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.     

Spatial heterogeneity and plant species richness at different spatial scales under rabbit grazing

Herbivores influence spatial heterogeneity in soil resources and vegetation in ecosystems. Despite increasing recognition that spatial heterogeneity can drive species richness at different spatial scales, few studies have quantified the effect of grazing on spatial heterogeneity and species richness simultaneously. Here we document both these variables in a rabbit-grazed grassland. We measured mean values and spatial patterns of grazing intensity, rabbit droppings, plant height, plant biomass, soil water content, ammonia and nitrate in sites grazed by rabbits and in matched, ungrazed exclosures in a grassland in southern England. Plant species richness was recorded at spatial scales ranging between 0.0001 and 150 m(2). Grazing reduced plant height and plant biomass but increased levels of ammonia and nitrate in the soil. Spatial statistics revealed that rabbit-grazed sites consisted of a mixture of heavily grazed patches with low vegetation and nutrient-rich soils (lawns) surrounded by patches of high vegetation with nutrient-poor soils (tussocks). The mean patch size (range) in the grazed controls was 2.1 +/- 0.3 m for vegetation height, 3.8 +/- 1.8 m for soil water content and 2.8 +/- 0.9 m for ammonia. This is in line with the patch sizes of grazing (2.4 +/- 0.5 m) and dropping deposition (3.7 +/- 0.6 m) by rabbits. In contrast, patchiness in the ungrazed exclosures had a larger patch size and was not present for all variables. Rabbit grazing increased plant species richness at all spatial scales. Species richness was negatively correlated with plant height, but positively correlated to the coefficient of variation of plant height at all plot sizes. Species richness in large plots (<25 m(2)) was also correlated to patch size. This study indicates that the abundance of strong competitors and the nutrient availability in the soil, as well as the heterogeneity and spatial pattern of these factors may influence species richness, but the importance of these factors can differ across spatial scales.     

Surveying carnivores at large spatial scales: a comparison of four broad-applied methods

Reliable methods to estimate species richness are very important to managers and conservationists because they provide key data to make the right decisions in conservation programmes. In the case of carnivore mammals, traditional methods, such as direct count censuses, are not useful since these animals are usually scarce, elusive and nocturnal. Difficulties in carnivore sampling are compounded when monitoring programmes are developed at large spatial scales, where high economic costs and field efforts are necessary to achieve reliable richness or abundance estimates. These problems have highlighted the need to find more effective carnivore survey methods, especially in regions with high rates of landscape change, such as the Mediterranean basin. The present study, performed in a typical Mediterranean area, was the first in Europe to test simultaneously the relative efficiencies of four broad-applied sampling methods to detect carnivore species at large spatial scales. Sign surveys based on scat detection, scent stations, camera-trapping and live-trapping were investigated. We compared efficiencies using biological parameters and by considering both the logistic and economic costs of each method. Overall, scent stations and sign surveys were the most efficient methods both in economic and logistic terms. In addition, the use of scent stations may be necessary to detect species rarely detected by scats. Detailed and extensive training programmes for conducting sign surveys and scent stations may overcome perceived problems thus enhancing the widespread use of both methods. Our results are applicable not only to other Mediterranean areas, but also to other habitats and regions of the world. More research into the suitability of these and other methods in relation to different landscapes, seasons and species is required.     

祁连山区不同海拔草地群落的物种多样性

依据植物在各海拔带出现的频率和不同海拔带植物种的生活型,对祁连山区草地群落海拔2500～3000m的物种α多样性变化特征进行了研究．结果表明,随海拔升高,Shahnon-Wiener指数从1．432上升到1．832,Simpson指数的变化较为复杂．海拔2701～2800m带是物种多样性的一个重要转折点,是物种多样性较丰富的地带．在祁连山草地群落中,有毒的与食性较差的物种在各海拔带所占比例均超过50％,生物量占到整个样地生物量的60％以上,物种多样性已呈现衰退．     

Effects of the expansion by Hippophaë rhamnoides on plant species richness in coastal dunes

