Small-scale patterns of species richness in Swedish semi-natural grasslands: the effects of community species pools

Temperate semi-natural grasslands are known for their high plant species richness at small spatial scales. We examined the variation in small-scale species richness in a sample of 63 sites from Swedish semi-natural grasslands, located as fragments in the modem landscape dominated by forest and agricultural land. Data were obtained from two spatial scales at each site. 1 dm² and 4 m². Using an analysis based on a Monte Carlo simulation, we found support for the species-pool hypothesis: a high species richness at the I dm- scale was associated with high species richness at the 4 m2 scale. The conclusion from this pattern analysis would, however, be considerably strengthened if we could reduce the likelihood that other mechanisms than sampling from species pools of unequal size influence the pattern of small-scale species richness. Additional analyses were made in order to identify such mechanisms. We examined whether four putative key traits: seed size, seed production, plant size and reproductive allocation were different among species at comparatively species-rich vs species-poor I dm' plots. We found only a little evidence for such differences. There was a weak tendency that species in the plots with high species richness possessed larger (and fewer) seeds than species from species-poor plots. Our results are congruent with the main prediction of the species pool model: variation in small-scale species richness (1 dm²- scale) is basically a result of sampling from unequally sized community species pools (4 m² scale). Variation in species richness between the 4 nr semi-natural grassland "patches" may thus be sought for among mechanisms operating al larger spatial scales than 4 m². We briefly discuss such mechanisms, based on other studies performed in the same study area.     

Grain-dependent relationships between plant productivity and invertebrate species richness and biomass in calcareous grasslands

The relationships among productivity, species richness and consumer biomass are of fundamental importance for understanding determinants of biodiversity. These relationships may depend on grain size. We examined the relationships between productivity (above-ground phytomass) and plant species richness and between productivity and species richness and biomass of gastropods and grasshoppers using sampling units of different sizes (0.5, 2.75 and 23 m²) in nutrient-poor, calcareous grasslands in north-western Switzerland in two successive years. Species richness of forbs had a unimodal relationship with productivity in sampling units of 0.5 m² and was negatively correlated with productivity at the other two plot sizes in one year. In the other year, forb species richness tended to decrease with productivity in sampling units of 23 m². No similar relationship was found for grasses. Gastropod biomass had a unimodal relationship with productivity at 0.5 m² in the first year. Grasshopper species richness was correlated with forb species richness at plot sizes of 2.75 and 23 m². This study demonstrates that patterns detected between productivity and diversity and between productivity and biomass of consumers depend on the grain size used in the investigation and vary among years.Die Zusammenhänge zwischen Produktivität, Artenreichtum und Biomasse von Konsumenten sind wichtig, um zu verstehen, was Biodiversität beeinflußt. Diese Zusammenhänge können von der Größe der Untersuchungsfläche abhängig sein. Wir untersuchten während zwei aufeinanderfolgenden Jahren die Zusammenhänge zwischen Produktivität (oberirdische Pflanzenbiomasse) und Artenreichtum von Gefäßpflanzen, sowie zwischen Produktivität und Artenreichtum und Biomasse von Schnecken und Heuschrecken bezüglich dreier räumlicher Skalen (0,5, 2,75 und 23 m²) in Kalkmagerrasen in der Nordwestschweiz. Der Zusammenhang zwischen dem Artenreichtum von Kräutern und der Produktivität war unimodal in Flächeneinheiten von 0,5 m² und negativ in Flächeneinheiten von 2,75 und 23 m² im ersten Jahr und war tendenziell negativ in Flächeneinheiten von 23 m² im zweiten Jahr, während kein solcher Zusammenhang bei Gräsern gefunden wurde. Der Zusammenhang zwischen Produktivität und Biomasse von Schnecken war unimodal in Flächeneinheiten von 0,5 m² im ersten Jahr. Außerdem bestand ein Zusammenhang zwischen dem Artenreichtum von Kräutern und Heuschrecken in Flächeneinheiten von 2,75 und 23 m². Diese Arbeit zeigt, daß Zusammenhänge zwischen Produktivität und Diversität sowie zwischen Produktivität und Biomasse von Konsumenten von der Größe der Untersuchungsfläche abhängen und zwischen Jahren variieren.     

