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Southern sea otter (Enhydra lutris nereis) population biology at Big Sur and Monterey, California --Investigating the consequences of resource abundance and anthropogenic stressors for sea otter recovery

The range of the southern sea otter ( Enhydra lutris nereis ) spans most of the central California coast from Half Moon Bay to Gaviota. Some coastal areas within this range are heavily developed and highly impacted by humans, while other areas are wild and largely pristine. Determining the relative importance of food resource abundance, environmental conditions, and anthropogenic increases in pathogens and pollutants to population change in sea otters is critical to understanding limitations to population growth. To investigate the causal links between the sluggish population growth of sea otters in central California and factors that could be driving variation in survival and reproduction, we designed a study to compare two distinct subpopulations—one in an area of low human impact (Big Sur) and one in an area of high human impact (Monterey). Between 2008 and 2011, the U.S. Geological Survey and collaborators conducted a telemetry-based study of sea otters at these two locations. The results of this study were not consistent with the hypothesis that sea otters adjacent to human population centers (Monterey) experience higher exposure to pollutants and pathogens than those in lower impacted areas (Big Sur). In fact, based on serological analysis, female sea otters from Big Sur showed higher exposure rates to Toxoplasma gondii than did female otters from Monterey, while domoic acid exposure appeared to be similar at both sites. Gene expression (specifically transcription) analysis did not indicate any consistent differences between the two populations that would have suggested a response to pathogen or toxin exposure, although there were temporal changes in gene transcription for sea otters at Big Sur following potential exposure to run-off from wildfires that occurred during the study. Together, these metrics suggest that variation in exposure to environmental stressors occurred, but patterns were not clearly attributable to differences in human population densities or land-use patterns. When compared to Monterey, sea otters in Big Sur spent more time feeding, had a higher degree of dietary specialization, were in poorer body condition, and had lower survival rates (both pups and adults). Together, these metrics suggest that otters at Big Sur had greater nutritional stress, consistent with lower per-capita resource abundance. Overall, study results indicate that density-dependent population regulation, mediated by per-capita resource abundance, is the most significant factor currently limiting population growth in the center part of the range. Additionally, spatial and temporal variation in environmental and anthropogenic stressors also can affect sea otter health, although patterns of variation are complex and are not simply a function of proximity to human populations. We also found that exposure to environmental stressors (either natural or anthropogenic in origin) often is associated with resource limitation. Finally, our results indicate that sea otter populations are structured at relatively small spatial scales, and the processes that regulate population abundance (including density-dependent resource abundance) also occur at these smaller, more local scales.

California↗

The status and conservation needs of the Micronesian Megapode (Megapodius laperouse laperouse) across the Mariana archipelago

Context Accurate baseline data for wildlife populations are important to track trends of these populations over time and to identify threats to their long-term persistence. Aims We aimed to assess the status and distribution of the little studied megapode ( Megapodius laperouse laperouse ) across the Mariana Islands. Methods Using passive and call playback facilitated surveys in 2008 through 2010, we employed point–transect distance sampling to assess island-level and archipelago-wide status of this megapode. To assess conservation needs, we defined human presence as the current, recent, or intermittent occurrence of humans on islands. Key results We recorded 657 megapode detections and estimated an archipelago level abundance of 11,542 individuals (95% CI: 5456–17,623) from 699 sampling points across 10 islands. Three islands supported 86% of the megapode population, but cumulatively comprise only 2% of the archipelago’s land area. Conclusions Micronesian Megapodes preferred native forest. Human presence and the availability of native forest may limit their abundance and distribution in the Mariana Islands. Although the probability of detecting megapodes was significantly greater on islands without high human presence, significantly more detections were recorded in forests with dense or closed understory on those islands that supported greater human populations. Implications Given their status and confined distribution in the Mariana Islands, additional studies investigating megapode incubation sites and movement within and between islands would provide fundamental information on megapode ecology and enhance conservation efforts. Continued and expanded ungulate removal, predator control, and habitat restoration would further enhance the likelihood of megapode persistence in the archipelago.

