Plant species richness in continental southern Siberia: effects of pH and climate in the context of the species pool hypothesis

Aim Many high‐latitude floras contain more calcicole than calcifuge vascular plant species. The species pool hypothesis explains this pattern through an historical abundance of high‐pH soils in the Pleistocene and an associated opportunity for the evolutionary accumulation of calcicoles. To obtain insights into the history of calcicole/calcifuge patterns, we studied species richness–pH–climate relationships across a climatic gradient, which included cool and dry landscapes resembling the Pleistocene environments of northern Eurasia. Location Western Sayan Mountains, southern Siberia. Methods Vegetation and environmental variables were sampled at steppe, forest and tundra sites varying in climate and soil pH, which ranged from 3.7 to 8.6. Species richness was related to pH and other variables using linear models and regression trees. Results Species richness is higher in areas with warmer winters and at medium altitudes that are warmer than the mountains and wetter than the lowlands. In treeless vegetation, the species richness–pH relationship is unimodal. In tundra vegetation, which occurs on low‐pH soils, richness increases with pH, but it decreases in steppes, which have high‐pH soils. In forests, where soils are more acidic than in the open landscape, the species richness–pH relationship is monotonic positive. Most species occur on soils with a pH of 6–7. Main conclusions Soil pH in continental southern Siberia is strongly negatively correlated with precipitation, and species richness is determined by the opposite effects of these two variables. Species richness increases with pH until the soil is very dry. In dry soils, pH is high but species richness decreases due to drought stress. Thus, the species richness–pH relationship is unimodal in treeless vegetation. Trees do not grow on the driest soils, which results in a positive species richness–pH relationship in forests. If modern species richness resulted mainly from the species pool effects, it would suggest that historically common habitats had moderate precipitation and slightly acidic to neutral soils.     

Mollusc diversity patterns in Central European fens: hotspots and conservation priorities

Aim To assess mollusc species composition and diversity patterns of treeless fen sites and to find simple environmental parameters that characterize diversity hotspots and priority sites for conservation. Location Western Carpathian Mountains, Europe. Methods Mollusc communities were sampled quantitatively from a homogeneous area of 16 m² in the central part of each of 145 treeless fen sites. Water conductivity and pH, geographical coordinates, altitude and habitat size of the sites studied, mean annual rainfall, mean annual temperature and mean January temperature were compiled for each plot. Nestedness in species composition was tested using the binmatnest program to confirm the ‘nested habitat‐quality hypothesis’. Patterns in species diversity were analysed using the regression trees method to isolate the main predictors of species diversity. Results Nested subset patterns of species composition were found along the gradient of mineral richness. Species distribution was highly nested (T_obs = 11.21, P << 0.001) in mineral‐poor sites (with water conductivity < 300 μS cm^?1, n = 42) and was highly correlated with the site’s mineral richness (r_s = 0.76, P << 0.001). By contrast, species distribution and richness of mineral‐rich sites (c.≥ 300 μS cm^?1, n = 103) were not controlled by mineral richness. Variation in species richness was further explained by January temperature, landscape geomorphology, and total habitat area. The southern mineral‐rich low altitude fens were the most species rich, especially those of larger total area (23 species on average). These 24 sites (17% of all sites) harboured 90% of all recorded species, including all highly endangered ones. Mineral‐rich fens in montane valleys were the second most important group because they hosted the majority of populations of two rare glacial relict species (Vertigo geyeri and Pupilla alpicola). Main conclusions The significant nestedness raises the possibility of conserving the whole fen‐mollusc species pool within the most species‐rich sites. Thus, to select the conservation priority sites, easily available site characteristics for the prediction of species richness are needed. This knowledge can help us maintain fen biodiversity, which has become closely dependent on conservation management practices after the cessation of traditional mowing of fens for haymaking.     

