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Managing climate change refugia for climate adaptation

Refugia have long been studied from paleontological and biogeographical perspectives to understand how populations persisted during past periods of unfavorable climate. Recently, researchers have applied the idea to contemporary landscapes to identify climate change refugia, here defined as areas relatively buffered from contemporary climate change over time that enable persistence of valued physical, ecological, and socio-cultural resources. We differentiate historical and contemporary views, and characterize physical and ecological processes that create and maintain climate change refugia. We then delineate how refugia can fit into existing decision support frameworks for climate adaptation and describe seven steps for managing them. Finally, we identify challenges and opportunities for operationalizing the concept of climate change refugia. Managing climate change refugia can be an important option for conservation in the face of ongoing climate change.

PLoS ONE↗

Columbia Environmental Research Center

The U.S. Geological Survey Columbia Environmental Research Center performs research to solve challenging environmental problems related to contaminants and habitat alterations in aquatic and terrestrial ecosystems. The research is interdisciplinary and pursued through partnerships within the U.S. Geological Survey and with national, international, state, and local agencies; nongovernmental organizations; and universities. Research is prioritized to provide science to the U.S. Department of the Interior and other natural resource management agencies to inform rehabilitation of degraded habitats and imperiled fish and wildlife populations. The Columbia Environmental Research Center was established in 1966 in Columbia, Missouri, as the U.S. Fish and Wildlife Service’s Fish Pesticide Research Laboratory; the Columbia Environmental Research Center was incorporated into the U.S. Geological Survey in 1996. The U.S. Geological Survey’s staff of 130 includes 90 scientists of which one-half have advanced degrees in ecology, toxicology, biology, biochemistry, chemistry, hydrology, geology, and information technology.

Missouri↗

Road impacts on the Baca National Wildlife Refuge, Colorado, with emphasis on effects to surface- and shallow ground-water hydrology - A literature review

A review of published research on unpaved road effects on surface-water and shallow ground-water hydrology was undertaken to assist the Baca National Wildlife Refuge, Colorado, in understanding factors potentially influencing refuge ecology. Few studies were found that addressed hydrological effects of roads on a comparable area of shallow slope in a semiarid region. No study dealt with road effects on surface- and ground-water supplies to ephemeral wetlands, which on the refuge are sustained by seasonal snowmelt in neighboring mountains. Road surfaces increase runoff, reduce infiltration, and serve as a sediment source. Roadbeds can interfere with normal surface- and ground-water flows and thereby influence the quantity, timing, and duration of water movement both across landscapes and through the soil. Hydrologic effects can be localized near the road as well as widespread and distant. The number, arrangement, and effectiveness of road-drainage structures (culverts and other devices) largely determine the level of hydrologic alteration produced by a road. Undesirable changes to natural hydrologic patterns can be minimized by considering potential impacts during road design, construction, and maintenance. Road removal as a means to restore desirable hydrologic conditions to landscapes adversely affected by roads has yet to be rigorously evaluated.

Colorado↗

Federal collaboration in science for invasive mammal management in U.S. National Parks and Wildlife Refuges of the Pacific Islands

Some of the most isolated islands in the Pacific Ocean are home to US National Parks and Wildlife Refuges. These islands are known for flora and fauna that occur nowhere else, but also for invasive species and other factors which have resulted in the disproportionate extinction of native species. The control of invasive mammals is the single most expensive natural resource management activity essential for restoring ecological integrity to parks in the Hawaiian Islands, American Samoa, and the islands of Guam and Saipan. Science-based applications supporting management efforts have been shaped by longstanding collaborative federal research programs over the past four decades. Consequently, feral goats (Capra hircus) have been removed from >690 km2 in National Parks, and feral pigs (Sus scrofa) have been removed from >367 km2 of federal lands of Hawai‘i, bringing about the gradual recovery of forest ecosystems. The exclusion of other non-native ungulates and invasive mammals is now being undertaken with more sophisticated control techniques and fences. New fence designs are now capable of excluding feral cats (Felis catus) from large areas to protect endangered native waterfowl and nesting seabirds. Rodenticides which have been tested and registered for hand and aerial broadcast in Hawai‘i have been used to eradicate rats from small offshore islands to protect nesting seabirds and are now being applied to montane environments of larger islands to protect forest birds. Forward-looking infrared radar (FLIR) is also being applied to locate wild ungulates which were more recently introduced to some islands. All invasive mammals have been eradicated from some remote small islands, and it may soon be possible to manage areas on larger islands to be free of invasive mammals at least during seasonally important periods for native species.

Conference Paper↗

Ciscoes ( Coregonus , subgenus Leucichthys ) of the Laurentian Great Lakes and Lake Nipigon

This study of the ciscoes ( Coregonus , subgenus Leucichthys ) of the Great Lakes and Lake Nipigon represents a furtherance through 2015 of field research initiated by Walter Koelz in 1917 and continued by Stanford Smith in the mid-1900s—a period spanning nearly a century. Like Koelz’s study, this work contains information on taxonomy, geographical distribution, ecology, and status of species (here considered forms). Of the seven currently recognized forms ( C. artedi , C. hoyi , C. johannae , C. kiyi , C. nigripinnis , C. reighardi , and C. zenithicus ) described by Koelz as major in his 1929 monograph, two ( C. johannae and C. reighardi ) are extinct. In addition, C. alpenae , described by Koelz but subsequently synonymized with C. zenithicus , although extinct, is recognized as valid making a total of eight major forms. Six of these forms, all but C. artedi and C. hoyi , have been lost from Lake Michigan, and seven have been lost from Lake Huron, leaving in Lake Huron only C. artedi and an introgressed deepwater form that we term a hybrid swarm. C. artedi appears, like its sister form C. alpenae , to have been lost from Lake Erie. Only C. artedi remains extant in Lake Ontario, its three sister forms ( C. hoyi , C. kiyi , and C. reighardi ) having disappeared long ago. Lakes Superior and Nipigon have retained their original species flocks consisting of four forms each: C. artedi , C. hoyi , and C. zenithicus in both lakes; C. kiyi in Lake Superior; and C. nigripinnis in Lake Nipigon. Morphological deviations from the morphotypes described by Koelz have been modest in contemporary samples. Overall, C. kiyi and C. artedi were the most morphologically stable forms while C. hoyi , C. nigripinnis , and C. zenithicus were the least stable. Although contemporary populations of C. artedi from Lakes Michigan and Huron are highly diverged from the morphotypes described by Koelz, the contemporary samples were of undescribed deep-bodied forms unlikely to have been sampled by Koelz because of their association with bays. Of the two intact species flocks, Lake Nipigon’s was much less stable morphologically than Lake Superior’s even though Lake Nipigon is far less disturbed. Two priorities for research are determining the role of developmental plasticity in morphological divergence, especially within C. zenithicus of Lake Superior, and the basis for morphological divergence in C. artedi .

Lake Nipigon, Laurentian Great Lakes↗

Minimizing effects of methodological decisions on interpretation and prediction in species distribution studies: An example with background selection

Evaluating the conditions where a species can persist is an important question in ecology both to understand tolerances of organisms and to predict distributions across landscapes. Presence data combined with background or pseudo-absence locations are commonly used with species distribution modeling to develop these relationships. However, there is not a standard method to generate background or pseudo-absence locations, and method choice affects model outcomes. We evaluated combinations of both model algorithms (simple and complex generalized linear models, multivariate adaptive regression splines, Maxent, boosted regression trees, and random forest) and background methods (random, minimum convex polygon, and continuous and binary kernel density estimator (KDE)) to assess the sensitivity of model outcomes to choices made. We evaluated six questions related to model results, including five beyond the common comparison of model accuracy assessment metrics (biological interpretability of response curves, cross-validation robustness, independent data accuracy and robustness, and prediction consistency). For our case study with cheatgrass in the western US, random forest was least sensitive to background choice and the binary KDE method was least sensitive to model algorithm choice. While this outcome may not hold for other locations or species, the methods we used can be implemented to help determine appropriate methodologies for particular research questions.

Ecological Modelling↗

Optimal sampling design for spatial capture‐recapture

Spatial capture‐recapture (SCR) has emerged as the industry standard for estimating population density by leveraging information from spatial locations of repeat encounters of individuals. The precision of density estimates depends fundamentally on the number and spatial configuration of traps. Despite this knowledge, existing sampling design recommendations are heuristic and their performance remains untested for most practical applications. To address this issue, we propose a genetic algorithm that minimizes any sensible, criteria‐based objective function to produce near‐optimal sampling designs. To motivate the idea of optimality, we compare the performance of designs optimized using three model‐based criteria related to the probability of capture. We use simulation to show that these designs out‐perform those based on existing recommendations in terms of bias, precision, and accuracy in the estimation of population size. Our approach, available as a function in the R package oSCR, allows conservation practitioners and researchers to generate customized and improved sampling designs for wildlife monitoring.

Ecology↗

Use of dynamic occupancy models to assess the response of Darters (Teleostei: Percidae) to varying hydrothermal conditions in a southeastern United States tailwater

During the past 100 years, most large rivers in North America have been altered for flood control, hydropower, navigation or water supply development. Although these activities clearly provide important human services, their associated environmental disturbances can profoundly affect stream-dwelling organisms. We used dynamic multi-species occupancy models combined with a trait-based approach to estimate the influence of site-level and species-level characteristics on patch dynamic rates for 15 darter species native to the Elk River, a large, flow-regulated Tennessee River tributary in Tennessee and Alabama. Dynamic occupancy modelling results indicated that for every 2.5 °C increase in stream temperature, darters were 3.94 times more likely to colonize previously unoccupied stream reaches. Additionally, large-bodied darter species were 3.72 times more likely to colonize stream reaches compared with small-bodied species, but crevice-spawning darter species were 5.24 times less likely to colonize previously unoccupied stream reaches. In contrast, darters were 2.21 times less likely to become locally extinct for every 2.5 °C increase in stream temperature, but high stream discharge conditions elevated the risk of local extinction. Lastly, the presence of populations in neighbouring upstream study reaches contributed to a lower risk of extinction, whereas the presence of populations in neighbouring downstream study reaches contributed to higher rates of colonization. Our study demonstrates the application of a trait-based approach combined with a metapopulation framework to assess the patch dynamics of darters in a regulated river. Results from our study will provide a baseline for evaluating the ecological consequences of alternative dam operations.

River Research and Applications↗

Thermal characteristics of amphibian microhabitats in a fire-disturbed landscape

Disturbance has long been a central issue in amphibian conservation, often regarding negative effects of logging or other forest management activities, but some amphibians seem to prefer disturbed habitats. After documenting increased use of recently burned forests by boreal toads (Bufo boreas), we hypothesized that burned habitats provided improved thermal opportunities in terrestrial habitats. We tested this hypothesis by conducting a radio telemetry study of habitat use (reported previously) and by using physical models that simulated the temperature of adult toads. We deployed 108 physical models in and adjacent to a 1-year old burn using a fully-replicated design with three burn severities (unburned, partial, high severity) and four microhabitats (open surface, under vegetation, under log, in burrow). Model temperatures were compared to a range of preferred temperatures in published studies. We found 70% more observations within the preferred temperature range of B. boreas in forests burned with high severity than in unburned areas. Burned forest was warmer than unburned forest across all microhabitats, but the largest relative difference was in burrows, which averaged 3 ??C warmer in high-severity burn areas and remained warmer though the night. More than twice as many observations were within the preferred temperature range in high-severity burrows than in unburned burrows. Areas burned with high severity were still warmer than unburned forest 3 years after the fire. Habitat use of toads during the concurrent radio telemetry study matched that predicted by the physical models. These results suggest there are fitness-linked benefits to toads using burned habitats, such as increased growth, fertility, and possibly disease resistance. However, increased soil temperatures that result from wildfire may be detrimental to other amphibian species that prefer cooler temperatures and stable environments. More broadly, our data illustrate the use of physical models to measure and interpret changes that amphibians may experience from disturbance, and highlight the need for research linking vital rates such as growth and survival to disturbance.

Forest Ecology and Management↗

Reconstructing the response of C3 and C4 plants to decadal-scale climate change during the late Pleistocene in southern Illinois using isotopic analyses of calcified rootlets

The ??13C and ??18O values of well-preserved carbonate rhizoliths (CRs) provide detailed insights into changes in the abundance of C3 and C4 plants in response to approximately decadal-scale changes in growing-season climate. We performed stable isotope analyses on 35-40 CRs sampled at 1-cm intervals from an 18-cm-thick paleosol formed in southern Illinois during Wisconsin interstadial 2. Minimum ??13C values show little variation with depth, whereas maximum values vary dramatically, and average values show noticeable variability; maximum ??18O values vary less than the minimum ??18O values. These findings indicate that a diverse and stable C3 flora with a limited number of C4 grass species prevailed during this interval, and suggest that the maximum growing-season temperatures were relatively stable, but minimum growing-season temperatures varied considerably. Two general patterns characterize the relationships between the ??13C and ??18O values obtained from the 1-cm samples. In some cases, low ??13C values correspond to low ??18O values and high ??13C values correspond to high ??18O values, suggesting that cooler growing-season temperatures favored C3 and warmer growing-season temperatures favored C4 plants. In other cases, low ??13C values correspond to high ??18O values, likely suggesting that wetter growing-season conditions were favorable to C3 plants. The high density of well-preserved CRs in this paleosol provides a unique opportunity to study detailed ecological responses to high-resolution variability in growing-season climate. ?? 2006 University of Washington.

Quaternary Research↗

Effective discharge analysis of ecological processes in streams

Discharge is a master variable that controls many processes in stream ecosystems. However, there is uncertainty of which discharges are most important for driving particular ecological processes and thus how flow regime may influence entire stream ecosystems. Here the analytical method of effective discharge from fluvial geomorphology is used to analyze the interaction between frequency and magnitude of discharge events that drive organic matter transport, algal growth, nutrient retention, macroinvertebrate disturbance, and habitat availability. We quantify the ecological effective discharge using a synthesis of previously published studies and modeling from a range of study sites. An analytical expression is then developed for a particular case of ecological effective discharge and is used to explore how effective discharge varies within variable hydrologic regimes. Our results suggest that a range of discharges is important for different ecological processes in an individual stream. Discharges are not equally important; instead, effective discharge values exist that correspond to near modal flows and moderate floods for the variable sets examined. We suggest four types of ecological response to discharge variability: discharge as a transport mechanism, regulator of habitat, process modulator, and disturbance. Effective discharge analysis will perform well when there is a unique, essentially instantaneous relationship between discharge and an ecological process and poorly when effects of discharge are delayed or confounded by legacy effects. Despite some limitations the conceptual and analytical utility of the effective discharge analysis allows exploring general questions about how hydrologic variability influences various ecological processes in streams.

Water Resources Research↗

Ecogeomorphologic response of a tidally restricted salt marsh to tidal restoration and sea-level rise

Diked marshes with decades of restricted tidal flow exhibit fundamentally altered hydrologic and ecologic dynamics that result in limited vertical accretion, land subsidence, and underdeveloped tidal channels. These changes complicate marsh restoration efforts and constrain the marsh's ability to build elevation in response to future sea levels. In this study, we developed an integrated hydrodynamic and marsh accretion model to conduct the first assessment of the restoration potential of a diked marsh in response to reintroduction of tidal flow and sea-level rise (SLR). We applied our model to evaluate changes in biomass production in the Herring River estuary, Cape Cod, MA, USA—which has experienced significant ecological shifts due to more than a century of tidal restriction—in response to a range of SLR scenarios, tidal flow conditions, management interventions, and biomass production rates through 2100. We found that with restoration of tidal flow, the simulated marsh extent expanded in areas with efficient drainage, increasing biomass production by up to 4.4 times. However, when SLR was imposed, parts of the marsh gradually transitioned to open water, leading to a decline in marsh coverage from 19% to as low as 12%. Management interventions aimed at enhancing marsh elevation and improving the drainage capacity of tidal channels slowed marsh loss, supporting marsh area through 2100. Our findings provide insight into the key drivers of marsh resilience and loss, and our approach can be applied across tidally restricted and restored systems to develop targeted, adaptive restoration strategies that achieve desired ecological outcomes.

Massachusetts↗

Parasite (Ribeiroia ondatrae) infection linked to amphibian malformations in the western United States

