Landscape evaluation in conservation: molecular sampling and habitat modeling for the Iberian lynx.

Conservation of endangered species requires comprehensive understanding of their distribution and habitat requirements, in order to implement better management strategies. Unfortunately, this understanding is often difficult to gather at the short term required by rapidly declining populations of many rare vertebrates. We present a spatial habitat modeling approach that integrates a molecular technique for species detection with landscape information to assess habitat requirements of a critically endangered mammalian carnivore, the Iberian lynx (Lynx pardinus), in a poorly known population in Spain. We formulated a set of model hypotheses for habitat selection at the spatial scale of home ranges, based on previous information on lynx requirements of space, vegetation, and prey. To obtain the required data for model selection, we designed a sampling protocol based on surveys of feces and their molecular analysis for species identification. After comparing candidate models, we selected a parsimonious one that allowed (1) reliable assessment of lynx habitat requirements at the scale of home ranges, (2) prediction of lynx distribution and potential population size, and (3) identification of landscape management priorities for habitat conservation. This model predicted that the species was more likely to occur in landscapes with a higher percentage of rocky areas and higher cover of bushes typical of mature mediterranean shrubland mosaics. Its accuracy for discriminating lynx presence was approximately 85%, indicating high predictive performance. Mapping model predictions showed that only 16% of the studied areas constitute potential habitat for lynx, even though the region is dominated by large extents of well-preserved native vegetation with low human interference. Habitat was mostly clumped in two nearby patches connected by vegetation adequate for lynx dispersal and had a capacity for 28-62 potential breeding territories. The lynx population in Sierra Morena is probably the largest persisting today, but it is still critically small for optimism about its long-term persistence. Model results suggest habitat conservation and restoration actions needed for preserving the species, including reconciliation of hunting management with preservation of mature shrubland over large areas (particularly in rocky landscapes). The approach presented here can be applied to many other species for which the ecological information needed to develop sound habitat conservation strategies is lacking.     

Managing Boreal Forest Landscapes for Flying Squirrels

Abstract: Flying squirrel (Pteromys volans) populations have declined severely during the past few decades, and the species has become a focal species in forest management and the conservation debate in Finland. We compared landscape structure around known flying squirrel home ranges with randomly chosen forest sites to determine which landscape patterns characterize the areas occupied by the species in northern Finland. We sought to identify the key characteristics of the landscape that support the remaining flying squirrel populations. We analyzed landscape structure within circular areas with 1- and 3-km radii around 63 forest sites occupied by flying squirrels, and around 96 random sites. We applied stepwise analysis of the landscape structure where landscapes were built up step-by-step by adding patch types in order of their suitability for the flying squirrel. The land-use and forest-resource data for the analysis were derived from multisource national forest inventory and imported to a geographical information system. Landscape patch types were divided into three suitability categories: breeding habitat (mixed spruce-deciduous forests); dispersal habitat ( pine and young forests); and unsuitable habitat ( young sapling stands, open habitats, water). Flying squirrel landscapes contained more suitable breeding habitat patches and were better connected by dispersal habitats than random landscapes. Our results suggest that for the persistence of the flying squirrel, forest managers should 1) maintain a deciduous mixture, particularly in spruce-dominated forests; 2) maintain physical connectivity between optimal breeding habitats; and 3) impose coarse-grained structures on northeastern Finnish landscapes at current levels of habitat availability.     

Differences in gypsum plant communities associated with habitat fragmentation and livestock grazing.