Pacific Conservation Biology↗

Representing plant diversity in land models: An evolutionary approach to make ‘Functional Types’ more functional

Plants are critical mediators of terrestrial mass and energy fluxes, and their structural and functional traits have profound impacts on local and global climate, biogeochemistry, biodiversity, and hydrology. Yet Earth System Models (ESMs), our most powerful tools for predicting the effects of humans on the coupled biosphere-atmosphere system, simplify the incredible diversity of land plants into a handful of coarse categories of ‘Plant Functional Types’ (PFTs) that often fail to capture ecological dynamics such as biome distributions. The inclusion of more realistic functional diversity is a recognized goal for ESMs, yet there is currently no consistent, widely accepted way to add diversity to models, i.e. to determine what new PFTs to add and with what data to constrain their parameters. We review approaches to representing plant diversity in ESMs and draw on recent ecological and evolutionary findings to present an evolution-based functional type approach for further disaggregating functional diversity. Specifically, the prevalence of niche conservatism, or the tendency of closely related taxa to retain similar ecological and functional attributes through evolutionary time, reveals that evolutionary relatedness is a powerful framework for summarizing functional similarities and differences among plant types. We advocate that Plant Functional Types based on dominant evolutionary lineages (‘Lineage Functional Types’) will provide an ecologically defensible, tractable, and scalable framework for representing plant diversity in next-generation ESMs, with the potential to improve parameterization, process representation, and model benchmarking. We highlight how the importance of evolutionary history for plant function can unify the work of disparate fields to improve predictive modeling of the Earth system.

Global Change Biology↗

Environmental covariates associated with Cambarus veteranus (Decapoda: Cambaridae), an imperiled Appalachian crayfish endemic to West Virginia, USA

Cambarus veteranus Faxon, 1914 , a narrow endemic crayfish native to the Upper Guyandotte River Basin (UGB) in West Virginia, USA, was petitioned in 2014 by the United States Fish and Wildlife Service to be listed as endangered, but a status survey was recommended to determine if listing was warranted. During May and June 2015, surveys were undertaken across the UGB to determine the current distribution of the species. A total of 71 sites were sampled, including all streams where the species was previously recorded, as well as semi-randomly selected streams, with 1-9 125 m long sites sampled per wadeable stream. Physiochemical and physical habitat data (based on the Qualitative Habitat Evaluation Index, QHEI) were obtained at each site to determine abiotic factors that were associated with the presence of C. veteranus . Site detection or non-detection of C. veteranus and associated site covariates were modeled using logistic regression to determine covariates associated with the presence of the species. Cambarus veteranus was present in both the Pinnacle Creek and Clear Fork/Laurel Fork watersheds at 10 sites, but it was not observed in the remaining 61 sites. An additive effects model with conductivity and QHEI was selected as the best approximating model. Cambarus veteranus was associated with lower than average UGB conductivity (379 µS) and high (>80) QHEI score. All sites where C. veteranus was not detected had higher conductivity and/or lower QHEI scores.

West Virginia↗

Disease emergence in birds: Challenges for the twenty-first century

The paper by Hartup et al. (2001) on House Finch ( Carpodacus mexicanus ) conjunctivitis is an example of the rapid geographic spread that can result from disease emergence in naïve populations. That event was neither novel nor transient relative to its occurrence or effects. Disease emergence and reemergence are hallmarks of the latter part of the twentieth century (Center for Disease Control 1994, Levins et al. 1994, DaSilva and Laccarino 1999, Gratz 1999). Current examples involving domestic animals include the problems in Europe with bovine spongiform encephalopathy (BSE, or “mad cow disease”) (Brown 2001) and foot-and-mouth disease (FMD) (Kitching 1999). Human health has been affected by diseases caused by an array of viruses (Morse 1993, Nichol et al. 1993, Murphy and Nathanson 1994), bacteria (Dennis 1998, DaSilva and Laccarino 1999), rickettsia (Walker and Dumier 1996, Azad et al. 1997), protozoans (Tuerrant 1997, Saini et al. 2000), and metazoan parasites (Hildreth et al. 1991, Gubler 1998), as well as other causes. Acquired immune deficiency syndrome (AIDS) has received the most notoriety of those diseases (Hahn et al. 2000, Schwartlander et al. 2000). A similar pattern exists on a global scale for free-ranging wildlife populations (Table 1) (Friend 1994, 1995; Epstein et al. 1998, Daszak et al. 2000). However, in comparison to disease emergence affecting humans and domestic animals, response to emerging diseases of wildlife is generally superficial. We present concepts and data to support our contention that failure to adequately address disease emergence in free-ranging wildlife is resulting in a diminished capability to achieve and sustain desired geographic distributions and population abundance for species of wild birds, including some threatened and endangered avifauna. For clarity, we define disease and disease emergence in the context of our use of those terms because they are the focus of our comments. Disease is any departure from health (Guralnik 1982); that is, dysfunction contributing to physiological, physical, reproductive, behavioral, or other impairment that reduces the probability of survival of individuals. If enough individuals are affected, the collective effects can reduce the sustainability of the population. Although disease can result from exposure to a wide variety of physical, chemical, and biological agents and other conditions, we focus this paper on microbes and parasites and to overt mortality caused by them. Thus, disease effects presented only represent the proverbial “tip of the iceberg” relative to the challenges wild avifauna face from disease. Our perspective of disease emergence expands the earlier definitions of emerging diseases by others (Centers for Disease Control and Prevention 1994, Morse 1995) to include all species. Our comments are defined by the context of disease occurrences that have increased within the past three decades, or threaten to increase in the near future relative to populations affected, geographic distribution, or magnitude of effects.

