Effectiveness of population-based recovery actions for threatened southern mountain caribou.

Habitat loss is affecting many species, including the southern mountain caribou (Rangifer tarandus caribou) population in western North America. Over the last half century, this threatened caribou population's range and abundance have dramatically contracted. An integrated population model was used to analyze 51 years (1973-2023) of demographic data from 40 southern mountain caribou subpopulations to assess the effectiveness of population-based recovery actions at increasing population growth. Reducing potential limiting factors on threatened caribou populations offered a rare opportunity to identify the causes of decline and assess methods of recovery. Southern mountain caribou abundance declined by 51% between 1991 and 2023, and 37% of subpopulations were functionally extirpated. Wolf reduction was the only recovery action that consistently increased population growth when applied in isolation, and combinations of wolf reductions with maternal penning or supplemental feeding provided rapid growth but were applied to only four subpopulations. As of 2023, recovery actions have increased the abundance of southern mountain caribou by 52%, compared to a simulation with no interventions. When predation pressure was reduced, rapid population growth was observed, even under contemporary climate change and high levels of habitat loss. Unless predation is reduced, caribou subpopulations will continue to be extirpated well before habitat conservation and restoration can become effective.

Mesocarnivores vary in their spatiotemporal avoidance strategies at communications hubs of an apex carnivore.

Mesocarnivores face interspecific competition and risk intraguild predation when sharing resources with apex carnivores. Within a landscape, carnivores across trophic levels may use the same communication hubs, which provide a mix of risks (injury/death) and rewards (gaining information) for subordinate species. We predicted that mesocarnivores would employ different strategies to avoid apex carnivores at shared communication hubs, depending on their trophic position. To test our prediction, we examined how different subordinate carnivore species in the Santa Cruz Mountains of California, USA, manage spatial overlap with pumas (Puma concolor), both at communication hubs and across a landscape-level camera trap array. We estimated species-specific occurrence, visitation rates, temporal overlap, and Avoidance-Attraction Ratios from camera traps and tested for differences between the two types of sites. We found that mesocarnivores generally avoided pumas at communication hubs, and this became more pronounced when pumas scent-marked during their most recent visit. Coyotes (Canis latrans), the pumas' closest subordinate competitor in our system, exhibited the strongest avoidance at communication hubs. Gray foxes (Urocyon cinereoargenteus) avoided pumas the least, which may suggest possible benefits from pumas suppressing coyotes. Overall, mesocarnivores exhibited various spatiotemporal avoidance strategies at communication hubs rather than outright avoidance, likely because they benefit from information gained while 'eavesdropping' on puma activity. Variability in avoidance strategies may be due to differential predation risks, as apex carnivores often interact more aggressively with their closest competitors. Combined, our results show how apex carnivores trigger complex species interactions across the entire carnivore guild and how trophic position determines behavioral responses and subsequent space use of subordinate mesocarnivores across the landscape.

Season, prey availability, sex, and age explain prey size selection in a large solitary carnivore.

Prey selection is a fundamental aspect of ecology that drives evolution and community structure, yet the impact of intraspecific variation on the selection for prey size remains largely unaccounted for in ecological theory. Here, we explored puma (Puma concolor) prey selection across six study sites in North and South America. Our results highlighted the strong influence of season and prey availability on puma prey selection, and the smaller influence of puma age. Pumas in all sites selected smaller prey in warmer seasons following the ungulate birth pulse. Our top models included interaction terms between sex and age, suggesting that males more than females select larger prey as they age, which may reflect experiential learning. When accounting for variable sampling across pumas in our six sites, male and female pumas killed prey of equivalent size, even though males are larger than females, challenging assumptions about this species. Nevertheless, pumas in different study sites selected prey of different sizes, emphasizing that the optimal prey size for pumas is likely context-dependent and affected by prey availability. The mean prey weight across all sites averaged 1.18 times mean puma weight, which was less than predicted as the optimal prey size by energetics and ecological theory (optimal prey = 1.45 puma weight). Our results help refine our understanding of optimal prey for pumas and other solitary carnivores, as well as corroborate recent research emphasizing that carnivore prey selection is impacted not just by energetics but by the effects of diverse ecology.

Landscape-wide pulse events predict trait-based responses among wetland birds in perennial channels of a dryland wetland.

