Response of a desert lizard community to habitat degradation: Do ideas about habitat specialists/generalists hold?

We report the impact of human-induced desertification on the species richness, abundance, and composition of sand dune flora and herpetofauna of North Sinai, Egypt. Our hypothesis was that degraded habitats would have reduced vegetation complexity, richness, and abundance, and consequently lower reptile species richness and abundance. We also hypothesized that desert lizards would not follow the typical generalist/specialist responses to habitat degradation found in other biomes. Instead, we predicted that because vegetation loss intensifies the environmental extremity of deserts, those species specialized for open and sandy environments would be more likely to persist in desertified habitats than would desert generalists. Our results showed that areas protected from vegetation loss did not have significantly higher vegetation richness or abundance except for only a few species. However, protected sites did have significantly higher percent vegetation cover and height. Habitat protection clearly had strong effects on the reptile community as species richness and abundances were significantly higher in protected sites. The composition of the reptile community between protected and unprotected sites differed significantly. Contrary to past studies in other environments, desert generalist species were not able to persist in degraded sites and were only found in protected sites. Specialist species were ubiquitous in that they occurred in both areas protected and unprotected from vegetation loss. We propose that the effects of disturbance on species composition (specialists or generalists) depends on whether the disturbance exacerbates or reduces environmental harshness and the conditions that favor specialization. In extreme environments, specialist and generalist responses to habitat degradation are opposite to that of more productive environments.     

Rabbits as a keystone species in southern Europe

A keystone species is one that is crucial in maintaining the organization and diversity of their ecological communities. We tested the idea that rabbits act as a keystone species in southern Europe by exploring relationships between rabbit abundance and the abundance and diversity of raptor species. At 20 sites in southern Spain we assessed rabbit abundance through counts of animals along transects and assessed the number of raptor individuals and species through watches from vantage points. In a further 120 locations we also derived an index of rabbit abundance, from pellets and compared this to the presence or absence of the critically endangered Spanish imperial eagle. Rabbit abundance was positively associated with the number of observations of raptors, the number of raptor species and the number of species of conservation concern. Sites with the most rabbits had higher conservation value. Moreover, the presence of Spanish imperial eagle was strongly associated with sites where rabbits were at high density. We conclude that rabbits do act as a keystone species and we suggest that conservation efforts should focus on improving the status of this small-game species in southern Europe.     

Land use changes and raptor conservation in steppe habitats of Eastern Kazakhstan

Steppe habitats in central Asia have suffered important land use changes during this century which are similar to those that have been pointed out as the causes of the decline of steppe birds in western Europe. During June 1999 we conducted road surveys of raptors in Eastern Kazakhstan to detect specific and community responses to land use changes. We detected 11 species of raptors. Kestrels (Falco naumanni and tinnunculus) were the most common species in grasslands and agricultural landscapes, harriers (Circus pygargus and macrourus) were dominant in saline steppes and steppe eagles (Aquila nipalensis) were dominant in dry steppes. There were fewer species in agricultural habitats than in grassland and steppe habitats. Ground-nesting raptors were negatively affected by land use changes and four species were never detected in agricultural zones. Raptor abundance patterns differed between natural steppe habitats and human-transformed habitats, where a patchy distribution was detected. The future of raptor communities in Kazakhstan seems uncertain although the progressive abandonment of intensive agriculture may benefit species sensitive to human presence. The long-term conservation of vertebrate communities may depend upon the maintenance of ecologically and socially sustainable grazing systems.     

Land use and habitat gradients determine bird community diversity and abundance in suburban, rural and reserve landscapes of Minnesota, USA