The Auk↗

Skeletal variation and taxonomic boundaries among mainland and island populations of the common treeshrew (Mammalia: Scandentia: Tupaiidae)

Treeshrews (order Scandentia) include 23 currently recognized species of small-bodied mammals from South and Southeast Asia. The taxonomy of the common treeshrew, Tupaia glis , which inhabits the Malay Peninsula south of the Isthmus of Kra, as well as a variety of offshore islands, has an extremely complicated history resulting from its wide distribution and subtly variable pelage. In our ongoing investigation of species boundaries in Tupaia , we compared island and mainland populations of T. glis using multivariate analyses. Specifically, we compared the skull and hand morphology of 13 island populations, most of which have been recognized as separate species or subspecies, to that of the mainland population. Island populations generally average smaller body size than those on the mainland, but none of the island populations are sufficiently distinct from the mainland population to warrant species recognition. This has important conservation implications for this widespread and morphologically variable species. It also highlights the potential role that ecogeographic explanations can play in understanding intraspecific variation, a role that should be considered in taxonomic studies and investigated further in T. glis and other treeshrews.

Malay Peninsula↗

Characterizing fish community diversity across Virginia landscapes: Prerequisite for conservation

The number of community types occurring within landscapes is an important, but often unprotected, component of biological diversity. Generally applicable protocols for characterizing community diversity need to be developed to facilitate conservation. We used several multivariate techniques to analyze geographic variation in the composition of fish communities in Virginia streams. We examined relationships between community composition and six landscape variables: drainage basin, physiography, stream order, elevation, channel slope, and map coordinates. We compared patterns at two scales (statewide and subdrainage-specific) to assess sensitivity of community classification to spatial scale. We also compared patterns based on characterizing communities by species composition vs. ecological composition. All landscape variables explained significant proportions of the variance in community composition. Statewide, they explained 32% of the variance in species composition and 48% of the variance in ecological composition. Typical communities in each drainage or physiography were statistically distinctive. Communities in different combinations of drainage, physiography, and stream size were even more distinctive, but composition was strongly spatially autocorrelated. Ecological similarity and species similarity of community pairs were strongly related, but replacement by ecologically similar species was common among drainage–physiography combinations. Landscape variables explained significant proportions of variance in community composition within selected subdrainages, but proportions were less than at the statewide scale, and the explanatory power of individual variables varied considerably among subdrainages. Community variation within subdrainages appeared to be much more closely related to environmental variation than to replacement among ecologically similar species. Our results suggest that taxonomic and ecological characterizations of community composition are complementary; both are useful in a conservation context. Landscape features such as drainage, physiography, and water body size generally may provide a basis for assessing aquatic community diversity, especially in regions where the biota is poorly known. Systematic conservation of community types would be a major advance relative to most current conservation programs, which typically focus narrowly on populations of imperiled species. More effective conservation of aquatic biodiversity will require new approaches that recognize the value of both species and assemblages, and that emphasize protection of key landscape-scale processes.

