Resource tracking by avian frugivores in mountain habitats of northern Spain

Systems comprising avian frugivores and fleshy‐fruited plants are commonly used as models to study how animals respond to temporal and spatial variations in food abundance and distribution. Results have been far from conclusive, partly because of methodological constraints. In this study we investigated the fulfilment of a necessary condition for food tracking: correlation over time and space between the abundance of food and the abundance of food trackers. We used a paired‐sample design in which fruit sites were compared with fruitless sites on two different spatial scales (landscape and habitat patch) in eight different river basins in the Cantabrian Mountains (NW Spain). Fruit and bird abundances were recorded monthly (October to February) at all sites during three consecutive fruiting seasons. Fruiting seasons (“years”) could be ranked by fruit abundance as low (2003), intermediate (2002) and high (2004). Since fruit sites were located at considerably higher elevations than fruitless sites, the possible effects of altitude, together with those of other uninvestigated factors, were taken into account by using non‐frugivorous birds. Our results apparently suggest a temporal and spatial association between frugivores (avian seed dispersers) and fruit availability at the landscape scale but not at the smaller (habitat patch) scale. This, together with the absence of a similar pattern in non‐frugivores, suggests that fruit availability is an important ecological factor affecting the autumn and winter abundance patterns of frugivorous birds and that this factor operates at the landscape scale but not at the habitat patch scale. Moreover, the negative elevational gradient in bird abundance that is typical of temperate mountain ecosystems was clearly reversed for frugivores in the intermediate and high fruiting years (suggesting that the negative effect of higher altitude may have been counterbalanced by the greater availability of food). As in other European upland areas, traditional farming practices in the Cantabrian Mountains are declining and, as a result, the proportion of fruitless habitats is increasing. Our study suggests that in these upland ecosystems frugivorous bird abundances in autumn and winter are associated to fruit supply, thus this habitat change tendency will probably have, in the long‐term, a negative effect on bird populations.     

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.     

Sexual segregation by Masai giraffes at two spatial scales

In this paper alternative explanations for observed patterns of sexual segregation by giraffes are examined at two spatial scales: within-habitats and within-landscape. Habitats are defined as recognizable plant associations and the landscape as the collection of all available habitat types. The study was conducted in Mikumi National Park, Tanzania. At the within-landscape level, all sex and age classes of giraffes exhibited high degrees of preference for riverine habitats. Sex differences in habitat selection were mostly due to females with young, who tended to select open floodplain habitats in which their vigilance time was lowest. Males, and females without young, preferred more heavily-wooded habitat. Habitat preferences were not related to observed habitat-specific forage intake rates for either males or females. Within habitats, male and female giraffes selected different feeding heights, males feeding higher in the canopy than females. Females showed a strong tendency to generalize with respect to feeding height. It is suggested that a sexual dimorphism–body size hypothesis provides a parsimonious explanation for the observed feeding height selection patterns, whereas a reproductive strategy hypothesis can explain sex-differences in habitat selection patterns within the landscape.     

Grains of connectivity: analysis at multiple spatial scales in landscape genetics

Landscape genetic analyses are typically conducted at one spatial scale. Considering multiple scales may be essential for identifying landscape features influencing gene flow. We examined landscape connectivity for woodland caribou (Rangifer tarandus caribou) at multiple spatial scales using a new approach based on landscape graphs that creates a Voronoi tessellation of the landscape. To illustrate the potential of the method, we generated five resistance surfaces to explain how landscape pattern may influence gene flow across the range of this population. We tested each resistance surface using a raster at the spatial grain of available landscape data (200 m grid squares). We then used our method to produce up to 127 additional grains for each resistance surface. We applied a causal modelling framework with partial Mantel tests, where evidence of landscape resistance is tested against an alternative hypothesis of isolation-by-distance, and found statistically significant support for landscape resistance to gene flow in 89 of the 507 spatial grains examined. We found evidence that major roads as well as the cumulative effects of natural and anthropogenic disturbance may be contributing to the genetic structure. Using only the original grid surface yielded no evidence for landscape resistance to gene flow. Our results show that using multiple spatial grains can reveal landscape influences on genetic structure that may be overlooked with a single grain, and suggest that coarsening the grain of landcover data may be appropriate for highly mobile species. We discuss how grains of connectivity and related analyses have potential landscape genetic applications in a broad range of systems.     

