Biological Connectivity of Seasonally Ponded Wetlands across Spatial and Temporal Scales

Many species that inhabit seasonally ponded wetlands also rely on surrounding upland habitats and nearby aquatic ecosystems for resources to support life stages and to maintain viable populations. Understanding biological connectivity among these habitats is critical to ensure that landscapes are protected at appropriate scales to conserve species and ecosystem function. Biological connectivity occurs across a range of spatial and temporal scales. For example, at annual time scales many organisms move between seasonal wetlands and adjacent terrestrial habitats as they undergo life‐stage transitions; at generational time scales, individuals may disperse among nearby wetlands; and at multigenerational scales, there can be gene flow across large portions of a species’ range. The scale of biological connectivity may also vary among species. Larger bodied or more vagile species can connect a matrix of seasonally ponded wetlands, streams, lakes, and surrounding terrestrial habitats on a seasonal or annual basis. Measuring biological connectivity at different spatial and temporal scales remains a challenge. Here we review environmental and biological factors that drive biological connectivity, discuss implications of biological connectivity for animal populations and ecosystem processes, and provide examples illustrating the range of spatial and temporal scales across which biological connectivity occurs in seasonal wetlands.     

Biota Connect Aquatic Habitats throughout Freshwater Ecosystem Mosaics

Freshwater ecosystems are linked at various spatial and temporal scales by movements of biota adapted to life in water. We review the literature on movements of aquatic organisms that connect different types of freshwater habitats, focusing on linkages from streams and wetlands to downstream waters. Here, streams, wetlands, rivers, lakes, ponds, and other freshwater habitats are viewed as dynamic freshwater ecosystem mosaics (FEMs) that collectively provide the resources needed to sustain aquatic life. Based on existing evidence, it is clear that biotic linkages throughout FEMs have important consequences for biological integrity and biodiversity. All aquatic organisms move within and among FEM components, but differ in the mode, frequency, distance, and timing of their movements. These movements allow biota to recolonize habitats, avoid inbreeding, escape stressors, locate mates, and acquire resources. Cumulatively, these individual movements connect populations within and among FEMs and contribute to local and regional diversity, resilience to disturbance, and persistence of aquatic species in the face of environmental change. Thus, the biological connections established by movement of biota among streams, wetlands, and downstream waters are critical to the ecological integrity of these systems. Future research will help advance our understanding of the movements that link FEMs and their cumulative effects on downstream waters.     

A Review of Habitat Connectivity Research for Pacific Salmon in Marine,Estuary, and Freshwater Environments

Long‐term conservation planning for diadromous fishes would benefit from a better understanding of both the role of connectivity among environments and habitat variability in the expression of life‐history diversity. Most of the scientific knowledge on habitat fragmentation and connectivity has been developed in terrestrial systems in the discipline of landscape ecology. Research on habitat connectivity in aquatic systems (e.g., salmonid research that spans the spectrum of habitats from freshwater to the sea) is uncommon and largely focused on barriers to fish passage. Here, we present a review of the literature characterizing current research patterns on habitat connectivity within and among environments for Pacific salmon. We found this topic is still incipient: the literature is dominated by studies of freshwaters, with few articles focusing on habitat needs in estuary and marine systems. Pan‐environment studies are rare, pointing to a gap in our understanding of complex habitat relationships that might be significant in the development of long‐term conservation and restoration plans for Pacific salmon, particularly in light of the potential impact of climate change.     

Hydrologic Connectivity and the Contribution of Stream Headwaters to Ecological Integrity at Regional Scales1

