Using Age Structure to Detect Impacts on Threatened Populations: a Case Study with Steller Sea Lions

Abstract:   A delayed response to change is often a characteristic of long-lived species and presents a major challenge to monitoring their status. However, rapid shifts in age structure can occur even while population size remains relatively static. We used time-varying matrix models to study age-structure information as a tool for improving detection of survivorship and fecundity change and status. We applied the methods to Steller sea lions (  Eumetopias jubatus ), a long-lived endangered marine mammal found throughout the North Pacific Rim. Population and newborn counts were supplemented with information on the fraction of the population that was juvenile, obtained by measuring animals in aerial photographs taken during range-wide censuses. By fitting the model to 1976–1998 data, we obtained maximum-likelihood estimates and 95% confidence intervals for juvenile survivorship, adult survivorship, and adult fecundity in the mid-1980s, late 1980s, and 1990s. We used a series of nested models to test whether the data were best fit by a model with one, two, or three temporal changes in demographic rates, and we fit the models to different lengths of data to test the number of years of data needed to detect a demographic change. The declines in the early 1980s were associated with severely low juvenile survivorship, whereas declines in the 1990s were associated with disproportionately low fecundity. We repeated these analyses, fitting only to the count data without the juvenile-fraction information, to determine whether the age-structure information changed the conclusions and/or changed the certainty and speed with which demographic-rate changes could be detected. The juvenile-fraction data substantially improved the degree to which estimates from the model were consistent with field data and significantly improved the speed and certainty with which changes in demographic rates were detected.     

Group size in wedge-capped capuchin monkeys Cebus olivaceus and the reproductive success of males and females

Summary The effect of variation in group size on age-specific survivorship and fecundity rates were examined in a population of wedge-capped capuchin monkeys Cebus olivaceus during a 10 year study. Life tables were constructed separately for four large (15 individuals) and four small groups (<15 individuals). Female reproductive success, and its relative contribution to population growth, was much higher in large groups, primarily through higher age-specific fecundity. Age-specific survivorship was similar in groups of different sizes. The reproductive success of the single breeding male in a group was much higher in large than small groups. Compared to small groups, breeding males in large groups had a longer breeding tenure, and access to greater numbers of reproductive females with a higher average fecundity. Differences in female reproductive success apparently resulted from variation in access to monopolizable fruit trees. Large groups predictably displaced small groups during intergroup encounters. Group rank depended on the number of males resident in groups. The large number of non-breeding males in large groups results from their longer average residency time. I explain the longer residency of males in large groups by the higher average reproductive success of breeding males in these groups.     

Generation time and the maximum growth rate for populations with age-specific fecundities and unknown juvenile survival

In age-classified population models where all parameters are known, the generation time and growth rate are calculated in a straightforward manner. For many populations, some parameters, such as juvenile survival, are difficult to estimate accurately. In a simplified population model where fecundity and survival are constant from the onset of breeding, it is known that generation time may be calculated given only adult survival, age at first reproduction, and the population growth rate. However, the assumption of constant fecundity from the onset of breeding does not hold for many populations. An extended population model allows calculation of generation time with the additional knowledge of the ratio of age-specific fecundities compared to a maximum fecundity rate. When these relative fecundities are unknown, an ad hoc adjustment to the simplified model performs well.When the study population is in an ideal environment, the optimal generation time and maximum growth rate are linked, and both may be approximated knowing only adult survival, age at first reproduction, and the relative fecundities. The maximum growth rate has important conservation implications, and calculating it correctly is therefore important. Improper use of the simplified population model to calculate the maximum growth rate, combined with a simple decision rule, leads to an average overharvest of 36%, and >60% for three of six bird species studied, compared to the full population model. By comparison, using the approximation from the extended or adjusted models results in average overharvests of only 8% (extended model) and 5% (adjusted model), and <50% for all six species (either model).     

