Apex Carnivores Research Articles

Reconciling biome-wide conservation of an apex carnivore with land-use economics in the increasingly threatened Pantanal wetlands

Conservation of carnivores involves finding solutions to minimize habitat loss and human-wildlife conflict. Understanding the nature of land-use economics can allow us to mitigate both threats. In the Pantanal, the two main economic activities are cattle ranching and ecotourism, each of which directly and indirectly affect the persistence of jaguars (Panthera onca). To understand how the geography of these economic activities is related to jaguar populations, we developed a jaguar distribution model (JDM), livestock density model, and ecotourism lodge density model for the Pantanal. Due to the recent wildfires within the Pantanal, we also assess the impact of burnt areas that are suitable for jaguars, cattle ranching, and tourism. Our JDM indicate that 64% of the Pantanal holds suitable habitat for jaguars. However, jaguar habitat suitability was positively correlated with ecotourism, but negatively correlated with areas most suitable for intensive cattle-ranching. This demonstrates a biome-wide scenario compatible with jaguar conservation. Of particular concern, recent wildfires overlap most suitable areas for jaguars. If wildfires become increasingly frequent, this would represent a serious threat to jaguars and many other wildlife populations. We emphasize the global importance of the Pantanal wetland ecoregion as a key stronghold for long-term jaguar conservation.

Competition versus opportunism: Diet and trophic niche relationship of two sympatric apex carnivores in a tropical forest

There is a little understanding of how apex carnivores partition their diet to coexist. We studied food habit and trophic niche overlap of two apex carnivores, tiger and leopard in the tropical forest of Similipal tiger reserve, eastern India. We used line transect and scat analysis method, to estimate the prey availability and determine the diet and prey selection of two apex carnivores. Tigers consumed mostly large and medium-sized prey, whereas leopards mostly consumed medium and small-sized prey. Both carnivores were not random in their consumption of prey, sambar and wild pigs were selectively consumed by tigers, whereas leopards selectively consumed wild pigs, barking deer and mouse deer. Dietary overlap between two carnivore species was moderate (Pianka's niche overlap index: 0.55), and trophic niche breadth (Levin's standardized niche breadth) of the leopard (0.52) was greater than that of the tiger (0.37). Overall, tigers exhibited specialized feeding habits, whereas leopards showed generalist feeding habits. Our study highlights the opportunistic nature of leopards and probably a reason for the species successfully coexist with tigers.

Perceived risk structures the space use of competing carnivores

Abstract Competition structures ecological communities. In carnivorans, competitive interactions are disproportionately costly to subordinate carnivores who must account for the risk of interspecific killing when foraging. Accordingly, missed opportunity costs for meso-carnivores imposed by risk can benefit the smallest-bodied competitors. However, the extent to which the risk perpetuates into spatial partitioning in hierarchically structured communities remains unknown. To determine how risk-avoidance behaviors shape the space-use of carnivore communities, we studied a simple community of carnivores in northern Patagonia, Argentina: pumas (Puma concolor; an apex carnivore), culpeo foxes (Lycalopex culpaeus; a meso-carnivore), and chilla foxes (Lycalopex griseus; a small carnivore). We used multi-species occupancy models to quantify the space use within the carnivore community and giving-up densities to understand the behaviors that structure space use. Notably, we applied an analytical framework that tests whether the actual or perceived risk of predation most strongly influences the space use of subordinate carnivores although accounting for their foraging and vigilance behaviors. We found that there was a dominance hierarchy from the apex carnivore through the meso-carnivore to the subordinate small carnivore, which was reflected in space. Although both meso- and small carnivores exhibited similar predator avoidance behavioral responses to apex carnivores, the habitat associations of apex carnivores only altered meso-carnivore space use. The biases in risk management we observed for meso-carnivores likely translates into stable co-existence of this community of competing carnivores. We believe our analytical framework can be extended to other communities to quantify the spatial-behavioral tradeoffs of risk.

