Effects of Dam‐Induced Landscape Fragmentation on Amazonian Ant–Plant Mutualistic Networks

Mutualistic networks are critical to biological diversity maintenance; however, their structures and functionality may be threatened by a swiftly changing world. In the Amazon, the increasing number of dams poses a large threat to biological diversity because they greatly alter and fragment the surrounding landscape. Tight coevolutionary interactions typical of tropical forests, such as the ant–myrmecophyte mutualism, where the myrmecophyte plants provide domatia nesting space to their symbiotic ants, may be jeopardized by the landscape changes caused by dams. We analyzed 31 ant–myrmecophyte mutualistic networks in undisturbed and disturbed sites surrounding Balbina, the largest Central Amazonian dam. We tested how ant–myrmecophyte networks differ among dam‐induced islands, lake edges, and undisturbed forests in terms of species richness, composition, structure, and robustness (number of species remaining in the network after partner extinctions). We also tested how landscape configuration in terms of area, isolation, shape, and neighborhood alters the structure of the ant–myrmecophyte networks on islands. Ant–myrmecophytic networks were highly compartmentalized in undisturbed forests, and the compartments had few strongly connected mutualistic partners. In contrast, networks at lake edges and on islands were not compartmentalized and were negatively affected by island area and isolation in terms of species richness, density, and composition. Habitat loss and fragmentation led to coextinction cascades that contributed to the elimination of entire ant–plant compartments. Furthermore, many myrmecophytic plants in disturbed sites lost their mutualistic ant partners or were colonized by opportunistic, nonspecialized ants. Robustness of ant–myrmecophyte networks on islands was lower than robustness near lake edges and in undisturbed forest and was particularly susceptible to the extinction of plants. Beyond the immediate habitat loss caused by the building of large dams in Amazonia, persistent edge effects and habitat fragmentation associated with dams had large negative effects on animal–plant mutualistic networks. Efectos de la Fragmentación del Paisaje Inducida por Presas sobre Redes Mutualistas Hormiga‐Planta Amazónicas     

Fragmentation effects on an endangered species across a gradient from the interior to edge of its range

Understanding how habitat fragmentation affects individual species is complicated by challenges associated with quantifying species-specific habitat and spatial variability in fragmentation effects within a species’ range. We aggregated a 29-year breeding survey data set for the endangered marbled murrelet (Brachyramphus marmoratus) from >42,000 forest sites throughout the Pacific Northwest (Oregon, Washington, and northern California) of the United States. We built a species distribution model (SDM) in which occupied sites were linked with Landsat imagery to quantify murrelet-specific habitat and then used occupancy models to test the hypotheses that fragmentation negatively affects murrelet breeding distribution and that these effects are amplified with distance from the marine foraging habitat toward the edge of the species’ nesting range. Murrelet habitat declined in the Pacific Northwest by 20% since 1988, whereas the proportion of habitat comprising edges increased by 17%, indicating increased fragmentation. Furthermore, fragmentation of murrelet habitat at landscape scales (within 2 km of survey stations) negatively affected occupancy of potential breeding sites, and these effects were amplified near the range edge. On the coast, the odds of occupancy decreased by 37% (95% confidence interval [CI] –54 to 12) for each 10% increase in edge habitat (i.e., fragmentation), but at the range edge (88 km inland) these odds decreased by 99% (95% CI 98 to 99). Conversely, odds of murrelet occupancy increased by 31% (95% CI 14 to 52) for each 10% increase in local edge habitat (within 100 m of survey stations). Avoidance of fragmentation at broad scales but use of locally fragmented habitat with reduced quality may help explain the lack of murrelet population recovery. Further, our results emphasize that fragmentation effects can be nuanced, scale dependent, and geographically variable. Awareness of these nuances is critical for developing landscape-level conservation strategies for species experiencing broad-scale habitat loss and fragmentation.     

