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Application of the new keystone-species concept to prairie dogs: How well does it work?

It has been suggested that the keystone-species concept should be dropped from ecology and conservation, primarily because the concept is poorly defined. This prompted Power et al. (1996) to refine the definition: keystone species have large effects on community structure or ecosystem function (i.e., high overall importance), and this effect should be large relative to abundance (i.e., high community importance). Using prairie dogs ( Cynomys spp.) as an example, I review operational and conceptual difficulties encountered in applying this definition. As applied to prairie dogs, the implicit assumption that overall importance is a linear function of abundance is invalid. In addition, community importance is sensitive to abundance levels, the definition of community, and sampling scale. These problems arise largely from the equation for community importance, as used in conjunction with removal experiments at single abundance levels. I suggest that we shift from the current emphasis on the dualism between keystone and nonkeystone species and instead examine how overall and community importance vary (1) with abundance, (2) across spatial and temporal scales, and (3) under diverse ecological conditions. In addition, I propose that a third criterion be incorporated into the definition: keystone species perform roles not performed by other species or processes. Examination of how these factors vary among populations of keystone species should help identify the factors contributing to, or limiting, keystone-level functions, thereby increasing the usefulness of the keystone-species concept in ecology and conservation. Although the quantitative framework of Power et al. falls short of being fully operational, my conceptual guidelines may improve the usefulness of the keystone-species concept. Careful attention to the factors that limit keystone function will help avoid misplaced emphasis on keystone species at the expense of other species.

Conservation Biology

Social network models predict movement and connectivity in ecological landscapes

Network analysis is on the rise across scientific disciplines because of its ability to reveal complex, and often emergent, patterns and dynamics. Nonetheless, a growing concern in network analysis is the use of limited data for constructing networks. This concern is strikingly relevant to ecology and conservation biology, where network analysis is used to infer connectivity across landscapes. In this context, movement among patches is the crucial parameter for interpreting connectivity but because of the difficulty of collecting reliable movement data, most network analysis proceeds with only indirect information on movement across landscapes rather than using observed movement to construct networks. Statistical models developed for social networks provide promising alternatives for landscape network construction because they can leverage limited movement information to predict linkages. Using two mark-recapture datasets on individual movement and connectivity across landscapes, we test whether commonly used network constructions for interpreting connectivity can predict actual linkages and network structure, and we contrast these approaches to social network models. We find that currently applied network constructions for assessing connectivity consistently, and substantially, overpredict actual connectivity, resulting in considerable overestimation of metapopulation lifetime. Furthermore, social network models provide accurate predictions of network structure, and can do so with remarkably limited data on movement. Social network models offer a flexible and powerful way for not only understanding the factors influencing connectivity but also for providing more reliable estimates of connectivity and metapopulation persistence in the face of limited data.

Proceedings of the National Academy of Sciences of

Discontinuities concentrate mobile predators: Quantifying organism-environment interactions at a seascape scale

Understanding environmental drivers of spatial patterns is an enduring ecological problem that is critical for effective biological conservation. Discontinuities (ecologically meaningful habitat breaks), both naturally occurring (e.g., river confluence, forest edge, drop-off) and anthropogenic (e.g., dams, roads), can influence the distribution of highly mobile organisms that have land- or seascape scale ranges. A geomorphic discontinuity framework, expanded to include ecological patterns, provides a way to incorporate important but irregularly distributed physical features into organism–environment relationships. Here, we test if migratory striped bass ( Morone saxatilis ) are consistently concentrated by spatial discontinuities and why. We quantified the distribution of 50 acoustically tagged striped bass at 40 sites within Plum Island Estuary, Massachusetts during four-monthly surveys relative to four physical discontinuities (sandbar, confluence, channel network, drop-off), one continuous physical feature (depth variation), and a geographic location variable (region). Despite moving throughout the estuary, striped bass were consistently clustered in the middle geographic region at sites with high sandbar area, close to channel networks, adjacent to complex confluences, with intermediate levels of bottom unevenness, and medium sized drop-offs. In addition, the highest striped bass concentrations occurred at sites with the greatest additive physical heterogeneity (i.e., where multiple discontinuities co-occurred). The need to incorporate irregularly distributed features in organism–environment relationships will increase as high-quality telemetry and GIS data accumulate for mobile organisms. The spatially explicit approach we used to address this challenge can aid both researchers who seek to understand the impact of predators on ecosystems and resource managers who require new approaches for biological conservation.

