Does resource availability,resource heterogeneity or species turnover mediate changes in plant species richness in grazed grasslands?

Grazing by large ungulates often increases plant species richness in grasslands of moderate to high productivity. In a mesic North American grassland with and without the presence of bison (Bos bison), a native ungulate grazer, three non-exclusive hypotheses for increased plant species richness in grazed grasslands were evaluated: (1) bison grazing enhances levels of resource (light and N) availability, enabling species that depend on higher resource availability to co-occur; (2) spatial heterogeneity in resource availability is enhanced by bison, enabling coexistence of a greater number of plant species; (3) increased species turnover (i.e. increased species colonization and establishment) in grazed grassland is associated with enhanced plant species richness. We measured availability and spatial heterogeneity in light, water and N, and calculated species turnover from long-term data in grazed and ungrazed sites in a North American tallgrass prairie. Both regression and path analyses were performed to evaluate the potential of the three hypothesized mechanisms to explain observed patterns of plant species richness under field conditions. Experimental grazing by bison increased plant species richness by 25% over an 8-year period. Neither heterogeneity nor absolute levels of soil water or available N were related to patterns of species richness in grazed and ungrazed sites. However, high spatial heterogeneity in light and higher rates of species turnover were both strongly related to increases in plant species richness in grazed areas. This suggests that creation of a mosaic of patches with high and low biomass (the primary determinant of light availability in mesic grasslands) and promotion of a dynamic species pool are the most important mechanisms by which grazers affect species richness in high productivity grasslands.     

Plant species richness and composition of heathland relics in north‐western Belgium: evidence for a rescue‐effect?

Aim To study the effects of the degree of fragmentation of heathland patches on their species richness and species composition, and to infer the ecological mechanisms behind the observed patterns. Location The heathland patches of the north‐western part of Flanders, Belgium. During the last 200 years, the heathland area in this study area has been reduced from c. 10,000 to c. 40 ha, yielding c. 150 small and highly isolated relic fragments. Methods Different isolation measures were calculated for each of the 153 inventoried heathland patches. The influence of isolation, area and habitat diversity on species richness was investigated using correlation techniques. Community composition of the patches was tested for nestedness, and the mechanisms potentially underlying this pattern were determined. Results and main conclusions Both the analyses at the species richness and community composition level yielded evidence for a positive species–area relation. This relation was not caused by higher habitat heterogeneity in larger patches. Patch isolation, measured in different ways, however seemed much more important in explaining species richness and community composition than fragment area. Our results indicate that area effects are overcompensated by the rescue‐effect: if a patch is close to other patches, species can disperse between them and prevent the species from going extinct. Species having a short living seed bank were also more sensitive to isolation than species with a long living seed bank, indicating that the latter most probably depend on the seed bank to survive periods when environmental conditions are harsh. Analogously to the spatial rescue‐effect, the existence of a persistent seed bank may lead to a sort of temporal rescue‐effect, where the extinction of a plant species is prevented through survival in the seed bank of a patch.     

What explains patterns of species richness? The relative importance of climatic‐niche evolution,morphological evolution,and ecological limits in salamanders

A major goal of evolutionary biology and ecology is to understand why species richness varies among clades. Previous studies have suggested that variation in richness among clades might be related to variation in rates of morphological evolution among clades (e.g., body size and shape). Other studies have suggested that richness patterns might be related to variation in rates of climatic‐niche evolution. However, few studies, if any, have tested the relative importance of these variables in explaining patterns of richness among clades. Here, we test their relative importance among major clades of Plethodontidae, the most species‐rich family of salamanders. Earlier studies have suggested that climatic‐niche evolution explains patterns of diversification among plethodontid clades, whereas rates of morphological evolution do not. A subsequent study stated that rates of morphological evolution instead explained patterns of species richness among plethodontid clades (along with “ecological limits” on richness of clades, leading to saturation of clades with species, given limited resources). However, they did not consider climatic‐niche evolution. Using phylogenetic multiple regression, we show that rates of climatic‐niche evolution explain most variation in richness among plethodontid clades, whereas rates of morphological evolution do not. We find little evidence that ecological limits explain patterns of richness among plethodontid clades. We also test whether rates of morphological and climatic‐niche evolution are correlated, and find that they are not. Overall, our results help explain richness patterns in a major amphibian group and provide possibly the first test of the relative importance of climatic niches and morphological evolution in explaining diversity patterns.     

