Large‐scale dark diversity estimates: new perspectives with combined methods

Large‐scale biodiversity studies can be more informative if observed diversity in a study site is accompanied by dark diversity, the set of absent although ecologically suitable species. Dark diversity methodology is still being developed and a comparison of different approaches is needed. We used plant data at two different scales (European and seven large regions) and compared dark diversity estimates from two mathematical methods: species co‐occurrence (SCO) and species distribution modeling (SDM). We used plant distribution data from the Atlas Florae Europaeae (50 × 50 km grid cells) and seven different European regions (10 × 10 km grid cells). Dark diversity was estimated by SCO and SDM for both datasets. We examined the relationship between the dark diversity sizes (type II regression) and the overlap in species composition (overlap coefficient). We tested the overlap probability according to the hypergeometric distribution. We combined the estimates of the two methods to determine consensus dark diversity and composite dark diversity. We tested whether dark diversity and completeness of site diversity (log ratio of observed and dark diversity) are related to various natural and anthropogenic factors differently than simple observed diversity. Both methods provided similar dark diversity sizes and distribution patterns; dark diversity is greater in southern Europe. The regression line, however, deviated from a 1:1 relationship. The species composition overlap of two methods was about 75%, which is much greater than expected by chance. Both consensus and composite dark diversity estimates showed similar distribution patterns. Both dark diversity and completeness measures exhibit relationships to natural and anthropogenic factors different than those exhibited by observed richness. In summary, dark diversity revealed new biodiversity patterns which were not evident when only observed diversity was examined. A new perspective in dark diversity studies can incorporate a combination of methods.     

Observed and dark diversity of alien plant species in Europe: estimating future invasion risk

Invasion by alien species is a growing concern for nature conservation. We estimated the level of invasion by alien plant species and future invasion risks at the European scale. We used a pan-European atlas and eight regional plant atlases to determine the distribution of alien and native plant richness. In addition, we estimated alien and native dark diversity (species currently absent from a site but present in the surrounding region and able to colonize the site). We used relative diversity metrics to indicate current and future risks by alien species: relative alien richness (compared to native species), alien and native completeness (log-ratio of observed to dark diversity) and completeness difference between alien and native species. Observed and relative richness of alien species were greatest in NW Europe; this suggests that sites in NW Europe could be more disturbed. Observed alien and native species richness show clear regional hotspots; the distribution of completeness values is dispersed, indicating local hotspots. Northern Europe has relatively lower alien completeness, likely because potential invaders inhabit the region but have not yet reached many localities, thereby suggesting a risk of future invasion. A greater number of potential alien species in the region increases the probability that some alien species could have detrimental impacts. Both alien richness and completeness are positively correlated with native richness and completeness, respectively, indicating that both groups share similar distribution patterns. Alien species diversity metrics in Europe are related positively to human population density and agricultural land-use. We suggest that the dark diversity concept can broaden our understanding of alien species diversity and future invasion risks.     

Plio-Pleistocene climate change and geographic heterogeneity in plant diversity–environment relationships

Plio‐Pleistocene climate change may have induced geographic heterogeneity in plant species richness–environment relationships in Europe due to greater in situ species survival and speciation rates in southern Europe. We formulate distinct hypotheses on how Plio‐Pleistocene climate change may have affected richness–topographic heterogeneity and richness–water‐energy availability relationships, causing steeper relationships in southern Europe. We investigated these hypotheses using data from Atlas Florae Europaeae on the distribution of 3069 species and geographically weighted regression (GWR). Our analyses showed that plant species richness generally increased with topographic heterogeneity (ln‐transformed altitudinal range) and actual evapotranspiration (AET). We also found evidence for strong geographic heterogeneity in the species richness–environment relationship, with a greater increase in species richness with increasing topographic heterogeneity in southern Europe (mean standardized local slope 0.610±0.245 SD in southern Europe, but only 0.270±0.175 SD in northern Europe). However, the local AET slopes were, at most, weakly different between the two regions, and their pattern did not conform to predictions, as there was a band of high local slopes across southern‐central northern Europe. This band broadly matches the transition between the temperate and boreal zones and may simply reflect the fact that few species tolerate the boreal climate. We discuss the potential explanations for the contrasting findings for the two richness–environment relationships. In conclusion, we find support for the idea that Plio‐Pleistocene climate change may sometimes affect current species richness–environment relationships via its effects on regional species pools. However, further studies integrating information on species ages and clade differentiation rates will be needed to substantiate this interpretation. On a general level, our results indicate that although strong richness–environment relationships are often found in macroecological studies, these can be contingent upon the historical constraints on the species pool.     

Dark diversity in dry calcareous grasslands is determined by dispersal ability and stress‐tolerance

Temperate calcareous grasslands are characterized by high levels of species richness at small spatial scales. Nevertheless, many species from a habitat‐specific regional species pool may be absent from local communities and represent the ‘dark diversity’ of these sites. Here we investigate dry calcareous grasslands in northern Europe to determine what proportion of the habitat‐specific species pool is realized at small scales (i.e. how the community completeness varies) and which mechanisms may be contributing to the relative sizes of the observed and dark diversity. We test whether the absence of particular species in potentially suitable grassland sites is a consequence of dispersal limitation and/or a low ability to tolerate stress (e.g. drought and grazing). We analysed a total of 1223 vegetation plots (1 × 1 m) from dry calcareous grasslands in Sweden, Estonia and western Russia. The species co‐occurrence approach was used to estimate the dark diversity for each plot. We calculated the maximum dispersal distance for each of the 291 species in our dataset by using simple plant traits (dispersal syndrome, growth form and seed characteristics). Large seed size was used as proxy for small seed number; tall plant height and low S‐strategy type scores were used to characterise low stress‐tolerance. Levels of small‐scale community completeness were relatively low (more species were absent than present) and varied between the grasslands in different geographic areas. Species in the dark diversity were generally characterized by shorter dispersal distances and greater seed weight (fewer seeds) than species in the observed diversity. Species within the dark diversity were generally taller and had a lower tolerance of stressful conditions. We conclude that, even if temperate grasslands have high levels of small‐scale plant diversity, the majority of potentially suitable species in the regional species pool may be absent as a result of dispersal limitation and low stress‐tolerance.     

DarkDivNet – A global research collaboration to explore the dark diversity of plant communities

DarkDivNet is a global research collaboration which explores dark diversity — the set of species that are absent from a site despite being suitable under the site conditions and present in the region. Participants of the network survey vascular plant diversity both at local (10  m × 10 m) and regional scales (radius 10 km) using a standardized approach. They also measure simple plant traits and collect soil samples. Observed and dark diversity together form the site‐specific species pool, and the ratio of observed diversity and dark diversity describes community completeness. We shall explore how observed and dark diversity, site‐specific species pool and community completeness vary across natural and anthropogenic gradients. We link plant diversity measures to the information obtained from environmental DNA: soil biota and plant taxa that occurred at the site before. We will refine existing dark diversity methods and use large vegetation databases to infer species habitat suitability. We expand the dark diversity concept from a purely taxonomy‐based approach to include the functional and phylogenetic aspects of diversity. DarkDivNet currently includes 161 planned sampling areas globally, but new participants are welcome. The main vegetation sampling period is scheduled until September 2020, with the first research papers being produced after that.     

Community Completeness: Linking Local and Dark Diversity within the Species Pool Concept

Biodiversity of ecological communities has been examined widely. However, comparisons of observed species richness are limited because they fail to reveal what part of the differences are caused by natural variation in species pool size and what part is due to dark diversity – the absence of suitable species from a species pool. In other words, conventional biodiversity inventories do not convey information about how complete local plant communities are. We therefore propose the community completeness concept – a new perspective on the species pool framework. In order to ascertain community completeness, we need to estimate the extent of dark diversity, for which several methods are under development. We recommend the Community Completeness Index based on a log-ratio (or logistic) expression: ln(observed richness/dark diversity). This metric offers statistical advantages over other methods (e.g. the proportion of observed richness from the species pool). We discuss how community completeness can be related to long-term and successional community stability, landscape properties and disturbance patterns as well as to a variety of biotic interactions within and among trophic levels. The community completeness concept is related to but distinctive from the alpha-beta-gamma diversity approach and the community saturation phenomenon. The Community Completeness Index is a valuable metric for comparing biodiversity of different ecosystems for nature conservation. It can be used to measure the success of ecological restoration and vulnerability to invasion by alien species. In summary, community completeness is an interface between observed local observed species richness and dark diversity, which can be useful both in theoretical and applied biodiversity research.     

Geographical patterns in prediction errors of species distribution models

Aim To describe and explain geographical patterns of false absence and false presence prediction errors that occur when describing current plant species ranges with species distribution models. Location Europe. Methods We calibrated species distribution models (generalized linear models) using a set of climatic variables and gridded distribution data for 1065 vascular plant species from the Atlas Florae Europaeae. We used randomly selected subsets for each species with a constant prevalence of 0.5, modelled the distribution 1000 times, calculated weighted averages of the model parameters and used these to predict the current distribution in Europe. Using a threshold of 0.5, we derived presence/absence maps. Comparing observed and modelled species distribution, we calculated the false absence rates, i.e. species wrongly modelled as absent, and the false presence rates, i.e. species wrongly modelled as present, on a 50 × 50 km grid. Subsequently, we related both error rates to species range properties, land use and topographic variability within grid cells by means of simultaneous autoregressive models to correct for spatial autocorrelation. Results Grid‐cell‐specific error rates were not evenly distributed across Europe. The mean false absence rate was 0.16 ± 0.12 (standard deviation) and the mean false presence rate was 0.22 ± 0.13. False absence rates were highest in central Spain, the Alps and parts of south‐eastern Europe, while false presence rates were highest in northern Spain, France, Italy and south‐eastern Europe. False absence rates were high when range edges of species accumulated within a grid cell and when the intensity of human land use was high. False presence rates were positively associated with relative occurrence area and accumulation of range edges. Main conclusions Predictions for various species are not only accompanied by species‐specific but also by grid‐cell‐specific errors. The latter are associated with characteristics of the grid cells but also with range characteristics of occurring species. Uncertainties of predictive species distribution models are not equally distributed in space, and we would recommend accompanying maps of predicted distributions with a graphical representation of predictive performance.     

The distribution of Salix species in Fennoscandia - a numerical analysis

This paper aims to detect, describe, and interpret the major distributional patterns of species within the genus Salix in Fennoscandia, and to compare these results with a European-scale analysis of Salix distributional patterns
The occurrences of the 24 native Salix species in the 558.50 km grid squares that comprise Fennoscandia, were recorded on the basis of distnbutional maps in Atlas Florae Europaeae Distnbutional patterns were summarised numencally by means of two-way indicator species analysis and detrended correspondence analysis, and were related numerically to 42 climatic, geographical, topographical, geological, land-use and edaphic variables by means of canonical correspondence analysis The possible importance of the relationships between species distributions and the environment was tested statistically
Regional climate, the most important components being related to summer temperatures, largely accounts for the regional distributional patterns within Salix in Fennoscandia This result agrees with the European-scale results for Salix , and suggests that these two scales may belong to the same domain
The regionalisation based on Salix is similar to earlier phytogeographical regionalisations of Fennoscandia, suggesting that the climatic factors proposed as being important for Salix distributions may also be important for regional vegetation patterns at the scale of Fennoscandia     

Species pools,community completeness and invasion: disentangling diversity effects on the establishment of native and alien species

Invasion should decline with species richness, yet the relationship is inconsistent. Species richness, however, is a product of species pool size and biotic filtering. Invasion may increase with richness if large species pools represent weaker environmental filters. Measuring species pool size and the proportion realised locally (completeness) may clarify diversity‐invasion relationships by separating environmental and biotic effects, especially if species’ life‐history stage and origin are accounted for. To test these relationships, we added seeds and transplants of 15 native and alien species into 29 grasslands. Species pool size and completeness explained more variation in invasion than richness alone. Although results varied between native and alien species, seed establishment and biotic resistance to transplants increased with species pool size, whereas transplant growth and biotic resistance to seeds increased with completeness. Consequently, species pools and completeness represent multiple independent processes affecting invasion; accounting for these processes improves our understanding of invasion.     

Plant species diversity and grazing in the Scandinavian mountains - patterns and processes at different spatial scales

There is a long tradition of grazing by semi‐domestic reindeer and sheep in alpine and sub‐alpine Scandinavian habitats, but present management regimes are questioned from a conservation point of view. In this review we discuss plant diversity patterns in the Scandinavian mountains in a global, regional and local perspective. The main objective was to identify processes that influence diversity at different spatial scales with a particular focus on grazing. In a global perspective the species pool of the Scandinavian mountains is limited. partly reflecting the general latitudinal decline of species but also historical and ecological factors operating after the latest glaciation. At the local scale, both productivity and disturbance are primary factors structuring diversity, but abiotic factors such as soil pH, snow distribution and temperature are also important. Although evidence is scarce, grazing favours local species richness in productive habitats, whereas species richness decreases with grazing when productivity is low. Regional patterns of plant diversity is set by, 1) the species pool. 2) the heterogeneity and fragmentation of communities, and 3) local diversity of each plant community. We suggest that local shifts in community composition depend both on the local grazing frequency and the return‐time of the plant community after a grazing session. In addition, an increasing number of grazing‐modified local patches homogenises the vegetation and is likely to reduce the regional plant diversity. The time scale of local shifts in community composition depends on plant colonisation and persistence, From a mechanistic point of view, diversity patterns at a regional scale also depend on the regional dynamics of single species. Colonisation is usually a slow and irregular process in alpine environments, whereas the capacity for extended local persistence is generally high. Although the poor knowledge of plant regional dynamics restricts our understanding of how grazing influences plant diversity, we conclude that grazing is a key process for maintaining biodiversity in the Scandinavian mountains.     

Temporal lags in observed and dark diversity in the Anthropocene

Understanding biodiversity changes in the Anthropocene (e.g. due to climate and land‐use change) is an urgent ecological issue. This important task is challenging because global change effects and species responses are dependent on the spatial scales considered. Furthermore, responses are often not immediate. However, both scale and time delay issues can be tackled when, at each study site, we consider dynamics in both observed and dark diversity. Dark diversity includes those species in the region that can potentially establish and thrive in the local sites’ conditions but are currently locally absent. Effectively, dark diversity connects biodiversity at the study site to the regional scales and defines the site‐specific species pool (observed and dark diversity together). With dark diversity, it is possible to decompose species gains and losses into two space‐related components: one associated with local dynamics (species moving from observed to dark diversity and vice versa) and another related to gains and losses of site‐specific species pool (species moving to and from the pool after regional immigration, regional extinction or change in local ecological conditions). Extinction debt and immigration credit are useful to understand dynamics in observed diversity, but delays might happen in species pool changes as well. In this opinion piece we suggest that considering both observed and dark diversity and their temporal dynamics provides a deeper understanding of biodiversity changes. Considering both observed and dark diversity creates opportunities to improve conservation by allowing to identify species that are likely to go regionally extinct as well as foreseeing which of the species that newly arrive to the region are more likely to colonize local sites. Finally, by considering temporal lags and species gains and losses in observed and dark diversity, we combine phenomena at both spatial and temporal scales, providing a novel tool to examine biodiversity change in the Anthropocene.     

The biogeography and filtering of woody plant functional diversity in North and South America

Aim In recent years evidence has accumulated that plant species are differentially sorted from regional assemblages into local assemblages along local‐scale environmental gradients on the basis of their function and abiotic filtering. The favourability hypothesis in biogeography proposes that in climatically difficult regions abiotic filtering should produce a regional assemblage that is less functionally diverse than that expected given the species richness and the global pool of traits. Thus it seems likely that differential filtering of plant traits along local‐scale gradients may scale up to explain the distribution, diversity and filtering of plant traits in regional‐scale assemblages across continents. The present work aims to address this prediction. Location North and South America. Methods We combine a dataset comprising over 5.5 million georeferenced plant occurrence records with several large plant functional trait databases in order to: (1) quantify how several critical traits associated with plant performance and ecology vary across environmental gradients; and (2) provide the first test of whether the woody plants found within 1° and 5° map grid cells are more or less functionally diverse than expected, given their species richness, across broad gradients. Results The results show that, for many of the traits studied, the overall distribution of functional traits in tropical regions often exceeds the expectations of random sampling given the species richness. Conversely, temperate regions often had narrower functional trait distributions than their smaller species pools would suggest. Main conclusion The results show that the overall distribution of function does increase towards the equator, but the functional diversity within regional‐scale tropical assemblages is higher than that expected given their species richness. These results are consistent with the hypothesis that abiotic filtering constrains the overall distribution of function in temperate assemblages, but tropical assemblages are not as tightly constrained.     