Abstract: Cumulatively, headwater streams contribute to maintaining hydrologic connectivity and ecosystem integrity at regional scales. Hydrologic connectivity is the water‐mediated transport of matter, energy and organisms within or between elements of the hydrologic cycle. Headwater streams compose over two‐thirds of total stream length in a typical river drainage and directly connect the upland and riparian landscape to the rest of the stream ecosystem. Altering headwater streams, e.g., by channelization, diversion through pipes, impoundment and burial, modifies fluxes between uplands and downstream river segments and eliminates distinctive habitats. The large‐scale ecological effects of altering headwaters are amplified by land uses that alter runoff and nutrient loads to streams, and by widespread dam construction on larger rivers (which frequently leaves free‐flowing upstream portions of river systems essential to sustaining aquatic biodiversity). We discuss three examples of large‐scale consequences of cumulative headwater alteration. Downstream eutrophication and coastal hypoxia result, in part, from agricultural practices that alter headwaters and wetlands while increasing nutrient runoff. Extensive headwater alteration is also expected to lower secondary productivity of river systems by reducing stream‐system length and trophic subsidies to downstream river segments, affecting aquatic communities and terrestrial wildlife that utilize aquatic resources. Reduced viability of freshwater biota may occur with cumulative headwater alteration, including for species that occupy a range of stream sizes but for which headwater streams diversify the network of interconnected populations or enhance survival for particular life stages. Developing a more predictive understanding of ecological patterns that may emerge on regional scales as a result of headwater alterations will require studies focused on components and pathways that connect headwaters to river, coastal and terrestrial ecosystems. Linkages between headwaters and downstream ecosystems cannot be discounted when addressing large‐scale issues such as hypoxia in the Gulf of Mexico and global losses of biodiversity.     

Landscape-level ecological regions: Linking state-level ecoregion frameworks with stream habitat classifications

Regionalization is a form of spatial classification, where boundaries are drawn around areas that are relatively homogeneous in landscape characteristics. The process of delineating ecological regions, or ecoregions, includes the analysis of ecosystem structure. To date, ecoregions have been developed at national and state scales for research and resource management. Stream classification is another method to order the variability of aquatic habitats that spans spatial scales from microhabitat to valley segment. In this study, landscape-level ecoregions are developed for the upper Grande Ronde River basin in northeastern Oregon, 3000 sq km in area. The ecoregion framework presented here is proposed to bridge the gap between stream habitat and state-level ecoregion classifications. Classification at this scale is meant to address issues of management at local scales: to aid in sampling design, in extrapolation of the results of site-specific studies, and in the development of best management practices that are more predictive of ecosystem response than current methods.     

Assessing the Potential for Salmon Recovery via Floodplain Restoration: A Multitrophic Level Comparison of Dredge-Mined to Reference Segments

Pre-restoration studies typically focus on physical habitat, rather than the food-base that supports aquatic species. However, both food and habitat are necessary to support the species that habitat restoration is frequently aimed at recovering. Here we evaluate if and how the productivity of the food-base that supports fish production is impaired in a dredge-mined floodplain within the Yankee Fork Salmon River (YFSR), Idaho (USA); a site where past restoration has occurred and where more has been proposed to help recover anadromous salmonids. Utilizing an ecosystem approach, we found that the dredged segment had comparable terrestrial leaf and invertebrate inputs, aquatic primary producer biomass, and production of aquatic invertebrates relative to five reference floodplains. Thus, the food-base in the dredged segment did not necessarily appear impaired. On the other hand, we observed that off-channel aquatic habitats were frequently important to productivity in reference floodplains, and the connection of these habitats in the dredged segment via previous restoration increased invertebrate productivity by 58%. However, using a simple bioenergetic model, we estimated that the invertebrate food-base was at least 4× larger than present demand for food by fish in dredged and reference segments. In the context of salmon recovery efforts, this observation questions whether additional food-base productivity provided by further habitat restoration would be warranted in the YFSR. Together, our findings highlight the importance of studies that assess the aquatic food-base, and emphasize the need for more robust ecosystem models that evaluate factors potentially limiting fish populations that are the target of restoration.     

Basic Principles and Ecological Consequences of Altered Flow Regimes for Aquatic Biodiversity

The flow regime is regarded by many aquatic ecologists to be the key driver of river and floodplain wetland ecosystems. We have focused this literature review around four key principles to highlight the important mechanisms that link hydrology and aquatic biodiversity and to illustrate the consequent impacts of altered flow regimes: Firstly, flow is a major determinant of physical habitat in streams, which in turn is a major determinant of biotic composition; Secondly, aquatic species have evolved life history strategies primarily in direct response to the natural flow regimes; Thirdly, maintenance of natural patterns of longitudinal and lateral connectivity is essential to the viability of populations of many riverine species; Finally, the invasion and success of exotic and introduced species in rivers is facilitated by the alteration of flow regimes. The impacts of flow change are manifest across broad taxonomic groups including riverine plants, invertebrates, and fish. Despite growing recognition of these relationships, ecologists still struggle to predict and quantify biotic responses to altered flow regimes. One obvious difficulty is the ability to distinguish the direct effects of modified flow regimes from impacts associated with land-use change that often accompanies water resource development. Currently, evidence about how rivers function in relation to flow regime and the flows that aquatic organisms need exists largely as a series of untested hypotheses. To overcome these problems, aquatic science needs to move quickly into a manipulative or experimental phase, preferably with the aims of restoration and measuring ecosystem response.     

FORUM: Restoration of Stream Habitats in the Western United States: Restoration as Reexpression of Habitat Capacity

/ Ecological restoration is increasingly invoked as a tool for the maintenance and regeneration of biodiversity. Yet the conceptual foundations and assumptions underlying many restoration management activities are frequently unclear or unstated. Unforeseen, undesirable consequences of restoration activities may emerge as a result. A general conceptual framework for restoration is needed to better accommodate dynamic habitat systems and evolving biota in restoration strategies. A preliminary framework for stream habitat restoration emphasizing stream habitat-biota development is proposed. As developing systems, streams and stream biota exhibit temporal behaviors that change with stream environments. Underlying the dynamic development of streams is potential capacity. Streams express this capacity as an array of habitats over time and across the landscape. Human land uses in the western United States have rapidly altered aquatic habitats and the processes that shape habitat. As a result, the diversity of native fishes and their habitats has been suppressed. Restoration is fundamentally about allowing stream systems to reexpress their capacities. Several steps are provided to guide stream restoration activities. Key tasks include: identification of the historic patterns of habitat development; identification of developmental constraints; relief of those constraints; classification of sensitive, critical, or refuge habitats; protection of the developmental diversity that remains; and monitoring of biotic responses to habitat development. KEY WORDS: Stream habitat; Stream biota; System capacity; System development; Restoration; Classification     

Investigating habitat value to inform contaminant remediation options: case study

Habitat valuation methods were implemented to support remedial decisions for aquatic and terrestrial contaminated sites at the East Tennessee Technology Park (ETTP) on the US Department of Energy (DOE) Oak Ridge Reservation in Oak Ridge, TN, USA. The habitat valuation was undertaken for six contaminated sites: Contractor's Spoil Area, K-901-N Disposal Area, K-770 Scrapyard, K-1007-P1 pond, K-901 pond, and the Mitchell Branch stream. Four of these sites are within the industrial use area of ETTP and two are in the Black Oak Ridge Conservation Easement. These sites represent terrestrial and aquatic habitat for vertebrates, terrestrial habitat for plants, and aquatic habitat for benthic invertebrates. Current and potential future, no-action (no remediation) scenarios were evaluated primarily using existing information. Valuation metrics and scoring criteria were developed in a companion paper, this volume. The habitat valuation consists of extensive narratives, as well as scores for aspects of site use value, site rarity, and use value added from spatial context. Metrics for habitat value were expressed with respect to different spatial scales, depending on data availability. There was significant variation in habitat value among the six sites, among measures for different taxa at a single site, between measures of use and rarity at a single site, and among measures for particular taxa at a single site with respect to different spatial scales. Most sites had aspects of low, medium, and high habitat value. Few high scores for current use value were given. These include: wetland plant communities at all aquatic sites, Lepomid sunfish and waterbirds at 1007-P1 pond, and Lepomid sunfish and amphibians at K-901 pond. Aquatic sites create a high-value ecological corridor for waterbirds, and the Contractor's Spoil Area and possibly the K-901-N Disposal Site have areas that are part of a strong terrestrial ecological corridor. The only example of recent observations of rare species at these sites is the gray bat observed at the K-1007-P1 pond. Some aspects of habitat value are expected to improve under no-action scenarios at a few of the sites. Methods are applicable to other contaminated sites where sufficient ecological data are available for the site and region.     

Prioritizing Stream Barrier Removal to Maximize Connected Aquatic Habitat and Minimize Water Scarcity

Instream barriers, such as dams, culverts, and diversions, alter hydrologic processes and aquatic habitat. Removing uneconomical and aging instream barriers is increasingly used for river restoration. Historically, selection of barrier removal projects used score‐and‐rank techniques, ignoring cumulative change and the spatial structure of stream networks. Likewise, most water supply models prioritize either human water uses or aquatic habitat, failing to incorporate both human and environmental water use benefits. Here, a dual‐objective optimization model identifies barriers to remove that maximize connected aquatic habitat and minimize water scarcity. Aquatic habitat is measured using monthly average streamflow, temperature, channel gradient, and geomorphic condition as indicators of aquatic habitat suitability. Water scarcity costs are minimized using economic penalty functions while a budget constraint specifies the money available to remove barriers. We demonstrate the approach using a case study in Utah's Weber Basin to prioritize removal of instream barriers for Bonneville cutthroat trout, while maintaining human water uses. Removing 54 instream barriers reconnects about 160 km of quality‐weighted habitat and costs approximately US$10 M. After this point, the cost‐effectiveness of removing barriers to connect river habitat decreases. The modeling approach expands barrier removal optimization methods by explicitly including both economic and environmental water uses.     

Safeguarding Wetlands and Their Connections within Wetlandscapes to Improve Conservation Outcomes for Threatened Amphibian Species

Wetlands should not be considered as independent objects but as dynamically connected objects, collectively known as wetlandscapes. We developed a framework that analyzes the influences of wetland suitability and connectivity on amphibian distributions. We defined two indices: a Wetland Suitability Index describing wetland quality and a Movement Permeability Index characterizing wetland connectivity for amphibian population dynamics. These indices were calculated from raster datasets and time‐varying inundation estimates. The indices were used to define a wetlandscape and an amphibian model was used to simulate population dynamics within the wetlandscape. The framework was applied to the Nose Creek watershed, a highly modified wetlandscape in Alberta, Canada. Two amphibian species were selected with different habitat preferences: the Northern Leopard Frog that prefers wet habitats and has high mobility over land, and the Great Plains Toad that prefers terrestrial habitats and has low mobility over land. We found each amphibian species had a “preferred” wetlandscape, reflecting their life cycle traits and migration strategies which in turn were dependent on the hydrological and ecological connections within the wetlandscape. This study highlights the importance of investigating both individual wetlands and the wetlandscape and considering both wetland habitat quality and connectivity as non‐substitutable properties that act jointly, but differently, on population dynamics.     

Patterns of biodiversity and habitat sensitivity in agricultural landscapes

Design of landscape is the process of the arrangement of spatial features with the objective of sustaining ecosystem services, and maintaining ecological functionality to meet societal needs. Along a gradient of cultivation intensity, the functional quality of agricultural landscape was explored and the relationships between landscape metrics and functional quality were analyzed, in order to make effective recommendations for landscape design aimed at sustainable land use schemes. The functional quality of landscape was calculated using the InVest model for 20 farm landscapes (North-Eastern Italy) where biodiversity (plant taxa) and sensitivity to disturbance (hemeroby) were used as model inputs. Results highlighted the importance of specific habitat types such as meadows and woodlands rather than other habitats for improving the biodiversity of agricultural landscapes. A high proportion of these habitats enhanced the functional quality of the landscape when the habitats were organized in large and not isolated patches in heterogeneous landscapes.     

Lakes,Wetlands, and Streams as Predictors of Land Use/Cover Distribution

The importance of the surrounding landscape to aquatic ecosystems has been well established. Most research linking aquatic ecosystems to landscapes has focused on the one-way effect of land on water. However, to understand fully the complex interactions between aquatic and terrestrial ecosystems, aquatic ecosystems must be seen not only as receptors of human modification of the landscape, but also as potential drivers of these modifications. We hypothesized that the presence of aquatic ecosystems influences the spatial distribution of human land use/cover of the nearby landscape (≤1 km) and that this influence has changed through time from the 1930s to the 1990s. To test this hypothesis, we compared the distribution of residential, agricultural, and forested land use/cover around aquatic ecosystems (lakes, wetlands, and streams) to the overall regional land use/cover proportion in an area in southeast Michigan, USA; we also compared the distribution of land use/cover around county roads/highway and towns (known determinants of many land use/cover patterns) to the regional proportion. We found that lakes, wetlands, and streams were strongly associated with the distribution of land use/cover, that each ecosystem type showed different patterns, and that the magnitude of the association was at least as strong as the association with human features. We also found that the area closest to aquatic ecosystems (<500 m) was more strongly associated with land use/cover distribution than areas further away. Finally, we found that the strength of the association between aquatic ecosystems and land use/cover increased from 1938 to 1995, although the overall patterns were similar through time. Our results show that a more complete understanding is needed of the role of aquatic ecosystems on the distribution of land use/cover.     

Correspondence between spatial patterns in fish assemblages in ohio streams and aquatic ecoregions

Land classification systems can be useful for assessing aquatic ecosystems if relationships among them exist. Because the character of an aquatic ecosystem depends to a large extent upon the character of the landscape it drains, spatial patterns in aquatic ecosystems should correspond to patterns in the landscape. To test this hypothesis, the US state of Ohio was divided into four aquatic ecoregions based on an analysis of spatial patterns in the combination of land-surface form, land use, potential natural vegetation, and soil parent material. During the period July–October 1983, fish assemblages were sampled in 46 streams that were representative of the ecoregions, and that had watersheds relatively undisturbed by human activities. Spatial patterns of the fish assemblages were examined relative to the ecoregions; distinct regional differences were identified. The assemblages differed most between the Huron/Erie Lake Plain region and the Western Allegheny Plateau region; assemblages in the Eastern Corn Belt Plains and the Erie/Ontario Lake Plain-Interior Plateau regions were intermediate. This pattern also reflects the gradient in landscape character as one moves from the northwest to the southeast of Ohio.     

Identification of Putative Geographically Isolated Wetlands of the Conterminous United States

Geographically isolated wetlands (GIWs) are wetlands completely surrounded by uplands. While common throughout the United States (U.S.), there have heretofore been no nationally available, spatially explicit estimates of GIW extent, complicating efforts to understand the myriad biogeochemical, hydrological, and habitat functions of GIWs and hampering conservation and management efforts at local, state, and national scales. We used a 10‐m geospatial buffer as a proxy for hydrological or ecological connectivity of National Wetlands Inventory palustrine and lacustrine wetland systems to nationally mapped and available stream, river, and lake data. We identified over 8.3 million putative GIWs across the conterminous U.S., encompassing nearly 6.5 million hectares of wetland resources (average size 0.79 ± 4.81 ha, median size 0.19 ha). Putative GIWs thus represent approximately 16% of the freshwater wetlands of the conterminous U.S. The water regime for the majority of the putative GIWs was temporarily or seasonally flooded, suggesting a vulnerability to ditching or hydrologic abstraction, sedimentation, or alterations in precipitation patterns. Additional analytical applications of this readily available geospatially explicit mapping product (e.g., hydrological modeling, amphibian metapopulation, or landscape ecological analyses) will improve our understanding of the abundance and extent, effect, connectivity, and relative importance of GIWs to other aquatic systems of the conterminous U.S.     

城市湿地生态系统的结构特征及现存问题

对我国城市湿地生态系统的结构特征和存在问题进行了综述。目前我国城市湿地生态系统的结构特征表现为面积小、分布不均、空间结构和营养结构简单、稳定性低、脆弱性强,同时其形态结构具有显著的景观美学特征,但受水环境因素的制约性强。目前的主要问题是：湿地面积迅速减少、空间结构脆弱性加剧、环境污染加剧、生物多样性降低、生物入侵严重、生态系统稳定性下降、水土流失与湖泊淤积严重、规划设计不当、人为干预强烈。通过综述,将增加人们对城市湿地生态系统结构的深入认识,为开展科学保护、建设、开发、利用活动和促进城市可持续发展提供依据。     

Effects of Habitat Disturbance from Residential Development on Breeding Bird Communities in Riparian Corridors

This study assessed the relationship among land use, riparian vegetation, and avian populations at two spatial scales. Our objective was to compare the vegetated habitat in riparian corridors with breeding bird guilds in eight Rhode Island subwatersheds along a range of increasing residential land use. Riparian habitats were characterized with fine-scale techniques (used field transects to measure riparian vegetation structure and plant species richness) at the reach spatial scale, and with coarse-scale landscape techniques (a Geographic Information System to document land-cover attributes) at the subwatershed scale. Bird surveys were conducted in the riparian zone, and the observed bird species were separated into guilds based on tolerance to human disturbance, habitat preference, foraging type, and diet preference. Bird guilds were correlated with riparian vegetation metrics, percent impervious surface, and percent residential land use, revealing patterns of breeding bird distribution. The number of intolerant species predominated below 12% residential development and 3% impervious surface, whereas tolerant species predominated above these levels. Habitat guilds of edge, forest, and wetland bird species correlated with riparian vegetation. This study showed that the application of avian guilds at both stream reach and subwatershed scales offers a comprehensive assessment of effects from disturbed habitat, but that the subwatershed scale is a more efficient method of evaluation for environmental management.     

