Determinants of avian species richness at different spatial scales

ABSTRACT. Studies of factors influencing avian biodiversity yield very different results depending on the spatial scale at which species richness is calculated. Ecological studies at small spatial scales (plot size 0.0025–0.4 km²) emphasize the importance of habitat diversity, whereas biogeographical studies at large spatial scales (quadrat size 400–50,000 km²) emphasize variables related to available energy such as temperature. In order to bridge the gap between those two approaches the bird atlas data set of Lake Constance was used to study factors determining avian species diversity at the intermediate spatial scales of landscapes (quadrat size 4–36 km²). At these spatial scales bird species richness was influenced by habitat diversity and not by variables related to available energy probably because, at the landscape scale, variation in available energy is small. Changing quadrat size between 4 and 36 km², but keeping the geographical extension of the study constant resulted in profound changes in the degree to which the amount of different habitat types was correlated with species richness. This suggests that high species diversity is achieved by different management regimes depending on the spatial scale at which species richness is calculated. However, generally, avian species diversity seems to be determined by spatial heterogeneity at the corresponding spatial scale. Thus, protecting the diversity of landscapes and ecosystems appears to ensure also high levels of species diversity.     

From plant neighbourhood to landscape scales: how grazing modifies native and exotic plant species richness in grassland

The interactive effect of grazing and soil resources on plant species richness and coexistence has been predicted to vary across spatial scales. When resources are not limiting, grazing should reduce competitive effects and increase colonisation and richness at fine scales. However, at broad scales richness is predicted to decline due to loss of grazing intolerant species. We examined these hypotheses in grasslands of southern Australia that varied in resources and ungulate grazing intensity since farming commenced 170 years ago. Fine-scale species richness was slightly greater in more intensively grazed upper slope sites with high nutrients but low water supply compared to those that were moderately grazed, largely due to a greater abundance of exotic species. At broader scales, exotic species richness declined with increasing grazing intensity whether nutrients or water supply were low or high. Native species richness declined at all scales in response to increasing grazing intensity and greater resource supply. Grazing also reduced fine-scale heterogeneity in native species richness and although exotics were also characterised by greater heterogeneity at fine scales, grazing effects varied across scales. In these grasslands patterns of plant species richness did not match predictions at all scales and this is likely to be due to differing responses of native and exotic species and their relative abundance in the regional species pool. Over the past 170 years intolerant native species have been eliminated from areas that are continually and heavily grazed, whereas transient, light grazing increases richness of both exotics and natives. The results support the observation that the processes and scales at which they operate differ between coevolved ungulate—grassland systems and those in transition due to recent invasion of herbivores and associated plant species.     

Effects of restoration with cattle grazing on plant species composition and richness of semi-natural grasslands

Restoration of semi-natural grasslands by cattle grazing is among the most practical options for reversing the decline of northern European floristic diversity, but no studies on this subject are available. In this work the success of restoration of abandoned, privately owned mesic semi-natural grasslands by farmers receiving support from the EU agri-environmental support scheme was studied in southwestern Finland. Three kinds of grasslands were compared: old (continuously cattle grazed), new (cattle grazing restarted 3–8 years ago) and abandoned pastures (grazing terminated >10 years ago). Plant species composition of the three pasture types was floristically different in multivariate analyses (non-metric multidimensional scaling). Total species richness, richness of grassland plants, indicator plants and rare plants were highest in old and lowest in abandoned pastures in all studied spatial scales (0.25–0.8 ha, 1 and 0.01 m²). The results were congruent with different scales and species list definitions, suggesting that species density scale (1 m²) can be used as a partial surrogate for large scale species richness. Species richness of new pastures was 20% higher on 0.25–0.8 ha, 40–50% higher on 1 m² and 30% higher on the 0.01 m² scale compared to abandoned grasslands. Rare species showed insignificant response to resumed grazing. Despite problems in management quality, this study showed promising results of restoration of abandoned grasslands by cattle grazing on private farms. However, populations of several rare grassland plants may not recover with present cattle grazing regimes. Management regulations in the agri-environmental support scheme need to be defined more precisely for successful restoration.     