Ontogeny of scent marking behaviours in an apex carnivore

Abstract Puma (Puma concolor) communication with conspecifics is via indirect scent marking behaviours that are important for individuals to advertise their territory and reproductive status, but little is known about how the behaviours develop with age. To examine the development of scent marking behaviours, we monitored the behaviours of adult pumas and dependent kittens. Based on video recordings, we found that the frequency of puma communication behaviours significantly changed over time. Kittens exhibited olfactory investigation more frequently as they aged, but kittens generally did not exhibit scent marking behaviours. Kittens travel with their mothers until they disperse, so there is no need to establish territories or advertise availability to mate, but kittens are at risk of injury or mortality from other pumas. It is possible that there is no functional need for dependent kittens to scent mark until they mature, but there is a need for frequent use of investigative behaviours.

Low apex carnivore density does not release a subordinate competitor when driven by prey depletion

Conservation of competitively subordinate carnivores presents a difficult challenge because they are limited by dominant competitors. Prey depletion is one of the leading causes of large carnivore decline worldwide, but little is known about the net effect of prey depletion on subordinate carnivores when their dominant competitors are also reduced. African wild dogs are often limited by high densities of dominant competitors, particularly lions. We measured African wild dog density and survival, using mark-recapture models fit to 8 years of data from 425 known individuals in the Greater Kafue Ecosystem, Zambia. The GKE is affected by prey depletion, particularly of large herbivores, and thus the density of lions is significantly lower than ecologically comparable ecosystems. Counter to expectations from mesopredator release theory, wild dog density in GKE was far lower than comparable ecosystems with higher lion and prey density, though annual survival rates were comparable to large and stable populations. Average pack size was small and home range size was among the largest recorded. Our results show that low lion density did not competitively release the GKE wild dog population and we infer that the low density of wild dogs was a product of low prey density. Our results suggest that there is an optimal ratio of prey and competitors at which wild dogs achieve their highest densities. This finding has immediate implications for the conservation of the endangered African wild dog, and broad implications for the conservation of subordinate species affected by resource depletion and intraguild competition.

An apex carnivore's life history mediates a predator cascade.

Apex predators can shape communities via cascading top-down effects, but the degree to which such effects depend on predator life history traits is largely unknown. Within carnivore guilds, complex hierarchies of dominance facilitate coexistence, whereby subordinate species avoid dominant counterparts by partitioning space, time, or both. We investigated whether a major life history trait (hibernation) in an apex carnivore (black bears Ursus americanus) mediated its top-down effects on the spatio-temporal dynamics of three sympatric mesocarnivore species (coyotes Canis latrans, bobcats Lynx rufus, and gray foxes Urocyon cinereoargenteus) across a 15,000km2 landscape in the western USA. We compared top-down, bottom-up, and environmental effects on these mesocarnivores using an integrated modeling approach. Black bears exerted top-down effects that varied as a function of hibernation and were stronger than bottom-up or environmental impacts. High black bear activity in summer and fall appeared to buffer the most subordinate mesocarnivore (gray foxes) from competition with dominant mesocarnivores (coyotes and bobcats), which were in turn released by black bear hibernation in winter and early spring. The mesocarnivore responses occurred in space (i.e., altered occupancy and site visitation intensity) rather than time (i.e., diel activity patterns unaffected). These results suggest that the spatio-temporal dynamics of mesocarnivores in this system were principally shaped by a spatial predator cascade of interference competition mediated by black bear hibernation. Thus, certain life history traits of apex predators might facilitate coexistence among competing species over broad time scales, with complex implications for lower trophic levels.