Ecosphere

The diamondback terrapin: The biology, ecology, cultural history, and conservation status of an obligate estuarine turtle

Ranging from Cape Cod to nearly the Texas-Mexico border, the diamondback terrapin (Malaclemys terrapin) is the only species of North American turtle restricted to estuarine systems. Despite this extensive distribution, its zone of occurrence is very linear, and in places fragmented, resulting in a relatively small total area of occupancy. On a global scale, excluding marine species, few turtles even venture into brackish water on a regular basis, and only two Asian species approach the North American terrapin's dependency on estuarine habitats. Here we describe some of the biological and behavioral adaptations of terrapins that allow them to live in the rather harsh estuarine environment. In this chapter we review the natural and cultural history of this turtle, discuss conservation issues, and provide information on the types of research needed to make sound management decisions for terrapin populations in peril.

Studies in Avian Biology

Towards an urgent yet deliberate conservation strategy: Sustaining social-ecological systems in rangelands of the Northern Great Plains, Montana

Urgency and deliberateness are often at odds when executing conservation projects, especially as the scale and complexity of objectives increases. The pace of environmental degradation supports immediate and measurable action. However, best practices for adaptive governance and building resilient social-ecological systems call for more deliberate efforts and participatory processes, which can be slow. We explore conflicts between urgency and deliberateness and the potential for their reconciliation through a case study of the challenges of conserving native rangelands in North America’s Northern Great Plains, an ecoregion targeted for global conservation initiatives. This region is undergoing a significant social-ecological transition, which underscores a need to rethink conservation strategies in light of the social-ecological system dynamics and potential future trajectories. Based on a structured narrative literature review process and iterative engagement with key regional stakeholders, we identify three interrelated factors critical to the system’s future outcomes that illustrate system complexity as well as trade-offs between urgent and deliberate action and unilateral and multilateral approaches to conservation: (1) influences of land management on biodiversity, (2) economic restructuring and shifting land use priorities, and (3) changing climate and disturbance regimes. We identify key gaps in the literature for each factor and across the factors—an effort that informs our call for research and practice agendas that address uncertainty and complexity at regional scales through more inclusive and future-oriented approaches.

Montana

Analysis of ecological context for identifying vegetation and animal conservation planning foci: An example from the arid South-western USA

In developing conservation strategies, it is important to maximize effects of conservation within a specified land tract and to maximize conservation effects on surrounding area (ecological context). The authors proposed two criteria to select biotic entities for conservation foci: (1) the relative occurrence of fauna or flora in a tract is greater than that of an ecological context region; and (2) occurrence of the fauna or flora is relatively limited in the ecological context region. Using extensive spatial data on vegetation and wildlife habitat distribution, the authors identified strategic vegetation and fauna conservation foci for the 400 000 ha Fort Bliss military reservation in New Mexico and Texas relative to a 164 km radius ecological context region intersecting seven ecological zones and the predicted habitat distribution of 616 animal species. The authors set two specific criteria: (1) predicted area of a species' occurrence is <50% of the ecological context region; and (2) percentage of Fort Bliss intersecting the species' or vegetation community predicted areas in the ecological context region is >5% (Fort Bliss is 4.2% of the region). These criteria selected one vegetation class and 40 animal species. Further, these vegetation and animal foci were primarily located in two areas of Fort Bliss. Sensitivity analyses with other analytical radii corroborated the context radius used. Conservation of the two areas and associated taxa will maximize the contribution of Fort Bliss's conservation efforts in its ecological proximity. This relatively simple but information-rich process represents economical and defensible preliminary contextual analysis for detailed conservation planning.

Journal of Environmental Planning and Management

The conservation genetics juggling act: Integrating genetics and ecology, science and policy

The field of conservation genetics, when properly implemented, is a constant juggling act integrating molecular genetics, ecology, and demography with applied aspects concerning managing declining species or implementing conservation laws and policies. This young field has grown substantially since the 1980&rsquo;s following development of the polymerase chain reaction and now into the genomics era. Our lab has &ldquo;grown up&rdquo; with the field, having worked on these issues for over three decades. Our multi-disciplinary approach entails understanding the behavior and ecology of species as well as the underlying processes that contribute to genetic viability. Taking this holistic approach provides a comprehensive understanding of factors that influence species persistence and evolutionary potential while considering annual challenges that occur throughout their life cycle. As a federal lab, we are often addressing the needs of the U.S. Fish and Wildlife Service in their efforts to list, de-list or recover species. Nevertheless, there remains an overall communication gap between research geneticists and biologists who are charged with implementing their results. Therefore, we outline the need for a National Center for Small Population Biology to ameliorate this problem and provide organizations charged with making status decisions firmer ground from which to make their critical decisions.

Evolutionary Applications

A mosaic-based approach is needed to conserve biodiversity in disturbed freshwater ecosystems

Conserving native biodiversity in the face of human‐ and climate‐related impacts is a challenging and globally important ecological problem that requires an understanding of spatially connected, organismal‐habitat relationships. Globally, a suite of disturbances (e.g., agriculture, urbanization, climate change) degrades habitats and threatens biodiversity. A mosaic approach (in which connected, interacting collections of juxtaposed habitat patches are examined) provides a scientific foundation for addressing many disturbance‐related, ecologically based conservation problems. For example, if specific habitat types disproportionately increase biodiversity, these keystones should be incorporated into research and management plans. Our sampling of fish biodiversity and aquatic habitat along ten 3‐km sites within the Upper Neosho River subdrainage, KS, from June‐August 2013 yielded three generalizable ecological insights. First, specific types of mesohabitat patches (i.e., pool, riffle, run, and glide) were physically distinct and created unique mosaics of mesohabitats that varied across sites. Second, species richness was higher in riffle mesohabitats when mesohabitat size reflected field availability. Furthermore, habitat mosaics that included more riffles had greater habitat diversity and more fish species. Thus, riffles (<5% of sampled area) acted as keystone habitats. Third, additional conceptual development, which we initiate here, can broaden the identification of keystone habitats across ecosystems and further operationalize this concept for research and conservation. Thus, adopting a mosaic approach can increase scientific understanding of organismal‐habitat relationships, maintain natural biodiversity, advance spatial ecology, and facilitate effective conservation of native biodiversity in human‐altered ecosystems.

Kansas

A review of supervised learning methods for classifying animal behavioural states from environmental features

Accurately predicting behavioural modes of animals in response to environmental features is important for ecology and conservation. Supervised learning (SL) methods are increasingly common in animal movement ecology for classifying behavioural modes. However, few examples exist of applying SL to classify polytomous animal behaviour from environmental features especially in the context of millions of animal observations. We review SL methods (weighted k -nearest neighbours; neural nets; random forests; and boosted classification trees with XGBoost) for classifying polytomous animal behaviour from environmental predictors. We also describe tuning parameter selection and assessment strategies, approaches for visualizing relationships between predictors and class outputs, and computational considerations. We demonstrate these methods by predicting three categories of risk to bald eagles from colliding with wind turbines using, as predictors, 12 environmental state features associated with 1.7 million GPS telemetry data points from 57 eagles. Of the SL methods we considered, XGBoost yielded the most accurate model with 86.2% classification accuracy and pairwise-averaged area under the ROC curve of 90.6. Computational time of XGBoost scaled better to large data than any other SL method. We also show how SHAP values integrated in the R package ( xgboost ) facilitate investigation of variable relationships and importance. For big data applications, XGBoost appears to provide superior classification accuracy and computational efficiency. Our results suggest XGBoost should be considered as an early modelling option in situations where the intent is to classify millions of animal behaviour observations from environmental predictors and to understand relationships between those predictors and movement behaviours. We also offer a tutorial to assist researchers in implementing this method.

