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.     

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.     

Woody riparian plant distributions in western Oregon,USA: comparing landscape and local scale factors

We studied riparian forests along mountain streams in four large watersheds of western Oregon and far northern California, USA, to better understand the multiscale controls on woody riparian vegetation in a geographically complex region. In each of the four-study watersheds, we sampled woody riparian vegetation in161-ha sampling reaches that straddled the stream channel. Within each hectare, we sampled riparian vegetation and local environmental factors in 40 m² sampling plots arrayed along topographic transects. We also surveyed natural disturbance gaps in 6 ha in each watershed to explore the effects of fine scale disturbance on species distributions. We compared species composition across our study watersheds and used Nonmetric Multidimensional Scaling (NMS) and chi-squared analyses to compare the relative importance of landscape scale climate variables and local topographic and disturbance variables in explaining species distributions at sampling plot and hectare scales. We noted substantial turnover in the riparian flora across the region, with greatest numbers of unique species in watersheds at the ends of the regional gradient. In NMS ordinations at both scales, variation in woody riparian species composition showed strongest correlations with climatic variables and Rubus spectabilis cover, but the latter was only an important factor in the two northern watersheds. At the smaller scale, topographic variables were also important. Chi-squared analyses confirmed that more species showed landscape scale habitat preferences (watershed associations) than associations with topographic position (94.7% vs. 42.7% of species tested) or gap versus forest setting (94.7% vs. 24.6% of species tested). The woody riparian flora of western Oregon shows important biogeographic variation; species distributions showed strong associations with climatic variables, which were the primary correlates of compositional change between riparian sites at both scales analyzed. Additional local variation in composition was explained by measures of topography and disturbance.     

Tree diversity patterns in successive vegetation types along an elevation gradient in the Mountains of Eastern Mexico

Tree species richness changes along elevation gradients in response to underlying environmental conditions. Our hypothesis was that richness is associated with climatic variables and decreases with elevation. The objective was to identify trends in species, genus and family richness, diversity and vegetation structure in relation to climate variables along an elevation gradient with successive types of forest in Veracruz, Mexico. Trees were identified and measured in 0.1 ha at 15 sites located from 140 to 4000 m a.s.l. Generalized linear models were used to fit richness, diversity, basal area and density as a function of elevation; the best model was selected using Akaike’s Information Criterion. Multivariate analyses were used to explore climatic variables associated to composition of groups of sites along the gradient. Along the entire elevation gradient, species, genus and family richness decreased unimodally, and diversity decreased monotonically. Richness was positively correlated with temperature but not with precipitation. Basal area increased monotonically and highest basal area was associated with high humidity and certain tree species (Quercus and Abies). Ordinations indicated three groups of sites: lower elevation dry forest associated with temperature seasonality, mid-elevation cloud forest associated with precipitation-related variables, and coniferous forest at the top of the gradient associated with elevation. Our study shows that different plant communities are associated with certain climatic conditions and harbour different tree species, genera and families. The results support the hypothesis that species richness is associated with climate, and decreases with elevation.     

Determinants of nocturnal Lepidopteran diversity and community structure in a conifer-dominated forest

Prediction of community response of fauna to anthropogenic or environmental disturbance requires knowledge of faunal distribution and abundance as well as an understanding of the mechanisms underlying community organization. We investigated linkages between Lepidoptera, one of the most influential insect taxa in forested ecosystems, and vegetation in riparian areas of conifer-dominated forests of western Oregon, USA. Using model selection techniques, we found that canopy cover explained variability in patterns of moth species dominance and diversity better than all other factors investigated, whereas elevation best explained patterns of moth species richness. Using canonical correspondence analysis, we determined that gradients in shrub species richness, elevation, and understory species richness accounted for the most variation in regional moth community structure. Results suggest that reductions of riparian forest canopy are likely to have the greatest impact on the variety and relative abundance of moths. Due to the relative rarity and patchy distribution of the majority of individual species, we predict that lower intensity vegetation manipulations distributed across larger spatial scales are likely to be less damaging to regional moth biodiversity than more intensive alterations at smaller scales. Finally, if global warming trends continue, upward elevational shifts in moth species distributions will make forested regions at higher elevation worthy of concerted protection.     

