Importance of Habitat Quality and Landscape Connectivity for the Persistence of Endangered Natterjack Toads

Abstract: The natterjack toad (Bufo calamita) is endangered in several parts of its distribution, including Belgium, where it occurs mainly in artificial habitats. We parameterized a general model for natterjack population viability analysis (PVA) and tested its sensitivity to changes in the values of basic parameters. Then we assessed the relative efficiency of various conservation measures in 2 situations: a small isolated population and a system of 4 populations connected by rare dispersal movements. We based the population viability analysis on a stage‐structured model of natterjack population dynamics. We parameterized the model in the RAMAS GIS platform with vital rates obtained from our own field experience and from published studies. Simulated natterjack populations were highly sensitive to habitat quality (particularly pond drying), to dispersal from surrounding local populations, and to a lesser extent to values of fecundity and survival of terrestrial stages. Population trajectories were nearly insensitive to initial abundances, carrying capacities, and the frequency of extreme climatic conditions. The simulations showed that in habitats with highly ephemeral ponds, where premetamorphosis mortality was high, natterjack populations nearly always had a very high extinction risk. We also illustrated how low dispersal rates (<1 dispersing individual/generation) efficiently rescued declining local populations. Such source‐sink dynamics demonstrate that the identification and management of source populations should be a high priority.     

Dispersal, density dependence, and population dynamics of a fungal microbe on leaf surfaces

Despite the ubiquity and importance of microbes in nature, little is known about their natural population dynamics, especially for those that occupy terrestrial habitats. Here we investigate the dynamics of the yeast-like fungus Aureobasidium pullulans (Ap) on apple leaves in an orchard. We asked three questions. (1) Is variation in fungal population density among leaves caused by variation in leaf carrying capacities and strong density-dependent population growth that maintains densities near carrying capacity? (2) Do resident populations have competitive advantages over immigrant cells? (3) Do Ap dynamics differ at different times during the growing season? To address these questions, we performed two experiments at different times in the growing season. Both experiments used a 2 x 2 factorial design: treatment 1 removed fungal cells from leaves to reveal density-dependent population growth, and treatment 2 inoculated leaves with an Ap strain engineered to express green fluorescent protein (GFP), which made it possible to track the fate of immigrant cells. The experiments showed that natural populations of Ap vary greatly in density due to sustained differences in carrying capacities among leaves. The maintenance of populations close to carrying capacities indicates strong density-dependent processes. Furthermore, resident populations are strongly competitive against immigrants, while immigrants have little impact on residents. Finally, statistical models showed high population growth rates of resident cells in one experiment but not in the other, suggesting that Ap experiences relatively "good" and "bad" periods for population growth. This picture of Ap dynamics conforms to commonly held, but rarely demonstrated, expectations of microbe dynamics in nature. It also highlights the importance of local processes, as opposed to immigration, in determining the abundance and dynamics of microbes on surfaces in terrestrial systems.     

Source-Sink Models and the Problem of Habitat Degradation: General Models and Applications to the Yellowstone Grizzly

I used source-sink population models to explore the consequences of habitat degradation for populations living on good and degraded habitats linked by movement. In particular, I modeled the conversion of land from good habitat quality supporting positive population growth to a degraded condition in which there was population decline. I found that with high rates of movement between good and bad quality areas populations require relatively large amounts of good habitat to remain stable. However, low movement rates resulted in greater sensitivity of population growth to habitat loss. Even small amounts of habitat degradation could result in rapid changes in overall population growth rates depending upon the rates of population increase and decline in the two habitat types. I also developed and simulated an age-structured model for grizzly bears ( Ursus arctos horribilis ) existing in good and degraded habitats and fit this model to data from the Yellowstone grizzly population. I used this model to predict the ability to detect crucial amounts of habitat degradation from census data and found that when degradation is slow (e.g., 1% conversion of good to poor habitat per year), more than a decade may pass between crucial amounts of degradation—beyond which populations begin long-term decline—and its detection, even if census data were unrealistically good. Thus these simple models indicate that, at least in some circumstances, habitat degradation can have rapid and severe impacts on population dynamics and traditional monitoring programs may not be adequate to detect the consequences of degradation.     

Can we measure carrying capacity with foraging behavior?

Carrying capacity is one of the most important, yet least understood and rarely estimated, parameters in population management and modeling. A simple behavioral metric of carrying capacity would advance theory, conservation, and management of biological populations. Such a metric should be possible because behavior is finely attuned to variation in environment including population density. We connect optimal foraging theory with population dynamics and life history to develop a simple model that predicts this sort of adaptive density-dependent change in food consumption. We then confirm the model's unexpected and manifold predictions with field experiments. The theory predicts reproductive thresholds that alter the marginal value of energy as well as the value of time. Both effects cause a pronounced discontinuity in quitting-harvest rate that we revealed with foraging experiments. Red-backed voles maintained across a range of high densities foraged at a lower density-dependent rate than the same animals exposed to low-density treatments. The change in harvest rate is diagnostic of populations that exceed their carrying capacity. Ecologists, conservation biologists, and wildlife managers may thus be able to use simple and efficient foraging experiments to estimate carrying capacity and habitat quality.     

Conservation of Species in Dynamic Landscapes: Divergent Fates of Silene tatarica Populations in Riparian Habitats

Abstract:  In transient environments, where local extinctions occur as a result of destruction or deterioration of the local habitat, the long-term persistence of a species requires successful colonizations at new, suitable sites. This kind of habitat tracking should be associated with the asynchronous dynamics of local populations, and it can be especially important for the conservation of rare plant species in riparian habitats. We determined spatiotemporal variation in the demography of the perennial Silene tatarica (L.) Pers. in 15 populations (1998–2003) located in periodically disturbed riparian habitats. The habitats differed according to their morphology (flat shores, slopes) and the amount of bare ground (open, intermediate, closed) along a successional gradient. We used elasticity and life-table response analyses and stochastic simulations to study the variation in population demography. Finite population growth rate was higher in intermediate habitats than in open and closed habitats. In stochastic simulations population size increased in most cases, but four populations were projected to become extinct within 12–70 years. The viability of local populations depended most on the survival and growth of juvenile individuals and on the fecundity of large fertile individuals. On a regional scale, the persistence of this species will require a viable network of local populations as protection against local extinctions caused by natural disturbances and succession. Accordingly, the long-term persistence of riparian species may depend on habitat changes; thus, their conservation requires maintenance of natural disturbance dynamics. Along regulated rivers, management activities such as the creation of open habitats for new colonization should be implemented. Similarly, these activities can be rather general requirements for the conservation of endangered species dependent on transient habitats along successional gradients.     

Role of Patch Size, Disease, and Movement in Rapid Extinction of Bighorn Sheep

Abstract: The controversy (  Berger 1990, 1999 ; Wehausen 1999 ) over rapid extinction in bighorn sheep ( Ovis canadensis ) has focused on population size alone as a correlate to persistence time. We report on the persistence and population performance of 24 translocated populations of bighorn sheep. Persistence in these sheep was strongly correlated with larger patch sizes, greater distance to domestic sheep, higher population growth rates, and migratory movements, as well as to larger population sizes. Persistence was also positively correlated with larger average home-range size ( p = 0.058, n = 10 translocated populations) and home-range size of rams ( p = 0.087, n = 8 translocated populations). Greater home-range size and dispersal rates of bighorn sheep were positively correlated to larger patches. We conclude that patch size and thus habitat carrying capacity, not population size per se, is the primary correlate to both population performance and persistence. Because habitat carrying capacity defines the upper limit to population size, clearly the amount of suitable habitat in a patch is ultimately linked to population size. Larger populations (250+ animals) were more likely to recover rapidly to their pre-epizootic survey number following an epizootic ( p = 0.019), although the proportion of the population dying in the epizootic also influenced the probability of recovery ( p = 0.001). Expensive management efforts to restore or increase bighorn sheep populations should focus on large habitat patches located ≥23 km from domestic sheep, and less effort should be expended on populations in isolated, small patches of habitat.     

Large Herbivores and the Design of Large-Scale Nature Reserves in Western Europe

Nature conservation in densely populated areas, as in western Europe, faces the continuous loss and fragmentation of suitable habitats for wild species. Current conservation measures have not been successful in countering this threat to biological diversity, so a new concept for nature conservation is required. I propose using the habitat requirements of large herbivores as a key to the design of large-scale nature reserves. Large herbivores can have a significant influence on vegetation composition and thus act as keystone species. Moreover, they require large tracts of land and can be considered an umbrella species group for the preservation of other plants and animals. I review the present status of nature reserves in the European Union concerning size distribution and the occurrence of large herbivores and their predators. I argue that habitat fragmentation and the increased management costs of maintaining seminatural areas necessitate an enlargement of existing reserves, a process-oriented management approach, and a minimization of human influence for the long-term preservation of biological diversity.     

Habitat complexity modifies post-settlement mortality and recruitment dynamics of a marine fish

For species that have an open population structure, local population size may be strongly influenced by a combination of propagule supply and post-settlement survival. While it is widely recognized that supply of larvae (or recruits) is variable and that variable recruitment may affect the relative contribution of pre- and post-settlement factors, less effort has been made to quantify how variation in the strength of post-settlement mortality (particularly density-dependent mortality) will affect the importance of processes that determine population size. In this study, I examined the effects of habitat complexity on mortality of blue rockfish (Sebastes mystinus) within nearshore reefs off central California. I first tested whether variation in habitat complexity (measured as three-dimensional complexity of rocky substrate) affected the magnitude of both density-independent and density-dependent mortality. I then used limitation analysis to quantify how variation in habitat complexity alters the relative influence of recruitment, density-independent mortality, and density-dependent mortality in determining local population size. Increased habitat complexity was associated with a reduction in both density-independent and density-dependent mortality. At low levels of habitat complexity, limitation analysis revealed that mortality was strong and recruitment had relatively little influence on population size. However, as habitat complexity increased, recruitment became more important. At the highest levels of habitat complexity, limitation by recruitment was substantial, although density-dependent mortality was ultimately the largest constraint on population size. In high-complexity habitats, population dynamics may strongly reflect variation in recruitment even though fluctuations may be dampened by density-dependent mortality. By affecting both density-independent and density-dependent mortality, variation in habitat complexity may result in qualitative changes in the dynamics of populations. These findings suggest that the relative importance of pre- vs. post-settlement factors may be determined by quantifiable habitat features, rather than ambient recruitment level alone. Because the magnitude of recruitment fluctuations can affect species coexistence and the persistence of populations, habitat-driven changes in population dynamics may have important consequences for both community structure and population viability.     

Demographic Viability of a Relict Population of the Critically Endangered Plant Borderea chouardii

Abstract:   In addition to human-caused changes in the environment, natural stochasticity may threaten species persistence, and its impact must be taken into account when priorities are established and management plans are designed. Borderea chouardii is a Tertiary relict at risk of extinction that occurs in only one location in the world, where the probability of human disturbance is low. Its persistence, therefore, is mainly linked to its response to natural threats such as stochasticity. Over 8 years I monitored up to 25% of this rupicolous small geophyte. The population had an unbalanced size structure and 90% failure in seed arrival at appropriate microhabitats, which suggests a problem with recruitment. I used matrix models to describe its population dynamics, conducted hand sowings, and performed stochastic simulations to investigate the effect of environmental stochasticity on population trend and viability. I modeled several scenarios to represent a variety of ecological situations, such as population reduction, episodic or persistent disease, and enhancement or decrease of recruitment. Population growth rate (λ) was never significantly different from unity over the study period. The risk of extinction was null over the next five centuries under current conditions. Increase of mortality and decrease of recruitment reduced stochastic population growth rate, but no factor except a persistent increase of 10% mortality resulted in extinction. These results are the consequence of the plant's extremely long life span (over 300 years) and low temporal variability of key vital rates. Even though hand sowing significantly increased the stochastic population growth rate, other approaches may be more important for the persistence of this species. The extremely slow capacity for recovery following disturbances renders habitat preservation essential. In addition, the founding of new populations would reduce the risk associated with habitat destruction.     