Sensitivity of Marine-Reserve Design to the Spatial Resolution of Socioeconomic Data

Abstract:  Socioeconomic considerations should have an important place in reserve design. Systematic reserve-selection tools allow simultaneous optimization for ecological objectives while minimizing costs but are seldom used to incorporate socioeconomic costs in the reserve-design process. The sensitivity of this process to biodiversity data resolution has been studied widely but the issue of socioeconomic data resolution has not previously been considered. We therefore designed marine reserves for biodiversity conservation with the constraint of minimizing commercial fishing revenue losses and investigated how economic data resolution affected the results. Incorporating coarse-resolution economic data from official statistics generated reserves that were only marginally less costly to the fishery than those designed with no attempt to minimize economic impacts. An intensive survey yielded fine-resolution data that, when incorporated in the design process, substantially reduced predicted fishery losses. Such an approach could help minimize fisher displacement because the least profitable grounds are selected for the reserve. Other work has shown that low-resolution biodiversity data can lead to underestimation of the conservation value of some sites, and a risk of overlooking the most valuable areas, and we have similarly shown that low-resolution economic data can cause underestimation of the profitability of some sites and a risk of inadvertently including these in the reserve. Detailed socioeconomic data are therefore an essential input for the design of cost-effective reserve networks.     

Incorporating consumer–resource spatial interactions in reserve design

The persistence of species in reserves depends in large part on the persistence of functional ecological interactions. Despite their importance, however, ecological interactions have not yet been explicitly incorporated into conservation prioritization methods. We develop here a general method for incorporating consumer–resource interactions into spatial reserve design. This method protects spatial consumer–resource interactions by protecting areas that maintain the connectivity between the distribution of consumers and resources. We illustrate our method with a conservation planning case study of a mammalian predator, American marten (Martes americana), and its two primary prey species, Red-backed vole (Clethrionomys rutilus) and Deer mouse (Peromyscus maniculatus). The conservation goal was to identify a reserve for marten that comprised 12% of a forest management unit in the boreal forest in Québec, Canada. We compared reserves developed using analysis variants that utilized different levels of information about predator and prey habitat distributions, species-specific connectivity requirements, and interaction connectivity requirements. The inclusion of consumer–resource interactions in reserve-selection resulted in spatially aggregated reserves that maintained local habitat quality for the species. This spatial aggregation was not induced by applying a qualitative penalty for the boundary length of the reserve, but rather was a direct consequence of modelling the spatial needs of the interacting consumer and resources. Our method for maintaining connectivity between consumers and their resources within reserves can be applied even under the most extreme cases of either complete spatial overlap or complete spatial segregation of consumer–resource distributions. The method has been made available via public software.     

Adaptive Decision Rules for the Acquisition of Nature Reserves

Abstract:  Although reserve-design algorithms have shown promise for increasing the efficiency of conservation planning, recent work casts doubt on the usefulness of some of these approaches in practice. Using three data sets that vary widely in size and complexity, we compared various decision rules for acquiring reserve networks over multiyear periods. We explored three factors that are often important in real-world conservation efforts: uncertain availability of sites for acquisition, degradation of sites, and overall budget constraints. We evaluated the relative strengths and weaknesses of existing optimal and heuristic decision rules and developed a new set of adaptive decision rules that combine the strengths of existing optimal and heuristic approaches. All three of the new adaptive rules performed better than the existing rules we tested under virtually all scenarios of site availability, site degradation, and budget constraints. Moreover, the adaptive rules required no additional data beyond what was readily available and were relatively easy to compute.     