放牧干扰下高寒草甸物种和生活型丰富度与地上及地下生物量的关系

明晰放牧干扰下高寒草甸植物丰富度与生物量的相关关系,为草地植物不同生长时期生物量的预测提供依据。设置6个放牧强度样地,连续3a放牧,2014年进行3个季节(6月、8月、10月)的植物丰富度和地上、地下生物量调查,对比分析放牧干扰下物种和生活型丰富度(生活型的种类)分别与地上、地下生物量的相关关系。结果表明:(1)物种和生活型丰富度与地上生物量均受放牧强度的显著影响,物种丰富度仅在8月与放牧强度显著负相关,生活型丰富度在10月随放牧强度单峰变化,地上生物量在不同季节均与放牧强度显著负相关,而地下生物量与放牧强度无关。(2)物种丰富度与地上和地下生物量均受季节的显著影响,物种丰富度和地上生物量仅在低强度放牧区随季节呈单峰变化,地下生物量在中等强度放牧区随季节呈单峰变化;生活型丰富度与季节无关。(3)放牧干扰前物种和生活型丰富度与地上和地下生物量均显著正相关。3a放牧后仅在8月,物种丰富度只与地上生物量显著正相关,生活型丰富度与地上和地下生物量均显著正相关。(4)对于不同放牧强度,物种丰富度仅在低强度放牧区与地上生物量显著正相关,而生活型丰富度在所有放牧强度区均与地上生物量显著正相关。综上所述,放牧干扰扰乱了高寒草甸丰富度与生物量之间的关系,尤其影响了物种丰富度与地下生物量之间的相关关系。生活型丰富度与地上生物量之间的显著关系不受放牧强度干扰,使生活型丰富度在预测生物量方面表现出优势。     

Environmental heterogeneity,species diversity and co‐existence at different spatial scales

The positive relationship between spatial environmental heterogeneity and species diversity is a widely accepted concept, generally associated with niche limitation. However, niche limitation cannot account for negative heterogeneity–diversity relationships (HDR) revealed in several case studies. Here we explore how HDR varies at different spatial scales and provide novel theories for small‐scale species co‐existence that explain both positive and negative HDR. At large spatial scales of heterogeneity (e.g. landscape level), different communities co‐exist, promoting large regional species pool size and resulting in positive HDR. At smaller scales within communities, species co‐existence can be enhanced by increasing the number of different patches, as predicted by the niche limitation theory, or alternatively, restrained by heterogeneity. We conducted meta‐regressions for experimental and observational HDR studies, and found that negative HDRs are significantly more common at smaller spatial scales. We propose three theories to account for niche limitation at small spatial scales. (1) Microfragmentation theory: with increasing spatial heterogeneity, large homogeneous patches lose area and become isolated, which in turn restrains the establishment of new plant individuals and populations, thus reducing species richness. (2) Heterogeneity confounded by mean: when heterogeneity occurs at spatial scales smaller than the size of individual plants, which forage through the patches, species diversity can be either positively or negatively affected by a change in the mean of an environmental factor. (3) Heterogeneity as a separate niche axis: the ability of species to tolerate heterogeneity at spatial scales smaller than plant size varies, affecting HDR. We conclude that processes other than niche limitation can affect the relationship between heterogeneity and diversity.     

Enhancement of habitat heterogeneity and species richness on rocky shores inundated by sand

Summary Many rocky shores are subject to periodic inundation by sand, which is often thought to reduce species richness by eliminating organisms intolerant of sand scour or sand smothering. However, regular disturbance (e.g. inundation) should promote richness by preventing the development of low diversity climax communities. A study of faunal richness on 10 regularly inundated shores showed that inundation does promote richness, but by increasing habitat heterogeneity. Some species are excluded from parts of the shore by sand, but because of the patchiness of sand deposits they are rarely excluded from the entire shore. Other species are found only on rocks associated with sand, while typically sandy shore animals occur in the sand deposits themselves. Total richness (281 species) was greater than for local noninundated shores and sandy beaches combined.     