Bird species’ community responses to land use in the suburbanizing Twin Cities, Minnesota, USA, were contrasted among reserves, rural lands, and suburbs. For each land use type, bird composition, diversity, and abundance were recorded for 2 years in ≈99 plots in three sampling units (each ≈4500 ha). A habitat gradient defined by canopy structure (grasslands to savannas to forests) was influenced by land use, so ≈300 plots were used to characterize simultaneous variation in bird communities along land use and habitat gradients. At broad scales (aggregate of 33 plots covering ≈4500 ha) suburbs supported the lowest bird richness and diversity and rural landscapes the most, with reserves slightly below rural. Although reserves were like rural lands in diversity of bird communities, they supported more species of conservation concern, particularly of grasslands and savannas. Differences among land use types varied with habitat structure. Suburbs, rural lands, and reserves had similar forest bird communities, but differed in grassland and savanna bird communities. The extensive rural forests are important for the region’s forest birds. Suburban grasslands and savannas had low shrub abundance, low native bird richness and high non-native bird richness and abundance. However, total bird richness and diversity were as high in suburban as in rural and reserve plots because high native richness in suburban forests and high non-native species richness in suburban grasslands and savannas compensated for lower native richness in suburban grasslands and savannas. Bird conservation here and in the Midwest USA should protect rural forests, expand grasslands and savannas in reserves, and improve habitat quality overall.     

Avian assemblages along a gradient of urbanization in a highly fragmented landscape

Our goal was to evaluate how avian assemblages varied along a gradient of urbanization in the highly fragmented landscape of coastal southern California. We measured species richness and abundance of birds within continuous blocks of habitat, within urban habitat fragments that varied in landscape and local habitat variables, and within the urban matrix at different distances from the wildland interface. These comparisons allowed us to characterize patterns of avifaunal response to a gradient of urban fragmentation. At the fragment scale, we found that fragment area was a strong, positive predictor of the total number of breeding species detected per fragment; total bird abundance per point count also increased with fragment size. Tree cover was higher in small fragments, as was the abundance of birds that typically occupy wooded habitats. Comparisons between core, fragment, and urban transects revealed differing patterns of response of individual bird species to urbanization. In unfragmented habitat, we recorded a relatively high diversity of urbanization-sensitive birds. In urban transects, these species were rare, and a relatively few species of non-native and anthropophilic birds were common. These urbanization-enhanced birds were also recorded in previous urban gradient studies in northern California and Ohio. Bird communities along the urban gradient reached their highest richness and abundance in fragments. The marked difference in vegetation structure between urban and natural landscapes in this arid shrubland system likely contributed to this pattern; the presence of native shrubs and exotic trees in fragments enabled both shrub and arboreal nesters to co-occur. As is characteristic of biotic homogenization, urban fragmentation in coastal southern California may increase local diversity but decrease overall regional avifaunal diversity.     

Riparian zones provide for distinct bird assemblages in forest mosaics of south-east Australia

Riparian zones are a characteristic component of many landscapes throughout the world and increasingly are recognised as key areas for biodiversity conservation. Their importance for bird communities has been well recognised in semi-arid environments and in modified landscapes where there is a marked contrast between riparian and adjacent upslope vegetation. The value of riparian zones in largely intact landscapes with continuous vegetation cover is less well understood. In this study, birds were surveyed at 30 pairs of riparian and adjacent non-riparian sites in extensive mesic forests of the Victorian Highlands, Australia. Riparian sites were floristically distinct from non-riparian sites and had a more complex vegetation structure, including a mid-storey tree layer mostly absent from non-riparian sites. Bird assemblages at riparian sites had significantly greater richness, abundance and diversity of species than was recorded at adjacent non-riparian sites. Species composition also differed significantly between these habitat types. Compositional differences in assemblages were due to a suite of distinctive species in each habitat and to significant contrasts in the densities of species that occurred in both habitat types. Many species (36%) attained a significantly greater abundance in riparian habitats. The distinctiveness and richness of the riparian avifauna contribute to the diversity of continuous forest landscapes. The spatial patterning of the avifauna, the occurrence of complementary assemblages, the presence of rare species and the potential for riparian habitats to serve as refuges, all point to the value of riparian zones and highlight the importance of landscape-level planning and management for avifaunal conservation.     

From multi-criteria approach to simple protocol: Assessing habitat patches for conservation value using species rarity

