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Multiple stressors in amphibian communities: Effects of chemical contamination, bullfrogs, and fish

A leading hypothesis of amphibian population declines is that combinations of multiple stressors contribute to declines. We examined the role that chemical contamination, competition, and predation play singly and in combination in aquatic amphibian communities. We exposed larvae of American toads (Bufo americanus), southern leopard frogs (Rana sphenocephala), and spotted salamanders (Ambystoma maculatum) to overwintered bullfrog tadpoles (R. catesbeiana), bluegill sunfish (Lepomis macrochirus), the insecticide carbaryl, and ammonium nitrate fertilizer in 1000-L mesocosms. Most significantly, our study demonstrated that the presence of multiple factors reduced survival of B. americanus and A. maculatum and lengthened larval periods of R. sphenocephala. The presence of bluegill had the largest impact on the community; it eliminated B. americanus and A. maculatum and reduced the abundance of R. sphenocephala. Chemical contaminants had the second strongest effect on the community with the insecticide, reducing A. maculatum abundance by 50% and increasing the mass of anurans (frogs and toads) at metamorphosis; the fertilizer positively influenced time and mass at metamorphosis for both anurans and A. maculatum. Presence of overwintered bullfrogs reduced mass and increased time to metamorphosis of anurans. While both bluegill and overwintered bullfrog tadpoles had negative effects on the amphibian community, they performed better in the presence of one another and in contaminated habitats. Our results indicate that predicting deleterious combinations from single-factor effects may not be straightforward. Our research supports the hypothesis that combinations of factors can negatively impact some amphibian species and could contribute to population declines. ?? 2007 by the Ecological Society of America.

Ecological Applications↗

Impacts of agricultural land use on biological integrity: A causal analysis

Agricultural land use has often been linked to nutrient enrichment, habitat degradation, hydrologic alteration, and loss of biotic integrity in streams. The U.S. Geological Survey's National Water Quality Assessment Program sampled 226 stream sites located in eight agriculture‐dominated study units across the United States to investigate the geographic variability and causes of agricultural impacts on stream biotic integrity. In this analysis we used structural equation modeling (SEM) to develop a national and set of regional causal models linking agricultural land use to measured instream conditions. We then examined the direct, indirect, and total effects of agriculture on biotic integrity as it acted through multiple water quality and habitat pathways. In our nation‐wide model, cropland affected benthic communities by both altering structural habitats and by imposing water quality‐related stresses. Region‐specific modeling demonstrated that geographic context altered the relative importance of causal pathways through which agricultural activities affected stream biotic integrity. Cropland had strong negative total effects on the invertebrate community in the national, Midwest, and Western models, but a very weak effect in the Eastern Coastal Plain model. In the Western Arid and Eastern Coastal Plain study regions, cropland impacts were transmitted primarily through dissolved water quality contaminants, but in the Midwestern region, they were transmitted primarily through particulate components of water quality. Habitat effects were important in the Western Arid model, but negligible in the Midwest and Eastern Coastal Plain models. The relative effects of riparian forested wetlands also varied regionally, having positive effects on biotic integrity in the Eastern Coastal Plain and Western Arid region models, but no statistically significant effect in the Midwest. These differences in response to cropland and riparian cover suggest that best management practices and planning for the mitigation of agricultural land use impacts on stream ecosystems should be regionally focused.

Ecological Applications↗

Could residual oil from the Exxon Valdez spill create a long-term population "sink" for sea otters in Alaska?