Some ecophysiological and historical approaches to species richness and calcicole/calcifuge behaviour — contribution to a debate

Species richness in vascular plants was related to the plants’ calcifuge or calcicole behaviour using documentation from forests and open-land vegetation at about one thousand sites in the southern parts of Sweden. It is concluded that vegetation of strongly acid soils (pH-KCl < 4.5) have fewer vascular plant species than comparable vegetation of other soils, whereas there are no consistent differences in species richness between slightly-moderately acid and calcareous sites. Low species richness is particularly related to high concentrations of Al³⁺ and H⁺ ions (either soil solution concentrations or concentrations of exchangeable ions), not to a lack of calcium carbonate. The majority of plant species are able to render the sparingly soluble phosphate, iron and manganese compounds of high-pH soils available, but they are unable to tolerate much Al³⁺ or H⁺. Acidicole (calcifuge) species have developed the power of tolerating Al³⁺ and H⁺, which may be considered a secondary property of plants, but they have lost the power of solubilizing critical mineral nutrients in high-pH soils. The reasons why these ecophysiological properties are inversely related in the current flora are obscure, difficult to account for experimentally and a main ecological problem. In areas with cool-temperate climates the flora was partly or mainly extinguished by the Pleistocene glaciations. Comparatively fewer calcifuge than calcicole species have, since then, had enough time to develop, and the number of calcifuges is lower, in spite of the fact that most natural and seminatural soils of these areas are nowadays acidic.     

Iron and phosphate uptake explains the calcifuge–calcicole behavior of the terricolous lichens Cladonia furcata subsp. furcata and C. rangiformis

Mechanisms causing the calcifuge–calcicole behavior of lichens are largely unexplored. Studying the case examples of two closely related terricolous lichens, the calcifuge Cladonia furcata subsp. furcata and the calcicole C. rangiformis, we found that preference for acidic or calcareous soils in these lichens is related to iron and phosphate uptake as in vascular plants. In laboratory studies, the calcicole species was more efficient in the intracellular uptake of Fe³⁺ and phosphate at pH 8 than the calcifuge species. At pH 3, intracellular uptake of Fe²⁺ in the calcicole species significantly exceeded that in the calcifuge species suggesting that calcicole lichens suffer from toxicity symptoms by excess Fe²⁺ at acidic sites. Though these observations parallel findings from calcifuge and calcicole vascular plants, mechanisms leading to the different iron and phosphate uptake characteristics in the studied calcifuge and calcicole lichens may differ from those in vascular plants and should be the topic of future research. A role of the depside atranorin in facilitating iron uptake by reducing Fe³⁺ in the apoplast is hypothesized.     

Patterns of bryophyte and vascular plant richness in European subalpine springs

The diversity of spring habitats can be determined not only by local environmental conditions, but also by large-scale biogeographical effects. The effects can differ across various groups of organisms. We compared α-, β- and γ-diversity patterns of bryophytes and vascular plants of (sub)alpine springs in three contrasting mountain ranges: Alps (Switzerland), Balkans (Bulgaria), Western Carpathians (Slovakia, Poland). We used univariate and multivariate statistics to test for the effects of pH, conductivity, altitude, slope, mean annual temperature and annual precipitation on diversity patterns of both taxonomic groups and compared diversity patterns among the regions for particular pH and conductivity classes. We identified acidophyte and basiphyte, calcifuge and calcicole species using species response modelling. All regions displayed significant relationship between conductivity and α-diversity of vascular plants. Bulgaria showed the highest α-diversity of vascular plants for the middle part of the conductivity gradient. For both taxonomic groups, the β-diversity in the middle part of gradient was highest in Swiss Alps. The total species pool was lowest in Bulgaria. The percentage of basiphyte and calcicole species was highest in the Alps. In (sub)alpine springs, mineral richness was a better determinant of vascular plant α-diversity than pH, and the extent of the alpine area did not coincide with α-diversity. Observed inter-regional differences in diversity patterns could be explained by the different proportion of limestone bedrock and different biogeographic history. The differences in α-diversity between both taxonomic groups are presumably result of the different rates of adaptation processes.     

Local and regional patterns of species richness in Central European vegetation types along the pH/calcium gradient

We investigated the relationship between soil pH/calcium content and species richness of vascular plants in seven broadly defined Central European vegetation types, using Ellenberg indicator values for soil reaction and a phytosociological data set of 11,041 vegetation sample plots from the Czech Republic. The vegetation types included (A) broad-leaved deciduous forests, (B) meadows, (C) dry grasslands, (D) reed-bed and tall-sedge vegetation, (E) fens and transitional mires, (F) perennial synanthropic vegetation and (G) annual synanthropic vegetation. Relationships between local species richness (alpha diversity) and pH/calcium were positive for vegetation types A and C, negative for D and G, unimodal for E, and insignificant for B and F. Ellenberg soil reaction values explained 37% of variation in local species richness for vegetation type E, 24% for A, 13% for D, but only less than 4% for the others. Species pool size, i.e., the number of species that can potentially occur in a given habitat, was calculated for each plot using Beals index of sociological favourability applied to a large phytosociological database. For most vegetation types, the relationships between species pool size and pH/calcium were similar to the relationships between local species richness and pH/calcium, with the exception of meadows (weak unimodal) and perennial synanthropic vegetation (weak negative).These patterns suggest that for those types of Central European vegetation that developed independently of human influence in the Pleistocene or early Holocene (dry grasslands, deciduous forests), there are larger pools of calcicole than calcifuge species. This pattern is also found at the level of local species richness, where it is, however, less clearly pronounced, possibly due to the predominance of a few widespread and generalist calcifuges in acidic habitats. The unimodal pattern found in mires may result from similar underlying mechanisms, but in high pH environments mineral-rich spring waters probably decrease species richness by having toxic effects on plant growth. By contrast, vegetation types developed under direct human influence (meadows, synathropic vegetation) show weak negative or no relationships of local species richness or species pool to pH/calcium gradient. These results support the hypothesis ofPärtel (Ecology 83: 2361–2366, 2002) andEwald (Folia Geobot. 38: 357–366, 2003), that the modern calcicole/calcifuge disparity in the species pool of Central European flora has resulted from historical and evolutionary processes that took place on high pH soils. In the Pleistocene, calcareous soils dominated both the dry continental landscapes of Central Europe and glacial refugia of temperate flora, which were mostly situated in southern European mountain ranges with abundant limestone and dolomite. The negative pattern of species richness along the pH/calcium gradient found in reed-bed and tall-sedge vegetation, however, is not consistent with this historical explanation.     

Immobilization of tissue iron on calcareous soil: differences between calcicole and calcifuge plants

Deficiency of P and sometimes of micronutrients, especially Fe, is of importance to the calcicole–calcifuge behaviour of plants. Calcifuge species are unable to solubilize these elements or keep them metabolically active in sufficient amounts on calcareous soils. To demonstrate if calcicole, calcifuge and ‘soil indifferent’ species differ in Fe nutrition dynamics, samples of such species were transplanted on a slightly acid silicate soil (pH BaCl₂ ca 4.0) and on a calcareous soil (pH BaCl₂ ca 7.2). Plants were grown in a computer‐controlled greenhouse at a soil moisture content of 50–60% water holding capacity and with additional light (ca 160 μE s^?1 m^?2, 12 h d^?1) if ambient light was <120 μE s^?1 m^?2.
The calcifuge species developed chlorosis when grown on the calcareous soil, whereas the other species did not. Calcareous‐soil grown plants had less 1,10‐phenanthroline extractable Fe in their leaf tissues than the silicate‐grown plants whereas total leaf Fe showed more species specific properties. The ratio of 1,10‐phenanthroline extractable to total Fe in the leaves was significantly lower in the calcifuges than in the calcicoles when grown on the calcareous soil. ‘Soil indifferent’ species did not differ much from the calcicoles. Root Fe, fractioned as DCB extractable ‘plaque’ on the root surface and Fe remaining in the root after DCB extraction, showed no distinct pattern of DCB‐Fe related to the different categories, but remaining root Fe tended to be lower in the calcifuges compared to the two other categories. Leaf colour estimated by a colour scale correlated well with chlorophyll a+b content measured in the leaves of two calcifuges. Leaf P concentrations did not differ between the different categories but were more species dependent.
We conclude that chlorosis in calcifuge species is related to an immobilization of Fe in physiologically less active forms in the tissue, if plants are forced to grow on a calcareous soil, whereas calcicole and ‘soil indifferent’ species are able to retain a much higher share of their leaf Fe in metabolically active form. This probably decreases the vitality and may exclude calcifuge plants from calcareous soil. We consider this property, previously almost unconsidered in an ecological context, as important to the calcifuge–calcicole behaviour of plants.     

pH optima for Danish forest species compared with Ellenberg reaction values

Species distribution depends on the physiological and ecological niche where a species can exist and regenerate in resource competition with other species (niche limitation). The realized niche is influenced by local biotic processes that influence species behaviour and the shape of the response curves relative to environmental gradients. Processes on larger scales also influence the species niche through source-sink mechanisms (dispersal limitation) and the species richness of an area (pool limitation). Despite the growing evidence of skewed or irregular species response curves along gradients, many ecologists still assume symmetric, unimodal response curves along gradients in ecological interpretation. Ellenberg’s indicator system is probably the most common example. However, the assumption is not ecologically or statistically valid, due to the many different processes affecting the distribution of plant species. Here I present the results of Huisman-Olff-Fresco (HOF) regressions for 209 Danish forest species. HOF modelling is chosen to avoid the classical drawbacks of assuming symmetric, unimodal response patterns. I calculate the optima for all species with unimodal responses to soil pH and compare these with the Ellenberg indicator values for reaction (R), which are often used as a substitute for soil pH measurements. I demonstrate that the assumption of symmetric, unimodal species behaviour is violated in 54% of the cases and that pH optima and R indicator values for species are not always compatible. Ellenberg reaction scale has been used byEwald (Folia Geobot. 38: 357–366, 2003) as an indicator of which species are calcicole, i.e., whether they can grow and reproduce on calcareous soils. Such affinities of species, however, are related to both local niche properties and processes on large scales and cannot be generalized from a single empirical variable such as pH, nor from Ellenberg semi-ordinal indicator scale. I conclude that while the determination of whether species are calcicole or calcifuge requires more research, it is evident that Denmark contains a fairly balanced number of calciphytic and acidophytic species. This is probably due to the nearly equal areas with acidic and alkaline soils in Denmark, which also contribute to the high species richness of more than 500 vascular plant species in Danish forests.     

Plant species richness and diversity along an altitudinal gradient in the Sierra Nevada, Mexico

This article presents an analysis of plant species richness and diversity and its association with climatic and soil variables along a 1300‐m elevation gradient on the Cerro Tláloc Mountain in the northern Sierra Nevada in Mexico. Two 1000‐m² tree sampling plots were created at each of 21 selected sampling sites, as well as two 250‐m² plots for shrubs and six 9‐m² plots for herbaceous plants. Species richness and diversity were estimated for each plant life form, and beta diversity between sites was estimated along the gradient. The relationship between species richness and diversity and environmental variables was modelled using simple linear correlation and regression trees. Species richness and diversity showed a unimodal pattern with a bias towards high values in the lower half of the elevation gradient under study. This response was consistent for all three life forms. Beta diversity increased steadily along the elevation gradient, being lower between contiguous sites at intermediate elevations and high – the species replacement rate was nearly 100%– between sites at the extremes of the gradient. Few species were adapted to the full spectrum of environmental variation along the elevation gradient studied. The regression tree suggests that differences in species richness are mainly influenced by elevation (temperature and humidity) and soil variables, namely A₂ permanent wilting point, organic matter and horizon field capacity and A₁ horizon Mg²⁺.     

Natives and non‐natives plants show different responses to elevation and disturbance on the tropical high Andes of Ecuador

The aim was to assess patterns of plant diversity in response to elevation and disturbance in a tropical mountain. The study area was located in north‐central portion of the Eastern Cordillera of the Ecuadorian Andes, on a road from 1,150 m a.s.l. (Osayacu) to 4,000 (Papallacta). Along a mountain road spanning a wide altitudinal gradient, at 20 elevations we sampled three plots: one at the roadside and two perpendicular to the roadside. The relationship between elevation and species richness was assessed using linear and quadratic regressions, the effect of disturbance on species richness was determined by ANCOVA and a t test with parameters obtained from quadratic equations. Similarity of species composition among the roadside and sites distant was evaluated with the Chao‐Jaccard and classic Jaccard similarity indices, the distribution of non‐native species according to their origin were analyzed with linear and quadratic regression. The native species showed a linearly monotonic decrease with elevation, whereas non‐natives showed a quadratic distribution. Disturbed areas had the greatest number of non‐native species and lower native species richness, showing also a high floristic similarity; less disturbed areas showed the opposite. The non‐native species of temperate origin were more numerous and showed unimodal elevational distribution, while species of tropical origin were few and decreased linearly with elevation. We conclude that in a tropical highland mountain range, native and non‐native plant species respond differently to elevation: native species exhibit a monotonically linear decrease, and non‐native species show a unimodal trend. Disturbance positively affects non‐native species showing higher richness and fewer species turnover. In addition, the non‐native species are located along of the elevational gradient in relation to their biogeographic origin.     

Mollusc and plant assemblages controlled by different ecological gradients at Eastern European fens

Ecological patterns of mollusc assemblages and vegetation in relation to water chemistry, water regime, nutrient availability and climate were studied in eastern Polish lowland fens. Our goal was to examine if major compositional changes differ for molluscs and vegetation under the joint influence of multiple ecological gradients. Altogether 32 fen sites were investigated in 2010–2011, and analyzed using metric multidimensional scaling, cluster analysis and generalized additive models. Two major gradients driving the differences in mollusc species composition were revealed. The main direction of compositional changes was associated with the water table gradient, governing a species turnover from inundated and strongly water-logged sites occupied mostly by aquatic mollusc species, to moderately wet sites with the predominance of fen and meadow species. The second most important gradient for molluscs was that of mineral richness. For vegetation, three major gradients explained the changes in species composition. The highest importance was assigned to the nitrogen-to-phosphorus availability gradient (defined as a shift from N-limited to P-limited vegetation), followed by the water table gradient, and the mineral richness gradient. Our results demonstrate that the impact of mineral richness gradient, which has been often reported as the major determinant of compositional changes of fen molluscs and vegetation, can be exceeded by other ecological gradients of comparable variation. We also document for the first time that the main species turnover of fen vegetation is not accompanied by the analogous change in species composition of mollusc assemblages, due to a different sensitivity of these taxa to particular environmental factors (i.e. water level dynamics and type of nutrient limitation).     

The richness–heterogeneity relationship differs between heterogeneity measures within and among habitats

The positive monotonic relationship between habitat heterogeneity and species richness is a cornerstone of ecology. Recently, it was suggested that this relationship should be unimodal rather than monotonic due to a tradeoff between environmental heterogeneity and population sizes, which increases local species extinctions at high heterogeneity levels. Here, we studied the richness–heterogeneity relationship for an avian community using two different environmental variables, foliage‐height diversity and cover type diversity. We analyzed the richness–heterogeneity within different habitat types (grasslands, savannas, or woodlands) and at the landscape scale. We found strong evidence that both positive and unimodal relationships exist at the landscape scale. Within habitats we found positive relationships between richness and heterogeneity in grasslands and woodlands, and unimodal relationships in savannas. We suggest that the length of the environmental heterogeneity gradient (which is affected by both spatial scale and the environmental variable being analyzed) affects the type of the richness–heterogeneity relationship. We conclude that the type of the relationship between species richness and environmental heterogeneity is non‐ubiquitous, and varies both within and among habitats and environmental variables.     

Vascular plant species richness and rarity across a minerotrophic gradient in wetlands of St. Lawrence County,New York,USA

Thirteen wetlands in St. Lawrence County, NY were sampled to examine the effect of a minerotrophic gradient on vascular plant species richness and rarity. Wetlands ranged from organic soil based poor fens (average conductivity 46.40 microsemens, average Ca 3.55 ppm) to mineral soil based rich fens (average conductivity 342.10 microsemens, Ca 23.00 ppm). Vascular plant species richness was sampled during 1990 in randomly located 1.0 m² quadrats. Specific conductivity, presence or absence of hummocks, and water depth predicted 62% of the variation in richness. Richness increased as conductivity increased until 413 microsemens at which a down trend became obvious. The negative curvilinear relation between conductivity and richness is in accordance with the hump-backed model of Grime but occurs at high rather than intermediate conductivity values. State-listed rare species were found in species-rich wetlands only and had a mean associated richness value of 14.50 species m^-2. This relationship should be taken into consideration when selecting wetlands for protection or managing wetlands for maximum plant diversity.     

Shifts in the ecological behaviour of plant species between two distant regions: evidence from the base richness gradient in mires

Aim Water pH and conductivity are known to be major environmental factors controlling the species composition of nutrient‐poor wetlands. Based on the analysis of two large data sets of species co‐occurrence, sampled along the entire pH/calcium gradient, we explored whether species exhibit similar or different ecological behaviour in the two regions. Location West Carpathians (central Europe) and Bulgaria (south‐eastern Europe), situated 800 km apart. Bulgaria represents a range margin for many mire species. Methods The probability of occurrence of the 41 most common species along the pH and conductivity gradients was assessed using logistic regression fitted by means of generalized additive models. The species optimum and amplitude were determined. To check the possible effect of competitive release, we estimated where the potential maximum number of species (maximum overlap in realized niches) occurs along the base richness gradient. Results Most of the 41 frequently occurring species showed a significant response to water pH and ln‐transformed conductivity (approximating total mineral richness) in both regions. Eight species showed a shift in pH optimum greater than one unit, while 12 species showed the same or a larger shift along the conductivity gradient. Nearly all these striking shifts were connected to an extension of species tolerance towards mineral‐poor acid habitats in Bulgaria, which causes links between species and measured factors to be conspicuously weaker in Bulgaria than in the West Carpathians. Regarding ecological amplitude, 24 species exhibited a wider tolerance to water conductivity in the West Carpathians, whereas 17 species exhibited a wider tolerance in Bulgaria. Main conclusions A distinctive variation in the realized niche was observed in a large portion of the species examined. Niche shifts between local populations of the same species were similar to those of closely related vicariant species. Ecotypic adaptation within species is a possible explanation for this pattern. Other possible explanations (competitive release, specific habitat conditions, compensation for climate) seem to be less justified. The local populations of rich‐fen species may have adapted to mineral‐poor acid conditions in the high crystalline mountains of Bulgaria during dry periods of pleniglacials. Nomenclature Marhold & Hindák (1998) ; for Balkan elements not included in this source, Andreev et al. (1992) .     

Floristic diversity of an eastern Mediterranean dwarf shrubland: the importance of soil pH

Questions: Does plant species richness and composition of eastern Mediterranean dwarf shrubland (phrygana) correlate with soil pH? How important is the effect of pH on species diversity in relation to other environmental factors in this ecosystem? What is the evolutionary background of the diversity–pH relationship? Location: Western Crete, Greece. Methods: Species composition of vascular plants, soil and other environmental variables were sampled in 100‐m² plots on acidic and basic bedrock in phrygana vegetation. The relationships between species composition and environmental variables (including climate) were tested using canonical correspondence analysis, and relationships between species richness and environment using correlation and regression analyses. Data were analysed separately for different plant functional types based on life form and life span. Results: Although soil pH varied across a narrow range (5.9‐8.1), species composition changed significantly along the pH gradient within all plant functional types. For most functional types, the effect of soil pH on species composition was stronger than that of other environmental variables. Species richness of annuals, geophytes and suffruticose chamaephytes increased with soil pH, while richness of hemicryptophytes and shrubs was not correlated with pH. Conclusions: The results are consistent with the evolutionary species pool hypothesis. High numbers of calcicole annuals, geophytes and suffruticose chamaephytes may be a result of the evolution of these groups on base‐rich dry soils in the Mediterranean climate. In contrast, hemicryptophytes, a life form typical of the temperate zone, evolved on both acidic and basic soils and therefore their species numbers do not respond to soil pH across the narrow range studied. The lack of a relationship between shrub species richness and pH is difficult to explain: it may reflect the more diverse or older origin of Mediterranean woody species and their conservative niches.     

Calcicole–calcifuge plant strategies limit restoration potential in a regional semi‐arid flora

AimTo examine calcicole and calcifuge plant strategies, as well as nutrient‐acquisition strategies, as drivers of the distribution of species in response to edaphic factors, and the degree to which these strategies may act as filters to species establishment in ecological restoration on heavily altered or reconstructed substrates.LocationAn 82,000‐ha area within a major mining province in the Mid‐West region of Western Australia, harboring vegetation communities ranging from species‐poor halophytic scrub on saline flats to dense biodiverse shrubland on the skeletal soils of ancient Banded Ironstone Formations (BIF).MethodsUnivariate and multivariate analyses were employed to examine how variation in soil chemistry and landscape position (undulating plains, slopes, and BIF crests and ridges) influenced patterns of floristic diversity, calcifuge plant strategies, and nutrient‐acquisition strategies in 538 plant species from 830 relevés.ResultsLandscape position was the strongest driver of species richness and vegetation functional composition. Soils became increasingly acidic and P‐impoverished along an increasing elevational gradient. Vegetation from different landscape positions was not compositionally dissimilar, but vegetation of BIF crests and ridges was up to twice as biodiverse as vegetation from adjacent lower‐relief areas and harbored higher proportions of calcifuge species and species with mycorrhizal associations.Main conclusionsTopographic and edaphic complexity of BIF landforms in an otherwise relatively homogenous landscape has likely facilitated species accumulation over long time periods. They represent musea of regional floristic biodiversity, excluding only species that cannot establish or are inferior competitors in heavily weathered, acidic, skeletal, and nutrient‐impoverished soils. Plant strategies likely represent a major filter in establishing biodiverse, representative vegetation on postmining landforms in geologically ancient regions.     

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.     

Inter-relationships between standing crop, biodiversity and trait attributes of hydrophytic vegetation in artificial waterways

1. The extensive British canal system potentially provides a favourable habitat for aquatic plants and, because of its uniformity, it is ideal for studying relationships between disturbance intensity (largely from boat traffic) and communities of aquatic macrophytes. 2. The standing crop and species composition of aquatic plants were measured in summer at 396 sites distributed randomly over the canal network. We also quantified the number of rare species and ‘attribute groups’ (groups of species sharing similar suites of biological traits). These data were analysed in relation to standing crop, assumed to indicate a disturbance gradient. 3. Consistent with unimodal models and the Intermediate Disturbance Hypothesis, maximum hydrophyte species richness occurred at an intermediate biomass (50–200 g DWm^?2). This corresponded to a low frequency of low magnitude disturbance (light boat traffic) on navigable sections, or occasional high magnitude disturbance (the post‐interventionist phase) on sections currently unnavigable and subject to active vegetation management. The frequency of rare species was also the greatest under these conditions, reflecting the availability of regeneration niches. 4. Sorting of species into attribute groups revealed that the overall relationship between species richness and standing crop comprised of closely overlapping unimodal responses of nine attribute groups, superimposed on a core vegetation of Potamogeton pectinatus, together with greater representation of filamentous algae, lemnids and elodeids with increasing standing crop (i.e. assumed low disturbance). High species richness was associated with the overlap of functionally different groups of species, rather than with disturbance‐mediated coexistence of functionally similar plants. 5. The analysis of a matrix of sites and the representation of plant traits weighted by the biomass of species that displayed them, in relation to different aspects of disturbance, highlighted a shift from traits associated with resilience (turions, unanchored floating or submerged leaves, low body flexibility, budding, small body size), or competitiveness (entire leaves, low reproductive output, high biomass density, large body size) at high standing crop, through to attributes more associated with resistance to disturbance (rhizomes, tubers, streamlining of foliage, low biomass density) at low standing crop. 6. Comparison with a stochastic null model of change in species number along a constrained gradient, after correction for variation in sampling effort, indicated that sites towards the tails of the gradient (excluding those with extremely low biomass) supported more species than might be expected from chance alone, while the most species‐rich sites in mid‐gradient generally supported many fewer species than expected. 7. We suggest that a disturbance regime that maintains intermediate standing crops would be appropriate for the conservation of species‐rich aquatic vegetation. Precise definition of this regime, under a range of circumstances, requires the study of temporal change at representative sites.     

