Effect of habitat complexity on population density and species richness in tropical intertidal predatory gastropod assemblages

Summary Structural complexity of the habitat significantly increases population density and number of species in assemblages of predatory gastropod molluscs (families Conidae, Muricidae, Mitridae and Vasidae) on intertidal, generally smooth, horizontal limestone platforms fringing tropical Pacific islands. The important topographic features are physical (depressions partly filled with coral rubble) and biotic (thick algal turf binding sand). Higher population density and species richness in areas with than without such natural refuges, and following experimental addition of artificial refuges on portions of habitat lacking them support this hypothesis. Two species of Drupa differ from the other species present in not utilizing refuges during times of physical stress; this is attributed to their depressed shell and broad, tenacious foot. Highest gastropod densities occur in steep-sided depressions and those containing much coral rubble and sand, suggesting that these are important qualities of refuges. We believe this is the first demonstration of how specific environmental factors affect population density and species diversity of benthic invertebrates in a coral reef-associated habitat.     

Statutory protected areas and avian species richness in Britain

An effective portfolio of protected areas should, all else being equal, give rise to positive relationships between the amount of protected land in a region and the numbers of species present. Tests of this prediction are, however, extremely scarce, and most do not control for the potentially confounding effects of environmental factors that influence broad geographic trends in biodiversity. Here, we document the form of the relationship between species richness and coverage by protected areas using the British avifauna as a case study. We contrast relationships that arise for breeding and wintering assemblages, considering both all species collectively and threatened species only. We use spatially explicit multiple regression analyses that take into account environmental factors previously shown to exert a marked influence on avian species richness in Britain (temperature and altitude). Avian species richness and the amount of protected land are consistently positively correlated with each other, and the slopes of these relationships do not differ between assemblages (breeding/wintering and all species/threatened species). Explanatory power is, however, very weak which may be indicative of the ability of conservation measures in the wider landscape to maintain avian species richness, reducing any distinctive influence of protected areas. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The role of historical processes in determining tree species richness in the forests of Western Caucasus

To estimate the role of history in variation of tree species richness in the forests of the Western Caucasus we analyzed correlation between their local richness (S--the mean number of species per 300 m2) and size of actual species pool (N--the number of species per 1 ha). If compared communities are differently distant from the point of evolutionary equilibrium one should expect a significant variation in correlation between S and N (determined with the greater sensitivity of N than S in respect of historical factors). The lower value of N/S corresponds to less saturated level of historically determined species richness. A mean N/S ratio in Japana temperate broadleaved forests (Masaki et al., 1999) provided the basis for analysis. The present tree species richness of the forest communities in the 1 ha plots seem essentially determined by the historical processes. The mountain forest communities of Western Caucasus are characterized on the average with lower saturation level of the actual tree species pool in comparison with the Japan temperate broad-leaved forest communities. On the Western Caucasus the middle mountain beech and coniferous-broadleaved communities (400-1600 m a.s.l.) are characterized with the higher saturation level of the actual species pool in comparison with communities located lower and higher. These results confirm published historical reconstructions, according to which the middle mountain forest communities in the Western Caucasus are older than forests located higher or lower. Present low mountain forests of the southern (to Black Sea) and the northern macroslopes of the Western Caucasus are characterized with similar saturation level of the actual species pool. These data agree with the assumption of Dolukhanov (1980) that low mountain zone of the southern macroslope was not a refuge for tree species in Pleistocene.     

Lizard species richness patterns in China and its environmental associations

Many ecological hypotheses have been widely used to explain species richness variation across the globe. We investigated lizard species richness patterns in China, and identified areas of high species richness. Furthermore, we tested hypotheses concerning the relationships between lizard richness and environmental variables. A large data including 30,902 records of point locality data for 151 lizard species occurring in China were retrieved from Herpetology museums of CIB/CAS and other museums through HerpNET, and published sources, and then predicted distributions maps were generated using ecological niche modeling. We overlaid all species prediction maps into a composite map to describe species richness patterns. A multiple regression analysis using eigenvector-based spatial filtering (SEVM) was performed to examine the best environmental predictors of species richness. Richness peaked mainly in southern China located in the Oriental realm. Our best multiple regression models explained a total of 80.1% variance of lizard richness (r² = 0.801; F = 203.47; P < 0.001). Among related factors in shaping species richness distribution, the best environmental predictors of species richness were: frost-day frequency, elevation, vegetation, and wet-day frequency. Based on models selection, our results revealed that underlying mechanisms related to different ecological hypotheses might work together and best explain lizard richness in China. We are in an initial step to develop a large data set on species richness, and provide the necessary conservation implications from habitat loss. Additional studies that test species richness at different geographical scale are required to better understand the factors that may influence the species richness distribution in East Asia.     

