Error propagation in spatially explicit population models: A reassessment
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Management of amphibian populations to reverse recent declines will require defining high-quality habitat for individual species or groups of species, followed by efforts to retain or restore these habitats on the landscape. We examined landscape-level habitat relationships for frogs and toads by measuring associations between relative abundance and species richness based on survey data derived from anuran calls and features of land-cover maps for Iowa and Wisconsin. The most consistent result across all anuran guilds was a negative association with the presence of urban land. Upland and wetland forests and emergent wetlands tended to be positively associated with anurans. Landscape metrics that represent edges and patch diversity also had generally positive associations, indicating that anurans benefit from a complex of habitats that include wetlands. In Iowa the most significant associations with relative abundance were the length of the edge between wetland and forest ( positive) and the presence of urban land (negative). In Wisconsin the two most significant associations with relative abundance were forest area and agricultural area ( both positive). Anurans had positive associations with agriculture in Wisconsin but not in Iowa. Remnant forest patches in agricultural landscapes may be providing refuges for some anuran species. Differences in anuran associations with deep water and permanent wetlands between the two states suggest opportunities for management action. Large-scale maps can contribute to predictive models of amphibian habitat use, but water quality and vegetation information collected from individual wetlands will likely be needed to strengthen those predictions. Landscape habitat analyses provide a framework for future experimental and intensive research on specific factors affecting the health of anurans.
Metapopulation theory has proven useful for understanding the population structure and dynamics of many species of conservation concern. The metapopulation concept has been applied almost exclusively to nonmigratory species, however, for which subpopulation demographic independence—a requirement for a classically defined metapopulation - is explicitly related to geographic distribution and dispersal probabilities. Defining the degree of demographic independence among subpopulations of migratory animals, and thus the applicability of metapopulation theory as a conceptual framework for understanding population dynamics, is much more difficult. Unlike nonmigratory species, subpopulations of migratory animals cannot be defined as synonymous with geographic areas. Groups of migratory birds that are geographically separate at one part of the annual cycle may occur together at others, but co-occurrence in time and space does not preclude the demographic independence of subpopulations. I suggest that metapopulation theory can be applied to migratory species but that understanding the degree of subpopulation independence may require information about both spatial distribution throughout the annual cycle and behavioral mechanisms that may lead to subpopulation demographic independence. The key for applying metapopulation theory to migratory animals lies in identifying demographically independent subpopulations, even as they move during the annual cycle and potentially co-occur with other subpopulations. Using examples of migratory bird species, I demonstrate that spatial and temporal modes of subpopulation independence can interact with behavioral mechanisms to create demographically independent subpopulations, including cases in which subpopulations are not spatially distinct in some parts of the annual cycle.
The Florida torreya ( Torreya taxifolia ) is a coniferous tree endemic to a 35-km stretch of bluffs and ravines along the east side of the Apalachicola River in northern Florida and southern Georgia. This formerly locally abundant tree has declined as a result of disease during the 1950s and is on the U.S. endangered species list. With no seed production in the wild, this species is headed toward extinction. We conducted a survey on roughly 200 trees from 1988 to 1996 and used these data to estimate the likelihood of population persistence during the next several decades. We compared a stage-class transition matrix model ( RAMAS) and an individual-based stochastic model ( TORSIM) of growth and mortality to project future populations. Given the current lack of seed production in the wild, all models predict extinction. The question of concern is the imminence of this predicted extinction. Differing predicted times to extinction would suggest different immediate management recommendations. Both models predicted an over 90% likelihood of persistence during the next 50 years. Predictions differed in that the transition matrix model was less optimistic than the individual-based model regarding persistence. Model sensitivity analysis showed that the results were robust to significant decreases in growth and sprouting probabilities. Submodels identified different persistence likelihoods in different populations. Nonetheless, unless management of the population can facilitate maturation and seed production, extinction of this species in the wild is inevitable.