Species richness in deciduous forests: Effects of species pools and environmental variables

Abstract. The study was conducted in deciduous forests of two Swedish regions, Öland and Uppland. It had two objectives: to (1) test the species pool hypothesis by examining if differences in small‐scale species richness are related to differences in large‐scale species richness and the size of the regional species pool, and (2) to examine the relationship between species richness and productivity and its scale‐dependence. The first data set comprised 36 sites of moderate to high productivity. In each site, we recorded the presence of vascular plant species in nested plots ranging from 0.001 to 1000 m² and measured several environmental variables. Soil pH and Ellenberg site indicator scores for nitrogen were used as estimators of productivity. The second data set included 24 transects (each with 20 1‐m² plots) on Öland in sites with low to high productivity. Species number, soil pH and relative light intensity were determined in each plot. The forest sites on Öland were more species‐rich than the Uppland sites on all spatial scales, although environmental conditions were similar. Small‐scale and large‐scale species richness were positively correlated. The results present evidence in favour of the species pool hypothesis. In the nested‐plots data set, species number was negatively correlated with pH and nitrogen indicator scores, whereas a unimodal relationship between species number and pH was found for the transect data set. These results, as well as previously published data, support the hump‐shaped relationship between species richness and productivity in Swedish deciduous forests. Two explanations for the higher species richness of the sites with moderate productivity are given: first, these sites have a higher environmental heterogeneity and second, they have a larger ‘habitat‐specific’ species pool.     

Species response curves along environmental gradients. A case study from SE Norwegian swamp forests

Abstract. Vegetation science has relied on untested paradigms relating to the shape of species response curves along environmental gradients. To advance in this field, we used the HOF approach to model response curves for 112 plant species along six environmental gradients and three ecoclines (as represented by DCA ordination axes) in SE Norwegian swamp forests. Response curve properties were summarized in three binary response variables: (1) model unimodal or monotonous (determinate) vs. indeterminate; (2) for determinate models, unimodal vs. monotonous and (3) for unimodal models, skewed vs. symmetric. We used logistic regression to test the influence, singly and jointly, of seven predictor variables on each of three response variables. Predictor variables included gradient type (environmental or ecocline) and length (compositional turnover); species category (vascular plant, moss, Sphagnum or hepatic), species frequency and richness, tolerance (the fraction of the gradient along which the species occurs) and position of species along each gradient. The probability for fitting a determinate model increased as the main occurrence of species approached gradient extremes and with increasing species tolerance and frequency and gradient length. Appearance of unimodal models was favoured by low species tolerance and disfavoured by closeness of species to gradient extremes. Appearance of skewed models was weakly related to predictors but was slightly favoured by species optima near gradient extremes. Contrary to the results of previous studies, species category, gradient type and variation in species richness along gradients did not contribute independently to model prediction. The overall best predictors of response curve shape were position along the gradient (relative to extremes) and tolerance; the latter also expressing gradient length in units of compositional turnover. This helps predicting species responses to gradients from gradient specific species properties. The low proportion of skewed response curves and the large variation of species response curves along all gradients indicate that skewed response curves is a smaller problem for the performance of ordination methods than often claimed. We find no evidence that DCA ordination increases the unimodality, or symmetry, of species response curves more than expected from the higher compositional turnover along ordination axes. Thus ordination axes may be appropriate proxies for ecoclines, applicable for use in species response modelling.