Height-growth response to climatic changes differs among populations of Douglas-fir: a novel analysis of historic data

Projected climate change will affect existing forests, as substantial changes are predicted to occur during their life spans. Species that have ample intraspecific genetic differentiation, such as Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), are expected to display population-specific growth responses to climate change. Using a mixed-effects modeling approach, we describe three-year height (HT) growth response to changes in climate of interior Douglas-fir populations. We incorporate climate information at the population level, yielding a model that is specific to both species and population. We use data from provenance tests from previous studies that comprised 236 populations from Idaho, Montana, and eastern Washington, USA. The most sensitive indicator of climate was the mean temperature of the coldest month. Population maximum HT and HT growth response to changes in climate were dependent on seed source climate. All populations had optimum HT growth when transferred to climates with warmer winters; those originating in sites with the warmest winters were taller across sites and had highest HT growth at transfer distances closest to zero; those from colder climates were shortest and had optimum HT growth when transferred the farthest. Although this differential response damped the height growth differences among populations, cold-climate populations still achieved their maximum growth at lower temperatures than warm-climate populations. The results highlight the relevance of understanding climate change impacts at the population level, particularly in a species with ample genetic variation among populations.     

"Mobbing" in Hawaiian Monk Seals (Monachus schauinslani): The Value of Simulation Modeling in the Absence of Apparently Crucial Data

Several small populations of Hawaiian monk seals ( Monachus schauinslani ) exhibit male-biased adult sex ratios and "mobbing," an aggressive behavior in which adult males injure and often kill adult females and immature seals of both sexes during mating attempts. Mobbing appears to be limiting the growth of some populations of this endangered species. The frequency of mobbing deaths appears to increase as a population's sex ratio becomes increasingly male-biased, although the exact relationship between these two variables (the mobbing response) is unknown. We developed a stochastic demographic model of a small Hawaiian monk seal population using several different assumptions about the mobbing response. We used the model to explore the origins of male-biased sex ratios in monk seal populations and to determine whether it was possible, given the lack of data on the mobbing response, to evaluate the probable effects of alternative management strategies to address the mobbing problem. Small populations (100 to 200 seals) and those with slower growth rates were more likely to develop male-biased adult sex ratios. Almost all of our modeling scenarios supported the immediate removal of males from populations where mobbing occurs. Our conclusions were relatively unaffected when the assumptions regarding the mobbing response were varied. Thus, a model was helpful even when apparently crucial data were unavailable.     

Estimating age from recapture data: integrating incremental growth measures with ancillary data to infer age-at-length

Estimating the age of individuals in wild populations can be of fundamental importance for answering ecological questions, modeling population demographics, and managing exploited or threatened species. Significant effort has been devoted to determining age through the use of growth annuli, secondary physical characteristics related to age, and growth models. Many species, however, either do not exhibit physical characteristics useful for independent age validation or are too rare to justify sacrificing a large number of individuals to establish the relationship between size and age. Length-at-age models are well represented in the fisheries and other wildlife management literature. Many of these models overlook variation in growth rates of individuals and consider growth parameters as population parameters. More recent models have taken advantage of hierarchical structuring of parameters and Bayesian inference methods to allow for variation among individuals as functions of environmental covariates or individual-specific random effects. Here, we describe hierarchical models in which growth curves vary as individual-specific stochastic processes, and we show how these models can be fit using capture-recapture data for animals of unknown age along with data for animals of known age. We combine these independent data sources in a Bayesian analysis, distinguishing natural variation (among and within individuals) from measurement error. We illustrate using data for African dwarf crocodiles, comparing von Bertalanffy and logistic growth models. The analysis provides the means of predicting crocodile age, given a single measurement of head length. The von Bertalanffy was much better supported than the logistic growth model and predicted that dwarf crocodiles grow from 19.4 cm total length at birth to 32.9 cm in the first year and 45.3 cm by the end of their second year. Based on the minimum size of females observed with hatchlings, reproductive maturity was estimated to be at nine years. These size benchmarks are believed to represent thresholds for important demographic parameters; improved estimates of age, therefore, will increase the precision of population projection models. The modeling approach that we present can be applied to other species and offers significant advantages when multiple sources of data are available and traditional aging techniques are not practical.     