Protected areas and regional avian species richness in South Africa

Protected areas are generally regarded as essential for the long-term maintenance of biodiversity. Evidence for their effectiveness in this regard is, however, somewhat equivocal. Here, we document the relationship between the proportion of protected land and species richness in a region, both with and without taking spatial variation in environmental energy availability into account. Using the South African avifauna as a case study, we find that total and threatened species richness exhibit modest increases with the proportion of protected land. While the protected area network should be expanded, it is essential that conservation efforts also focus on maintaining biodiversity in the wider unprotected landscape that supports high species richness.     

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.     

Predicting distributions of species richness and species size in regional floras: Applying the species pool hypothesis to the habitat templet model

The species pool hypothesis is applied here to the interpretation of ‘hump-shaped’ (unimodal) species richness patterns along gradients of both habitat fertility and disturbance level (the habitat templet). A ‘left-wall’ effect analogous to that proposed for the evolution of organismal complexity predicts a right-skewed unimodal distribution of historical habitat commonness on both gradients. According to the species pool hypothesis, therefore, the distribution of opportunity for net species accumulation (speciation minus extinction) should also have a corresponding unimodal central tendency on both habitat gradients. Two assumptions of this hypothesis are illustrated with particular reference to highly fertile, relatively undisturbed habitats: (i) such habitats have been relatively uncommon in space and time, thus providing relatively little historical opportunity for the origination of species with the traits necessary for effective competitive ability under these habitat conditions; and (ii) species that have evolved adaptation to these habitats are relatively large, thus imposing fundamental ‘packing’ limitations on the number of species that can ‘fit’ within such habitats. Based on these assumptions, the species pool hypothesis defines two associated predictions that are both supported by available data: (a) resident species richness will be relatively low in highly fertile, relatively undisturbed contemporary habitats; and (b) species sizes within regional floras should display as a right-skewed unimodal (log-normal) distribution. The latter is supported here by an analysis of data for 2,715 species in the vascular flora of northeastern North America.     

Landscape composition and habitat area affects butterfly species richness in semi-natural grasslands

During the last 50 years, the distribution and abundance of many European butterfly species associated with semi-natural grasslands have declined. This may be the result of deteriorating habitat quality, but habitat loss, resulting in decreasing area and increasing isolation of remaining habitat, is also predicted to result in reduced species richness. To investigate the effects of habitat loss on species richness, we surveyed butterflies in semi-natural grasslands of similar quality and structure, but situated in landscapes of different habitat composition. Using spatially explicit habitat data, we selected one large (6–10 ha) and one small (0.5–2 ha) grassland site (pasture) in each of 24 non-overlapping 28.2 km² landscapes belonging to three categories differing in the proportion of the area that consisted of semi-natural grasslands. After controlling for local habitat quality, species richness was higher in grassland sites situated in landscapes consisting of a high proportion of grasslands. Species richness was also higher in larger grassland sites, and this effect was more pronounced for sedentary than for mobile species. However, the number of species for a given area did not differ between large and small grasslands. There was also a significant relationship between butterfly species richness and habitat quality in the form of vegetation height and abundance of flowers. In contrast, butterfly density was not related to landscape composition or grassland size. When species respond differently to habitat area or landscape composition this leads to effects on community structure, and nestedness analysis showed that depauperate communities were subsets of richer ones. Both grassland area and landscape composition may have contributed to this pattern, implying that small habitat fragments and landscapes with low proportions of habitat are both likely to mainly contain common generalist species. Based on these results, conservation efforts should aim at preserving landscapes with high proportions of the focal habitat.     

The role of landscape structure in determining palynological and floristic richness

The associations between floristic and palynological richness and landscape structure were studied based on modern pollen?Cvegetation data from a patchy cultural landscape in southern Estonia (northern temperate vegetation zone). Nine study sites (small lakes and their surrounding vegetation) represent land cover gradient from closed forest to semi-open vegetation. Floristic richness (number of species) and floristic richness of pollen types (number of pollen-equivalent taxa) were used to describe the vegetation within the radius of 250?m from the pollen sampling sites. Palynological richness was calculated to describe the modern pollen samples diversity. Landscape structure was estimated on the basis of landscape openness and three landscape diversity measures: richness of community patches, Simpson evenness of community patches and Simpson diversity of community patches. To study the effect of the spatial scale of landscapes on the vegetation?Clandscape and pollen?Clandscape associations, landscape structure was estimated within eight radii (250?C2,000?m) around each lake. The results showed that landscape openness was the most important determinant of both floristic richness and palynological richness in southern Estonia and that landscape diversity estimated by Simpson diversity index was also significantly associated with the richness estimates. Floristic and palynological richness were significantly positively correlated with landscape structure within the radii greater than 1,000?m from the pollen sampling sites, which is similar to the estimated Relevant Source Area of Pollen in southern Estonia. We conclude that within one floristic or climatic region, palynological richness gives reliable estimates about the variation in floristic richness and landscape structure; however, caution must be taken when comparing pollen-inferred vegetation diversities from different regions or when interpreting fossil pollen records from times with highly different vegetation associations.     