放牧和刈割条件下草山草坡群落空间异质性分析

采用变异矩分析和分形方法,研究了草山草坡群落在放牧和刈割条件下的空间异质性及空间自相关性,结果表明,群落空间格局有尺度依赖性,刈割条件下空物异质性及空间相关性弱,多样性梯度即β多样性小,放牧消除地形引起的样地差异,因而使空间异质性简单化。     

不同放牧强度下短花针茅荒漠草原植被的空间异质性

分析不同放牧强度下植物群落中物种的空间分布特征, 有助于阐明群落在放牧胁迫下的演替规律。该研究基于幂函数法则, 探讨了不同放牧强度下短花针茅(Stipa breviflora)荒漠草原群落植物的频率和空间异质性。结果表明: 不同放牧强度下物种空间分布与幂函数法则能很好地吻合; 不同物种空间异质性具有特异性, 随着放牧强度的增加, 提高群落空间异质性的物种分别由无芒隐子草(Cleistogenes songorica)、冷蒿(Artemisia frigida)、短花针茅、银灰旋花(Convolvulus ammannii)等多个物种逐渐转变为以无芒隐子草、短花针茅为主的少数物种, 同时, 物种空间异质性大于群落空间异质性的物种数逐渐减少。     

Effects of spatial heterogeneity on butterfly species richness in Rocky Mountain National Park, CO, USA

We investigated butterfly responses to plot-level characteristics (plant species richness, vegetation height, and range in NDVI [normalized difference vegetation index]) and spatial heterogeneity in topography and landscape patterns (composition and configuration) at multiple spatial scales. Stratified random sampling was used to collect data on butterfly species richness from seventy-six 20 × 50 m plots. The plant species richness and average vegetation height data were collected from 76 modified-Whittaker plots overlaid on 76 butterfly plots. Spatial heterogeneity around sample plots was quantified by measuring topographic variables and landscape metrics at eight spatial extents (radii of 300, 600 to 2,400 m). The number of butterfly species recorded was strongly positively correlated with plant species richness, proportion of shrubland and mean patch size of shrubland. Patterns in butterfly species richness were negatively correlated with other variables including mean patch size, average vegetation height, elevation, and range in NDVI. The best predictive model selected using Akaike’s Information Criterion corrected for small sample size (AIC_c), explained 62% of the variation in butterfly species richness at the 2,100 m spatial extent. Average vegetation height and mean patch size were among the best predictors of butterfly species richness. The models that included plot-level information and topographic variables explained relatively less variation in butterfly species richness, and were improved significantly after including landscape metrics. Our results suggest that spatial heterogeneity greatly influences patterns in butterfly species richness, and that it should be explicitly considered in conservation and management actions.     

Fine-scale spatial variation in plant species richness and its relationship to environmental conditions in coastal marshlands

Previous studies have shown that variations in environmental conditions play a major role in explaining variations in plant species richness at community and landscape scales. In this study, we considered the degree to which fine-scale spatial variations in richness could be related to fine-scale variations in abiotic and biotic factors. To examine spatial variation in richness, grids of 1 m² plots were laid out at five sites within a coastal riverine wetland landscape. At each site, a 5 × 7 array of plots was established adjacent to the river’s edge with plots one meter apart. In addition to the estimation of species richness, environmental measurements included sediment salinity, plot microelevation, percent of plot recently disturbed, and estimated community biomass. Our analysis strategy was to combine the use of structural equation modeling (path modeling) with an assessment of spatial association. Mantel’s tests revealed significant spatial autocorrelation in species richness at four of the five sites sampled, indicating that richness in a plot correlated with the richness of nearby plots. We subsequently considered the degree to which spatial autocorrelations in richness could be explained by spatial autocorrelations in environmental conditions. Once data were corrected for environmental correlations, spatial autocorrelation in residual species richness could not be detected at any site. Based on these results, we conclude that in this coastal wetland, there appears to be a fine-scale mapping of diversity to microgradients in environmental conditions.     