Ecological Applications↗

Population ecology of variegate darter (Etheostoma variatum) in Virginia

Variegate darters ( Etheostoma variatum ) were listed as endangered in Virginia in 1992. Reasons for listing included habitat degradation and concerns about current and future impacts of coal mining throughout their Virginia range. Prior to this research, little was known about variegate darter distribution, habitat use, or populations in Virginia. Two primary goals of this research were to gain knowledge about the current population ecology and the relationship between landscape-level factors (e.g., land cover changes, watershed size, isolation from other populations) on current and past variegate darter population sizes. We investigated distribution, habitat suitability, population genetics, and population size and structure of variegate darters in the upper Big Sandy River drainage, Buchanan, Dickenson, and Wise Co., Virginia. Our results indicate variegate darters are primarily found in the Levisa Fork, with highest densities and abundances between its confluence with Dismal Creek and the Virginia-Kentucky border. Sporadic occurrences in smaller tributaries to the Levisa and Tug forks indicate they exist more widely in low densities, especially near the confluence with the Tug and Levisa mainstems. Detection of variegate darters in smaller tributaries was inconsistent, with reach-level occupancy estimates varying among years. We detected young-of-year variegate darters every year we sampled, but age 1 + darters were indistinguishable from older darters based on standard length. Variegate darter population size and stability in Virginia were estimated via multiple methods, including site occupancy surveys, mark-recapture studies, and population genetic analysis. Using mark-recapture methods at five sites, we estimated overall population size in 2011 to be approximately 12,800 individuals in the 35-km reach between the Levisa Fork - Dismal Creek confluence and the Virginia-Kentucky border. Age structure seemed stable, with breeding adults and young-of-year collected annually during 2008-2011. Population genetic analysis indicated variegate darters in the Levisa Fork and its tributaries are part of a single genetic population. Historical and current genetic stability were seen in our analysis of the variegate darter population, with no genetic differentiation among riffles across the upper Levisa Fork watershed, indicating dispersal among these sites is enough to overcome random genetic drift. This population is genetically isolated from downstream populations by the dam at Fishtrap Lake, Pike Co., Kentucky, and is beginning to show genetic isolation from other nearby populations. As expected, the Virginia population is most closely related to those in the Russell Fork and Levisa Fork downstream of the dam. Regular monitoring of variegate darters in the Levisa Fork mainstem from the Dismal Creek confluence to the Virginia-Kentucky border would facilitate better understanding of normal fluctuations of population size and distribution, as well as assessments of population status. This reach encompasses the core of the variegate darter population in Virginia, and its persistence will determine long-term viability of this species. Given that little is known about long-term population trends, we suggest that annual site-occupancy and population size estimates be made at ten randomly selected riffles for at least ten years to understand normal levels of variability. Thereafter, these population parameters could be monitored bi-annually as a way to detect shrinking distribution or abundance, especially after any fish kill or other pollution event in the Levisa Fork. We further suggest that the sites upstream and downstream of the saline diffusor pipe be monitored to detect changes in the extent of the impact zone. Overall, the variegate darter population in Virginia appears stable, although primarily confined to the lower 35 km of the Levisa Fork. Nevertheless, variegate darters in Virginia remain susceptible to extirpation due to catastrophic events, both physical (chemical spill) and biological (disease outbreak or invasive species introduction).

Virginia, West Virginia↗

Contaminants as viral cofactors: assessing indirect population effects

Current toxicological methods often miss contaminant effects, particularly when immune suppression is involved. The failure to recognize and evaluate indirect and sublethal effects severely limits the applicability of those methods at the population level. In this study, the Vitality model is used to evaluate the population level effects of a contaminant exerting only indirect, sublethal effects at the individual level. Juvenile rainbow trout ( Oncorhynchus mykiss ) were injected with 2.5 or 10.0 mg/kg doses of the model CYP1A inducer, β-naphthoflavone (BNF) as a pre-stressor, then exposed to a challenge dose of 10 2 or 10 4 pfu/fish of infectious hematopoietic necrosis virus (IHNV), an important viral pathogen of salmonids in North America. At the end of the 28-d challenge, the mortality data were processed according to the Vitality model which indicated that the correlation between the average rate of vitality loss and the pre-stressor dose was strong: R 2 = 0.9944. Average time to death and cumulative mortality were dependent on the BNF dose, while no significant difference between the two viral dosages was shown, implying that the history of the organism at the time of stressor exposure is an important factor in determining the virulence or toxicity of the stressor. The conceptual framework of this model permits a smoother transfer of results to a more complex stratum, namely the population level, which allows the immunosuppressive results generated by doses of a CYP1A inducer that more accurately represent the effects elicited by environmentally-relevant contaminant concentrations to be extrapolated to target populations. The indirect effects of other environmental contaminants with similar biotransformation pathways, such as polycyclic aromatic hydrocarbons (PAH), could be assessed and quantified with this model and the results applied to a more complex biological hierarchy.

