Long-term effects of deer browsing and trampling on the vegetation of peatlands

Overabundance of wild ungulates, especially exotic species, is a major threat to several ecosystems worldwide. While the response of forest vegetation to high density of herbivores has been well studied, far less is known about peatland vegetation. In this paper, we assessed the long term impact of white-tailed deer (Odocoileus virginianus) on plant communities of ombrotrophic (bog) and minerotrophic (fen) peatlands in eastern North America. Vegetation of five peatlands that have experienced high deer densities for at least 75 years was compared with that of five peatlands situated at proximity but on deer-free islands. We investigated deer impacts on plant species composition and cover, shrub height and cloudberry (Rubus chamaemorus) fruit/flower production and morphology. In bogs, white-tailed deer had no long-term impact on plant species assemblages, but reduced lichen cover and increased sedges and grasses cover as well as the surface area of bare peat. On the other hand, the floristic composition of fens differed significantly between sites where deer were present or absent. Plant diversity was greater in undisturbed fens than in disturbed fens, especially for shrubs, sedges and liverworts. No detrimental effects of browsing on shrub height were observed. Conversely, deer browsing seemed to have deleterious impacts on cloudberry fruit/flower production as well as on the number of leaves per individual. Overall, our results suggest that white-tailed deer had some important impacts on the vegetation of peatlands that could be harmful for the long-term conservation of peatland plant diversity.     

Long-term response of spring flora to chronic herbivory and deer exclusion in Great Smoky Mountains National Park, USA

Chronic herbivory by white-tailed deer (Odocoileus virginianus Zimmerman) can have profound impacts on the function and structure of forest ecosystems. We examined the combined influence of intense herbivory associated with a deer population eruption and chronic herbivory by the post eruption population on the spring flora of Cades Cove, Great Smoky Mountains National Park. During the 1970s the deer population reached a peak of 43 deer per km², from which it has slowly declined in recent decades. To examine the influence of intense herbivory, we compared the abundance and flowering rates of early flowering plants in Cades Cove to a nearby reference site with similar bedrock geology, vegetation, and disturbance history but contrasting history of deer abundance. Our results suggest that significant changes (p ? 0.05) in the diversity, evenness, and species richness of the spring flora occurred during the eruptive phase. Trillium spp. and other liliaceous species appeared to be disproportionately impacted. Comparisons between control and exclosure plots established after the deer population eruption indicate that recovery has been largely restricted to species that were able to persist under intense herbivory. These species have increased in number in exclosures, suggesting continued impacts by deer on the plant community outside the exclosures. Little to no recolonization by browse sensitive species was observed. Consequently, to restore the natural diversity of early flowering plants once present in Cades Cove, active restoration may be necessary in addition to maintaining deer densities below current levels.     

Conservation status and causes of decline of musk deer (Moschus spp.) in China

Five species of musk deer (Forest musk deer Moschus berezovskii, Alpine musk deer M. sifanicus, Black musk deer M. fuscus, Himalayan musk deer M. chrysogaster and Siberian musk deer M. moschiferus) occur throughout 17 provinces in China, and all were listed in the second category of the Chinese State Key Protected Wildlife List in 1998. Among these species, the Forest musk deer has the widest distribution, and its population is estimated at about 100,000-200,000 individuals in the wild. Additionally, there are about 100,000 Alpine musk deer and 2000 Siberian musk deer. As for the Black musk deer and Himalayan musk deer, which occur only in parts of Yunnan Province and Tibet, their populations remain unknown but they can be estimated to be rare due to their narrow and limited distribution. Furthermore, there are some 1900 musk deer kept in farms in China. Generally, the musk deer population has been declining due to over-hunting for musk and to loss of habitats. However, the Chinese government is greatly concerned with wildlife protection today. A series of laws and regulations have been launched and many natural reserves and national parks have been established as refuges for this animal, most of which, are within the musk deer's range, and should have positive effects for musk deer conservation.     