Use of Inverse Spatial Conservation Prioritization to Avoid Biological Diversity Loss Outside Protected Areas

Globally expanding human land use sets constantly increasing pressure for maintenance of biological diversity and functioning ecosystems. To fight the decline of biological diversity, conservation science has broken ground with methods such as the operational model of systematic conservation planning (SCP), which focuses on design and on‐the‐ground implementation of conservation areas. The most commonly used method in SCP is reserve selection that focuses on the spatial design of reserve networks and their expansion. We expanded these methods by introducing another form of spatial allocation of conservation effort relevant for land‐use zoning at the landscape scale that avoids negative ecological effects of human land use outside protected areas. We call our method inverse spatial conservation prioritization. It can be used to identify areas suitable for economic development while simultaneously limiting total ecological and environmental effects of that development at the landscape level by identifying areas with highest economic but lowest ecological value. Our method is not based on a priori targets, and as such it is applicable to cases where the effects of land use on, for example, individual species or ecosystem types are relatively small and would not lead to violation of regional or national conservation targets. We applied our method to land‐use allocation to peat mining. Our method identified a combination of profitable production areas that provides the needed area for peat production while retaining most of the landscape‐level ecological value of the ecosystem. The results of this inverse spatial conservation prioritization are being used in land‐use zoning in the province of Central Finland.     

Reserve design to optimize functional connectivity and animal density

Ecological distance-based spatial capture–recapture models (SCR) are a promising approach for simultaneously estimating animal density and connectivity, both of which affect spatial population processes and ultimately species persistence. We explored how SCR models can be integrated into reserve-design frameworks that explicitly acknowledge both the spatial distribution of individuals and their space use resulting from landscape structure. We formulated the design of wildlife reserves as a budget-constrained optimization problem and conducted a simulation to explore 3 different SCR-informed optimization objectives that prioritized different conservation goals by maximizing the number of protected individuals, reserve connectivity, and density-weighted connectivity. We also studied the effect on our 3 objectives of enforcing that the space-use requirements of individuals be met by the reserve for individuals to be considered conserved (referred to as home-range constraints). Maximizing local population density resulted in fragmented reserves that would likely not aid long-term population persistence, and maximizing the connectivity objective yielded reserves that protected the fewest individuals. However, maximizing density-weighted connectivity or preemptively imposing home-range constraints on reserve design yielded reserves of largely spatially compact sets of parcels covering high-density areas in the landscape with high functional connectivity between them. Our results quantify the extent to which reserve design is constrained by individual home-range requirements and highlight that accounting for individual space use in the objective and constraints can help in the design of reserves that balance abundance and connectivity in a biologically relevant manner.     

Decoupling fragmentation from habitat loss for spiders in patchy agricultural landscapes

Habitat loss reduces species diversity, but the effect of habitat fragmentation on number of species is less clear because fragmentation generally accompanies loss of habitat. We compared four methods that aim to decouple the effects of fragmentation from the effects of habitat loss. Two methods are based on species-area relations, one on Fisher's alpha index of diversity, and one on plots of cumulative number of species detected against cumulative area sampled. We used these methods to analyze the species diversity of spiders in 2, 3.2 × 4 km agricultural landscapes in Southern Judea Lowlands, Israel. Spider diversity increased as fragmentation increased with all four methods, probably not because of the additive within-patch processes, such as edge effect and heterogeneity. The positive relation between fragmentation and species diversity might reflect that most species can disperse through the fields during the wheat-growing season. We suggest that if a given area was designated for the conservation of spiders in Southern Judea Lowlands, Israel, a set of several small patches may maximize species diversity over time.     

Potential Spatial Overlap of Heritage Sites and Protected Areas in a Boreal Region of Northern Canada

Abstract:  Under article 8-J of the Convention on Biological Diversity, governments must engage indigenous and local communities in the designation and management of protected areas. A better understanding of the relationship between community heritage sites and sites identified to protect conventional conservation features could inform conservation-planning exercises on indigenous lands. We examined the potential overlap between Gwich'in First Nations' (Northwest Territories, Canada) heritage sites and areas independently identified for the protection of conventional conservation targets. We designed nine hypothetical protected-area networks with different targets for woodland caribou ( Rangifer tarandus caribou ) habitat, high-quality wetland areas, representative vegetation types, water bodies, environmentally significant area, territorial parks, and network aggregation. We compared the spatial overlap of heritage sites to these nine protected-area networks. The degree of spatial overlap (Jaccard similarity) between heritage sites and the protected-area networks with moderate or high aggregation was significantly higher ( p < 0.001) than random spatial overlap, whereas the overlap between heritage sites and the protected-area networks with no aggregation was not significant or significantly lower ( p < 0.001) than random spatial overlap. Our results suggest that protected-area networks designed to capture conventional conservation features may protect key heritage sites but only if the underlying characteristics of these sites are considered. The Gwich'in heritage sites are highly aggregated and only protected-area networks that had moderate and high aggregation had significant overlap with the heritage sites. We suggest that conventional conservation plans incorporate heritage sites into their design criteria to complement conventional conservation targets and effectively protect indigenous heritage sites .     

