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Hierarchical modeling of an invasive spread: The eurasian collared-dove streptopelia decaocto in the United States

Invasive species are regularly claimed as the second threat to biodiversity. To apply a relevant response to the potential consequences associated with invasions (e.g., emphasize management efforts to prevent new colonization or to eradicate the species in places where it has already settled), it is essential to understand invasion mechanisms and dynamics. Quantifying and understanding what influences rates of spatial spread is a key research area for invasion theory. In this paper, we develop a model to account for occupancy dynamics of an invasive species. Our model extends existing models to accommodate several elements of invasive processes; we chose the framework of hierarchical modeling to assess site occupancy status during an invasion. First, we explicitly accounted for spatial structure and how distance among sites and position relative to one another affect the invasion spread. In particular, we accounted for the possibility of directional propagation and provided a way of estimating the direction of this possible spread. Second, we considered the influence of local density on site occupancy. Third, we decided to split the colonization process into two subprocesses, initial colonization and recolonization, which may be ground-breaking because these subprocesses may exhibit different relationships with environmental variations (such as density variation) or colonization history (e.g., initial colonization might facilitate further colonization events). Finally, our model incorporates imperfection in detection, which might be a source of substantial bias in estimating population parameters. We focused on the case of the Eurasian Collared-Dove (Streptopelia decaocto) and its invasion of the United States since its introduction in the early 1980s, using data from the North American BBS (Breeding Bird Survey). The Eurasian Collared-Dove is one of the most successful invasive species, at least among terrestrial vertebrates. Our model provided estimation of the spread direction consistent with empirical observations. Site persistence probability exhibits a quadratic response to density. We also succeeded at detecting differences in the relationship between density and initial colonization vs. recolonization probabilities. We provide a map of sites that may be colonized in the future as an example of possible practical application of our work. ?? 2011 by the Ecological Society of America.

Ecological Applications

Niche evolution and adaptive radiation: Testing the order of trait divergence

In the course of an adaptive radiation, the evolution of niche parameters is of particular interest for understanding modes of speciation and the consequences for coexistence of related species within communities. We pose a general question: In the course of an evolutionary radiation, do traits related to within-community niche differences (?? niche) evolve before or after differentiation of macrohabitat affinity or climatic tolerances (?? niche)? Here we introduce a new test to address this question, based on a modification of the method of independent contrasts. The divergence order test (DOT) is based on the average age of the nodes on a tree, weighted by the absolute magnitude of the contrast at each node for a particular trait. The comparison of these weighted averages reveals whether large divergences for one trait have occurred earlier or later in the course of diversification, relative to a second trait; significance is determined by bootstrapping from maximum-likelihood ancestral state reconstructions. The method is applied to the evolution of Ceanothus, a woody plant group in California, in which co-occurring species exhibit significant differences in a key leaf trait (specific leaf area) associated with contrasting physiological and life history strategies. Co-occurring species differ more for this trait than expected under a null model of community assembly. This ?? niche difference evolved early in the divergence of two major subclades within Ceanothus, whereas climatic distributions (?? niche traits) diversified later within each of the subclades. However, rapid evolution of climate parameters makes inferences of early divergence events highly uncertain, and differentiation of the ?? niche might have taken place throughout the evolution of the group, without leaving a clear phylogenetic signal. Similar patterns observed in several plant and animal groups suggest that early divergence of ?? niche traits might be a common feature of niche evolution in many adaptive radiations. ?? 2006 by the Ecological Society of America.