The negative consequences of habitat fragmentation for plant communities have been documented in many regions of the world. In some fragmented habitats, livestock grazing has been proposed to be a dispersal mechanism reducing isolation between fragments. In others, grazing acts together with fragmentation in a way that increases habitat degradation. Iberian gypsum plant communities have been grazed and fragmented by agricultural practices for centuries. Although their conservation is considered a priority by the European Community, the effects of fragmentation on gypsum plant communities and the possible role of livestock grazing remain unknown. In addition, a substantial proportion of plant species growing in gypsum environments are gypsum specialists. They could be particularly affected by fragmentation, as was found for other habitat specialists (i.e., serpentine and calcareous specialists). In this study (1) we investigated the effect of fragmentation and grazing on gypsum plant community composition (species and life-forms), and (2) we tested to see if gypsum specialists were differently affected by fragmentation and grazing than habitat generalists. A vegetation survey was conducted in the largest gypsum outcrop of Europe (Middle Ebro Valley, northeast Spain). Fragmented and continuous sites in grazed and ungrazed areas were compared. Measurements related to species and composition of life-forms were contrasted first for the whole gypsum plant community and then specifically for the gypsum specialists. In the whole community, our results showed lower plant species diversity in fragmented sites, mainly due to the larger dominance of species more tolerant to fragmented habitat conditions. With livestock grazing, the plant species richness and the similarity in plant species composition between remnants was larger, suggesting that animals were acting as dispersal agents between fragments. As expected, gypsum specialists were less abundant in fragmented areas, and grazing led to the disappearance of the rare gypsum specialist Campanula fastigiata. According to our results, conservation strategies for gypsum plant communities in human-dominated landscapes should consider that fragmentation and grazing modify plant community composition affecting gypsum specialists in particular.     

Effects of Artificial Roosts for Frugivorous Bats on Seed Dispersal in a Neotropical Forest Pasture Mosaic

Abstract:  In the Neotropics ongoing deforestation is producing open and heavily fragmented landscapes dominated by agriculture, mostly plantations and cattle pastures. After some time agriculture often becomes uneconomical and land is abandoned. Subsequent habitat regeneration may be slow because seed inputs are restricted by a lack of incentives—such as suitable roost sites—for seed dispersers to enter deforested areas. Increasing environmental awareness has fostered growing efforts to promote reforestation. Practical and cost-efficient methods for kick-starting forest regeneration are, however, lacking. We investigated whether artificial bat roosts for frugivorous bat species can attract these key seed dispersers to deforested areas, thereby increasing seed rain. We installed artificial bat roosts in a forest-pasture mosaic in the Costa Rican Atlantic lowlands and monitored bat colonization and seed dispersal. Colonization occurred within a few weeks of installation, and 10 species of bats occupied the artificial roosts. Five species of frugivorous or nectarivorous bats colonized artificial roosts permanently in both primary habitat and in deforested areas, in numbers similar to those found in natural roosts. Seed input around artificial roosts increased significantly. Sixty-nine different seed types, mostly of early-successional plant species, were transported by bats to artificial roosts in disturbed habitats. The installation of artificial bat roosts thus successfully attracted frugivorous bats and increased seed inputs into degraded sites. This method is likely to speed up early-vegetation succession, which in turn will attract additional seed dispersers, such as birds, and provide a microhabitat for seeds of mid- and late-successional plants. As well as supporting natural forest regeneration and bat conservation, this cost-efficient method can also increase environmental awareness among landowners.     

Defining and Evaluating the Umbrella Species Concept for Conserving and Restoring Landscape Connectivity

Conserving or restoring landscape connectivity between patches of breeding habitat is a common strategy to protect threatened species from habitat fragmentation. By managing connectivity for some species, usually charismatic vertebrates, it is often assumed that these species will serve as conservation umbrellas for other species. We tested this assumption by developing a quantitative method to measure overlap in dispersal habitat of 3 threatened species—a bird (the umbrella), a butterfly, and a frog—inhabiting the same fragmented landscape. Dispersal habitat was determined with Circuitscape, which was parameterized with movement data collected for each species. Despite differences in natural history and breeding habitat, we found substantial overlap in the spatial distributions of areas important for dispersal of this suite of taxa. However, the intuitive umbrella species (the bird) did not have the highest overlap with other species in terms of the areas that supported connectivity. Nevertheless, we contend that when there are no irreconcilable differences between the dispersal habitats of species that cohabitate on the landscape, managing for umbrella species can help conserve or restore connectivity simultaneously for multiple threatened species with different habitat requirements. Definición y Evaluación del Concepto de Especie Paraguas para Conservar y Restaurar la Conectividad de Paisajes     

Managing Aquatic Species of Conservation Concern in the Face of Climate Change and Invasive Species

Abstract:  The difficult task of managing species of conservation concern is likely to become even more challenging due to the interaction of climate change and invasive species. In addition to direct effects on habitat quality, climate change will foster the expansion of invasive species into new areas and magnify the effects of invasive species already present by altering competitive dominance, increasing predation rates, and enhancing the virulence of diseases. In some cases parapatric species may expand into new habitats and have detrimental effects that are similar to those of invading non-native species. The traditional strategy of isolating imperiled species in reserves may not be adequate if habitat conditions change beyond historic ranges or in ways that favor invasive species. The consequences of climate change will require a more active management paradigm that includes implementing habitat improvements that reduce the effects of climate change and creating migration barriers that prevent an influx of invasive species. Other management actions that should be considered include providing dispersal corridors that allow species to track environmental changes, translocating species to newly suitable habitats where migration is not possible, and developing action plans for the early detection and eradication of new invasive species.     

An Experimental Test of Matrix Permeability and Corridor Use by an Endemic Understory Bird

Abstract:  Because of widespread habitat fragmentation, maintenance of landscape connectivity has become a major focus of conservation planning, but empirical tests of animal movement in fragmented landscapes remain scarce. We conducted a translocation experiment to test the relative permeability of three landscape elements (open habitat, shrubby secondary vegetation, and wooded corridors) to movement by the Chucao Tapaculo ( Scelorchilus rubecula ), a forest understory bird endemic to South American temperate rainforest. Forty-one radio-tagged subjects were translocated (individually) to three landscape treatments consisting of small release patches that were either entirely surrounded by open habitat (pasture), entirely surrounded by dense shrubs, or linked to other patches by wooded corridors that were otherwise surrounded by open matrix. The number of days subjects remained in release patches before dispersal (a measure of habitat resistance) was significantly longer for patches surrounded by open habitat than for patches adjoining corridors or surrounded by dense shrubs. These results indicate that open habitat significantly constrains Chucao dispersal, in accord with expectation, but dispersal occurs equally well through wooded corridors and shrub-dominated matrix. Thus, corridor protection or restoration and management of vegetation in the matrix (to encourage animal movement) may be equally feasible alternatives for maintaining connectivity.     

Conservation planning on China's borders with Myanmar,Laos, and Vietnam

Transboundary conservation is playing an increasingly important role in maintaining ecosystem integrity and halting biodiversity loss caused by anthropogenic activities. However, lack of information on species distributions in transboundary regions and understanding of the threats in these areas impairs conservation. We developed a spatial conservation plan for the transboundary areas between Yunnan province, southwestern China, and neighboring Myanmar, Laos, and Vietnam in the Indo-Burma biodiversity hotspot. To identify priority areas for conservation and restoration, we determined species distribution patterns and recent land-use changes and examined the spatiotemporal dynamics of the connected natural forest, which supports most species. We assessed connectivity with equivalent connected area (ECA), which is the amount of reachable habitat for a species. An ECA incorporates the presence of habitat in a patch and the amount of habitat in other patches within dispersal distance. We analyzed 197,845 locality records from specimen collections and monographs for 21,004 plant and vertebrate species. The region of Yunnan immediately adjacent to the international borders had the highest species richness, with 61% of recorded species and 56% of threatened vertebrates, which suggests high conservation value. Satellite imagery showed the area of natural forest in the border zone declined by 5.2% (13,255 km²) from 1995 to 2018 and monoculture plantations increased 92.4%, shrubland 10.1%, and other cropland 6.2%. The resulting decline in connected natural forest reduced the amount of habitat, especially for forest specialists with limited dispersal abilities. The most severe decline in connectivity was along the Sino-Vietnamese border. Many priority areas straddle international boundaries, indicating demand and potential for establishing transboundary protected areas. Our results illustrate the importance of bi- and multilateral cooperation to protect biodiversity in this region and provide guidance for future conservation planning and practice.     