Distorted perception of the spatial distribution of plant diversity through uneven collecting efforts: the example of Asteraceae in Australia

Aim Our aims were (1) to compare observed, estimated and predicted patterns of species richness using the Australian native Asteraceae as an example, (2) to identify candidates for hotspots of diversity for the study group, and (3) to examine the distortion of our perception of the spatial distribution of species richness through uneven or misdirected sampling efforts. Location Australia. Methods Based on data from Australia’s Virtual Herbarium, we calculated and visualized observed species richness, the Chao1 estimate of richness, the C index of collecting completeness, and an estimate of richness derived from environmental niche modelling for grid cells at a resolution of 1°. The 20 cells with the highest diversity values were used to define hotspots of diversity. Results Uneven collecting activity results in misleading diversity patterns for the family Asteraceae. While observed species richness is much higher in central Australia than in other parts of the arid interior, this is an artefact resulting from the area being a hotspot of collecting activity. The mountain ranges of south‐eastern Australia and Tasmania are candidates for unbiased hotspots of species richness. Main conclusions Vast areas of the Australian interior are insufficiently sampled on a local scale, although most of them can be expected to be relatively species poor. Some areas in the south‐east and south‐west of the continent remain undersampled relative to their high species richness. Observed species numbers, estimators and environmental niche‐modelling all have their unique advantages and disadvantages for the inference of patterns of diversity.     

A probabilistic approach to estimating species pools from large compositional matrices

Abstract. Species pools are increasingly recognized as important controls of local plant community structure and diversity. While existing approaches to estimate their content and size either rely on phytosociological expert knowledge or on simple response models across environmental gradients, the proposed application of phytosociological smoothing according to Beals exploits the full information of plant co‐occurrence patterns statistically. Where numerous representative compositional data are available, the new method yields robust estimates of the potential of sites to harbour plant species. To test the new method, a large phytosociological databank covering the forested regions of Oregon (US) was subsampled randomly and evenly across strata defined by geographic regions and elevation belts. The resulting matrix of species presence/absence in 874 plots was smoothed by calculating Beals' index of sociological favourability, which estimates the probability of encountering each species at each site from the actual plot composition and the pattern of species co‐occurrence in the matrix. In a second step, the resulting lists of sociologically probable species were intersected with complete species lists for each of 14 geographical subregions. Species pools were compared to observed species composition and richness. Species pool size exhibited much clearer spatial trends than plot richness and could be modelled much better as a function of climatic factors. In this framework the goal of modelling species pools is not to test a hypothesis, but to bridge the gap between manageable scales of empirical observation and the spatio‐temporal hierarchy of diversity patterns.     

Multiscale Patterns of Riparian Plant Diversity and Implications for Restoration

Planning riparian restoration to resemble historic reference conditions requires an understanding of both local and regional patterns of plant species diversity. Thus, understanding species distributions at multiple spatial scales is essential to improve restoration planting success, to enhance long‐term ecosystem functioning, and to match restoration planting designs with historic biogeographic distributions. To inform restoration planning, we examined the biogeographic patterns of riparian plant diversity at local and regional scales within a major western U.S.A. drainage, California's Sacramento—San Joaquin Valley. We analyzed patterns of species richness and complementarity (β‐diversity) across two scales: the watershed scale and the floodplain scale. At the watershed scale, spatial patterns of native riparian richness were driven by herbaceous species, whereas woody species were largely cosmopolitan across the nearly 38,000 km² study area. At the floodplain scale, riparian floras reflected species richness and dissimilarity patterns related to hydrological and disturbance‐driven successional sequences. These findings reinforce the importance of concurrently evaluating both local and regional processes that promote species diversity and distribution of native riparian flora. Furthermore, as restoration activities become more prevalent across the landscape, strategies for restoration outcomes should emulate the patterns of species diversity and biogeographic distributions found at regional scales.     