Aquatic Toxicology↗

Vision for a worldwide fluvial-sediment information network

The nations of the world suffer both from the deleterious effects of some natural and human-altered fluxes of fluvial sediment and a lack of consistent and reliable information on the temporal and spatial occurrence of fluvial sediments. Decades ago, this difficulty was unavoidable due to a lack of understanding of the magnitude and scope of environmental influences exerted by fluvial sediment coupled with a dearth of tools for monitoring and studying the data. Such is no longer the case. Fluvial sediment has a broad influence on the environment and humanity. Data needs that were once limited primarily to reservoir and channel maintenance now include issues associated with public water supply; contaminated sediment management; productivity of agricultural lands; stream restoration and watershed health; in-stream biotic stability; post-wildfire channel morphology; dam decommissioning, rehabilitation, or removal; and legal requirements for sediment management (Gray and Glysson, 2005). The adverse effects of poorly managed or unmanaged sediment movement related to these and other issues are well-known qualitatively, and in some cases quantitatively. For example, physical, chemical, and biological damages attributable to fluvial sediment in North America alone are now estimated to range between $20 billion and $50 billion annually (Pimental and others, 1995; Osterkamp and others, 1998; 2004). Capabilities for monitoring, analyzing, storing, and sharing fluvial-sediment data have been developed and, in many cases, are sufficiently mature for consideration for global utilization. Hence, there is not only a strong and expanding need for a global effort to gauge and understand fluvial-sediment characteristics and processes better, but the knowledge and tools to achieve these ends are largely available and ready for their applicability to be evaluated. Given the increasing importance of erosion and sediment processes for water-resources management, an International Sedimentation Initiative (ISI, 2007a), under the United Nations Educational, Scientific, and Cultural Organization’s International Hydrologic Programme (IHP, 2007) was adopted in 2004. The ISI, the focus of which is on sustainable water-resources management on the global scale, features six major activities and projects, which are listed as part of the section entitled, “Relation of the WoFSIN concept to the thrusts of the International Sedimentation Initiative,” that precedes the “Conclusions” section of this paper. Based on the need for more, and more consistent and reliable fluvial-sediment information and on the existence of the ISI and other international and national sediment programs, we envision the need for a Worldwide Fluvial Sediment-Information Network (WoFSIN) with a focus on data acquisition, storage, and dissemination globally. Envisioned components of a WoFSIN, administered largely via the Internet and relying mostly on the benefits derived from existing resources and programs, follow that summary. The goal of the WoFSIN is to maximize the availability and usefulness of the world’s historical and current fluvial-sediment and ancillary data through collaboration with existing programs so as to require few additional resources in the long-term. Thus, the WoFSIN concept was developed recognizing that informed resource management is predicated on the availability of adequate and reliable information. The WoFSIN is described in the ensuing sections in stand-alone fashion, followed by a section that describes the complementary aspects of the WoFSIN and the International Sediment Initiative. Thus, our first objective is to describe the fundamental components of a WoFSIN. Our second objective is to identify overlap or gaps between the WoFSIN and ISI concepts that might be useful in refining the ISI’s ability to meet its global mission to develop decision support for sediment management at the global scale more fully, cost-effectively, and (or) with enhanced quality.

Conference Paper↗

Establishing endangered species recovery criteria using predictive simulation modeling