Issues and approaches in assessing cumulative impacts on waterbird habitat in wetlands

Wetlands are attractive to vertebrates because of their abundant nutrient resources and habitat diversity. Because they are conspicuous, vertebrates commonly are used as indicators of changes in wetlands produced by environmental impacts. Such impacts take place at the landscape level where extensive areas are lost; at the wetland complex level where some (usually small) units of a closely spaced group of wetlands are drained or modified; or at the level of the individual wetland through modification or fragmentation that impacts its habitat value. Vertebrates utilize habitats differently according to age, sex, geographic location, and season, and habitat evaluations based on isolated observations can be biased. Current wetland evaluation systems incorporate wildlife habitat as a major feature, and the habitat evaluation procedure focuses only on habitat. Several approaches for estimating bird habitat losses are derived from population curves based on natural and experimentally induced population fluctuations. Additional research needs and experimental approaches are identified for addressing cumulative impacts on wildlife habitat values.     

Testing hypotheses about management to enhance habitat for feeding birds in a freshwater wetland

The level of water was manipulated in a freshwater wetland, with the aim of enhancing abundances of benthic animals and, ultimately, improving habitat for feeding birds (Japanese Snipe, Gallinago hardwickii). We tested whether these actions had the predicted and desired effects on benthic animals, by contrasting changes in two managed locations to one control location which was left unmanipulated. The number of taxa and abundances of chironomids decreased strongly and significantly in the manipulated locations, while the abundance of oligochaetes appeared to vary in a seasonal manner. Temporal variability of the structure and composition of assemblages was also increased in manipulated locations. Such effects have previously been suggested to indicate stress in benthic assemblages. Therefore, in contrast to what was predicted, managerial actions made benthic fauna less abundant and thus, less suitable as habitat for feeding birds. Several general lessons can be learned from these results. (1) Effects of managerial actions like these are difficult to predict a priori and can only be reliably evaluated with an experimental framework. (2) Because abundances of animals vary naturally, evaluations of managerial actions must include appropriate spatial replication. (3) Sampling at hierarchical temporal scales is important, because abundances of animals may vary in an unpredictable manner at short temporal scales and because changes in temporal variability may be a symptom of stress. (4) Combined use of uni- and multivariate techniques provides a comprehensive set of tools to assess the effects of restoration and creation of new habitats. Finally, these results emphasise the need for clear predictions about desired outcomes and specific experimental plans about how to test whether the desired results were achieved, before managerial actions are taken. Although this is often very difficult to achieve in real situations, it is necessary for practices of management to evolve on the basis of sound empirical experience.     

ENVIRONMENTAL DESIGNS FOR STREAMBANK PROTECTION PROJECTS1

ABSTRACT: Streambank protection projects are intended to prevent streambank erosion, thereby preventing streambank failure and maintaining a desirable channel alignment. Streambank erosion is a natural process of unaltered, dynamic river systems, and protection projects seek to impose stability on this natural system. The environmental impacts of such projects are primarily changes to terrestrial and aquatic habitats and to aesthetics. Adverse environmental impacts have been minimized and enhancement of existing habitat and aesthetics have been achieved through the development of new, innovative designs or modifications to existing designs and through use of construction and maintenance practices that promote habitat and aesthetics. Designs based on channel flow characteristics, e.g., revetments using a variety of structural materials, can result in preservation of wildlife habitat by reducing the use of structural protection by matching the erosion potential of flow at the bank with the protection capability of the materials used. Designs based on streambed stabilization prevent bank failure caused by bank undermining, result in preservation or establishment of streamside vegetation, and enhance aesthetics. Protection schemes that manage and preserve floodplains, berms, and riparian areas preserve the natural condition of the floodplain area. Designs based on deflection of erosive flows, e.g., dikes, minimize disturbance to the bank vegetation and create low-velocity aquatic habitats. Use of vegetation for bank protection is most effective when used in combination with structural components. Construction and maintenance practices can be scheduled and modified to minimize impacts to floodplain areas and to enhance wildlife habitat while preserving the integrity of the protection structure.