Effects of law enforcement and community outreach on mammal diversity in a biodiversity hotspot

Management activities such as law enforcement and community outreach are thought to affect conservation outcomes in protected areas, but their importance relative to intrinsic environmental characteristics of the parks and extrinsic human pressures surrounding the parks have not been explored. Furthermore, it is not clear which is more related to conservation outcomes—the management itself or local people's perceptions of the management. We measured objective (reports by park staff) and subjective (reports by local people) levels of community outreach and law enforcement based on responses to 374 questionnaires. We estimated mammal abundance and diversity of 6 protected areas based on data from 115 camera traps in Xishuangbanna, southwest China, a biodiversity hotspot with high hunting and land-conversion pressures. We then examined correlations among them and found that local people's perception of law enforcement was positively related to the local abundance of 2 large, hunted species, wild boar (Sus scrofa) (β = 15.22) and muntjac (Muntiacus vaginalis) (β = 14.82), but not related to the abundance of smaller mammals or to objective levels of enforcement. The subjective frequency of outreach by park staff to local communities (β = 3.42) and park size (β = 3.28) were significantly and positively related to mammal species richness, whereas elevation, human population density, and subjective frequency of law enforcement were not. We could not conclude that community outreach and law enforcement were directly causing increased mammal abundance and diversity. Nevertheless, the patterns we detected are some of the first empirical evidence consistent with the idea that biodiversity in protected areas may be more positively and strongly related to local perceptions of the intensity of park management than to either intrinsic (e.g., elevation, park size) or extrinsic (e.g., human population density) environmental factors.     

Insularization of Tanzanian Parks and the Local Extinction of Large Mammals

Island biogeography theory predicts that species will be lost on habitat "islands" created by the fragmentation of continental regions. Many Tanzanian parks are rapidly becoming habitat islands as a result of human settlement, agricultural development, and the active elimination of wildlife on adjacent lands. The rate of extinction of mammals in six Tanzanian parks over the last 35–83 years is significantly and inversely related to park area, suggesting that increasing insularization of the parks has been an important contributory factor in large mammal extinctions. I compared observed patterns of persistence of mammals in Tanzanian parks to predictions derived from earlier extinction models. The predictions of the S ¹ models of Soulé et al. (1979) and Burkey (1994) and the S ² and S ³ models of Soulé et al. (1979) match very closely the observed pattern of persistence of mammals in Tanzanian parks. The loss of mammal species will probably continue, particularly in the smaller parks. Establishment of wildlife corridors linking the parks in northern Tanzania could help to reduce the potential loss of species in the future.     

Area-based assessment of extinction risk

Underpinning the International Union for Conservation of Nature (IUCN) Red List is the assessment of extinction risk as determined by the size and degree of loss of populations. The IUCN system lists a species as Critically Endangered, Endangered, or Vulnerable if its population size declines 80%, 50%, or 30% within a given time frame. However, effective implementation of the system faces substantial challenges and uncertainty because geographic scale data on population size and long-term dynamics are scarce. I develop a model to quantify extinction risk using a measure based on a species' distribution, a much more readily obtained quantity. The model calculates the loss of the area of occupancy that is equivalent to the loss of a given proportion of a population. It is a very simple yet general model that has no free parameters and is independent of scale. The model predicted well the distributions of 302 tree species at a local scale and the distributions of 348 species of North American land birds. This area-based model provides a solution to the long-standing problem for IUCN assessments of lack of data on population sizes, and thus it will contribute to facilitating the quantification of extinction risk worldwide.     

Life-history correlates of extinction risk and recovery potential

Extinction risk is inversely associated with maximum per capita population growth rate (r(max)). However, this parameter is not known for most threatened species, underscoring the value in identifying correlates of r(max) that, in the absence of demographic data, would indirectly allow one to identify species and populations at elevated risk of extinction and their associated recovery potential. We undertook a comparative life-history analysis of 199 species from three taxonomic classes: Chondrichthyes (e.g., sharks; n = 82), Actinopterygii (teleost or bony fishes; n = 47), and Mammalia (n = 70, including 16 marine species). Median r(max) was highest for (and similar between) terrestrial mammals (0.71) and teleosts (0.43), significantly lower among chondrichthyans (0.26), and lower still in marine mammals (0.07). Age at maturity was the primary (and negative) correlate of r(max). In contrast, although body size was negatively correlated with r(max) in chondrichthyans and mammals, evidence of an association in teleosts was equivocal, and fecundity was not related to r(max) in fishes, despite recurring assertions to the contrary. Our analyses suggest that age at maturity can serve as a universal predictor of extinction risk in fishes and mammals when r(max) itself is unknown. Moreover, in contrast to what is generally expected, the recovery potential of teleost fishes does not differ from that of terrestrial mammals. Our findings are supportive of the application of extinction-risk criteria that are based on generation time and that are independent of taxonomic affinity.     