We investigated conservation value (CV) related to the quality of spider communities in different non-wooded habitat patches - ranging from arable land to natural grasslands. The study was conducted in two ecologically distinct regions of Hungary: the Hungarian Great Plain and the Buda Hills. We used seven variables to indicate CV which together formed a multi-criteria space of spider community characteristics. These variables were either related to species characters obtained from an extensive background database: abundance and frequency based rarity, specialist status, association to natural habitats; or were calculated for the community at the given patch: species richness, functional diversity and species evenness. Using the variables in an ordination analysis we could establish a gradient of the patches in the multi-criteria space of the spider community characteristics. Position of patches along the first axis of the ordination was taken as the multi-criteria measure of CV. CVs established this way were strongly and positively correlated with an independent botanical CV assessment. We also sought a simpler measure of spider CV by: (a) calculating only one variable out of the seven and using it as a surrogate for the multi-criteria CV measurement; by (b) calculating this variable only for a short time period or (c) for only one spider family. Average abundance based rarity value of the species proved to be the best surrogate of the multi-criteria CV measure for both regions, and it also performed very well when sample size was restricted to two sampling occasions per patch or to a single family. This adds further evidence to, what has been found in other studies, that species rarity is a sensitive and reliable measure of the ecological and conservational status of communities.     

Small isolated aspen stands enrich bird communities in southwestern ponderosa pine forests

Small aspen stands are disappearing from the landscape in the Southwest, so it is important to understand their contribution to the avian community. We sampled birds in 53 small, isolated aspen stands and 53 paired plots within the ponderosa pine forest in northern Arizona, during the 1996 and 1997 breeding seasons. Bird species richness and abundance were higher in aspen than in pine. However, bird species richness and abundance did not vary with size of the aspen patch or isolation index. In addition, direct ordination of species distributions with habitat factors suggested no distinct avian communities. This suggests that aspen stands do not harbor separate populations, but rather are locations where the regional avifauna reaches high local density and richness and may be crucial to birds in years of resource scarcity. Thus it is important for avian conservation to maintain many aspen stands across the landscape, encompassing a diversity of vegetation structure and composition.     

Endemic species: Contribution to community uniqueness, effect of habitat alteration, and conservation priorities

The biodiversity crisis, particularly dramatic in freshwaters, has prompted further setting of global and regional conservation priorities. Species rarity and endemism are among the most fundamental criteria for establishing these priorities. We studied the patterns of rarity and the role of rare species in community uniqueness using data on freshwater bivalve molluscs (family Unionidae) in Texas. Due to the large size and gradients in landscape and climate, Texas has diverse and distinct unionid communities, including numerous regional and state endemic species. Analysis of the state-wide distribution and abundance of Unionidae allowed us to develop a non-arbitrary method to classify species rarity based on their range size and relative density. Of the 46 Unionidae species currently present in Texas, 65% were classified as rare and very rare, including all state and regional endemics. We found that endemic species were a critical component in defining the uniqueness of unionid communities. Almost all endemics were found exclusively in streams and rivers, where diversity was almost double that of lentic waters. Man’s ongoing alteration of lotic with lentic waterbodies favors common species, and dramatically reduces habitat for endemics, contributing to homogenization of unionid fauna. We identified hotspots of endemism, prioritized species in need of protection, estimated their population size, and recommended changes to their current conservation status.     

Disentangling effects of habitat diversity and area on orthopteran species with contrasting mobility

Loss of semi-natural grasslands and reduction of habitat diversity are considered major potential threats to arthropod diversity in agricultural landscapes. The main aim of this study was to investigate how area and habitat diversity, mediated by shrub encroachment after grassland abandonment, affect species richness of orthopterans in island-like grasslands, and how contrasting mobility might alter species richness response to both factors. We selected 35 isolated patches in landscapes dominated by arable land (durum wheat) in order to obtain two statistically uncorrelated gradients: (i) one in habitat area ranging from 0.2 to 55 ha and (ii) one in habitat diversity ranging from patches dominated by one habitat (either open grasslands or shrublands) to patches with a mosaic of different habitats. Habitat loss due to land-use conversion into arable fields was associated with a substantial loss of species with a positive species-area relationship (SAR), with sedentary species having a steeper and stronger SAR than mobile species. Halting habitat loss is, therefore, needed to avoid further species extinctions. Shrub encroachment, triggered by abandonment, presented a hump-shaped relationship with habitat diversity. An increase in habitat diversity enhanced species richness irrespective of patch area and mobility. Maintaining or enhancing habitat diversity, by cutting or burning small sectors and by reintroducing extensive sheep grazing into abandoned grassland, are suggested as complementary strategies to mitigate further decline of orthopteran diversity in the remnant patches. This would be equally important in both small and large patches.     