Question: Is the expansion of Hippophaë rhamnoides in coastal dunes associated with a decline in plant species richness, and is this decline best described by a hump‐backed relationship between species number and shrub cover? Location: Grey and yellow dunes on the East Frisian islands Spiekeroog and Norderney. Methods: Total plant species richness as well as the number of herbaceous and cryptogam species were determined in 2001 using plots of 16 m² size. We compared shrubland plots with varying cover of Hippophaë with neighbouring dune grassland plots without shrubs as reference sites. Soil samples were collected to determine the values of some important edaphic variables (pH, organic matter, nitrogen). Results: The shrubland plots with Hippophaë had or tended to have lower soil pH and C/N ratios and higher contents of organic matter and nitrogen than the grassland plots. Total species richness was marginally significantly related to the cover of Hippophaë in a hump‐backed manner on both islands. The pattern was more pronounced for mosses and lichens than for herbaceous species. For all species groups on Spiekeroog and for the herbaceous species on Norderney, the hump‐backed relationship was much improved when using the difference in species number between shrubland and grassland plot as a dependent variable. Relationships could be improved by including the soil parameters as co‐variables. Species richness was highest at moderate levels of shrub expansion, while it was much reduced in very dense shrubland. The decrease in species number is caused by the decline in grassland species typical of the open dunes, including some rare taxa. Conclusions: The expansion of Hippophaë rhamnoides is a serious threat to the plant species richness of open coastal dunes, and needs to be counteracted by management measures.     

Variation in invertebrate–bryophyte community structure at different spatial scales along altitudinal gradients

Aim This study assessed changes in diversity and assemblage composition in bryophytes and their associated invertebrates along altitudinal gradients in Australia and New Zealand. The importance of altitude in shaping these communities and for the diversity of both invertebrates and bryophytes was examined at different spatial scales, including local, altitudinal, regional and biogeographical. Location Samples were taken from four Australasian mountain ranges between 42° and 43°S: Mt Field and Mt Rufus, Tasmania, Australia, and Otira Valley and Seaward Kaikoura Mountains, South Island, New Zealand. Methods On both Tasmanian mountains, five altitudes were assessed (250, 500, 750, 1000 and 1250 m). At each location (mountain/altitude combination) two sites were chosen and six samples were taken. Six altitudes were assessed on New Zealand mountains (Otira: 250, 500, 750, 1000, 1250 and 1500 m; Kaikoura: 1130, 1225, 1325, 1425, 1525 and 2000 m). Bryophyte substrate was collected, and all samples were stored in 70% ethanol. Invertebrates were extracted from bryophytes using kerosene‐phase separation and all invertebrates were identified to family. At each location in Tasmania, all bryophyte species within six 25‐cm² grids per site were collected and identified to species. Bryophytes from New Zealand were identified to species from the invertebrate sample substrate because of sampling constraints. Results Altitude did have a significant effect on diversity, however, no general trend was found along the altitudinal gradient on the four mountains. There were distinct differences in diversity between biogeographical regions, mountains, altitudes and sites. In Tasmania, Mt Field had the highest diversity in invertebrates and bryophytes at 750 m. In contrast, Mt Rufus had consistent low invertebrate and bryophyte diversity along the entire altitudinal gradient. There were also distinctive differences between locations in the composition of invertebrate and bryophyte communities in Tasmania. Along the two altitudinal gradients in New Zealand, Otira had highest diversity for both invertebrates and bryophytes at low altitudes, whereas Kaikoura had highest invertebrate and lowest bryophyte diversity at the highest altitude. Main conclusions There was an effect of altitude, however, there were no consistent changes in diversity or composition on the four different mountains. There was considerable local and regional variation, and, despite a strong sampling design, no underlying altitudinal trends were detectable. This study demonstrates the importance of examining a range of spatial scales if patterns in community structure along altitudinal gradients are to be studied. The implications of this study are discussed with reference to survey design, taxonomic resolution, climate change and conservation of habitat.     