Intergenerational effects of climate generate cohort variation in lizard reproductive performance

An evaluation of the link between climate and population dynamics requires understanding of climate effects both within and across generations. In ectothermic vertebrates, demographic responses to climate changes should crucially depend on balancing needs for heat and water. Here, we studied how temperature and rainfall regimes experienced before and during adulthood influenced reproductive performances (litter size, offspring size, and survival) in a natural population of the live-bearing common lizard, Lacerta vivipara, monitored continuously from 1989 to 2004. Rainfall regime, but not temperature, had both immediate and delayed effects on these reproductive performances. Rainfall during the first month of life was positively correlated with juvenile survival. Females experiencing more rainfall during gestation produced smaller neonates that showed greater survival when controlling for the positive effect of body size on survival. Furthermore, females that experienced heavier rainfall when in utero produced fewer but longer neonates during adulthood. These demographic effects of rainfall on adult reproductive traits may come from maternal effects of climate conditions and/or from delayed effects of rainfall on the environment experienced early in life. Irrespective of the precise mechanism, however, this study provides evidence of intergenerational climate effects in natural populations of an ectothermic vertebrate.     

Cost‐Effectiveness of Translocation Options for a Threatened Waterbird

Abstract: Reintroduction of captive‐reared animals has become increasingly popular in recent decades as a conservation technique, but little is known of how demographic factors affect the success of reintroductions. We believe whether the increase in population persistence associated with reintroduction is sufficient to warrant the cost of rearing and relocating individuals should be considered as well. We examined the trade‐off between population persistence and financial cost of a reintroduction program for Crested Coots (Fulica cristata). This species was nearly extirpated from southern Europe due to unsustainable levels of hunting and reduction in amount and quality of habitat. We used a stochastic, stage‐based, single‐sex, metapopulation model with site‐specific parameters to examine the demographic effects of releasing juveniles or adults in each population for a range of durations. We parameterized the model with data from an unsuccessful reintroduction program in which juvenile captive‐bred Crested Coots were released between 2000 and 2009. Using economic data from the captive‐breeding program, we also determined whether the strategy that maximized abundance coincided with the least expensive strategy. Releasing adults resulted in slightly larger final abundance than the release of nonreproductive juveniles. Both strategies were equally poor in achieving a viable metapopulation, but releasing adults was 2–4 times more expensive than releasing juveniles. To obtain a metapopulation that would be viable for 30 years, fecundity in the wild would need to increase to the values observed in captivity and juvenile survival would need to increase to almost unity. We suggest that the most likely way to increase these vital rates is by increasing habitat quality at release sites.     

Life history consequences of variation in age at primiparity in northern elephant seals

Summary The age when female northern elephant seals, Mirounga angustirostris, bear their first young varies from 2 to 6 years. At Año Nuevo, California, a group of 77 females, primiparous at age 3, had a lower survivorship rate to each successive year up to age 8 than a group of 98 females that deferred initial pupping until age 4. The difference in survivorship appears to be due to the greater relative energetic costs of gestation and lactation incurred by the earlier breeding females during a period in their development when growth is rapid. An alternate hypothesis for the difference in survivorship — that young primiparous females are in poor condition from birth-is untenable; females that pupped early in life were larger at weaning age (a correlate of condition) than females that were primiparous 1 year later.Models based on the data show that differential survival of seals that vary in age at primiparity has important consequences for population growth and life history strategies. The effect of age at primiparity on the rate of increase of populations varies with colony density and juvenile survivorship. The optimal life history strategy for female elephant seals under most conditions existing today, including those at Ano Nuevo during the study period, is to bear the first offspring at age 4. Primiparity at age 3 is projected to be favored when harem density is very low and weaning success and juvenile survivorship are high; postponement of first breeding to age 5 is expected at high harem densities with intense competition for breeding space. Offprint requests to: B.J. Le Boeuf     

Recovery of the Kakerori: An Endangered Forest Bird of the Cook Islands

The Kakerori, or Rarotonga Flycatcher (Pomarea dimidiata) , is an endangered monarch flycatcher endemic to the island of Rarotonga, Cook Islands. This bird was reported to be common until the middle of the nineteenth century, but it declined before 1885 and was thought to be extinct early this century. A small population persisted in the rugged interior of Rarotonga; in 1987 the population stood at 38 birds but was in decline. We determined that introduced predators, especially ship rats (Rattus rattus) , were affecting breeding success. Through a program of experimental management, aimed mainly at rodent control, the population increased from a low of 29 birds in Spring 1989 to 60 birds in Spring 1993. The recovery can be attributed to both improved breeding success and increased adult survivorship.     

Effects of Multiple Levels of Social Organization on Survival and Abundance

Abstract: Identifying how social organization shapes individual behavior, survival, and fecundity of animals that live in groups can inform conservation efforts and improve forecasts of population abundance, even when the mechanism responsible for group‐level differences is unknown. We constructed a hierarchical Bayesian model to quantify the relative variability in survival rates among different levels of social organization (matrilines and pods) of an endangered population of killer whales (Orcinus orca). Individual killer whales often participate in group activities such as prey sharing and cooperative hunting. The estimated age‐specific survival probabilities and survivorship curves differed considerably among pods and to a lesser extent among matrilines (within pods). Across all pods, males had lower life expectancy than females. Differences in survival between pods may be caused by a combination of factors that vary across the population's range, including reduced prey availability, contaminants in prey, and human activity. Our modeling approach could be applied to demographic rates for other species and for parameters other than survival, including reproduction, prey selection, movement, and detection probabilities.     

Age-structured modeling reveals long-term declines in the natality of western Steller sea lions.

Since the mid-1970s, the western Steller sea lion (Eumetopias jubatus), inhabiting Alaskan waters from Prince William Sound west through the Aleutian Islands, has declined by over 80%. Changing oceanographic conditions, competition from fishing operations, direct human-related mortality, and predators have been suggested as factors driving the decline, but the indirect and interactive nature of their effects on sea lions have made it difficult to attribute changes in abundance to specific factors. In part, this is because only changes in abundance, not changes in vital rates, are known. To determine how vital rates of the western Steller sea lion have changed during its 28-year decline, we first estimated the changes in Steller sea lion age structure using measurements of animals in aerial photographs taken during population surveys since 1985 in the central Gulf of Alaska (CGOA). We then fit an age-structured model with temporally varying vital rates to the age-structure data and to total population and pup counts. The model fits indicate that birth rate in the CGOA steadily declined from 1976 to 2004. Over the same period, survivorship first dropped severely in the early 1980s, when the population collapsed, and then survivorship steadily recovered. The best-fitting model indicates that in 2004, the birth rate in the central Gulf of Alaska was 36% lower than in the 1970s, while adult and juvenile survivorship were close to or slightly above 1970s levels. These predictions and other model predictions concerning population structure match independent field data from mark-recapture studies and photometric analyses. The dominant eigenvalue for the estimated 2004 Leslie matrix is 1.0014, indicating a stable population. The stability, however, depends on very high adult survival, and the shift in vital rates results in a population that is more sensitive to changes in adult survivorship. Although our modeling analysis focused exclusively on the central Gulf of Alaska, the western Gulf of Alaska and eastern Aleutians show a similar pattern of declining pup fraction with no increase in the juvenile, or pre-breeding, fraction. This suggests that declining birth rate may be a problem for western Steller sea lions across the Gulf of Alaska and into the Aleutian Islands.     

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.     

Space use,longevity, and reproductive success in meadow voles

Summary We addressed the question of how reproductive success (RS) was limited in the shortlived but highly fecund meadow vole, Microtus pennsylvanicus. In so doing, we asked how differential space use patterns could affect longevity and hence RS in each sex. The sample comprised all voles achieving sexual competency over the course of a 40-week breeding season in a live-trapped population in Manomet, MA USA. Matrilineal families were determined using a radionuclide labelling technique; paternity was estimated using a maximum likelihood model. Individual RS was defined as the number of offspring successfully recruited into the trappable population per adult. We found that the variance in RS among female meadow voles was greater than the variance among males. In an attempt to explain this pattern, reproductively successful individuals were compared to reproductively unsuccessful individuals with regard to survivorship, maximum body weight achieved, and spatial mobility. The only difference between fathers and reproductively unsuccessful males was that fathers were heavier. In contrast, mothers differed from unsuccessful females in every measurement. Females lived longer than males, and mothers lived longer than either fathers or reproductively unsuccessful females. The observed differences in longevity may have been largely the result of differences in levels of mobility, assuming more mobile voles were more susceptible to predation. Mothers were significantly more site tenacious than were either males or unsuccessful females. These patterns explain the distribution of RS in our population if predation differentially affects male and female meadow voles. The meadow vole is the only non-polyandrous vertebrate reported to date in which the variance in RS among females exceeds the variance in RS among males.     