Dark diversity reveals importance of biotic resources and competition for plant diversity across habitats

Species richness is the most commonly used metric to quantify biodiversity. However, examining dark diversity, the group of missing species which can potentially inhabit a site, can provide a more thorough understanding of the processes influencing observed biodiversity and help evaluate the restoration potential of local habitats. So far, dark diversity has mainly been studied for specific habitats or large‐scale landscapes, while less attention has been given to variation across broad environmental gradients or as a result of local conditions and biotic interactions. In this study, we investigate the importance of local environmental conditions in determining dark diversity and observed richness in plant communities across broad environmental gradients. Using the ecospace concept, we investigate how these biodiversity measures relate to abiotic gradients (defined as position), availability of biotic resources (defined as expansion), spatiotemporal extent of habitats (defined as continuity), and species interactions through competition. Position variables were important for both observed diversity and dark diversity, some with quadratic relationships, for example, plant richness showing a unimodal response to soil fertility corresponding to the intermediate productivity hypothesis. Interspecific competition represented by community mean Grime C had a negative effect on plant species richness. Besides position‐related variables, organic carbon was the most important variable for dark diversity, indicating that in late‐succession habitats such as forests and shrubs, dark diversity is generally low. The importance of highly competitive species indicates that intermediate disturbance, such as grazing, may facilitate higher species richness and lower dark diversity.     

The species pool concept as a framework for studying patterns of plant diversity

Co‐existence theories fail to adequately explain observed community patterns (diversity and composition) because they mainly address local extinctions. For a more complete understanding, the regional processes responsible for species formation and geographic dispersal should also be considered. The species pool concept holds that local variation in community patterns is dependent primarily on the availability of species, which is driven by historical diversification and dispersal at continental and landscape scales. However, empirical evidence of historical effects is limited. This slow progress can be attributed to methodological difficulties in determining the characteristics of historical species pools and how they contributed to diversity patterns in contemporary landscapes. A role of landscape‐scale dispersal limitation in determining local community patterns has been demonstrated by numerous seed addition experiments. However, disentangling general patterns of dispersal limitation in communities still requires attention. Distinguishing habitat‐specific species pools can help to meet both applied and theoretical challenges. In conservation biology, the use of absolute richness may be uninformative because the size of species pools varies between habitats. For characterizing the dynamic state of individual communities, biodiversity relative to species pools provides a balanced way of assessing communities in different habitats. Information about species pools may also be useful when studying community assembly rules, because it enables a possible mechanism of trait convergence (habitat filtering) to be explicitly assessed. Empirical study of the role of historic effects and dispersal on local community patterns has often been restricted due to methodological difficulties in determining habitat‐specific species pools. However, accumulating distributional, ecological and phylogenetic information, as well as use of appropriate model systems (e.g. archipelagos with known biogeographic histories) will allow the species pool concept to be applied effectively in future research.     

Exotic plant species invade diversity hot spots: the alien flora of northwestern Kenya

We analysed the distribution of native and alien plant species across 20 ecogeographic zones of northwestern Kenya. The source pool for the majority of aliens was Europe and America. Thus, the source pool has a biogeographic bias which explains the low proportion of aliens in the tropics: most species in the European or American source pool are not well adapted to tropical conditions. As expected, native and alien plant species showed an area effect. Correcting for this area effect. species rich zones showed a higher proportion of alien plant species in their flora. At the analysed scale, species richness of native plant communities does not increase the resistance to invasions and alien plant species invade diversity hotspots. Compared to the other ecogeographic zone, the urban area around Nairobi showed an increased richness in alien and native plant species. This is very similar to findings in Europe, although the history of urbanisation is much shorter in Kenya. The species turnover between zones (β-diversity) shows a similar pattern in native and alien plant species. Within a very short time scale the alien plant species mapped the biogeographic patterns of natives, although the geography of human activities influences the propagule pressure.     

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.