Complementary habitat use by wild bees in agro-natural landscapes

Human activity causes abrupt changes in resource availability across the landscape. In order to persist in human-altered landscapes organisms need to shift their habitat use accordingly. Little is known about the mechanisms by which whole communities persist in human-altered landscapes, including the role of complementary habitat use. We define complementary habitat use as the use of different habitats at different times by the same group of species during the course of their activity period. We hypothesize that complementary habitat use is a mechanism through which native bee species persist in human-altered landscapes. To test this idea, we studied wild bee communities in agro-natural landscapes and explored their community-level patterns of habitat and resource use over space and time. The study was conducted in six agro-natural landscapes in the eastern United States, each containing three main bee habitat types (natural habitat, agricultural fields, and old fields). Each of the three habitats exhibited a unique seasonal pattern in amount, diversity, and composition of floral resources, and together they created phenological complementarity in foraging resources for bees. Individual bee species as well as the bee community responded to these spatiotemporal patterns in floral availability and exhibited a parallel pattern of complementary habitat use. The majority of wild bee species, including all the main crop visitors, used fallow areas within crops early in the season, shifted to crops in mid-season, and used old-field habitats later in the season. The natural-forest habitat supported very limited number of bees, mostly visitors of non-crop plants. Old fields are thus an important feature in these arable landscapes for maintaining crop pollination services. Our study provides a detailed examination of how shifts in habitat and resource use may enable bees to persist in highly dynamic agro-natural landscapes, and points to the need for a broad cross-habitat perspective in managing these landscapes.     

Bird conservation and the land sharing-sparing continuum in farmland-dominated landscapes of lowland England

Empirical evidence from many regions suggests that most species would be least negatively affected if human food demand were met through high-yield agricultural production and conservation of nonfarm ecosystems (land sparing), rather than through wildlife-friendly farming over a larger area (land sharing). However, repeated glaciation and a long history of agriculture may lead to different results in regions such as western Europe. We compared the consequences of land sparing and land sharing on breeding bird species in 2 lowland regions of England, The Fens, with 101 species, and Salisbury Plain, with 83. We derived density–yield responses for each species and then estimated regional population size under regional food production strategies, including land sharing and land sparing, a range of intermediate strategies, and a novel mixed strategy. In both regions, more species achieved maximum regional population size under land sparing than land sharing. In The Fens, the majority of birds were loser species (estimated to have smaller populations under all food production strategies than in the preagricultural baseline scenario), whereas in Salisbury Plain the majority were winners (smaller populations in the preagricultural baseline scenario). Loser species overwhelmingly achieved maximum regional population size under land sparing, whereas winner species achieved maximum regional population size under either land sharing or an intermediate strategy, highlighting the importance of defining which groups of species are the target of conservation. A novel 3-compartment strategy (combining high-yield farming, natural habitat, and low-yield farming) often performed better than either land sharing or land sparing. Our results support intermediate or 3-compartment land-sparing strategies to maximize bird populations across lowland agricultural landscapes. To deliver conservation outcomes, any shift toward land sparing must, however, ensure yield increases are sustainable in the long term, do not entail increased negative effects on surrounding areas, and are linked to allocation of land for nature.     

Agricultural production and bird conservation in complex landscapes of the dry Chaco

The South American dry Chaco is a mosaic of woody vegetation and grasslands with high deforestation rates in recent decades. Considering forests and grasslands as the main natural habitats, we assessed the trade-offs between bird populations and agricultural production to compare the potential consequences of different land use strategies (‘sharing’, ‘sparing’, and intermediate) for populations of bird species sensitive to agriculture, while attaining a regional production target. We evaluated how populations responded to scenarios with different proportions of forest and grasslands, considering three reference states (100% forest, 80:20% and 50:50% forest and grasslands, respectively); and scenarios capable of meeting three after-farming scenarios, with land destined to reach a regional production target with three variations of forest:grasslands within spared land. We fitted curves to relate bird abundance to agricultural yield along a gradient of meat production intensity; and we classified bird species as ‘losers’ (if their populations were lower than the baseline population in the reference state, at any level of production) and ‘winners’ (if their current populations were higher than the baseline population). At the ‘current’ (c. 2010) level of regional agricultural production, we found a similar number of loser species maximized by land-sparing and land-sharing strategies; while intermediate strategies were the least favourable to balance production and bird populations. Under the most probable scenarios of increases in regional meat production, most loser bird species populations were maximized by a land-sparing strategy, suggesting that if meat production targets are going to increase in the region, this can be more efficiently achieved by combining well-protected forests and grasslands, and high-yielding mechanized agriculture (e.g. soybean). Our results highlight the importance of assessing all the important natural habitats (e.g. forests and grasslands) of a region to explore conservation strategies at a regional scale.     