A natural experiment on the effects of high deer densities on the native flora of coastal temperate rain forests

The introduction of Sitka black-tailed deer (Odocoileus hemionus sitkensis Merriam) to Haida Gwaii (Queen Charlotte Islands, BC, Canada) in the late 19th century, provided an opportunity to understand the long-term effects of deer populations on the vegetation of temperate rain forests in the absence of their natural predators wolves (Canis lupus L.), and cougars (Puma concolor L.). Using seven small islands with different browsing histories (no deer, deer for <20 years, deer for >50 years), we tested the long-term effects of high deer densities on plant cover and species richness in the understorey of forest interior and forest edge habitats. Overall vegetation cover exceeded 80% in the lower vegetation layers on islands without deer and was less than 10% on the islands with deer for more than 50 years. Although overall plant species richness was similar on islands with or without deer, plant species richness at the plot scale (314 m²) was reduced by 20-50% on islands with deer for >50 years. The differences were most pronounced for the species-rich edge communities and among herb and shrub species. These results suggest that in the absence of predators, deer have the potential to greatly simplify the forest ecosystem.     

Trophic cascades involving cougar, mule deer, and black oaks in Yosemite National Park

Using a historical reconstruction, we discovered a potential trophic cascade involving cougar (Puma concolor), mule deer (Odocoileus hemionus), and California black oak (Quercus kelloggii) in the Yosemite Valley of Yosemite National Park in California. Our objective was to determine whether large deer populations in the absence of a top-level carnivore were suppressing tree regeneration. As human visitation increased in the early 1900s and cougar became increasingly scarce, the mule deer population irrupted in the 1920s. In August 2006, we undertook a retrospective study of black oak recruitment (i.e., growth of seedling/sprouts into tall saplings and trees) by inventorying all black oaks in stands accessible to deer and greater than 0.5 ha in size (18 stands, 2921 trees). We similarly inventoried oaks within sites representing refugia from deer browsing (4 stands, 481 trees). While significantly diminished oak recruitment has occurred since the 1920s in stands accessible to deer (p < 0.01), continuous recruitment of oaks was found in refugia sites. We also found less oak recruitment in areas of high human activity near the park’s visitor center (p < 0.01), possibly due to behaviorally-mediated effects of lower cougar and higher deer densities. Overall our results are consistent with trophic cascade theory involving large predators, herbivores, and plants. The long-term lack of oak recruitment is also an indicator of a probable loss of biodiversity.     

Effects of community-level grassland management on the non-target rare annual Agalinis auriculata

The objective of grassland management is to maintain keystone species, as well as species diversity, to promote a particular community structure, or to preserve specific ecosystem processes. Studies of grassland management rarely ascertain the effects on rare plant species, although conservation plans for rare herbaceous plants often recommend habitat management as well as restoration activities. Indeed, conservation biology has evolved from a relatively species-specific discipline into one more focused on larger scale issues such as ecosystem function, community composition and habitat restoration. Few studies have tested whether species of concern are adequately managed with a community or ecosystem-level approach. In this study, we evaluate whether community-level management of white-tailed deer and woody brush improves the viability of the rare annual plant species, Agalinis auriculata (Earleaf false foxglove; Orobanchaceae). Reducing deer browse and removing brush each positively affect plant vital rates. Brush removal increases the proportion of plants that reach the largest size class. Reducing deer browse increases the fertility of plants, particularly those in the largest size class. We report on demographic matrix models created with data from five populations of A. auriculata studied across 4 years. We find that both management activities positively affect the non-target plant species and are, in fact, necessary for A. auriculata to persist. Conservation of A. auriculata at our study sites requires both reducing the density of white-tailed deer and brush removal. Our study demonstrates that management at the community and ecosystem levels can be compatible with conservation goals at the species and population levels.     

Conservation of the red-winged grasshopper, Oedipoda germanica (Latr.): the influence of reproductive behaviour