Ecology

Simulating the effects of frequent fire on southern california coastal shrublands

Fire disturbance is a primary agent of change in the mediterranean-climate chaparral shrublands of southern California, USA. However, fire frequency has been steadily increasing in coastal regions due to ignitions at the growing wildland-urban interface. Although chaparral is resilient to a range of fire frequencies, successively short intervals between fires can threaten the persistence of some species, and the effects may differ according to plant functional type. California shrublands support high levels of biological diversity, including many endangered and endemic species. Therefore, it is important to understand the long-term effects of altered fire regimes on these communities. A spatially explicit simulation model of landscape disturbance and succession (LANDIS) was used to predict the effects of frequent fire on the distribution of dominant plant functional types in a study area administered by the National Park Service. Shrubs dependent on fire-cued seed germination were most sensitive to frequent fire and lost substantial cover to other functional types, including drought-deciduous subshrubs that typify coastal sage scrub and nonnative annual grasses. Shrubs that resprout were favored by higher fire frequencies and gained in extent under these treatments. Due to this potential for vegetation change, caution is advised against the widespread use of prescribed fire in the region. ?? 2006 by the Ecological Society of America.

Ecological Applications

Additive effects of vertebrate predators on insects in a Puerto Rican coffee plantation

A variety of studies have established the value of shaded coffee plantations as habitat for birds. While the value of birds as biological controls in coffee has received some attention, the interactions between birds and other predators of insects have not been tested. We used exclosures to examine the effects of vertebrate predators on the arthropods associated with coffee, in particular the coffee leafminer (Leucoptera coffeella) and the flatid planthopper Petrusa epilepsis, in a shaded coffee plantation in Puerto Rico. We used a 2 x 2 factorial design with four treatments: exclusion of birds, lizards, birds and lizards, and control (no exclusion). Abundance of insects >5 mm increased when birds or both birds and lizards were removed. Birds and lizards had an additive effect for insects <5 mm and for all insects combined. Coffee leafminers showed a weak response to removal of predators while planthopper abundance increased significantly in the absence of avian predators. Arthropod predators and parasitoids did not differ significantly between treatments. Our findings suggest that vertebrate insectivores have an additive effect on insects in coffee and may help control abundances of some coffee pests. Equally important, we present evidence suggesting that they do not interfere with other known natural enemies of coffee pests. ?? 2006 by the Ecological Society of America.

Ecological Applications

Book review: Behavioral ecology of the eastern red-backed salamander: 50 years of research

In commemoration of the 100th anniversary of the British Ecological Society, Sutherland et al. (2013) identified 100 questions of fundamental significance in “pure” (i.e., not applied) ecology. A somewhat unexpected outcome of these authors’ exercise was the realization that, after 100 years of comprehensive, intensive scientific research, there remained “profound knowledge gaps” in ecology, such as a clear understanding of “the central mechanisms driving ecosystems…communities…, and even population dynamics.” Animal behavior (along with other attributes such as physiology and genetics) is such a mechanism that can structure ecological interactions, and the study of behavioral ecology provides important insights into many fundamental ecological phenomena. For example, the well-known historical characterization of ecology as the study of the distribution and abundance of organisms (Andrewartha and Birch 1954) invokes numerous questions, such as: what factors influence coexistence among competing species, or between predators and their prey? Ultimately, the answers to these and other questions are best addressed with fine-scale, mechanistic studies of habitat selection, foraging behavior/prey selection, and movement/dispersal behavior. Similarly, at the population level, insight into the spatial distribution of individuals could be gained with studies of territoriality, dominance hierarchies, and even mate choice.

Herpetological Review

Quantifying restoration success and recovery in a metal-polluted stream: A 17-year assessment of physicochemical and biological responses

Evaluating the effectiveness of stream restoration is often challenging because of the lack of pre-treatment data, narrow focus on physicochemical measures and insufficient post-restoration monitoring. Even when these fundamental elements are present, quantifying restoration success is difficult because of the challenges associated with distinguishing treatment effects from seasonal variation, episodic events and long-term climatic changes.2. We report results of one of the most comprehensive and continuous records of physical, chemical and biological data available to assess restoration success for a stream ecosystem in North America. Over a 17 year period we measured seasonal and annual changes in metal concentrations, physicochemical characteristics, macroinvertebrate communities, and brown trout Salmo trutta populations in the Arkansas River, a metal-contaminated stream in Colorado, USA.3. Although we observed significant improvements in water quality after treatment, the effectiveness of restoration varied temporally, spatially and among biological response variables. The fastest recovery was observed at stations where restoration eliminated point sources of metal contamination. Recovery of macroinvertebrates was significantly delayed at some stations because of residual sediment contamination and because extreme seasonal and episodic variation in metal concentrations prevented recolonization by sensitive species. Synthesis and applications. Because recovery trajectories after the removal of a stressor are often complex or nonlinear, long-term studies are necessary to assess restoration success within the context of episodic events and changes in regional climate. The observed variation in recovery among chemical and biological endpoints highlights the importance of developing objective criteria to assess restoration success. Although the rapid response of macroinvertebrates to reduced metal concentrations is encouraging, we have previously demonstrated that benthic communities from the Arkansas River remained susceptible to other novel anthropogenic stressors. We suggest that the resistance or resilience of benthic macroinvertebrate communities to novel stressors may be effective indicators of restoration success that can account for the non-additive (e.g. synergistic) nature of compound perturbations. ?? 2010 The Authors. Journal compilation ?? 2010 British Ecological Society.

Journal of Applied Ecology

Scaling the effects of moose browsing on forage distribution, from the geometry of plant canopies to landscapes

Landscape heterogeneity influences large herbivores by altering their feeding rates, but as herbivores attempt to maximize feeding rates they also create spatial heterogeneity by altering plant growth. Herbivore feeding rates thus provide a quantitative link between the causes and consequences of spatial heterogeneity in herbivore-dominated ecosystems. The fractal geometry of plant canopies determines both the density and mass of twigs available to foraging herbivores. These properties determine a threshold distance between plants (d * ) that distinguishes the mechanisms regulating herbivore intake rates. When d * is greater than the actual distance between plants (d), intake is regulated by the rate of food processing in the mouth. But when d * < d, intake is regulated by the rate at which the herbivore encounters new plants. Alterations to plant geometry due to past browsing could change the rate at which herbivores encounter and process bites of plant tissue, modify d * relative to d, and thus change intake rates and the distribution of mechanisms regulating it across landscapes. We measured changes in the geometry of aspen (Populus tremuloides) and balsam fir (Abies balsamea) saplings along gradients of moose browsing from 2001 to 2005 at Isle Royale National Park, Michigan, USA. For aspen saplings, fractal dimension of bite density, bite mass, and forage biomass responded quadratically to increasing moose browsing and were greatest at -3-4 g-g.m-2.yr" 1 consumption. For balsam fir, in contrast, these same measures declined steadily with increasing moose browsing. The different responses of plant canopies to increased browsing altered d * around plants. In summer, d * > d for aspen saplings at all prior consumption levels. Food processing therefore regulated summer moose feeding rates across our landscapes. In winter, changes in bite mass due to past browsing were sufficient to cause d * < d for aspen and balsam fir. Therefore, travel velocity and food processing jointly regulated intake rate during winter. Browsing-induced changes in the small-scale geometry of plant canopies can determine intake rate at larger spatial scales by changing d * relative to d and, hence, which mechanisms determine intake rate, essentially altering how herbivores sense the distribution of their food resources. ?? 2009 by the Ecological Society of America.

Ecological Monographs

Spatial and temporal patterns of chronic wasting disease: Fine-scale mapping of a wildlife epidemic in Wisconsin