Resource allocation and growth rates in the sea urchin Evechinus chloroticus (Echinoidea: Echinometridae)

The morphometry of the sea urchin Evechinus chloroticus from habitats of contrasting algal abundance but of similar urchin density was examined at two localities in southern New Zealand during 1993. Populations from habitats of high algal abundance (Dusky Sound) had similar relationships of demipyramid (jaws) to test diameter, but individuals had significantly smaller jaws relative to their test diameter than those from a locality where algal abundance was low (Arapawa Island). The body wall mass (in relation to total wet weight) was similar for populations from both localities but, for Dusky Sound populations, individuals from exposed sites had greater relative body mass than those from sheltered sites. The ratios of gonad weight:total weight were similar between localities. However, E. chloroticus from Arapawa Island reached reproductive maturity at a smaller size than those from Dusky Sound. Growth rates of E. chloroticus varied among localities in Dusky Sound. Growth was modelled by the Tanaka function, which allows for rapid early growth until reproductive maturity is reached and declining growth rates thereafter. The results show that sea urchins respond to low resource availability by increasing the size of the food-gathering apparatus, maturing at a smaller size, and growing to a smaller size than individuals from food-rich habitats. Received: 3 December 1996 / Accepted: 29 January 1997     

The effect of summer drought on the predictability of local extinctions in a butterfly metapopulation

The ecological impacts of extreme climatic events on population dynamics and community composition are profound and predominantly negative. Using extensive data of an ecological model system, we tested whether predictions from ecological models remain robust when environmental conditions are outside the bounds of observation. We observed a 10-fold demographic decline of the Glanville fritillary butterfly (Melitaea cinxia) metapopulation on the Åland islands, Finland in the summer of 2018 and used climatic and satellite data to demonstrate that this year was an anomaly with low climatic water balance values and low vegetation productivity indices across Åland. Population growth rates were strongly associated with spatiotemporal variation in climatic water balance. Covariates shown previously to affect the extinction probability of local populations in this metapopulation were less informative when populations were exposed to severe drought during the summer months. Our results highlight the unpredictable responses of natural populations to extreme climatic events.     

The length of growing season and adult sex ratio affect sexual size dimorphism in moose

While factors affecting body growth have been extensively studied, very little is known about the factors likely to affect the sexual size dimorphism (SSD) in polygynous mammals. Based on the carcass mass of 24420 male and female moose recorded in 14 Norwegian populations, we examine three hypotheses to explain geographical variation in SSD. First, SSD is expected to decrease when the relative density of animals (for a given habitat quality) increases, because resource limitation at high population densities is assumed to affect body growth of males more than females. Second, because males are selected to invest in growth more than females, environmental seasonality and related improvement of the forage quality during the short and intense growing season are expected to increase SSD. Third, by decreasing the proportion of adult males in the population, resulting in start of rutting earlier in life, hunting may decrease the SSD by increasing the reproductive cost of young males. We found that males grew faster and for a longer time of their life than did females and thus were heavier (-24%) when they reached adulthood. Sexual size dimorphism was independent of density but was higher in areas with short growing seasons. The low SSD in populations with largely adult female-biased sex ratios (males per female) shows that male body growth decreases with a decreasing proportion of adult males in the population. Our results indicate that geographical variation in moose SSD is influenced by divergent responses in the sexes to ecological factors affecting body growth.     

Patterns of Genetic Diversity and Its Loss in Mammalian Populations

Abstract:  Policy aimed at conserving biodiversity has focused on species diversity. Loss of genetic diversity, however, can affect population persistence, evolutionary potential, and individual fitness. Although mammals are a well-studied taxonomic group, a comprehensive assessment of mammalian genetic diversity based on modern molecular markers is lacking. We examined published microsatellite data from populations of 108 mammalian species to evaluate background patterns of genetic variability across taxa and body masses. We tested for loss of genetic diversity at the population level by asking whether populations that experienced demographic threats exhibited lower levels of genetic diversity. We also evaluated the effect of ascertainment bias (a reduction in variability when microsatellite primers are transferred across species) on our assessment of genetic diversity. Heterozygosity did not vary with body mass across species ranging in size from shrews to whales. Differences across taxonomic groupings were noted at the highest level, between populations of marsupial and placental mammals. We documented consistently lower heterozygosity, however, in populations that had experienced demographic threats across a wide range of mammalian species. We also documented a significant ( p = 0.01) reduction in heterozygosity as a result of ascertainment bias. Our results suggest that populations of both rare and common mammals are currently losing genetic diversity and that conservation efforts focused above the population level may fail to protect the breadth of persisting genetic diversity. Conservation policy makers may need to focus their efforts below the species level to stem further losses of genetic resources.     