Extinction Rate Estimates for Plant Populations in Revisitation Studies: Importance of Detectability

Abstract:  Many researchers have obtained extinction-rate estimates for plant populations by comparing historical and current records of occurrence. A population that is no longer found is assumed to have gone extinct. Extinction can then be related to characteristics of these populations, such as habitat type, size, or species, to test ideas about what factors may affect extinction. Such studies neglect the fact that a population may be overlooked, however, which may bias estimates of extinction rates upward. In addition, if populations are unequally detectable across groups to be compared, such as habitat type or population size, comparisons become distorted to an unknown degree. To illustrate the problem, I simulated two data sets, assuming a constant extinction rate, in which populations occurred in different habitats or habitats of different size and these factors affected their detectability. The conventional analysis implicitly assumed that detectability equalled 1 and used logistic regression to estimate extinction rates. It wrongly identified habitat and population size as factors affecting extinction risk. In contrast, with capture-recapture methods, unbiased estimates of extinction rates were recovered. I argue that capture-recapture methods should be considered more often in estimations of demographic parameters in plant populations and communities.     

Wildlife Disease in U.S. National Parks: Historical and Coevolutionary Perspectives

Diseases of wildlife have significant management implications in a number of lands of the U.S. National Park Service due to increasing interactions between wildlife and domestic animals. We review the paleontology, history, and coevolution of infectious diseases in North American ungulates. We provide two examples related to bovine brucellosis in bison in Yellowstone National Park and lungworm-pneumonia complex in bighorn sheep in several western national parks. These examples illustrate the difficulty of managing wild populations and their diseases in national parks and other protected areas. In some instances, human intervention may be justifiable in order to protect native populations, domestic animals, and humans from acquiring a disease.     

Effects of Human Activity on Global Extinction Risk

Both natural and anthropogenic factors are important in determining a species' risk of extinction. Little work has been done, however, to quantify the magnitude of current anthropogenic influences on the extinction process. The purpose of this study is to determine the extent to which measures of the intensity of human activity are related to the global variability of two measures of species' susceptibility to extinction. We observed six indices of human activities in 90 countries, and we tested their relationships to the proportion of threatened bird and mammal species in each country, as well as to mammalian population density. After correcting for area effects, latitudinal diversity gradients, and body size (for population density), 28 to 50% of the remaining variation was statistically attributable to anthropogenic variables. Different measures of anthropogenic influence were most closely related to extinction risk in birds and mammals. Human population density was the variable most closely related to the proportion of threatened bird species per country, whereas per capita GNP was more important for mammal species. Mammalian population density strongly correlates with the extent of protected area per country. Contrary to suggestions in earlier literature, our work does not support the hypothesis that habitat loss is a prime contributor to species loss because frequencies of threatened birds and mammals are not closely related to patterns of land use.     

Changing Species Richness and Composition in Canadian National Parks

Abstract: Canada's national parks and their surrounding areas differ markedly in size, climate, vegetation, and extent of human development. We tested the extent to which total species richness, native species richness, and the number of extirpations and introductions of terrestrial vertebrates were correlated with each of these factors. To do this, we used surveys of park fauna from the present and from the time of park establishment. Richness, extirpations, and introductions were all strongly related to climate. After we controlled for climate, smaller parks had higher rates of species loss than larger parks. Land-use patterns (forest cover and fragmentation, roads, etc.) within parks were strongly correlated with land use in the regions surrounding the parks, showing that parks have not been isolated from regional development. Richness and extirpations within parks were generally more strongly related to regional characteristics than to the characteristics of the parks themselves. Species richness and numbers of introduced species were higher in parks found in landscapes with greater fragmentation. Frequencies of extirpations were less clearly related to human-influenced habitat characteristics. Introductions and extinctions most often involved game species or species directly associated with human activities. There is little evidence of subtle ecological effects being responsible for species loss. Our results suggest that management should focus on direct human interventions, such as hunting, introduction of game species, and habitat fragmentation, in parks and surrounding areas.     