Wetlands in arid or semiarid zones are vital for maintaining biodiversity but face growing threats. Flooding regime variability is a key driver of ecological dynamism in these systems, dictating primary productivity on a large spatial scale. The functional composition or diversity of wetland-dependent bird species has been found to be sensitive to fluctuations in hydrological regimes and can thus be indicative of cascading ecosystem responses associated with climate change. In this paper, we investigate whether large-scale changes in inundation and fire-a significant additional biodiversity determinant in (semi-)arid landscapes-are reliable predictors of functional group responses of wetland-dependent birds along a perennial channel of the Okavango Delta, Botswana. We fit generalized additive models (GAMs) to 6 years of bird survey data collected along ~190-km-long annual transects and use remotely sensed landscape-level inundation estimates, as well as spatiotemporal distance to fire, to predict the responsiveness of seven trait-based functional group abundances. During the surveys, a total of 89 different wetland-dependent bird species were recorded, including 76 residents, across all years, with below-surface feeding waders consistently the most abundant functional group. Including estimated spatiotemporal variability in flooding and fire, as well as their interactions, improved model fit for all seven functional groups, explaining between 46.8% and 68.3% of variability in functional group abundances. Covariates representing longer-term variability in inundation generally performed better than shorter-term ones. For example, variability in inundation over the 5 months preceding a survey best predicted the responses of all functional groups, which also all exhibited responsiveness to the interaction between flooding and fire. We were able to interpret the responses of individual functional groups, based on the resource exploitation assumption. Overall, our results suggest that perennial waters in dryland wetlands offer functional refugia to wetland-dependent birds and highlight the indicative power of large-scale trait-based bird monitoring. Our findings demonstrate the potential utility of such a monitoring regime for dryland wetland ecosystems vulnerable to industrial-scale anthropogenic pressure and associated climate change.

The ecology of human-caused mortality for a protected large carnivore.

Mitigating human-caused mortality for large carnivores is a pressing global challenge for wildlife conservation. However, mortality is almost exclusively studied at local (within-population) scales creating a mismatch between our understanding of risk and the spatial extent most relevant to conservation and management of wide-ranging species. Here, we quantified mortality for 590 radio-collared mountain lions statewide across their distribution in California to identify drivers of human-caused mortality and investigate whether human-caused mortality is additive or compensatory. Human-caused mortality, primarily from conflict management and vehicles, exceeded natural mortality despite mountain lions being protected from hunting. Our data indicate that human-caused mortality is additive to natural mortality as population-level survival decreased as a function of increasing human-caused mortality and natural mortality did not decrease with increased human-caused mortality. Mortality risk increased for mountain lions closer to rural development and decreased in areas with higher proportions of citizens voting to support environmental initiatives. Thus, the presence of human infrastructure and variation in the mindset of humans sharing landscapes with mountain lions appear to be primary drivers of risk. We show that human-caused mortality can reduce population-level survival of large carnivores across large spatial scales, even when they are protected from hunting.

Implications of large-scale infrastructure development for biodiversity in Indonesian Borneo.

Indonesia is embarking on an ambitious relocation of its capital city to Kalimantan, Borneo, bringing with it major urban and road infrastructure. Yet, despite being one of the world's most biologically diverse regions, the potential implications of this development for wildlife have yet to be fully assessed. We explored the potential impacts of the capital relocation, and road expansion and upgrades to critical habitat for medium-large mammals (>1 kg) using camera trap data from 11 forested landscapes. We applied Bayesian multi-species occupancy models to predict community and species-level responses to anthropogenic and environmental factors. We extrapolated spatial patterns of occupancy and species diversity across the forests of Kalimantan and identified "critical habitats" as the top 20th percentile of occupancy and species richness values. We subsequently overlapped these critical habitat layers with infrastructure impact zones to estimate the area that could potentially be affected by direct or secondary impacts. At both the community and species-level, distance to primary roads had the strongest negative influence on habitat-use. Occupancy was also influenced by forest quality and multidimensional poverty conditions in adjacent villages, demonstrating the sensitivity of biodiversity to socio-ecological pressures. Less than 1 % of the critical habitat for the threatened mammal community lay within the direct impact zone (30 km radius) of the capital relocation. However, approximately 16 % was located within 200 km and could potentially be affected by uncontrolled secondary impacts such as urban sprawl and associated regional development. The often-overlooked secondary implications of upgrading existing roads could also intersect a large amount of critical habitat for lowland species. Mitigating far-reaching secondary impacts of infrastructure development should be fully incorporated into environmental impact assessments. This will provide Indonesia with an opportunity to set an example of sustainable infrastructure development in the tropics.