Reproductive isolation can function as a mechanism to maintain locally adapted gene complexes while decreasing the heterozygosity in distinct populations. As a result, reproductive behaviour should be considered a fundamental factor influencing reproductive isolation. This is of interest to conservation biology when one desires to regulate gene flow between two populations either by creating opportunities for increased dispersal, by relocation of individuals, or by re-introduction of a species to its natural habitat. Reproductive behaviour can also influence the effective population size and the actual population size through the Allee effect. We investigated the reproductive behaviour of individuals from two isolated populations of the red winged grasshopper, Oedipoda germanica, an endangered species in Central Europe. We detail several methods to show how several aspects of the reproductive behaviour of this species interact with the conservation of this species. Foreign males were not disfavoured in mate choice and male body size was also unimportant in mating success. Heterogamic matings were as productive as homogamic matings in terms of total number of eggs per female, egg hatching rate, or nymph survival. Therefore, we suggest that cross-matings of individuals from different populations do not positively influence population size by heterosis effects nor act they negatively in the form of outbreeding depression. We found that female O. germanica were able to store viable sperm for extended periods but egg pods showed a decrease in hatching rate when these females were deprived of additional mating opportunities. Multiple mated females laid more eggs than once-mated females. Hence, females are capable of founding new populations even after only one mating but their reproductive output may be reduced. Present-day gene flow between populations of O. germanica probably does not occur. Consequently, there is an urgent need to pursue habitat management and release programmes that maintain current population sizes of this species. With regard to the reproductive behaviour of the red-winged grasshopper, we suggest that relocation programmes release males and female together and at an early adult stage. Also, because female reproductive output increases with mating activity, the initial release should involve excess females. Because males have a shorter lifespan than females a subsequent release of males at a later time may enhance the overall success of local conservation efforts by resulting in more offspring per female.     

Effects of sika deer on vegetation in Japan: A review

The Japanese sika deer Cervus nippon has expanded its range by nearly 70% during the last two decades. Browsing by sika deer affects vegetation in both agricultural and forested habitats. Effects of sika deer on vegetation are conspicuous on deer-inhabited islands: forest structure and composition are altered by deer grazing and browsing, and consequently regeneration is prevented. By felling of old trees, forest gaps are formed, but since sapling recruitment is prevented, shade intolerant plants invade. Unpalatable forbs like ragwort Senecio cannabifolius and ferns like bracken Pteridium aquilinum (Dennstaedtiaceae) become dominants in open habitats. At the places of highest deer density, the Zoysia japonica community, a low growing grass mat, develops. Sika deer function as a seed dispersal agent of this grass. Indirect effects of sika deer are not well studied, but some studies have shown reduction of understory bamboo cover, which in turn improves the survival of tree seedlings and declines of wood mice Apodemus spp. Japanese forests in lower mountainous areas were widely logged during the 1940s and 1950s. These areas were not well planted during the war and until 1950, but thereafter intensively planted to alter to conifer plantation as a nationwide campaign. For several decades after the war, rodents and hares grazed planted trees. According to growth of the planted trees to form canopy, which is unfavorable for rodents and hares, their damage declined. After the 1960s, old-growth forests in high mountainous areas were logged, and conifers were planted. Animals causing forestry damage were replaced by sika deer during the 1980s. Sika deer eat a wide variety of foods, and are gregarious, which causes heavy impacts on vegetation. Effects of sika deer are expanding to natural forests, alpine vegetation, and marshes. To reduce damages on forestry and natural vegetation, as many as 100,000 sika deer are culled every year. However, the hunter population is rapidly declining, and it is expected population control by culling will be insufficient. Although sika deer are an important component of Japanese forests, current population densities exceed the capacity of many ecosystems to tolerate herbivory.     

Demographic effects of road mortality in black ratsnakes (Elaphe obsoleta)

Roads negatively affect animal populations by presenting barriers to movement and gene flow and by causing mortality. We investigated the impact of a secondary road on a population of black ratsnakes (Elaphe obsoleta) in Ontario, Canada by radio-tracking 105 individuals over 8 years. The road was not a significant barrier to movement and none of the reproductive classes examined (male, non-reproductive female, reproductive female) avoided crossing the road. However, the road was a significant source of mortality. From a total of 115 road crossings by radio-implanted snakes, 3 individuals were killed by cars, resulting in a mortality rate of 0.026 deaths per crossing. We multiplied this mortality rate by the total number of expected road crossings by all individuals in the population in an active season (340) to estimate the number of road kills (9 individuals) each year. This estimate was higher than the actual number of road kills found, but half the number estimated from road kill models. Population viability analysis revealed that our estimate of road mortality was enough to increase the extinction probability for this population from 7.3% to 99% over 500 years. Road mortality of more than 3 adult females per year increased the extinction probability to >90%. Our results strengthen the view that road mortality can have a pronounced negative effect on populations of long-lived species.     

Pampas deer conservation with respect to habitat loss and protected area considerations in San Luis, Argentina

Ozotoceros bezoarticus celer is the most endangered subspecies of pampas deer. Although common in the Argentine Pampas 100 years ago, it persists in only two small populations. The largest population has survived due to the rarity of roads, internal farm subdivisions, and the low cattle density. However, habitat condition for this population has changed dramatically in the last 16 years. Five Landsat images (1985, 1992, 1997, 1999, 2001), covering 4608 km², were used to quantify pampas deer habitat loss due to the replacement of natural grassland by exotic pastures and crops. Image classification showed that natural grassland cover was reduced from 84.5 to 37.8% between 1985 and 2001. The annual transformation rate increased significantly from 1.4 to 10.9%. Average paddock size was significantly reduced from 1470 to 873 ha, and the number of paddocks increased from 129 to 227. The land within this area proposed for a national park has not escaped these habitat changes. In the last 6 years the amount of replaced area within the proposed park has increased from 9.1 to 51.1% due to actions by ranchers to avoid inclusion within park boundaries. Three patches of natural grassland still remain within the pampas deer distribution, one of which is the proposed national park. The implementation of a national park is a decisive challenge for the survival of pampas deer and its habitat in Argentina.     

Impacts of urbanization on Florida Key deer behavior and population dynamics

Rapid human population growth and urbanization have had a negative impact on species biodiversity. As competition for resources between man and wildlife continues, it is important to understand the effects of urbanization on species. Endangered Key deer (Odocoileus virginianus clavium) are endemic to the Florida Keys which have undergone rapid human population growth and development over the past 30 years. Our study objectives were to evaluate the impacts of urban development on Key deer habitat use, population dynamics, behavior, and body mass. We used data from two comprehensive studies on Key deer spanning 30 years to evaluate these changes. Our results suggest that Key deer have become more urbanized, using urban areas more today than they did 30 years ago. Contrary to our predictions, survival was higher for more urban deer than for less urban deer. Problems still exist with mortality factors heavily impacting some portions of the deer population including lower survival associated with less urban male deer. Analysis of Key deer body mass also was converse to our predictions as deer weights appear to have increased over time. Collectively, our results suggest that over the past 30 years Key deer have become more urbanized and that deer plasticity has allowed them to adapt and persist in an urbanizing environment. However, the future ability of Key deer to persist in an environment with continued urban development is unknown. At some threshold, urban development would become unsustainable and unlike other forms of habitat change or environmental disturbances, urban development is in most cases irreversible requiring careful planning in habitat conservation strategies.     

Two genetically distinct units of the Chinese sika deer (cervus nippon): analyses of mitochondrial DNA variation

Chinese sika deer, Cervus nippon, are currently threatened by habitat loss, fragmentation and human hunting, which has led to the extinction of three subspecies in the wild. The remaining subspecies subsist in the narrow regions of Jilin, Heilongjiang, Sichuan, Gansu, Jiangxi, Anhui, and Zhejiang provinces. In order to design effective conservation strategies for the Chinese sika deer, we have investigated genetic diversity, population structure and gene flow in the Chinese sika deer populations by analyzing ≈995 base pairs of the mitochondrial DNA (mtDNA) control region in 59 individuals sampled from the northeast of China, Sichuan, Jiangxi and Zhejiang. Chinese sika deer exhibited low mtDNA diversity and high gene flow among the four populations, and showed no strong geographical structure. The analyses of mtDNA variation among individuals of sika deer identified only two main phylogenetic groups even though three subspecies were sampled. These data singles out the Zhejiang population as being highly genetically distinct and worthy of separate conservation consideration. Therefore, it is recommended that a breeding program for the Zhejiang population be established.     

Female biased mortality caused by anthropogenic nest loss contributes to population decline and adult sex ratio of a meadow bird

Meadow breeding birds such as the whinchat Saxicola rubetra have been declining due to increased farming intensity. In modern grassland management, the first mowing and the bird’s breeding cycle coincide, causing high nest destruction rates and low productivity of grassland bird populations. However, it is virtually unknown whether the mowing process directly affects adult survival by accidentally killing incubating females. We studied adult survival of an Alpine whinchat population during two breeding seasons using either colour-ringing or radio-tracking of 71 adults. Assessing territories, mowing phenology and nest destruction from 1988 to 2007 allowed changes in the factors associated with female mowing mortality to be estimated. Adult survival over 5-day-periods was Φ = 0.986, but during the period of mowing female survival was strongly reduced (Φ = 0.946). As a result, 80.6% of the males, but only 68.4% of the females survived the breeding season. Mowing undoubtedly killed two of 20 radio-tagged females when they were laying or incubating. In the 20-year period, an increasing proportion of nests were destroyed before the chicks hatched and this change was associated with an increased distortion of the adult sex ratio. Modelling the population growth rate showed that including the additional effect of mowing on female mortality resulted in a 1.7 times faster local population decline. These results are consistent with the hypothesis that the extinction of whinchat populations in the lowlands of central Europe was caused not only by habitat degradation and low productivity, but also by increased man-made female mortality.     