Detrimental effects of introduced fishes on native amphibian populations have prompted removal of introduced cutthroat (Oncorhynchus clarki), rainbow (Oncorhynchus mykiss), and brook trout (Salvelinus fontinalis) from naturally fishless lakes at Mt. Rainier National Park, Washington (U.S.A.). Using paleolimnological indicators (diatoms, invertebrates, and sediment characteristics) in eight 480-year-old sediment cores from eight lakes, we (1) derived estimates of baseline environmental conditions and natural variation, (2) assessed the effects of stocking naturally fishless lakes, and (3) determined whether lakes returned to predisturbance conditions after fish removal (restoration). Diatom floras were relatively stable between 315 and 90 years before present in all lakes; we used this time period to define lake-specific "baseline" conditions. Dissimilarity analyses of diatoms revealed sustained, dramatic changes in diatom floras that occurred approximately 80 years ago (when fish were introduced) in four of five stocked lakes, whereas the diatom floras in two unstocked lakes had not changed significantly in the last 315 years. Diatoms were not preserved in an eighth lake. State changes also occurred in two lakes over 200 years before European settlement of the Pacific Northwest. Preserved invertebrate densities fluctuated dramatically over time in all cores, providing a poor reference for assessing the effects of fishes. Nevertheless, fish-invertebrate interactions have been demonstrated in other paleolimnological studies and may be useful for lower-elevation or more productive lakes. Because diatom communities have not returned to predisturbance assemblages in restored lakes, even 20-30 years after fish removal, we conclude that Mt. Rainier lakes were not successfully restored by the removal of fishes.
There is a significant disparity in the protection of terrestrial and marine environments in the United States. Despite the considerable literature dedicated to the subject of protected areas, both terrestrial and marine, in the United States, we are not aware of work explicitly describing this dichotomy. We compared marine and terrestrial areas under federal jurisdiction to provide a quantitative assessment of the differences between the conservation of land and sea in the United States. Specifically, we compared national marine sanctuaries (including sanctuary preservation areas and ecological reserves) with national parks, national forests, and national wildlife refuges (including national wilderness preservation areas). Our results suggest that marine sanctuaries are fewer in number, smaller in total area, and smaller in percentage of area covered than are terrestrial protected areas.
Historical variability in fire regime is a conservative indicator of ecosystem sustainability, and thus understanding the natural role of fire in chaparral ecosystems is necessary for proper fire management. It has been suggested that the “natural” fire regime was one of frequent small fires that fragmented the landscape into a fine-grained mixture of age classes that precluded large, catastrophic fires. Some researchers claim that this regime was lost because of highly effective fire suppression and conclude that if fire managers could “restore” a regime of frequent fires with widespread prescription burning, they could eliminate the hazard of catastrophic fires. The primary evidence in support of this model is a study that compared contemporary burning patterns in southern California, U.S.A., a region subject to fire suppression, with patterns in northern Baja California, Mexico, where there is less effective fire suppression. We found that differences in fire regime between these two regions are inconclusive and could not be ascribed conclusively to differences in fire suppression. Historical records suggest that the natural fire regime in southern California shrublands was rather coarse-grained and not substantively different from the contemporary regime. There is no evidence that fire-management policies have created the contemporary fire regime dominated by massive Santa Ana wind-driven fires. Increased expenditures on fire suppression and increased loss of property and lives are the result of human demographic patterns that place increasing demand on fire-suppression forces.