Methods in Ecology and Evolution

Forecasting sea otter recolonization: Insights from isotopic analysis of modern and zooarchaeological populations

Retrospective datasets offer essential context for conservation by revealing species’ ecological roles before industrial-era human impacts. We analysed isotopic compositions of pre-industrial and modern sea otters ( Enhydra lutris ) to reconstruct pre-extirpation ecology and offer insights for management. Our study focuses on southeast Alaska (SEAK), where sea otters are recolonizing, and northern Oregon, where translocations are being considered. We measured bulk bone collagen δ 13 C and δ 15 N values and essential amino acid δ 13 C values of extirpated sea otters from archaeological contexts, and bulk isotopic values from vibrissae of modern SEAK sea otters. We compare these results with published isotopic data of potential prey and additional archaeological datasets. In SEAK, our data show pre-industrial sea otter populations consumed infaunal bivalves and used soft-sediment (33%) and kelp forest habitats (67%), with sub-regional variation. We anticipate current populations will expand into this historical niche, and conflict with regional traditional/subsistence bivalve fisheries will persist. In northern Oregon, isotopic data from extirpated sea otters indicate past consumption of low trophic level invertebrates and a stronger reliance on kelp forests (88%) rather than soft-sediment habitats, highlighting the importance of kelp forests for future translocations. Our work exemplifies the value of historical ecology in informing conservation strategies for recovering species.

Alaska, Oregon

Monitoring for conservation

Human-mediated environmental changes have resulted in appropriate concern for the conservation of ecological systems and have led to the development of many ecological monitoring programs worldwide. Many programs that are identified with the purpose of `surveillance? represent an inefficient use of conservation funds and effort. Here, we revisit the 1964 paper by Platt and argue that his recommendations about the conduct of science are equally relevant to the conduct of ecological monitoring programs. In particular, we argue that monitoring should not be viewed as a stand-alone activity, but instead as a component of a larger process of either conservation-oriented science or management. Corresponding changes in monitoring focus and design would lead to substantial increases in the efficiency and usefulness of monitoring results in conservation.

Trends in Ecology and Evolution

Integrating human impacts and ecological integrity into a risk-based protocol for conservation planning

Conservation planning aims to protect biodiversity by sustainng the natural physical, chemical, and biological processes within representative ecosystems. Often data to measure these components are inadequate or unavailable. The impact of human activities on ecosystem processes complicates integrity assessments and might alter ecosystem organization at multiple spatial scales. Freshwater conservation targets, such as populations and communities, are influenced by both intrinsic aquatic properties and the surrounding landscape, and locally collected data might not accurately reflect potential impacts. We suggest that changes in five major biotic drivers—energy sources, physical habitat, flow regime, water quality, and biotic interactions—might be used as surrogates to inform conservation planners of the ecological integrity of freshwater ecosystems. Threats to freshwater systems might be evaluated based on their impact to these drivers to provide an overview of potential risk to conservation targets. We developed a risk-based protocol, the Ecological Risk Index (ERI), to identify watersheds with least/most risk to conservation targets. Our protocol combines risk-based components, specifically the frequency and severity of human-induced stressors, with biotic drivers and mappable land- and water-use data to provide a summary of relative risk to watersheds. We illustrate application of our protocol with a case study of the upper Tennessee River basin, USA. Differences in risk patterns among the major drainages in the basin reflect dominant land uses, such as mining and agriculture. A principal components analysis showed that localized, moderately severe threats accounted for most of the threat composition differences among our watersheds. We also found that the relative importance of threats is sensitive to the spatial grain of the analysis. Our case study demonstrates that the ERI is useful for evaluating the frequency and severity of ecosystemwide risk, which can inform local and regional conservation planning.