Effects of habitat heterogeneity at multiple spatial scales on fish community assembly

Habitat variability at multiple spatial scales may affect community structure within a given habitat patch, even within seemingly homogenous landscapes. In this context, we tested the importance of habitat variables at two spatial scales (patch and landscape) in driving fish community assembly using experimental artificial reefs constructed across a gradient of seagrass cover in a coastal bay of The Bahamas. We found that species richness and benthic fish abundance increased over time, but eventually reached an asymptote. The correlation between habitat variables and community structure strengthened over time, suggesting deterministic processes were detectable in community assembly. Abundance of benthic fishes, as well as overall community structure, were predicted by both patch- and landscape-scale variables, with the cover of seagrass at the landscape-scale emerging as the most important explanatory variable. Results of this study indicate that landscape features can drive differences in community assembly even within a general habitat type (i.e., within seagrass beds). A primary implication of this finding is that human activities driving changes in seagrass cover may cause significant shifts in faunal community structure well before complete losses of seagrass habitat.     

景观格局与植物多样性的关系及其空间尺度效应——以浑善达克沙地为例

景观格局是植物多样性的重要影响因素,为了探明沙地景观格局与植物多样性的关系及其空间尺度效应,以浑善达克沙地腹地为研究区,采用样方法进行植物群落学调查,基于GIS和遥感分析技术,分析景观格局指数,计算了10个尺度(100,200,300,400,500,600,700,800,900,1000m)下33个景观格局指数,利用双变量相关性分析法筛选出不同尺度下与植物多样性显著相关的指数;运用逐步回归的方法识别不同尺度下对植物多样性贡献显著的景观格局指数,并进行拟合检验。研究结果表明:景观格局指数对物种多样性影响具有显著的尺度依赖性。在所有空间尺度,景观格局指数主要对Shannon多样性指数和β指数影响较大;在100m尺度下,Shannon指数逐步回归R~2值达到最大(0.138);β指数在900m尺度下R2值达到最大值(0.2)。拟合检验发现,在500m尺度下,均匀度指数、Shannon多样性指数和β多样性指数实测值和预测值拟合均达到显著水平(P0.05),故认为500m尺度下,景观格局对植物多样性影响较大。因此,在不同尺度下管理和保护沙地植物多样性,应综合考虑景观格局指数的尺度效应,根据需要建立和优化有利于植物多样性的景观格局。     

Habitat selection at multiple spatial scales by foraging Glossy Black‐cockatoos

Abstract: Habitat selection among vertebrates entails decision making at a number of spatial scales. An understanding of factors influencing decisions at each of these scales is required for the effective management of wildlife populations. This study investigates the foraging ecology of a population of Glossy Black‐cockatoos in central New South Wales. We took advantage of the characteristic feeding sign produced by Glossy Black‐cockatoos to examine factors influencing habitat selection at multiple spatial scales. Birds preferred to forage at sites where food was abundant and avoided open sites where the predation risk may be greater. Their two food species, Allocasuarina diminuta and Allocasuarina gymnanthera, differed in profitability (kernel intake rate as measured by the ratio of seed weight to total seed and cone weight), as did trees within a species. Both species were utilized extensively, although foraging intensity was greater at sites where the more profitable species was present. In order to maximize their food intake, birds selected individual trees on the basis of cone abundance and profitability. Cones produced in the previous year were preferred.     

Foraging across a variable landscape: behavioral decisions made by woodland caribou at multiple spatial scales

We examined the foraging behavior of woodland caribou (Rangifer tarandus caribou) relative to the spatial and temporal heterogeneity of their environment. We assessed (1) whether caribou altered their behavior over time while making trade-offs between forage abundance and accessibility; and (2) whether foraging decisions were consistent across spatial scales (i.e., as scale increased, similar decision criteria were used at each scale). We discuss whether caribou adjusted their behavior to take advantage of changing forage availability through time and space. At the scale of the feeding site (as revealed by discriminant function analyses), caribou in both forested and alpine (above tree-line) environments selected sites where the biomass of particular lichen species was greatest and snow the least deep. Caribou did not select those species with the highest nutritional value (i.e., digestible protein and energy) in either area. Where snow depth, density, and hardness limited access to terrestrial lichens in the forest, caribou foraged instead at those trees with the greatest amount of arboreal lichen. Selection of lichen species and the influence of snow differed across time, indicating that in this system the abundance or accessibility of forage temporally influenced foraging behavior. A path analysis of forest data and multiple regression analysis of alpine data were used to test the hypothesis that variables important at the scale of the feeding site explained foraging effort at the scale of the patch. For forest patches, our hypothesized model reliably explained foraging effort, but not all variables that were statistically important at the scale of the feeding site were significant predictors at the scale of the patch. For alpine patches, our hypothesized model did not explain a statistically significant portion of the variation in the number of feeding sites within the patch, and none of the individual variables from the feeding site remained statistically significant at the patch scale. The incongruity between those variables important at the scale of the feeding site and those important at the patch showed that spatial scale affects the foraging decisions of woodland caribou. At the scale of the landscape, there was a trade-off between forage abundance and accessibility. Relative to the alpine environment, caribou in the forest foraged at feeding sites and patches with greater amounts of less variably distributed lichen, but deeper less variable snow depths. Considering the behavioral plasticity of woodland caribou, there may be no distinct advantage to foraging in one landscape over the other.     

Patterns of alien plant distribution at multiple spatial scales in a large national park: implications for ecology, management and monitoring

Aim Spatial scale is critical for understanding and managing biological invasions. In providing direction to managing alien plant invasions, much emphasis is placed on collecting spatially explicit data. However, insufficient thought is often given to how the data are to be used, frequently resulting in the incompatibility of the data for different uses. This paper explores the role of spatial scale in interpreting, managing and monitoring alien plant invasions in a large protected area. Location Kruger National Park, South Africa. Methods Using 27,000 spatially‐explicit records of invasive alien plants for the Kruger National Park (> 20,000 km²) we assessed alien plant species richness per cell at nine different scales of resolution. Results When assessing the patterns of alien plants at the various scales of resolution, almost identical results are obtained when working at scales of quarter‐degree grids and quaternary watersheds (the fourth level category in South Africa's river basin classification system). Likewise, insights gained from working at resolutions of 0.1–0.5 km and 1–5 km are similar. At a scale of 0.1 × 0.1 km cells, only 0.4% of the Kruger National Park is invaded, whereas > 90% of the park is invaded when mapped at the quarter‐degree cell resolution. Main conclusions Selecting the appropriate scale of resolution is crucial when evaluating the distribution and abundance of alien plant invasions, understanding ecological processes, and operationalizing management applications and monitoring strategies. Quarter‐degree grids and quaternary watersheds are most useful at a regional or national scale. Grid cells of 1 to 25 km² are generally useful for establishing priorities for and planning management interventions. Fine‐scale data are useful for informing management in areas which are small in extent; they also provide the detail appropriate for assessing patterns and rates of invasion.     

Consequences of forest fragmentation for polyporous fungi at two spatial scales

Greatly reduced area of old-growth forests and the very low amount of dead wood in managed forests in northern Europe have caused a marked decline in the populations of saproxylic species. It is less clear at which spatial and temporal scales these adverse changes are taking place, and more information is needed to reliably predict which species are especially sensitive to loss and fragmentation of habitat. Here we compare species richness, incidence of occurrence in forest fragments, and abundance of polyporous fungal species and species groups between two regions in Finland with contrasting histories of forestry and a marked difference in the amount and spatial configuration of old-growth forests. We also analyse the consequences of increasing loss of connectivity on the presence and abundance of polypores in a study region with a documented short-term history of old-growth fragmentation. Our results show that the species number, incidence of occurrence, and abundance of especially the rare, threatened, and near-threatened species are much lower in the old-growth fragments in Häme in southern Finland in comparison with Kuhmo in eastern Finland, most probably because of the longer history of intensive forestry in Häme. Among the rare species, the species that show the greatest difference between the two regions (at the scale of 500 km) also tended to respond most strongly to the more recent forest fragmentation within the study region in Kuhmo (at the scale of 50 km). Polypores associated with spruce seem to be more strongly affected by forestry than species associated with pine, possibly reflecting the differences in the natural dynamics of spruce-dominated and pine-dominated forests.     

