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.     

Slow response of plant species richness to habitat loss and fragmentation

We examined the response of vascular plant species richness to long-term habitat loss and fragmentation of Estonian calcareous grasslands (alvars). The current number of habitat specialist species in 35 alvars was not explained by their current areas and connectivities but it was explained by their areas and connectivities 70 years ago ( R ² = 0.27). We estimated the magnitude of extinction debt in local communities by assuming an equilibrium species richness in 14 alvars that had lost only a small amount of area and by applying this model to the remaining alvars, in which the average area has declined from 3.64 km² in the 1930s to 0.21 km² at present. The extinction debt estimated for individual alvars was around 40% of their current species number. Our conclusions are applicable to temperate grasslands in general, which have lost much area because of agricultural intensification and cessation of traditional management.     

Estimating species richness at large spatial scales using data from small discrete plots

Estimating species richness in large biomes is a central challenge in ecology and conservation biology. However, accurate census data is often available only from small discrete plots distributed within the biome. Using tree species richness data collected from 48 plots (0.25 ha each) widely distributed through 60 000 km² in the rainforests of the Western Ghats of southern India, we test the application of a proposed method for estimating species richness at large scales from measured species commonalities between pairs of censused plots. We show that the method allows extrapolation of species richness from a scale of 0.25 ha plots to that of the entire biome, or 10⁵ km².     

Integrating multiple landscape-scale drivers in the lichen epiphyte response: climatic setting, pollution regime and woodland spatial-temporal structure

Aim To quantify the role of multiple biodiversity drivers – pollution, woodland structure and climate – controlling lichen epiphyte composition and diversity. Location  Scotland, north‐west Europe. Methods Four compatible datasets were assembled: site‐scale species distribution data (response) and base‐line modelled data on climate, pollution loads and extent of old‐growth woodland (explanatory variables). First, partial‐canonical correspondence analysis was used: (1) to compare the importance of environmental variables to pure spatial effects and (2) to partition the importance of environmental variables in explaining species composition. Secondly, patterns of species richness were investigated using multiple least‐squares regression. Results Old‐growth woodland was the most important control of species richness. Pollution was the most important explanatory variable for species composition. The impact of pollution on composition (and to a lesser extent on richness) is explained: (1) By recovery of lichens with declining SO₂ pollution, although with epiphyte composition shifted by the recent effects of N‐pollution and (2) By the limited spatial extent of severe pollution, and generally low‐to‐moderate pollution loads across our study area, combined with the positive effect of old‐growth woodland extent in controlling species richness. The effect of climate and old‐growth woodland on species composition covaried, supporting an interaction between habitat quality and climatic setting, which may be important in understanding the epiphyte response to climate change. Conclusions Advances in conservation planning will likely require an integrated approach to understanding simultaneous effects of multiple drivers, providing opportunities for integrated management strategies. Our study provides a preliminary example of this approach by combining three key biodiversity drivers into a single framework for lichen epiphytes. Thus, reducing pollution loads may make old‐growth woodland that currently exists in a polluted landscape available for colonization, thereby extending the available habitat for epiphytes, and facilitating an effective species response to climate change.     

Determinants of local spider (Araneidae and Thomisidae) species richness on a regional scale: climate and altitude vs. habitat structure

Abstract.  1. The habitat heterogeneity hypothesis states that the more complex the habitat, the higher the species richness. The present study analyzes the effect of local factors on regional spider ( Araneidae and Thomisidae ) richness. The main objective is to disentangle the relative importance of habitat structure and other environmental variables.
2. Fifteen territorial units of 1 km² were sampled to obtain reliable and comparable inventories of the two spider families. Richness values were modelled using general regression models and a set of climate, topographic and vegetation structure variables. Pure and joint effects were computed via variation partitioning.
3. The results highlight the great importance of vegetation complexity, especially of grass and sub-shrub cover, in determining spider species richness.
4. The maximum temperature is the only climate variable significantly related to species richness, although its effect is combined with that of spatial and vegetation structure variables.
5. These results support the habitat heterogeneity hypothesis, and highlight the importance of taking vegetation complexity into account when managing habitats and where spider conservation is desired.     

Changing climate and historic‐woodland structure interact to control species diversity of the ‘Lobarion’ epiphyte community in Scotland

Question: How will changing climate and habitat structure interact to control the species diversity of lichen epiphytes? Location: Scotland. Method: Species richness (=diversity) of the epiphyte lichen community known as Lobarion (named after Lobaria pulmonaria) was quantified for 94 Populus tremula stands across Scotland, and compared in a predictive model to seven climate variables and eight measures of woodland structure. An optimum model was selected and used to project Lobarion diversity over the geographic range of the study area, based on IPCC climate change scenarios and hypothetical shifts in woodland structure. Results: Species diversity of the Lobarion community was best explained by three climate variables: (1) average annual temperature; (2) autumn and winter precipitation; in combination with (3) historic‐woodland extent. Projections indicate a positive effect of predicted climate change on Lobarion diversity, consistent with the physiological traits of cyanobac‐terial lichens comprising the Lobarion. However, the general response to climate is modified significantly by the effect on diversity of historic‐woodland extent. Conclusions: Historic‐woodland extent may exert an important control over local climate, as well as impacting upon the metapopulation dynamics of species in the Lobarion. In particular, a temporal delay in the response of Lobarion species to changed woodland structure is critical to our understanding of future climate change effects. Future Lobarion diversity (e.g. in the 2050s) may depend upon the interaction of contemporary climate (e.g. 2050s climate) and historic habitat structure (e.g. 1950s woodland extent). This is supported by previous observations for an extinction debt amongst lichen epiphytes, but suggests an extension of simple climate‐response models is necessary, before their wider application to lichen epiphyte diversity.     

Brazilian Tapir Density in the Pantanal: A Comparison of Systematic Camera-Trapping and Line-Transect Surveys

The density of Brazilian tapirs ( Tapirus terrestris) was studied in the northeastern part of the Pantanal wetlands of Brazil using two simultaneous and independent methods: (1) systematic camera trapping combined with capture–recapture analysis, with camera traps spaced 1 km apart and distributed over 54 km²; and (2) line-transect sampling using an array of 12 linear transects, from 3.8 to 7.2 km long, covering the principal open and forest habitat types across the entire 1063 km² SESC Pantanal Reserve. The two methods yielded conservative density estimates of 0.58 ± 0.11 tapirs/km² (camera trapping) and 0.55 (95% CI 0.30–1.01) tapirs/km² (line transects). The study suggests that certain Pantanal habitats and sites can sustain relatively high population densities of tapirs when these animals are protected from hunting. Further testing of the camera-trapping methodology as applied to tapirs is required, particularly focusing on extending the survey period. As it represents a relatively rapid method for estimating population density, in comparison to line-transect surveys, and as it generates information simultaneously on multiple species that are conservation priorities, we recommend that camera-trapping surveys be applied more widely across a variety of Pantanal habitats and land-use categories in order to confirm the value of the vast 140,000 km² wilderness region for this vulnerable species.     

Ranges of males of the Great Hornbill Buceros bicornis, Brown Hornbill Ptilolaemus tickelli and Wreathed Hornbill Rhyticeros undulatus in Khao Yai National Park, Thailand

Ranges of individual males of three hornbill species were determined by radio telemetry during the breeding and non-breeding seasons in Khao Yai National Park, Thailand. Two Great Hornbills Buceros bicornis and two Brown Hornbills Ptilolaemus tickelli were studied in both 1988 and 1989, and two Wreathed Hornbills Rhyticeros undulatus were studied in 1989, 1990 and 1991.
In the breeding season, the home range of the Great Hornbill was 3.7 km², similar to that of the Brown Hornbill (4.3 km²), while the Wreathed Hornbill occupied the largest home range (10.0 km²). In the non-breeding season, the range size of the Wreathed Hornbill (28.0 km²) was greater than that of the Great Hornbill (14.7 km²). Differences in range sizes of different species may be related to differences in diet and breeding strategy. Ranges overlapped within and between the species, and this has implications for the estimation of the minimum area required for the conservation of hornbills within the Khao Yai National Park.     

