Christopher Wolf

Collapse of the world's largest herbivores.

The collapsing populations of large herbivores will have extensive ecological and social consequences. Large wild herbivores are crucial to ecosystems and human societies. We highlight the 74 largest terrestrial herbivore species on Earth (body mass ≥100 kg), the threats they face, their important and often overlooked ecosystem effects, and the conservation efforts needed to save them and their predators from extinction. Large herbivores are generally facing dramatic population declines and range contractions, such that ~60% are threatened with extinction. Nearly all threatened species are in developing countries, where major threats include hunting, land-use change, and resource depression by livestock. Loss of large herbivores can have cascading effects on other species including large carnivores, scavengers, mesoherbivores, small mammals, and ecological processes involving vegetation, hydrology, nutrient cycling, and fire regimes. The rate of large herbivore decline suggests that ever-larger swaths of the world will soon lack many of the vital ecological services these animals provide, resulting in enormous ecological and social costs.

Saving the World's Terrestrial Megafauna

From the late Pleistocene to the Holocene, and now the so called Anthropocene, humans have been driving an ongoing series of species declines and extinctions (Dirzo et al. 2014). Large-bodied mammals are typically at a higher risk of extinction than smaller ones (Cardillo et al. 2005). However, in some circumstances terrestrial megafauna populations have been able to recover some of their lost numbers due to strong conservation and political commitment, and human cultural changes (Chapron et al. 2014). Indeed many would be in considerably worse predicaments in the absence of conservation action (Hoffmann et al. 2015). Nevertheless, most mammalian megafauna face dramatic range contractions and population declines. In fact, 59% of the world’s largest carnivores (≥ 15 kg, n = 27) and 60% of the world’s largest herbivores (≥ 100 kg, n = 74) are classified as threatened with extinction on the International Union for the Conservation of Nature (IUCN) Red List (supplemental table S1 and S2). This situation is particularly dire in sub-Saharan Africa and Southeast Asia, home to the greatest diversity of extant megafauna (figure 1). Species at risk of extinction include some of the world’s most iconic animals—such as gorillas, rhinos, and big cats (figure 2 top row)—and, unfortunately, they are vanishing just as science is discovering their essential ecological roles (Estes et al. 2011). Here, our objectives are to raise awareness of how these megafauna are imperiled (species in supplemental table S1 and S2) and to stimulate broad interest in developing specific recommendations and concerted action to conserve them.

Bushmeat hunting and extinction risk to the world’s mammals

Terrestrial mammals are experiencing a massive collapse in their population sizes and geographical ranges around the world, but many of the drivers, patterns and consequences of this decline remain poorly understood. Here we provide an analysis showing that bushmeat hunting for mostly food and medicinal products is driving a global crisis whereby 301 terrestrial mammal species are threatened with extinction. Nearly all of these threatened species occur in developing countries where major coexisting threats include deforestation, agricultural expansion, human encroachment and competition with livestock. The unrelenting decline of mammals suggests many vital ecological and socio-economic services that these species provide will be lost, potentially changing ecosystems irrevocably. We discuss options and current obstacles to achieving effective conservation, alongside consequences of failure to stem such anthropogenic mammalian extirpation. We propose a multi-pronged conservation strategy to help save threatened mammals from immediate extinction and avoid a collapse of food security for hundreds of millions of people.

What is a Trophic Cascade

Few concepts in ecology have been so influential as that of the trophic cascade. Since the 1980s, the term has been a central or major theme of more than 2000 scientific articles. Despite this importance and widespread usage, basic questions remain about what constitutes a trophic cascade. Inconsistent usage of language impedes scientific progress and the utility of scientific concepts in management and conservation. Herein, we offer a definition of trophic cascade that is designed to be both widely applicable yet explicit enough to exclude extraneous interactions. We discuss our proposed definition and its implications, and define important related terms, thereby providing a common language for scientists, policy makers, conservationists, and other stakeholders with an interest in trophic cascades.