Quantifying Relative Extinction Risks and Targeting Intervention for the Orchid Flora of a Natural Park in the European Prealps

Abstract:  Conservation currently relies largely on hindsight because demographic studies identify population decline after the event. Nevertheless, the degree of aggregation within a population is an "instantaneous" characteristic with the potential to identify populations presently at greatest risk of genetic impoverishment (via Allee effects and in-breeding depression) and local decline. We sought to determine the relative extinction risk for sympatric orchid species throughout Monte Barro natural park (Lecco, Italy), based on an index of dispersion ( I ) calculated from the size and location of subpopulations (recorded with GPS and mapped with GIS). Three population dispersion types were identified: (1) highly aggregated and locally abundant (large subpopulations restricted to particular sites; e.g., Gymnadenia conopsea [L.] R.Br.; I = 54.5); (2) widespread and moderately aggregated (opportunistic throughout the elevational range of the mountain; e.g., Listera ovata [L.] R.Br.; I = 18.9); and (3) weakly aggregated and locally rare (small, highly diffuse subpopulations; e.g., endemic Ophrys benacensis [Reisigl] O. & E. Danesch & Ehrend.; I = 4.4). Type 1 populations are more likely to respond to in situ intervention, whereas type 2 are relatively invasive species for which conservation intervention is not necessary, and type 3 are rare species that are least likely to respond to habitat management, for which ex situ conservation and population reinforcement would be most appropriate. Although our methodology provides only a "snapshot" of aboveground patterns of population dispersion, it can help target the application of in situ and ex situ conservation activities proactively and is of particular utility for parks for which a rapid assessment of local extinction risks is needed.     

Conservation Implications of the Natural Loss of Lineages in Wild Mammals and Birds

Because populations in zoological parks and nature reserves often are derived from only a few individuals, conservationists have attempted to minimize founder effects by equalizing family group sizes and increasing the reproductive contributions of all individuals. Although such programs reduce potential losses of genetic diversity, information is rarely available about the actual persistence of family groups or genetic lineages in natural populations. In the absence of such data, it can be difficult to weigh the importance of human intervention in the conservation of small populations. Separate long-term studies of two mammals, the North American bison (Bison bison) and the white-nosed coati (Nasua narica), and a bird, the Acorn Woodpecker (Melanerpes formicivorus), demonstrate differential extinction of genetic lineages. Irrespective of the mechanisms affecting population structure, which may range from stochastic environmental events to such behavioral phenomena as poor intrasexual competitive abilities, our results show that lineages can be lost at rapid rates from natural populations. A survey of comparable studies from the literature indicates that the loss of matrilines over the course of the study varies from 3% to 87% in wild mammals and from 30% to 80% in birds, with several small mammals losing approximately 20% of matrilines per year of study. These lineage extinctions were not an artifact of the length of the study or the generation time of the species. Such rapid losses of lineages in less than 20-year periods in natural populations suggest that efforts to maintain maximal genetic diversity within populations may not always reflect processes that occur in the wild. Conservation biologists need to give further thought to the extent to which parity among genetic lines should be a primary goal of management of captive and small wild populations.     

Changes in size and trends of North American sea duck populations associated with North Pacific oceanic regime shifts

Broad-scale multi-species declines in populations of North American sea ducks for unknown reasons is cause for management concern. Oceanic regime shifts have been associated with rapid changes in ecosystem structure of the North Pacific and Bering Sea. However, relatively little is known about potential effects of these changes in oceanic conditions on marine bird populations at broad scales. I examined changes in North American breeding populations of sea ducks from 1957 to 2011 in relation to potential oceanic regime shifts in the North Pacific in 1977, 1989, and 1998. There was strong support for population-level effects of regime shifts in 1977 and 1989, but little support for an effect of the 1998 shift. The continental-level effects of these regime shifts differed across species groups and time. Based on patterns of sea duck population dynamics associated with regime shifts, it is unclear if the mechanism of change relates to survival or reproduction. Results of this analysis support the hypothesis that population size and trends of North American sea ducks are strongly influenced by oceanic conditions. The perceived population declines appear to have halted >20 years ago, and populations have been relatively stable or increasing since that time. Given these results, we should reasonably expect dramatic changes in sea duck population status and trends with future oceanic regime shifts.     