Terrestrial salamanders of the family Plethodontidae have unique attributes that make them excellent indicators of biodiversity and ecosystem integrity in forested habitats. Their longevity, small territory size, site fidelity, sensitivity to natural and anthropogenic perturbations, tendency to occur in high densities, and low sampling costs mean that counts of plethodontid salamanders provide numerous advantages over counts of other North American forest organisms for indicating environmental change. Furthermore, they are tightly linked physiologically to microclimatic and successional processes that influence the distribution and abundance of numerous other hydrophilic but difficult-to-study forest-dwelling plants and animals. Ecosystem processes such as moisture cycling, food-web dynamics, and succession, with their related structural and microclimatic variability, all affect forest biodiversity and have been shown to affect salamander populations as well. We determined the variability associated with sampling for plethodontid salamanders by estimating the coefficient of variation (CV ) from available time-series data. The median coefficient of variation indicated that variation in counts of individuals among studies was much lower in plethodontids (27%) than in lepidoptera (93%), passerine birds (57%), small mammals (69%), or other amphibians (37–46%), which means plethodontid salamanders provide an important statistical advantage over other species for monitoring long-term forest health.
We examined nesting success and levels of nest predation and cowbird parasitism among five different silvicultural treatments: regenerating (3-6 years old), mid-rotation (12-15 years old), and thinned (17-23 years old) pine plantations, single-tree selection, and late-rotation pine-hardwood stands in the Ouachita Mountains of Arkansas from 1993 to 1995. We monitored 1674 nests. Differences in daily mortality and daily predation rate among two or more treatments were found for 4 and 3 of 12 species, respectively. These differences were lost following Bonferroni adjustments, but thinned stands had higher levels of predation than single-tree selection stands when predation levels were averaged across species. Daily predation rates were positively correlated with the relative abundance of birds, suggesting that nest predators respond to prey availability (i.e., nests) in a density-dependent manner. The relative abundance of cowbirds differed among treatments, with the highest densities in regenerating, thinned, and single-tree selection stands. Field Sparrows (Spizella pusilla) and Yellow-breasted Chats (Icteria virens) experienced higher levels of parasitism in thinned than regenerating plantations, whereas White-eyed Vireos (Vireo griseus) experienced higher parasitism in regenerating plantations than in mid-rotation or thinned plantations. Several shrub-nesting and 1 ground-nesting species had lower nesting success in thinned and regenerating plantations than has been reported in previously published studies. Thus, some seral stages of even-aged management may provide low-quality nesting habitat for several early-successional bird species. In contrast, many species nesting in mid-rotation and single-tree selection stands had nesting success similar to or greater than that found in previous studies, suggesting that some silvicultural treatments, when embedded in a largely forested landscape, may provide suitable habitat for forest land birds without affecting their reproductive success.
Some ecosystems, such as yellow pine forests, have had a long history of frequent surface fires, but because of fire suppression policy, fires have been largely excluded from them during the last century (Covington 2000). Unnatural fuel accumulation in these forests has increased the potential for large, catastrophic crown fires, and re-introduction of prescribed fire is one remedy for this critical fire hazard. But fire ecologists and fire managers need to be cautious in transferring this model to all western ecosystems (Anderson et al. 1999; Gutsell et al. 2001). Although large, catastrophic crown fires are apparently unnatural in yellow pine forests (but cf. Shinneman & Baker 1997 ), this is not so in other western forests and shrub-lands, and widespread prescription burning is not warranted everywhere.
Genetic analysis of introduced populations, especially in morphologically conservative taxa, can clarify introduction histories, identify management units and source populations, provide a more realistic estimate of the frequency of successful invasion, and suggest strategies for preventing further introductions. In the last 7 years, populations of swamp eels, referred to the Asian genus Monopterus (Family Synbranchidae) on the basis of external morphology, have been discovered in aquatic habitats near Atlanta, Georgia; Tampa, Florida; North Miami, Florida; and most recently in close proximity to Everglades National Park in Homestead, Florida. Swamp eels are large predators capable of dispersal over land and have the potential to disrupt already threatened ecosystems. We analyzed mitochondrial DNA sequences from four known populations in the continental United States and samples from Malaysia, Indonesia, Vietnam, and two locations in China to determine introduction histories, source populations, genetic diversity, and relationships among populations. Our results indicate that there have been at least three independent introductions of genetically distinct forms. Introduced populations in close proximity (separated by <40 km) are genetically distinct. The level of sequence difference among introduced populations reaches levels seen among sister families of teleost fishes for the same region of the mitochondrial genome. These genetically distinct introduced populations in all likelihood represent at least two and possibly three species. Regardless of species status, these genetically distinct lineages may be expected to vary in ecological or life-history traits, representing different potential threats to the ecosystems where they have been introduced. Given the success of swamp eels in invading many habitats around the world, further study of these eels is warranted to elucidate the characteristics of successful invaders and invasions.