Assessment of mammal reproduction for hunting sustainability through community‐based sampling of species in the wild

Wildlife subsistence hunting is a major source of protein for tropical rural populations and a prominent conservation issue. The intrinsic rate of natural increase. (r_max) of populations is a key reproductive parameter in the most used assessments of hunting sustainability. However, researchers face severe difficulties in obtaining reproductive data in the wild, so these assessments often rely on classic reproductive rates calculated mostly from studies of captive animals conducted 30 years ago. The result is a flaw in almost 50% of studies, which hampers management decision making. We conducted a 15‐year study in the Amazon in which we used reproductive data from the genitalia of 950 hunted female mammals. Genitalia were collected by local hunters. We examined tissue from these samples to estimate birthrates for wild populations of the 10 most hunted mammals. We compared our estimates with classic measures and considered the utility of the use of r_max in sustainability assessments. For woolly monkey (Lagothrix poeppigii) and tapir (Tapirus terrestris), wild birthrates were similar to those from captive populations, whereas birthrates for other ungulates and lowland‐paca (Cuniculus paca) were significantly lower than previous estimates. Conversely, for capuchin monkeys (Sapajus macrocephalus), agoutis (Dasyprocta sp.), and coatis (Nasua nasua), our calculated reproductive rates greatly exceeded often‐used values. Researchers could keep applying classic measures compatible with our estimates, but for other species previous estimates of r_max may not be appropriate. We suggest that data from local studies be used to set hunting quotas. Our maximum rates of population growth in the wild correlated with body weight, which suggests that our method is consistent and reliable. Integration of this method into community‐based wildlife management and the training of local hunters to record pregnancies in hunted animals could efficiently generate useful information of life histories of wild species and thus improve management of natural resources.     

Costs and benefits of pocket gopher foraging: linking behavior and physiology

Animals can attain fitness benefits by maintaining a positive net energy balance, including costs of movement during resource acquisition and the profits from foraging. Subterranean rodent burrowing provides an excellent system in which to examine the effects of movement costs on foraging behavior because it is energetically expensive to excavate burrows. We used an individual-based modeling approach to study pocket gopher foraging and its relationship to digging cost, food abundance, and food distribution. We used a unique combination of an individual-based foraging-behavior model and an energetic model to assess survival, body mass dynamics, and burrow configurations. Our model revealed that even the extreme cost of digging is not as costly as it appears when compared to metabolic costs. Concentrating digging in the area where food was found, or area-restricted search (ARS), was the most energetically efficient digging strategy compared to a random strategy. Field data show that natural burrow configurations were more closely approximated by the animals we modeled using ARS compared to random diggers. By using behavior and simple physiological principles in our model, we were able to observe realistic body mass dynamics and recreate natural movement patterns.     

Using Demographic Invariants to Detect Overharvested Bird Populations from Incomplete Data

Abstract:  Conservation biology must be able to provide guidelines even when available data are incomplete, because data on rare and endangered species are usually limited. For instance, data on the effect of additional—human-induced—sources of mortality on vertebrate populations, such as bycatch of seabirds by longline fisheries, are typically incomplete. The importance of an additional source of mortality can be evaluated by comparing it with the maximum annual growth rate of the species of concern, and various authors have attempted to determine the maximum growth rate from incomplete data. We developed a procedure we call the "demographic invariant method" (DIM). First we determined that the maximum growth rate per generation does not vary, by recalling that it is a dimensionless number primarily independent of body weight and also by empirically establishing its near constancy over a restricted set of bird species for which reliable demographic information was available. This first step provided an implicit function linking generation time and maximum annual growth rate, from which we obtained the maximum annual growth rate as a simple function of generation time. From several different ways of obtaining estimates of generation time, we derived in turn several ways to estimate the maximum annual growth rate of a bird species from incomplete demographic data set. We applied our approach to the Black-footed Albatross (Phoebastria nigripes) and determined from incomplete data that longline fishery bycatch has a biologically significant impact on the growth potential of Black-footed Albatross populations. Our method can be applied broadly to the conservation and management of harvested bird populations.     