From population sources to sieves: the matrix alters host-parasitoid source-sink structure

Field experiments that examine the impact of immigration, emigration, or landscape structure (e.g., the composition of the matrix) on the source sink dynamics of fragmented populations are scarce. Here, planthoppers (Prokelisia crocea) and egg parasitoids (Anagrus columbi) were released among host-plant patches that varied in structural (caged, isolated, or in a network of other patches) and functional (mudflat matrix that impedes dispersal vs. brome-grass matrix that facilitates dispersal) connectivity. Planthoppers and parasitoids on caged patches exhibited density-dependent growth rates, achieved high equilibrium densities, and rarely went extinct. Therefore, experimental cordgrass patches were classified as population sources. Because access to immigrants did not result in elevated population densities, source populations were not also pseudosinks, i.e., patches whose densities occur above carrying capacity due to high immigration. Planthoppers and parasitoids in open patches in mudflat had dynamics similar to those in caged patches, but went extinct in 4-5 generations in open patches in brome. Brome-embedded patches leaked emigrants at a rate that exceeded the gains from reproduction and immigration; populations of this sort are known as population sieves. For species whose suitable patches are becoming smaller and more isolated as a result of increased habitat fragmentation, emigration losses are likely to become paramount, a condition favoring the formation of population sieves. An increase in the proportion of patches that are sieves is predicted to destabilize regional population dynamics.     

A multispecies test of source–sink indicators to prioritize habitat for declining populations

For species at risk of decline or extinction in source–sink systems, sources are an obvious target for habitat protection actions. However, the way in which source habitats are identified and prioritized can reduce the effectiveness of conservation actions. Although sources and sinks are conceptually defined using both demographic and movement criteria, simplifications are often required in systems with limited data. To assess the conservation outcomes of alternative source metrics and resulting prioritizations, we simulated population dynamics and extinction risk for 3 endangered species. Using empirically based habitat population models, we linked habitat maps with measured site‐ or habitat‐specific demographic conditions, movement abilities, and behaviors. We calculated source–sink metrics over a range of periods of data collection and prioritized consistently high‐output sources for conservation. We then tested whether prioritized patches identified the habitats that most affected persistence by removing them and measuring the population response. Conservation decisions based on different source–sink metrics and durations of data collection affected species persistence. Shorter time series obscured the ability of metrics to identify influential habitats, particularly in temporally variable and slowly declining populations. Data‐rich source–sink metrics that included both demography and movement information did not always identify the habitats with the greatest influence on extinction risk. In some declining populations, patch abundance better predicted influential habitats for short‐term regional persistence. Because source–sink metrics (i.e., births minus deaths; births and immigrations minus deaths and emigration) describe net population conditions and cancel out gross population counts, they may not adequately identify influential habitats in declining populations. For many nonequilibrium populations, new metrics that maintain the counts of individual births, deaths, and movement may provide additional insight into habitats that most influence persistence.     

Fragmentation of Phragmites Habitats, Minimum Viable Population Size, Habitat Suitability, and Local Extinction of Moths, Midges, Flies, Aphids, and Birds

Results from populations of insects and birds inhabiting Phragmites habitats were used to analyze effects of fragmentation. Flush-crash cycles of the stem-boring moth Archanara geminipuncta (Lepidoptera, Noctuidae) showed regionally concurrent, local extinctions despite an originally enormous population size (more than 180,000 adults), emphasizing the importance of metapopulation dynamics. Further, A. geminipuncta could be considered a keystone species, since shoot damage facilitated more than twenty species of herbivores, saprovores (of the caterpillars' feces), and their parasitoids. The gall midge Lasioptera arundinis could survive only in side shoots induced by shoot damage of A. geminipuncta .
Small Phragmites stands had thinner shoots (due to a water or nutrient deficiency) and shoots with more leaves (due to a better light supply) than large stands, thereby influencing species-specific demands for habitat suitability and nutritiousness of reed tissue. In other words significance of habitat fragmentation could not be assessed by area alone. For example, two chloropid flies depending on thin, yellowish shoots survived only in small habitats or in the unmown edges of large habitats.
Local persistence of Phragmites herbivores depended on much larger population sizes than could be expected from a population size sufficient to maintain genetic variation. At least 11,000 adults of the gall midge Giraudiella inclusa (or more than 84,000 galls) were necessary to avoid local extinction.
With regard to conservation management of reed habitats, nature reserves should consist of old and unmown reeds, have fewer disturbed (particularly, fewer mown) habitat edges, measure more than two hectares (priority should go to the largest remaining fragments), and be surrounded by nearby reed habitats providing reservoir populations and diverse shoot types.     

Four Factors Modifying the Effect of Competition on Carnivore Population Dynamics as Illustrated by African Wild Dogs

Abstract: Limitation of predator populations by prey availability and the effects of predators on prey populations are widely recognized as important ecological processes that affect carnivore conservation. Interspecific competition can also be a strong limiting factor for carnivore populations, and the effects of competition help explain why some carnivore species are prone to extinction. Competition among carnivores is unusual in some ways, so some predictions from traditional models of competition do not hold. For example, an increase in the density of prey can increase the effect of competition among carnivores, rather than weakening it. I used published data from African wild dogs (    Lycaon pictus ) to highlight four complexities that can modify the effects of competition on the population dynamics of carnivores: habitat fragmentation, counterintuitive effects of prey density, predator-prey size ratios, and habitat type.     

Wood Thrush Populations Are Not All Sinks in the Agricultural Midwestern United States

Abstract: Conservation strategies for forest-breeding, Neotropical migratory birds focus on the identification and preservation of source habitats. In sources, populations maintain relatively high levels of reproductive success, whereas in sinks, reproductive success is consistently below net replacement rate. In agricultural regions of the midwestern United States, forest patches of various sizes appear to act as sinks for Neotropical migrants. In 1994–1996 I monitored Wood Thrush (   Hylocichla mustelina ) nests in four forest patches in an agricultural landscape in northern Indiana. Given the prevailing estimates of adult and juvenile survival, I report productivity data that show substantial annual variation and that are sufficient to suggest the maintenance of temporal sources. Indeed, some Wood Thrush populations may act as source habitats in regions with relatively high levels of Brown-headed Cowbird (   Molothrys ater ) parasitism and nest predation because pairs produce multiple clutches and have the ability to fledge cowbirds and their own young from parasitized nests. My results and those from other studies of Wood Thrushes in landscapes with fragmented forests illustrate the variable ways in which source-sink dynamics are affected by habitat fragmentation. In some agricultural regions, populations in small forest patches may act as sources for Wood Thrushes and may be a legitimate focus of conservation efforts aimed at some migratory songbirds.     

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.     

Modeling the spatio-temporal dynamics and interactions of households,landscapes, and giant panda habitat

Human modification of land-cover has been a leading cause of floral and faunal species extirpation and loss of local and global biodiversity. As natural areas are impacted, habitat and populations can become fragmented and isolated. This is particularly evident in the mountainous areas of southwestern China that support the remaining populations of giant pandas (Ailuropoda melanoleuca). Giant panda populations have been restricted to remnants of habitat from extensive past land use and land-cover change. Households are a basic socio-economic unit that continues to impact the remaining habitat through activities such as fuelwood consumption and new household creation. Therefore, we developed a spatio-temporal model of human activities and their impacts by directly integrating households into the landscape. The integrated model allows us to examine the landscape factors influencing the spatial distribution of household activities and household impacts on habitat. As an example application, we modeled household activities in a giant panda reserve in China and examined the spatio-temporal dynamics of households, the landscape, and their mutual interactions. Human impacts are projected to result in the loss of up to 16% of all existing habitat within the reserve over the next 30 years. In addition, we found that accessibility largely controls the spatial distribution of household activities and considerable changes in management and household activities will be required to maintain the current level of habitat within the reserve.     

Modeling the Effects of Habitat Fragmentation on Source and Sink Demography of Neotropical Migrant Birds

Many songbird populations in the midwestem United States are structured as a network of sources and sinks that are linked by dispersal. We used a modeling approach to examine explicitly how populations respond to incremental fragmentation of source habitat and how this response may vary depending upon two life-history attributes: fidelity to natal habitat type and reproductive strength of the source. Fragmentation of source habitat led to a predictable decline in population for both attributes examined, but the manner in which populations declined varied depending upon the reproductive strength of the source and the level of fidelity. When the source was weak and produced few excess individuals, fragmentation of source habitats resulted in a predictable and parallel population decline of adults in both the source and the sink. In this situation high fidelity to natal habitats was important for maintenance of population size and structure. Low fidelity to weak sources resulted in population extinction; populations experienced a demographic cost by dispersing from high quality source habitat to low quality sink habitat. In contrast, when the source was strong and produced many excess individuals, fragmentation of the source led to population declines in both the source and the sink, but this decline was more abrupt in sink habitats. When the source was strong and produced a large excess of individuals, nonfidelity to natal habitats had little effect on metapopulation size and structure.     

Resource distributions among habitats determine solitary bee offspring production in a mosaic landscape.

Within mosaic landscapes, many organisms depend on attributes of the environment that operate over scales ranging from a single habitat patch to the entire landscape. One such attribute is resource distribution. Organisms' reliance on resources from within a local patch vs. those found among habitats throughout the landscape will depend on local habitat quality, patch quality, and landscape composition. The ability of individuals to move among complementary habitat types to obtain various resources may be a critical mechanism underlying the dynamics of animal populations and ultimately the level of biodiversity at different spatial scales. We examined the effects that local habitat type and landscape composition had on offspring production and survival of the solitary bee Osmia lignaria in an agri-natural landscape in California (U.S.A.). Female bees were placed on farms that did not use pesticides (organic farms), on farms that did use pesticides (conventional farms), or in seminatural riparian habitats. We identified pollens collected by bees nesting in different habitat types and matched these to pollens of flowering plants from throughout the landscape. These data enabled us to determine the importance of different plant species and habitat types in providing food for offspring, and how this importance changed with landscape and local nesting-site characteristics. We found that increasing isolation from natural habitat significantly decreased offspring production and survival for bees nesting at conventional farms, had weaker effects on bees in patches of seminatural habitat, and had little impact on those at organic farm sites. Pollen sampled from nests showed that females nesting in both farm and seminatural habitats relied on pollen from principally native plant species growing in seminatural habitat. Thus connectivity among habitats was critical for offspring production. Females nesting on organic farms were buffered to isolation effects by switching to floral resources growing at the farm site when seminatural areas were too distant. Overall local habitat conditions (farm management practices) can help bolster pollinators, but maintaining functional connectivity among habitats will likely be critical for persistence of pollinator populations as natural habitats are increasingly fragmented by human activities.     

Simulated Responses of a Forest-Interior Bird Population to Forest Management Options in Central Hardwood Forests of the United States

I used estimates of carrying capacity, survival, fecundity, and edge effects to simulate the responses of a forest-interior bird population to selection cutting clearcutting, and no timber harvest. I also modeled population sensitivity to changes in fecundity, survival, K , and edge relationships. Because model parameters are based on scant data, results should he regarded as hypotheses to be further investigated or measures of the relative impact or sensitivity (given model assumptions). Simulated population size was greater with no timber harvest than with clearcutting and greater with clearcutting than with group selection when edge effects were included in the model. Without edge effects, population levels were only slightly lower under group selection than under no timber harvest, and greater than clearcutting. Edge effects had only a small impact on population levels under clearcutting. Clearcut size did not have much effect on population levels, but longer and shorter rotation ages resulted in higher and lower population levels, respectively. The model was very sensitive to declines in mean fecundity and survival, suggesting that factors affecting mean demographic rates could be more important than local edge effects. Some methods of timber harvest may be compatible with the conservation of forest-interior birds, but better demographic data and information on habitat suitability of selectively cut forests and young even-aged stands is needed to adequately evaluate management options.     

Limitations of Captive Breeding in Endangered Species Recovery

The use of captive breeding in species recovery has grown enormously in recent years, but without a concurrent growth in appreciation of its limitations. Problems with (1) establishing self-sufficient captive populations, (2) poor success in reintroductions, (3) high costs, (4) domestication, (5) preemption of other recovery techniques, (6) disease outbreaks, and (7) maintaining administrative continuity have all been significant. The technique has often been invoked prematurely and should not normally be employed before a careful field evaluation of costs and benefits of all conservation alternatives has been accomplished and a determination made that captive breeding is essential for species survival. Merely demonstrating that a species' population is declining or has fallen below what may be a minimum viable size does not constitute enough analysis to justify captive breeding as a recovery measure. Captive breeding should be viewed as a last resort in species recovery and not a prophylactic or long-term solution because of the inexorable genetic and phenotypic changes that occur in captive environments. Captive breeding can play a crucial role in recovery of some species for which effective alternatives are unavailable in the short term. However, it should not displace habitat and ecosystem protection nor should it be invoked in the absence of comprehensive efforts to maintain or restore populations in wild habitats. Zoological institutions with captive breeding programs should operate under carefully defined conditions of disease prevention and genetic/behavioral management. More important, these institutions should help preserve biodiversity through their capacities for public education, professional training, research, and support of in situ conservation efforts.