Support for the habitat amount hypothesis from a global synthesis of species density studies

Decades of research suggest that species richness depends on spatial characteristics of habitat patches, especially their size and isolation. In contrast, the habitat amount hypothesis predicts that (1) species richness in plots of fixed size (species density) is more strongly and positively related to the amount of habitat around the plot than to patch size or isolation; (2) habitat amount better predicts species density than patch size and isolation combined, (3) there is no effect of habitat fragmentation per se on species density and (4) patch size and isolation effects do not become stronger with declining habitat amount. Data on eight taxonomic groups from 35 studies around the world support these predictions. Conserving species density requires minimising habitat loss, irrespective of the configuration of the patches in which that habitat is contained.     

Richness and composition of plants and birds on land‐bridge islands: effects of island attributes and differential responses of species groups

Aim To test relationships between the richness and composition of vascular plants and birds and attributes of habitat fragments using a model land‐bridge island system, and to investigate whether the effects of fragmentation differ depending on species natural history traits. Location Thousand Island Lake, China. Methods We compiled presence/absence data of vascular plant and bird species through exhaustive surveys of 41 islands. Plant species were assigned to two categories: shade‐intolerant and shade‐tolerant species; bird species were assigned to three categories: edge, interior, and generalist species. We analysed the relationships between island attributes (area, isolation, elevation, shape complexity, and perimeter to area ratio) and species richness using generalized linear models (GLMs). We also investigated patterns of composition in relation to island attributes using ordination (redundancy analysis). Results We found that island area explained a high degree of variation in the species richness of all species groups. The slope of the species–area relationship (z) was 0.16 for all plant species and 0.11 for all bird species. The lowest z‐value was for generalist birds (0.04). The species richness of the three plant species groups was associated with island area per se, while that of all, generalist, and interior birds was explained mainly by elevation, and that of edge bird species was associated primarily with island shape. Patterns of species composition were most strongly related to elevation, island shape complexity, and perimeter to area ratio rather than to island area per se. Species richness had no significant relationship with isolation, but species composition did. We also found differential responses among the species groups to changes in island attributes. Main conclusions Within the Thousand Island Lake system, the effects of fragmentation on both bird and plant species appear to be scale‐dependent and taxon‐specific. The number of plant species occurring on an island is strongly correlated with island area, and the richness of birds and the species composition of plants and birds are associated with variables related to habitat heterogeneity. We conclude that the effects of fragmentation on species diversity and composition depend not only on the degree of habitat loss but also on the specific patterns of habitat fragmentation.     

At the landscape level,birds respond strongly to habitat amount but weakly to fragmentation

Aim

It is usually thought that habitat fragmentation acts negatively on species survival, and consequently, on biodiversity. Recent literature challenges whether habitat fragmentation per se affects species richness, beyond the effect of habitat area. Theoretical studies have suggested that fragmentation may matter most when the amount of available habitat is small or at intermediate levels. However, a recent review suggests that the effect of fragmentation on species richness is usually positive. Here, we dissect the richness–fragmentation relationship. What is the effect size? Does it depend upon the amount of habitat cover? How do individual species respond to fragmentation?

Methods

Applying a macroecological approach, we empirically related avian richness and the probability of occurrence (p_occ) of individual species to fragmentation (number of patches), after controlling for habitat amount in 991 landscapes, each 100‐km², in southern Ontario, Canada.

Results

Species richness was strongly related to total habitat amount, but habitat fragmentation had no detectable additional effect. Individual species’ p_occ related strongly to habitat amount. For some species, p_occ also related secondarily to habitat fragmentation within landscapes. Logistic models revealed that p_occ related significantly negatively to fragmentation after controlling for habitat amount for only ~13% of forest‐ and 18% of open‐habitat species bird species. However, p_occ related significantly positively to fragmentation for even greater proportions of species, including some red‐listed species. Fragmentation effects were not stronger at low or intermediate levels of habitat amount within landscapes.