Effects of Future Infrastructure Development on Threat Status and Occurrence of Amazonian Birds

Abstract:  Researchers predict that new infrastructure development will sharply increase the rate and extent of deforestation in the Brazilian Amazon. There are no predictions, however, of which species it will affect. We used a spatially explicit model that predicts the location of deforestation in the Brazilian Amazon by 2020 on the basis of historical patterns of deforestation following infrastructure development. We overlaid the predicted deforested areas onto maps of bird ranges to estimate the amount of habitat loss within species ranges. We also estimated the amount of habitat loss within modified ecoregions, which were used as surrogates for areas of bird endemism. We then used the extent of occurrence criterion of the World Conservation Union to predict the future conservation status of birds in the Brazilian Amazon. At current rates of development, our results show that at least 16 species will qualify as threatened or will lose more than half of their forested habitat. We also identified several subspecies and isolated populations that would also qualify as threatened. Most of the taxa we identified are not currently listed as threatened, and the majority are associated with riverine habitats, which have been largely ignored in bird conservation in Amazonia. These habitats and the species they hold will be increasingly relevant to conservation as river courses are altered and hydroelectric dams are constructed in the Brazilian Amazon.     

The Near Extinction of Two Large European Predators: Super Specialists Pay a Price

Abstract:  Despite recovery plans, the Spanish Imperial Eagle ( Aquila adalberti ) and the Iberian lynx ( Lynx pardinus ) are in danger of extinction. These two flagship species tend to occur in pristine Mediterranean forests, and both prey preferentially on the rabbit ( Oryctologus cuniculus ). Spanish lynxes and eagles have sister species in continental Europe, the Eurasian lynx (  Lynx europaeus ) and the Eastern Imperial Eagle ( A. heliaca ), respectively. Recent genetic evidence indicates that these two pairs of species started to diverge from their ancestor species slightly less than 1 million years ago, when the longest-lasting Pleistocene glaciations covered Europe. We hypothesize that the Iberian lynx and the Spanish Imperial Eagle emerged as separate species in the Pleistocene refugia of southern Spain, where they hunted yet another locally evolved species, the rabbit, on which they have become dependent for survival. Two large predators that emerged at the same time may go extinct simultaneously because of their inability to shift to alternative prey. Many other relict species, including numerous species from oceanic islands, have naturally small populations because of evolutionary constraints and are permanently threatened with extinction. Recovery plans aimed at putting these species out of danger are unrealistic, as their populations are and have been chronically scarce. We suggest that what these species need are maintenance plans designed to buffer population declines due to either stochastic or human-induced events. A metaphor for this would be "emergency care units" for conservation.     

Desert-dwelling Mountain Sheep: Conservation Implications of a Naturally Fragmented Distribution

Abstract: Mountain sheep (Ovis canadensis) are closely associated with steep, mountainous, open terrain. Their habitat consequently occurs in a naturally fragmented pattern, often with substantial expanses of unsuitable habitat between suitable patches; the sheep have been noted to be slow colonizers of vacant suitable habitat. As a result, resource managers have focused on (1) conserving "traditional" mountainous habitats, and (2) forced colonization through reintroduction. Telemetry studies in desert habitats have recorded more intermountain movement by desert sheep than was previously thought to OCCUT. Given the heretofore unrecognized vagility of mountain sheep, we argue that existing corridors of "nontraditional" habitat connecting mountain ranges be given adequate conservation consideration. Additionally, small areas of mountainous habitat that an? not permanently occupied but that may serve as "stepping stones" within such corridors must be recognized for their potential importance to relatively isolated populations of mountain sheep. We discuss the potential importance of such corridors to other large, vagile species.     

Cougar Dispersal Patterns, Metapopulation Dynamics, and Conservation

Abstract: We examined cougar (   Puma concolor ) dispersal, emigration, and immigration in the San Andres Mountains, New Mexico, from 1985 to 1995 to quantify the effects of dispersal on the local population and surrounding subpopulations. We captured, tagged, and radio-collared animals to detect the arrival of new immigrants and dispersal characteristics of progeny. We found that cougars in southern New Mexico exhibited a metapopulation structure in which cougar subpopulations were separated by expanses of noncougar habitat and linked by dispersers. Of 43 progeny (n = 20 males , 23 females ) studied after independence, only 13 females exhibited philopatric behavior. Males dispersed significantly farther than females, were more likely to traverse large expanses of noncougar habitat, and were probably most responsible for nuclear gene flow between habitat patches. We estimated that an average of 8.5 progeny (i.e., cougars born in the study area) successfully emigrated from and 4.3 cougars successfully immigrated to the San Andres Mountains each year. Concurrently, an average of 4.1 progeny were recruited into the San Andres cougar population. Protected cougar subpopulations can contribute to metapopulation persistence by supplying immigrants to surrounding subpopulations that are affected by fragmentation or offtake by humans. Cougar population dynamics and dispersal behavior dictate that cougar management and conservation should be considered on a regional scale.     

The Value of Primary, Secondary, and Plantation Forests for a Neotropical Herpetofauna

Abstract:  Plantation forests and second-growth forests are becoming dominant components of many tropical forest landscapes. Yet there is little information available concerning the consequences of different forestry options for biodiversity conservation in the tropics. We sampled the leaf-litter herpetofauna of primary, secondary, and Eucalyptus plantation forests in the Jari River area of northeastern Brazilian Amazonia. We used four complementary sampling techniques, combined samples from 2 consecutive years, and collected 1739 leaf-litter amphibians (23 species) and 1937 lizards (30 species). We analyzed the data for differences among forest types regarding patterns of alpha and beta diversity, species-abundance distributions, and community structure. Primary rainforest harbored significantly more species, but supported a similar abundance of amphibians and lizards compared with adjacent areas of second-growth forest or plantations. Plantation forests were dominated by wide-ranging habitat generalists. Secondary forest faunas contained a number of species characteristic of primary forest habitat. Amphibian communities in secondary forests and Eucalyptus plantations formed a nested subset of primary forest species, whereas the species composition of the lizard community in plantations was distinct, and was dominated by open-area species. Although plantation forests are relatively impoverished, naturally regenerating forests can help mitigate some negative effects of deforestation for herpetofauna. Nevertheless, secondary forest does not provide a substitute for primary forest, and in the absence of further evidence from older successional stands, we caution against the optimistic claim that natural forest regeneration in abandoned lands will provide refuge for the many species that are currently threatened by deforestation .     

Combining Measures of Dispersal to Identify Conservation Strategies in Fragmented Landscapes

Abstract: Understanding the way in which habitat fragmentation disrupts animal dispersal is key to identifying effective and efficient conservation strategies. To differentiate the potential effectiveness of 2 frequently used strategies for increasing the connectivity of populations in fragmented landscapes—corridors and stepping stones—we combined 3 complimentary methods: behavioral studies at habitat edges, mark‐recapture, and genetic analyses. Each of these methods addresses different steps in the dispersal process that a single intensive study could not address. We applied the 3 methods to the case study of Atrytonopsis new species 1, a rare butterfly endemic to a partially urbanized stretch of barrier islands in North Carolina (U.S.A.). Results of behavioral analyses showed the butterfly flew into urban and forested areas, but not over open beach; mark‐recapture showed that the butterfly dispersed successfully through short stretches of urban areas (<500 m); and genetic studies showed that longer stretches of forest (>5 km) were a dispersal barrier, but shorter stretches of urban areas (≤5 km) were not. Although results from all 3 methods indicated natural features in the landscape, not urbanization, were barriers to dispersal, when we combined the results we could determine where barriers might arise: forests restricted dispersal for the butterfly only when there were long stretches with no habitat. Therefore, urban areas have the potential to become a dispersal barrier if their extent increases, a finding that may have gone unnoticed if we had used a single approach. Protection of stepping stones should be sufficient to maintain connectivity for Atrytonopsis new species 1 at current levels of urbanization. Our research highlights how the use of complementary approaches for studying animal dispersal in fragmented landscapes can help identify conservation strategies.     