Effects of age and sex ratios on offspring recruitment rates in translocated black rhinoceros

Success of animal translocations depends on improving postrelease demographic rates toward establishment and subsequent growth of released populations. Short‐term metrics for evaluating translocation success and its drivers, like postrelease survival and fecundity, are unlikely to represent longer‐term outcomes. We used information theory to investigate 25 years of data on black rhinoceros (Diceros bicornis) translocations. We used the offspring recruitment rate (ORR) of translocated females—a metric integrating survival, fecundity, and offspring recruitment at sexual maturity—to detect determinants of success. Our unambiguously best model (AICω = 0.986) predicted that ORR increases with female age at release as a function of lower postrelease adult rhinoceros sex ratio (males:females). Delay of first postrelease reproduction and failure of some females to recruit any calves to sexual maturity most influenced the pattern of ORRs, and the leading causes of recruitment failure were postrelease female death (23% of all females) and failure to calve (24% of surviving females). We recommend translocating older females (≥6 years old) because they do not exhibit the reproductive delay and low ORRs of juveniles (<4 years old) or the higher rates of recruitment failure of juveniles and young adults (4–5.9 years old). Where translocation of juveniles is necessary, they should be released into female‐biased populations, where they have higher ORRs. Our study offers the unique advantage of a long‐term analysis across a large number of replicate populations—a science‐by‐management experiment as a proxy for a manipulative experiment, and a rare opportunity, particularly for a large, critically endangered taxon such as the black rhinoceros. Our findings differ from previous recommendations, reinforce the importance of long‐term data sets and comprehensive metrics of translocation success, and suggest attention be shifted from ecological to social constraints on population growth and species recovery, particularly when translocating species with polygynous breeding systems.     

Use of fecundity measured directly throughout the breeding season to test a source-sink demographic model

Populations of landbirds (bird species that occupy terrestrial habitats for most of their life cycle) are declining throughout North America (north of Mexico) and Europe, yet little is known about how demography is driving this trend. A recent model of 5 geographically separated populations of Cerulean Warblers (Dendroica cerulea) that was based on within-season sampling of nest survival and fledgling success shows that all populations are sinks (annual reproduction is consistently less than annual adult mortality). I tested this indirect model by directly measuring fecundity (number of female fledglings/female) during the breeding season for 2 years in a Cerulean Warbler population occupying a mature forest in southwestern Michigan (U.S.A.) I determined territories of male birds on the basis of male plumage characters and phases of the nesting cycle (2007) and on uniquely color-banded males (2008). I transferred locations of identified males to topographic maps. I counted all fledglings in territories from May to July each year. The model I tested may apply only to single-brooded species; therefore, I searched the literature to estimate the percentage of single-brooded species in North America. The breeding season of Cerulean Warblers was short- nearly all nests were initiated from mid-May to late June. Nest predation and brood parasitism were primary and rare causes of nest failure, respectively. Significantly fewer Cerulean Warblers fledged from parasitized than from nonparasitized nests. Fledgling survival required to maintain the population size was well above previously published values for Neotropical migrants. Single-brooded species comprise 62% of North American breeding bird species for which the number of broods per year is known; I believe my results may apply to these species. The consistency between identification of populations as sources or sinks on the basis of either model estimates or direct measurements suggests that a demographic model relying on within-season sampling of fecundity is adequate to determine population status of single-brooded avian populations. In addition, on the basis of results of previous studies, annual adult survival rate of the Cerulean Warbler is typical of parulid warblers that are not declining. Thus, low fecundity, here determined with different quantitative methods, can drive status of landbird species with high-observed survival.     

Using Modeling to Improve Monitoring of Structured Populations: Are We Collecting the Right Data?