Emerging infectious diseases threaten wildlife populations and human health. Understanding the spatial distributions of these new diseases is important for disease management and policy makers; however, the data are complicated by heterogeneities across host classes, sampling variance, sampling biases, and the space-time epidemic process. Ignoring these issues can lead to false conclusions or obscure important patterns in the data, such as spatial variation in disease prevalence. Here, we applied hierarchical Bayesian disease mapping methods to account for risk factors and to estimate spatial and temporal patterns of infection by chronic wasting disease (CWD) in white-tailed deer (Odocoileus virginianus) of Wisconsin, USA. We found significant heterogeneities for infection due to age, sex, and spatial location. Infection probability increased with age for all young deer, increased with age faster for young males, and then declined for some older animals, as expected from disease-associated mortality and age-related changes in infection risk. We found that disease prevalence was clustered in a central location, as expected under a simple spatial epidemic process where disease prevalence should increase with time and expand spatially. However, we could not detect any consistent temporal or spatiotemporal trends in CWD prevalence. Estimates of the temporal trend indicated that prevalence may have decreased or increased with nearly equal posterior probability, and the model without temporal or spatiotemporal effects was nearly equivalent to models with these effects based on deviance information criteria. For maximum interpretability of the role of location as a disease risk factor, we used the technique of direct standardization for prevalence mapping, which we develop and describe. These mapping results allow disease management actions to be employed with reference to the estimated spatial distribution of the disease and to those host classes most at risk. Future wildlife epidemiology studies should employ hierarchical Bayesian methods to smooth estimated quantities across space and time, account for heterogeneities, and then report disease rates based on an appropriate standardization. ?? 2009 by the Ecological Society of America.

Wisconsin

Testing alternative models of climate-mediated extirpations

Biotic responses to climate change will vary among taxa and across latitudes, elevational gradients, and degrees of insularity. However, due to factors such as phenotypic plasticity, ecotypic variation, and evolved tolerance to thermal stress, it remains poorly understood whether losses should be greatest in populations experiencing the greatest climatic change or living in places where the prevailing climate is closest to the edge of the species' bioclimatic envelope (e.g., at the hottest, driest sites). Research on American pikas (Ochotona princeps) in montane areas of the Great Basin during 1994-1999 suggested that 20th-century population extirpations were predicted by a combination of biogeographic, anthropogenic, and especially climatic factors. Surveys during 2005-2007 documented additional extirpations and within-site shifts of pika distributions at remaining sites. To evaluate the evidence in support of alternative hypotheses involving effects of thermal stress on pikas, we placed temperature sensors at 156 locations within pika habitats in the vicinity of 25 sites with historical records of pikas in the Basin. We related these time series of sensor data to data on ambient temperature from weather stations within the Historical Climate Network. We then used these highly correlated relationships, combined with long-term data from the same weather stations, to hindcast temperatures within pika habitats from 1945 through 2006. To explain patterns of loss, we posited three alternative classes of direct thermal stress: (1) acute cold stress (number of days below a threshold temperature); (2) acute heat stress (number of days above a threshold, temperature); and. (3) chronic heat stress (average summer temperature). Climate change was defined as change in our thermal metrics between two 31-y.r periods: 1945-1975 and 1976-2006. We found that patterns of persistence were well predicted by metrics of climate. Our best models suggest some effects of climate change; however, recent and long-term metrics of chronic heat stress and acute cold stress, neither previously recognized as sources of stress for pikas, were some of the best predictors of pika persistence. Results illustrate that extremely rapid distributional shifts can be explained by climatic influences and have implications for conservation topics such as reintroductions and early-warning indicators. ?? 2010 by the Ecological society of America.

Ecological Applications

When can efforts to control nuisance and invasive species backfire?

Population control through harvest has the potential to reduce the abundance of nuisance and invasive species. However, demographic structure and density-dependent processes can confound removal efforts and lead to undesirable consequences, such as overcompensation (an increase in abundance in response to harvest) and instability (population cycling or chaos). Recent empirical studies have demonstrated the potential for increased mortality (such as that caused by harvest) to lead to overcompensation and instability in plant, insect, and fish populations. We developed a general population model with juvenile and adult stages to help determine the conditions under which control harvest efforts can produce unintended outcomes. Analytical and simulation analyses of the model demonstrated that the potential for overcompensation as a result of harvest was significant for species with high fecundity, even when annual stage-specific survivorship values were fairly low. Population instability as a result of harvest occurred less frequently and was only possible with harvest strategies that targeted adults when both fecundity and adult survivorship were high. We considered these results in conjunction with current literature on nuisance and invasive species to propose general guidelines for assessing the risks associated with control harvest based on life history characteristics of target populations. Our results suggest that species with high per capita fecundity (over discrete breeding periods), short juvenile stages, and fairly constant survivorship rates are most likely to respond undesirably to harvest. It is difficult to determine the extent to which overcompensation and instability could occur during real-world removal efforts, and more empirical removal studies should be undertaken to evaluate population-level responses to control harvests. Nevertheless, our results identify key issues that have been seldom acknowledged and are potentially generic across taxa. ?? 2009 by the Ecological Society ot America.