Assessment of the Conservation Efforts to Prevent Extinction of the Iberian Lynx

Abstract: The Iberian lynx (Lynx pardinus) may be the first charismatic felid to become extinct in a high‐income country, despite decades of study and much data that show extinction is highly probable. The International Union for Conservation of Nature categorizes it as critically endangered; about 200 free‐ranging individuals remain in two populations in southern Spain. Conservation measures aimed at averting extirpation have been extensively undertaken with 4 of the former 10 Iberian lynx populations recorded 25 years ago. Two of the four populations have been extirpated. The number of individuals in the third population have declined by 83%, and in the fourth the probability of extirpation has increased from 34% to 95%. Major drivers of the pending extinction are the small areas to which conservation measures have been applied; lack of incorporation of evidence‐based conservation, scientific monitoring, and adaptive management into conservation efforts; a lack of continuity in recovery efforts, and distrust by conservation agencies of scientific information. In contrast to situations in which conservation and economic objectives conflict, in the case of the Iberian lynx all stakeholders desire the species to be conserved.     

Birds as Suppliers of Seed Dispersal in Temperate Ecosystems: Conservation Guidelines from Real‐World Landscapes

Abstract: Seed dispersal by animals is considered a pivotal ecosystem function that drives plant‐community dynamics in natural habitats and vegetation recovery in human‐altered landscapes. Nevertheless, there is a lack of suitable ecological knowledge to develop basic conservation and management guidelines for this ecosystem service. Essential questions, such as how well the abundance of frugivorous animals predicts seeding function in different ecosystems and how anthropogenic landscape heterogeneity conditions the role of dispersers, remain poorly answered. In three temperate ecosystems, we studied seed dispersal by frugivorous birds in landscape mosaics shaped by human disturbance. By applying a standardized design across systems, we related the frequency of occurrence of bird‐dispersed seeds throughout the landscape to the abundance of birds, the habitat features, and the abundance of fleshy fruits. Abundance of frugivorous birds in itself predicted the occurrence of dispersed seeds throughout the landscape in all ecosystems studied. Even those landscape patches impoverished due to anthropogenic disturbance received some dispersed seeds when visited intensively by birds. Nonetheless, human‐caused landscape degradation largely affected seed‐deposition patterns by decreasing cover of woody vegetation or availability of fruit resources that attracted birds and promoted seed dispersal. The relative role of woody cover and fruit availability in seed dispersal by birds differed among ecosystems. Our results suggest that to manage seed dispersal for temperate ecosystem preservation or restoration one should consider abundance of frugivorous birds as a surrogate of landscape‐scale seed dispersal and an indicator of patch quality for the dispersal function; woody cover and fruit resource availability as key landscape features that drive seedfall patterns; and birds as mobile links that connect landscape patches of different degrees of degradation and habitat quality via seed deposition.     

Connected fragmented habitats facilitate stable coexistence dynamics

In this paper we endeavor to test the controversial ideas that exist about the role of fragmentation in a conservation context. In line with earlier understanding, we find that habitat fragmentation alone results in a strong detrimental effect (especially for the predator population). Connecting the fragmented habitats facilitates predator survival and hence prey survival as compared to the unconnected fragmented case. Our main result is counterintuitive: in the presence of a high quality predator, connected fragmented habitats ensure a better chance for coexistence than does even the unfragmented case. We explain why a connected fragmented habitat might thus be beneficial for the stabilization of the system, and how connections between sub-habitats are able to protect prey population from over-exploitation. In the model, habitat fragmentation is separated from the effects of habitat destruction, in order to better understand how populations react to habitat transformation.     