Listing a species under the Endangered Species Act (ESA) and developing a recovery plan requires U.S. Fish and Wildlife Service to establish specific and measurable criteria for delisting. Generally, species are listed because they face (or are perceived to face) elevated risk of extinction due to issues such as habitat loss, invasive species, or other factors. Recovery plans identify recovery criteria that reduce extinction risk to an acceptable level. It logically follows that the recovery criteria, the defined conditions for removing a species from ESA protections, need to be closely related to extinction risk. Extinction probability is a population parameter estimated with a model that uses current demographic information to project the population into the future over a number of replicates, calculating the proportion of replicated populations that go extinct. We simulated extinction probabilities of piping plovers in the Great Plains and estimated the relationship between extinction probability and various demographic parameters. We tested the fit of regression models linking initial abundance, productivity, or population growth rate to extinction risk, and then, using the regression parameter estimates, determined the conditions required to reduce extinction probability to some pre-defined acceptable threshold. Binomial regression models with mean population growth rate and the natural log of initial abundance were the best predictors of extinction probability 50 years into the future. For example, based on our regression models, an initial abundance of approximately 2400 females with an expected mean population growth rate of 1.0 will limit extinction risk for piping plovers in the Great Plains to less than 0.048. Our method provides a straightforward way of developing specific and measurable recovery criteria linked directly to the core issue of extinction risk. Published by Elsevier Ltd.

Biological Conservation↗

Space use of wintering waterbirds in India: Influence of trophic ecology on home-range size

Relationship between species' home range and their other biological traits remains poorly understood, especially in migratory birds due to the difficulty associated with tracking them. Advances in satellite telemetry and remote sensing techniques have proved instrumental in overcoming such challenges. We studied the space use of migratory ducks through satellite telemetry with an objective of understanding the influence of body mass and feeding habits on their home-range sizes. We marked 26 individuals, representing five species of migratory ducks, with satellite transmitters during two consecutive winters in three Indian states. We used kernel methods to estimate home ranges and core use areas of these waterfowl, and assessed the influence of body mass and feeding habits on home-range size. Feeding habits influenced the home-range size of the migratory ducks. Carnivorous ducks had the largest home ranges, herbivorous ducks the smallest, while omnivorous species had intermediate home-ranges. Body mass did not explain variation in home-range size. To our knowledge, this is the first study of its kind on migratory ducks, and it has important implications for their conservation and management.

Current Zoology↗

Mechanistic variables can enhance predictive models of endotherm distributions: The American pika under current, past, and future climates

How climate constrains species’ distributions through time and space is an important question in the context of conservation planning for climate change. Despite increasing awareness of the need to incorporate mechanism into species distribution models (SDMs), mechanistic modeling of endotherm distributions remains limited in this literature. Using the American pika ( Ochotona princeps ) as an example, we present a framework whereby mechanism can be incorporated into endotherm SDMs. Pika distribution has repeatedly been found to be constrained by warm temperatures, so we used Niche Mapper, a mechanistic heat-balance model, to convert macroclimate data to pika-specific surface activity time in summer across the western United States. We then explored the difference between using a macroclimate predictor (summer temperature) and using a mechanistic predictor (predicted surface activity time) in SDMs. Both approaches accurately predicted pika presences in current and past climate regimes. However, the activity models predicted 8–19% less habitat loss in response to annual temperature increases of ~3–5 °C predicted in the region by 2070, suggesting that pikas may be able to buffer some climate change effects through behavioral thermoregulation that can be captured by mechanistic modeling. Incorporating mechanism added value to the modeling by providing increased confidence in areas where different modeling approaches agreed and providing a range of outcomes in areas of disagreement. It also provided a more proximate variable relating animal distribution to climate, allowing investigations into how unique habitat characteristics and intraspecific phenotypic variation may allow pikas to exist in areas outside those predicted by generic SDMs. Only a small number of easily obtainable data are required to parameterize this mechanistic model for any endotherm, and its use can improve SDM predictions by explicitly modeling a widely applicable direct physiological effect: climate-imposed restrictions on activity. This more complete understanding is necessary to inform climate adaptation actions, management strategies, and conservation plans.

Global Change Biology↗

Toxicity of copper sulfate and rotenone to Chinese mystery snail ( Bellamya chinensis )

The Chinese mystery snail ( Bellamya chinensis ) is a freshwater snail native to Southeast Asia, Japan, and Russia and is currently classified as an invasive species in at least 27 states in the USA. The species tolerates a wide range of environmental conditions, making management of established populations difficult. We tested the efficacy of two traditional chemical treatments, rotenone and copper sulfate, on the elimination of adult Chinese mystery snails in laboratory experiments. All snails (N=50) survived 72-hour exposure to rotenone-treated lake water, and 96% (N=25) survived 72-hour exposure to pre-determined rotenone concentrations of 0.25, 2.5, and 25.0 mg/L. All snails (N=10) survived exposure to 1.25 mg/L copper sulfate solution, 90% (N=10) survived exposure to 2.50 mg/L copper sulfate solution, and 80% (N=5) survived exposure to 5.0 mg/L copper sulfate solution. Neither rotenone nor copper sulfate effectively killed adult Chinese mystery snails in laboratory experiments, most likely due to their relatively large size, thick shell, and operculum. Therefore, it appears that populations will be very difficult to control once established, and management should focus on preventing additional spread or introductions of this species.