Global forest loss disproportionately erodes biodiversity in intact landscapes

Global biodiversity loss is a critical environmental crisis, yet the lack of spatial data on biodiversity threats has hindered conservation strategies. Theory predicts that abrupt biodiversity declines are most likely to occur when habitat availability is reduced to very low levels in the landscape (10–30%). Alternatively, recent evidence indicates that biodiversity is best conserved by minimizing human intrusion into intact and relatively unfragmented landscapes. Here we use recently available forest loss data to test deforestation effects on International Union for Conservation of Nature Red List categories of extinction risk for 19,432 vertebrate species worldwide. As expected, deforestation substantially increased the odds of a species being listed as threatened, undergoing recent upgrading to a higher threat category and exhibiting declining populations. More importantly, we show that these risks were disproportionately high in relatively intact landscapes; even minimal deforestation has had severe consequences for vertebrate biodiversity. We found little support for the alternative hypothesis that forest loss is most detrimental in already fragmented landscapes. Spatial analysis revealed high-risk hot spots in Borneo, the central Amazon and the Congo Basin. In these regions, our model predicts that 121–219 species will become threatened under current rates of forest loss over the next 30 years. Given that only 17.9% of these high-risk areas are formally protected and only 8.9% have strict protection, new large-scale conservation efforts to protect intact forests are necessary to slow deforestation rates and to avert a new wave of global extinctions.

Prey depletion as a threat to the world's large carnivores

Large terrestrial carnivores are an ecologically important, charismatic and highly endangered group of species. Here, we assess the importance of prey depletion as a driver of large carnivore endangerment globally using lists of prey species for each large carnivore compiled from the literature. We consider spatial variation in prey endangerment, changes in endangerment over time and the causes of prey depletion, finding considerable evidence that loss of prey base is a major and wide-ranging threat among large carnivore species. In particular, the clouded leopard (Neofelis nebulosa), Sunda clouded leopard (Neofelis diardi), tiger (Panthera tigris), dhole (Cuon alpinus) and Ethiopian wolf (Canis simensis) all have at least 40% of their prey classified as threatened on the International Union for the Conservation of Nature (IUCN) Red List and, along with the leopard (Panethra pardus), all of these species except the Ethiopian wolf have at least 50% of their prey classified as declining. Of the 494 prey species in our analysis, an average of just 6.9% of their ranges overlap protected areas. Together these results show the importance of a holistic approach to conservation that involves protecting both large carnivores directly and the prey upon which they depend.

Extinction risk is most acute for the world’s largest and smallest vertebrates

Significance Determining the drivers of extinction risk has been a key pursuit of conservation biology. Considering that body mass could be a strong predictor of extinction risk, we constructed a global database of body masses for 27,647 vertebrate species. Results show that the smallest- and largest-bodied vertebrates have elevated extinction risk. The largest vertebrates are mostly threatened by direct killing by humans, whereas the smallest species are more likely to have restricted geographic ranges—an important predictor of extinction risk—and be threatened by habitat degradation. Declines of large and small vertebrate species will truncate the size distributions characterizing these taxa, jeopardizing ecosystem services to humans, and generating cascading ecological and evolutionary effects on other species and processes. Extinction risk in vertebrates has been linked to large body size, but this putative relationship has only been explored for select taxa, with variable results. Using a newly assembled and taxonomically expansive database, we analyzed the relationships between extinction risk and body mass (27,647 species) and between extinction risk and range size (21,294 species) for vertebrates across six main classes. We found that the probability of being threatened was positively and significantly related to body mass for birds, cartilaginous fishes, and mammals. Bimodal relationships were evident for amphibians, reptiles, and bony fishes. Most importantly, a bimodal relationship was found across all vertebrates such that extinction risk changes around a body mass breakpoint of 0.035 kg, indicating that the lightest and heaviest vertebrates have elevated extinction risk. We also found range size to be an important predictor of the probability of being threatened, with strong negative relationships across nearly all taxa. A review of the drivers of extinction risk revealed that the heaviest vertebrates are most threatened by direct killing by humans. By contrast, the lightest vertebrates are most threatened by habitat loss and modification stemming especially from pollution, agricultural cropping, and logging. Our results offer insight into halting the ongoing wave of vertebrate extinctions by revealing the vulnerability of large and small taxa, and identifying size-specific threats. