones, such as dispersal of materials and cells between connected habitats. However, little is known about the importance of cell dispersal for phytoplankton community structure in lakes. 2. We investigated the abundance and dispersal of phytoplankton cells between connected rivers and lakes, and analysed whether similarities in phytoplankton community composition between rivers and lakes were primarily related to cell import rates or to characteristics of the local habitat. We focused on lakes along a gradient of theoretical water retention times (TWRT). Two data sets from Swedish lakes were used; a seasonal study of two connected boreal forest lakes, differing in TWRT, and a multi‐lake study of 13 lakes with a continuous range of TWRTs. 3. Phytoplankton cells were transported and dispersed in all investigated rivers. In the seasonal study, cell import rates and similarities in phytoplankton community composition between the lake and its inlet(s) were much higher in the lake with a shorter TWRT. Phytoplankton community structure in different habitats was associated with total organic carbon (TOC). This indicates that local habitat characteristics may be important in determining lake phytoplankton community composition, even in the presence of substantial cell import. 4. The multi‐lake study also showed a negative relationship between TWRT and similarities in phytoplankton community composition between inlets and lakes. Moreover, similarity in community structure was related to both cell import rates from inlet to lake and differences in habitat characteristics between inlet and lake. However, the variable most strongly correlated with community structure was TOC, indicating that species sorting rather than a mass effect was the most important mechanism underlying the correlation between community structure and retention time. 5. Overall, our data suggest that local habitat characteristics may play a key role in determining community similarity in this set of lakes covering a large range of habitat connectedness. Due to the strong co‐variations between cell dispersal and TOC, it was hard to unequivocally disentangle the different mechanisms; hence, there is a need for further studies of the role of dispersal for phytoplankton community structures.     

Distribution of alien vs. native plant species in roadside communities along an altitudinal gradient in Tenerife and Gran Canaria (Canary Islands)

Roadside plant communities were studied along two roads following an altitudinal gradient in Gran Canaria and Tenerife (Canary Islands). Our aim was to investigate variation in plant species richness, particularly of the alien flora, along a gradient from coastal shrubland to summit vegetation (1950 m a.s.l. in Gran Canaria, 2300 m in Tenerife) in relation to variation in habitat factors (altitude, habitat structure, roadside disturbance, distance to urban nuclei). We compared different species groups that were classified in terms of their biogeographical status, origin and life form. Altitude was the most important factor determining species richness and composition along both roadside transects. Alien plants showed a unimodal distribution pattern along the altitudinal gradient, with less species and lower abundance at low and high altitudes, and highest abundance at intermediate altitude. Alien plant species were also relatively more frequent near urban centres. The number of native and alien species was significantly positively correlated along the altitudinal gradient. Both alien and native, non-endemic species showed differences in their distribution along the altitudinal gradient according to their biogeographical affinities and climatic tolerances. Despite considerable differences in species pools these patterns were consistent among the two islands. Environmental (abiotic) stress is proposed as a primary, altitude-related factor acting as a filter against most alien plants at coastal and high-mountain altitudes. A higher frequency or intensity of disturbance at intermediate altitudes may be a further causal factor promoting alien plants in this zone. Future management efforts to control alien plants along roads should, therefore, concentrate on intermediate altitudinal zones of the higher Canary Islands.     

Colonization by Pinus sylvestris of a former middle-geolittoral habitat on the Umeälven river in northern Sweden, following river regulation for hydro-electric power

The colonization by Scots pine of a former middle-geolittoral habitat on the Umeälven river, northern Sweden, laid almost completely dry in 1958, was compared with that of a similar habitat, subject to natural changes in water-level, on the Vindelälven river. The geographical position, altitude, exposition, slope and type of vegetation were similar at both sites. The percentage cover of field and bottom layers of the middle geolittoral are considered to have decreased after the fall in water-level of the Umeälven river. The pines were dated by making annual ring counts. During the period 1962–67 a significant increase in pine colonization took place at the Umeälven site above the new, regulated, high-water level. The tree densities (pines m^-2) in 1978 on middle geolittoral sites of the Umeälven and Vindelälven rivers were 4.9 and 1.4, respectively.     