Conclusion

In earlier studies, negative effects of isolation were observed at the patch level in experimental manipulations. However, at the landscape level, avian species richness in southern Ontario apparently responds primarily to habitat amount and negligibly to fragmentation. We argue that the evidence is inconsistent with the hypothesis that reducing habitat fragmentation per se would be an effective conservation strategy for birds at the landscape level.

     

Habitat fragmentation and species richness

In a recent article in this journal, Fahrig (2013, Journal of Biogeography, 40 , 1649–1663) concludes that variation in species richness among sampling sites can be explained by the amount of habitat in the ‘local landscape’ around the sites, while the spatial configuration of habitat within the landscape makes little difference. This conclusion may be valid for small spatial scales and when the total amount of habitat is large, but modelling and empirical studies demonstrate adverse demographic consequences of fragmentation when there is little habitat across large areas. Fragmentation effects are best tested with studies on individual species rather than on communities, as the latter typically consist of species with dissimilar habitat requirements. The total amount of habitat and the degree of fragmentation tend to be correlated, which poses another challenge for empirical studies. I conclude that fragmentation poses an extra threat to biodiversity, in addition to the threat posed by loss of habitat area.     

A model for the species–area–habitat relationship

Aim To propose a model (the choros model) for species diversity, which embodies number of species, area and habitat diversity and mathematically unifies area per se and habitat hypotheses. Location Species richness patterns from a broad scale of insular biotas, both from island and mainland ecosystems are analysed. Methods Twenty‐two different data sets from seventeen studies were examined in this work. The r² values and the Akaike's Information Criterion (AIC) were used in order to compare the quality of fit of the choros model with the Arrhenius species–area model. The classic method of log‐log transformation was applied. Results In twenty of the twenty‐two cases studied, the proposed model gave a better fit than the classic species–area model. The values of z parameter derived from choros model are generally lower than those derived from the classic species–area equation. Main conclusions The choros model can express the effects of area and habitat diversity on species richness, unifying area per se and the habitat hypothesis, which as many authors have noticed are not mutually exclusive but mutually supplementary. The use of habitat diversity depends on the specific determination of the ‘habitat’ term, which has to be defined based on the natural history of the taxon studied. Although the values of the z parameter are reduced, they maintain their biological significance as described by many authors in the last decades. The proposed model can also be considered as a stepping‐stone in our understanding of the small island effect.     

Extinction,turnover and species diversity in an experimentally fragmented California annual grassland

Summary Many natural populations are subdivided among partially isolated habitat patches, but the influence of habitat patchiness per se on species immigration, extinction, and the resulting patterns of species diversity, has received virtually no experimental study. In an experiment designed to test the effects of habitat subdivision on local community structure, we compare the diversity and annual turnover of flowering plant species in 3 treatments of the same total area, but subdivided to different degrees. We experimentally fragmented a California winter annual grassland into isolated plots, two of 32 m², eight of 8 m², and 32 of 2 m², each treatment representing a combined area of 64 m². Insularization of the experimental habitat fragments is provided by grazing sheep. The effects of plot area on species diversity, extinction, and turnover are consistent with the MacArthur-Wilson model. Species richness increases with the degree of habitat subdivision. Extinction, immigration, and turnover, however, are relatively independent of the degree of subdivision. These experimental results contrast with predictions that habitat subdivision necessarily results in greater rates of extinction accompanied by reduced species diversity.     