Diversity of forest plant species at the community and landscape scales in Switzerland

Abstract

Conservation strategies increasingly refer to indicators derived from large biological data. However, such data are often unique with respect to scale and species groups considered. To compare richness patterns emerging from different inventories, we analysed forest species richness at both the landscape and the community scales in Switzerland. Numbers of forest species were displayed using nationwide distributional species data and referring to three different definitions of forest species. The best regression models on a level of four predictor variables ranged between adj. R ² = 0.50 and 0.66 and revealed environmental heterogeneity/energy, substrate (rocky outcrops) and precipitation as best explanatory variables of forest species richness at the landscape scale. A systematic sample of community data (n = 729; 30 m², 200 m², 500 m²) was examined with respect to nationwide community diversity and plot species richness. More than 50% of all plots were assigned to beech forests (Eu-Fagion, Cephalanthero-Fagion, Luzulo-Fagion and Abieti-Fagion), 14% to Norway spruce forests (Vaccinio-Piceion) and 13% to silver fir forests (Piceo-Abietion). Explanatory variables were derived from averaged indicator values per plot, and from biophysical and disturbance factors. The best models for plot species richness using four predictor variables ranged between adj. R ² = 0.31 and 0.34. Light (averaged L-indicator, tree canopy) and substrate (averaged R-indicator and pH) had the highest explanatory power at all community scales. By contrast, the influence of disturbance variables was very small, as only a small portion of plots were affected by this factor. The effects of disturbances caused by extreme events or by management would reduce the tree canopies and lead to an increase in plant species richness at the community scale. Nevertheless, such community scale processes will not change the species richness at the landscape scale. Instead, the variety of different results derived from different biological data confirms the diversity of aspects to consider. Therefore, conservation strategies should refer to value systems.     

Germination in Baltic coastal wetland meadows: similarities and differences between vegetation and seed bank

The abundance and variety of seedlings in Baltic coastal grasslands was studied in cattle grazed and ungrazed areas in seashore and delta on the western coast of Finland. The vegetation, seed bank and environmental conditions of the same sites were also studied. Altogether 4609 seedlings were observed in 79 field plots (20 cm× 20 cm) making an average of 1458.54 seedlings/m². The majority of the seedlings were dicots (67.5%) and perennials (96.8%). Altogether 44 species and an average of 4.14 species per plot were identified as seedlings. The most abundant taxon, Carex formed 23.2% of all the seedlings observed in the field. In the delta the seedling density was two times that found at the seashore. Also, the numbers of species of seedlings per plot were significantly higher in the delta than in the seashore, but in total, more species germinated from the seashore than from the delta even though the number of plots was higher in the delta (n = 51) than in the seashore (n = 28). The numbers of species and seedlings per plot were smaller in grazed sites than in ungrazed ones similar to the pattern earlier reported concerning the mature vegetation and seed bank. The mean numbers of seedlings were the highest at lower geolittoral plots (about 20 cm elevation), where the seed bank was also densest. About 60.0% of the mature species in the plots were also detected as seedlings and 91.4% of the seedling species were found in the mature vegetation of the plots. The number of seedlings was positively correlated with the cover of mature vegetation giving support to the nurse effect theory. Seedling numbers were negatively correlated with the abundance of litter indicating that litter restricts germination because it is a mechanical barrier but also decreases the amount of light. The number of seedlings and species, both in total and per unit area, were greater in the seed bank than in the field seedling population. The seedling population was more similar to the mature vegetation than to the seed bank. This revised version was published online in August 2006 with corrections to the Cover Date.     

Grazing without grasses: Effects of introduced livestock on plant community composition in an arid environment in northern Patagonia

Question: How does grazing intensity affect plant density, cover and species richness in an Patagonian arid ecosystem? Location: Monte steppe ecoregion, SW Argentina. Methods: I analysed the effect of grazing on plant density, cover and species richness using a stocking rate gradient within the same habitat. Six paddocks were used with stocking rates ranging between 0.002 – 0.038 livestock/ha. Plant density, species richness, plant cover and percentage of grazed branches were determined by sampling plots within each paddock. The percentage of grazed branches was used as an independent measurement of grazing intensity. Results: Higher stocking rates were related to lower plant density, species richness and plant cover. The paddock with the lowest grazing intensity had 86% more plants per unit area, 63% more plant cover and 48% higher species richness. The percentage of grazed branches and the quantity of dung increased with stocking rate. Conclusions: Introduced livestock seriously affect native vegetation in the Patagonian Monte. The damage observed in this xerophytic plant community suggests that plant adaptations to aridity do not provide an advantage to tolerate or avoid grazing by vertebrate herbivores in this region. Plant degradation in this arid environment is comparable to the degradation found in more humid ecosystems.     

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.     