Sex-specific energy requirements in nestlings of an extremely sexually size dimorphic bird, the European sparrowhawk (Accipiter nisus)

Allocation of parental investment is predicted to be equal at the population level between both sexes of offspring, and should lead to sex ratio biases in species that exhibit a sex-difference in parental care. Sex-differences in parental care are rarely quantified. We measured daily energy expenditure in free-living nestlings of the extremely sexually size dimorphic European sparrowhawk (Accipiter nisus), using the doubly labelled water method. These data were combined with measured growth characteristics to estimate daily and total metabolised energy intake of male and female young during the nestling stage. Females reached an asymptotic body mass 1.6 times higher than males. This resulted in a total metabolised energy an estimated 1.4 times higher for the nestling stage. Furthermore, we observed a decline in daily metabolised energy with an increase in brood size, which was significantly stronger for females than for males. These results are discussed in the context of Fishers equal allocation theory. Empirical evidence of a sex ratio bias at the end of parental care, with an overall excess of males, is lacking in this species. Consequently, our data do not support the idea of equal allocation between the sexes. The observed sex difference in daily metabolised energy in response to brood size may give scope for sex ratio bias at the level of the individual brood.     

Sexual signalling and viability in a wolf spider (Hygrolycosa rubrofasciata): measurements under laboratory and field conditions

This study examined the crucial prediction of the conditional-handicap theory, the relationship between male sexual trait size and male viability, in the wolf spider Hygrolycosa rubrofasciata. In this species, males court females by drumming dry leaves with their abdomen, and males with the highest drumming rate enjoy highest mating success. We determined male drumming rate, body mass, and mobility, which reflects mate-searching activity, in relation to male survival. Because it is often difficult to know how results obtained from laboratory studies reflect the natural world, particularly when the measured variable is survival, we repeated our study in both laboratory and field conditions. Males drumming at the highest rate survived better than males drumming at a lower rate in both laboratory and field conditions. These results are in accordance with the predictions of conditional-handicap models of sexual selection. Survival was independent of male body mass and there was no significant correlation between male drumming activity and body mass. However, large males moved further than smaller males, and males moving longer distances lost less mass than those moving shorter distances. These results suggest that, moving, and consequently mate-searching activity, may be a condition-dependent trait and that there may be a advantage for large males in mate searching. Received: 22 October 1998 / Received in revised form: 28 January 1999 / Accepted: 14 February 1999     

Asymmetric larval competition in the parasitoid wasp <Emphasis Type="Italic">Nasonia vitripennis</Emphasis>: a role in sex allocation?

Sex allocation theory offers excellent opportunities for testing how animals adjust their behaviour in response to environmental conditions. A major focus has been on instances of local mate competition (LMC), where female-biased broods are produced to maximise mating opportunities for sons. However, the predictions of LMC theory can be altered if there is both local competition for resources during development and an asymmetry between the competitive abilities of the sexes, as has been seen in animals ranging from wasps to birds. In this paper, we test the extent to which asymmetric larval competition alters the predictions of LMC theory in the parasitoid wasp Nasonia vitripennis. We found that the body size of both sexes was negatively correlated with the number of offspring developing within the host. Further, we found that when faced with high levels of competition, the body size of females, but not males, was influenced by the sex ratio of the competing offspring; females were smaller when a higher proportion of the brood was female. This asymmetric competition should favour less biased sex ratios than are predicted by standard LMC theory. We then develop a theoretical model that can be parameterised with our data, allowing us to determine the quantitative consequences of the observed level of asymmetric larval competition for sex allocation. We found that although asymmetric competition selects for less biased sex ratios, this effect is negligible compared to LMC. Furthermore, a similar conclusion is reached when we re-analyse existing data from another parasitoid species where asymmetric larval competition has been observed; Bracon hebetor. Consequently, we suspect that asymmetric larval competition will have its greatest influence on sex ratio evolution in species that have smaller clutches and where local mate competition is not an issue, such as birds and mammals.     

Seasonality of feeding and nutritional status during the austral winter in the Antarctic sea urchin Sterechinus neumayeri