Life history and viability of a long-lived marine invertebrate: the octocoral Paramuricea clavata

The red gorgonian Paramuricea clavata is a long-lived, slow-growing sessile invertebrate of ecological and conservation importance in the northwestern Mediterranean Sea. We develop a series of size-based matrix models for two Paramuricea clavata populations. These models were used to estimate basic life history traits for this species and to evaluate the viability of the red gorgonian populations we studied. As for many other slow-growing species, sensitivity and elasticity analysis demonstrate that gorgonian population growth is far more sensitive to changes in survival rates than to growth, shrinkage, or reproductive rates. The slow growth and low mortality of red gorgonians results in low damping ratios, indicating slow convergence to stable size structures (at least 50 years). The stable distributions predicted by the model did not differ from the observed ones. However, our simulations point out the fragility of this species, showing both populations in decline and high risk of extinction over moderate time horizons. These declines appear to be related to a recent increase in anthropogenic disturbances. Relative to their life span, the values of recruitment elasticity for Paramuricea clavata are lower than those reported for other marine organisms but are similar to those reported for some long-lived plants. These values and the delayed age of sexual maturity, in combination with the longevity of the species, show a clear fecundity/mortality trade-off. Full demographic studies of sessile marine species are quite scarce but can provide insight into population dynamics and life history patterns for these difficult and under-studied species. While our work shows clear results for the red gorgonian, the variability in some of our estimates suggest that future work should include data collection over longer temporal and spatial scales to better understand the long-term effects of natural and anthropogenic disturbances on red gorgonian populations.     

On the generality of a climate-mediated shift in the distribution of the American pika (Ochotona princeps)

Alpine species are among those most threatened by climatic shifts due to their physiological and geographic constraints. The American pika (Ochotona princeps), a small mammal found in mountainous, rocky habitats throughout much of western North America, has experienced recent population extirpations in the Great Basin linked to climatic drivers. It remains unclear whether these patterns of climate-related loss extend to other portions of the species' range. We investigated the distribution of the American pika and the climatic processes shaping this distribution within the Southern Rocky Mountain region. Results from a survey of 69 sites historically occupied by pikas indicate that only four populations have been extirpated within this region over the past few decades. Despite relatively few extirpations, low annual precipitation is implicated as a limiting factor for pika persistence in the Southern Rockies. Extirpations occurred only at sites that were consistently dry over the last century. While there was no climate change signal in our results, these data provide valuable insight into the potential future effects of climate change on O. princeps throughout its range.     

Optimism and Challenge for Science‐Based Conservation of Migratory Species in and out of U.S. National Parks

Public agencies sometimes seek outside guidance when capacity to achieve their mission is limited. Through a cooperative agreement and collaborations with the U.S. National Park Service (NPS), we developed recommendations for a conservation program for migratory species. Although NPS manages ～36 million hectares of land and water in 401 units, there is no centralized program to conserve wild animals reliant on NPS units that also migrate hundreds to thousands of kilometers beyond parks. Migrations are imperiled by habitat destruction, unsustainable harvest, climate change, and other impediments. A successful program to counter these challenges requires public support, national and international outreach, and flourishing migrant populations. We recommended two initial steps. First, in the short term, launch or build on a suite of projects for high‐profile migratory species that can serve as proof to demonstrate the centrality of NPS units to conservation at different scales. Second, over the longer term, build new capacity to conserve migratory species. Capacity building will entail increasing the limited knowledge among park staff about how and where species or populations migrate, conditions that enable migration, and identifying species’ needs and resolving them both within and beyond parks. Building capacity will also require ensuring that park superintendents and staff at all levels support conservation beyond statutory borders. Until additional diverse stakeholders and a broader American public realize what can be lost and do more to protect it and engage more with land management agencies to implement actions that facilitate conservation, long distance migrations are increasingly likely to become phenomena of the past. Optimismo y Retos para la Conservación Científicamente Basada de Especies Migratorias Dentro y Fuera de Parques Nacionales de E.U.A.     

The unified neutral theory of biodiversity: do the numbers add up?

Hubbell's unified neutral theory is a zero-sum ecological drift model in which population sizes change at random in a process resembling genetic drift, eventually leading to extinction. Diversity is maintained within the community by speciation. Hubbell's model makes predictions about the distribution of species abundances within communities and the turnover of species from place to place (beta diversity). However, ecological drift cannot be tested adequately against these predictions without independent estimates of speciation rates, population sizes, and dispersal distances. A more practical prediction from ecological drift is that time to extinction of a population of size N is approximately 2N generations. I test this prediction here using data for passerine birds (Passeriformes). Waiting times to speciation and extinction were estimated from genetic divergence between sister populations and a lineage-through-time plot for endemic South American suboscine passerines. Population sizes were estimated from local counts of birds in two large forest plots extrapolated to the area of wet tropical forest in South America and from atlas data on European passerines. Waiting times to extinction (ca. 2 Ma) are much less than twice the product of average population size (4.0 and 14.4 x 10(6) individuals in South America and Europe) and generation length (five and three years) for songbirds, that is, 40 and 86 Ma, respectively. Thus, drift is too slow to account for turnover in regional avifaunas. Presumably, other processes, involving external drivers, such as climate and physiographic change, and internal drivers, such as evolutionary change in antagonistic interactions, predominate. Hubbell's model is historical and geographic, and his perspective importantly links local and regional process and pattern. Ecological reality can be added to the mix while retaining Hubbell's concept of continuity of communities in space and time.     