Mammal density and patterns of ectoparasite species richness and abundance

Patterns of species richness, prevalence and abundance of ectoparasites have rarely been investigated at both the levels of populations and species of hosts. Here, we investigated the effects in changes in small mammal density on species richness, abundance and prevalence of ectoparasitic fleas. The comparative analyses were conducted for different small mammal species and among several populations during a long-term survey. We tested the hypothesis that an increase in host density should be linked with an increase in parasite species richness both among host species and among populations within host species, as predicted by epidemiological models. We also used host species density data from literature. We found that host density has a major influence on the species richness of ectoparasite communities of small mammals among host populations. We found no relationship between data of host density from the literature and parasite species richness. In contrast with epidemiological hypotheses, we found no relationships between abundance, or prevalence, and host density, either among host species or among host populations. Moreover, a decrease in abundance of fleas in relation with an increase in host density was observed for two mammal species (Apodemus agrarius and A. flavicollis). The decrease or the lack of increase in flea abundance in relation with an increase in host density suggests anti-parasitic behavioural activities such as grooming.     

Effects of habitat fragmentation and isolation on species richness: evidence from biogeographic patterns

Summary Habitat subdivision by geography or human activity may be an important determinant of regional species richness. Cumulative species-area relationships for vertebrates, land plants, and insects on island archipelagoes show that collections of small islands generally harbor more species than comparable areas composed of one or a few large islands. The effect of the degree of habitat subdivision in increasing species richness appears to increase with the distance from potential sources of colonists. Mountaintop biotas show no clear differences between species richness on large alpine areas and collections of smaller peaks. National park faunas generally have more species in collections of small parks than in the larger parks. In all cases where a consistent effect of subdivision is observed, the more subdivided collection of islands or isolates contains more species. To the degree that these data provide guidance for establishing nature reserves, they suggest that increasing the numbers of reserves may be an important component of conservation strategies.     

Dispersal capability in a habitat specialist bush cricket: the role of population density and habitat moisture

1. In fragmented landscapes many insect species depend on a regular exchange of individuals between subpopulations to ensure the persistence of the population. Thus, the ability to disperse is of particular relevance. 2. However, in some insect species mobility is not a fixed trait. Hence, knowing the causes of phenotypic plasticity is of great importance when evaluating whether a species is able to survive in fragmented landscapes or not. 3. A multi‐year field study was conducted to identify possible causes of macroptery in the wing‐dimorphic habitat specialist Metrioptera brachyptera L. and to quantify its dispersal capability (% macropters). Therefore, 746 individuals of the species were caught on 135 plots. Additionally, environmental variables that possibly induce the development of macropters (population density and habitat moisture) were recorded. 4. Dispersal capability of M. brachyptera was very low. Less than 3% were long‐winged. The statistical analysis revealed that the proportion of long‐winged M. brachyptera was strongly correlated with high bush‐cricket densities and not with habitat moisture. 5. The low dispersal capability of M. brachyptera leads to the conclusion that individual exchange between isolated populations is limited or even impossible. Habitat specialists, like M. brachyptera, may thus be unable to respond to rapid changes in the availability of suitable habitats by dispersing, and hence may be especially dependent on habitat management activities that promote the long‐term stability of existing habitat patches.     

Distinguishing area and habitat heterogeneity effects on species richness: Birds in Victorian buloke remnants

Abstract Resolving whether area per se or habitat heterogeneity has the greater influence in controlling species richness remains a controversial yet important question. Here we show that avian species richness of same-sized transects (1 ha) is independent of the remnant area (of buloke woodland) within which a transect is positioned. We also show that avi-faunal similarity of pairs of transects randomly placed within the largest remnants (≥ 48 ha) is not consistently related to either proximity (i. e. being within the same remnant) nor to physiognomic characteristics of the transects. We believe that much of the controversy over area/habitat heterogeneity effects is probably related to scalar issues and propose a protocol by which some resolution of the question might be reached. The protocol involves ‘zoom’ sampling in which successively larger transect sizes are used, and measures of faunal richness and habitat heterogeneity are made at these different grains of resolution. One of our intentions is to stimulate discussion on how heterogeneity might be measured when grains increase from typical transect sizes (ca 1 ha) up to much larger grains (ca 128 ha).     