A quantitative study of the epiphytic vegetation on pollarded trunks of Fraxinus excelsior at Havrå, Osterøy,western Norway

The epiphytic vegetation on 19 pollarded trees of Fraxinus excelsior at the farm Havrå, Osterøy, western Norway was investigated. Each trunk was divided into a basal zone, a middle zone and a top zone. In each zone the four different aspects were analysed by squares (12 sampling units from each trunk). Within a total of 225 sampling units, 173 taxa were recorded (84 lichens, 72 bryophytes, 17 vascular plants). The epiphytic vegetation was classified into seven TWINSPAN groups, and it is shown by detrended correspondence analysis (DCA) that these groups are mainly separated by the first two DCA axes. In an attempt to explain the floristic composition, several environmental variables were measured, and the floristic and environmental data were analysed by canonical correspondence analysis (CCA). A pollarded tree, depending on its age, represents a continuum of perhaps several hundred years. Some of the recorded epiphytic lichens are very rare in Norway, and the pollarded trees thus contribute greatly to the biodiversity of the old cultural landscape. Several epiphytic species take advantage of favourable light conditions on trunks in the open fields. At Havrå, woodland has developed by tree colonization of old meadows and open fields during the last 40 years after the cessation of earlier managements. Some floristic differences between the epiphytic vegetation on trunks in the open fields and in the woodland are found. This suggests a change in the epiphytic vegetation because of the creation of a more shady habitat in parts of the area during a few decades.     

Ecological impacts of tropical forest fragmentation: how consistent are patterns in species richness and nestedness?

Large areas of tropical forest now exist as remnants scattered across agricultural landscapes, and so understanding the impacts of forest fragmentation is important for biodiversity conservation. We examined species richness and nestedness among tropical forest remnants in birds (meta-analysis of published studies) and insects (field data for fruit-feeding Lepidoptera (butterflies and moths) and ants). Species-area relationships were evident in all four taxa, and avian and insect assemblages in remnants typically were nested subsets of those in larger areas. Avian carnivores and nectarivores and predatory ants were more nested than other guilds, implying that the sequential loss of species was more predictable in these groups, and that fragmentation alters the trophic organization of communities. For butterflies, the ordering of fragments to achieve maximum nestedness was by fragment area, suggesting that differences among fragments were driven mainly by extinction. In contrast for moths, maximum nestedness was achieved by ordering species by wing length; species with longer wings (implying better dispersal) were more likely to occur at all sites, including low diversity sites, suggesting that differences among fragments were driven more strongly by colonization. Although all four taxa exhibited high levels of nestedness, patterns of species turnover were also idiosyncratic, and thus even species-poor sites contributed to landscape-scale biodiversity, particularly for insects.     

The effects of reindeer grazing on the composition and species richness of vegetation in forest–tundra ecotone

We conducted an 8-year exclosure experiment (1999–2006) in a forest–tundra ecotonal area in northwestern Finnish Lapland to study the effects of reindeer grazing on vegetation in habitats of variable productivity and microhabitat structure. The experimental sites included tundra heath, frost heath and riparian habitats, and the two latter habitats were characterized by hummock-hollow ground forms. The total cover of vegetation, cover of willow (Salix spp.), dwarf birch (Betula nana), dwarf shrubs, forbs and grasses (Poaceae spp.) increased in exclosures in all habitats. The increase in the total cover of vegetation and in the covers of willow and dwarf birch tended to be greatest in the least productive tundra heath. Opposing to the increase in the dominant vascular plant groups, the cover and species number of bryophytes decreased in exclosures. We conclude that the effects of reindeer grazing on vegetation composition depend on environmental heterogeneity and the responses vary among plant groups. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Fungal species and their boundaries matter – Definitions,mechanisms and practical implications

Recent scientific and technological advances have improved and streamlined our ability to recognise and describe fungal species. Detailed comparative genomics studies have also expanded our understanding of species boundaries. Against this background, we explore the nature of fungal species and consider how this impacts our understanding of their genetics and evolution. The current body of evidence suggests that fungal species are unique evolutionary units that are separated from one another by boundaries that are “porous” under certain conditions (“semipermeable” in analogy to the differential permeability of membranes). Overall, the penetrability of these boundaries depends on the relatedness between donor and recipient species, the spatial proximity of related species to one another during their evolution, and the evolutionary potential associated with the breach of a boundary. Furthermore, the semipermeable nature of species boundaries fundamentally affects the population genetics of a species, with potentially profound effects on its overall evolution and biology. This also influences the methodologies used in taxonomy, because some species appear capable of maintaining their genetic isolation despite extensive penetrability of their boundaries. Most analytical procedures are also not able to distinguish the signals of species boundary permeability from those associated with incomplete lineage sorting or intraspecific diversity. Collectively, these issues greatly complicate how we study and name fungi. An awareness of the nature of species, their boundaries and the biological and genomic signatures of boundary breaches, will enhance our ability to identify them and, perhaps more importantly, to develop realistic strategies to manage and manipulate their growth and distribution.     

3,000 species and no end – species richness and community pattern of woodland macrofungi in Mecklenburg-Western Pomerania,Germany

In addition to newly generated and continuously growing datasets in mycological research, existing compilations are of high value to assess the fungi of a whole region. In the present study, a private database with ca. 65,000 entries of macromycetous fruit body observations in Mecklenburg-Western Pomerania, Germany, was analysed. Observed species richness of tree-associated mycorrhizal and saprobic fungi exceeded 3,000 taxa. The total fungal species richness could not be determined with confidence but will possibly exceed 4,000. Distinct species turnover with respect to host trees was observed. However, the rate of community overlap clearly differed between mycorrhizal and saprobic fungi and deciduous and coniferous trees. By separating the data into abundant core species and rare satellite taxa potential indicator species are presented, whose preservation will be beneficial to many other fungi and the entire ecosystems they live in.     

What predicts the richness of seeder and resprouter species in fire‐prone Cape fynbos: Rainfall reliability or vegetation density?

In ecosystems subject to regular canopy fires, woody species have evolved two general strategies of post‐fire regeneration. Seeder species are killed by fire and populations regenerate solely by post‐fire recruitment from a seed bank. Resprouter species survive fire and regenerate by vegetative regrowth from protected organs. Interestingly, the abundance of these strategies varies along environmental gradients and across regions. Two main hypotheses have been proposed to explain this spatial variation: the gap dependence and the environmental‐variability hypotheses. The gap‐dependence model predicts that seeders are favoured in sparse vegetation (vegetation gaps allowing effective post‐fire recruitment of seedlings), while resprouters are favoured in densely vegetated sites (seedlings being outcompeted by the rapid crown regrowth of resprouters). The environmental‐variability model predicts that seeders would prevail in reliable rainfall areas, whereas resprouters would be favoured in areas under highly variable rainfall that are prone to severe dry events (leading to high post‐fire seedling mortality). We tested these two models using distribution data, captured at the scale of quarter‐degree cells, for seeder and resprouter species of two speciose shrub genera (Aspalathus and Erica) common in fire‐prone fynbos ecosystems of the mediterranean‐climate part of the Cape Floristic Region. Contrary to the predictions of the gap‐dependence model, species number of both resprouters and seeders increased with values of the Normalized Difference Vegetation Index (a widely used surrogate for vegetation density), with a more marked increase for seeders. The predictions of the environmental‐variability hypothesis, by contrast, were not refuted by this study. Seeder and resprouter species of both genera showed highest richness in environments with high rainfall reliability. However, with decreasing reliability, seeder numbers dropped more quickly than those of resprouters. We conclude that the environmental‐variability model is better able to explain the abundance of woody seeder and resprouter species in Southern Hemisphere fire‐prone shrublands (fynbos and kwongan) than the gap‐dependence model.     

What drives invasibility? A multi‐model inference test and spatial modelling of alien plant species richness patterns in northern Portugal

Understanding and predicting biological invasions can focus either on traits that favour species invasiveness or on features of the receiving communities, habitats or landscapes that promote their invasibility. Here, we address invasibility at the regional scale, testing whether some habitats and landscapes are more invasible than others by fitting models that relate alien plant species richness to various environmental predictors. We use a multi‐model information‐theoretic approach to assess invasibility by modelling spatial and ecological patterns of alien invasion in landscape mosaics, and by testing competing hypotheses of environmental factors that may control invasibility. Because invasibility may be mediated by particular characteristics of invasiveness, we classified alien species according to their C‐S‐R plant strategies. We illustrate this approach with a set of 86 alien species in northern Portugal. We first focus on predictors influencing species richness and expressing invasibility, and then evaluate whether distinct plant strategies respond to the same or different groups of environmental predictors. We confirmed climate as a primary determinant of alien invasions, and as a primary environmental gradient determining landscape invasibility. The effects of secondary gradients were detected only when the area was sub‐sampled according to predictions based on the primary gradient. Then, multiple predictor types influenced patterns of alien species richness, with some types (landscape composition, topography and fire regime) prevailing over others. Alien species richness responded most strongly to extreme land management regimes, suggesting that intermediate disturbance induces biotic resistance by favouring native species richness. Land‐use intensification facilitated alien invasion, whereas conservation areas hosted few invaders, highlighting the importance of ecosystem stability in preventing invasions. Plants with different strategies exhibited different responses to environmental gradients, particularly when the variations of the primary gradient were narrowed by sub‐sampling. Such differential responses of plant strategies suggest using distinct control and eradication approaches for different areas and alien plant groups.     

Do seasonal temperatures,species traits and nearby timber harvest predict variation in moth species richness and abundance in unlogged deciduous forests?

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The relationship between diatoms and surface water quality in the Høylandet area of Nord-Trøndelag, Norway

Although the ecological effects of surface wateracidification are now well researched, factorscontrolling the abundance and occurrence of aquaticorganisms in unpolluted acid-sensitive systems arepoorly known. The Høylandet region in central Norwayexperiences relatively low levels of atmosphericpollution and its surface waters, although acid, arenot significantly acidified. Hence lakes and streamsin this region were selected to study the influence ofwater chemistry on diatom algae. Relationships betweenthe two were explored using the multivariate techniqueof canonical correspondence analysis (CCA). Theprincipal water chemistry variables influencingspecies composition of periphytic diatoms were foundto be pH and water colour. Furthermore, therelationship between species abundance and pH wassufficiently strong to enable reconstruction of wateracidity from diatom data. Establishing the nature ofaquatic communities in atmospherically clean butgeologically sensitive regions is an important meansof identifying control systems against which therecovery of acidified lakes in polluted regions can beassessed. The Høylandet region has the potential toprovide a Europe-wide control system of this naturebut much further work is required to follow up andextend the results of this preliminary study.     

Vegetation and environment of the Rødalen alpine area, Central Norway, with emphasis on the early Holocene

Detailed pollen analysis and pine megafossils from the immediate area of Rødalen in Central Norway have revealed new knowledge of Holocene alpine environments. A period of about 1,000 years characterised by pioneer herbs, dwarf-shrubs (Betula nana, Empetrum) and Juniperus followed the Holocene climatic amelioration. Local birch forest became established around 10.3 ka b.p., ca 150 years earlier than the local pine rise. Pine dominated at 1,100 m a.s.l. from 9.9 to 8.5 ka b.p., followed by birch forests until 1.3 ka b.p. when deforestation occurred. Slightly after 6 ka b.p., pine forests disappeared from the valley floor (930 m a.s.l.), an area that today is dominated by birch forest. Three short-lasting vegetational set-backs at ca 10.7, 10.5 and 10.3 ka b.p. may indicate climate oscillations. A temporary reduction of local forests reflects the Erdalen 2/9.7 ka b.p. event. The influence of the 8.2 event, superimposed on a cooling trend, lasted ca 400 years and involved a two-step vegetational regression: (1) A strong reduction of pine forests due to cooling and (2) reduction of alder due to cold and drought. Winter stress preventing pine regeneration may have caused scarcity of pine megafossils from the latter period. In the early Holocene, vegetation in the present alpine region was not in equilibrium with temperature development. It is suggested that the birch forest establishment lagged by about 1,000 years due to drought, whereas winter stress may have delayed the establishment of pine even longer.     

Small-scale species patterns and associations in an alvar limestone grassland on Öland, Sweden

This paper reports on the results of a transect analysis carried out in order to detect small-scale patterns ofAvenetum species in a limestone grassland on the Great Alvar on the Baltic island of Öland, Sweden. Three transects consisting of 500 10×10 cm quadrats were used. The cover and number of flowering stalks (if any) of the 21 most common species were recorded. In addition the total cover of vascular plants, cover of mosses and lichens, cover of cow dung and soil depth in each quadrat were recorded. Ellenberg indicator values for light, moisture, reaction and nitrogen were calculated for each quadrat as weighted averages of the values for the species included in the analysis. Patch size frequency analysis, PASFRAN, a new method for detecting patches and their sizes, revealed that 42% of all significant species patches detected had mean patch sizes (diameter) in the range of 26–50 cm, which matched the size of dung patches—49 cm in transect 1 and 30 cm in transect 2. Results of association analysis revealed correlations for several species pairs and between species and environmental factors with similar patch sizes. Combination of PASFRAN and association analysis is recommended as an effective method for pattern detection.     

Correlated development, organism‐wide asymmetry and patterns of asymmetry in two moth species

Understanding the mechanisms that determine the development of a bilaterally symmetrical trait is crucial to the interpretation of patterns of fluctuating asymmetry (FA). Experimental and theoretical studies have indicated that feedback mechanisms both within and between developing traits, may participate in the developmental control of asymmetry. This study provides evidence that naturally occurring patterns of FA are affected by interactions between different traits. We found positive between‐trait correlations in signed FA values for tibia lengths on different legs, but not between wing and tibia FA in two moth species. Further research should investigate if trait functionality is related to this presumed correlated development. An extension of the Rashevsky–Turing model of morphogenesis further showed that correlations between the signed FA values can be generated by feedback mechanisms that regulate growth patterns between traits. We argue that such feedback mechanisms can be expected to be widespread and show that between‐trait correlations in the unsigned FA then become confounded with correlations in the signed FA. In addition, correlated development appeared to invalidate the use of the hypothetical repeatability to translate correlations between the unsigned FA values into correlations in the presumed underlying developmental instability. In conclusion, the presence of an organism‐wide asymmetry, which are most frequently found in morphologically integrated traits, may be even less common than previously thought. This revised version was published online in July 2006 with corrections to the Cover Date.     

Altitudinal patterns of plant species richness on the Baekdudaegan Mountains, South Korea: mid-domain effect, area, climate, and Rapoport’s rule

We studied the altitudinal patterns of plant species richness and examined the effects of geometric constraints, area, and climatic factors on the observed richness patterns along the ridge of the Baekdudaegan Mountains, South Korea. Rapoport’s altitudinal rule was evaluated by examining the relationship between altitudinal range size and midpoint. We also examined the latitudinal effect on species richness. Plant data were collected from 1,100 plots along a 200–1,900 m altitudinal gradient along the ridge of the Baekdudaegan. A total of 802 plant species from 97 families and 342 genera were found. The altitudinal patterns of plant species richness along the ridge of the Baekdudaegan depicted distinctly hump-shaped patterns, although the absolute altitudes of the richness peaks vary somewhat among plant groups. While the mid-domain effect (MDE) was the most powerful explanatory variable in simple regression models, species richness was also associated with climatic factors, especially mean annual precipitation (MAP) and temperature (MAT) in multiple regression models. The relative importance of the MDE and climatic factors were different among plant groups. The MDE was more important for woody plants and for large-ranged species, whereas climatic factors were better predictors for total and herbaceous plants and for small-ranged species. Rapoport’s altitudinal rule and a latitudinal effect on species richness were not supported. Our study suggests that a combined interaction of the MDE and climatic factors influences species richness patterns along the altitudinal gradient of the Baekdudaegan Mountains, South Korea.     

Reconstruction of the Allerød vegetation of the Neuwied Basin, western Germany, and its surroundings at 12,900 cal b.p.

During the late-glacial eruption of the Laacher See volcano 12,900 b.p. the whole area of the Neuwied Basin, middle Rhine region, was covered by several metres of the Laacher See Tephra (LST) damaging but also partly conserving the former vegetation. This gave the unique opportunity to study plant remains in situ and to estimate the spatial biodiversity of plant taxa within the area investigated at the end of the last glacial as if in a snapshot of 12,900 years ago. Different kinds of plant material can be found because of different preservation conditions namely imprints, charcoal and waterlogged remains. Investigations of this material from several places within the Neuwied Basin and the Brohl valley north of it allowed the reconstruction of the former vegetation at different scales from local to regional. This resulted in a large scale draft of a vegetation map of the studied area also based on the distribution and the stage of soil development at that time. As to whether thermophilous trees were already present is discussed but largely rejected. Electronic Supplementary Material Supplementary material is available for this article at     

How many species of Diacyclops? New taxonomic characters and species richness in a freshwater cyclopid genus (Copepoda,Cyclopoida)

The genus Diacyclops Kiefer, 1927 emend. Morton (1985) includes more than 100 species, widely distributed in all kinds of freshwater environments, and is the richest genus in the family Cyclopidae. Traditionally, Diacyclops species were defined according to differences in a few morphological characters; some characters (number of antennulary segments, segmentation pattern of swimming legs) are useful only in the separation of species groups, others (length of caudal rami and caudal setae) are highly variable even within the same population. During the study of the D. languidoides-group, minute morphological characters were used to differentiate between species (spinulation pattern and setation of antennary basis, setation and aesthetasc shape of male antennule, setation of mandibulary palp and maxilliped, shape of leg 4 basis) which allow to identify several valid species up to now concealed under the name `Diacyclops languidoides' (Lilljeborg, 1901). The coexistence of up to six congeneric species in the same sampling area: (a) supports the validity of the proposed taxonomic characters, (b) demonstrates that species richness may be highly underestimated in freshwater cyclopoid assemblages in absence of good taxonomic practice; and (c) requires an ecological explanation of species coexistence. The role of morphologically based taxonomy in order to solve general problems of distributional ecology and theoretical biology is explored.