Standardizing protocols for determining the cause of mortality in wildlife studies.

Mortality site investigations of telemetered wildlife are important for cause-specific survival analyses and understanding underlying causes of observed population dynamics. Yet, eroding ecoliteracy and a lack of quality control in data collection can lead researchers to make incorrect conclusions, which may negatively impact management decisions for wildlife populations. We reviewed a random sample of 50 peer-reviewed studies published between 2000 and 2019 on survival and cause-specific mortality of ungulates monitored with telemetry devices. This concise review revealed extensive variation in reporting of field procedures, with many studies omitting critical information for the cause of mortality inference. Field protocols used to investigate mortality sites and ascertain the cause of mortality are often minimally described and frequently fail to address how investigators dealt with uncertainty. We outline a step-by-step procedure for mortality site investigations of telemetered ungulates, including evidence that should be documented in the field. Specifically, we highlight data that can be useful to differentiate predation from scavenging and more conclusively identify the predator species that killed the ungulate. We also outline how uncertainty in identifying the cause of mortality could be acknowledged and reported. We demonstrate the importance of rigorous protocols and prompt site investigations using data from our 5-year study on survival and cause-specific mortality of telemetered mule deer (Odocoileus hemionus) in northern California. Over the course of our study, we visited mortality sites of neonates (n = 91) and adults (n = 23) to ascertain the cause of mortality. Rapid site visitations significantly improved the successful identification of the cause of mortality and confidence levels for neonates. We discuss the need for rigorous and standardized protocols that include measures of confidence for mortality site investigations. We invite reviewers and journal editors to encourage authors to provide supportive information associated with the identification of causes of mortality, including uncertainty.

Functional traits driving species role in the structure of terrestrial vertebrate scavenger networks.

Species assemblages often have a non-random nested organization, which in vertebrate scavenger (carrion-consuming) assemblages is thought to be driven by facilitation in competitive environments. However, not all scavenger species play the same role in maintaining assemblage structure, as some species are obligate scavengers (i.e., vultures) and others are facultative, scavenging opportunistically. We used a database with 177 vertebrate scavenger species from 53 assemblages in 22 countries across five continents to identify which functional traits of scavenger species are key to maintaining the scavenging network structure. We used network analyses to relate ten traits hypothesized to affect assemblage structure with the "role" of each species in the scavenging assemblage in which it appeared. We characterized the role of a species in terms of both the proportion of monitored carcasses on which that species scavenged, or scavenging breadth (i.e., the species "normalized degree"), and the role of that species in the nested structure of the assemblage (i.e., the species "paired nested degree"), therefore identifying possible facilitative interactions among species. We found that species with high olfactory acuity, social foragers, and obligate scavengers had the widest scavenging breadth. We also found that social foragers had a large paired nested degree in scavenger assemblages, probably because their presence is easier to detect by other species to signal carcass occurrence. Our study highlights differences in the functional roles of scavenger species and can be used to identify key species for targeted conservation to maintain the ecological function of scavenger assemblages.

Biotic and abiotic drivers of dispersion dynamics in a large-bodied tropical vertebrate, the Western Bornean orangutan.

Understanding of animal responses to dynamic resource landscapes is based largely on research on temperate species with small body sizes and fast life histories. We studied a large, tropical mammal with an extremely slow life history, the Western Bornean orangutan (Pongo pygmaeus wurmbii), across a heterogeneous natural landscape encompassing seven distinct forest types. Our goals were to characterize fluctuations in abundance, test hypotheses regarding the relationship between dispersion dynamics and resource availability, and evaluate how movement patterns are influenced by abiotic conditions. We surveyed abundance in Gunung Palung National Park, West Kalimantan, Indonesia, for 99 consecutive months and simultaneously recorded weather data and assessed fruit availability. We developed a Bayesian hierarchical distance sampling model to estimate population dispersion and assess the roles of fruit availability, rainfall, and temperature in driving movement patterns across this heterogeneous landscape. Orangutan abundance varied dramatically over space and time. Each forest type was important in sustaining more than 40% of the total orangutans on site during at least one month, as animals moved to track asynchronies in fruiting phenology. We conclude that landscape-level movements buffer orangutans against fruit scarcity, peat swamps are crucial fallback habitats, and orangutans' use of high elevation forests is strongly dependent on abiotic conditions. Our results show that orangutans can periodically occupy putative-sink habitats and be virtually absent for extended periods from habitats that are vitally important in sustaining their population, highlighting the need for long-term studies and potential risks in interpreting occurrence or abundance measures as indicators of habitat importance.