Abstract:  Population monitoring is central to most demographic studies and conservation efforts, but it may not always be directed at the most appropriate life stage. We used stochastic simulation modeling to evaluate the effectiveness of a monitoring program for a well-studied population of Eastern Imperial Eagles ( Aquila heliaca ) in Kazakhstan. Specifically, we asked whether the most appropriate data were being collected to understand system state and population dynamics. Our models were parameterized with data collected over the course of 25 years of study of this population. We used the models to conduct simulation experiments to evaluate relationships between monitored or potentially monitored parameters and the demographic variables of interest—population size ( N ) and population growth (λ). Static analyses showed that traditional territory-based monitoring was a poor indicator of eagle population size and growth and that monitoring survivorship would provide more information about these parameters. Nevertheless, these same traditionally monitored territory-based parameters had greater power to detect long-term changes in population size than did survivorship or population structure. Regardless of the taxa considered, threats can have immediate impacts on population size and growth or longer-term impacts on population dynamics. Prudently designed monitoring programs for any species will detect the demographic effects of both types of threats.     

Cooperative breeding by the Galápagos mockingbird,Nesomimus parvulus

Summary The costs and benefits of helping behavior were analyzed for 36 pairs of the Galápagos mockingbird, Nesomimus parvulus, and their associates. Helping at the nest is usually done by sons or males suspected to be offspring of the breeders. Costs and benefits to breeders were assessed by comparison of pairs with and without helpers, and costs and benefits to helpers were assessed by comparison of birds which help and those which establish themselves as novice breeders.Helping behavior benefits breeders by increasing fledging success and by reducing the adult energy load in territory defense and feeding of nestlings. Breeders assisted by helpers may also benefit by decreased nest predation. Helpers enhance their inclusive fitness by helping, and gain directly by increasing their chances of securing a territory. Helpers do not appear to gain any fitness advantage from the experience of assisting, nor do they increase their survivorship by remaining on natal territory.Ecological and demographic features such as saturated territories and low territory turn-over rates due to high adult survival may be primarily responsible for the evolution of the helping behavior, with kinselection reinforcing it. Associated features of this system are a male-biased population sex ratio, a greater energetic benefit to breeding males than to breeding females in having helpers, earlier dispersal and breeding by females than by males, and much more frequent helping by males than by females. These are interpreted as consequences of brothersister aggression that indirectly minimizes the chances of inbreeding.     

Management and Recovery Options for Ural River Beluga Sturgeon

Abstract: Management of declining fisheries of anadromous species sometimes relies heavily on supplementation of populations with captive breeding, despite evidence that captive breeding can have negative consequences and may not address the root cause of decline. The beluga sturgeon (Huso huso), a species threatened by the market for black caviar and reductions in habitat quality, is managed through harvest control and hatchery supplementation, with an emphasis on the latter. We used yield per recruit and elasticity analyses to evaluate the population status and current levels of fishing and to identify the life‐history stages that are the best targets for conservation of beluga of the Ural River. Harvest rates in recent years were four to five times higher than rates that would sustain population abundance. Sustainable rates of fishing mortality are similar to those for other long‐lived marine species such as sharks and mammals. Yield per recruit, which is maximized if fish are first harvested at age 31 years, would be greatly enhanced by raising minimum size limits or reducing illegal take of subadults. Improving the survival of subadult and adult females would increase population productivity by 10 times that achieved by improving fecundity and survival from egg to age 1 year (i.e., hatchery supplementation). These results suggest that reducing mortality of subadults and adult wild fish is a more effective conservation strategy than hatchery supplementation. Because genetics is not factored into hatchery management practices, supplementation may even reduce the viability of the beluga sturgeon.     

Aspects of population ecology in two populations of fiddler crabs, <Emphasis Type="Italic">Uca pugnax</Emphasis>

Fiddler crabs, Uca pugnax, were collected from a highly contaminated site and a relatively clean site, both in New Jersey to determine if and how environments with varying levels of pollutants may impact aspects of population biology including individual size, morphology (major cheliped size), population density, fecundity, recruitment and survivorship of early benthic phases. Crabs from the highly contaminated site were significantly larger in size, but had lower population density, lower recruitment, reduced reproductive season and lower survivorship of early benthic phases. Our study suggests that contamination may play a role in population ecology of U. pugnax. This study also determined that the reproductive season for U. pugnax in New Jersey is much longer than reported in the literature and could potentially be impacted by global climate change.     