Does management of a top carnivore influence the response of mesopredators and prey to rainfall in arid ecosystems? Evidence for a Baseline Density theory

The removal of apex carnivores from ecosystems can impact the abundance and diversity of species in lower trophic levels. In arid ecosystems, where “bottom up” forces of primary productivity and resource availability strongly affect trophic interactions, the role of “top down” effects is still much debated. This study explored the potential role of an apex predator, the dingo, as a “top down” trophic regulator in Australian arid ecosystems under different levels of primary productivity and dingo management regimes. Consistent with the theory of top down regulation, strong relationships were found between dingo management, dingo activity and fox activity. Dingoes appeared to suppress fox activity where dingoes were uncontrolled or only opportunistically controlled. At sites where dingoes were absent or in low numbers, fox activity was higher, and this inverse relationship persisted regardless of rainfall. The activity of rabbits and small mammals was lower where dingoes were absent and fox activity was high, while the activity of macropods was higher in the absence of dingoes. Feral cat activity did not differ significantly between sites under different dingo management or between years. These results suggest that management of dingoes is a key determinant of fox activity and the activity of some prey under varying levels of productivity. Evidence from this research showed that while the strength of trophic regulation by dingoes may fluctuate, top down effects occurred both prior to and post significant rainfall events. Following this, top down regulation of fox populations during dry periods at sites where dingoes are retained may enable higher and more stable “baseline” densities of small vertebrates, from which a larger and more rapid rate of increase of these prey during the “boom” periods can occur. Understanding the relative strength and interactions of top down and bottom up forces in regulating populations, and under what ecological states the importance of each changes, is important for the long-term conservation of biodiversity in arid regions.

Migration dynamics of polar bears (Ursus maritimus) in western Hudson Bay

Abstract Migration is predicted to change both spatially and temporally as climate change alters seasonal resource availability. Species in extreme environments are especially susceptible to climate change; hence, it is important to determine environmental and biological variables that influence their migration. Polar bears (Ursus maritimus) are an Arctic apex carnivore whose migration phenology has been affected by climate change and is vulnerable to future changes. Here, we used satellite-linked telemetry collar data from adult female polar bears in western Hudson Bay from 2004 to 2016 and multivariate response regression models to demonstrate that 1) spatial and temporal migration metrics are correlated, 2) ice concentration and wind are important environmental variables that influence polar bear migration in seasonal ice areas, and 3) migration did not vary across the years of our study, highlighting the importance of continued monitoring. Specifically, we found that ice concentration, wind speed, and wind direction affected polar bear migration onto ice during freeze-up and ice concentration and wind direction affected migration onto land during breakup. Bears departed from land earlier with increased wind speed and the effect of wind direction on migration may be linked to prey searching and ice drift. Low ice concentration was associated with higher movement during freeze-up and breakup. Our findings suggest that migration movement may increase in response to climate change as ice concentration and access to prey declines, potentially increasing nutritional stress on bears.

Spatial ecology of lions in a small, semi-fenced park surrounded by dense human populations: the case study of Nairobi National Park, Kenya

Abstract Human population growth near protected areas often results in detrimental edge effects for apex carnivores, such as the African lion. Urbanization leads to new scenarios of the human–lion conflict, thus understanding ranging patterns close to urban environments is crucial to inform future management strategies. We collected GPS data from 12 collared lions between 2014 and 2018 in Nairobi National Park, which borders the capital city of Kenya, Nairobi city. We estimated home ranges, calculated daily distance traveled and tested for differences between sex, season and pride. Additionally, we investigated how disturbance from Nairobi and surrounding human settlements affected space-use of lions, and tested for differences between sex, season and time of day. Lions showed restricted movements (4.5 km/day) and had small home ranges (49 km2). Male lions had larger ranges than females, but avoidance behavior of disturbed areas was similar. Lions took advantage during times of low human activity, i.e., during the night, to extend ranging behavior in search for resources. Risk for livestock depredation also increased during the wet season when lions roamed longer, more frequently, and deeper into the community lands. We recommend the establishment of buffer zones to maintain a viable lion population and reduced risk for conflict.