Vital rates of two small populations of brown bears in Canada and range-wide relationship between population size and trend.

Identifying mechanisms of population change is fundamental for conserving small and declining populations and determining effective management strategies. Few studies, however, have measured the demographic components of population change for small populations of mammals (<50 individuals). We estimated vital rates and trends in two adjacent but genetically distinct, threatened brown bear (Ursus arctos) populations in British Columbia, Canada, following the cessation of hunting. One population had approximately 45 resident bears but had some genetic and geographic connectivity to neighboring populations, while the other population had <25 individuals and was isolated. We estimated population-specific vital rates by monitoring survival and reproduction of telemetered female bears and their dependent offspring from 2005 to 2018. In the larger, connected population, independent female survival was 1.00 (95% CI: 0.96-1.00) and the survival of cubs in their first year was 0.85 (95% CI: 0.62-0.95). In the smaller, isolated population, independent female survival was 0.81 (95% CI: 0.64-0.93) and first-year cub survival was 0.33 (95% CI: 0.11-0.67). Reproductive rates did not differ between populations. The large differences in age-specific survival estimates resulted in a projected population increase in the larger population (λ = 1.09; 95% CI: 1.04-1.13) and population decrease in the smaller population (λ = 0.84; 95% CI: 0.72-0.95). Low female survival in the smaller population was the result of both continued human-caused mortality and an unusually high rate of natural mortality. Low cub survival may have been due to inbreeding and the loss of genetic diversity common in small populations, or to limited resources. In a systematic literature review, we compared our population trend estimates with those reported for other small populations (<300 individuals) of brown bears. Results suggest that once brown bear populations become small and isolated, populations rarely increase and, even with intensive management, recovery remains challenging.

Reintroduced wolves and hunting limit the abundance of a subordinate apex predator in a multi-use landscape.

Top-down effects of apex predators are modulated by human impacts on community composition and species abundances. Consequently, research supporting top-down effects of apex predators occurs almost entirely within protected areas rather than the multi-use landscapes dominating modern ecosystems. Here, we developed an integrated population model to disentangle the concurrent contributions of a reintroduced apex predator, the grey wolf, human hunting and prey abundances on vital rates and abundance of a subordinate apex predator, the puma. Increasing wolf numbers had strong negative effects on puma fecundity, and subadult and adult survival. Puma survival was also influenced by density dependence. Overall, puma dynamics in our multi-use landscape were more strongly influenced by top-down forces exhibited by a reintroduced apex predator, than by human hunting or bottom-up forces (prey abundance) subsidized by humans. Quantitatively, the average annual impact of human hunting on equilibrium puma abundance was equivalent to the effects of 20 wolves. Historically, wolves may have limited pumas across North America and dictated puma scarcity in systems lacking sufficient refugia to mitigate the effects of competition.

Scavenging in the Anthropocene: Human impact drives vertebrate scavenger species richness at a global scale.

Understanding the distribution of biodiversity across the Earth is one of the most challenging questions in biology. Much research has been directed at explaining the species latitudinal pattern showing that communities are richer in tropical areas; however, despite decades of research, a general consensus has not yet emerged. In addition, global biodiversity patterns are being rapidly altered by human activities. Here, we aim to describe large-scale patterns of species richness and diversity in terrestrial vertebrate scavenger (carrion-consuming) assemblages, which provide key ecosystem functions and services. We used a worldwide dataset comprising 43 sites, where vertebrate scavenger assemblages were identified using 2,485 carcasses monitored between 1991 and 2018. First, we evaluated how scavenger richness (number of species) and diversity (Shannon diversity index) varied among seasons (cold vs. warm, wet vs. dry). Then, we studied the potential effects of human impact and a set of macroecological variables related to climatic conditions on the scavenger assemblages. Vertebrate scavenger richness ranged from species-poor to species rich assemblages (4-30 species). Both scavenger richness and diversity also showed some seasonal variation. However, in general, climatic variables did not drive latitudinal patterns, as scavenger richness and diversity were not affected by temperature or rainfall. Rainfall seasonality slightly increased the number of species in the community, but its effect was weak. Instead, the human impact index included in our study was the main predictor of scavenger richness. Scavenger assemblages in highly human-impacted areas sustained the smallest number of scavenger species, suggesting human activity may be overriding other macroecological processes in shaping scavenger communities. Our results highlight the effect of human impact at a global scale. As species-rich assemblages tend to be more functional, we warn about possible reductions in ecosystem functions and the services provided by scavengers in human-dominated landscapes in the Anthropocene.