Assessing Risks to Biodiversity from Future Landscape Change

We examined the impacts of possible future land development patterns on the biodiversity of a landscape. Our landscape data included a remote sensing derived map of the current habitat of the study area and six maps of future habitat distributions resulting from different land development scenarios. Our species data included lists of all bird, mammal, reptile, and amphibian species in the study area, their habitat associations, and area requirements for each. We estimated the area requirements using home ranges, sampled population densities, or genetic area requirements that incorporate dispersal distances. Our measures of biodiversity were species richness and habitat abundance. We calculated habitat abundance in two ways. First, we computed the total habitat area for each species in each landscape. Second, we calculated the number of habitat units for each species in each landscape by dividing the size of each habitat patch in the landscape by the area requirement and summing over all patches. Species richness was based on presence of habitat. Species became extinct in the landscape if they had no habitat area or no habitat units, respectively. We then computed ratios of habitat abundance in each future landscape to habitat abundance in the present for each species. We also computed the ratio of future to present species richness. We then calculated summary statistics across all species. Species richness changed little from present to future. There were distinctly greater risks to habitat abundance in landscapes that extrapolated from present trends or zoning patterns, however, as opposed to landscapes in which land development activities followed more constrained patterns. These results were stable when tested using Monte Carlo simulations and sensitivity tests on the area requirements. We conclude that this methodology can begin to discriminate the effects of potential changes in land development on vertebrate biodiversity.     

Optimizing conservation in species-specific agricultural landscapes

Recovery of grassland birds in agricultural landscapes is a global imperative. Agricultural landscapes are complex, and the value of resource patches may vary substantially among species. The spatial extent at which landscape features affect populations (i.e., scale of effect) may also differ among species. There is a need for regional-scale conservation planning that considers landscape-scale and species-specific responses of grassland birds to environmental change. We developed a spatially explicit approach to optimizing grassland conservation in the context of species-specific landscapes and prioritization of species recovery and applied it to a conservation program in Kentucky (USA). We used a hierarchical distance-sampling model with an embedded scale of effect predictor to estimate the relationship between landscape structure and abundance of eastern meadowlarks (Sturnella magna), field sparrows (Spizella pusilla), and northern bobwhites (Colinus virginianus). We used a novel spatially explicit optimization procedure rooted in multi-attribute utility theory to design alternative conservation strategies (e.g., prioritize only northern bobwhite recovery or assign equal weight to each species’ recovery). Eastern meadowlarks and field sparrows were more likely to respond to landscape-scale resource patch adjacencies than landscape-scale patch densities. Northern bobwhite responded to both landscape-scale resource patch adjacencies and densities and responded strongly to increased grassland density. Effects of landscape features on local abundance decreased as distance increased and had negligible influence at 0.8 km for eastern meadowlarks (0.7–1.2 km 95% Bayesian credibility intervals [BCI]), 2.5 km for field sparrows (1.5–5.8 km 95% BCI), and 8.4 km for bobwhite (6.4–26 km 95% BCI). Northern bobwhites were predicted to benefit greatly from future grassland conservation regardless of conservation priorities, but eastern meadowlark and field sparrow were not. Our results suggest similar species can respond differently to broad-scale conservation practices because of species-specific, distance-dependent relationships with landscape structure. Our framework is quantitative, conceptually simple, customizable, and predictive and can be used to optimize conservation in heterogeneous ecosystems while considering landscape-scale processes and explicit prioritization of species recovery.     