Spatial heterogeneity in parasite infections at different spatial scales in an intertidal bivalve

Spatial heterogeneities in the abundance of free-living organisms as well as in infection levels of their parasites are a common phenomenon, but knowledge on parasitism in invertebrate intermediate hosts in this respect is scarce. We investigated the spatial pattern of four dominant trematode species which utilize a common intertidal bivalve, the cockle Cerastoderma edule, as second intermediate host in their life cycles. Sampling of cockles from the same cohort at 15 sites in the northern Wadden Sea (North Sea) over a distance of 50 km revealed a conspicuous spatial heterogeneity in infection levels in all four species over the total sample as well as among and within sampling sites. Whereas multiple regression analyses indicated the density of first intermediate upstream hosts to be the strongest determinant of infection levels in cockles, the situation within sites was more complex with no single strong predictor variable. However, host size was positively and host density negatively correlated with infection levels and there was an indication of differential susceptibility of cockle hosts. Small-scale differences in physical properties of the habitat in the form of residual water at low tide resulted in increased infection levels of cockles which we experimentally transferred into pools. A complex interplay of these factors may be responsible for within-site heterogeneities. At larger spatial scales, these factors may be overridden by the strong effect of upstream hosts. In contrast to first intermediate trematode hosts, there was no indication for inter-specific interactions. In other terms, the recruitment of trematodes in second intermediate hosts seems to be largely controlled by pre-settlement processes both among and within host populations.     

Habitat patchiness and plant species richness

The pattern of woody species richness decline with a decrease in woody vegetation cover was studied within a tallgrass prairie. The decline in species richness is highly non-linear, with a well-defined threshold below which species richness collapses. This relationship can be understood after considering information on how landscape structure changes with woody vegetation cover, and how species richness is related to landscape structure.     

Landscape heterogeneity metrics as indicators of bird diversity: Determining the optimal spatial scales in different landscapes

Species distribution models are often used to study the biodiversity of ecosystems. The modelling process uses a number of parameters to predict others, such as the occurrence of determinate species, population size, habitat suitability or biodiversity. It is well known that the heterogeneity of landscapes can lead to changes in species’ abundance and biodiversity. However, landscape metrics depend on maps and spatial scales when it comes to undertaking a GIS analysis.We explored the goodness of fit of several models using the metrics of landscape heterogeneity and altitude as predictors of bird diversity in different landscapes and spatial scales. Two variables were used to describe biodiversity: bird richness and trophic level diversity, both of which were obtained from a breeding bird survey by means of point counts. The relationships between biodiversity and landscape metrics were compared using multiple linear regressions. All of the analyses were repeated for 14 different spatial scales and for cultivated, forest and grassland environments to determine the optimal spatial scale for each landscape typology.Our results revealed that the relationships between species’ richness and landscape heterogeneity using 1:10,000 land cover maps were strongest when working on a spatial scale up to a radius of 125–250 m around the sampled point (circa 4.9–19.6 ha). Furthermore, the correlation between measures of landscape heterogeneity and bird diversity was greater in grasslands than in cultivated or forested areas. The multi-spatial scale approach is useful for (a) assessing the accuracy of surrogates of bird diversity in different landscapes and (b) optimizing spatial model procedures for biodiversity mapping, mainly over extensive areas.     