Secondary forest development during urbanization sustains apex carnivore populations of Eurasian otters (Lutra lutra)

Better understanding of the ecological and socioeconomic determinants of land-use change is a key to achieve more sustainable development and conservation objectives. Long-term studies on the ability of regenerated secondary forests to sustain biodiversity compared with primary forests have been little explored. South Korea has experienced rapid urbanization and rapid forest regeneration. In this study, we asked two questions: (1) Does secondary forest development alleviate the negative impacts of urbanization on an apex predator; and (2) Can we elucidate a land use planning balance between urbanization and biodiversity conservation using scenario analysis? We studied the Eurasian otter (Lutra lutra) population in South Korea. We applied generalized linear mixed models to five national otter surveys from 1994 to 2019 with 18 environmental variables. Based on the most parsimonious model, we developed 10 scenarios with three patterns of urbanization (dispersed, clustered, and no further urbanization) with or without forest development or with forest degeneration. The most parsimonious model showed that otter occurrences steadily increased with greater forest density, but were negatively correlated with the percentage of urban areas. The scenarios suggested that diffused urban development had a less negative impact on otter distribution than clustered urban areas. Developed forests considerably alleviated the negative impacts of urbanization. In sample sub-basins, scenarios showed that even without further urbanization, otters will take more than six years to occur with the current forest development trends. Hence, sustainable urban planning should include strategic spatial planning for forest development and preservation.

Artificial nightlight alters the predator–prey dynamics of an apex carnivore

Artificial nightlight is increasingly recognized as an important environmental disturbance that influences the habitats and fitness of numerous species. However, its effects on wide‐ranging vertebrates and their interactions remain unclear. Light pollution has the potential to amplify land‐use change, and as such, answering the question of how this sensory stimulant affects behavior and habitat use of species valued for their ecological roles and economic impacts is critical for conservation and land‐use planning. Here, we combined satellite‐derived estimates of light pollution, with GPS‐data from cougars Puma concolor (n = 56), mule deer Odocoileus hemionus (n = 263) and locations of cougar‐killed deer (n = 1562 carcasses), to assess the effects of light exposure on mammal behavior and predator–prey relationships across wildland–urban gradients in the southwestern United States. Our results indicate that deer used the anthropogenic environments to access forage and were more active at night than their wildland conspecifics. Despite higher nightlight levels, cougars killed deer at the wildland–urban interface, but hunted them in the relatively darkest locations. Light had the greatest effect of all covariates on where cougars killed deer at the wildland–urban interface. Both species exhibited functional responses to light pollution at fine scales; individual cougars and deer with less light exposure increasingly avoided illuminated areas when exposed to greater radiance, whereas deer living in the wildland–urban interface selected elevated light levels. We conclude that integrating estimates of light pollution into ecological studies provides crucial insights into how the dynamic human footprint can alter animal behavior and ecosystem function across spatial scales.

Using spatial characteristics of apex carnivore communication and reproductive behaviors to predict responses to future human development

A growing body of evidence has documented how wildlife alter their behavior in response to human encroachment. For carnivores, behaviors related to reproduction and communication are particularly sensitive to human disturbance and can provide an early warning indicator of development’s negative impacts. Despite the important role carnivores play in an ecosystem, few tools have been developed to anticipate how future human development impacts these behaviors. We developed a set of models to understand spatial relationships between anthropogenic development and puma (Puma concolor) habitat selection for two critical reproductive behaviors: nursery habitat for raising young, and sites for communication with mates. Using geospatial location data from the Santa Cruz Mountains in California, USA, we found that female pumas use small nursery home ranges (9 km2 ± 1.72 SE) of predominantly natural habitat, potentially with low levels of human development (< 1 housing unit per 40 acres), when supporting kittens < 8 weeks old. Areas immediately surrounding (≤ 600 m) puma communication sites were also almost entirely composed of undeveloped habitat or low-density development. When modeling projected human development compared to current land use, we found that increases in human development may eliminate 20% of current puma nursery habitat and nearly 50% of current communication site habitat. Future development will also increase the patchiness of suitable habitat, intensifying the difficulty of locating and accessing suitable areas for nurseries and communication. Focusing on the habitat needed to support reproductive and communication behaviors may be an effective way to prioritize conservation planning for pumas and other apex carnivores.