Plant diversity in sacred forest fragments of Western Ghats: a comparative study of four life forms

The effect of fragmentation on different life forms within tropical forest plant communities is poorly understood. We studied the effect of degree of fragmentation and surrounding matrix on trees, lianas, shrubs and epiphytes in tropical forest fragments of Kodagu, Western Ghats, India. These fragments exist as sacred groves amidst a highly modified agricultural landscape, and have been preserved by the religious sentiments of local communities. Plants were sampled at two sites in continuous forests and 11 forest fragments. A total of 122 species of trees, 29 species of lianas, 60 species of shrubs and 66 species of epiphytes were recorded. Trees exhibited a significant species–area relationship (R ² = 0.74). Richness estimates after controlling for stem density (rarefaction) revealed that observed species–area relationship was not an artefact of passive sampling. Variation in species richness of the other three groups was explained by stem density and structural diversity. Linear distance from the reserve forest did not explain any variation in species richness. All life forms exhibited significant nested pattern. Trees were nested along the area gradient while nestedness in the other three groups showed evidence in support of habitat nestedness. The four life forms thus responded variably to degree of fragmentation. Our study revealed that 74% of the regional diversity for trees was contributed by diversity among plots, highlighting the importance of inter-patch habitat diversity in maintaining the total regional species pool. We conclude that trees alone cannot serve as good indicator for taking appropriate conservation measures to mitigate species loss resulting from habitat fragmentation.     

Fate of epiphytes on phorophytes with different architectural characteristics along the perturbation gradient of Sabal mexicana forests in Veracruz,Mexico

Question: Vascular epiphytes and hemiepiphytes (E/HE) in neotropical forests account for a large fraction of plant richness, but little is known of how the interplay between phorophyte architectural characteristics and habitat perturbation affect communities of E/HE. Location: Sabal mexicana forests in a coastal area of Veracruz, Mexico. Methods: We compared communities of E/HE on phorophytes with different architectural characteristics – the palm S. mexicana and non‐palm phorophytes – in three environments: conserved sites, perturbed sites and small regenerated forest fragments. We combined traditional (abundance, species richness, similarity and complementarity indices) and more recent (phylogenetic diversity) metrics to describe the communities of E/HE. Results: Overall, we recorded 924 E/HE individuals (nine families, 16 genera and 21 species). The abundance and species richness of E/HE was higher on palms than on non‐palm phorophytes. Abundance‐based complementarities between phorophytes and sites were high. We detected clear changes in community structure of E/HE with habitat perturbation, but there were no effects on the phylogenetic diversity of the E/HE community. Palm phorophytes hosted a more phylogenetically diverse community of E/HE than did non‐palm phorophytes. Conclusions: Palm phorophytes are key elements supporting the conservation of resilient communities of E/HE in S. mexicana forest. Habitat fragmentation has a strong effect on the structure of the E/HE community in S. mexicana forests. Ferns are the group of epiphytes most severely affected by habitat perturbation, but we detected no significant effect on the phylogenetic diversity of the community.     

系统发育多样性与系统发育结构在岛屿植物群落保护中的意义——以蜈支洲岛为例

为探讨系统发育多样性和系统发育结构在岛屿植物群落保护决策制定中的作用,以海南三亚蜈支洲岛被子植物群系为例,分析物种丰富度和系统发育多样性的相关性。结果表明,蜈支洲岛植被可分为10种群系类型,22个样方的物种丰富度与系统发育多样性呈显著正相关,但系统发育多样性与物种丰富度对植物多样性保护优先顺序的指示有差异。森林群系的物种丰富度和系统发育多样性普遍高于灌木和草本群系,4个森林群系样方表现为系统发育结构发散,3个森林群系样方表现为系统发育结构聚集。灌木群系中露兜树(Pandanus tectorius)灌丛和草海桐(Scaevola sericea)灌丛系统发育多样性较高,草本群系中厚藤(Vitex trifolia)灌草丛系统发育多样性较高,灌木和草本群系大多倾向于系统发育结构聚集。距海或人类活动区较近的群系可能受环境过滤的作用,而远离海和人类活动区的群系主要受竞争排斥作用影响。因此,在岛屿植物多样性保护策略制定中应当综合考虑物种和系统发育维度,以及环境、演替阶段及用岛方式等因素的影响。     