Scale dependence of effective specialization: its analysis and implications for estimates of global insect species richness

Estimates of global insect species richness are sometimes based on effective specialization, a calculation used to estimate the number of insect species that is restricted to a particular tree species. Yet it is not clear how effective specialization is influenced by spatial scale or characteristics of the insect community itself (e.g. species richness). We investigated scale dependence and community predictors of effective specialization using 15,907 beetles (583 species) collected by insecticide fogging from the crowns of 96 trees (including 32 Quercus trees) located in Ohio and Indiana. Trees were distributed across 24 forest stands (∼1 ha) nested within six sites (∼10–100 km²) and two ecoregions (> 1000 km²). Using paired-sample randomization tests, we found that effective specialization ( f _k ) exhibited negative scale-dependence in early (May–June 2000) and late (August–September 2000) sampling periods. Our average effective specialization ( F ) values — those that are comparable to Erwin's (1982) estimates — ranged from 19% to 97%, and increased as spatial scale decreased. We also found that beetle species richness and the number of shared beetle species across host trees were significant and consistent negative predictors of F . This shows that increases in spatial scale, species richness, and the number of trees (and/or tree species) all coincide with decreases in effective specialization. Collectively, our results indicate that estimates of global insect species richness based on effective specialization at a single spatial scale are overestimating the magnitude of global insect species richness. We propose that scale dependence should be promoted to a central concept in the research program on global estimates of species richness.     

Scale dependence of the correlation between human population presence and vertebrate and plant species richness

Human presence is generally negatively related to species richness locally, but the relationship is positive at coarse scales. An increase in the strength of the latter correlation with increasing study resolution has been documented within studies, but it is not known whether such a scale dependence is present across different studies. We test this with data on the spatial co-occurrence of human beings and the species richness of plants and vertebrates from a continuum of scales. The correlation coefficient between human presence and species richness is positively related to study grain and extent. The correlation turns from positive to negative below a study grain of c. 1 km and below a study extent of c. 10 000 km². The broad-scale positive correlation between human presence and species richness suggests that people have preferentially settled and generally flourished in areas of high biodiversity and/or have contributed to it with species introductions and habitat diversification. The scale dependency of the correlation between people and biodiversity's presence emphasizes the importance of the preservation of green areas in densely populated regions.     

Effects of topographic variability on the scaling of plant species richness in gradient dominated landscapes

It is commonly assumed that variation in abiotic site conditions influences the number of niches, which in turn affects the potential species richness in an area. Based on theoretical considerations, abiotic variation is often used as an estimator of species richness at broad scales, while at finer landscape scales the diversity of habitat types is used. However, habitat estimators assume the landscape to be composed of discrete, homogeneous patches with sharp boundaries, and such a concept is hard to apply in gradient-dominated landscapes. The aim of this study was therefore to investigate the influence of topographic variability (TV) on species richness at the landscape level (gamma (γ) diversity) and on its components (alpha (α) and beta (β) diversity) at microsite and habitat group levels. Using floristic data from 12 "landscapes" of 1 km² we investigated the influence on diversity components of two simple and one complex measures of TV. While the standard deviation (SD) of altitude explained a high proportion of the variation in γ diversity (linear regression model, R²=0.63), the complex measure, SD of solar radiation explained it even better (R²=0.82). There were strong effects of TV on α and β diversity components at the microsite level, but only marginal increases of the diversity components at the habitat level. Further analyses revealed that the missing increase of the habitat level components was caused by differences between habitat groups and that only grassland diversity components increased significantly with TV. We conclude that TV at a landscape scale has strong effects on niche or microsite diversity and is an appropriate estimator of relative species richness in landscapes that are topographically heterogeneous and gradient dominated.     

Detecting decline in a formerly widespread species: how common is the common blue butterfly Polyommatus icarus?

Grid square distribution maps have been used widely to measure rates of decline and target conservation resources. However, common species that may have many populations per grid square can decline substantially from within squares without being lost from entire squares. In order to quantify this process, fine scale population and habitat data have been collected for the common blue butterfly Polyommatus icarus in a 35 km² area of fragmented landscape in north Wales. Present day habitat associations, determined from over 2000 transect walks, combined with data on historical and present day habitat distributions reveal that the species has declined by about 74% since 1901. Similar data concerning the species major host plant Lotus corniculatus indicate a decline of 46%. Based on 1 km² grid maps, neither species have been assessed as declining at all. These results suggest that apparently 'common' species may have declined just as much as many of Britain's rare species: using present methods of assessment these declines are undetected.     