Grazing promotes dung beetle diversity in the xeric landscape of a Mexican Biosphere Reserve

We analyze the impact of grazing on dung beetle diversity at the Barranca de Metztitlán Biosphere Reserve, a xeric ecosystem in central Mexico with a long history of use by humans. We compared the community structure, as well as the alpha and beta diversity between two cover conditions (open and closed vegetation) that represent the impact of grazing within a habitat, and between habitat types (submountainous and crassicaule scrublands). From 576 samples we collected 75,605 dung beetles belonging to 20 taxa. While mean species richness and diversity were different between habitat types, cumulative species richness was not. The effects of grazing on vegetation structure influenced the cumulative species richness and diversity of dung beetles in the submountainous scrubland, where grazing has created land mosaics of a grassland matrix with scrubland patches. This was not the case in the crassicaule scrubland where the impact of grazing is not as evident. Beta diversity significantly responds to the effects of grazing on habitat conditions. We discuss the ecological factors that may promote these responses by landscape diversity components. We also identify the species that could act as useful indicators to monitor the effect of land management on biodiversity. Our results indicate cattle farming maintains a diversified land mosaic, and these areas support more diverse dung beetle ensembles than homogeneous areas of closed, shrubby vegetation cover. Thus, controlled grazing activity could certainly favour the conservation of dung beetle biodiversity and improve ecosystem functioning by maintaining dung decomposition rates.     

Toward more concern for specialisation and less for species diversity in conserving farmland biodiversity

In order to face the large and worrying biodiversity decline in agricultural landscapes, important policy instruments like agri-environmental schemes (AES) have been implemented. Studies that have examined the ecological effects of AES are now numerous and generally use indicators of biodiversity such as species richness and diversity as well as species abundance. Yet, it has been shown that simple metrics such as species richness or abundance may give misleading messages about biodiversity status and fate. Moreover, those indicators cannot detect another important source of biodiversity loss, biotic homogenisation. In this context, taking into account to a wider extent ecological difference among species would be more relevant, as well as focusing on the species specialisation which is known to be linked to higher species vulnerability. A bibliographic review investigating the criteria generally used to assess the success of AES showed that 55% of studies used species richness and/or abundance exclusively without any consideration of specialisation or other ecological traits in their evaluation of AES effectiveness. Based on data from the French breeding bird survey and studies at regional scale in France on farmland birds, we show that: (i) species richness and specialisation are generally negatively correlated in agricultural areas, (ii) habitat heterogeneity does not benefit specialist species, and (iii) monitoring of species diversity should be coupled with the monitoring of specialist species to improve conservation strategies in farming systems. Overall, this study emphasizes the need to account for both community richness and composition when assessing AES or similar conservation planning.     

Bird community structure across riparian buffer strips of varying width in a coastal temperate forest

Buffer strips are strips of forest retained along streamsides after harvesting to mitigate negative impacts of forestry on aquatic and riparian fauna and water quality. The capacity of riparian buffer strips of old-growth forest to maintain species richness and abundance of natural bird communities was explored in coastal montane forest on Vancouver Island, Canada. Breeding bird communities in buffer strips of varying widths along rivers were compared with controls of equivalent area in uncut old-growth riparian forest to identify shifts in species richness, diversity, abundance and composition. We observed that effects on riparian bird communities were greatest in very narrow buffers with high amounts of edge habitat. Several forest-interior species were found almost exclusively in wider buffers and abundances dropped dramatically between wide (125 m) and medium (41 m) width buffers with replacement by open-edge species in narrow buffers. Species composition of communities in wide buffers were very similar to controls while narrow buffers shared less than half of their species with controls. Species richness and diversity increased in buffers over the three years while remaining constant in controls. Increases in species richness and abundance within buffers were positively correlated with similar increases in the adjacent clearcuts, suggesting that regeneration in clearcuts may facilitate recolonization of forest in remnant buffers. For the forest-interior species found primarily in wide buffers, buffers >100 m may need to be retained.     