Effects of habitat area, isolation, and landscape diversity on plant species richness of calcareous grasslands

Calcareous grasslands harbour a high biodiversity, but are highly fragmented and endangered in central Europe. We tested the relative importance of habitat area, habitat isolation, and landscape diversity for species richness of vascular plants. Plants were recorded on 31 calcareous grasslands in the vicinity of the city of Göttingen (Germany) and were divided into habitat specialist and generalist species. We expected that habitat specialists were more affected by area and isolation, and habitat generalists more by landscape diversity. In multiple regression analysis, the species richness of habitat specialists (n = 66 species) and habitat generalists (n = 242) increased with habitat area, while habitat isolation or landscape diversity did not have significant effects. Contrary to predictions, habitat specialists were not more affected by reduced habitat area than generalists. This may have been caused by delayed extinction of long-living plant specialists in small grasslands. Additionally, non-specialists may profit more from high habitat heterogeneity in large grasslands compared to habitat specialists. Although habitat isolation and landscape diversity revealed no significant effect on local plant diversity, only an average of 54% of habitat specialists of the total species pool were found within one study site. In conclusion, habitat area was important for plant species conservation, but regional variation between habitats contributed also an important 46% of total species richness.     

Plant species richness in calcareous grasslands as affected by dispersability in space and time

Abstract. Species richness in calcareous grassland is discussed against the background of historical dispersal processes associated with traditional land-use management such as grazing, but also the artificial establishment by hayseed. Important vectors in the traditionally man-made landscape were sheep and cattle or other livestock such as goats. Calcareous grasslands were not only connected to each other but also to other habitats such as villages, forests, arable fields and heathlands by these vectors which could cover large distances (e.g. transhumance shepherding), which is not the case in the current man-made landscape. Species richness after restoration management of abandoned and afforested calcareous grasslands was predicted by using characters of dispersability in space and time. These were the persistence of the species in the vegetation and the diaspore bank after abandonment or afforestation and the dispersal capacity through wind and sheep. The results reveal that reintroduction of sheep grazing is necessary to reestablish the original species richness. The first validation of the prediction of the succession on clear-cut sites and a comparison with data of species composition in abandoned quarries and the surroundings made it obvious that a species' own dispersal capacity in space is very low except for some well wind-dispersed species. Therefore, it is recommended to include and to simulate dispersal processes in future management to be able to restore the original species richness.     

Multi-scale analysis of plant species richness in Serengeti grasslands

Aim To assess scale dependence between environmental factors and plant species richness. Additionally, we aimed to identify the scales at which niche relations and habitat heterogeneity, as hypothesized by A. Shmida & M.V. Wilson (1985) Journal of Biogeography, 12 , 1–20, operate in the savanna grasslands that were the focus of this study. Location Savanna grassland plant communities of Serengeti National Park, Tanzania. Methods Plant species richness was sampled in 102 modified Whittaker plots and tested for associations with two climate factors, mean annual rainfall (MAP) and potential evapotranspiration (PET), and two landscape variables, plot aspect (ASP) and topographic variation (TOPO), using multiple regressions. Scale dependence was assessed by conducting regressions after altering three aspects of spatial scale: grain, extent and focus. Grain was altered by analysing plant richness at 1, 10, 10² and 10³ m²; extent was investigated by restricting the maximum distance between samples to 75, 100, 125 and 150 km; and focus was manipulated by averaging samples spatially according to geographical land regions. Within the context of our data, we assumed that niche relations were represented by climate factors and habitat heterogeneity by landscape factors. Results Across all 102 plots, plant species richness between 1 and 10² m² had a negative relation to PET and a weak positive relation to MAP. Plant species richness at 10³ m² had a positive association with TOPO and weaker associations with climate factors. ASP stayed in the model between grains of 10 and 10³ m², but had a very weak positive association with richness. When the focus was changed to land regions, associations between plant species richness and explanatory variables strengthened, but were not qualitatively different. At spatial extents of 75 and 100 km, PET was the strongest correlate of plant species richness across all spatial grains. At spatial extents ≥ 125 km, PET explained the majority of the model variance at spatial grains ≤ 10² m², whereas TOPO explained equal amounts or more of the model variance at spatial grains of 10³ m². Main conclusions Both climate and topographic variation explained plant species richness in Serengeti grasslands, but specific patterns depended on grain, extent and, to a lesser degree, focus. Consistent with the ideas of Shmida & Wilson (1985) , determinants of plant species richness shifted from niche relations to habitat heterogeneity between spatial grains of 1 and 10³ m², although this occurred only at relatively large spatial extents (≥ 150 km). Finally, the signs, strength and shape of plant species richness relationships in Serengeti closely match those that describe macro‐scale patterns of woody plant species richness across the entire African continent.     