Tropical tree species diversity in a mountain system in southern Mexico: local and regional patterns and determinant factors

Mechanisms explaining patterns of biodiversity along elevation gradients in tropical mountain systems remain controversial. We use a set of climatic, topographic, and soil variables encompassing regional, landscape, and local‐level spatial scales to explain the spatial variation of tree species diversity in the Sierra Madre of Chiapas, Mexico. We sampled 128 circular plots (0.1‐ha each) in four elevational bands along four elevation gradients or transects encompassing 100–2200 m. A total of 12,533 trees belonging to 444 species were recorded. Diversity patterns along the elevation gradient and the explanatory power of independent variables were dependent on spatial scale (regional vs transect) and functional group (total vs late‐successional or pioneer species). Diversity of all species and late‐successional species (1 – proportion of pioneer species) showed a constant pattern at the regional and transect scales, with low predictive power of climatic variables and/or elevation. A linear decrease in either number or proportion of pioneer species diversity was observed with increasing elevation, which was correlated with temperature, rainfall, and human disturbance trends. Total species diversity showed an increase with rainfall of the warmest quarter, indicating a regional‐level limiting effect of seasonality (drought duration). Yet the explanatory power of climatic and topographic variables was higher at the individual transect level than at the regional scale, suggesting the parallel but differential influence of evolutionary and geological history factors on diversification not so far studied to explain elevation patterns of species diversity in tropical mountain systems.     

Macroecological patterns of spider species richness across Europe

We analysed the pattern of covariation of European spider species richness with various environmental variables at different scales. Four layers of perception ranging from single investigation sites to the whole European continent were selected. Species richness was determined using published data from all four scales. Correlation analyses and stepwise multiple linear regression were used to relate richness to topographic, climatic and biotic variables. Up to nine environmental variables were included in the analyses (area, latitude, elevation range, mean annual temperature, local variation in mean annual temperature, mean annual precipitation, mean July temperature, local variation in mean July temperature, plant species richness). At the local and at the continental scale, no significant correlations with surface area were found, whereas at the landscape and regional scale, surface area had a significant positive effect on species richness. Factors that were positively correlated with species richness at both broader scales were plant species richness, elevation range, and specific temperature variables (regional scale: local variation in mean annual, and mean July temperature; continental scale: mean July temperature). Latitude was significantly negatively correlated with the species richness at the continental scale. Multiple models for spider species richness data accounted for up to 77% of the total variance in spider species richness data. Furthermore, multiple models explained variation in plant species richness up to 79% through the variables mean July temperature and elevation range. We conclude that these first continental wide analyses grasp the overall pattern in spider species richness of Europe quite well, although some of the observed patterns are not directly causal. Climatic variables are expected to be among the most important direct factors, although other variables (e.g. elevation range, plant species richness) are important (surrogate) correlates of spider species richness.     

Scaling Disturbance Instead of Richness to Better Understand Anthropogenic Impacts on Biodiversity

A primary impediment to understanding how species diversity and anthropogenic disturbance are related is that both diversity and disturbance can depend on the scales at which they are sampled. While the scale dependence of diversity estimation has received substantial attention, the scale dependence of disturbance estimation has been essentially overlooked. Here, we break from conventional examination of the diversity-disturbance relationship by holding the area over which species richness is estimated constant and instead manipulating the area over which human disturbance is measured. In the boreal forest ecoregion of Alberta, Canada, we test the dependence of species richness on disturbance scale, the scale-dependence of the intermediate disturbance hypothesis, and the consistency of these patterns in native versus exotic species and among human disturbance types. We related field observed species richness in 1 ha surveys of 372 boreal vascular plant communities to remotely sensed measures of human disturbance extent at two survey scales: local (1 ha) and landscape (18 km²). Supporting the intermediate disturbance hypothesis, species richness-disturbance relationships were quadratic at both local and landscape scales of disturbance measurement. This suggests the shape of richness-disturbance relationships is independent of the scale at which disturbance is assessed, despite that local diversity is influenced by disturbance at different scales by different mechanisms, such as direct removal of individuals (local) or indirect alteration of propagule supply (landscape). By contrast, predictions of species richness did depend on scale of disturbance measurement: with high local disturbance richness was double that under high landscape disturbance.     