两种不同生境中国沙棘种子内生细菌的多样性

植物内生菌广泛分布于植物的各种组织及器官中,对植物的生长表现出各种作用,而植物种子中的内生菌对植物的作用也越来越受到人们的关注。以榆中县和秦安县两种不同生境的中国沙棘种子为材料,分析中国沙棘种子内生细菌多样性,以期探究生境对种子内生菌多样性的影响,并为进一步研究种子内生菌与沙棘的相互作用提供依据。研究利用纯培养方法和高通量测序方法分别进行中国沙棘种子内生细菌多样性分析。对纯培养分离得到的内生细菌,利用16S rRNA基因序列分析法结合形态学特征进行内生细菌的鉴定;对高通量测序得到的数据进行基于OTUs(Operational Taxonomic units,可操作分类单元)的物种注释分析。通过纯培养方法从榆中县中国沙棘种子中分离得到4株内生细菌,分属于芽孢杆菌属(Bacillus)、葡萄球菌属(Staphylococcus)和假单胞菌属(Pseudomonas);秦安县中国沙棘种子中分离得到5株内生细菌,分属于芽孢杆菌属(Bacillus)、葡萄球菌属(Staphylococcus)和不动杆菌属(Acinetobacter)。采用高通量测序方法检测到榆中县中国沙棘种子内生细菌分属于7个...     

空间尺度对地形异质性-物种多样性关系的影响:粒度和幅度

空间尺度是影响我们理解生态学格局和过程的关键因素.目前已有多种关于物种多样性分布格局形成机制的假说且研究者未达成共识,原因之一是空间尺度对物种多样性分布格局的环境影响因子的解释力和相对重要性有重要影响.地形异质性是物种多样性分布格局的重要影响因素.本文综述了在地形异质性-物种多样性关系的研究中,不同空间粒度和幅度对研究...     

Effects of hedgerows on bats and bush crickets at different spatial scales

Biodiversity is threatened by the loss and fragmentation of habitats. The role of hedgerows in maintaining biodiversity is well established, but few studies have addressed the importance for biodiversity of the intrinsic characteristics of hedgerows and the quality of hedgerow networks along a spatial scale. We examined three quality indices providing information at different territorial levels: density in the landscape, structural diversity and wood production. We performed an acoustic survey in a grassland to estimate the species abundance and community composition of bats (9 taxa) and bush crickets (11 species). Using an approach based on species and traits, we assessed how hedgerow quality influenced the activity of these taxa at different spatial scales (from 50 to 1000 m) and focused on three types of traits: bush cricket mobility ability, bat foraging strategy and habitat specialization. In general, our results showed the importance of hedgerow quality for bats and bush crickets, but the strength of the association between taxa and hedgerows varied substantially among the species and the spatial scales. Although it depends on the taxa, the production, density and structural diversity of hedgerows each had an overall positive effect. Our results suggested that these effects were generally more important at large scales. The scale effect of the production index is the best predictor of activity for bat and bush cricket taxa and traits. Our results showed the importance of hedgerow quality for the ecology of bat and bush cricket communities and could be used to improve conservation management.     

Comparative biodiversity of crustaceans and aquatic insects from various water body types in coastal Mediterranean wetlands

Coastal wetlands are characterized by a high biodiversity. At the same time, biodiversity is one of the main criteria used to establish protection policy priorities, or to propose management actions. In this study, crustacean and aquatic insect species richness in the Empordà wetlands was investigated. These two groups contribute in an important way to the total biodiversity, and still they are seldom taken into account in the management of natural areas. Representative samples (38 points) of all aquatic water body types in the Empordà wetlands were taken monthly (dip net with 250 μm mesh). Sampling was carried out between 1996 and 2000, but until present, only qualitative data have been extracted. A rich fauna of 125 crustacean taxa and 295 aquatic insect taxa were found. Some environments were characterized by low richness and high singularity (isolated artesian freshwater springs), some by high richness and high singularity (estuarine waters, brackish and meso-eutrophic freshwater wetlands), and others by low richness and low singularity (hypertrophic freshwater wetlands and hyperhaline wetlands). Factors determining singularity and richness are discussed. Comparison with crustacean richness of other western Mediterranean wetlands showed a similar high species richness in our study sites, probably due to high spatial heterogeneity of these areas.     