Florida panther dispersal and conservation

We studied dispersal in 27 radio-collared Florida panthers Puma concolor coryi in southern Florida from 1986 to 2000. Male dispersal was longer (mean=68.4 km) than that of females (mean=20.3 km), tended to be circular, frustrated, and of insufficient length to ameliorate inbreeding. Females were philopatric and established home ranges that were less than one home range width away from their natal ranges. All females were successful in establishing territories, whereas males were successful 63% of the time. Dispersing panthers avoided moving toward the southeast and into an area of limited forest cover. Independence and the initiation of dispersal occurred at about 14 months of age and lasted for an average of 7.0-9.6 months for females and males, respectively. On average, Florida panthers disperse shorter distances than are typical for western populations of Puma concolor. A recent increase in long distance male dispersal events may be related to an increase in reproduction and population density resulting from the introduction of female cougars P. c. stanleyana into south Florida. Although the population exhibits the behavioral ability to colonize nearby vacant range, females have yet to do so. Successful dispersal to these areas could be facilitated by habitat restoration and translocation of females.     

Effects of Key deer herbivory on forest communities in the lower Florida Keys

Ungulate herbivory can have strong impacts on plant communities, but these impacts are rarely considered in recovery plans of endangered species. This study examined the effects of the endangered Key deer (Odocoileus virginianus clavium) on its environment in the lower Florida Keys. The Key deer population has increased to over 700 deer from approximately 50 deer in the 1950s; however, approximately 75% of the population resides on only a few islands (Big Pine, No Name, Big Munson) where Key deer herbivory on forest communities may be substantial. Effects of deer herbivory on plant densities were estimated on these islands using vegetation quadrats in hardwood hammock, buttonwood transition, and mangrove wetlands and compared to nine other islands with intermediate or low deer densities. On islands with high deer density, densities of preferred woody plant species <1.2 m tall (within Key deer reach) were significantly lower than islands with lower deer densities, while densities of some nonpreferred species were significantly higher. Deer exclosures established in hardwood hammock on a high-density deer island revealed a mean increase in abundance/height of preferred woody species inside exclosures, while nonpreferred species significantly increased in open plots. We conclude that on high deer density islands, highly preferred plant species might eventually fail to regenerate and unpalatable plant species may become dominant. Careful criteria need to be developed to maintain Key deer numbers above an endangered species status yet below levels that are destructive to local forest species.     

Genetic diversity and local population structure of fragmented populations of Trillium camschatcense (Trilliaceae)

Trillium camschatcense, a long-lived common woodland herb, has been experiencing intensive habitat fragmentation over the last century in eastern Hokkaido, Japan. We examined the genetic diversity and population genetic structure of 12 fragmented populations with different population sizes using allozyme electrophoresis. The percentage of polymorphic loci and mean number of alleles per locus were positively related to population size, probably due to the stochastic loss of rare alleles (frequency of q<0.1) in small populations. Populations with 350 flowering plants or fewer had lost almost all of their rare alleles. While the heterozygosity and inbreeding coefficient were not related to population size, some small populations showed relatively high inbreeding coefficients. In spite of the low genetic differentiation among overall populations (F_ST=0.130), local population structuring was recognized between the two geographically discontinuous population groups. Within groups, sufficient historical gene flow was inferred, whereas a low dispersal ability of this species and geographical separation could produce apparent differentiation between groups.     