Parasites and pathogens can influence the survivorship, behavior, and very structure of their host species. For example, experimental studies have shown that trematode parasites can cause high frequencies of severe limb malformations in amphibians. In a broad-scale field survey covering parts of California, Oregon, Washington, Idaho, and Montana, we examined relationships between the frequency and types of morphological abnormalities in amphibians and the abundance of trematode parasite infection, pH, concentrations of 61 pesticides, and levels of orthophosphate and total nitrate. We recorded severe malformations at frequencies ranging from 1% to 90% in nine amphibian species from 53 aquatic systems. Infection of larvae by the trematode Ribeiroia ondatrae was associated with, and functionally related to, higher frequencies of amphibian limb malformations than found in uninfected populations (≤5%). Parasites were concentrated around the basal tissue of hind limbs in infected anurans, and malformations associated with infection included skin webbings, supernumerary limbs and digits, and missing or malformed hind limbs. In the absence of Ribeiroia, amphibian populations exhibited low (0-5%) frequencies of abnormalities involving missing digits or distal portions of a hind limb. Species were affected differentially by the parasite, and Ambystoma macrodactylum, Hyla regilla, Rand aurora, R. luteiventris, and Taricha torosa typically exhibited the highest frequencies of abnormalities. None of the water-quality variables measured was associated with malformed amphibians, but aquatic snail hosts (Planorbella spp.) were significant predictors of the presence and abundance of Ribeiroia infection. Morphological comparisons of adult specimens of Ribeiroia collected from different sites and raised in experimental definitive hosts suggested that all samples represented the same species - R. ondatrae. These field results, coupled with experimental research on the effects of Ribeiroia on amphibians, demonstrate that Ribeiroia infection is an important and widespread cause of amphibian limb malformations in the western United States. The relevance of trematode infection to declines of amphibian populations and the influence of habitat modification on the pathology and life cycle of Ribeiroia are emphasized as areas requiring further research.

Ecological Monographs↗

Fire severity and ecosytem responses following crown fires in California shrublands

Chaparral shrublands burn in large high-intensity crown fires. Managers interested in how these wildfires affect ecosystem processes generally rely on surrogate measures of fire intensity known as fire severity metrics. In shrublands burned in the autumn of 2003, a study of 250 sites investigated factors determining fire severity and ecosystem responses. Using structural equation modeling we show that stand age, prefire shrub density, and the shortest interval of the prior fire history had significant direct effects on fire severity, explaining >50% of the variation in severity. Fire severity per se is of interest to resource managers primarily because it is presumed to be an indicator of important ecosystem processes such as vegetative regeneration, community recovery, and erosion. Fire severity contributed relatively little to explaining patterns of regeneration after fire. Two generalizations can be drawn: fire severity effects are mostly short-lived, i.e., by the second year they are greatly diminished, and fire severity may have opposite effects on different functional types. Species richness exhibited a negative relationship to fire severity in the first year, but fire severity impacts were substantially less in the second postfire year and varied by functional type. Much of this relationship was due to alien plants that are sensitive to high fire severity; at all scales from 1 to 1000 m 2 , the percentage of alien species in the postfire flora declined with increased fire severity. Other aspects of disturbance history are also important determinants of alien cover and richness as both increased with the number of times the site had burned and decreased with time since last fire. A substantial number of studies have shown that remote-sensing indices are correlated with field measurements of fire severity. Across our sites, absolute differenced normalized burn ratio (dNBR) was strongly correlated with field measures of fire severity and with fire history at a site but relative dNBR was not. Despite being correlated with fire severity, absolute dNBR showed little or no relationship with important ecosystem responses to wildfire such as shrub resprouting or total vegetative regeneration. These findings point to a critical need for further research on interpreting remote sensing indices as applied to postfire management of these shrublands.

Ecological Applications↗

Activity, aggression, and habitat use of ruffe ( Gymnocephalus cernuus ) and round goby ( Apollonia melanostoma ) under laboratory conditions

Potential negative ecological interactions between ruffe Gymnocephalus cernuus and round goby Apollonia melanostoma (formerly Neogobius melanostomus ) might affect the colonization dynamics of these invasive species where they are sympatric in the Great Lakes. In order to determine the potential for ecological interactions between these species, we examined the activity, aggression, and habitat use of round gobies and ruffe in single species and mixed species laboratory experiments. Trials included conditions in which food was concentrated (in light or darkness) or scattered. Results showed that ruffe were more active than gobies, particularly when food was scattered. Activity of both species was significantly lower during darkness. Round gobies were significantly more aggressive than ruffe, and total aggression was lower in mixed species trials. Habitat use by ruffe and round gobies overlapped considerably, but we observed significant differences between species in their use of specific habitats that depended on experimental conditions. Overall, ruffe used open habitats more often than did round gobies, primarily when food was scattered. Round gobies used rocks significantly more frequently than did ruffe, but their use of rock habitat decreased during dark conditions. Ruffe were found more often in plant habitats and less often near the wall of the pool in trials during daylight with concentrated food. Activity and habitat use of ruffe and round goby did not significantly differ between single and mixed species trials. Overall, we found little evidence for negative ecological interactions between ruffe and round goby in these laboratory experiments.