Ecological Applications

Risk of Myxobolus cerebralis infection to rainbow trout in the Madison River, Montana, USA

Myxobolus cerebralis, the parasite that causes salmonid whirling disease, has had detrimental effects on several salmonid populations in the Intermountain West, including the rainbow trout in the Madison River, Montana, USA. The goal of this study was to examine relationships among characteristics of the environment, Tubifex tubifex (the alternate host) populations, and rainbow trout whirling disease risk in the Madison River. Environmental characteristics were measured in side channels of the Madison River, and differences were described with a principal components analysis. The density of T. tubifex, the prevalence of infection in T. tubifex, and the density of infected T. tubifex were determined for the side channels using benthic core samples and examination of live tubificids for infection. The site-specific contribution to whirling disease risk in the side channels was determined using in situ exposures of sentinel rainbow trout. Regression analyses were used to determine correlations among these characteristics. Side channels differed in site-specific contribution to rainbow trout whirling disease risk, which was positively correlated to the density of infected T. tubifex. Side channels with fine sediments and lower water temperatures made greater site-specific contribution to whirling disease risk and had higher densities of infected T. tubifex than side channels with coarser sediments and higher temperatures. The ability to characterize areas of high whirling disease risk is essential for improving our understanding of the dynamics of M. cerebralis such that appropriate management strategies can be implemented. In addition, this study provides a model of how the disease ecology of complex aquatic parasites can be examined when the influential processes operate on different spatial scales. ?? 2006 by the Ecological Society of America.

Montana

Carryover aquatic effects on survival of metamorphic frogs during pond emigration

In organisms with complex life cycles, physiological stressors during early life stages may have fitness-level impacts that are delayed into later stages or habitats. We tested the hypothesis that body size and date of metamorphosis, which are highly responsive to aquatic stressors, influence post-metamorphic survival and movement patterns in the terrestrial phase of an ephemeral pond-breeding frog by examining these traits in two populations of northern red-legged frogs (Rana aurora aurora). To increase variation of body size at metamorphosis, we manipulated food availability for 314 of 1045 uniquely marked tadpoles and estimated the probability that frogs survived and emigrated using concentric rings of drift fencing surrounding ponds and Bayesian capture-recapture modeling. The odds of surviving and emigrating from the ponds to the innermost drift fences, ???12 m, increased by factors of 2.20 (95% credibility intervals 1.39-4.23) and 2.54 (0.94-4.91) with each millimeter increase in snout-vent length and decreased by factors of 0.91 (0.85-0.96) and 0.89 (0.80-1.00) with each day's delay in metamorphosis for the two ponds. The odds of surviving and moving to the next ring of fencing, 12 m to ???40 m from the ponds, increased by a factor of 1.20 (0.45-4.06) with each millimeter increase in size. Our results demonstrated that body size and timing of metamorphosis relate strongly to the performance of newly metamorphosed frogs during their initial transition into terrestrial habitat. Carryover effects of aquatic stressors that reduce size and delay metamorphosis may have population-level impacts that are not expressed until terrestrial stages. Since changes in both aquatic and terrestrial systems are implicated in many amphibian declines, quantifying both immediate and delayed effects of stressors on demographic rates is critical to sound management. ?? 2006 by the Ecological Society of America.

Ecological Applications

Body size and predatory performance in wolves: Is bigger better?