Can initial intraspecific spatial aggregation increase multi-year coexistence by creating temporal priority?

Both intraspecific spatial aggregation and temporal priority effects have the potential to increase long-term species coexistence. Theory and models suggest that intraspecific aggregation can facilitate coexistence via limited dispersal or asymmetric interaction distances. During community assembly, intraspecific aggregation may also delay interactions between more and less competitive species, thus creating opportunities for priority effects to facilitate longer-term coexistence. Few empirical studies have tested predictions about aggregation and coexistence, especially in the context of community assembly or ecological restoration. We investigated (1) impacts of intraspecific aggregation on the assembly of eight-species communities over three years, (2) the scale dependence of these impacts, and (3) implications for California prairie restoration. We planted eight native species in each of 19, 5 m wide, octagonal plots. Species were either interspersed throughout the plot or aggregated into eight, 2.2-m(2), wedge-shaped, monospecific sectors. Over three years, species diversity declined more quickly in interspersed plots than in aggregated plots. Two species had higher cover or increased more in interspersed than aggregated plots and were identified as "aggressives." Four species had higher cover or increased more in aggregated than interspersed plots and were identified as "subordinates." Within aggregated plots, aggressive species expanded beyond the sector in which they were originally seeded. Cover of aggressive species increased faster and reached higher values in sectors that were adjacent to the originally planted sector, compared to nonadjacent sectors. Cover of aggressive species also increased more and faster near plot centers, compared to plot edges. Areas near plot centers were representative of smaller aggregation patches since species were planted closer to heterospecific neighbors. Two subordinate species maintained higher cover near plot edges than near plot centers. Moreover, two subordinate species maintained higher cover when seeded in sectors farther away from aggressive species. These results suggest that initial intraspecific aggregation can facilitate species coexistence for at least three years, and larger aggregation patches may be more effective than smaller ones in the face of dispersing dominants. The creation of temporal priority effects may represent an underappreciated pathway by which intraspecific aggregation can increase coexistence. Restorationists may be able to maintain more diverse communities by planting in a mosaic of monospecific patches.     

Habitat Fragmentation Effects on Trophic Processes of Insect-Plant Food Webs

Abstract:  Habitat fragmentation is the transformation of once-extensive landscapes into smaller, isolated remnants surrounded by new types of habitat. There is ample evidence of impoverished biodiversity as a consequence of habitat fragmentation, but its most profound effects may actually result from functional changes in ecological processes such as trophic interactions. We studied the trophic processes of herbivory and parasitism in insect-plant food webs composed of hundreds of species in a fragmented woodland landscape. We recorded all plant species, collected mined leaves, and reared leafminers and parasitoids from 19 woodland remnants. Herbivory and parasitism rates were then analyzed in relation to woodland size and edge or interior location. Herbivory by leaf-mining insects and their overall parasitism rates decreased as woodland remnants became smaller. For each remnant the intensity of both processes differed between edge and interior. Our results provide novel evidence of the magnitude of habitat fragmentation effects, showing they can be so pervasive as to affect trophic processes of highly complex food webs and suggesting a response associated with trophic specialization of the involved organisms as much as with their trophic level.     