The area-independent effects of habitat complexity on biodiversity vary between regions

Potential explanatory variables often co‐vary in studies of species richness. Where topography varies within a survey it is difficult to separate area and habitat‐diversity effects. Topographically complex surfaces may contain more species due to increased habitat diversity or as a result of increased area per se. Fractal geometry can be used to adjust species richness estimates to control for increases in area on complex surfaces. Application of fractal techniques to a survey of rocky shores demonstrated an unambiguous area‐independent effect of topography on species richness in the Isle of Man. In contrast, variation in species richness in south‐west England reflected surface availability alone. Multivariate tests and variation in limpet abundances also demonstrated regional variation in the area‐independent effects of topography. Community composition did not vary with increasing surface complexity in south‐west England. These results suggest large‐scale gradients in the effects of heterogeneity on community processes or demography.     

Independent effects of habitat loss,habitat fragmentation and structural connectivity on forest‐dependent birds

Aim Habitat loss and fragmentation are amongst the greatest threats to biodiversity world‐wide. However, there is still little evidence on the relative influence of these two distinct processes on biodiversity, and no study, to date, has investigated the independent contribution of structural connectivity in addition to habitat loss and fragmentation. The aim of this study is to evaluate the independent effects of habitat loss (the decrease in total amount of habitat), habitat fragmentation per se (habitat subdivision) and structural connectivity (in the form of hedgerow networks) on the distribution of seven resident forest‐dependent birds in central Italy. Location Central Italy. Methods We strategically selected 30 landscapes (each of 16 km² in size) with decreasing total amount of forest cover and with contrasting configuration of patches and contrasting lengths of hedgerow networks. Presence/absence of birds in each landscape unit was studied through point counts. Results The amount of forest cover in the landscape had the strongest relative influence on birds’ occupancy, whilst habitat subdivision played a negligible role. Structural connectivity and the geographic position of the landscape unit played a relatively important role for four species. Main conclusions Our study shows the importance of disentangling the contribution of different landscape properties in determining distribution patterns. Our results are consistent with the fact that halting habitat loss and carrying out habitat restoration should be conservation priorities, since habitat loss is the main factor affecting the distribution of the target species; implementation of structural connectivity through hedgerows, instead, should be evaluated with caution since its contribution is secondary to the predominant role of habitat loss.     

Island biogeography of bats in Baja California, Mexico: patterns of bat species richness in a near-shore archipelago

Aim We studied the relationship between the size and isolation of islands and bat species richness in a near‐shore archipelago to determine whether communities of vagile mammals conform to predictions of island biogeography theory. We compared patterns of species richness in two subarchipelagos to determine whether area per se or differences in habitat diversity explain variations in bat species richness. Location Islands in the Gulf of California and adjacent coastal habitats on the Baja California peninsula in northwest Mexico. Methods Presence–absence surveys for bats were conducted on 32 islands in the Gulf of California using acoustic and mist‐net surveys. We sampled for bats in coastal habitats of four regions of the Baja peninsula to characterize the source pool of potential colonizing species. We fitted a semi‐log model of species richness and multiple linear regression and used Akaike information criterion model selection to assess the possible influence of log₁₀ area, isolation, and island group (two subarchipelagos) on the species richness of bats. We compared the species richness of bats on islands with greater vegetation densities in the southern gulf (n = 20) with that on drier islands with less vegetation in the northern gulf (n = 12) to investigate the relationship between habitat diversity and the species richness of bats. Results Twelve species of bats were detected on islands in the Gulf of California, and 15 species were detected in coastal habitats on the Baja peninsula. Bat species richness was related to both area and isolation of islands, and was higher in the southern subarchipelago, which has denser vegetation. Log₁₀ area was positively related to bat species richness, which increased by one species for every 5.4‐fold increase in island area. On average, richness declined by one species per 6.25 km increase in isolation from the Baja peninsula. Main conclusions Our results demonstrate that patterns of bat species richness in a near‐shore archipelago are consistent with patterns predicted by the equilibrium theory of island biogeography. Despite their vagility, bats may be more sensitive to moderate levels of isolation than previously expected in near‐shore archipelagos. Differences in vegetation and habitat xericity appear to be associated with richness of bat communities in this desert ecosystem. Although observed patterns of species richness were consistent with those predicted by the equilibrium theory, similar relationships between species richness and size and isolation of islands may arise from patch‐use decision making by individuals (optimal foraging strategies).     