Multiple anthropogenic interventions drive puma survival following wolf recovery in the Greater Yellowstone Ecosystem.

Humans are primary drivers of declining abundances and extirpation of large carnivores worldwide. Management interventions to restore biodiversity patterns, however, include carnivore reintroductions, despite the many unresolved ecological consequences associated with such efforts. Using multistate capture-mark-recapture models, we explored age-specific survival and cause-specific mortality rates for 134 pumas (Puma concolor) monitored in the Greater Yellowstone Ecosystem during gray wolf (Canis lupus) recovery. We identified two top models explaining differences in puma survivorship, and our results suggested three management interventions (unsustainable puma hunting, reduction in a primary prey, and reintroduction of a dominant competitor) have unintentionally impacted puma survival. Specifically, puma survival across age classes was lower in the 6-month hunting season than the 6-month nonhunting season; human-caused mortality rates for juveniles and adults, and predation rates on puma kittens, were higher in the hunting season. Predation on puma kittens, and starvation rates for all pumas, also increased as managers reduced elk (Cervus elaphus) abundance in the system, highlighting direct and indirect effects of competition between recovering wolves and pumas over prey. Our results emphasize the importance of understanding the synergistic effects of existing management strategies and the recovery of large, dominant carnivores to effectively conserve subordinate, hunted carnivores in human-dominated landscapes.

Analyses of phenotypic differentiations among South Georgian Diving Petrel (Pelecanoides georgicus) populations reveal an undescribed and highly endangered species from New Zealand.

Unresolved taxonomy of threatened species is problematic for conservation as the field relies on species being distinct taxonomic units. Differences in breeding habitat and results from a preliminary molecular analysis indicated that the New Zealand population of the South Georgian Diving Petrel (Pelecanoides georgicus) was a distinct, yet undescribed, species. We measured 11 biometric characters and scored eight plumage characters in 143 live birds and 64 study skins originating from most populations of P. georgicus, to assess their taxonomic relationships. We analysed differences with principal component analyses (PCA), factorial ANOVAs, and Kruskal-Wallis rank sum tests. Results show that individuals from New Zealand differ significantly from P. georgicus from all other populations as following: 1) longer wings, 2) longer outer tail feathers, 3) deeper bills, 4) longer heads, 5) longer tarsi, 6) limited collar extent, 7) greater extent of contrasting scapulars, 8) larger contrasting markings on the secondaries, 9) paler ear coverts, 10) paler collars, and 11) paler flanks. Furthermore, we used a species delimitation test with quantitative phenotypic criteria; results reveal that the New Zealand population of P. georgicus indeed merits species status. We hereby name this new species Pelecanoides whenuahouensis sp. nov. Due to severe reductions in its range and the very low number of remaining birds (~150 individuals limited to a single breeding colony on Codfish Island/Whenua Hou) the species warrants listing as 'Critically Endangered'. An abstract in the Maori language/Te Reo Maori can be found in S1 File.

The importance of motivation, weapons, and foul odors in driving encounter competition in carnivores.

Encounter competition is interference competition in which animals directly contend for resources. Ecological theory predicts the trait that determines the resource holding potential (RHP), and hence the winner of encounter competition, is most often body size or mass. The difficulties of observing encounter competition in complex organisms in natural environments, however, has limited opportunities to test this theory across diverse species. We studied the outcome of encounter competition contests among mesocarnivores at deer carcasses in California to determine the most important variables for winning these contests. We found some support for current theory in that body mass is important in determining the winner of encounter competition, but we found that other factors including hunger and species-specific traits were also important. In particular, our top models were "strength and hunger" and "size and hunger," with models emphasizing the complexity of variables influencing outcomes of encounter competition. In addition, our wins above predicted (WAP) statistic suggests that an important aspect that determines the winner of encounter competition is species-specific advantages that increase their RHP, as bobcats (Lynx rufus) and spotted skunks (Spilogale gracilis) won more often than predicted based on mass. In complex organisms, such as mesocarnivores, species-specific adaptations, including strategic behaviors, aggressiveness, and weapons, contribute to competitive advantages and may allow certain species to take control or defend resources better than others. Our results help explain how interspecific competition shapes the occurrence patterns of species in ecological communities.