Tennessee

The migratory bird treaty and a century of waterfowl conservation

In the final decades of the nineteenth century, concern was building about the status of migratory bird populations in North America. In this literature review, we describe how that concern led to a landmark conservation agreement in 1916, between the United States and Great Britain (on behalf of Canada) to conserve migratory birds shared by Canada and the United States. Drawing on published literature and our personal experience, we describe how subsequent enabling acts in both countries gave rise to efforts to better estimate population sizes and distributions, assess harvest rates and demographic impacts, design and fund landscape-level habitat conservation initiatives, and organize necessary political and regulatory processes. Executing these steps required large-scale thinking, unprecedented regional and international cooperation, ingenuity, and a commitment to scientific rigor and adaptive management. We applaud the conservation efforts begun 100 years ago with the Migratory Bird Treaty Convention. The agreement helped build the field of wildlife ecology and conservation in the twentieth century but only partially prepares us for the ecological and social challenges ahead.

Journal of Wildlife Management

Cerulean Warbler Technical Group: Coordinating international research and conservation

Effective conservation for species of concern requires interchange and collaboration among conservationists and stakeholders. The Cerulean Warbler Technical Group (CWTG) is a consortium of biologists and managers from government agencies, non-governmental organizations, academia, and industry, who are dedicated to finding pro-active, science-based solutions for conservation of the Cerulean Warbler (Setophaga cerulea). Formed in the United States in 2001, CWTG’s scope soon broadened to address the species’ ecology and conservation on both the breeding and non-breeding ranges, in partnership with biologists from South and Central America. In 2004, CWTG launched the Cerulean Warbler Conservation Initiative, a set of activities aimed at addressing information and conservation needs for the species. These include (1) studies in the core breeding range to assess Cerulean Warbler response to forest management practices and to identify mined lands that could be reforested to benefit the species, (2) ecological and demographic studies on the winter range, and (3) surveys of Cerulean Warbler distribution on the breeding and winter ranges and during migration. A rangewide conservation action plan has been completed, along with a more detailed conservation plan for the non-breeding range. CWTG and partners now move forward with on-the-ground conservation, while still addressing unmet information needs.

Ornitologia Neotropical

Linking animals aloft with the terrestrial landscape

Despite using the aerosphere for many facets of their life, most flying animals (i.e., birds, bats, some insects) are still bound to terrestrial habitats for resting, feeding, and reproduction. Comprehensive broad-scale observations by weather surveillance radars of animals as they leave terrestrial habitats for migration or feeding flights can be used to map their terrestrial distributions either as point locations (e.g., communal roosts) or as continuous surface layers (e.g., animal densities in habitats across a landscape). We discuss some of the technical challenges to reducing measurement biases related to how radars sample the aerosphere and the flight behavior of animals. We highlight a recently developed methodological approach that precisely and quantitatively links the horizontal spatial structure of birds aloft to their terrestrial distributions and provides novel insights into avian ecology and conservation across broad landscapes. Specifically, we present case studies that (1) elucidate how migrating birds contend with crossing ecological barriers and extreme weather events, (2) identify important stopover areas and habitat use patterns of birds along their migration routes, and (3) assess waterfowl response to wetland habitat management and restoration. These studies aid our understanding of how anthropogenic modification of the terrestrial landscape (e.g., urbanization, habitat management), natural geographic features, and weather (e.g., hurricanes) can affect the terrestrial distributions of flying animals.

Book chapter