青海湖流域典型草地物种丰富度与生产力的关系

通过2009-2010年青海湖流域的样地实测数据,研究了不同尺度物种丰富度与生产力的关系,并初步探讨了其形成机制.结果表明:青海湖流域典型草地物种丰富度与生产力的关系在小尺度上(群落内和群落间)表现为线性正相关、负相关,在大尺度上(区域)则以单峰相关为主.沼泽草甸和高寒草甸的物种丰富度随生产力的增加而降低,高寒草原和温性草原的物种丰富度随生产力的增加而增加.在整个研究区内则表现为单峰型相关.研究还表明,研究区群落生产力的变化范围为65～585 g·m-2·a-1,物种丰富度为2～9种;生产力从高到低的顺序为沼泽草甸＞高寒草甸＞高寒草原＞温性草原.     

Small-scale dynamics and species richness in successional alvar plant communities

Species richness and small-scale spatial dynamics (turnover) were compared in four sites during 1990-1993 1) former alvar grassland, now young Pinus sylvestris forest, 2) clear-cut in 1990, 3) clear-cut and grazed by sheep, 4) nearby grassland It was hypothesized that in communities which show large-scale dynamics, small-scale dynamics is also more rapid to support succession Area-based species richness and shoot density increased in the cut and grazed site No significant dynamics was found in shoot-based species richness Therefore the forest was considered to be stable and the cut and grazed site large-scale dynamical (successional) Almost equally high small-scale turnover values were found in all sites, which supports the carousel model for successional communities, but means we had to reject the initial hypothesis The successional and intrinsic small-scale dynamics of the communities studied were two separate processes which acted at different scales, but the scale could not be only spatially determined, it depended also on the shoot density The overgrowing of this former alvar grassland has resulted in the decrease of shoot density and thus the decrease of species richness per unit area, but the total number of species has not changed much Consequently the possibility to restore grassland communities remains     

Species richness,alien species and plant traits in Central Argentine mountain grasslands

Abstract. Pattern of native vegetation, distribution of alien species and variation of environmental parameters were studied in mountain grasslands in a lithologically homogeneous Córdoba mountain range in Central Argentina. CCA showed that altitude was the most important factor determining the compositional variation of the vegetation, with soil nutrient status and stoniness as additional factors. Short‐grass communities, associated with the driest habitats on plateaus, showed higher small‐scale native species richness than wet‐turf communities in valleys and tall‐grass communities on slopes. Species richness was negatively correlated with soil parameters that indicate nutrient status and water availability. Also, there was a negative correlation between soil Ca‐ and Mg‐ content and richness. High native species richness coincided with high alien species richness. When smaller units – community types – were considered, it became evident that within short‐grass vegetation, the three most species‐rich community types contained significant numbers of alien species, while the other two did not. Even within one community type, the same quadrats that contained the highest number of native species, were also characterized by the highest numbers of alien species. Evidently, the same mechanism was responsible for high richness of both native and alien species. Alien species were distinguished by a greater proportion of annuals and prostrate stoloniferous plants, by lower palatability and by smaller proportion of zoochory. DCA ordination of quadrats on the basis of plant traits as attributes resulted in a clear distinction of three main vegetation types. Short‐grass vegetation was distinguished by a predomination of late flowering species, tall‐grass vegetation by the presence of high herbaceous plants and bushes, and wet‐turf vegetation by the presence of plants with storage organs, the lack of hairy leaves, and by a high proportion of cryptophytes. Quadrats with and without alien species were distinguished as well, indicating that the occurrence of aliens may be dependent on plant traits in a particular patch of a community.     