Preliminary estimates of the population density of the Sokoke scops owl Otus ireneae Ripley in the East Usambara lowlands, Tanzania

Preliminary population density estimates are presented for a recently discovered population of the Sokoke scops owl Otus ireneae in the lowlands of the East Usambara mountains, Tanzania. Calling birds were mapped at two sites totalling 6·3 km². Approximate densities were 3–4 territories/km² in Kwamgumi Forest Reserve and <1·5 territories/km² in Manga Forest Reserve, a much more heavily logged site. Densities in prime habitat in Arabuko‐Sokoke forest, Kenya, the only other locality for the species, exceed seven territories/km². The total population in the East Usambaras cannot be calculated, but it is probably markedly smaller than in Arabuko‐Sokoke. Suggestions for more precise surveys are made.     

The relationships among three habitat scales and stream benthic invertebrate community structure

1. The relationships between three habitat scales and lotic invertebrate species composition were investigated for the 15 540 km² Yakima River basin in south-central Washington, U.S.A.
2. The three spatial scales were sample (the sampled riffle), reach (a length of ten–twenty stream widths) and segment (a length of stream of nearly uniform slope and valley form having no change in stream order).
3. Physical variables were highly correlated between scales and expressed a relationship between altitude, basin form and small-scale physical structure.
4. Multiple discriminant function analyses indicated that segment- and reach-scale variables discriminated among species-defined groups better than sample-scale variables.
5. Species composition varied along a complex altitudinal gradient of changing basin form and resultant land use.
6. There was no clear relationship between species richness and altitude on a site basis. However, when viewed at the basin scale, maximum richness was observed at the transition between montane and valley sites.     

Wintering site fidelity and movement patterns of Western Sandpipers Calidris mauri in the San Francisco Bay estuary

Western Sandpipers Calidris mauri are the most numerous shorebird species in the San Francisco Bay estuary during winter. A sample of 106 Western Sandpipers was captured in mist nets and radio-marked with 1-g transmitters to examine their wintering site fidelity and movements. Differences in distances moved, home range extent and core area size were examined by age, sex, season, site, time of day and tide. All birds remained in the south San Francisco Bay region during winter and exhibited strong site fidelity, with a mean home range of 22.0 km² or only 8% of the study area. First-year birds had larger home ranges (26.6 ± 3.6 km²) than adults (17.2 ± 2.5 km²) in winter, but home range sizes of males and females were not significantly different in any period. Home range sizes were similar between seasons, but core areas were smaller in spring (6.3 ± 1.2 km²) than in early (9.6 ± 4.0 km²) or late (11.6 ± 1.6 km²) winter. Movements and home range size were similar for radio-marked birds located during day and night. The high degree of regional and local site fidelity demonstrated that the mixture of natural mud fiats and artificial salt ponds in southern San Francisco Bay provided sufficient resources for large wintering populations of Western Sandpipers.     

The relationships among three habitat scales and stream benthic invertebrate community structure

1. The relationships between three habitat scales and lotic invertebrate species composition were investigated for the 15 540 km² Yakima River basin in south-central Washington, U.S.A.
2. The three spatial scales were sample (the sampled riffle), reach (a length of ten–twenty stream widths) and segment (a length of stream of nearly uniform slope and valley form having no change in stream order).
3. Physical variables were highly correlated between scales and expressed a relationship between altitude, basin form and small-scale physical structure.
4. Multiple discriminant function analyses indicated that segment- and reach-scale variables discriminated among species-defined groups better than sample-scale variables.
5. Species composition varied along a complex altitudinal gradient of changing basin form and resultant land use.
6. There was no clear relationship between species richness and altitude on a site basis. However, when viewed at the basin scale, maximum richness was observed at the transition between montane and valley sites.     

Home range and habitat use of wolverines Gulo gulo in Yukon, Canada

Home ranges and habitat use are described for three adult female, one adult male, and one subadult male wolverines in the Kluane Game Sanctuary, Yukon. When long distance excursions are not included, home ranges of wolverines in the Kluane Game Sanctuary were between 76 and 269 km² for females and 209 and 269 km² for males. Habitat use of females was similar to habitat availability. Males used subalpine coniferous habitats more frequently than other habitat types during winter. Although individual variation in the use of forest cover types, aspects, slopes and elevations was apparent, seasonal use did not differ from availability for each sex. Within the 1590 km² study area, three adult males and six adult females were present, corresponding to a density of one resident wolverine 177 km^-2.     