Landscape geometry determines community response to disturbance

Ecological communities are impacted by anthropogenic changes in both habitat geometry (i.e. amount, shape, fragmentation and connectivity of habitat) and disturbance regime. Although the effect of each of these drivers on diversity is well-documented, few studies have considered how habitat geometry and disturbance interact to affect diversity. We used a miniature landscape of moss patches to experimentally manipulate both habitat geometry and disturbance frequency on microarthropod communities. Species richness and abundance in local patches declined linearly with disturbance rate in all experimental landscapes, but the speed of this decline (a measure of ecological resilience) depended on the size and connectivity of the surrounding region. Reductions in region size had little effect on community resilience to disturbance until habitat loss resulted in complete loss of connectivity between patches, suggesting a threshold in community response to habitat loss. Beyond this threshold, repeated disturbance resulted in rapid declines in patch species richness and abundance and substantial changes in community composition. These effects of habitat geometry and disturbance on diversity were scale-dependent. Gamma (regional-scale) diversity was unaffected by habitat geometry, suggesting experimental reductions in alpha (local-scale) diversity were offset by increases in beta diversity. There was no effect of body size, abundance, or trophic position in determining species response to disturbance. Taxonomic grouping had a weak effect, with oribatids less affected by drought. We conclude that, in this system, dispersal from the surrounding metacommunity is vital in allowing recovery of local communities from disturbance. When habitat loss and fragmentation disrupt this process, extinctions result. Studies that examine separately the effects of habitat alterations and disturbance on diversity may therefore underestimate their combined effects.     

Local native plant diversity responds to habitat loss and fragmentation over different time spans and spatial scales

Separating the threats that habitat loss and habitat fragmentation pose to biodiversity is challenging because these processes usually occur simultaneously. Additionally, their importance may be underestimated due to time-delayed extinction. In central Texas savannas, woody plant encroachment reduces the amount of habitat available to herbaceous species while fragmenting remaining habitat. We examined the relationships between present species richness and present and past habitat amount and fragmentation (measured as fractal dimension) using a series of aerial photographs taken over nearly 60 years. We show that woody plant encroachment, a common phenomenon in savannas worldwide, reduces the diversity of herbaceous vegetation through both habitat loss and fragmentation. Habitat loss has the strongest impact on species richness over short time spans and small spatial scales in these savannas. Habitat fragmentation, however, has the strongest impact over longer time spans and larger spatial scales and generates long-term extinction debts. We also demonstrate that examining habitat loss and habitat fragmentation across different time periods and at different spatial scales is essential for understanding their joint and individual effects on plant community composition.     

Effects of area,environmental status and environmental variation on species richness per unit area in Mediterranean wetlands

Abstract. We propose a mechanistic model to relate α- and γ-diversity to area per se, moisture status and environmental variation (local and total), and explored the effects these abiotic variables have on species richness per unit area (α-diversity) for plant communities in a network of wetland habitats located in a Mediterranean mountainous region of central Spain. In this study, environmental status is measured as actual evapotranspiration (as an expression of energy), slope and soil wetness, and environmental variation refers to slope variation and soil wetness variation. Species richness per unit area was related to soil wetness, soil wetness variation, ground slope and ground slope variation. There were also positive correlations among moisture status and environmental variation variables. There is a joint effect of slope and soil wetness variation in explaining species richness per unit area of these wetland habitats, but area effects and energy are relatively unimportant. We conclude that species richness per unit area of wetland vegetation can be explained by moisture status and local environmental variation, and that habitat area may not have an important effect. Area could affect γ-diversity directly through random sampling and/or indirectly through increasing β-diversity, and energy may be important in areas with larger energy ranges. Complete surveys of environmental status, local and total environmental variation, and their associated species assemblages are needed to explain the processes that give rise to the rule that larger areas have larger species richness.     

