To select appropriate recovery strategies for endangered populations, we must understand the dynamics of small populations and distinguish between the possible causes that drive such populations to low numbers. It has been suggested that the pattern of population decline may be inversely density-dependent with population growth rates decreasing as populations become very small; however, empirical evidence of such accelerated declines at low densities is rare. Here we analyzed the pattern of decline of a threatened population of woodland caribou (Rangifer tarandus caribou) in British Columbia, Canada. Using information on the instantaneous rate of increase relative to caribou density in suitable winter foraging habitat, as well as on pregnancy rates and on causes and temporal distribution of mortalities from a sample of 349 radiocollared animals from 15 subpopulations, we tested 3 hypothesized causes of decline: (a) food regulation caused by loss of suitable winter foraging habitat, (b) predation-sensitive foraging caused by loss of suitable winter foraging habitat and (c) predation with caribou being secondary prey. Population sizes of caribou subpopulations ranged from < 5 to > 500 individuals. Our results showed that the rates of increase of these subpopulations varied from -0.1871 to 0.0496 with smaller subpopulations declining faster than larger subpopulations. Rates of increase were positively related to the density of caribou in suitable winter foraging habitat. Pregnancy rates averaged 92.4% +/-2.24 and did not differ among subpopulations. In addition, we found predation to be the primary cause of mortality in 11 of 13 subpopulations with known causes of mortality and predation predominantly occurred during summer. These results are consistent with predictions that caribou subpopulations are declining as a consequence of increased predation. Recovery of these woodland caribou will thus require a multispecies perspective and an appreciation for the influence of inverse density dependence on population trajectories.

We used census results and radiotelemetry locations of >380 collared individuals sampled over the entire distribution of the endangered mountain ecotype of woodland caribou (Rangifer tarandus caribou (Gmelin, 1788)) in British Columbia, Canada, to delineate population structure and document the size and trend of the identified populations. We also describe the spatial pattern of decline and the causes and timing of adult mortality and provide estimates of vital rates necessary to develop a population viability analysis. Our results indicate that the abundance of mountain caribou in British Columbia is declining. We found adult female annual survival rates below annual survival rates commonly reported for large ungulates. The major proximate cause of population decline appears to be predation on adult caribou. Spatial patterns of population dynamics revealed a continuous range contraction and an increasing fragmentation of mountain caribou into smaller, isolated subpopulations. The population fragmentation process predominantly occurs at the outer boundaries of the current distribution. Our results indicate that recovery strategies for mountain caribou should be directed at factors contributing to the fragmentation and isolation of mountain caribou populations as well as management strategies aimed at increasing adult survival.

Summary1. Large-scale habitat loss is frequently identified with loss of biodiversity, but examples of the direct effect of habitat alterations on changes in vital rates remain rare. Quantifying and understanding the relationship between habitat composition and changes in vital rates, however, is essential for the development of effective conservation strategies. 2. It has been suggested that the decline of woodland caribou Rangifer tarandus caribou populations in North America is precipitated by timber harvesting that creates landscapes of early seral forests. Such habitat changes have altered the predator-prey system resulting in asymmetric predation, where predators are maintained by alternative prey (i.e. apparent competition). However, a direct link between habitat condition and caribou population declines has not been documented. 3. We estimated survival probabilities for the threatened arboreal lichen-feeding ecotype of woodland caribou in British Columbia, Canada, at two different spatial scales. At the broader scale, observed variation in adult female survival rates among 10 distinct populations (range = 0·67-0·93) was best explained by variation in the amount of early seral stands within population ranges and population density. At the finer scale, home ranges of caribou killed by predators had lower proportions of old forest and more mid-aged forest as compared with multi-annual home ranges where caribou were alive. 4. These results are consistent with predictions from the apparent competition hypothesis and quantify direct fitness consequences for caribou following habitat alterations. We conclude that apparent competition can cause rapid population declines and even extinction where changes in species composition occur following large scale habitat change.