Assessing the performance of index calibration survey methods to monitor populations of wide-ranging low-density carnivores.

Apex carnivores are wide‐ranging, low‐density, hard to detect, and declining throughout most of their range, making population monitoring both critical and challenging. Rapid and inexpensive index calibration survey (ICS) methods have been developed to monitor large African carnivores. ICS methods assume constant detection probability and a predictable relationship between the index and the actual population of interest. The precision and utility of the resulting estimates from ICS methods have been questioned. We assessed the performance of one ICS method for large carnivores—track counts—with data from two long‐term studies of African lion populations. We conducted Monte Carlo simulation of intersections between transects (road segments) and lion movement paths (from GPS collar data) at varying survey intensities. Then, using the track count method we estimated population size and its confidence limits. We found that estimates either overstate precision or are too imprecise to be meaningful. Overstated precision stemmed from discarding the variance from population estimates when developing the method and from treating the conversion from tracks counts to population density as a back‐transformation, rather than applying the equation for the variance of a linear function. To effectively assess the status of species, the IUCN has set guidelines, and these should be integrated in survey designs. We propose reporting the half relative confidence interval width (HRCIW) as an easily calculable and interpretable measure of precision. We show that track counts do not adhere to IUCN criteria, and we argue that ICS methods for wide‐ranging low‐density species are unlikely to meet those criteria. Established, intensive methods lead to precise estimates, but some new approaches, like short, intensive, (spatial) capture–mark–recapture (CMR/SECR) studies, aided by camera trapping and/or genetic identification of individuals, hold promise. A handbook of best practices in monitoring populations of apex carnivores is strongly recommended.

Spatio-temporal coexistence of sympatric mesocarnivores with a single apex carnivore in a fine-scale landscape

Mesocarnivores uniquely and profoundly impact ecosystem function, structure, and dynamics. Sympatric species tend to spatially and temporally partition limited resources to facilitate coexistence. We investigated the seasonal spatial and temporal cooccurrences among six mesocarnivores, the leopard cat (Prionailurus bengalensis), red fox (Vulpes vulpes), Asian badger (Meles leucurus), Siberian weasel (Mustela sibirica), masked palm civet (Paguma larvata) and yellow-throated marten (Martes flavigula), as well as a single apex predator (Northern Chinese leopard, Panthera pardus japonensis). We used a camera trapping dataset collected from June 2016 to May 2017 (16,636 camera trapping days, 52 camera locations). The activity patterns varied among seasons and species. Most species were most active during summer. Leopards were most active in winter. Siberian weasels and yellow-throated martens were mainly diurnal, tended to spatially avoid each other, and were temporally segregated from the other mesocarnivores. Leopard cats, Asian badgers, red foxes and masked palm civets were nocturnal and showed high overlap in every season, but their highest peaks of activity were staggered. Mesocarnivores may be affected by the threat of the apex carnivore; they mainly avoided leopards spatially, showing low spatial overlap with leopards in all seasons. Interestingly, we found that the animals may engage in temporal-spatial coordination to facilitate coexistence, as increased temporal overlap in a given season was associated with decreased spatial overlap. Our results provide new insight into the carnivore community of terrestrial mammals in northern China and will facilitate future studies on the mechanisms determining the coexistence of animal species within the trophic cascade.

FIRST OBSERVATION OF A PUMA (PUMA CONCOLOR) FEEDING ON A PINNIPED NEAR DEEP COVE, BRITISH COLUMBIA

Apex carnivores, including Pumas (Puma concolor), play a key role in ecological communities. Pumas have variable foraging patterns across their extensive range and are known scavengers. A recent review of Puma diet and predation studies documented a total of 48 mammal prey types representing 9 of the 13 orders found in North America, but excluding marine mammals (Orders Cetacea and Sirenia, and Clade Pinnipedimorpha) or tapirs (Order Perissodactyla). This report documents what is, to the best of our knowledge, the 1st observation of a Puma feeding on a marine mammal, a Harbor Seal (Phoca vitulina). Pinnipeds, as hunted or scavenged prey, could be a significant resource for Pumas in coastal areas and should be a focus of future studies. Novel observations of prey items, such as this, are important to document as they expand our understanding of apex carnivores and the effects they have on ecological communities.