Journal of Great Lakes Research↗

Bridging theory and practice to inform seed selection for restoration

Introduction Land managers often face the critical decision of what plant materials to select for ecological restoration. Selection depends on factors that span ecological conditions and human interventions in the restoration process, which can strongly influence the recovery of a degraded ecosystem. Objectives To improve the seed selection process, we compared scientific literature and guidance from five federal agencies to bridge seed selection “theory” and “practice” in the interior western and central United States. Methods We extracted recurring themes and keywords associated with abiotic, biotic, land management, and producer practices that influence seed selection and analyzed their proportional use between agency guidance and scientific literature. Results We identified significant overlap between key concepts in guidance and literature, highlighting the interconnectedness between science and management. However, nuanced patterns emerged across seed selection themes, including more focus in the literature on local adaptation and genetic structure, ramifications of introducing gene flow and artificial selection during plant production, and provenancing strategies to address changing climatic conditions. In contrast, agency guidance focused on genetically appropriate and environmentally suitable seed selection and practical concerns, including seed cost, availability, and diversity. Seed selection studies in the literature were concentrated in the Great Basin and Colorado Plateau, scattered in the Great Plains, and were largely experimental. Conclusions Our review suggests new opportunities for co-production of science and decision-making frameworks among researchers, land managers, and seed producers to inform seed selection guidance for restoration.

Restoration Ecology↗

Nutritional ecology of ursids: A review of newer methods and management implications

The capability to understand the nutritional ecology of free-ranging bears has increased dramatically in the last 20 years. Advancements have occurred because (1) managers and biologists recognized the need to link habitat quality, productivity, and variability with bear movements, home ranges, and demographic parameters like reproductive output, survival, and population growth, and (2) several research teams are using new methods to build on the results of earlier field studies. Our ability to couple new field methods and empirical field research with controlled experiments using captive bears has been central to our increased understanding of bear nutrition. Newer methods include the use of stable isotopes to quantify assimilated diet and nutrient flows within ecosystems, bioelectrical impedance to measure body composition, and naturally occurring mercury to estimate fish intake. Controlled experiments using captive bears have been integral to developing methods, isolating specific variables by controlling the environment, and providing additional nutritional understanding necessary to interpret field observations. We review new methods and apply our increased understanding of bear nutritional ecology to 3 management issues: (1) the importance of salmon ( Oncorhynchus spp.) to brown bears ( Ursus arctos ) in the Pacific Northwest, (2) the consequences of the closure of the Yellowstone garbage dumps to grizzly bears, and (3) the relocation of problem bears.

Ursus↗

Considering transient population dynamics in the conservation of slow life-history species: An application to the sandhill crane

The importance of transient dynamics of structured populations is increasingly recognized in ecology, yet these implications are not largely considered in conservation practices. We investigate transient and long-term population dynamics to demonstrate the process and utility of incorporating transient dynamics into conservation research and to better understand the population management of slow life-history species; these species can be theoretically highly sensitive to short- and long-term transient effects. We are specifically interested in the effects of anthropogenic removal of individuals from populations, such as caused by harvest, poaching, translocation, or incidental take. We use the sandhill crane ( Grus canadensis ) as an exemplar species; it is long-lived, has low reproduction, late maturity, and multiple populations are subject to sport harvest. We found sandhill cranes to have extremely high potential, but low likelihood for transient dynamics, even when the population is being harvested. The typically low population growth rate of slow life-history species appears to buffer against many perturbations causing large transient effects. Transient dynamics will dominate population trajectories of these species when stage structures are highly biased towards the younger and non-reproducing individuals, a situation that may be rare in established populations of long-lived animals. However, short-term transient population growth can be highly sensitive to vital rates that are relatively insensitive under equilibrium, suggesting that stage structure should be known if perturbation analysis is used to identify effective conservation strategies. For populations of slow life-history species that are not prone to large perturbations to their most productive individuals, population growth may be approximated by equilibrium dynamics.

Biological Conservation↗