Tropical amphibians in shifting thermal landscapes under land‐use and climate change

Land‐cover and climate change are both expected to alter species distributions and contribute to future biodiversity loss. However, the combined effects of land‐cover and climate change on assemblages, especially at the landscape scale, remain understudied. Lowland tropical amphibians may be particularly susceptible to changes in land cover and climate warming because many species have narrow thermal safety margins resulting from air and body temperatures that are close to their critical thermal maxima (CT_max). We examined how changing thermal landscapes may alter the area of thermally suitable habitat (TSH) for tropical amphibians. We measured microclimates in 6 land‐cover types and CT_max of 16 frog species in lowland northeastern Costa Rica. We used a biophysical model to estimate core body temperatures of frogs exposed to habitat‐specific microclimates while accounting for evaporative cooling and behavior. Thermally suitable habitat area was estimated as the portion of the landscape where species CT_max exceeded their habitat‐specific maximum body temperatures. We projected changes in TSH area 80 years into the future as a function of land‐cover change only, climate change only, and combinations of land‐cover and climate‐change scenarios representing low and moderate rates of change. Projected decreases in TSH area ranged from 16% under low emissions and reduced forest loss to 30% under moderate emissions and business‐as‐usual land‐cover change. Under a moderate emissions scenario (A1B), climate change alone contributed to 1.7‐ to 4.5‐fold greater losses in TSH area than land‐cover change only, suggesting that future decreases in TSH from climate change may outpace structural habitat loss. Forest‐restricted species had lower mean CT_max than species that occurred in altered habitats, indicating that thermal tolerances will likely shape assemblages in changing thermal landscapes. In the face of ongoing land‐cover and climate change, it will be critical to consider changing thermal landscapes in strategies to conserve ectotherm species.     

Effects of Land-Use and Tsetse Fly Control on Bird Species Richness in Southwestern Ethiopia

Successful control of tsetse (Glossina spp.)-transmitted trypanosomiasis in the Ghibe Valley, Ethiopia, appears to have accelerated conversion of wooded grassland into cropland. Land conversion, in turn, may have fragmented wildlife habitat. Our objective was to assess the influence of the expansion of agricultural land-use, brought about by tsetse control, on ecological properties by using bird species richness and composition as indicators of environmental impacts. We sampled bird species richness and composition (using Timed-Species counts) and habitat structure (using field sampling and remote sensing) in four land cover/land-use types in areas subjected to tsetse fly control and adjacent areas without control. At the height of the growing season bird species numbers and vegetative complexity were greater in the small-holder, oxen-plowed fields and riparian woodlands than in wooded grasslands or in large-holder, tractor-plowed fields. Species composition was highly dissimilar (40–70% dissimilarity) comparing among land-use types, with many species found only in a single type. This implies that trypanosomiasis control that results in land conversion from wooded grasslands to small-holder farming in this region may have no adverse impacts on bird species numbers but will alter composition. These results also suggest that moderate land-use by humans (e.g., small-holder field mosaics) increases habitat heterogeneity and bird species richness relative to high levels of use (e.g., tractor-plowed fields). Tsetse control may be indirectly maintaining species richness in the valley by encouraging the differential spread of these small-scale, heterogeneous farms in place of large-scale, homogeneous farms. Nevertheless, if the extent of small-holder farms significantly exceeds that of present levels, negative impacts on bird species richness and large shifts in species composition may occur.     

A Comparison of Landscapes Occupied by Increasing and Decreasing Populations of Grassland Birds

Abstract:  For several decades, many grassland bird species have been declining in abundance throughout the Midwest and Great Plains regions of the United States, possibly due to loss of natural grassland habitat and increasing urbanization. I used 20 years of data from the North American Breeding Bird Survey to identify increasing, decreasing, and stable populations of 36 grassland-nesting bird species. I characterized the immediate landscape (circle with radius = 30 km) surrounding each population based on data from the National Resources Inventory. For each landscape, I calculated the proportion of eight different land-cover types: restored grassland, rangeland, cultivated cropland, pasture, noncultivated cropland, forest, urban land, and water. Using a null model, I compared landscape composition of increasing, decreasing, and stable populations. As predicted on the basis of the habitat preferences of grassland birds, increasing populations inhabited landscapes that contained significantly more restored grassland and rangeland but significantly less forest land and urban land than landscapes inhabited by decreasing populations. There was no significant difference in the proportion of cropland within the landscapes of increasing and decreasing populations, although cropland composed a large proportion (>30%) of many landscapes. In contrast, restored grassland typically composed a very small proportion (<3.5%) of total land cover, yet it was significantly more common in the landscapes of increasing than decreasing populations. These results suggest that grassland birds may benefit from government initiatives, such as the Conservation Reserve Program, that promote the restoration of grassland at a landscape scale.     