Over 20 years ago, the Exxon Valdez oil tanker spilled 42 million L of crude oil into the waters of Prince William Sound, Alaska, USA. At the time of the spill, the sea otter (Enhydra lutris) population inhabiting the spill area suffered substantial acute injuries and loss. Subsequent research has resulted in one of the best-studied species responses to an oil spill in history. However, the question remains: Is the spill still influencing the Prince William Sound sea otter population? Here we fit time-varying population models to data for the sea otter population of western Prince William Sound to quantify the duration and extent of mortality effects from the spill. We hypothesize that the patchy nature of residual oil left in the environment has created a source-sink population dynamic. We fit models using the age distributions of both living and dying animals and estimates of sea otter population size to predict the number of sea otters in the hypothesized sink population and the number lost to this sink due to chronic exposure to residual oil. Our results suggest that the sink population has remained at just over 900 individuals (95% CI: 606-960) between 1990 and 2009, during which time prime-age survival remained 2-6% below pre-spill levels. This reduced survival led to chronic losses of ???900 animals over the past two decades, which is similar in magnitude to the number of sea otter deaths documented in western Prince William Sound during the acute phase of the spill. However, the unaffected source population appears to be counterbalancing these losses, with the model indicating that the sea otter population increased from ???2150 individuals in 1990 to nearly 3000 in 2009. The most optimistic interpretation of our results suggests that mortality effects dissipated between 2005 and 2007. Our results suggest that residual oil can affect wildlife populations on time scales much longer than previously believed and that cumulative chronic effects can be as significant as acute effects. Further, source-sink population dynamics can explain the slow recovery observed in the spill-affected western Prince William Sound sea otter population and are consistent with available data. ??2011 by the Ecological Society of America.

Ecological Applications↗

Ecosystem development in roadside grasslands: Biotic control, plant-soil interactions, and dispersal limitations

Roadside grasslands undergoing secondary succession are abundant, and represent ecologically meaningful examples of novel, human‐created ecosystems. Interactions between plant and soil communities (hereafter plant–soil interactions) are of major importance in understanding the role of biotic control in ecosystem functioning, but little is known about these links in the context of ecosystem restoration and succession. The assessment of the key biotic communities and interactions driving ecosystem development will help practitioners to better allocate the limited resources devoted to roadside grassland restoration. We surveyed roadside grasslands from three successional stages (0–2, 7–9, and >20 years) in two Mediterranean regions of Spain. Structural equation modeling was used to evaluate how interactions between plants, biological soil crusts (BSCs), and soil microbial functional diversity (soil microorganisms) affect indicators of ecosystem development and restoration: plant similarity to the reference ecosystem, erosion control, and soil C storage and N accumulation. Changes in plant community composition along the successional gradient exerted the strongest influence on these indicators. High BSC cover was associated with high soil stability, and high soil microbial functional diversity from late‐successional stages was associated with high soil fertility. Contrary to our expectations, the indirect effects of plants, mediated by either BSCs or soil microorganisms, were very weak in both regions, suggesting a minor role for plant–soil interactions upon ecosystem development indicators over long periods. Our results suggest that natural vegetation dynamics effectively improved ecosystem development within a time frame of 20 years in the grasslands evaluated. They also indicate that this time could be shortened if management actions focus on: (1) maintaining well‐conserved natural areas close to roadsides to enhance plant compositional changes towards late‐successional stages, (2) increasing BSC cover in areas under strong erosion risk, to avoid soil loss, and (3) enhancing soil microbial functional diversity in resource‐limited areas, to enhance soil C and N accumulation.

Ecological Applications↗

Weather effects on avian breeding performance and implications of climate change

The influence of recent climate change on the world’s biota has manifested broadly, resulting in latitudinal range shifts, advancing dates of arrival of migrants and onset of breeding, and altered community relationships. Climate change elevates conservation concerns worldwide because it will likely exacerbate a broad range of identified threats to animal populations. In the past few decades, grassland birds have declined faster than other North American avifauna, largely due to habitat threats such as the intensification of agriculture. We examine the effects of local climatic variations on the breeding performance of a bird endemic to the shortgrass prairie, the Lark Bunting ( Calamospiza melanocorys ) and discuss the implications of our findings relative to future climate predictions. Clutch size, nest survival, and productivity all positively covaried with seasonal precipitation, yet relatively intense daily precipitation events temporarily depressed daily survival of nests. Nest survival was positively related to average temperatures during the breeding season. Declining summer precipitation may reduce the likelihood that Lark Buntings can maintain stable breeding populations in eastern Colorado although average temperature increases of up to 38C (within the range of this study) may ameliorate declines in survival expected with drier conditions. Historic climate variability in the Great Plains selects for a degree of vagility and opportunism rather than strong site fidelity and specific adaptation to local environments. These traits may lead to northerly shifts in distribution if climatic and habitat conditions become less favorable in the drying southern regions of the Great Plains. Distributional shifts in Lark Buntings could be constrained by future changes in land use, agricultural practices, or vegetative communities that result in further loss of shortgrass prairie habitats.