Plant species richness and spatial organization at different small scales in western Mediterranean landscapes

The relationship between plant species richness and the space organization of the community at different small scales was studied. The study was based on 51 sites distributed along a belt from Central Spain to Portugal. Each site was analyzed with a transect cutting across the boundary between two neighboring patches of shrubland and grassland. Local spatial organization of vegetation was analyzed at different levels of detail and each transect was divided into successively smaller portions. The first division coincides with a physiognomic perception of the site in two patches (shrubland and grassland). The average spatial niche width of the species was used to calculate the spatial organization of the vegetation of each division in each site. The correlation between species richness and spatial organization depended on the block size under consideration. A physiognomic criterion, sectorizing the sites into patches of shrubland and grassland, determines noteworthy floristic changes but does not enable us to express satisfactorily the variability in plant richness. In order to account for this variation, other factors must be taken into account which act at a more detailed small-scale and which determine the internal variability of these patches. In the case studied, the species richness of the sites increases along with an increase in the percentage of species whose occupation of the space is relatively restricted within the site. Many of these species are, however, frequent within the whole of the territory studied. The results highlight the importance of the level of local scale at which the factors influencing occupation of the space, and consequentially, plant richness, preferentially act. This circumstance ought to be taken into consideration in strategies for the conservation of biological diversity, and based on the delimitation of protected spaces with criteria frequently linked to the physiognomy of the vegetation.Nomenclature: Follows T.G. Tutin et al. 1964-1980. Flora Europaea. Cambridge University Press, Cambridge     

Soil pH and species diversity in coastal dunes

Soil pH was measured at two different spatial scales in coastal dunes on Norderney, North Sea, and in Mecklenburg-Vorpommern, Baltic Sea, Germany. Relationships between the variability in soil pH, species richness and species diversity are presented. Species richness and diversity were highest in grey dunes, where soil pH was at intermediate levels; both variables were lower in yellow and brown dunes. The variability in pH increased with increasing species diversity and also with scale. Overall, soil pH variability decreased with increasing vegetation cover. The lowest pH heterogeneity was found in heath dominated by Empetrum nigrum L. and grey dunes dominated by Campylopus introflexus (Hedw.) Brid. Increasing abundance of dominant species and decreasing species diversity of vegetation apparently reduces soil heterogeneity. Decreasing species diversity of vegetation is likely to explain decreasing variability in soil pH.     

Vole mound effects and disturbance rate in a mediterranean plant community under different grazing and irrigation regimes

A factorial field experiment was used to assess the influence of soil-disturber mammals in the structure of a 9-year-old Mediterranean annual plant community subjected to different sheep grazing and irrigation regimes. We estimated the disturbance rate (mound building activity) by Mediterranean voles, their effects on vegetation and the mechanisms of these effects during a period of vole outbreak. The effects on vegetation were analysed at the levels of species, functional groups and plant community. Disturbance rate was high and voles can disturb the entire soil surface once every four or five years. The availability of certain trophic resources (perennial plants) appeared to drive vole expansion in the experimental plots and it was independent of the irrigation and grazing treatments. Mound building activities largely affected vegetation but conserved plot differences. Total vegetation cover, absolute cover of all functional groups, mean vegetation height and species richness were less on mounds than on undisturbed ground. These effects did not change the relative abundance of annuals, perennials, grasses and forbs. Only the relative abundance of small-seeded species decreased on mounds. As the proportion of these seeds was similar in both types of patches, we suggest that small-seeded species had more difficulties for germinating or emerging when they are buried during mound formation. Irrigation and sheep grazing promoted large changes in the vegetation parameters but these effects were, in general, similar on mounds and undisturbed ground. Our results show that the availability of germinable seeds may be the major limitation for mound revegetation, probably due to the scarcity of seeds existing at the depths from which soils are excavated. Our results also suggested a resource limitation on mounds. The results provide additional evidence that soil disturbances by small herbivore mammals exert relevant ecological effects on abandoned Mediterranean croplands. We discuss the ecological implications of vole mound-building activities for plant succession, plant species conservation and forage resource availability for livestock. This revised version was published online in June 2006 with corrections to the Cover Date.     