Control of plant species diversity and community invasibility by species immigration: seed richness versus seed density

Immigration rates of species into communities are widely understood to influence community diversity, which in turn is widely expected to influence the susceptibility of ecosystems to species invasion. For a given community, however, immigration processes may impact diversity by means of two separable components: the number of species represented in seed inputs and the density of seed per species. The independent effects of these components on plant species diversity and consequent rates of invasion are poorly understood. We constructed experimental plant communities through repeated seed additions to independently measure the effects of seed richness and seed density on the trajectory of species diversity during the development of annual plant communities. Because we sowed species not found in the immediate study area, we were able to assess the invasibility of the resulting communities by recording the rate of establishment of species from adjacent vegetation. Early in community development when species only weakly interacted, seed richness had a strong effect on community diversity whereas seed density had little effect. After the plants became established, the effect of seed richness on measured diversity strongly depended on seed density, and disappeared at the highest level of seed density. The ability of surrounding vegetation to invade the experimental communities was decreased by seed density but not by seed richness, primarily because the individual effects of a few sown species could explain the observed invasion rates. These results suggest that seed density is just as important as seed richness in the control of species diversity, and perhaps a more important determinant of community invasibility than seed richness in dynamic plant assemblages.     

Declining plant species richness in the tussock grasslands of Canterbury and Otago, South Island, New Zealand

We studied vegetation change on 142 permanently marked transects spread throughout tussock grasslands of Otago and Canterbury, in areas subject to both pastoral and conservation management. The transects were established between 1982 and 1986 and remeasured between 1993 and 1999, providing a record of vegetation change at each site over an interval varying from 10 to 15 years. Each transect consisted of 50 quadrats, each 0.25m(2), in which the presence of all vascular plant species had been recorded. For each transect, we calculated the change between measurements in the mean number of species recorded per quadrat, and the change in the total number of species recorded per transect. Averaged across all transects, there was a significant decline in species richness between measurements at both the quadrat and transect scales. Small herbs (those less than or equal to2 cm tall, excluding Hieracium species) showed the greatest decline. On average, more than one quarter of the small herb species present in a quadrat at the first measurement had disappeared within 10 years. Larger herbs, ferns, rushes, sedges and grasses (excluding Chionochloa species) also declined significantly in species richness, reflecting declines in the abundance of species in these groups. Woody species richness remained constant, while species in the genera Chionochloa and Hieracium increased significantly in mean quadrat species richness, reflecting increases in the abundance of these species along transects. The rate of decline in mean quadrat species richness was unrelated to changes in the abundance of either Chionochloa or Hieracium species, or to an overall increase in total vegetation cover on transects. The rate of decline in species richness was also unrelated to the level of grazing or burning between measurements. However, the rate of decline in species richness was greater at lower elevation, on schist rock and on yellow-brown and yellow-grey soils. Our results suggest that a major compositional change is occurring in these grasslands at a rate that is independent of local variation in management and independent of the widespread invasion of these grasslands by Hieracium species.     

Small-scale species richness in forest canopy gaps: the role of niche limitation versus the size of the species pool

Abstract. The form of the relationship between local species richness and the number of species in the surrounding region can be used as a test between competing theories of community structure. For 32 canopy gaps in New Zealand Nothofagus forest, we examined the relationship between the number of vascular plant species in 0.2-m² quadrats within the gap and the species richness of the whole gap. We found no evidence that competition for a limited number of niches placed an upper limit on the number of locally co-occurring species. Rather, the mean number of species in quadrats within canopy gaps increased in direct proportion to gap species richness. This relationship held after we controlled for potentially confounding factors, including variation in forest floor substrate, and gap size, age, shape and orientation. Our results suggest that even over relatively small spatial scales, local species richness can be constrained by the size of the species pool in the immediately surrounding region.     