Colorado↗

The efficacy of salvage logging in reducing subsequent fire severity in conifer-dominated forests of Minnesota, USA

Although primarily used to mitigate economic losses following disturbance, salvage logging has also been justified on the basis of reducing fire risk and fire severity; however, its ability to achieve these secondary objectives remains unclear. The patchiness resulting from a sequence of recent disturbances—blowdown, salvage logging, and wildfire—provided an excellent opportunity to assess the impacts of blowdown and salvage logging on wildfire severity. We used two fire‐severity assessments (tree‐crown and forest‐floor characteristics) to compare post‐wildfire conditions among three treatment combinations (Blowdown–Salvage–Fire, Blowdown–Fire, and Fire only). Our results suggest that salvage logging reduced the intensity (heat released) of the subsequent fire. However, its effect on severity (impact to the system) differed between the tree crowns and forest floor: tree‐crown indices suggest that salvage logging decreased fire severity (albeit with modest statistical support), while forest‐floor indices suggest that salvage logging increased fire severity. We attribute the latter finding to the greater exposure of mineral soil caused by logging operations; once exposed, soils are more likely to register the damaging effects of fire, even if fire intensity is not extreme. These results highlight the important distinction between fire intensity and severity when formulating post‐disturbance management prescriptions.

Minnesota↗

Carbon stocks across a chronosequence of thinned and unmanaged red pine ( Pinus resinosa ) stands

Forests function as a major global C sink, and forest management strategies that maximize C stocks offer one possible means of mitigating the impacts of increasing anthropogenic CO 2 emissions. We studied the effects of thinning, a common management technique in many forest types, on age-related trends in C stocks using a chronosequence of thinned and unmanaged red pine ( Pinus resinosa ) stands ranging from 9 to 306 years old. Live tree C stocks increased with age to a maximum near the middle of the chronosequence in unmanaged stands, and increased across the entire chronosequence in thinned stands. C in live understory vegetation and C in the mineral soil each declined rapidly with age in young stands but changed relatively little in middle-aged to older stands regardless of management. Forest floor C stocks increased with age in unmanaged stands, but forest floor C decreased with age after the onset of thinning around age 40 in thinned stands. Deadwood C was highly variable, but decreased with age in thinned stands. Total ecosystem C increased with stand age until approaching an asymptote around age 150. The increase in total ecosystem C was paralleled by an age-related increase in total aboveground C, but relatively little change in total belowground C. Thinning had surprisingly little impact on total ecosystem C stocks, but it did modestly alter age-related trends in total ecosystem C allocation between aboveground and belowground pools. In addition to characterizing the subtle differences in C dynamics between thinned and unmanaged stands, these results suggest that C accrual in red pine stands continues well beyond the 60–100 year management rotations typical for this system. Management plans that incorporate longer rotations and thinning in some stands could play an important role in maximizing C stocks in red pine forests while meeting other objectives including timber extraction, biodiversity conservation, restoration, and fuel reduction goals.

Ecological Applications↗

Comparing modern and presettlement forest dynamics of a subboreal wilderness: Does spruce budworm enhance fire risk?

Insect disturbance is often thought to increase fire risk through enhanced fuel loadings, particularly in coniferous forest ecosystems. Yet insect disturbances also affect successional pathways and landscape structure that interact with fire disturbances (and vice-versa) over longer time scales. We applied a landscape succession and disturbance model (LANDIS-II) to evaluate the relative strength of interactions between spruce budworm ( Choristoneura fumiferana ) outbreaks and fire disturbances in the Boundary Waters Canoe Area (BWCA) in northern Minnesota (USA). Disturbance interactions were evaluated for two different scenarios: presettlement forests and fire regimes vs. contemporary forests and fire regimes. Forest composition under the contemporary scenario trended toward mixtures of deciduous species (primarily Betula papyrifera and Populus spp.) and shade-tolerant conifers ( Picea mariana, Abies balsamea, Thuja occidentalis ), with disturbances dominated by a combination of budworm defoliation and high-severity fires. The presettlement scenario retained comparatively more “big pines” (i.e., Pinus strobus, P. resinosa ) and tamarack ( L. laricina ), and experienced less budworm disturbance and a comparatively less-severe fire regime. Spruce budworm disturbance decreased area burned and fire severity under both scenarios when averaged across the entire 300-year simulations. Contrary to past research, area burned and fire severity during outbreak decades were each similar to that observed in non-outbreak decades. Our analyses suggest budworm disturbances within forests of the BWCA have a comparatively weak effect on long-term forest composition due to a combination of characteristics. These include strict host specificity, fine-scaled patchiness created by defoliation damage, and advance regeneration of its primary host, balsam fir ( A. balsamea ) that allows its host to persist despite repeated disturbances. Understanding the nature of the three-way interaction between budworm, fire, and composition has important ramifications for both fire mitigation strategies and ecosystem restoration initiatives. We conclude that budworm disturbance can partially mitigate long-term future fire risk by periodically reducing live ladder fuel within the mixed forest types of the BWCA but will do little to reverse the compositional trends caused in part by reduced fire rotations.