Do conservation strategies that increase tiger populations have consequences for other wild carnivores like leopards?

Most large carnivore populations are declining across their global range except in some well managed protected areas (PA’s). Investments for conserving charismatic apex carnivores are often justified due to their umbrella effect on biodiversity. We evaluate population trends of two large sympatric carnivores, the tiger and leopard through spatially-explicit-capture-recapture models from camera trap data in Kanha PA, India, from 2011 to 2016. Our results show that the overall density (100 km−2) of tigers ranged between 4.82 ± 0.33 to 5.21 ± 0.55SE and of leopards between 6.63 ± 0.71 to 8.64 ± 0.75SE, with no detectable trends at the PA scale. When evaluated at the catchment scale, Banjar catchment that had higher prey density and higher conservation investments, recorded significant growth of both carnivores. While Halon catchment, that had lower prey and conservation investments, populations of both carnivores remained stable. Sex ratio of both carnivores was female biased. As is typical with large carnivores, movement parameter sigma (an index for range size), was larger for males than for females. However, sigma was surprisingly similar for the same genders in both carnivores. At home-range scale, leopards achieved high densities and positive growth rates in areas that had low, medium or declining tiger density. Our results suggest that umbrella-species conservation value of tigers is likely to be compromised at very high densities and therefore should not be artificially inflated through targeted management.

Africa's apex predator, the lion, is limited by interference and exploitative competition with humans

Apex predators are crucial for maintaining ecological patterns and processes, yet humans hinder their ability to fulfil this role by displacing them from the landscape. Many apex predator species such as African lions (Panthera leo) are experiencing catastrophic declines as a result of competition with growing human populations. Increasing our understanding of the competitive interactions between lions and humans, as well as identifying thresholds of lion tolerance to human activities are important both for lion conservation and our understanding of apex predator ecology in the Anthropocene. We investigated the relative and cumulative influences of anthropogenic pressures on lion occurrence across a 73 000 km2 multi-use landscape in southern Africa. We developed occupancy models from replicated detection/non-detection spoor surveys across gradients of anthropogenic and biotic features. We tested the two hypotheses that African lions were most limited by 1) interference competition with humans or 2) exploitative competition with humans and evaluated the relative contribution of individual anthropogenic and biotic variables to lion occurrence. Our models predicted that lions occupied 49% of the landscape. The strongest determinants of lion occupancy were negative associations with pastoralism and bushmeat poaching, and a positive association with preferred prey. Thus, lions in this landscape are limited by a combination of interference and exploitative competition with poachers and pastoralists. However, interference competition with pastoralism was the biggest driver limiting lion occupancy, with a clear disturbance threshold for lions cumulating in a near complete loss of lions from the landscape when cattle surpass 21% occurrence. This study provides a predictive understanding of the top-down impacts of humans on the world's vulnerable apex carnivores.

Cats, canines, and coexistence: dietary differentiation between the sympatric Snow Leopard and Grey Wolf in the western landscape of Nepal Himalaya

Understanding the dietary habits of sympatric apex carnivores advances our knowledge of ecological processes and aids their conservation. We compared the diets of the sympatric Snow Leopard Panthera uncia and Grey Wolf Canis lupus using standard micro-histological analyses of scats collected from the western complex of Nepal Himalaya. Our study revealed one of the highest recorded contributions of livestock to the diet of top predators (55% for Grey Wolf and 39% for Snow Leopard) and high dietary overlap (0.82) indicating potential exploitative or interference competition. Their diet composition, however, varied significantly based on their consumption of wild and domestic prey. Limitation in data precludes predicting direction and outcome of inter-specific interactions between these predators. Our findings suggest a high rate of negative interaction with humans in the region and plausibly retaliatory killings of these imperilled predators. To ensure the sustained survival of these two apex carnivores, conservation measures should enhance populations of their wild prey species while reducing livestock losses of the local community through preventive and mitigative interventions.