局域和区域过程共同控制着群落的物种多样性:种库假说

解释群落的物种多样性大小是生态学研究的一个重要的理论和实践问题。人们提出了群落物种多样性的多种假说, Zobel等人提出的种库假说(species pool hypothesis)是生物多样性理论研究的重要发展。该假说认为, 一个群落的物种多样性不仅与环境条件和生态过程(ecological process)(如竞争、捕食)有关, 也受区域种库(regional species pool)的限制。区域种库是指一个地区可进入某一群落的潜在物种数量, 它由地史过程(如冰期、地质年代)和区域过程(物种形成、迁移扩散以及消亡)所决定。按照种库假说, 某一生境类型的面积越大, 地质年代越古老, 物种形成的机会也就越多, 因而能适应和分布于该生境的物种也就越多, 实际群落中的物种丰富度也就越高。种库在空间上主要有两个层次: 区域种库和实际种库, 前者指某一生境所拥有的潜在物种数量, 主要由生物地理过程(biogeographic processes)所决定; 后者则为调查的群落中实际出现的物种数量, 主要由竞争等生态过程和区域种库共同决定。本文对种库假说的基本概念、主要内容、种库确定方法等作了介绍, 并阐述了作者对这些问题的理解和认识。     

On the possible role of local effects on the species richness of acidic and calcareous rock grasslands in northern Hungary

Ewald (Folia Geobot. 38: 357–366, 2003, this issue) stated that in Central Europe the number of calcifrequent species is higher than the number of acidofrequent species, while the range of acidofrequent communities is larger than that of the calcifrequent ones. All the explanations considered in his paper are based on an evolutionary spatial and temporal scale. In this paper we are trying to prove that local effects might also be important. Five open rock grassland communities on different bedrocks, viz. rhyolite (acidic), andesite (slightly acidic), calcareous sandstone (slightly calcareous), limestone (calcareous) and dolomite (calcareous) were chosen for the analysis. Two parameters of the species-area curve (i.e., local richness and the slope of log area-species richness line) were estimated based on all species and on rock specialist species separately. With this method we could simultaneously study three attributes of diversity: local species number, the slope of log area-species richness line, and species pool size.     

Changes in plant species richness in a calcareous grassland following changes in environmental conditions

This study presents results of a three and a half year experiment in a calcareous grassland on the Baltic island of Öland, south-eastern Sweden. Environmental conditions, viz. nutrient regime, water availability, light conditions and grazing pressure were changed in a randomised experimental design with ten different combinations of these factors, including a control. Weather conditions during the study period fluctuated considerably. Only the combination of reduced light and increased soil nutrient availability decreased species richness significantly, presumably by inducing asymmetric competition for light by competitively strong species. Physical space also became less available for weaker competitors. In cases where only nutrients or only water were supplied in sufficient amounts, competitive exclusion was inhibited by either low nutrient or low water availability. Under conditions which combined high levels of nutrient and water supply with regular disturbance (i.e. simulated grazing) high species richness was maintained. Although there was a significant negative correlation between species richness and amount of above-ground biomass, the number of species was not significantly reduced in the treatment which combined fertilisation and irrigation. One reason for this may be that the heterogeneous light regime in this treatment still allowed species persistence and coexistence.     

Kin selection,species richness and community

Can evolutionary and ecological dynamics operating at one level of the biological hierarchy affect the dynamics and structure at other levels? In social insects, strong hostility towards unrelated individuals can evolve as a kin-selected counter-adaptation to intraspecific social parasitism. This aggression in turn might cause intraspecific competition to predominate over interspecific competition, permitting coexistence with other social insect species. In other words, kin selection—a form of intra-population dynamics—might enhance the species richness of the community, a higher-level structure. The converse effect, from higher to lower levels, might also operate, whereby strong interspecific competition may limit the evolution of selfish individual traits. If the latter effect were to prove more important, it would challenge the common view that intra-population dynamics (via individual or gene selection) is the main driver of evolution.