Benthic macrofauna in Tasmanian estuaries: scales of distribution and relationships with environmental variables

Information on the distribution of species richness, faunal density, biomass and estimated productivity of benthic invertebrates in Tasmanian estuaries was quantified at a variety of spatial and temporal scales to assess general hypothesis relating community metrics to such environmental variables as salinity, seagrass biomass and sediment particle size. An associated aim was to assess appropriate scales of investigation for soft-sediment biota distributed in estuaries, including whether patterns identified at individual sites, estuaries, tidal levels or times are likely to have more general relevance. Faunal biomass and productivity varied principally at between-estuary (10 to 1000 km) and replicate-sample (1 m) scales, indicating that these two community metrics were largely responding to estuary-wide effects, such as nutrient loading, and to microhabitat features, rather than to locality characteristics at intermediate scales such as salinity, anoxia or sediment particle size. By contrast, faunal density showed greater response to tidal height (1 to 100 m) and to factors distributed at the locality scale within estuary (10 km) than to factors between estuary. Both faunal density and species richness in estuaries declined over three- and fivefold ranges down the shore from high water mark to the shallow sublittoral, while estimated productivity and biomass showed highest overall levels at low water mark. The greatest component of variance in species richness was associated with tidal height, with variance then distributed approximately evenly between other spatial scales examined. At the low-tide and shallow subtidal levels, species richness, faunal biomass and estimated productivity were all highly correlated with salinity and biomass of macrophytes, whereas faunal density was highly correlated with biomass of macrophytes only. Relationships between environmental and biological variables examined were poorly defined at high tidal levels. Seasonal plus interannual variance was much lower than spatial variance—a clear indication that sampling effort in studies would generally be better directed across a range of localities than for a single locality to be repeatedly investigated over time.     

砒砂岩区不同立地类型人工沙棘林下草本物种多样性环境解释

砒砂岩区是黄土高原区水土流失最为严重的地区之一。砒砂岩区沙棘人工林有良好的水土保持功能,沙棘林下草本植物是该人工林生物多样性的重要组成部分。目的是探明砒砂岩区人工沙棘林下草本物种多样性与环境的关系,依据生境条件的差异,沙棘林被划分为7个类型。野外调查记录了各类型所处的地形条件,采用样方法调查了每个样地草本的生物多样性,并取土样测定其养分和水分状况。采用Patrick指数、Pielou指数、Shannon指数、Simpson指数对砒砂岩区不同立地类型人工沙棘林下草本植物进行了物种多样性分析,并通过CCA、PCCA方法研究了物种分布与环境的关系。结果表明:(1)砒砂岩区不同立地类型人工沙棘林下草本层优势种不同,阴坡混交林物种数最多为23种,阳坡混交林Pielou指数最高为0.89,阴坡纯林和混交林Shannon指数均为最高,Simpson指数没有明显差异。(2)影响砒砂岩区人工沙棘林下草本物种的分布的环境因子顺序为地形因子土壤因子灌木层因子,其中坡向、坡度、土壤含水量、土层厚度、郁闭度、混交结构所占解释量较大。本研究可为当地沙棘人工林生物多样性保护和植被建设提供基础数据。     

The relative importance of disturbance and environmental stress at local and regional scales in French coastal sand dunes

Questions: 1. Is there a primary role of disturbance at local scale and of environmental stress at regional scale? 2. Does disturbance increase or decrease environmental stress at local scale? Location: The Atlantic coastal dune system of the Aquitaine Region (France). Methods: Species biomass and 16 environmental variables were sampled in 128 quadrats along a local beach‐inland gradient and a regional North‐South gradient. Environmental data were analysed with ANOVAs and vegetation‐environment relationships with Canonical Correspondence Analysis. Results: At the local scale community composition was primarily driven by disturbance due to sand burial, whereas water and nutrient stress better explained regional differences. However, random biogeographical events are very likely to also affect community composition at the largest scale. The main interaction between environmental stress and disturbance was the mitigation of nutrient stress induced by disturbance at a local scale. This was due to a positive direct effect of sand burial and a positive indirect effect of wind (decrease in VPD by ocean spray). Although wind had also a significant effect on soil conductivity and pH, there was no evidence that these factors had any role in community composition. Conclusions: Our results support the hypothesis that disturbance had a primary role at local scale and environmental stress at regional scale but further research is needed to separate the effect of stress from that of dispersal at regional scale. We also demonstrated that environmental stress in primary succession may not always decline with decreasing disturbance.