Minimum viable metapopulation size, extinction debt, and the conservation of a declining species.

A key question facing conservation biologists is whether declines in species' distributions are keeping pace with landscape change, or whether current distributions overestimate probabilities of future persistence. We use metapopulations of the marsh fritillary butterfly Euphydryas aurinia in the United Kingdom as a model system to test for extinction debt in a declining species. We derive parameters for a metapopulation model (incidence function model, IFM) using information from a 625-km2 landscape where habitat patch occupancy, colonization, and extinction rates for E. aurinia depend on patch connectivity, area, and quality. We then show that habitat networks in six extant metapopulations in 16-km2 squares were larger, had longer modeled persistence times (using IFM), and higher metapopulation capacity (lambdaM) than six extinct metapopulations. However, there was a > 99% chance that one or more of the six extant metapopulations would go extinct in 100 years in the absence of further habitat loss. For 11 out of 12 networks, minimum areas of habitat needed for 95% persistence of metapopulation simulations after 100 years ranged from 80 to 142 ha (approximately 5-9% of land area), depending on the spatial location of habitat. The area of habitat exceeded the estimated minimum viable metapopulation size (MVM) in only two of the six extant metapopulations, and even then by only 20%. The remaining four extant networks were expected to suffer extinction in 15-126 years. MVM was consistently estimated as approximately 5% of land area based on a sensitivity analysis of IFM parameters and was reduced only marginally (to approximately 4%) by modeling the potential impact of long-distance colonization over wider landscapes. The results suggest a widespread extinction debt among extant metapopulations of a declining species, necessitating conservation management or reserve designation even in apparent strongholds. For threatened species, metapopulation modeling is a potential means to identify landscapes near to extinction thresholds, to which conservation measures can be targeted for the best chance of success.     

Epiphyte Biodiversity in the Coffee Agricultural Matrix: Canopy Stratification and Distance from Forest Fragments

Abstract: Quality of the agricultural matrix profoundly affects biodiversity and dispersal in agricultural areas. Vegetatively complex coffee agroecosystems maintain species richness at larger distances from the forest. Epiphytes colonize canopy trees and provide resources for birds and insects and thus effects of agricultural production on epiphytes may affect other species. We compared diversity, composition, and vertical stratification of epiphytes in a forest fragment and in two coffee farms differing in management intensity in southern Mexico. We also examined spatial distribution of epiphytes with respect to the forest fragment to examine quality of the two agricultural matrix types for epiphyte conservation. We sampled vascular epiphytes in a forest fragment, a shade polyculture farm, and a shade monoculture farm at 100 m, 200 m, and 400 m from the forest. Epiphyte and orchid richness was greater in the forest than in the monoculture but richness was similar in the forest and polyculture farm. Epiphyte species composition differed with habitat type, but not with distance from the forest. In the forest, epiphytes were distributed throughout tree canopies, but in the farms, epiphytes were primarily found on trunks and larger branches. Epiphyte richness and species similarity to forest species declined with distance from the forest fragment in the monoculture, but richness and similarity to forest species did not decline with distance from forest in the polyculture. This suggests polyculture coffee has greater conservation value. In contrast, monoculture coffee is likely a sink habitat for epiphytes dispersing from forests into coffee. Coffee farms differ from forests in terms of the habitat they provide and species composition, thus protecting forest fragments is essential for epiphyte conservation. Nonetheless, in agricultural landscapes, vegetatively complex coffee farms may contribute to conservation of epiphytes more than other agricultural land uses.     