Management of Biological Invasions↗

New insight into the spawning behavior of lake trout, Salvelinus namaycush , from a recovering population in the Laurentian Great Lakes

Spawning behavior of lake trout, Salvelinus namaycush , is poorly understood, relative to stream-dwelling salmonines. Underwater video records of spawning in a recovering population from the Drummond Island Refuge (Lake Huron) represent the first reported direct observations of lake trout spawning in the Laurentian Great Lakes. These observations provide new insight into lake trout spawning behavior and expand the current conceptual model. Lake trout spawning consisted of at least four distinct behaviors: hovering, traveling, sinking, and gamete release. Hovering is a new courtship behavior that has not been previously described. The apparent concentration of hovering near the margin of the spawning grounds suggests that courtship and mate selection might be isolated from the spawning act (i.e., traveling, sinking, and gamete release). Moreover, we interpret jockeying for position displayed by males during traveling as a unique form of male-male competition that likely evolved in concert with the switch from redd-building to itinerant spawning in lake trout. Unlike previous models, which suggested that intra-sexual competition and mate selection do not occur in lake trout, our model includes both and is therefore consistent with evolutionary theory, given that the sex ratio on spawning grounds is skewed heavily towards males. The model presented in this paper is intended as a working hypothesis, and further revision may become necessary as we gain a more complete understanding of lake trout spawning behavior.

Environmental Biology of Fishes↗

Persistence and extirpation in invaded landscapes: patch characteristics and connectivity determine effects of non-native predatory fish on native salamanders

Studies have demonstrated negative effects of non-native, predatory fishes on native amphibians, yet it is still unclear why some amphibian populations persist, while others are extirpated, following fish invasion. We examined this question by developing habitat-based occupancy models for the long-toed salamander (Ambystoma macrodactylum) and nonnative fish using survey data from 1,749 water bodies across 470 catchments in the Northern Rocky Mountains, USA. We first modeled the habitat associations of salamanders at 468 fishless water bodies in 154 catchments where non-native fish were historically, and are currently, absent from the entire catchment. Wethen applied this habitat model to the complete data set to predict the probability of salamander occupancy in each water body, removing any effect of fish presence. Finally, we compared field-observed occurrences of salamanders and fish to modeled probability of salamander occupancy. Suitability models indicated that fish and salamanders had similar habitat preferences, possibly resulting in extirpations of salamander populations from entire catchments where suitable habitats were limiting. Salamanders coexisted with non-native fish in some catchments by using marginal quality, isolated (no inlet or outlet) habitats that remained fishless. They rarely coexisted with fish within individual water bodies and only where habitat quality was highest. Connectivity of water bodies via streams resulted in increased probability of fish invasion and consequently reduced probability of salamander occupancy.These results could be used to identify and prioritize catchments and water bodies where control measures would be most effective at restoring amphibian populations. Our approach could be useful as a framework for improved investigations into questions of persistence and extirpation of native species when non-native species have already become established.

Rocky Mountains↗

Forecasting sagebrush ecosystem components and greater sage-grouse habitat for 2050: learning from past climate patterns and Landsat imagery to predict the future