Long-term consequences of habitat fragmentation—highland birds in Oaxaca, Mexico

Studies of habitat fragmentation have been restricted primarily to anthropogenically-altered habitats, with most research conducted 60-90 years post-fragmentation. It is unclear whether patterns in older systems concur with results from these dynamic landscapes, and hence the long-term viability of populations inhabiting habitat fragments remains largely unexplored. I focused on resident birds in fragments of humid pine-oak forest in Oaxaca, southern Mexico, isolated over 5000 years ago by climate-change. Seventeen fragments, ranging from 2 ha to over 150,000 ha were sampled in 1997 and 1998 yielding 141 species, of which 60 residents were used for analysis. Avian assemblages exhibited a highly nested structure and, with several notable exceptions, assemblages of birds in low-richness fragments were predictable subsets of those in more diverse fragments. Patch-scale factors—area, shape, elevation, habitat diversity and fractal dimension of edge—all exerted strong univariate influence on avian richness but were so closely inter-related that none had a significant independent effect. Thus, larger fragments were more complex in shape, included higher peaks, supported more diverse forests, and contained higher diversities of resident species. In contrast, the landscape-scale index used—distance from nearest large fragment (>50,000 ha)—had little effect on richness. This was reinforced by species-level analyses—one species was significantly influenced by isolation, compared with 31 species that displayed significant minimum-area distributions, restricted to patches larger than a particular threshold value. In terms of autecology, vagility, relative abundance and elevational breadth were closely related to distribution—those species with greater mobility, higher abundances and broader elevational tolerances were consistently more widespread. I suggest that more abundant species were less prone to extinction initially, more vagile species were better dispersers and species with broader elevational tolerances more likely to be successful colonists. As with previous research from older landscapes, patch-scale factors were consistently found to be influential, with high quality fragments supporting diverse communities regardless of landscape context. This suggests that the influence of landscape-scale factors noted in younger, anthropogenically fragmented systems may be transitory, overwhelmed by patch-scale factors with time. Which patch attributes are most influential could not be resolved, however, indicating that even thousands of years after fragmentation, they affect diversity patterns in concert. Rather than differentiating effects of area from habitat heterogeneity and other patch-level factors, I advocate resource-based approaches to understand and manage diversity in habitat fragments.     

Influence of resource quality on the composition of soil decomposer community in fragmented and continuous habitat

The aim of this field experiment was to explore the combined effects of two factors potentially affecting the local composition of soil decomposer community: resource quality and habitat fragmentation. We created humus (habitat) patches with three different resource quality: (1) pure homogenised humus; (2) humus enriched with needle litter; and (3) humus enriched with needle and leaf litter. These patches were embedded either in a mineral soil matrix, thus representing fragmented habitat, or in natural forest soil, representing continuous (non-fragmented) habitat. The development of faunal (colonisations/extinctions of soil animal populations) and microbial communities in the patches was followed for 12 months. Our results partly supported the hypothesized strong influence of resource quality on the structure of local soil food webs: the abundances of practically all groups of soil fauna, together with biomass of fungi, were higher in the litter-enriched patches than in the pure humus patches. The manifestation and magnitude of the responses of fauna were, however, strongly affected by complex interactions between the characteristics (especially colonisation capacity) of the faunal group in question, habitat quality and time of sampling. In microarthropods and nematodes, the effect of resource quality cascaded up to the predatory level, rendering further support to the existence of strong bottom-up control in soil food webs. Contrary to our expectations, species richness of the communities was not unanimously affected by resource quality. Habitat fragmentation affected the communities only through different number and identity of patch-colonising species in the fragmented and continuous habitat: fragmentation induced no extinctions of species during the experiment at any resource quality level. Consequently, the results indicate that resource quality is more important factor than habitat fragmentation in determining the local structure of communities in soils. On the other hand, colonisation capacities of soil organisms appear to set limits to the exploitation of local resources.     

The effects of extreme forest fragmentation on the bird community of Singapore Island

Singapore Island suffered one of the highest known deforestation rates in the tropics from the mid-to-late 19th century when over 95% of its native lowland forest was cleared. We compared the current bird community structure and composition among three habitat types, i.e., old (>50 years, 7-935 ha) and young (?50 years, 29-49 ha) naturally regenerating secondary forests and abandoned wooded plantations (27-102 ha) dominated by exotic species. Forest patch area had the strongest influence on the current species richness. The overall bird richness was not greater in most mature forest patches, but 20 species were only found in the old secondary forests and five of these were found in <50% of these patches. The rapid decrease in the number of forest species in plantations was offset by an increase in the number of open habitat species. Comparisons with current bird communities in nearby mainland forest sites (Peninsular Malaysia) suggest that the forest avifauna of Singapore is depauperate. The preservation of larger mature and maturing forests is therefore required for conserving the extant forest avifauna in Singapore. Connecting isolated patches can also be envisioned to facilitate movements of forest birds that have low densities and restricted distribution.     