The Role of Translocation in Recovery of Woodland Caribou Populations

Abstract: Maintenance of viable populations of many endangered species will require conservation action in perpetuity. Efforts to conserve these species are more likely to be successful if their reliance on conservation actions is assessed at the population level. Woodland caribou (Rangifer tarandus caribou) were extirpated recently from Banff National Park, Canada, and translocations of caribou to Banff and neighboring Jasper National Park are being considered. We used population viability analysis to assess the relative need for and benefits from translocation of individuals among caribou populations. We measured stochastic growth rates and the probability of quasi extinction of four populations of woodland caribou with and without translocation. We used two vital rates in our analysis: mean adult female survival and mean number of calves per breeding‐age female as estimates of mean fecundity. We isolated process variance for each vital rate. Our results suggested the Tonquin caribou population in Jasper is likely to remain viable without translocation, but that translocation is probably insufficient to prevent eventual extirpation of the two other populations in Jasper. Simulated reintroductions of caribou into Banff resulted in a 53–98% probability of >8 females remaining after 20 years, which suggests translocation may be an effective recovery tool for some caribou populations.     

Phenotypic Alterations, Evolutionarily Significant Structures, and Rhino Conservation

To slow the impending loss of wild black ( Diceros bicornis ) and white ( Ceratotherium simum ) rhinos, three African countries have resorted to dehorning, a practice designed to remove the incentive for poachers to kill the hornless animals. The efficacy of this controversial conservation action remains unknown, in part because much uncertainty exists about the functional significance of rhino horns. We assessed the current utility of horns in Namibian black rhinos from phenotypically altered and intact populations in the Namib Desert, and we collated data on mortal fighting among horned females living in Etosha National Park. Infant mortality was 100% when dehorned mothers were sympatric with spotted hyenas ( Crocuta crocuta ). In contrast, infant survival was 100% for both horned mothers living with hyenas and occasional lions ( Panthera leo ) and 100% for dehorned mothers in the absence of dangerous carnivores. These data suggest that female horns can have direct fitness benefits in terms of calf survival. However, because lethal wounding due to fighting may account for up to 33% of the mortality of horned females, dehorning may improve adult survivorship. Our results (1) suggest that, where the aim of conservation programs is to improve population viability through juvenile recruitment, dehorning is unlikely to be a prudent strategy if practiced in areas with dangerous predators, and (2) illustrate the value of experimental approaches to onerous problems in conservation.     

Potential Effects of a Forest Management Plan on Bachman's Sparrows (Aimophila aestivalis): Linking a Spatially Explicit Model with GIS

By combining a spatially explicit, individual-based population simulation model with a geographic information system, we have simulated the potential effects of a U.S. Forest Service management plan on the population dynamics of Bachman's Sparrow ( Aimophila aestivalis ) at the Savannah River Site, a U.S. Department of Energy facility in South Carolina. Although the Forest Service's management plan explicitly sets management goals for many species, most of the prescribed management strategy deals with the endangered Red-cockaded Woodpecker ( Picoides borealis ) because of legal requirements. We explored how a species (the sparrow) that is not the target of specific management strategies but that shares some habitat requirements with the woodpecker, would fare under the management plan. We found that the major components of the proposed management plan may allow the sparrow population to reach and exceed the minimum management goal set for this species, but only after a substantial initial decline in sparrow numbers and a prolonged transition period. In the model, the sparrow population dynamics were most sensitive to demographic variables such as adult and juvenile survivorship and to landscape variables such as the suitability of young clearcuts and mature pine stands. Using various assumptions about habitat suitability, we estimated that the 50-year probability of population extinction is at least 5% or may be much higher if juvenile survivorship is low. We believe, however, that modest changes in the management plan might greatly increase the sparrow population and presumably decrease the probability of extinction. Our results suggest that management plans focusing on one or a few endangered species may potentially threaten other species of management concern. Spatially explicit population models are a useful tool in designing modifications of management plans that can reduce the impact on nontarget species of management concern.