Sagebrush ( Artemisia spp.) ecosystems constitute the largest single North American shrub ecosystem and provide vital ecological, hydrological, biological, agricultural, and recreational ecosystem services. Disturbances have altered and reduced this ecosystem historically, but climate change may ultimately represent the greatest future risk. Improved ways to quantify, monitor, and predict climate-driven gradual change in this ecosystem is vital to its future management. We examined the annual change of Daymet precipitation (daily gridded climate data) and five remote sensing ecosystem sagebrush vegetation and soil components (bare ground, herbaceous, litter, sagebrush, and shrub) from 1984 to 2011 in southwestern Wyoming. Bare ground displayed an increasing trend in abundance over time, and herbaceous, litter, shrub, and sagebrush showed a decreasing trend. Total precipitation amounts show a downward trend during the same period. We established statistically significant correlations between each sagebrush component and historical precipitation records using a simple least squares linear regression. Using the historical relationship between sagebrush component abundance and precipitation in a linear model, we forecasted the abundance of the sagebrush components in 2050 using Intergovernmental Panel on Climate Change (IPCC) precipitation scenarios A1B and A2. Bare ground was the only component that increased under both future scenarios, with a net increase of 48.98 km 2 (1.1%) across the study area under the A1B scenario and 41.15 km 2 (0.9%) under the A2 scenario. The remaining components decreased under both future scenarios: litter had the highest net reductions with 49.82 km 2 (4.1%) under A1B and 50.8 km 2 (4.2%) under A2, and herbaceous had the smallest net reductions with 39.95 km 2 (3.8%) under A1B and 40.59 km 2 (3.3%) under A2. We applied the 2050 forecast sagebrush component values to contemporary (circa 2006) greater sage-grouse ( Centrocercus urophasianus ) habitat models to evaluate the effects of potential climate-induced habitat change. Under the 2050 IPCC A1B scenario, 11.6% of currently identified nesting habitat was lost, and 0.002% of new potential habitat was gained, with 4% of summer habitat lost and 0.039% gained. Our results demonstrate the successful ability of remote sensing based sagebrush components, when coupled with precipitation, to forecast future component response using IPCC precipitation scenarios. Our approach also enables future quantification of greater sage-grouse habitat under different precipitation scenarios, and provides additional capability to identify regional precipitation influence on sagebrush component response.

Wyoming↗

Barrier displacement on a neutral landscape: Towards a theory of continental biogeography

Macroevolutionary theory posits three processes leading to lineage diversification and the formation of regional biotas: dispersal (species geographic range expansion), speciation (species lineage splitting), and extinction (species lineage termination). The Theory of Island Biogeography (TIB) predicts species richness values using just two of these processes; dispersal and extinction. Yet most species on Earth live on continents or continental shelves, and the dynamics of evolutionary diversification at regional and continental scales are qualitatively different from those that govern the formation of species richness on biogeographic islands. Certain geomorphological processes operating perennially on continental platforms displace barriers to gene flow and organismal dispersal, and affect all three terms of macroevolutionary diversification. For example, uplift of a dissected landscape and river capture both merge and separate portions of adjacent areas, allowing dispersal and larger geographic ranges, vicariant speciation and smaller geographic ranges, and extinction when range sizes are subdivided below a minimum persistence threshold. The TIB also does not predict many biogeographic and phylogenetic patterns widely observed in continentally distributed taxa, including: 1, power function-like species-area relationships; 2, log-normal distribution of species geographic range sizes, in which most species have restricted ranges (are endemic) and few species have broad ranges (are cosmopolitan); 3, mid-domain effects with more species towards the geographic center, and more early-branching, species-poor clades towards the geographic periphery; 4, exponential rates of net diversification with log-linear accumulation of lineages through geological time; and 5, power function-like relationships between species-richness and clade diversity, in which most clades are species-poor and few clades are species-rich. Current theory does not provide a robust mechanistic framework to connect these seemingly disparate patterns. Here we present SEAMLESS (Spatially-Explicit Area Model of Landscape Evolution by SimulationS) that generates clade diversification by moving geographic barriers on a continuous, neutral landscape. SEAMLESS is a neutral Landscape Evolution Model (LEM) that treats species and barriers as functionally equivalent with respect to model parameters. SEAMLESS differs from other model-based biogeographic methods (e.g. Lagrange, GeoSSE, BayArea, BioGeoBEARS) by modeling properties of dispersal barriers rather than areas, and by modeling the evolution of species lineages on a continuous landscape, rather than the evolution of geographic ranges along branches of a phylogeny. SEAMLESS shows how dispersal is required to maintain species richness and avoid clade-wide extinction, demonstrates that ancestral range size does not predict species richness, and provides a unified explanation for the suite of commonly observed biogeographic and phylogenetic patterns listed above. SEAMLESS explains how a simple barrier-displacement mechanism affects lineage diversification under neutral conditions, and is advanced here towards the formulation of a general theory of continental biogeography.

Systematic Biology↗