Taxonomic Considerations in Listing Subspecies Under the U.S. Endangered Species Act

Abstract:  The U.S. Endangered Species Act (ESA) allows listing of subspecies and other groupings below the rank of species. This provides the U.S. Fish and Wildlife Service and the National Marine Fisheries Service with a means to target the most critical unit in need of conservation. Although roughly one-quarter of listed taxa are subspecies, these management agencies are hindered by uncertainties about taxonomic standards during listing or delisting activities. In a review of taxonomic publications and societies, we found few subspecies lists and none that stated standardized criteria for determining subspecific taxa. Lack of criteria is attributed to a centuries-old debate over species and subspecies concepts. Nevertheless, the critical need to resolve this debate for ESA listings led us to propose that minimal biological criteria to define disjunct subspecies (legally or taxonomically) should include the discreteness and significance criteria of distinct population segments (as defined under the ESA). Our subspecies criteria are in stark contrast to that proposed by supporters of the phylogenetic species concept and provide a clear distinction between species and subspecies. Efforts to eliminate or reduce ambiguity associated with subspecies-level classifications will assist with ESA listing decisions. Thus, we urge professional taxonomic societies to publish and periodically update peer-reviewed species and subspecies lists. This effort must be paralleled throughout the world for efficient taxonomic conservation to take place.     

Reliability of Absolute and Relative Predictions of Population Persistence Based on Time Series

Abstract:  Conventional population viability analysis (PVA) is often impractical because data are scarce for many threatened species. For this reason, simple count-based models are being advocated. The simplest of these models requires nothing more than a time series of abundance estimates, from which variance and autocorrelation in growth rate are estimated and predictions of population persistence are generated. What remains unclear, however, is how many years of data are needed to generate reliable estimates of these parameters and hence reliable predictions of persistence. By analyzing published and simulated time series, we show that several decades of data are needed. Predictions based on short time series were very unreliable mainly because limited data yielded biased, unreliable estimates of variance in growth rate, especially when growth rate was strongly autocorrelated. More optimistically, our results suggest that count-based PVA is sometimes useful for relative risk assessment (i.e., for ranking populations by extinction risk), even when time series are only a decade long. However, some conditions consistently lead to backward rankings. We explored the limited conditions under which simple count-based PVA may be useful for relative risk assessment.     

Impact of Grazing Intensity during Drought in an Arizona Grassland

Abstract:  The ecological benefits of changing cattle grazing practices in the western United States remain controversial, due in part to a lack of experimentation. In 1997 we initiated an experimental study of two rangeland alternatives, cattle removal and high-impact grazing, and compared grassland community responses with those with more conventional, moderate grazing practices. The study was conducted in a high-elevation, semiarid grassland near Flagstaff, Arizona (U.S.A.). We conducted annual plant surveys of modified Whittaker plots for 8 years and examined plant composition shifts among treatments and years. High-impact grazing had strong directional effects that led to a decline in perennial forb cover and an increase in annual plants, particularly the exotic cheatgrass ( Bromus tectorum L.). A twofold increase in plant cover by exotic species followed a severe drought in the sixth year of the study, and this increase was greatest in the high-impact grazing plots, where native cover declined by one-half. Cattle removal resulted in little increase in native plant cover and reduced plant species richness relative to the moderate grazing control. Our results suggest that some intermediate level of cattle grazing may maintain greater levels of native plant diversity than the alternatives of cattle removal or high-density, short-duration grazing practices. Furthermore, episodic drought interacts with cattle grazing, leading to infrequent, but biologically important shifts in plant communities. Our results demonstrate the importance of climatic variation in determining ecological effects of grazing practices, and we recommend improving conservation efforts in arid rangelands by developing management plans that anticipate this variation.     

Long-Term Landscape Change and Bird Abundance in Amazonian Rainforest Fragments

Abstract:  The rainforests of the Amazon basin are being cut by humans at a rate >20,000 km²/year, leading to smaller and more isolated patches of forest, with remaining fragments often in the range of 1–100 ha. We analyzed samples of understory birds collected over 20 years from a standardized mist-netting program in 1– to 100-ha rainforest fragments in a dynamic Amazonian landscape near Manaus, Brazil. Across bird guilds, the condition of second growth immediately surrounding fragments was often as important as fragment size or local forest cover in explaining variation in abundance. Some fragments surrounded by 100 m of open pasture showed reductions in insectivorous bird abundance of over 95%, even in landscapes dominated by continuous forest and old second growth. These extreme reductions may be typical throughout Amazonia in small (≤10 ha), isolated fragments of rainforest. Abundance for some guilds returned to preisolation levels in 10- and 100-ha fragments connected to continuous forest by 20-year-old second growth. Our results show that the consequences of Amazonian forest loss cannot be accurately described without explicit consideration of vegetation dynamics in matrix habitat. Any dichotomous classification of the landscape into "forest" and "nonforest" misses essential information about the matrix.     

Effectiveness of a Regional Corridor in Connecting Two Florida Black Bear Populations

Abstract:  Corridors may mitigate the adverse effects of habitat fragmentation by restoring or maintaining connectivity between disjunct populations. The efficacy of corridors for large carnivores, however, has rarely been evaluated objectively. We used noninvasive sampling, microsatellite analysis, and population assignment tests to evaluate the effectiveness of a regional corridor in connecting two Florida black bear ( Ursus americanus floridanus ) populations (Osceola and Ocala). Bear movement was predominantly unidirectional, with a limited mixing of individuals from the two populations in one area of the corridor. We also documented bears in Osceola that were genetically assigned to Ocala and bears in Osceola that may be offspring from an Osceola-Ocala mating. Our results indicate that the Osceola-Ocala corridor is functional and provides a conduit for gene flow between these populations. Human development, however, may hinder the use of the Osceola-Ocala corridor by bears. The noninvasive sampling and genetic methods we used provide a means of evaluating corridor effectiveness that can help identify linkages necessary for maintaining metapopulation structure and population viability.     

Avian Community Response to Lowland Tropical Rainforest Isolation: 40 Years of Change at La Selva Biological Station, Costa Rica

Abstract:  Since 1960, most of the forest surrounding the La Selva Biological Station, an intensively studied tropical research facility in Costa Rica, has been converted to agricultural uses. We used quantitative censuses and analysis of previously published categorical abundances to assess changes in the bird community, and we evaluated potential causes of species-specific changes by assessing their association with habitat, diet, participation in mixed-species flocks, and nest type. Approximately the same percentage of species increased as decreased in abundance from 1960 to 1999 (10–20% of all species, depending on method of assessment). Diet was the single most important trait associated with declining species. At least 50% of the species that declined have insectivorous diets. Use of forest habitat and participation in mixed-species flocks were also significant factors associated with declines, but nest type was unrelated to change in abundance. The species that increased in abundance tended to occur in open habitats and have omnivorous diets. These results reinforce the importance of several population risk factors associated with tropical understory insectivory and mixed-species flocking: patchy spatial distribution, low population density, large home range, and dietary specialization. La Selva's protected area (1611 ha), despite a forested connection on one boundary with a higher elevation national park, is apparently too small to maintain at least one major guild (understory insectivores). This first quantitative assessment of bird community change at La Selva highlights the need to intensify study of the mechanisms and consequences of biological diversity change in tropical forest fragments.     

Assessing Habitat Quality for a Migratory Songbird Wintering in Natural and Agricultural Habitats

Abstract:  As tropical forests are cleared, a greater proportion of migratory songbirds are forced to winter in agricultural and disturbed habitats, which, if poorer in quality than natural forests, could contribute to population declines. We compared demographic indicators of habitat quality for a focal species, the American Redstart ( Setophaga ruticilla ), wintering in Jamaican citrus orchards and shade coffee plantations with those in four natural habitats: mangrove, coastal scrub, coastal palm, and dry limestone forests. Demographic measures of habitat quality included density, age and sex ratio, apparent survival, and changes in body mass. Measures of habitat quality for redstarts in citrus and coffee habitats were generally intermediate between the highest (mangrove) and lowest (dry limestone) measurements from natural habitats. The decline in mean body mass over the winter period was a strong predictor of annual survival rate among habitats, and we suggest that measures of body condition coupled with survival data provide the best measures of habitat quality for nonbreeding songbirds. Density, which is far easier to estimate, was correlated with these more labor-intensive measures, particularly in the late winter when food is likely most limiting. Thus, local density may be useful as an approximation of habitat quality for wintering migrant warblers. Our findings bolster those of previous studies based on bird abundance that suggest arboreal agricultural habitats in the tropics can be useful for the conservation of generalist, insectivorous birds, including many migratory passerines such as redstarts.     

Genetic Identification of Spotted Owls, Barred Owls, and Their Hybrids: Legal Implications of Hybrid Identity

Abstract:  Recent population expansion of Barred Owls (  Strix varia ) into western North America has led to concern that they may compete with and further harm the Northern Spotted Owl (  S. occidentalis caurina ), which is already listed as threatened under the U.S. Endangered Species Act (ESA). Because they hybridize, there is a legal need under the ESA for forensic identification of both species and their hybrids. We used mitochondrial control-region DNA and amplified fragment-length polymorphism (AFLP) analyses to assess maternal and biparental gene flow in this hybridization process. Mitochondrial DNA sequences (524 base pairs) indicated large divergence between Barred and Spotted Owls (13.9%). Further, the species formed two distinct clades with no signs of previous introgression. Fourteen diagnostic AFLP bands also indicated extensive divergence between the species, including markers differentiating them. Principal coordinate analyses and assignment tests clearly supported this differentiation. We found that hybrids had unique genetic combinations, including AFLP markers from both parental species, and identified known hybrids as well as potential hybrids with unclear taxonomic status. Our analyses corroborated the findings of extensive field studies that most hybrids genetically sampled resulted from crosses between female Barred Owls and male Spotted Owls. These genetic markers make it possible to clearly identify these species as well as hybrids and can now be used for research, conservation, and law enforcement. Several legal avenues may facilitate future conservation of Spotted Owls and other ESA-listed species that hybridize, including the ESA similarity-of-appearance clause (section 4[e]) and the Migratory Bird Treaty Act. The Migratory Bird Treaty Act appears to be the most useful route at this time.     

Flying Foxes as Strong Interactors in South Pacific Island Ecosystems: A Conservation Hypothesis

Abstract: The dependency of highly endemic island floras on few potential pollinators in depauperate island faunas suggests that pollinators and seed dispersers may be crucial in the preservation of biodiversity in isolated oceanic islands. We discuss the hypothesis that flying foxes are "strong interactors" in South Pacific islands where they serve as the principal pollinators and seed dispersers, This suggests that the ongoing decline and ultimate extinction of flying fox species on Pacific islands may lead to a cascade of linked plant extinctions. We propose an empirical test of this hypothesis: comparisons of plant reproductive success in Guam, which has virtually lost its flying fox populations, and Samoa, where significant populations remain.