Biogeographical gradients in galling species richness

Summary Five hypotheses were invoked to account for variation in galling species number per location on plants of different structural complexity, namely herbs, shrubs, and trees, both in Brazil and USA. The hypotheses were: 1) the altitudinal/latitudinal gradient hypothesis; 2) the harsh environment hypothesis; 3) the plant species richness hypothesis; 4) the host plant area hypothesis; 5) the plant structural complexity hypothesis. The altitudinal and the harsh environment hypotheses were correlated and supported with sample data in both localities, with increasing gall species number as altitude/latitude declined and as sites became hotter and drier. The two hypotheses were separated by studying riparian sites and dry hillside sites at the same elevation in Arizona. Galling species frequency was higher in dry sites than in riparian sites, supporting the harsh environment hypothesis. Of the five hypotheses tested only the harsh environment hypothesis predicted that galling insect species number should vary in response to environmental variables such as moisture and temperature. Temperate shrubs supported more galling species than did other plant types, both in dry and mesic sites. The overall difference between galling species richness for tropical and temperate latitudes was not statistically significant. Free-feeding insect herbivore species exhibited the opposite pattern of species richness to gallers, being more speciose in riparian sites. The present study corroborates the hypothesis that the gall forming habit is an adaptation to harsh or stressful environments, and we describe for the first time broad scale geographical patterns in galling insect species richness.     

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.     

Freshwater biodiversity at regional extent: determinants of macroinvertebrate taxonomic richness in headwater streams

We studied spatial variation of macroinvertebrate species richness in headwater streams at two spatial extents, within and across drainage systems, and assessed the relative importance of three groups of variables (local, landscape and regional) at each extent. We specifically asked whether the same variables proposed to control broad‐scale richness patterns of terrestrial organisms (temperature, topographic variability) are important determinants of species richness also in streams, or whether environmental factors effective at mainly local scales (in‐stream heterogeneity, potential productivity) constrain species richness in local communities. We used forward selection with two stopping criteria to identify the key environmental and spatial variables at each study extent. Eigenvector‐based spatial filtering was applied to evaluate spatial patterns in species richness, and variation partitioning was used to assess the amount of variation in richness attributable to purely environmental and spatial components. A prime regulator of richness variation at the bioregion extent was elevation range (increasing richness with higher topographic variability), whereas hydrological stability and temperature were unimportant. Water chemistry variables, particularly water color, exhibited strong spatially‐structured variation across drainage systems. Local environmental variables explained most of the variation in species richness at the drainage‐system extent, reflecting gradients in total phosphorus and water color (negative effect on richness). The importance of the pure spatial component was strongly region‐dependent, with a peak (60%) in one drainage system, suggesting the presence of unmeasured environmental factors. Our results emphasize the need for spatially‐explicit, regional studies to better understand geographical variation of freshwater biodiversity. Future studies need to relate species richness not only to local factors but also to broad‐scale climatic variables, recognizing the presence of spatially‐structured environmental variation.     

Small-scale plant species richness in calcareous grasslands determined by the species pool, community age and shoot density

Variation in small-scale plant species richness was examined between twenty discrete sites of calcareous alvar grasslands in the western Estonian coastal area. This community type is very species rich at the small-scale. The pattern of diversity has often been explained by variation in ecological conditions which makes it possible to avoid competitive exclusion. Here we test an alternative species pool hypothesis. The species pool was defined as the set of species present in a community, but excluding all those species whose presence may be connected with exceptional environmental conditions for this community. Monte Carlo modelling was used to determine the strengths of the relationships between the non-independent variables: richness vs species pool and average shoot density vs richness. Site age was characterized by its elevation above sea level, since there is a continuous land uplift in the study area. Species richness was positively correlated to the size of the species pool, community age and vegetation density. We conclude that historical processes on both the regional and local levels, which determine the arrival of any particular species to a target community, and not environmental heterogeneity, are responsible for the variations in species richness between communities of this type.     

Patterns of species richness for vascular plants in China's nature reserves

Explaining the heterogeneous distribution of biodiversity across the Earth has long been a challenge to ecologists and biogeographers. Here, we document the patterns of plant species richness for different taxonomic groups in China's nature reserves, and discuss their possible explanations at national and regional scales, using vascular plant richness data coupled with information on climate and topographical variables. We found that water deficit, energy and elevation range (a surrogate of habitat heterogeneity) represent the primary explanations for variation in plant species richness of the nature reserves across China. There are consistent relationships between species richness and climate and habitat heterogeneity for different taxonomic vascular plant groups at the national scale. Habitat heterogeneity is strongly associated with plant richness in all regions, whereas climatic constraints to plant diversity vary regionally. In the regions where energy is abundant or water is scarce, plant richness patterns were determined by water and habitat heterogeneity, whereas in the region with low energy inputs, water interacting with energy, and habitat heterogeneity determined its species richness pattern. Our results also suggest that energy variables alone do not represent the primary predictor of plant richness.     