Edge-Core Effects and the Effectiveness of Traditional Reserves in Conservation: Eurasian Badgers in Doñana National Park

Abstract: High rates of human-mediated mortality on animals are frequently reported inside protected areas, especially near their borders, so the overall effect of reserves on animal conservation is not usually known. To determine the effect of a traditional reserve (Doñana National Park, southwestern Spain) on the Eurasian badger (   Meles meles ), a medium sized carnivore, we studied mortality causes and rates, with the aid of radiotelemetry, in two populations inside the reserve, one closer to the border than the other. Human-related mortality ( poaching and road kills) was by far the most common cause of mortality (85% of deaths recorded). The average annual mortality rate due to poaching was high (0.48 ± 0.08) for the population close to the border of the park, whereas none of the radiomarked badgers in the core of the reserve died during the study period. A logistic model that included distance from the border of the park, sampling effort, and the local area (i.e., edge and core populations) indicated that the difference between both sites was due to the effect of distance from the border on survival probability. On a regional scale, badger density at the core of the park was 3.16 times higher than outside. Overall, the effect of the reserve was positive, but edge effects reduced reserve effectiveness by 36%. Edge effects in close proximity of the border were strong, reducing badger density even below the expected density outside the reserve. Edge effects should be considered carefully when reserves are implemented because they can greatly reduce reserve effectiveness and influence the viability of the populations inside. Enlargement of reserves and control of human activities that promote edge effects, both inside and outside the reserve, are the two management actions that can most effectively mitigate edge effects.     

Contribution of Roads to Forest Fragmentation in the Rocky Mountains

The contribution of roads to forest fragmentation has not been adequately analyzed. We quantified fragmentation due to roads in a 30,213-ha section of the Medicine Bow-Routt National Forest in sout heastern Wyoming with several indices of landscape structure using a geographic information system. The number of patches, mean patch area, mean interior area, mean area of edge influence, mean patch perimeter, total perimeter, and mean patch shape identified patch- and edge-related landscape changes. Shannon-Wiener diversity, dominance, contagion, contrast, and angular second moment indicated effects on landscape diversity and texture. Roads added to forest fragmentation more than clearcuts by dissecting large patches into smaller pieces and by converting forest interior habitat into edge habitat. Edge habitat created by roads was 1.54–1.98 times the edge habitat created by clearcuts. The total landscape area affected by clearcuts and roads was 2.5–3.5 times the actual area occupied by these disturbances. Fragmentation due to roads could be minimized if road construction is minimized or rerouted so that its fragmentation effects are reduced. Geographic information system technology can be used to quantify the potential fragmentation effects of individual roads and the cumulative effects of a road network on landscape structure.     

Planning for robust reserve networks using uncertainty analysis

Planning land-use for biodiversity conservation frequently involves computer-assisted reserve selection algorithms. Typically such algorithms operate on matrices of species presence–absence in sites, or on species-specific distributions of model predicted probabilities of occurrence in grid cells. There are practically always errors in input data—erroneous species presence–absence data, structural and parametric uncertainty in predictive habitat models, and lack of correspondence between temporal presence and long-run persistence. Despite these uncertainties, typical reserve selection methods proceed as if there is no uncertainty in the data or models. Having two conservation options of apparently equal biological value, one would prefer the option whose value is relatively insensitive to errors in planning inputs. In this work we show how uncertainty analysis for reserve planning can be implemented within a framework of information-gap decision theory, generating reserve designs that are robust to uncertainty. Consideration of uncertainty involves modifications to the typical objective functions used in reserve selection. Search for robust-optimal reserve structures can still be implemented via typical reserve selection optimization techniques, including stepwise heuristics, integer-programming and stochastic global search.     

Catastrophe management and inter-reserve distance for marine reserve networks

We consider the optimal spacing between marine reserves for maximising the viability of a species occupying a reserve network. The closer the networks are placed together, the higher the probability of colonisation of an empty reserve by an occupied reserve, thus increasing population viability. However, the closer the networks are placed together, the higher the probability that a catastrophe will cause extinction of the species in both reserves, thus decreasing population viability. Using a simple discrete-time Markov chain model for the presence or absence of the species in each reserve we determine the distance between the two reserves which provides the optimal trade-off between these processes, resulting in maximum viability of the species.     