The seasonal pattern of food intake and tissue energy status was measured over a 2 year period for two populations of the common Antarctic echinoid Sterechinus neumayeri living on contrasting substrata at Rothera Point, Adelaide Island, Antarctica. Food intake was estimated from faecal egestion, and both tissue mass and energy content assessed. Food availability was intensely seasonal, in that water column chlorophyll content and sediment pigment content varied markedly throughout the year. In response, both urchin populations showed an extremely seasonal cycle of faecal egestion, indicating a strong seasonality of feeding activity. Urchins from North Cove living on soft sediment fed at a considerably higher rate, and had a significantly larger Aristotle's lantern, than those in South Cove, 1 km away, living on hard substrata and taking a more cosmopolitan diet. Faecal egestion in both populations was zero for a 7 month period in the austral winter of 1997, and again for a 4 month period in 1998. During the austral summer of 1997/1998, 84% of the total annual energy intake took place in the period January to March. Significant decrease in gut tissue mass provided energy for maintenance in early winter, although progressive reduction of both gonad energy content (but without a detectable change in gonad mass) and body wall organic mass provided energy during the late winter period. The mass of reproductive tissue showed large differences between the two sites, but there was no marked decrease associated with spawning. Tissue proximate composition was assessed stoichiometrically from elemental composition and also checked by direct assay; gonad tissue was richer in lipid than gut tissue, though both were dominated by protein and contained only small amounts of carbohydrate. These data suggest that the very strong seasonality of food intake does not pose a significant energetic challenge for this species.     

Variation in plasma levels of insulin-like growth factor-I (IGF-I) in lingcod: relationships among season,size, and gonadal steroids

The physiological response of a fish to its environment is mediated through the endocrine axis controlling growth. Therefore, growth-regulating hormone levels can serve as ecologically relevant indicators of fish growth rate. We quantified variation in plasma insulin-like growth factor-I (IGF-I) to evaluate its potential as an indicator of growth in lingcod, an economically and ecologically important bottomfish species in the northeast Pacific. An information-theoretic model selection approach was used to test the hypothesis that variation in lingcod IGF-I is related to season, body size, and gonadal steroid concentration. Season and a length × season interaction were the most important predictors of plasma IGF-I among the variables we evaluated, suggesting that season and body size should be explicitly accounted for when interpreting endocrine patterns in wild fish populations. This is among the few studies that have measured and interpreted patterns of IGF-I in wild fish and the first to describe seasonal endocrine profiles in lingcod.     

Context-dependent effects of nestling growth trajectories on recruitment probability in the collared flycatcher

Nestling growth is known as an important determinant of fitness in altricial birds, but its evolutionary potential has been debated, and little is known about detailed patterns of current selection on growth. Relationships are often reported between nestling growth and attributes of nestlings and parents, but the interpretation of these depends on the advantages a given growth difference confers to the chicks. Increased growth may have positive, negative or context-dependent effects on offspring fitness, but these effects are largely unknown in natural populations. We measured growth trajectories of body mass in fostered broods of collared flycatchers (Ficedula albicollis) in 3 years of contrasting food conditions. We examined the growth of young and their recruitment probability to the breeding population in relation to year quality, hatching rank, sex, paternal age and paternal attractiveness. We also looked at the interactive effects of growth and intrinsic offspring attributes on recruitment probability. The predictors of nestling growth and those of recruitment did not agree. Moreover, the recruitment consequences of a given nestling growth rate were significantly influenced by nestling rank and paternal ornamentation. Differential recruitment effects of nestling growth in relation to parental traits and nestling attributes suggest that using growth as a generally applicable measure of nestling quality may not be justified. These findings also have implications for morphological evolution and the indicator value of sexual signals.     

Allometric Scaling of Minimal Mammal Densities

Minimum viable densities have rarely been determined directly. Theoretical analyses, based on empirical relationships between average mammal densities, suggest that minimum densities of viable populations are lower for larger mammals. This suggestion has been cast into doubt by other field studies showing populations of small insects and birds at very low densities. We collected 143 of the closest approximations of minimum viable density available, those of minimal, rare, and endangered mammal populations. We found that minimal density decreases as the −0.68 power of body mass. Minimal densities of small mammals are 1000 times those of the largest species. The correlation between minimum viable population density and body mass is negative in the majority of the mammalian taxonomic orders. Although minimum density is, on average, 10% of mean population density, viable population densities of herbivores are 13 times those of carnivores and insectivores. Populations in the wet tropics can apparently sustain themselves at densities much lower than those in temperate climates.     

Sink populations in carnivore management: cougar demography and immigration in a hunted population.