Impact of landscape pattern at multiple spatial scales on water quality: A case study in a typical urbanised watershed in China

Buffer zones along rivers and streams can provide water quality services by filtering nutrients, sediment and other contaminants from the surface. Redundancy analysis was used to determine the influence of the landscape pattern at the entire catchment scale and at multiple buffer zone scales (100 m, 300 m, 500 m, 1000 m and 1500 m) on the water quality in a highly urbanised watershed. Change-point analysis was further applied to estimate the specific locations along a gradient of landscape metric that result in a sudden change in the water quality variable. The landscape characteristics for 100 m buffer zones appeared to have a slightly greater influence on the water quality than the entire catchment. The patch density of urban land and the large patch index of water were recognised as the dominant variables influencing the water quality for a 100 m buffer zone. The result of change-point analysis indicated key interval values of the two landscape metrics within the 100 m buffer zone. When the patch density of urban land was >30–40 n/100 ha and the largest patch index of water was >2.5–3.5%, the watershed water quality appeared to be better protected.     

Primary productivity and species richness: relationships among functional guilds, residency groups and vagility classes at multiple spatial scales

One of the major determinants of species richness is the amount of energy available, often measured as primary productivity. Heterogeneity of environmental variables has also been found to influence species richness. Predicting species distributions across landscapes and identifying areas that have high species richness, or vulnerable groups of species, is useful for land management. Remotely sensed data may help identify such areas, with the Normalized Difference Vegetation Index (NDVI) providing an estimate of primary productivity. We examined the relationship between maximum productivity (NDVI), heterogeneity of productivity, and species richness of birds and butterflies at multiple spatial scales. We also explored relationships between productivity, functional guilds and residency groups of birds, and vagility classes of butterflies. Positive linear relationships between maximum NDVI and number of functional guilds of birds were found at two spatial scales. We also found positive linear relationships between maximum NDVI and species richness of neotropical migrant birds at two scales. Heterogeneity of NDVI, by contrast, was negatively associated with number of functional guilds of birds and species richness of resident birds. Maximum NDVI was associated with species richness of all butterflies and of the most vagile butterflies. No association was found between heterogeneity of NDVI and species richness of butterflies. In the Great Basin, where high greenness and availability of water correspond to areas of high species richness and maximum NDVI, our results suggest that NDVI can provide a reliable basis for stratifying surveys of biodiversity, by highlighting areas of potentially high biodiversity across large areas. Measures of heterogeneity of NDVI appear to be less useful in explaining species richness.     

Beta diversity of stream diatoms at two hierarchical spatial scales: implications for biomonitoring

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A test of spatial memory and movement patterns of bumblebees at multiple spatial and temporal scales

Naive bumblebee foragers appear to use movement rules at smallspatial and temporal scales, but it is not clear whether theserules determine movement patterns as the scales increase. Onestrategy for efficient foraging used by bumblebees is near-farsearch, involving short flights when in good patches of flowersand longer flights when in poor patches. Bumblebees also demonstratethe use of a spatial memory strategy by returning repeatedlyto patches of flowers, and even following the same route betweenflowers, over periods of days. We attempted to determine atwhat spatial scales bumblebees use spatial memory while foragingwithin a patch and after how many flower visits spatial memoryoutweighs near-far search. Bumblebees in the laboratory foragedon a 4 x 4 array of artificial flowers with distances rangingfrom 10 to 80 cm between flowers in two simple spatial patterns.The proportion of visits to flowers containing a sucrose rewardwas monitored for either 100 or 400 flower visits in two separateexperiments, after which the locations of the rewarding andnonrewarding flowers were interchanged, producing a mirror image.A drop in accuracy after the mirror image switch would indicatethat the bees had memorized the location of rewarding flowers.Mirror image tests, and comparisons to a simulation model ofnear-far search based on actual flight distances, indicate thatnaive bumblebees used near-far search on flowers 10 cm apartbut increasingly used spatial memory as experience and spatialseparation increased. Bumblebees thus have multiple tacticsavailable to forage efficiently in different environments.     

Multitrophic influences on egg distribution in a specialized leaf beetle at multiple spatial scales

Egg distribution in herbivorous beetles can be affected by bottom-up (host plant), and by top-down factors (parasitoids and predators), as well as by other habitat parameters. The importance of bottom-up and top-down effects may change with spatial scale.