Minnesota↗

Geologic processes influence the effects of mining on aquatic ecosystems

Geologic processes strongly influence water and sediment quality in aquatic ecosystems but rarely are geologic principles incorporated into routine biomonitoring studies. We test if elevated concentrations of metals in water and sediment are restricted to streams downstream of mines or areas that may discharge mine wastes. We surveyed 198 catchments classified as “historically mined” or “unmined,” and based on mineral-deposit criteria, to determine whether water and sediment quality were influenced by naturally occurring mineralized rock, by historical mining, or by a combination of both. By accounting for different geologic sources of metals to the environment, we were able to distinguish aquatic ecosystems limited by metals derived from natural processes from those due to mining. Elevated concentrations of metals in water and sediment were not restricted to mined catchments; depauperate aquatic communities were found in unmined catchments. The type and intensity of hydrothermal alteration and the mineral deposit type were important determinants of water and sediment quality as well as the aquatic community in both mined and unmined catchments. This study distinguished the effects of different rock types and geologic sources of metals on ecosystems by incorporating basic geologic processes into reference and baseline site selection, resulting in a refined assessment. Our results indicate that biomonitoring studies should account for natural sources of metals in some geologic environments as contributors to the effect of mines on aquatic ecosystems, recognizing that in mining-impacted drainages there may have been high pre-mining background metal concentrations.

Ecological Applications↗

Carbon storage, timber production, and biodiversity: comparing ecosystem services with multi-criteria decision analysis

Increasingly, land managers seek ways to manage forests for multiple ecosystem services and functions, yet considerable challenges exist in comparing disparate services and balancing trade-offs among them. We applied multi-criteria decision analysis (MCDA) and forest simulation models to simultaneously consider three objectives: (1) storing carbon, (2) producing timber and wood products, and (3) sustaining biodiversity. We used the Forest Vegetation Simulator (FVS) applied to 42 northern hardwood sites to simulate forest development over 100 years and to estimate carbon storage and timber production. We estimated biodiversity implications with occupancy models for 51 terrestrial bird species that were linked to FVS outputs. We simulated four alternative management prescriptions that spanned a range of harvesting intensities and forest structure retention. We found that silvicultural approaches emphasizing less frequent harvesting and greater structural retention could be expected to achieve the greatest net carbon storage but also produce less timber. More intensive prescriptions would enhance biodiversity because positive responses of early successional species exceeded negative responses of late successional species within the heavily forested study area. The combinations of weights assigned to objectives had a large influence on which prescriptions were scored as optimal. Overall, we found that a diversity of silvicultural approaches is likely to be preferable to any single approach, emphasizing the need for landscape-scale management to provide a full range of ecosystem goods and services. Our analytical framework that combined MCDA with forest simulation modeling was a powerful tool in understanding trade-offs among management objectives and how they can be simultaneously accommodated.