Scent marking in Sunda clouded leopards (Neofelis diardi): novel observations close a key gap in understanding felid communication behaviours.

Intraspecific communication is integral to the behavioural ecology of solitary carnivores, but observing and quantifying their communication behaviours in natural environments is difficult. Our systematic literature review found that basic information on scent marking is completely lacking for 23% of all felid species, and information on 21% of other felid species comes solely from one study of captive animals. Here we present results of the first systematic investigation of the scent marking behaviours of Sunda clouded leopards in the wild. Our observations using motion-triggered video cameras in Indonesian Borneo are novel for clouded leopards, and contrary to previous descriptions of their behaviour. We found that clouded leopards displayed 10 distinct communication behaviours, with olfaction, scraping, and cheek rubbing the most frequently recorded. We also showed that males make repeated visits to areas they previously used for marking and that multiple males advertise and receive information at the same sites, potentially enhancing our ability to document and monitor clouded leopard populations. The behaviours we recorded are remarkably similar to those described in other solitary felids, despite tremendous variation in the environments they inhabit, and close a key gap in understanding and interpreting communication behaviours of clouded leopards and other solitary felids.

The Role of Scent Marking in Mate Selection by Female Pumas (Puma concolor).

Mate selection influences individual fitness, is often based on complex cues and behaviours, and can be difficult to study in solitary species including carnivores. We used motion-triggered cameras at 29 community scrapes (i.e. scent marking locations used by multiple individuals) and home range data from 39 GPS-collared pumas (Puma concolor) to assess the relevance of communication behaviours for mate selection by female pumas in California. Female pumas visited community scrapes irregularly and visitation bouts appeared to be correlated with oestrus. Female pumas on average selected from 1.7 collared males, and selection was based on multiple cues that varied among the different time periods measured (i.e. the female's visitation bout and in 90 days previous to the consorting event). Female mate selection over the course of a visitation bout was based on frequency of the male visitation, mass, and age. In the 90 days previous to consorting, the number of scrapes a male created was the most important contributor to selection, which was likely related to his residency status. We also found that at least 14% of females mated with multiple males, thus possibly confusing paternity. Our findings provide a mechanistic understanding of how female pumas use scent and auditory communication at community scrapes to select dominant resident males to mate with.

The comparative effects of large carnivores on the acquisition of carrion by scavengers.

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.

Using predator-prey theory to predict outcomes of broadscale experiments to reduce apparent competition.

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.

Trophic facilitation or limitation? Comparative effects of pumas and black bears on the scavenger community.

Scavenging is a widespread behaviour and an important process influencing food webs and ecological communities. Large carnivores facilitate the movement of energy across trophic levels through the scavenging and decomposition of their killed prey, but competition with large carnivores is also likely to constrain acquisition of carrion by scavengers. We used an experimental approach based on motion-triggered video cameras at black-tailed deer (Odocoileus hemionus columbianus) carcasses to measure the comparative influences of two large carnivores in the facilitation and limitation of carrion acquisition by scavengers. We found that pumas (Puma concolor) and black bears (Ursus americanus) had different effects on their ecological communities. Pumas, as a top-level predator, facilitated the consumption of carrion by scavengers, despite significantly reducing their observed sum feeding times (165.7 min ± 21.2 SE at puma kills 264.3 min ± 30.1 SE at control carcasses). In contrast, black bears, as the dominant scavenger in the system, limited consumption of carrion by scavengers as evidenced by the observed reduction of scavenger species richness recorded at carcasses where they were present (mean = 2.33 ± 0.28 SE), compared to where they were absent (mean = 3.28 ± 0.23 SE). Black bears also had large negative effects on scavenger sum feeding times (88.5 min ± 19.8 SE at carcasses where bears were present, 372.3 min ± 50.0 SE at carcasses where bears were absent). In addition, we found that pumas and black bears both increased the nestedness (a higher level of order among species present) of the scavenger community. Our results suggest that scavengers have species-specific adaptions to exploit carrion despite large carnivores, and that large carnivores influence the structure and composition of scavenger communities. The interactions between large carnivores and scavengers should be considered in future studies of food webs and ecological communities.