Temporal shifts and niche overlapping in <Emphasis Type="Italic">Copestylum</Emphasis> (Diptera,Syrphidae) communities reared in cactus species in a central Mexican scrubland

Semiarid scrubland communities are highly dynamic in terms of their species composition, abundance, and functioning, given the drastic changes in climate among seasons. Spatiotemporal patterns of saprophagous Copestylum (Diptera: Syrphidae) communities in different cactus species richness have not yet been studied, although seasonal changes and plant species richness have been shown to strongly impact the diversity and distribution of many insect communities in scrublands. We analyzed the impact of seasonality and of habitat type (disturbed and undisturbed) on Copestylum communities reared from cactus species at the Barranca de Metztitlán Biosphere Reserve, in central Mexico, by comparing their community structure between seasons and habitats, and assessing the contribution of diversity components for the total diversity of this genus. We also measured patterns of temporal niche overlap among hoverfly species considering their breeding medium. Seasonal variation influenced Copestylum community composition most significantly. Species richness and abundance of Copestylum were higher in the rainy season. Additive partitioning of diversity showed that the main component for species richness is beta diversity between seasons. We detected high niche overlap during the dry season and low overlap during the rainy season. This study provides evidence of temporal shifts in xeric hoverfly communities and suggests that the Copestylum species partition resources over time.     

Direct versus indirect effects of habitat fragmentation on community patterns in experimental landscapes

Habitat area and fragmentation are confounded in many ecological studies investigating fragmentation effects. We thus devised an innovative experiment founded on fractal neutral landscape models to disentangle the relative effects of habitat area and fragmentation on arthropod community patterns in red clover (Trifolium pratense). The conventional approach in experimental fragmentation studies is to adjust patch size and isolation to create different landscape patterns. We instead use fractal distributions to adjust the overall amount and fragmentation of habitat independently at the scale of the entire landscape, producing different patch properties. Although habitat area ultimately had a greater effect on arthropod abundance and diversity in this system, we found that fragmentation had a significant effect in clover landscapes with ≤40 % habitat. Landscapes at these lower habitat levels were dominated by edge cells, which had fewer arthropods and lower richness than interior cells. Fragmentation per se did not have a direct effect on local-scale diversity, however, as demonstrated by the lack of a broader landscape effect (in terms of total habitat area and fragmentation) on arthropods within habitat cells. Fragmentation—through the creation of edge habitat—thus had a strong indirect effect on morphospecies richness and abundance at the local scale. Although it has been suggested that fragmentation should be important at low habitat levels (≤20–30 %), we show that fragmentation per se is significant only at intermediate (40 %) levels of habitat, where edge effects were neither too great (as at lower levels of habitat) nor too weak (as at higher levels of habitat).     

Pervasive effects of dispersal limitation on within- and among-community species richness in agricultural landscapes