Ecological Applications↗

Emerging prion disease drives host selection in a wildlife population

Infectious diseases are increasingly recognized as an important force driving population dynamics, conservation biology, and natural selection in wildlife populations. Infectious agents have been implicated in the decline of small or endangered populations and may act to constrain population size, distribution, growth rates, or migration patterns. Further, diseases may provide selective pressures that shape the genetic diversity of populations or species. Thus, understanding disease dynamics and selective pressures from pathogens is crucial to understanding population processes, managing wildlife diseases, and conserving biological diversity. There is ample evidence that variation in the prion protein gene (PRNP) impacts host susceptibility to prion diseases. Still, little is known about how genetic differences might influence natural selection within wildlife populations. Here we link genetic variation with differential susceptibility of white-tailed deer to chronic wasting disease (CWD), with implications for fitness and disease-driven genetic selection. We developed a single nucleotide polymorphism (SNP) assay to efficiently genotype deer at the locus of interest (in the 96th codon of the PRNP gene). Then, using a Bayesian modeling approach, we found that the more susceptible genotype had over four times greater risk of CWD infection; and, once infected, deer with the resistant genotype survived 49% longer (8.25 more months). We used these epidemiological parameters in a multi-stage population matrix model to evaluate relative fitness based on genotype-specific population growth rates. The differences in disease infection and mortality rates allowed genetically resistant deer to achieve higher population growth and obtain a long-term fitness advantage, which translated into a selection coefficient of over 1% favoring the CWD-resistant genotype. This selective pressure suggests that the resistant allele could become dominant in the population within an evolutionarily short time frame. Our work provides a rare example of a quantifiable disease-driven selection process in a wildlife population, demonstrating the potential for infectious diseases to alter host populations. This will have direct bearing on the epidemiology, dynamics, and future trends in CWD transmission and spread. Understanding genotype-specific epidemiology will improve predictive models and inform management strategies for CWD-affected cervid populations.

Ecological Applications↗

Cougar space use and movements in the wildland-urban landscape of western Washington

The wildland-urban interface lies at the confluence of human-dominated and wild landscapes, creating a number of management and conservation challenges. Because wildlife ecology, behavior, and evolution at this interface are shaped by both natural and human phenomena, this requires greater understanding of how diverse factors affect ecosystem and population processes. We illustrate the challenge of understanding and managing a frequent and often undesired inhabitant of the wildland-urban landscape, the cougar (Puma concolor). In wildland and residential areas of western Washington State, USA, we captured and radiotracked 27 cougars to model space use and understand the role of landscape features in interactions (sightings, encounters, and depredations) between cougars and humans. Resource utilization functions (RUFs) identified cougar use of areas with features that were probably attractive to prey, influential on prey vulnerability, and associated with limited or no residential development. Early-successional forest (+), conifer forest (+), distance to road (-), residential density (-), and elevation (-) were significant positive and negative predictors of use for the population, whereas use of other landscape features was highly variable. Space use and movement rates in wildland and residential areas were similar because cougars used wildland-like forest patches, reserves, and corridors in residential portions of their home range. The population RUF was a good predictor of confirmed cougar interactions, with 72% of confirmed reports occurring in the 50% of the landscape predicted to be medium-high and high cougar use areas. We believe that there is a threshold residential density at which the level of development modifies the habitat but maintains enough wildland characteristics to encourage moderate levels of cougar use and maximize the probability of interaction. Wildlife managers trying to reduce interactions between cougars and people should incorporate information on spatial ecology and landscape characteristics to identify areas with the highest overlap of human and cougar use to focus management, education, and landscape planning. Resource utilization functions provide a proactive tool to guide these activities for improved coexistence with wildlife using both wildland and residential portions of the landscape. ??2011 by the Ecological Society of America.

Ecological Applications↗

Insights and issues with simulating terrestrial DOC loading of Arctic river networks

Terrestrial carbon dynamics influence the contribution of dissolved organic carbon (DOC) to river networks in addition to hydrology. In this study, we use a biogeochemical process model to simulate the lateral transfer of DOC from land to the Arctic Ocean via riverine transport. We estimate that, over the 20th century, the pan-Arctic watershed has contributed, on average, 32 Tg C/yr of DOC to river networks emptying into the Arctic Ocean with most of the DOC coming from the extensive area of boreal deciduous needle-leaved forests and forested wetlands in Eurasian watersheds. We also estimate that the rate of terrestrial DOC loading has been increasing by 0.037 Tg C/yr2 over the 20th century primarily as a result of climate-induced increases in water yield. These increases have been offset by decreases in terrestrial DOC loading caused by wildfires. Other environmental factors (CO2 fertilization, ozone pollution, atmospheric nitrogen deposition, timber harvest, agriculture) are estimated to have relatively small effects on terrestrial DOC loading to Arctic rivers. The effects of the various environmental factors on terrestrial carbon dynamics have both offset and enhanced concurrent effects on hydrology to influence terrestrial DOC loading and may be changing the relative importance of terrestrial carbon dynamics on this carbon flux. Improvements in simulating terrestrial DOC loading to pan-Arctic rivers in the future will require better information on the production and consumption of DOC within the soil profile, the transfer of DOC from land to headwater streams, the spatial distribution of precipitation and its temporal trends, carbon dynamics of larch-dominated ecosystems in eastern Siberia, and the role of industrial organic effluents on carbon budgets of rivers in western Russia.