Patch Size and Connectivity Thresholds for Butterfly Habitat Restoration

Abstract:  Recovery of endangered species in highly fragmented habitats often requires habitat restoration. Selection of restoration sites typically involves too many options and too much uncertainty to reach a decision based on existing reserve design methods. The Fender's blue butterfly (  Icaricia icarioides fenderi ) survives in small, isolated patches of remnant prairie in Oregon's Willamette Valley—a habitat for which <0.5% of the original remains. Recovery of this species will require considerable habitat restoration. We investigated the potential of biologically based rules of thumb and more complex models to serve as tools in making land acquisitions. Based on Fender's blue dispersal behavior and demography, we have estimated that restored patches should be <1 km from existing habitat and at least 2 ha. We compared these rules to the results of two modeling approaches: an incidence function model and a spatially explicit simulation of demography and dispersal behavior. Not surprisingly, the simple rules and complex models all conclude that large (>2 ha) connected (<1 km) patches have the highest restoration value. The dispersal model, however, suggests that small, connected patches have more restoration value than large, isolated patches, whereas the incidence function model suggests that size and connectivity are equally important. These differences stem from model assumptions. We used incidence functions to predict long-term, stochastic, steady-state conditions and dispersal simulations to predict short-term (25-year) colonization dynamics. To apply our results in the context of selecting restoration sites on the ground, we recommend selecting nearby sites when short-term colonization dynamics are expected to be an important aspect of a species' biology.     

Habitat and landscape characteristics underlying anuran community structure along an urban-rural gradient

Urbanization has been cited as an important factor in worldwide amphibian declines, and although recent work has illustrated the important influence of broad-scale ecological patterns and processes on amphibian populations, little is known about the factors structuring amphibian communities in urban landscapes. We therefore examined amphibian community responses to wetland habitat availability and landscape characteristics along an urban-rural gradient in central Iowa, USA, a region experiencing rapid suburban growth. We conducted call surveys at 61 wetlands to estimate anuran calling activity, and quantified wetland habitat structure and landscape context. We used canonical correspondence analysis (CCA) to examine patterns in anuran community structure and identify the most important variables associated with those patterns. Urban density at the landscape scale had a significant negative influence on overall anuran abundance and diversity. While every species exhibited a decrease in abundance with increasing urban density, this pattern was especially pronounced for species requiring post-breeding upland habitats. Anurans most affected by urbanization were those associated with short hydroperiods, early breeding activity, and substantial upland habitat use. We suggest that broad-scale landscape fragmentation is an important factor underlying anuran community structure in this region, possibly due to limitations on the accessibility of otherwise suitable habitat in fragmented urban landscapes. This study underscores the importance of a regional approach to amphibian conservation in urban and urbanizing areas; in fragmented landscapes, a network of interconnected wetland and upland habitats may be more likely to support a successful, diverse anuran community than will isolated sites.     

Anthropogenic areas as incidental substitutes for original habitat

One speaks of ecological substitutes when an introduced species performs, to some extent, the ecosystem function of an extirpated native species. We suggest that a similar case exists for habitats. Species evolve within ecosystems, but habitats can be destroyed or modified by natural and human‐made causes. Sometimes habitat alteration forces animals to move to or remain in a suboptimal habitat type. In that case, the habitat is considered a refuge, and the species is called a refugee. Typically refugee species have lower population growth rates than in their original habitats. Human action may lead to the unintended generation of artificial or semiartificial habitat types that functionally resemble the essential features of the original habitat and thus allow a population growth rate of the same magnitude or higher than in the original habitat. We call such areas substitution habitats and define them as human‐made habitats within the focal species range that by chance are partial substitutes for the species’ original habitat. We call species occupying a substitution habitat adopted species. These are 2 new terms in conservation biology. Examples of substitution habitats are dams for European otters, wheat and rice fields for many steppeland and aquatic birds, and urban areas for storks, falcons, and swifts. Although substitution habitats can bring about increased resilience against the agents of global change, the conservation of original habitat types remains a conservation priority.