Large body size hinders locomotor performance in ways that may lead to trade-offs in predator foraging ability that limit the net predatory benefit of larger size. For example, size-related improvements in handling prey may come at the expense of pursuing prey and thus negate any enhancement in overall predatory performance due to increasing size. 2. This hypothesis was tested with longitudinal data from repeated observations of 94 individually known wolves (Canis lupus) hunting elk (Cervus elaphus) in Yellowstone National Park, USA. Wolf size was estimated from an individually based sex-specific growth model derived from body mass measurements of 304 wolves. 3. Larger size granted individual wolves a net predatory advantage despite substantial variation in its effect on the performance of different predatory tasks; larger size improved performance of a strength-related task (grappling and subduing elk) but failed to improve performance of a locomotor-related task (selecting an elk from a group) for wolves > 39 kg. 4. Sexual dimorphism in wolf size also explained why males outperformed females in each of the three tasks considered (attacking, selecting, and killing). 5. These findings support the generalization that bigger predators are overall better hunters, but they also indicate that increasing size ultimately limits elements of predatory behaviour that require superior locomotor performance. We argue that this could potentially narrow the dietary niche of larger carnivores as well as limit the evolution of larger size if prey are substantially more difficult to pursue than to handle. ?? 2009 British Ecological Society.

Journal of Animal Ecology

Hierarchical spatial models for predicting pygmy rabbit distribution and relative abundance

Conservationists routinely use species distribution models to plan conservation, restoration and development actions, while ecologists use them to infer process from pattern. These models tend to work well for common or easily observable species, but are of limited utility for rare and cryptic species. This may be because honest accounting of known observation bias and spatial autocorrelation are rarely included, thereby limiting statistical inference of resulting distribution maps. We specified and implemented a spatially explicit Bayesian hierarchical model for a cryptic mammal species (pygmy rabbit Brachylagus idahoensis). Our approach used two levels of indirect sign that are naturally hierarchical (burrows and faecal pellets) to build a model that allows for inference on regression coefficients as well as spatially explicit model parameters. We also produced maps of rabbit distribution (occupied burrows) and relative abundance (number of burrows expected to be occupied by pygmy rabbits). The model demonstrated statistically rigorous spatial prediction by including spatial autocorrelation and measurement uncertainty. We demonstrated flexibility of our modelling framework by depicting probabilistic distribution predictions using different assumptions of pygmy rabbit habitat requirements. Spatial representations of the variance of posterior predictive distributions were obtained to evaluate heterogeneity in model fit across the spatial domain. Leave-one-out cross-validation was conducted to evaluate the overall model fit. Synthesis and applications. Our method draws on the strengths of previous work, thereby bridging and extending two active areas of ecological research: species distribution models and multi-state occupancy modelling. Our framework can be extended to encompass both larger extents and other species for which direct estimation of abundance is difficult. ?? 2010 The Authors. Journal compilation ?? 2010 British Ecological Society.

Journal of Applied Ecology

Estimation of temporary emigration in male toads

Male boreal toads (Bufo boreas) are thought to return to the breeding site every year but, if absent in a particular year, will be more likely to return the following year. Using Pollock's robust design we estimated temporary emigration (the probability a male toad is absent from a breeding site in a given year) at three locations in Colorado, USA: two in Rocky Mountain National Park and one in Chaffee County. We present data that suggest that not all male toads return to the breeding site every year. Our analyses indicate that temporary emigration varies by site and time (for example, from 1992 to 1998, the probability of temporary emigration ranged from 10% to 29% and from 3% to 95% at Lost Lake and Kettle Tarn, respectively). Although the results provide weak evidence that males are more likely to return after a year's hiatus, a general pattern of state-dependent temporary emigration was not supported. We also hypothesized relationships between temporary emigration and a number of weather variables. While some competitive models included weather covariates, imprecise and variable estimates of the effects of these covariates precluded fully defining their impact on temporary emigration. ?? 2006 by the Ecological Society of America.