Ecological Applications↗

Predicted eelgrass response to sea level rise and its availability to foraging Black Brant in Pacific coast estuaries

Managers need to predict how animals will respond to habitat redistributions caused by climate change. Our objective was to model the effects of sea level rise on total eelgrass ( Zostera marina ) habitat area and on the amount of that area that is accessible to Brant geese ( Branta bernicla ), specialist grazers of eelgrass. Digital elevation models were developed for seven estuaries from Alaska, Washington, California (USA), and Mexico. Scenarios of future total eelgrass area were derived from combinations of estuarine specific sediment and tectonic rates (i.e., bottom change rate) with three rates of eustatic sea level rise (ESLR). Percentages of total eelgrass areas that were accessible to foraging Brant were determined for December when the birds overwinter at more southerly sites and in April as they move north to sites where they build body stores on their way to nesting areas in Alaska. The modeling showed that accessible eelgrass area could be lower than total area due to how daytime low-tide height, eelgrass shoot length, and the upper elevation of eelgrass determined Brant-reaching depth. Projections of future eelgrass area indicated that present-day ESLR (2.8 mm/yr) and bottom change rates should sustain the current pattern of estuarine use by Brant except in Morro Bay, where use should decrease because eelgrass is being ejected from this estuary by a positive bottom change rate. Higher ESLR rates (6.3 and 12.7 mm/yr) should result in less Brant use of estuaries at the northern and southern ends of the flyway, particularly during the winter, but more use of mid-latitude estuaries. The capacity of mid-latitude estuaries to function as Brant feeding refugia, or for these estuaries and Izembek Lagoon to provide drift rather than attached leaves, is eventually limited by the decrease in total eelgrass area, which is a result of a light extinction affect on the eelgrass, or the habitat being pushed out of the estuary by positive tectonic rates. Management responses are limited to the increase or decrease of sediment supply and the relocation of levees to allow for upslope migration of eelgrass habitat.

Alaska;California;Washington↗

Allowable levels of take for the trade in Nearctic songbirds

The take of Nearctic songbirds for the caged-bird trade is an important cultural and economic activity in Mexico, but its sustainability has been questioned. We relied on the theta-logistic population model to explore options for setting allowable levels of take for 11 species of passerines that were subject to legal take in Mexico in 2010. Because estimates of population size necessary for making periodic adjustments to levels of take are not routinely available, we examined the conditions under which a constant level of take might contribute to population depletion (i.e., a population below its level of maximum net productivity). The chance of depleting a population is highest when levels of take are based on population sizes that happen to be much lower or higher than the level of maximum net productivity, when environmental variation is relatively high and serially correlated, and when the interval between estimation of population size is relatively long (≥5 years). To estimate demographic rates of songbirds involved in the Mexican trade we relied on published information and allometric relationships to develop probability distributions for key rates, and then sampled from those distributions to characterize the uncertainty in potential levels of take. Estimates of the intrinsic rate of growth (r) were highly variable, but median estimates were consistent with those expected for relatively short-lived, highly fecund species. Allowing for the possibility of nonlinear density dependence generally resulted in allowable levels of take that were lower than would have been the case under an assumption of linearity. Levels of take authorized by the Mexican government in 2010 for the 11 species we examined were small in comparison to relatively conservative allowable levels of take (i.e., those intended to achieve 50% of maximum sustainable yield). However, the actual levels of take in Mexico are unknown and almost certainly exceed the authorized take. Also, the take of Nearctic songbirds in other Latin American and Caribbean countries ultimately must be considered in assessing population-level impacts.