Trophic ecology of large herbivores in a reassembling African ecosystem

Abstract Megafauna assemblages have declined or disappeared throughout much of the world, and many efforts are underway to restore them. Understanding the trophic ecology of such reassembling systems is necessary for predicting recovery dynamics, guiding management, and testing general theory. Yet, there are few studies of recovering large‐mammal communities, and fewer still that have characterized food‐web structure with high taxonomic resolution. In Gorongosa National Park, large herbivores have rebounded from near‐extirpation following the Mozambican Civil War (1977–1992). However, contemporary community structure differs radically from the prewar baseline: medium‐sized ungulates now outnumber larger bodied species, and several apex carnivores remain locally extinct. We used DNA metabarcoding to quantify diet composition of Gorongosa’s 14 most abundant large‐mammal populations. We tested five hypotheses: (i) the most abundant populations exhibit greatest individual‐level dietary variability; (ii) these populations also have the greatest total niche width (dietary diversity); (iii) interspecific niche overlap is high, with the diets of less‐abundant species nested within those of more‐abundant species; (iv) partitioning of forage species is stronger in more structurally heterogeneous habitats; and (v) selectivity for plant taxa converges within guilds and digestive types, but diverges across them. Abundant (and narrow‐mouthed) populations exhibited higher among‐individual dietary variation, but not necessarily the greatest dietary diversity. Interspecific dietary overlap was high, especially among grazers and in structurally homogenous habitat, whereas niche separation was more pronounced among browsers and in heterogeneous habitat. Patterns of selectivity were similar for ruminants—grazers and browsers alike—but differed between ruminants and non‐ruminants. Synthesis. The structure of this recovering food web was consistent with several hypotheses predicated on competition, habitat complexity, and herbivore traits, but it differed from patterns observed in more intact assemblages. We propose that intraspecific competition in the fastest‐recovering populations has promoted individual variation and a more nested food web, wherein rare species use subsets of foods eaten by abundant species, and that this scenario is reinforced by weak predation pressure. Future work should test these conjectures and analyse how the taxonomic dietary niche axis studied here interacts with other mechanisms of diet partitioning to affect community reassembly following wildlife declines.

Large species within carnivora are large carnivores.

Wolf & Ripple [1] is a global study that explored the potential for large carnivore reintroductions, identifying protected areas and other regions with low human footprint that warrant further investigation as possible locations for rewilding. In his comment, Miranda [2] raised a number of points related to Wolf & Ripple [1]. His primary points are that Wolf and Ripple (i) used terminology (carnivore, apex predator, and guild) incorrectly, (ii) selected an inappropriate group of species to study and (iii) listed some unsuitable sites for large carnivore reintroductions. However, each of these criticisms is fundamentally flawed and only acts to hinder conservation efforts. Miranda stated that the species selected by Wolf and Ripple ‘were neither necessarily apex predators nor carnivores, but a subset of the mammalian order Carnivora, […]'. However, Wolf and Ripple did not claim that the species being studied are apex predators. In fact, the term ‘apex predator' does not appear anywhere in the main text or supplement of Wolf and Ripple. The species selected included 25 large carnivore species with a body mass greater than or equal to 15 kg that have relatively accurate historic and current range maps. The scope and purpose of Wolf and Ripple was to identify potential reintroduction sites for large-bodied members of order Carnivora without regard for whether each individual species was an apex predator, which is in itself context dependent. For example, coyotes ( Canis latrans ) may be considered mesopredators when wolves are present, but apex predators when wolves are absent [3]. Had Wolf and Ripple focused on potential reintroductions of a different taxonomic order, such as Primates, would Miranda then critique that the paper did not focus enough on other groups of vertebrates? Miranda is incorrect in his claim that the species selected are …