Carnivores are widely hunted for both sport and population control, especially where they conflict with human interests. It is widely believed that sport hunting is effective in reducing carnivore populations and related human-carnivore conflicts, while maintaining viable populations. However, the way in which carnivore populations respond to harvest can vary greatly depending on their social structure, reproductive strategies, and dispersal patterns. For example, hunted cougar (Puma concolor) populations have shown a great degree of resiliency. Although hunting cougars on a broad geographic scale (> 2000 km2) has reduced densities, hunting of smaller areas (i.e., game management units, < 1000 km2), could conceivably fail because of increased immigration from adjacent source areas. We monitored a heavily hunted population from 2001 to 2006 to test for the effects of hunting at a small scale (< 1000 km2) and to gauge whether population control was achieved (lambda < or = 1.0) or if hunting losses were negated by increased immigration allowing the population to remain stable or increase (lambda > or = 1.0). The observed growth rate of 1.00 was significantly higher than our predicted survival/fecundity growth rates (using a Leslie matrix) of 0.89 (deterministic) and 0.84 (stochastic), with the difference representing an 11-16% annual immigration rate. We observed no decline in density of the total population or the adult population, but a significant decrease in the average age of independent males. We found that the male component of the population was increasing (observed male population growth rate, lambda(OM) = 1.09), masking a decrease in the female component (lambda(OF) = 0.91). Our data support the compensatory immigration sink hypothesis; cougar removal in small game management areas (< 1000 km2) increased immigration and recruitment of younger animals from adjacent areas, resulting in little or no reduction in local cougar densities and a shift in population structure toward younger animals. Hunting in high-quality habitats may create an attractive sink, leading to misinterpretation of population trends and masking population declines in the sink and surrounding source areas.     

Savviness of prey to introduced predators

The prey naivety hypothesis posits that prey are vulnerable to introduced predators because many generations in slow gradual coevolution are needed for appropriate avoidance responses to develop. It predicts that prey will be more responsive to native than introduced predators and less responsive to introduced predators that differ substantially from native predators and from those newly established. To test these predictions, we conducted a global meta-analysis of studies that measured the wariness responses of small mammals to the scent of sympatric mammalian mesopredators. We identified 26 studies that met our selection criteria. These studies comprised 134 experiments reporting on the responses of 36 small mammal species to the scent of six introduced mesopredators and 12 native mesopredators. For each introduced mesopredator, we measured their phylogenetic and functional distance to local native mesopredators and the number of years sympatric with their prey. We used predator and prey body mass as a measure of predation risk. Globally, small mammals were similarly wary of the scent of native and introduced mesopredators; phylogenetic and functional distance between introduced mesopredators and closest native mesopredators had no effect on wariness; and wariness was unrelated to the number of prey generations, or years, since first contact with introduced mesopredators. Small mammal wariness was associated with predator-prey body mass ratio, regardless of the nativity. The one thing animals do not seem to recognize is whether their predators are native.     

Irruptive population dynamics in Yellowstone pronghorn.

Irruptive population dynamics appear to be widespread in large herbivore populations, but there are few empirical examples from long time series with small measurement error and minimal harvests. We analyzed an 89-year time series of counts and known removals for pronghorn (Antilocapra americana) in Yellowstone National Park of the western United States during 1918-2006 using a suite of density-dependent, density-independent, and irruptive models to determine if the population exhibited irruptive dynamics. Information-theoretic model comparison techniques strongly supported irruptive population dynamics (Leopold model) and density dependence during 1918-1946, with the growth rate slowing after counts exceeded 600 animals. Concerns about sagebrush (Artemisia spp.) degradation led to removals of >1100 pronghorn during 1947-1966, and counts decreased from approximately 700 to 150. The best models for this period (Gompertz, Ricker) suggested that culls replaced intrinsic density-dependent mechanisms. Contrary to expectations, the population did not exhibit enhanced demographic vigor soon after the termination of the harvest program, with counts remaining between 100 and 190 animals during 1967 1981. However, the population irrupted (Caughley model with a one-year lag) to a peak abundance of approximately 600 pronghorn during 1982-1991, with a slowing in growth rate as counts exceeded 500. Numbers crashed to 235 pronghorn during 1992-1995, perhaps because important food resources (e.g., sagebrush) on the winter range were severely diminished by high densities of browsing elk, mule deer, and pronghorn. Pronghorn numbers remained relatively constant during 1996-2006, at a level (196-235) lower than peak abundance, but higher than numbers following the release from culling. The dynamics of this population supported the paradigm that irruption is a fundamental pattern of growth in many populations of large herbivores with high fecundity and delayed density-dependent effects on recruitment when forage and weather conditions become favorable after range expansion or release from harvesting. Incorporating known removals into population models that can describe a wide range of dynamics can greatly improve our interpretation of observed dynamics in intensively managed populations.