Mule deer and black‐tailed deer Odocoileus hemionus have exhibited marked population fluctuations throughout their range over the past century. The relative contributions of predation, forage availability and weather to observed population changes remain unclear and controversial. We reviewed 48 studies on Odocoileus hemionus survival and predation from the past 30 years and quantified age‐specific vital rates, population growth rates (λ) and causes of mortality. We also evaluated the effect of environmental variables on variation in vital rates and the contribution of age‐specific survival to population growth. Age‐specific survival (ϕ) was the most frequently studied population parameter. Odocoileus hemionus have lower and more variable fawn survival than other ungulate species (ϕsummer = 0.44, CV = 0.42; ϕannual = 0.29, CV = 0.67). Adult female survival conversely appeared to be high and stable throughout the geographical range of the species (ϕannual = 0.84, CV = 0.06). Observed low fawn survival appears to be compensated for by high fecundity rates. Predation was the primary proximate cause of mortality for all age classes, and was an important source of summer fawn mortality and of mortality in multi‐prey, multi‐predator systems. However, predator removal studies suggest that predation is compensatory, particularly at high deer densities, and that nutrition and weather shape population dynamics. We propose three models to explain local population dynamics of Odocoileus hemionus: (i) populations are limited by forage availability and weather; (ii) adult females are limited by forage availability, fawns are limited by forage availability and predation, and population growth is constrained by fecundity and fawn predation; and (iii) large changes in the abundance of predators or alternative prey change predation risk and destabilize population dynamics. Future research should be focused on: the effects of age‐specific survival on population growth; possible interactions between predation, forage availability and weather; and the importance of multiple predator and prey species in shaping the population dynamics of Odocoileus hemionus.

Pumas (Puma concolor) and black bears (Ursus americanus) are large carnivores that may influence scavenger population dynamics. We used motion-triggered video cameras deployed at deer carcasses to determine how pumas and black bears affected three aspects of carrion acquisition by scavengers: presence, total feeding time, and mean feeding-bout duration. We found that pumas were unable to limit acquisition of carrion by large carnivores but did limit aspects of carrion acquisition by both birds and mesocarnivores. Through their suppression of mesocarnivores and birds, pumas apparently initiated a cascading pattern and increased carrion acquisition by small carnivores. In contrast, black bears monopolized carrion resources and generally had larger limiting effects on carrion acquisition by all scavengers. Black bears also limited puma feeding behaviors at puma kills, which may require pumas to compensate for energetic losses through increasing their kill rates of ungulates. Our results suggest that pumas provide carrion and selectively influence species acquiring carrion, while black bears limit carrion availability to all other scavengers. These results suggest that the effects of large carnivores on scavengers depend on attributes of both carnivores and scavengers (including size) and that competition for carcasses may result in intraguild predation as well as mesocarnivore release.

Apparent competition is an important process influencing many ecological communities. We used predator-prey theory to predict outcomes of ecosystem experiments aimed at mitigating apparent competition by reducing primary prey. Simulations predicted declines in secondary prey following reductions in primary prey because predators consumed more secondary prey until predator numbers responded to reduced prey densities. Losses were exacerbated by a higher carrying capacity of primary prey and a longer lag time of the predator's numerical response, but a gradual reduction in primary prey was less detrimental to the secondary prey. We compared predictions against two field experiments where endangered woodland caribou (Rangifer tarandus caribou) were victims of apparent competition. First, when deer (Odocoileus sp.) declined suddenly following a severe winter, cougar (Puma concolor) declined with a 1-2-year lag, yet in the interim more caribou were killed by cougars, and caribou populations declined by 40%. Second, when moose (Alces alces) were gradually reduced using a management experiment, wolf (Canis lupus) populations declined but did not shift consumption to caribou, and the largest caribou subpopulation stabilized. The observed contrasting outcomes of sudden versus gradual declines in primary prey supported theoretical predictions. Combining theory with field studies clarified how to manage communities to mitigate endangerment caused by apparent competition that affects many taxa.

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Introduced species can threaten native taxa in multiple ways, including competition and hybridization, which can reduce fitness, alter ecological niches or swamp native genomes. Encroachment and hybridization by introduced species also provide opportunities to study the dynamics of invasiveness and hybridization during early stages following contact. We used 33 microsatellites, 51 single nucleotide polymorphisms and a mtDNA marker to characterize the extent and spatial pattern of encroachment and hybridization between a native, endemic subspecies of red fox (Vulpes vulpes patwin) and an introduced red fox population composed of highly admixed, phylogenetically divergent stock, resulting from a century of domestication. Both nuclear and mtDNA markers indicated that hybridization was primarily restricted to a narrow zone where the two populations came into contact. Although a few introgressed genotypes were detected in the interior of the native range, we found no immigrant foxes or F(1) or F(2) hybrids there, suggesting native foxes excluded introduced individuals. We speculate that the observed interbreeding at the periphery was facilitated by low densities. In total, 98% of mtDNA haplotypes in the native range were native and 96% of the nuclear ancestry was estimated to be native. Although the introduced range had expanded fivefold over the past four decades, native and non-native haplotypes from museum samples collected in and near the native range three decades earlier showed a similar geographic distribution as today, suggesting that the native range and hybrid zone were relatively stable. We hypothesize that the monogamous mating system of red foxes and other wild canids may enhance their resistance to hybridization because of greater fitness consequences associated with mate discrimination.