Extinction times in experimental populations

Predicting population extinctions is a key element of quantitative conservation biology and population ecology. Although stochastic population theories have long been used to obtain theoretical distributions of population extinction times, model-based predictions have rarely been tested. Here I report results from a quantitative analysis of extinction time in 281 experimental populations of water fleas (Daphnia magna) in variable environments. To my knowledge, this is the first quantitative estimate of the shape of the distribution of population extinction times based on extinction data for any species. The finding that the distribution of population extinction times was extraordinarily peaked is consistent with theoretical predictions for density-independent populations, but inconsistent with predictions for density-dependent populations. The tail of the extinction time distribution was not exponential. These results imply that our current theories of extinction are inadequate. Future work should focus on how demographic stochasticity scales with population size and effects of nonrandom variable environments on population growth and decline.     

Threshold to maturity in a long-lived reptile: interactions of age, size, and growth

Thresholds to sexual maturity—either age or size—are critical life history parameters. Usually investigated in short-lived organisms, these thresholds and interactions among age, size, and growth are poorly known for long-lived species. A 34-year study of captive green turtles (Chelonia mydas) that followed individuals from hatching to beyond maturity provided an opportunity to evaluate these parameters in a long-lived species with late maturity. Age and size at maturity are best predicted by linear growth rate and mass growth rate, respectively. At maturity, resource allocation shifts from growth to reproductive output, regardless of nutrient availability or size at maturity. Although captive turtles reach maturity at younger ages than wild turtles, the extensive variation in captive turtles under similar conditions provides important insights into the variation that would exist in wild populations experiencing stochastic conditions. Variation in age/size at maturity should be incorporated into population models for conservation and management planning.     

Avian Survival Rates and the Extinction Process on Barro Colorado Island, Panama

Abstract: Selective extinction following isolation of habitat patches may be due to biogeographical (e.g., island size or isolation) and ecological (species natural histories, interspecifc interactions) factors, or their interactions. Among the demographic and life history attributes commonly associated with high extinction probability are small populations, large size of individuals, and population variability. Long-term capture-recapture data from forest habitat in central Panama permit an examination of the association between mainland survival rates and extinction on a nearby land-bridge island Species of birds that no longer occur on Barro Colorado Island (BCI), Panama, have, on average, lower survival rates on the adjacent mainland than species that have persisted on BCI. Moreover, of the species that no longer occur on BCI, those with lower mainland survival rates generally disappeared earlier from the island. My analysis provides little evidence of a relationship between extinction and population size. Recolonization of BCI from the adjacent mainland by the forest undergrowth species studied here is unlikely. Reduced reproductive success on BCI combined with naturally low adult survival rates seems to be responsible for these BCI extinctions. High nest predation and/or altered landscape dynamics are probable agents in the low reproductive success. The methods used here could be employed in other circumstances to identify fragmentation-sensitive species.     

Topology of Extinction and Endangerment of Native Fishes in the Pacific Northwest and California (U.S.A.)

Recent studies have provided a broad data base on extinction and endangerment of species, subspecies, and distinct populations of inland fishes in western North America. Development of a synoptic, regional-scale image of extinction and risk of extinction is complicated by the small size and linear distribution of fluvial aquatic habitats and by interspecific variation in areal extent of populations. I developed a regional map of extinction-risk isopleths based on the number of extinct and persistently declining species in drainage basins of the Pacific Northwest and California. This topological synthesis is useful for delineating and monitoring areas of historic and ongoing loss of aquatic biodiversity, and for relating losses to patterns of land use and habitat modification, climate hydrology, and geomorphology. From an ecological perspective, endangerment of numerous indigenous populations of seven widely distributed species of anadromous salmonids in this region is as important as the more local, diffuse effects of declines in more than 60 endemic, nonanadromous species and subspecies. The simultaneous decline of numerous taxa in basins not afflicted with dams or diversions suggests that cumulative damage to aquatic habitats caused by logging, grazing, urbanization, and other land uses plays a major role in icthyofaunal impoverishment.