扎龙湿地丹顶鹤巢址空间分布变化及其对环境变化指征

扎龙国家级湿地保护区是我国一级保护鸟类丹顶鹤最重要的集中繁殖栖息地。以扎龙保护区为研究区,利用质心模型和最近邻体模型等方法,模拟1996至2004年间丹顶鹤（Grus japonensis）巢址的空间分布变化,分析这种变化对环境变化的指示作用。结果表明：丹顶鹤种群巢址分布出现向核心区集中的现象,鹤巢平均最近邻体距离大幅减少,巢址的空间分布格局由1996年的均匀分布转变为2004年的成群分布型;丹顶鹤巢址的空间分布变化表明该区环境发生巨大的变化：1996年核心区内栖息资源相对均匀、丰富,而缓冲区和实验区适宜生境呈零星分布格局,此后生境质量经历了由适宜一非适宜,适宜生境空间分布由均匀一斑块化的过程,到2004年各功能区适宜生境数量大幅减少,且核心区内适宜生境斑块化严重。为研究区丹顶鹤种群及其生境的保护和管理提供参考。     

Edge Effects on the Understory Bird Community in a Logged Forest in Uganda

Abstract: Understanding how the fauna of logged tropical rainforests responds to fragmentation and the creation of edges is vital to ensure conservation of biodiversity. We studied the composition of the understory bird community from the edge of a 15-ha clearing toward the interior of the forest in a part of Budongo Forest Reserve, Uganda, that was selectively logged about 45 years ago. Mist netting was conducted along five transects from the edge and 500 m into the interior. The total number of individuals captured did not change with distance from the edge, but there was a significant increase in the number of species. We sampled fewer, but more common species near the edge, whereas the interior of the forest had more, and less common species. Guild composition also changed with distance from the edge. Frugivore-insectivores and nectarivores were most common close to the edge. Among insectivores, ground foragers, bark-gleaners, and leaf-gleaners were most common in the interior of the forest, whereas sallying insectivores favored the edge. Graminivores were unaffected by the edge. Analysis of common species showed that Ispidina picta , Andropadus curvirostris , A. latirostris , Camaroptera brachyura , Terpsiphone rufiventer , and Nectarinia olivacea were associated with the edge, but no species showed significant avoidance of the edge. This finding may be explained by the generally low sample sizes of interior species. Our results show that even bird communities in logged forests respond to edges. Estimates of edge effects suggested that changes in bird densities may have occurred several hundred meters from the edge. In conclusion, logged forests provide habitat for bird species avoiding forest edges, and this should be considered in the management of such forests for conservation.     

Relative Importance of the Area and Shape of Patches to the Diversity of Multiple Taxa

Abstract: Although enhancing reserve shape has been suggested as an alternative to enlarging nature reserves, the importance of reserve shape relative to reserve area remains unclear. Here we examined the relative importance of area and shape of forest patches to species richness, species composition, and species abundance (abundance of each species) for 3 taxa (33 birds, 41 butterflies, and 91 forest‐floor plants) in a fragmented landscape in central Hokkaido, northern Japan. We grouped the species according to their potential edge responses (interior‐, neutral‐, and edge‐species groups for birds and forest‐floor plants, woodland‐ and open‐land‐species groups for butterflies) and analyzed them separately. We used a shape index that was independent of area as an index of shape circularization. Hierarchical partitioning and variation partitioning revealed that patch area was generally more important than patch shape for species richness and species composition of birds and butterflies. For forest‐floor plants, effects of patch area and shape were small, whereas effects of local forest structure were large. Patch area and circularization generally increased abundances of interior species of birds and forest‐floor plants and woodland species of butterflies. Nevertheless, only patch circularization increased abundances of 1 woodland species of butterfly and 2 and 6 interior species of birds and forest‐floor plants, respectively. We did not find any significant interaction effects between patch area and shape. Our results suggest that although reserves generally should be large and circular, there is a trade‐off between patch area and shape, which should be taken into consideration when managing reserves.