Landscape scale effects of disturbance on riparian vegetation

1. Differing responses in riparian species richness and composition to disturbance have been reported as a possible explanation for the differences along and between rivers. This paper explores the role of physical disturbance in shaping landscape‐scale patterns of species distribution in riparian vegetation along a free‐flowing river in northern Sweden. 2. To test whether sensitivity to disturbance varies across large landscapes, we experimentally disturbed riparian vegetation along an entire, free‐flowing river by scouring the soil and the vegetation turf, cutting vegetation, applying waterborne plant litter, and after a period of recovery we measured vegetation responses. The experiment was repeated for two consecutive years. 3. We found no significant effect of disturbance on species composition, but all three forms of disturbance significantly reduced species richness. There was no downstream variation in community responses to disturbance but morphological groups of species responded differently to different kinds of disturbance. Graminoids were most resistant, suppressed only by litter burial. All forms of disturbance except cutting reduced the density of herbaceous species, and species density of trees + shrubs and dwarf shrubs was negatively affected by both scouring and cutting. We also evaluated the effects of disturbance in relation to varying levels of species richness. In nearly all cases, responses were significantly negatively correlated with control plot species richness, and relative responses indicated that species‐rich plots were less resistant to scouring and cutting. 4. Our results suggest that although all disturbance treatments had an effect on species richness, variation in sensitivity to disturbance is not the most important factor shaping landscape‐scale patterns of riparian plant species richness along rivers.     

The stream as a habitat templet for bryophytes: species' distributions along gradients in disturbance and substratum heterogeneity

1. The structure of bryophyte communities in streams in relation to habitat characteristics, especially disturbance, is described. Disturbance in rivers is quantified as movement of the stream bed, whereas in small streams water level fluctuation is used as an indicator of disturbance frequency. 2. Canonical correspondence analysis differentiated frequently disturbed sites from more stable ones. The existence of a disturbance gradient was confirmed in a subset of the study sites using long-term records of discharge variation. A parallel change was detected in the species composition of bryophyte communities with low-statured, potentially fast colonizers dominating the disturbed end and large perennial species the stable end of the gradient. 3. A consistent pattern of zonation of bryophyte species was found along the gradient from continually submersed to persistently dry conditions in small streams and lake outlets. An abrupt increase in species richness occurred at or just above the water line, where facultatively aquatic species tolerant of both conditions formed the bulk of the community. 4. The relationship between species richness and standing crop in stream bryophyte communities was consistent with the hump-backed model of Grime (1979), especially at the within-habitat scale. Quadrats of low and very high standing crop were characterized by low species richness, while peak richness was observed at intermediate standing crops. 5. A few perennial species (e.g. Fontinalis spp. and Rhynchostegium riparioides) capable of monopolizing space dominated the most stable habitats. Species composition in low biomass sites was more variable, yet only one basic growth-form (small-statured species with high allocation to spore production) seemed possible in these highly disturbed streams. In habitats of intermediate biomass, small-scale disturbances apparently allow the formation of a more varied bryophyte community. 6. A habitat templet for stream bryophyte life strategies and community structure is presented. Disturbance is proposed to be the factor filtering out traits unsuitable for a given environment. The potential of stream bryophytes for testing and developing general ecological theory is emphasized.     

Plant species richness of nature reserves: the interplay of area, climate and habitat in a central European landscape

Aim To detect regional patterns of plant species richness in temperate nature reserves and determine the unbiased effects of environmental variables by mutual correlation with operating factors. Location The Czech Republic. Methods Plant species richness in 302 nature reserves was studied by using 14 explanatory variables reflecting the reserve area, altitude, climate, habitat diversity and prevailing vegetation type. Backward elimination of explanatory variables was used to analyse the data, taking into account their interactive nature, until the model contained only significant terms. Results A minimal adequate model with reserve area, mean altitude, prevailing vegetation type and habitat diversity (expressed as the number of major habitat types in the reserve) accounted for 53.9% of the variance in species number. After removing the area effect, habitat diversity explained 15.6% of variance, while prevailing vegetation type explained 29.6%. After removing the effect of both area and vegetation type, the resulting model explained 10.3% of the variance, indicating that species richness further increased with habitat diversity, and most obviously towards warm districts. After removing the effects of area, habitat diversity and climatic district, the model still explained 9.4% of the variance, and showed that species richness (i) significantly decreased with increasing mean altitude and annual precipitation, and with decreasing January temperature in the region of the mountain flora, and (ii) increased with altitudinal range in regions of temperate and thermophilous flora. Main conclusions We described, in quantitative terms, the effects of the main factors that might be considered to be determining plant species richness in temperate nature reserves, and evaluated their relative importance. The direct habitat effect on species richness was roughly equal to the direct area effect, but the total direct and indirect effects of area slightly exceeded that of habitat. It was shown that the overall effect of composite variables such as altitude or climatic district can be separated into particular climatic variables, which influence the richness of flora in a context‐specific manner. The statistical explanation of richness variation at the level of families yielded similar results to that for species, indicating that the system of nature conservation provides similar degrees of protection at different taxonomic levels.     