In this study, we investigated the influence of host plant factors and habitat structure on egg distribution in the leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae), a monophagous herbivore on Salvia pratensis L. (Lamiales: Lamiaceae), on four spatial scales: individual host plant, microhabitat, macrohabitat, and landscape. At the individual host plant scale we studied the correlation between egg clutch incidence and plant size and quality. On all other scales we analyzed the relationship between the egg clutch incidence of C. canaliculata and host plant percentage cover, host plant density, and the surrounding vegetation structure. Vegetation structure was examined as herbivores might escape egg parasitism by depositing their eggs on sites with vegetation factors unfavorable for host searching parasitoids.

The probability that egg clutches of C. canaliculata were present increased with an increasing size, percentage cover, and density of the host plant on three of the four spatial scales: individual host plant, microhabitat, and macrohabitat. There was no correlation between vegetation structure and egg clutch occurrence or parasitism on any spatial scale. A high percentage of egg clutches (38–56%) was parasitized by Foersterella reptans Nees (Hymenoptera: Tetracampidae), the only egg parasitoid, but there was no relationship between egg parasitism and the spatial distribution of egg clutches of C. canaliculata on any of the spatial scales investigated. However, we also discuss results from a further study, which revealed top-down effects on the larval stage.     

Patterns and monitoring of Sesia apiformis infestations in poplar plantations at different spatial scales

Poplar plantations are expanding worldwide but little is known about the hornet clearwing moth, Sesia apiformis, one of their most severe pests. Thirty‐two poplar plantations of I‐214 clone were sampled in Spain, according to a factorial design combining stand age, site condition and understorey management to investigate the main drivers of S. apiformis habitat selection at both tree and stand level. In each plantation, one pheromone trap was activated during the flight season to test the correlation between captures and percentage of attacked trees. The proportion of other poplar forests in surrounding landscapes was calculated and used as a covariate in predictive models of trap catches. There were significantly more attacked trees in older stands. In young stands, the percentage of infested trees increased with the percentage cover of understorey vegetation. There was no significant effect of site quality on the rate of infestation and no difference in tree height, diameter or crown condition between attacked and un‐attacked trees within each stand, suggesting that S. apiformis could behave as primary pest. We hypothesized that the critical stage in the life cycle of the moth was the first instar larvae, which may benefit from protection of deep bark cracks in older stands and understorey vegetation in younger stands. We observed a positive correlation between trap capture and percentage of attacked trees in a radius of 100 m around the trap. The regression of trap catches against percentage of attacked trees was improved when the area of other poplar plantations within a distance of 600 m was incorporated in the model. This suggests that surrounding poplar stands may act as sources of immigrating moths in monitored stands. Our findings confirm that S. apiformis should be considered as a potential threat to poplar plantations and that pheromone trapping provides a suitable monitoring tool.     

Relevance of multiple spatial scales in habitat models: A case study with amphibians and grasshoppers

Habitat models for animal species are important tools in conservation planning. We assessed the need to consider several scales in a case study for three amphibian and two grasshopper species in the post-mining landscapes near Leipzig (Germany). The two species groups were selected because habitat analyses for grasshoppers are usually conducted on one scale only whereas amphibians are thought to depend on more than one spatial scale.First, we analysed how the preference to single habitat variables changed across nested scales. Most environmental variables were only significant for a habitat model on one or two scales, with the smallest scale being particularly important. On larger scales, other variables became significant, which cannot be recognized on lower scales. Similar preferences across scales occurred in only 13 out of 79 cases and in 3 out of 79 cases the preference and avoidance for the same variable were even reversed among scales.Second, we developed habitat models by using a logistic regression on every scale and for all combinations of scales and analysed how the quality of habitat models changed with the scales considered. To achieve a sufficient accuracy of the habitat models with a minimum number of variables, at least two scales were required for all species except for Bufo viridis, for which a single scale, the microscale, was sufficient. Only for the European tree frog (Hyla arborea), at least three scales were required.The results indicate that the quality of habitat models increases with the number of surveyed variables and with the number of scales, but costs increase too. Searching for simplifications in multi-scaled habitat models, we suggest that 2 or 3 scales should be a suitable trade-off, when attempting to define a suitable microscale.