Colorado

Chinook salmon use of spawning patches: Relative roles of habitat quality, size, and connectivity

Declines in many native fish populations have led to reassessments of management goals and shifted priorities from consumptive uses to species preservation. As management has shifted, relevant environmental characteristics have evolved from traditional metrics that described local habitat quality to characterizations of habitat size and connectivity. Despite the implications this shift has for how habitats may be prioritized for conservation, it has been rare to assess the relative importance of these habitat components. We used an information-theoretic approach to select the best models from sets of logistic regressions that linked habitat quality, size, and connectivity to the occurrence of chinook salmon (Oncorhynchus tshawytscha) nests. Spawning distributions were censused annually from 1995 to 2004, and data were complemented with field measurements that described habitat quality in 43 suitable spawning patches across a stream network that drained 1150 km 2 in central Idaho. Results indicated that the most plausible models were dominated by measures of habitat size and connectivity, whereas habitat quality was of minor importance. Connectivity was the strongest predictor of nest occurrence, but connectivity interacted with habitat size, which became relatively more important when populations were reduced. Comparison of observed nest distributions to null model predictions confirmed that the habitat size association was driven by a biological mechanism when populations were small, but this association may have been an area-related sampling artifact at higher abundances. The implications for habitat management are that the size and connectivity of existing habitat networks should be maintained whenever possible. In situations where habitat restoration is occurring, expansion of existing areas or creation of new habitats in key areas that increase connectivity may be beneficial. Information about habitat size and connectivity also could be used to strategically prioritize areas for improvement of local habitat quality, with areas not meeting minimum thresholds being deemed inappropriate for pursuit of restoration activities. ?? 2007 by the Ecological Society of America.

Ecological Applications

Stream ecosystem response to limestone treatment in acid impacted watersheds of the allegheny plateau

Restoration programs are expanding worldwide, but assessments of restoration effectiveness are rare. The objectives of our study were to assess current acid-precipitation remediation programs in streams of the Allegheny Plateau ecoregion of West Virginia (USA), identify specific attributes that could and could not be fully restored, and quantify temporal trends in ecosystem recovery. We sampled water chemistry, physical habitat, periphyton biomass, and benthic macroinvertebrate and fish community structure in three stream types: acidic (four streams), naturally circumneutral (eight streams), and acidic streams treated with limestone sand (eight streams). We observed no temporal trends in ecosystem recovery in treated streams despite sampling streams that ranged from 2 to 20 years since initial treatment. Our results indicated that the application of limestone sand to acidic streams was effective in fully recovering some characteristics, such as pH, alkalinity, Ca2+, Ca:H ratios, trout biomass and density, and trout reproductive success. However, recovery of many other characteristics was strongly dependent upon spatial proximity to treatment, and still others were never fully recovered. For example, limestone treatment did not restore dissolved aluminum concentrations, macroinvertebrate taxon richness, and total fish biomass to circumneutral reference conditions. Full recovery may not be occurring because treated streams continue to drain acidic watersheds and remain isolated in a network of acidic streams. We propose a revised stream restoration plan for the Allegheny Plateau that includes restoring stream ecosystems as connected networks rather than isolated reaches and recognizes that full recovery of acidified watersheds may not be possible. ?? 2007 by the Ecological Society of America.

Ecological Applications

Population inertia and its sensitivity to changes in vital rates and population structure

Because the (st)age structure of a population may rarely be stable, studies of transient population dynamics and population momentum are becoming ever more popular. Yet, studies of "population momentum" are restricted in the sense that they describe the inertia of population size resulting from a demographic transition to the stationary population growth rate. Although rarely mentioned, inertia in population size is a general phenomenon and can be produced by any demographic transition or perturbation. Because population size is of central importance in demography, conservation, and management, formulas relating the sensitivity of population inertia to changes in underlying vital rates and population structure could provide much-needed insight into the dynamics of populations with unstable (st)age structure. Here, we derive such formulas, which are readily computable, and provide examples of their potential use in studies of life history and applied arenas of population study. ?? 2007 by the Ecological Society of America.

Ecology