Year-round cattle and horse grazing supports the restoration of abandoned,dry sandy grassland and heathland communities by supressing Calamagrostis epigejos and enhancing species richness

The spread of competitive grasses, changes in species composition and vegetation structure are direct consequences of grassland and heathland abandonment. As an alternative to more costly management measures such as traditional pastoralism, year-round low-intensity grazing with large herbivores is increasingly used to restore and maintain semi-open habitats. However, the suitability of this grazing regime has not yet been investigated for long-abandoned, highly degraded but nutrient-poor sandy grassland and heathland communities. In particular, it is unclear if year-round grazing is suitable for preventing the further spread of highly competitive grasses such as Calamagrostis epigejos while simultaneously maintaining or improving characteristic species richness and vegetation structure. Hence, we conducted a comprehensive field study on two spatial scales (plot-level: 25 m², macroplot-level: 1 ha) to analyse the impacts of year-round low-intensity cattle and horse grazing on the development of the highly competitive grass Calamagrostis epigejos, as well as the vegetation structure and plant species richness of long-abandoned but nutrient-poor dry sandy grassland and heathland communities, their mosaics and Calamagrostis stands within an 800 ha heathland between 2008 and 2015. Finally, we assessed the local conservation status of the habitat types after seven years of grazing in comparison to long-abandoned sites.Grazing successfully reduced the coverage of Calamagrostis epigejos, whereby Calamagrostis stands developed towards species-rich sandy grasslands after seven years of grazing. In addition, the quality of the vegetation structure was improved by enhancing the proportion of bare soil, while litter and grass cover, litter thickness and height of the field layer as well as the coverage of ruderal indicators were significantly reduced on grazed sites in comparison to ungrazed sites in 2015. Moreover, we found an overall positive grazing effect on species richness: Total species number, number of target species as well as subordinated target species significantly increased within the vegetation types over time.Thus, year-round low-intensity cattle and horse grazing is a suitable management tool for restoring, maintaining and even improving long-abandoned, nutrient-poor sandy grassland and heathland communities, and thus to enhance the local conservation status of the habitat types. However, if there is a high initial cover of woody species (e.g. shrubs, tree rejuvenation), then an extensive shrub and tree clearance will be necessary, with manual shrub cutting being crucial to reduce the heavy regrowth of the woody species. In addition, a one-time mowing should be implemented in highly degraded heaths to facilitate the vegetative rejuvenation of degenerate stands of Calluna vulgaris, thus improving its attractiveness for the grazing animals.     

The reliability of a composite biodiversity indicator in predicting bird species richness at different spatial scales

Several biodiversity features can be linked to landscape heterogeneity, that, in turn, can be informative for management and conservation purposes. Usually, the more the landscape is complex the more the biodiversity increases. Biodiversity indicators can be a useful tool to assess biodiversity status, in function of landscape heterogeneity. In this study, we developed a biodiversity indicator, based on Shannon diversity index and built from distribution maps of protected species. With such an approach, we seek to evaluate the feasibility of using a combination of target species as a surrogate for assessing the status of the whole bird community. Our approach was spread over multiple spatial scales, to determine which was the most informative. We selected four species protected by European regulation and generated a presence-absence map from species distribution modelling. We, therefore, used the FRAGSTATS biodiversity metric to calculate Shannon index for the overlapped presence-absence maps, at two spatial scales (500 m and 1000 m). Then, the relationships with the whole community was assessed through generalised least square models, at the spatial scale of 4 ha, 9 ha and 25 ha. Results showed that the higher rate of variability of community was explained by the biodiversity indicator at 1000 m scale. Indeed, the more informative spatial scale for the whole bird community was 9 ha. In addition, a pattern emerged about the relationships between biodiversity indicator and community richness, that is worth of further research. Our study demonstrates that the usefulness of surrogate species for biodiversity and community assessment can become clear only at a certain spatial scales. Indeed, they can be highly predictive of the whole community, and highly informative for conservation planning. Moreover, their use can optimize biodiversity monitoring and conservation, focusing on a small number of noteworthy species.