A strategy for wildlife management in depopulating rural areas of Japan

Former ranges of wild animals have been reestablished in many developed countries. However, this reestablishment has led to increasing human–wildlife conflict in agroforest ecosystems. In Japan, human–wildlife conflict, such as crop raiding by and ecological impacts of wild ungulates and primates, is a serious problem in depopulated rural areas due to these animal range expansions and increased abundances. Japan's human population is predicted to decline by 24% by 2050, and approximately 20% of agricultural settlements will become completely depopulated. In this scenario, anthropogenic pressures on wildlife (e.g., hunting and habitat alteration) will continue to decrease and human–wildlife conflict will increase due to increasing wildlife recovery. Japan's local governments plan to slow range recovery, prevent species reestablishment, or remove recolonizing large mammals through lethal control. This strategy, however, is not cost-effective, and workforce shortages in depopulated communities make it infeasible. Moreover, the suppression of wildlife prevents the recovery of ecological functions and thus would degrade regional biodiversity. The declining pressure on wildlife that accompanies human depopulation will prevent the restoration of any past states of human–wildlife interaction. We suggest human-used areas in rural landscapes be aggregated in compact cities and that in transition zones between human settlements and depopulated lands that land-sharing approaches be applied. Concentrating management efforts in compact cities may effectively decrease human–wildlife conflict, rather than intensifying human pressures. Reforestation of depopulated lands may lead to recovery of wildlife habitats, their ecosystem functions, and regional biodiversity due to minimization of negative anthropogenic effects (land-sparing approach). Balancing resolution of human–wildlife conflict and ecological rewilding could become a new, challenging task for regional wildlife managers.     

Thresholds in Songbird Occurrence in Relation to Landscape Structure

Abstract:  Theory predicts the occurrence of threshold levels of habitat in landscapes, below which ecological processes change abruptly. Simulation models indicate that below critical thresholds, fragmentation of habitat influences patch occupancy by decreasing colonization rates and increasing rates of local extinction. Uncovering such putative relationships is important for understanding the demography of species and in developing sound conservation strategies. Using segmented logistic regression, we tested for thresholds in occurrence of 15 bird species as a function of the amount of suitable habitat at multiple scales (150–2000-m radii). Suitable habitat was defined quantitatively based on previously derived, spatially explicit distribution models for each species. The occurrence of 10 out of 15 species was influenced by the amount of habitat at a landscape scale (≥500-m radius). Of these species all but one were best predicted by threshold models. Six out of nine species exhibited asymptotic thresholds; the effects of habitat loss intensified at low amounts of habitat in a landscape. Landscape thresholds ranged from 8.6% habitat to 28.7% (     = 18.5 ± 2.6%[95% CI]). For two species landscape thresholds coincided with sensitivity to fragmentation; both species were more likely to occur in large patches, but only when the amount of habitat in a landscape was low. This supports the fragmentation threshold hypothesis. Nevertheless, the occurrence of most species appeared to be unaffected by fragmentation, regardless of the amount of habitat present at landscape extents. The thresholds we identified may be useful to managers in establishing conservation targets. Our results indicate that findings of landscape-scale studies conducted in regions with relatively high proportions of habitat and low fragmentation may not be applicable in regions with low habitat proportions and high fragmentation.     

Multispecies and Multiscale Conservation Planning: Setting Quantitative Targets for Red‐Listed Lichens on Ancient Oaks

Abstract: Species occurrence in a habitat patch depends on local habitat and the amount of that habitat in the wider landscape. We used predictions from empirical landscape studies to set quantitative conservation criteria and targets in a multispecies and multiscale conservation planning effort. We used regression analyses to compare species richness and occurrence of five red‐listed lichens on 50 ancient oaks (Quercus robur; 120–140 cm in diameter) with the density of ancient oaks in circles of varying radius from each individual oak. Species richness and the occurrence of three of the five species were best explained by increasing density of oaks within 0.5 km; one species was best explained by the density of oaks within 2 km, and another was best predicted by the density of oaks within 5 km. The minimum numbers of ancient oaks required for “successful conservation” was defined as the number of oaks required to obtain a predicted local occurrence of 50% for all species included or a predicted local occurrence of 80% for all species included. These numbers of oaks were calculated for two relevant landscape scales (1 km² and 13 km²) that corresponded to various species responses, in such a way that calculations also accounted for local number of oaks. Ten and seven of the 50 ancient oaks surveyed were situated in landscapes that already fulfilled criteria for successful conservation when the 50% and 80% criteria, respectively, were used to define the level of successful conservation. For cost‐efficient conservation, oak stands in the landscapes most suitable for successful conservation should be prioritized for conservation and management (e.g., grazing and planting of new oaks) at the expense of oak stands situated elsewhere.     

Amphibian occurrence is influenced by current and historic landscape characteristics.