Aim To determine whether the effect of habitat fragmentation and habitat heterogeneity on species richness at different spatial scales depends on the dispersal ability of the species assemblages and if this results in nested species assemblages. Location Agricultural landscapes distributed over seven temperate Europe countries covering a range from France to Estonia. Methods We sampled 16 local communities in each of 24 agricultural landscapes (16 km²) that differ in the amount and heterogeneity of semi‐natural habitat patches. Carabid beetles were used as model organisms as dispersal ability can easily be assessed on morphological traits. The proximity and heterogeneity of semi‐natural patches within the landscape were related to average local (alpha), between local (beta) and landscape (gamma) species richness and compared among four guilds that differ in dispersal ability. Results For species assemblages with low dispersal ability, local diversity increased as the proximity of semi‐natural habitat increased, while mobile species showed an opposite trend. Beta diversity decreased equally for all dispersal classes in relation to proximity, suggesting a homogenizing effect of increased patch isolation. In contrast, habitat diversity of the semi‐natural patches affected beta diversity positively only for less mobile species, probably due to the low dispersal ability of specialist species. Species with low mobility that persisted in highly fragmented landscapes were consistently present in less fragmented ones, resulting in nested assemblages for this mobility class only. Main conclusions The incorporation of dispersal ability reveals that only local species assemblages with low dispersal ability show a decrease of richness as a result of fragmentation. This local species loss is compensated at least in part by an increase in species with high dispersal ability, which obscures the effect of fragmentation when investigated across dispersal groups. Conversely, fragmentation homogenizes the landscape fauna for all dispersal groups, which indicates the invasion of non‐crop habitats by similar good dispersers across the whole landscape. Given that recolonization of low dispersers is unlikely, depletion of these species in modern agricultural landscapes appears temporally pervasive.     

Competitive displacement or biotic resistance? Disentangling relationships between community diversity and invasion success of tall herbs and shrubs

Questions: Are negative invasion–diversity relationships due to biotic resistance of the invaded plant community or to post‐invasion displacement of less competitive species? Do invasion–diversity relationships change with habitat type or resident traits? Location/species: Lowlands and uplands of western and southern Germany, Heracleum mantegazzianum; mountain range in central Germany, Lupinus polyphyllus; and coastal dunes of northwest Germany, Rosa rugosa. Methods: We tested the significance and estimated regression slopes of invasion–diversity relationships using generalized linear (mixed effects) models relating invader cover and habitat type to species richness in different plant groups, stratified based on size, life cycle and community association. Results: We found negative, positive and neutral relationships between invader cover and species richness. There were negative linear correlations of invader cover with small plant species throughout, but no negative linear correlation with tall species. Invasion–diversity relationships tended to be more negative in early‐successional habitats, such as dunes or abandoned grasslands, than in late‐successional habitats. Conclusions: Invasion diversity–relationships are complex; they vary among habitat types and among different groups of resident species. Negative invasion–diversity relationships are due to asymmetric competitive displacement of inferior species and not due to biotic resistance. Small species are displaced in early‐successional habitats, while there is little effect on persistence of tall species.     

Effects of habitat fragmentation and black-tailed prairie dogs on urban avian diversity

Urbanization and habitat fragmentation have the potential to influence bird communities. In addition, these phenomena, as well as ongoing lethal control measures, have also greatly reduced the range of the black-tailed prairie dog (Cynomys ludovicianus) since the beginning of the 20th century. Although prairie dogs are highly interactive species that can influence avian communities, few studies have investigated whether these interactions persist in urban settings. Our goal was to investigate the relative impacts of habitat fragmentation and prairie dogs on bird communities within an urban matrix. We performed bird surveys on 20 habitat fragments (10 colonized by prairie dogs, 10 uncolonized by prairie dogs) distributed throughout the Denver metropolitan area, and calculated Shannon–Weiner diversity and richness of all birds and native species, as well as total counts of grassland birds and raptors. Diversity, richness, and counts of many species increased with increasing fragment connectivity, and decreased on fragments isolated for longer periods of time. Avian diversity and richness did not differ between fragments with and without prairie dogs, suggesting that this element of the ecological role of prairie dogs is not fully retained in urban habitat. Future studies of the role of prairie dogs as keystone species in urban systems should include other taxa as well as consider the influence of the urban matrix surrounding prairie dog habitat. Our results emphasize that conservation of urban avian diversity should focus on landscape connectivity as well as local habitat features.