Development of effective conservation plans for terrestrial animals will require some assessment of which human-modified and natural habitats can support populations of priority species. We examined bird communities associated with 11 natural and human-modified habitats in Panama and assessed the importance of those habitats for species of different vulnerability to disturbance. We calculated habitat importance scores using both relative habitat preferences and vulnerability scores for all species present. Species of moderate and high vulnerability were primarily those categorized as forest specialists or forest generalists. As expected, even species-rich nonforest habitats provided little conservation value for the most vulnerable species. However, shaded coffee plantations and gallery forest corridors were modified habitats with relatively high conservation value. Sugar cane fields and Caribbean pine plantations offered virtually no conservation value for birds. Our method of assessing the conservation importance of different habitats is useful because it considers the types of species present and the potential role of the habitat in the conservation of those species (i.e., habitat preference). This method of habitat evaluation could be tailored to other conservation contexts with any measure of species vulnerability desired.
Roads are believed to be a major contributing factor to the ongoing spread of exotic plants. We examined the effect of road improvement and environmental variables on exotic and native plant diversity in roadside verges and adjacent semiarid grassland, shrubland, and woodland communities of southern Utah (U.S.A.). We measured the cover of exotic and native species in roadside verges and both the richness and cover of exotic and native species in adjacent interior communities (50 m beyond the edge of the road cut) along 42 roads stratified by level of road improvement (paved, improved surface, graded, and four-wheel-drive track). In roadside verges along paved roads, the cover of Bromus tectorum was three times as great (27%) as in verges along four-wheel-drive tracks (9%). The cover of five common exotic forb species tended to be lower in verges along four-wheel-drive tracks than in verges along more improved roads. The richness and cover of exotic species were both more than 50% greater, and the richness of native species was 30% lower, at interior sites adjacent to paved roads than at those adjacent to four-wheel-drive tracks. In addition, environmental variables relating to dominant vegetation, disturbance, and topography were significantly correlated with exotic and native species richness and cover. Improved roads can act as conduits for the invasion of adjacent ecosystems by converting natural habitats to those highly vulnerable to invasion. However, variation in dominant vegetation, soil moisture, nutrient levels, soil depth, disturbance, and topography may render interior communities differentially susceptible to invasions originating from roadside verges. Plant communities that are both physically invasible (e.g., characterized by deep or fertile soils) and disturbed appear most vulnerable. Decision-makers considering whether to build, improve, and maintain roads should take into account the potential spread of exotic plants.
Predation by brown tree snakes ( Boiga irregularis ) devastated the avifauna of Guam in the Mariana Islands during the last half of the twentieth century, causing the extirpation or serious reduction of most of the island's 25 resident bird species. Past studies have provided qualitative descriptions of the decline of native forest birds but have not considered all species or presented quantitative analyses. We analyzed two sets of survey data gathered in northern Guam between 1976 and 1998 and reviewed unpublished sources to provide a comprehensive account of the impact of brown tree snakes on the island's birds. Our results indicate that 22 species, including 17 of 18 native species, were severely affected by snakes. Twelve species were likely extirpated as breeding residents on the main island, 8 others experienced declines of ≥90% throughout the island or at least in the north, and 2 were kept at reduced population levels during all or much of the study. Declines of ≥90% occurred rapidly, averaging just 8.9 years along three roadside survey routes combined and 1.6 years at a 100-ha forested study site. Declines in northern Guam were also relatively synchronous and occurred from about 1976 to 1986 for most species. The most important factor predisposing a species to coexistence with brown tree snakes was its ability to nest and roost at locations where snakes were uncommon. Large clutch size and large body size were also related to longer persistence times, although large body size appeared to delay, but not prevent, extirpation. Our results draw attention to the enormous detrimental impact that brown tree snakes are likely to have upon invading new areas. Increased containment efforts on Guam are needed to prevent further colonizations, but a variety of additional management efforts would also benefit the island's remaining bird populations.