Human-induced habitat loss and degradation are major threats to wetland species as reflected in the fact that wetlands have declined by more than 50% in Europe and North America during the last century. Both current and historic land-use patterns are likely to be significant determinants of wetland species' distributions; however their relative importance is often unknown. We studied the importance of local (study pond) and landscape (current and 18th-century landscape) characteristics in explaining the occurrence and species richness of amphibians (Rana arvalis, Bufo bufo, and Triturus vulgaris) on the Swedish island of Gotland, where more than 40% of wetlands have been lost since the 18th century. Current local habitat characteristics were the strongest determinants of occurrence for all study species. Additionally, species occurrence was related to current and historic landscape characteristics, which generally explained equal amounts of the variation in species-occurrence data. The proportions of both current and historic arable land were negative determinants of amphibian occurrence and species richness, indicating that agricultural land use may have an overall negative impact on amphibians, and that amphibians may occur less frequently in areas with a long agricultural history. Likewise, historic forest area was positively related to B. bufo occurrence and species richness, whereas current forests had no significant effects, suggesting that there may be a lag in the response of amphibians to agriculture-mediated habitat loss. Our results suggest that historic land-use patterns may influence current amphibian populations and that inclusion of historic land-use information could be a valuable tool in future studies on amphibian-habitat relations.     

若尔盖西部草甸的基本类型、特点及近几十年来的变化

2000年8月对若尔盖西部草地的调查，确定了该地18个草丛类型（不包括典型的沼泽植被）。经群丛间Pearson最大相关系数配合生境作图，将该地区的草地群丛分成3大类，即沼生草地，中生性草甸和旱生性草地。各群丛的SW指数，物种丰富度，地上生物量，裸地面积按照生境序列作图，显示中性草甸具有最高的物种丰富度和地上生物量。群丛C和J分别是沼生和旱生植被与中生性草甸之间的拐点式过渡类型，其物种丰富度明显比周期的群丛低，生境转换关键种分别为鹅绒委陵菜和垂穗披碱草。在草地物种组成上，杂类草占81．3％，莎草类仅占6．8％；群丛建群种方面，杂类草占61％，莎草类占28％；在次优势种方面，杂类草和禾草所占比重都有所增加。草原成分出现在丛中生至旱生的群丛中，在有些群丛中甚至成为次优势种。沼泽消退，黄河河道附近沙化以及过牧的阳坡山地生长带刺的旱生化植被，说明当地正经历着干旱化的过程，原因不仅有自然的还有人为的，特别是草地的过度使用。最后对当地草场的管理及发展提出了一些建议。图3表2参20     

Evaluating Effects of Habitat Loss and Land-Use Continuity on Ant Species Richness in Seminatural Grassland Remnants

Abstract:  Seminatural grasslands in Europe are susceptible to habitat destruction and fragmentation that result in negative effects on biodiversity because of increased isolation and area effects on extinction rate. However, even small habitat patches of seminatural grasslands might be of value for conservation and restoration of species richness in a landscape with a long history of management, which has been argued to lead to high species richness. We tested whether ant communities have been negatively affected by habitat loss and increased isolation of seminatural grasslands during the twentieth century. We examined species richness and community composition in seminatural grasslands of different size in a mosaic landscape in Central Sweden. Grasslands managed continuously over centuries harbored species-rich and ecologically diverse ant communities. Grassland remnant size had no effect on ant species richness. Small grassland remnants did not harbor a nested subset of the ant species of larger habitats. Community composition of ants was mainly affected by habitat conditions. Our results suggest that the abandonment of traditional land use and the encroachment of trees, rather than the effects of fragmentation, are important for species composition in seminatural grasslands. Our results highlight the importance of considering land-use continuity and dispersal ability of the focal organisms when examining the effects of habitat loss and fragmentation on biodiversity. Landscape history should be considered in conservation programs focusing on effects of land-use change.     

Is the effect of forest structure on bird diversity modified by forest productivity?