Spatial organization of leopards Panthera pardus in Taï National Park, Ivory Coast: is rainforest habitat a 'tropical haven'?

Between June 1992 and July 1994, two female leopards and one male were radio-tracked. Regular locations of the leopards, the use of a phototrap, and spoor data, provided the first detailed ecological data about this elusive felid in tropical rainforest habitat. The home range of the male was 86 km², those of the two females were 29 km² and 22 km², respectively. One female's home range was fully included within that of the male. Home ranges of neighbouring residents were not exclusive. Population density is estimated at one leopard per 9-14 km². Intraspecific interactions were rare and predominantly involved mating. The large size of the home ranges and a relatively high population density imply large overlap between adjacent resident leopards' ranges. Differences in the leopard's land tenure system between the rainforest and the savanna are discussed. Doubt is cast on the validity of the often-quoted estimate of one leopard per 1 km² in tropical rainforest habitat.     

Species richness and endemism of plant and bird communities along two gradients of elevation, humidity and land use in the Bolivian Andes

Abstract.  We studied the patterns of species richness and range–size rarity (as a measure of endemism) of two plant groups (Pteridophyta, Bromeliaceae) and birds along two gradients of elevation, humidity and human land use in a forested Andean valley. Both transects covered the transition from an arid valley bottom through a cloud forest zone to relictual high-elevation Polylepis forest, but transects differed in overall precipitation. Plants were surveyed in 88 plots of 400 m² each, while birds were detected primarily through visual observations and tape recordings over areas of 0.3–1.5 km². Global range sizes of all species were mapped on 1°-grids and range-size rarity was calculated as the mean inverse range size of all species recorded in elevational steps of 200 m. Patterns of species richness and range–size rarity were mainly unrelated between and within study groups. Monotonic increases and decreases and hump-shaped patterns were observed for species richness as well as range–size rarity. Several of these patterns can be interpreted in the light of the ecological requirements of each taxonomic group, e.g. dependence of fern species richness on humidity or of bird richness on habitat complexity. Species richness of ferns and birds peaked at higher elevations along the less rainy transect, possibly as a result of higher levels of solar radiation and ecosystem productivity. Patterns of species richness and endemism of the study groups are causally unrelated and cannot be used to predict those of other groups at the spatial scale of this study. Human impact was highest in areas of mostly low to intermediate species richness, but was often high in zones of high endemism.     

Micro-environmental characteristics of filamentous algal communities in flowing freshwaters

SUMMARY. 1. Filamentous algae in flowing freshwaters can represent a spatially and temporally distinct sub-habitat for epiphytic diatom communities. This sub-habitat is described in a low discharge, spring-fed stream with extensive filamentous green algal mats, and in a tuft of Cladophora glomerata (L.) Kützing from a large river.
2. Oxygen micro-electrodes, a thermistor current velocity probe, a standard pH probe and water chemistry were employed to assess spatial heterogeneity. Temporal patterns of epiphyte colonization were evaluated on filamentous artificial substrates.
3. There were steep spatial gradients in the low discharge stream. At mid-day, O₂ ranged from 0–1.5 times air-saturated O₂ concentrations, pH varied from 7.25–8.0, and current velocity spanned 0–0.5 m s⁻¹. Areas near the surface of algal mats had high O₂, pH and current velocity. These patterns were correlated with epiphyte community structure.
4. In the interior of C. glomerata tufts O₂ concentration was raised and current velocity depressed compared to the surrounding water, even when external current velocity was as high as 0.4 m s⁻¹.
5. After thirty-five days of colonization of artificial substrate in the low discharge environment, epiphyte communities were similar to those on filamentous atgae. Epiphyte diversity on artificial substrates subsequently decreased compared to natural substrates as did the similarity between the types on substrates, suggesting that microscale renewal of epiphyte habitat (growth of filamentous algae) maintains high epiphytic diversity.