The influence of patch area and connectivity on avian communities in urban revegetation

Landscape restoration through revegetation is being increasingly used in the conservation management of degraded landscapes. To effectively plan restoration programs information is required on how the landscape context of revegetation influences biodiversity gains. Here, we investigate the relative influence of patch area and connectivity on bird species richness and abundance within urban revegetation patches in Brisbane, Australia. We carried out bird surveys at 20 revegetation sites, and used hierarchical partitioning and model selection to test the relative importance of patch area (the area of revegetation including all directly connected remnant vegetation) and landscape connectivity (the vegetated area connected by less than 10 m, 20 m, 30 m, 40 m and 50 m cleared gaps). We controlled for a number of possible confounding variables within the hierarchical partitioning procedure. Both the hierarchical partitioning and model selection procedures indicated that connectivity had an important influence on bird species richness. Patch area in combination with connectivity were important influencing factors on overall bird abundance. We also carried out the hierarchical partitioning procedure for bird abundance data within a range of feeding guilds, yielding results specific to species groups. Overall our data suggest that greater connectivity enhances the habitat area that colonists can arrive from (resulting in greater species richness), whereas increased patch area allows for increased abundance by expanding the habitat available to species already present in a patch. A combined approach where connectivity and overall habitat area is enhanced across the landscape is likely to be necessary to meet long-term conservation objectives.     

Traditional salt-pans hold major concentrations of overwintering shorebirds in Southeast Asia

Shorebirds are declining worldwide due to loss and degradation of critical breeding and wintering habitats. Some human-modified habitats, particularly salt-pans which are used by shorebirds in many regions of the world, may help substitute for natural habitats lost for a wide range of species during migration. We studied the influence of landscape characteristics on species richness, abundance, and diversity of shorebirds at 20 sites covering most of the Inner Gulf of Thailand, a landscape with a long history of salt farming. Sites with salt-pans present held significantly higher species richness, abundance and diversity of shorebirds. Areas with larger proportions given over to aquaculture tended to have lower species richness, abundance and diversity. Generalized additive models indicated that landscapes with a larger proportion of tidal flats in conjunction with salt-pans were the best predictors of sites with higher species richness, abundance and diversity. Landscape configurations with higher richness, abundance and diversity of shorebirds also tended to be less fragmented and contained slightly larger patches. Shorebirds appeared to use ponds with exposed mud in salt-pans as both roosting sites and supplementary feeding grounds during high tide. Traditional salt-pans therefore proved to contribute significantly to maintenance of overwintering shorebird populations in this landscape and should be investigated elsewhere in Asian coastal zones. Collaboration between researchers, salt farmers and planning authorities as to how best to maintain salt-pans as potential shorebird roost sites such as in the Inner Gulf of Thailand is urgently needed in order to maintain habitat for shorebird populations in critical wintering and staging areas of this flyway.     

Global diversity and distribution of arbuscular mycorrhizal fungi

Arbuscular mycorrhizal (AM) fungi form associations with most land plants and can control carbon, nitrogen, and phosphorus cycling between above- and belowground components of ecosystems. Current estimates of AM fungal distributions are mainly inferred from the individual distributions of plant biomes, and climatic factors. However, dispersal limitation, local environmental conditions,and interactions among AM fungal taxa may also determine local diversity and global distributions. We assessed the relative importance of these potential controls by collecting 14,961 DNA sequences from 111 published studies and testing for relationships between AM fungal community composition and geography, environment, and plant biomes. Our results indicated that the global species richness of AM fungi was up to six times higher than previously estimated, largely owing to high beta diversity among sampling sites. Geographic distance, soil temperature and moisture, and plant community type were each significantly related to AM fungal community structure, but explained only a small amount of the observed variance. AM fungal species also tended to be phylogenetically clustered within sites, further suggesting that habitat filtering or dispersal limitation is a driver of AM fungal community assembly. Therefore, predicted shifts in climate and plant species distributions under global change may alter AM fungal communities.