Ecological Applications↗

Trajectory of the arctic as an integrated system

Although much remains to be learned about the Arctic and its component processes, many of the most urgent scientific, engineering, and social questions can only be approached through a broader system perspective. Here, we address interactions between components of the Arctic System and assess feedbacks and the extent to which feedbacks (1) are now underway in the Arctic; and (2) will shape the future trajectory of the Arctic system. We examine interdependent connections among atmospheric processes, oceanic processes, sea-ice dynamics, marine and terrestrial ecosystems, land surface stocks of carbon and water, glaciers and ice caps, and the Greenland ice sheet. Our emphasis on the interactions between components, both historical and anticipated, is targeted on the feedbacks, pathways, and processes that link these different components of the Arctic system. We present evidence that the physical components of the Arctic climate system are currently in extreme states, and that there is no indication that the system will deviate from this anomalous trajectory in the foreseeable future. The feedback for which the evidence of ongoing changes is most compelling is the surface albedo-temperature feedback, which is amplifying temperature changes over land (primarily in spring) and ocean (primarily in autumn-winter). Other feedbacks likely to emerge are those in which key processes include surface fluxes of trace gases, changes in the distribution of vegetation, changes in surface soil moisture, changes in atmospheric water vapor arising from higher temperatures and greater areas of open ocean, impacts of Arctic freshwater fluxes on the meridional overturning circulation of the ocean, and changes in Arctic clouds resulting from changes in water vapor content.

Ecological Applications↗

Per capita invasion probabilities: an empirical model to predict rates of invasion via ballast water

Ballast water discharges are a major source of species introductions into marine and estuarine ecosystems. To mitigate the introduction of new invaders into these ecosystems, many agencies are proposing standards that establish upper concentration limits for organisms in ballast discharge. Ideally, ballast discharge standards will be biologically defensible and adequately protective of the marine environment. We propose a new technique, the per capita invasion probability (PCIP), for managers to quantitatively evaluate the relative risk of different concentration-based ballast water discharge standards. PCIP represents the likelihood that a single discharged organism will become established as a new nonindigenous species. This value is calculated by dividing the total number of ballast water invaders per year by the total number of organisms discharged from ballast. Analysis was done at the coast-wide scale for the Atlantic, Gulf, and Pacific coasts, as well as the Great Lakes, to reduce uncertainty due to secondary invasions between estuaries on a single coast. The PCIP metric is then used to predict the rate of new ballast-associated invasions given various regulatory scenarios. Depending upon the assumptions used in the risk analysis, this approach predicts that approximately one new species will invade every 10–100 years with the International Maritime Organization (IMO) discharge standard of <10 organisms with body size >50 μm per m 3 of ballast. This approach resolves many of the limitations associated with other methods of establishing ecologically sound discharge standards, and it allows policy makers to use risk-based methodologies to establish biologically defensible discharge standards.

Ecological Applications↗

Approaches to setting organism-based ballast water discharge standards

As a vector by which foreign species invade coastal and freshwater waterbodies, ballast water discharge from ships is recognized as a major environmental threat. The International Maritime Organization (IMO) drafted an international treaty establishing ballast water discharge standards based on the number of viable organisms per volume of ballast discharge for different organism size classes. Concerns that the IMO standards are not sufficiently protective have initiated several state and national efforts in the United States to develop more stringent standards. We evaluated seven approaches to establishing discharge standards for the >50-μm size class: (1) expert opinion/management consensus, (2) zero detectable living organisms, (3) natural invasion rates, (4) reaction–diffusion models, (5) population viability analysis (PVA) models, (6) per capita invasion probabilities (PCIP), and (7) experimental studies. Because of the difficulty in synthesizing scientific knowledge in an unbiased and transparent fashion, we recommend the use of quantitative models instead of expert opinion. The actual organism concentration associated with a “zero detectable organisms” standard is defined by the statistical rigor of its monitoring program; thus it is not clear whether such a standard is as stringent as other standards. For several reasons, the natural invasion rate, reaction–diffusion, and experimental approaches are not considered suitable for generating discharge standards. PVA models can be used to predict the likelihood of establishment of introduced species but are limited by a lack of population vital rates for species characteristic of ballast water discharges. Until such rates become available, PVA models are better suited to evaluate relative efficiency of proposed standards rather than predicting probabilities of invasion. The PCIP approach, which is based on historical invasion rates at a regional scale, appears to circumvent many of the indicated problems, although it may underestimate invasions by asexual and parthenogenic species. Further research is needed to better define propagule dose–responses, densities at which Allee effects occur, approaches to predicting the likelihood of invasion from multi-species introductions, and generation of formal comparisons of approaches using standardized scenarios.

Ecological Applications↗