Assessing bee species richness in two Mediterranean communities: importance of habitat type and sampling techniques

The decline of bees has raised concerns regarding their conservation and the maintenance of ecosystem services they provide to bee-pollinated wild flowers and crops. Although the Mediterranean region is a hotspot for bee species richness, their status remains poorly studied. There is an urgent need for cost-effective, reliable, and unbiased sampling methods that give good bee species richness estimates. This study aims: (a) to assess bee species richness in two common Mediterranean habitat types: semi-natural scrub (phrygana) and managed olive groves; (b) to compare species richness in those systems to that of other biogeographic regions, and (c) to assess whether six different sampling methods (pan traps, variable and standardized transect walks, observation plots and trap nests), previously tested in other European biogeographic regions, are suitable in Mediterranean communities. Eight study sites, four per habitat type, were selected on the island of Lesvos, Greece. The species richness observed was high compared to other habitat types worldwide for which comparable data exist. Pan traps collected the highest proportion of the total bee species richness across all methods at the scale of a study site. Variable and standardized transect walks detected the highest total richness over all eight study sites. Trap nests and observation plots detected only a limited fraction of the bee species richness. To assess the total bee species richness in bee diversity hotspots, such as the studied habitats, we suggest a combination of transect walks conducted by trained bee collectors and pan trap sampling.     

Theoretical habitat templets, species traits, and species richness: floodplain vegetation in the Upper Rhône River

• 1 The floodplain vegetation at approximately 100 sites located in nine different habitat types of the Upper Rhône River, France, was surveyed three times over the past 27 years. Information on species traits of the higher plants comprising the Rhône floodplain vegetation was based on studies conducted between Geneva, Switzerland, and Lyon, France.
• 2 These data were structured using a ‘fuzzy coding’ technique and then examined using ordination analyses to investigate: (i) relationships among species traits; (ii) habitat utilization; (iii) the relationship between species traits and habitat utilization; and (iv) trends of species traits and species richness in the framework of spatial–temporal habitat variability to test predictions of the river habitat templet and the patch dynamics concept.
• 3 Size, number of descendants per reproductive cycle, number of reproductive cycles per individual, and the regeneration potential of an individual were positively related with each other, whereas the degree of attachment to the soil decreased, and the reproductive period shifted from autumn/late summer towards early summer/spring, as size increased.
• 4 The habitat utilization by the higher plants of the floodplain revealed a double lateral gradient: the first was from the banks of the temporary waters to terrestrial flats; the second from aggrading pebble to aggrading silt habitats. These gradients were related to gradients in water saturation, oxygen conditions, nutrient loading, and nutrient retention of the soils.
• 5 A significant relationship between species traits and habitat utilization was observed for the floodplain vegetation, i.e. plant communities used particular habitat types with a particular set of species trait modalities (= categories).
• 6 Patterns of species trait modalities were significantly related to temporal and spatial habitat variability but only modalities of the trait ‘parental care’ conformed to trends predicted from theory.
• 7 No trends were observed when species richness of different habitat types was considered in the framework of spatial–temporal habitat variability.
• 8 Although the habitats of the Upper Rhône clearly act as a templet for the species traits of the floodplain vegetation, the lack of agreement between observations and predictions on trends in species traits and richness in terms of habitat variability suggest that important elements of theory should be rejected. However, human-induced changes in these habitats are too recent when compared with the longer time periods required for floodplain vegetation to respond to such changes.

     

Invasive birds in a novel landscape: habitat associations and effects on established species

Studying the relationships among introduced species, their preferred habitats, and native species can be important for predicting the effects of invasions on native populations. Examining the colonization of North America by the Eurasian collared‐dove Streptopelia decaocto, we quantified the habitat characteristics of sites most likely to be occupied by this invasive bird species in the early stages of the invasion. Further, we studied the relationship between collared‐dove abundance and the abundance of other dove species in the study area, anticipating a negative effect on established species following the introduction of a potential competitor. Linking satellite‐derived landcover data with winter bird community data gathered from 444 study sites in Florida, USA from 1999 to 2008, we found that collared‐doves were more likely to occur in landscapes that had been highly‐modified by human activity than in forested landscapes. Collared‐dove abundance increased as the proportion of the landscape characterized as low‐intensity development and medium/high‐intensity development increased. The probability of collared‐doves occurring at a site was also related to the spatial proximity of other sites reporting doves (positive spatial autocorrelation). Contrary to our expectations, the site‐level abundance of four other dove species all increased with collared‐dove abundance throughout the sampling period. Interactions between collared‐doves and native species should be further studied in different environments as this invasive bird rapidly colonizes North America.