Effective conservation efforts for at-risk species require knowledge of the locations of existing populations. Species presence can be estimated directly by conducting field-sampling surveys or alternatively by developing predictive models . Direct surveys can be expensive and inefficient, particularly for rare and difficult-to-sample species, and models of species presence may produce biased predictions. We present a Bayesian approach that combines sampling and model -based inferences for estimating species presence. The accuracy and cost-effectiveness of this approach were compared to those of sampling surveys and predictive models for estimating the presence of the threatened bull trout (Salvelinus confluentus) via simulation with existing models and empirical sampling data. Simulations indicated that a sampling-only approach would be the most effective and would result in the lowest presence and absence misclassification error rates for three thresholds of detection probability. When sampling effort was considered, however, the combined approach resulted in the lowest error rates per unit of sampling effort. Hence, lower probability-of-detection thresholds can be specified with the combined approach, resulting in lower misclassification error rates and improved cost-effectiveness.
The Sea Turtle Stranding and Salvage Network has been monitoring turtle strandings for more than 20 years in the United States. High numbers of strandings in the early to mid-1980s prompted regulations to require turtle excluder devices (TEDs) on shrimping vessels (trawlers). Following year-round TED implementation in 1991, however, stranding levels in the Gulf of Mexico increased. We evaluated the efficacy of TEDs and other management actions (e.g., fisheries closures) on loggerhead (Caretta caretta) and Kemp's ridley (Lepidochelys kempii) turtle populations by analyzing a long-term, stranding data set from the western Gulf of Mexico. Our analyses suggest that both sea turtle population growth and shrimping activity have contributed to the observed increase in strandings. Compliance with regulations requiring turtle excluder devices was a significant factor in accounting for annual stranding variability: low compliance was correlated with high levels of strandings. Our projections suggest that improved compliance with TED regulations will reduce strandings to levels that, in conjunction with other protective measures, should promote population recoveries for loggerhead and Kemp's ridley turtles. Local, seasonal fisheries closures, concurrent with TED enforcement, could reduce strandings to even lower levels. A seasonal closure adjacent to a recently established Kemp's ridley nesting beach may also reduce mortality of nesting adults and thus promote long-term population persistence by fostering the establishment of a robust secondary nesting site.
The breeding phenologies of ectotherms are inextricably linked to weather, and amphibians in some temperate locations may have been breeding earlier in recent years in response to warmer spring temperatures (Beebee 1995: Forchhammer et al. 1998; Gibbs & Breisch 2001). Directional change in the timing of breeding resulting from climate change may have consequences for the fitness of individuals and may affect the persistence of amphibian populations (Ovaska 1997: Donnelly & Crump 1998). Blaustein et al. (2001) contribute valuable information to the small, but growing, data set of long-term observations of amphibian breeding phenology. As in other studies, Blaustein et al. found a significant relationship between air temperature and phenology, with earlier breeding associated with warmer air temperatures for boreal toads (Bufo boreas) and Cascades frogs (Rana cascadae) in Oregon and for spring peepers (Pseudacris crucifer) in Michigan. Contrary to other studies, however, there was no trend toward earlier breeding relative to year for any of these species or for Fowler's toads (B. fouleri) in Ontario. These results are important in demonstrating that changes in breeding phenology due to climate change are not universal among amphibians.