Currently, the most common strategy when managing forests for biodiversity at the landscape scale is to maintain structural complexity within stands and provide a variety of seral stages across landscapes. Advances in ecological theory reveal that biodiversity at continental scales is strongly influenced by available energy (i.e., climate factors relating to heat and light and primary productivity). This paper explores how available energy and forest structural complexity may interact to drive biodiversity at a regional scale. We hypothesized that bird species richness exhibits a hump-shaped relationship with energy at the regional scale of the northwestern United States. As a result, we hypothesized that the relationship between energy and richness within a landscape is positive in energy-limited landscapes and flat or decreasing in energy-rich landscapes. Additionally, we hypothesized that structural complexity explains less of the variation in species richness in energy-limited environments and more in energy-rich environments and that the slope of the relationship between structural complexity and richness is greatest in energy-rich environments. We sampled bird communities and vegetation across seral stages and biophysical settings at each of five landscapes arrayed across a productivity gradient from the Pacific Coast to the Rocky Mountains within the five northwestern states of the contiguous United States. We analyzed the response of richness to structural complexity and energy covariates at each landscape. We found that (1) richness had a hump-shaped relationship with available energy across the northwestern United States, (2) the landscape-scale relationships between energy and richness were positive or hump shaped in energy-limited locations and were flat or negative in energy-rich locations, (3) forest structural complexity explained more of the variation in bird species richness in energy-rich landscapes, and (4) the slope of the relationship between forest structural complexity and richness was steepest in energy-limited locations. In energy-rich locations, forest managers will likely increase landscape-scale bird diversity by providing a range of forest structural complexity across all seral stages. In low-energy environments, bird diversity will likely be maximized by managing local high-energy hotspots judiciously and adjusting harvest intensities in other locations to compensate for slower regeneration rates.     

Fragments as Islands: a Synthesis of Faunal Responses to Habitat Patchiness

Abstract:  Scientific interest in the impact of habitat fragmentation on biodiversity is increasing, but our understanding of fragmentation is clouded by a lack of appreciation for fundamental similarities and differences across studies representing a wide range of taxa and landscape types. In an effort to synthesize data describing ecological responses of animals to fragmentation across two classes of independent variables (taxonomic group and landscape), we sampled 148 studies of five major faunal groups from the primary literature and analyzed data on 13 variables extracted from those studies. We focused our analyses on three classes of dependent variables (effects of area and isolation on species richness, z values, and nestedness and species composition). Area ranged over more orders of magnitude than isolation and tended to explain more variation in species richness than isolation. There were few matrix or taxon effects on the patterns we investigated, although we did find that sky islands tended to manifest isolation effects on both species richness and nestedness more frequently than other patch types. Sky islands may offer insight into the future of habitat patches fragmented by contemporary habitat loss, and because they show a stronger effect of isolation than other patch types, we suggest that isolation will play an increasing role in the biology of habitat fragments. We use multiple lines of evidence to suggest that our understanding of the role of isolation on community assembly in fragmented landscapes is inadequate. Finally, our observation that consistent taxonomic differences in community patterns were minimal suggests that conservation actions intended to mitigate the negative effects of extinction may have far-reaching effects across taxonomic groups.     

Importance of Reserve Size and Landscape Context to Urban Bird Conservation

Abstract:  We tested whether reserve size, landscape surrounding the reserve, and their interaction affect forest songbirds in the metropolitan area of Seattle, Washington (U.S.A.), by studying 29 reserves of varying size (small, medium, large) and surrounding urbanization intensity (urban, suburban, exurban). Larger reserves contained richer and less even bird communities than smaller reserves. These size effects disappeared when we removed the positive correlation of shrub diversity with reserve size, suggesting that greater habitat diversity in large reserves supported additional species, some of which were rare. Standardizing the number of individuals detected among all reserve size classes reversed the effect of size on richness in exurban landscapes and reduced the magnitude of the effect in suburban or urban landscapes. The latter change suggested that richness increased with reserve size in most landscapes because larger areas also supported larger samples from the regional bird species pool. Most bird species associated with native forest habitat (native forest species) and with human activity (synanthropic species) were present in reserves larger than 42 ha and surrounded by >40% urban land cover, respectively. Thus, we recommend these thresholds as means for conserving the composition of native bird communities in this mostly forested region. Native forest species were least abundant and synanthropic species most abundant in urban landscapes, where exotic ground and shrub vegetation was most common. Therefore, control of exotic vegetation may benefit native songbird populations. Bird nests in shrubs were most dense in medium (suburban) and large reserves (urban) and tended to be most successful in medium (suburban) and large reserves (exurban), potentially supplying another mechanism by which reserve size increased retention of native forest species.