Woody plant diversity,composition and structure in relation to environmental variables and land-cover types in Lake Wanchi watershed,central highlands of Ethiopia

Knowledge of vegetation dynamics associated with human land-use change and environmental variables is crucial for sustainable watershed management. The objective of this study was to analyse woody plant diversity in response to the effects of anthropogenic disturbances, and topographic variables in different land-cover types. Woody plants ≥5 cm diameter at breast height (DBH) were recorded from 20-by-20 m plots, those <5 cm DBH were recorded from 10-by-10 m plots, and tree seedlings were recorded in 3-by-3 m plots. Nonmetric multidimensional scaling with Bray–Curtis distance was employed to analyse woody species composition. Woody species richness was analysed using generalised linear modelling. Tree diameter-class distributions were examined for population structure. The study identified 104 woody plants belonging to 52 families, where 74.5% were indigenous and 16.7% were endemic to Ethiopia. Asteraceae, Fabaceae and Rosaceae were the most species-rich families. There was evidence for the effect of anthropogenic disturbance and elevation on woody species composition. Wetlands contained the highest woody species richness, while woody species richness declined with increasing elevation and disturbance. Evidence of good regeneration was observed in wetlands. To sustain the ecological functions and services provided by the watershed, rehabilitation of the forest and shrub-cover should be a high priority for local management authorities.     

Variation in freshwater fish assemblages along a regional elevation gradient in the northern Andes,Colombia

Studies on elevation diversity gradients have covered a large number of taxa and regions throughout the world; however, studies of freshwater fish are scarce and restricted to examining their changes along a specific gradient. These studies have reported a monotonic decrease in species richness with increasing elevation, but ignore the high taxonomic differentiation of each headwater assemblage that may generate high β‐diversity among them. Here, we analyzed how fish assemblages vary with elevation among regional elevation bands, and how these changes are related to four environmental clines and to changes in the distribution, habitat use, and the morphology of fish species. Using a standardized field sampling technique, we assessed three different diversity and two structural assemblage measures across six regional elevation bands located in the northern Andes (Colombia). Each species was assigned to a functional group based on its body shape, habitat use, morphological, and/or behavioral adaptations. Additionally, at each sampling site, we measured four environmental variables. Our analyses showed: (1) After a monotonic decrease in species richness, we detected an increase in richness in the upper part of the gradient; (2) diversity patterns vary depending on the diversity measure used; (3) diversity patterns can be attributed to changes in species distribution and in the richness and proportions of functional groups along the regional elevation gradient; and (4) diversity patterns and changes in functional groups are highly correlated with variations in environmental variables, which also vary with elevation. These results suggest a novel pattern of variation in species richness with elevation: Species richness increases at the headwaters of the northern Andes owing to the cumulative number of endemic species there. This highlights the need for large‐scale studies and has important implications for the aquatic conservation of the region.     

Importance of scale,land‐use,and stream network properties for riparian plant communities along an urban gradient

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Compositional patterns of riparian woodlands in the Rift Valley of northern Kenya

Environmental factors governing the composition of riparian woodlands of the Gregory Rift Valley, northern Kenya were investigated. We collected tree species composition data on elevational transects along four active stream channels and two recently abandoned channels in Turkana District, Kenya and examined variation in riparian woodland species richness and composition at two scales: first, regional differences among channels and second, patterns along individual channels. The major factors governing woodland composition were: geographic location in relation to rainfall distribution, local heterogeneity in stream substrate, fluvial disturbance within individual channels and, in some cases, location along elevation gradients. Species richness increased with increasing rainfall. There were no significant differences in the composition of woodlands associated with active and recently abandoned channel segments. However, the species richness of riparian woodlands along abandoned channel segments seemed to decline with time since abandonment. The species richness of riparian woodlands in Turkana is an order of magnitude lower than that of comparable neotropical dry forests.     

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Predictability of plant and fungal species richness of old‐growth boreal forest islands

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