Expectations for population growth at new breeding locations for the vulnerable New Zealand sea lion (Phocarctos hookeri) using a simulation model

Management plans for threatened or recovering large vertebrate species that are increasing in population size and range focus on the establishment of viable populations within set temporal limits. New Zealand (Hookers) sea lions (Phocarctos hookeri) were declared a threatened species in 1997, and New Zealand legislation requires that threatened species of marine mammals must be managed to reduce human-induced mortality and achieve a non-threatened status within 20 years. The present breeding distribution of P. hookeri is highly localised, with over 95% of total annual pup production located at Auckland Islands and almost all of the remainder at Campbell Island. Breeding elsewhere has been ephemeral or restricted to <10 adult females. The only recorded sustainable breeding at a new location has been at Otago, South Island, New Zealand. This breeding population consisted of a total of four breeding females in 2002 and is derived from one immigrant female that gave birth to her first pup in the 1993/1994 breeding season. The New Zealand Department of Conservation management plan specifies that to achieve a non-threatened status P. hookeri (1) at Otago must increase in the number of breeding females to ?10, and (2) must establish ?two new breeding locations within the 20-year time frame, each with ?10 breeding females. This study 1) projects the population growth trends at a new location (Otago) to see if it will achieve ?10 breeding females within the legislated time frame, and (2) examines the likelihood that other breeding locations will establish elsewhere given the demographic information available for this species. We present 20 deterministic and three stochastic Leslie matrix model scenarios for female population growth for the initial years following the start of breeding at a new location. Our results indicate that (1) a new breeding population derived from one immigrant female is unlikely to reach 10 breeding females in 20 years; this duration is more likely to be 23-41 years (deterministic models) or 23-26 years (stochastic model), (2) the likelihood of two new sites establishing within 20 years is unquantifiable, but the probability is low, and (3) if the legislated outcome and time limit are not revised in the population management plan, the feasibility and effectiveness of re-locating young females could be investigated.     

Protecting more than the wetland: The importance of biased sex ratios and habitat segregation for conservation of the Hine’s emerald dragonfly, Somatochlora hineana Williamson

Within species habitat use may depend on age, season or sex of an individual. The distribution of males and females may vary both temporally and spatially due to differences in the costs of reproduction and the distribution of critical resources. Conservation of a species requires knowledge of the habitat use of both sexes in order to predict the population size and protect all habitats that a species requires. Adult dragonfly populations often have highly male-biased sex ratios at the breeding habitat. This bias has been attributed to females using alternative habitats to avoid male harassment, or to high female mortality. We monitored adult Hine’s emerald dragonfly (Somatochlora hineana Williamson) populations, in breeding and non-breeding habitats in Door County, Wisconsin and found significant differences in habitat use between males and females. Males primarily used wetland habitats, while females primarily used dry meadows and marginal breeding habitats, only coming into wetlands to lay-eggs or find mates. We assessed food resources in the different habitats and found that high quality insect prey (primarily adult Diptera) were more available in the wetland habitat, indicating that these areas were likely a more productive foraging area for adult dragonflies. The fact that females appear to avoid the wetland habitat is consistent with the hypothesis that male harassment alters female distribution patterns. Consideration of the patterns of habitat use by S. hineana indicates the need to develop a broader understanding of the importance of non-wetland areas in the conservation of wetland species.     

Viability of mountain caribou in British Columbia, Canada: Effects of habitat change and population density

Population viability analyses (PVA) are frequently employed to develop recovery plans and inform management of endangered species. Translating results from PVA into meaningful management recommendations often depends on an understanding of how population parameters change with environmental conditions as well as population density. The decline of mountain caribou (Rangifer tarandus caribou) in British Columbia, Canada, is believed to be caused by apparent competition with alternative prey species following changes to the forest age structure from timber harvest and wildfire. In addition, populations have been shown to decline at faster rates at low population density. To evaluate the potential effects of habitat change and population density on population persistence, we used stochastic projection models for 10 distinct populations varying in initial size from <10 to approximately 150 females. In an initial model, we used estimates of vital rates based on information sampled from >350 radiocollared caribou between 1984 and 2004. We then compared the results of the initial model to a set of models that evaluated the effects of habitat conditions and population density via their expected relationships to female adult survival. Assuming that vital rates remain constant over a 200-year time frame, only three populations have high probabilities (>0.95) of extinction. When models incorporate the declines in adult female survival know to occur with increasing proportions of young forest and declining population densities, all 10 populations are predicted to decline to extinction within <200 years. Based on our results, we suggest that PVA models that fail to incorporate the effects of changes in vital rates with habitat and population density may lead to overly optimistic assessments of the probability of population persistence in endangered species.