Enhancement of habitat heterogeneity and species richness on rocky shores inundated by sand

Summary Many rocky shores are subject to periodic inundation by sand, which is often thought to reduce species richness by eliminating organisms intolerant of sand scour or sand smothering. However, regular disturbance (e.g. inundation) should promote richness by preventing the development of low diversity climax communities. A study of faunal richness on 10 regularly inundated shores showed that inundation does promote richness, but by increasing habitat heterogeneity. Some species are excluded from parts of the shore by sand, but because of the patchiness of sand deposits they are rarely excluded from the entire shore. Other species are found only on rocks associated with sand, while typically sandy shore animals occur in the sand deposits themselves. Total richness (281 species) was greater than for local noninundated shores and sandy beaches combined.     

Rotifer species richness along an altitudinal gradient in the Alps

Aim Biodiversity patterns along altitudinal gradients are less studied in aquatic than terrestrial systems, even though aquatic sites provide a more homogeneous environment independent of moisture constraints. We studied the altitudinal species richness pattern for planktonic rotifers in freshwater lakes and identified the environmental predictors for which altitude is a proxy. Location Two hundred and eighteen lakes of Trentino–South Tyrol (Italy) in the eastern Alps; lakes covered 98% (range 65–2960 m above sea level) of the altitudinal gradient in the Alps. Methods We performed: (1) linear regression between species richness and altitude to evaluate the general pattern, (2) multiple linear regression between species richness and environmental predictors excluding altitude to identify the most important predictors, and (3) linear regression between the residuals of the best model of step (2) and altitude to investigate any additional explanatory power of altitude. Selection of environmental predictors was based on limnological importance and non‐parametric Spearman correlations. We applied ordinary least squares regression, generalized linear, and generalized least squares modelling to select the most statistically appropriate model. Results Rotifer species richness showed a monotonic decrease with altitude independent of scale effects. Species richness could be explained (R²= 51%) by lake area as a proxy for habitat diversity, reactive silica and total phosphorus as proxies for productivity, water temperature as a proxy for energy, nitrate as a proxy for human influence and north–south and east–west directions as covariates. These predictors completely accounted for the species richness–altitude pattern, and altitude had no additional effect on species richness. Main conclusions The linear decrease of species richness along the altitudinal gradient was related to the interplay of habitat diversity, productivity, heat content and human influence. These factors are the same in terrestrial and aquatic habitats, but the greater environmental stability of aquatic systems seems to favour a linear pattern.     

Delving into the variations in tree species composition and richness across South American subtropical Atlantic and Pampean forests

Aims We analyse here the variations in species composition and richness and the geographic ranges of the tree species occurring in South American subtropical Atlantic and Pampean forests. Our goals were to assess (i) the floristic consistency of usual classifications based on vegetation physiognomy, climate and elevation; (ii) the leading role of temperature-related variables on the variations in species composition and richness; (iii) the predominance of species with tropical–subtropical ranges, possibly as a result of forest expansion over grasslands after the Last Glacial Maximum (LGM); (iv) the restriction of most subtropical endemics to stressful habitats as a possible result of past forest refuges during the LGM.Methods The region was defined by the Tropic of Capricorn to the north, the Rio de la Plata to the south, the Atlantic shoreline to the east and the catchment areas of the upper Paraná and Uruguay Rivers to the west. Multivariate analyses, multiple regression modelling and variance partition analyses were performed on a database containing 63 994 occurrence records of 1555 tree species in 491 forest sites and 48 environmental variables. All species were also classified according to their known geographic range.Important findings A main differentiation in species composition and richness was observed between the eastern windward coastlands (rain and cloud forests) and western leeward hinterlands (Araucaria and semi-deciduous forests). Pre-defined forest types on both sides were consistent with variations in tree species composition, which were significantly related to both environmental variables and spatial proximity, with extremes of low temperature playing a chief role. Tree species richness declined substantially towards the south and also from rain to seasonal forests and towards the highland summits and sandy shores. Species richness was significantly correlated with both minimum temperature and actual evapotranspiration. About 91% of the subtropical flora is shared with the much richer tropical flora, probably extracting species that can cope with frost outbreaks. The 145 subtropical endemics were not concentrated in harsher habitats.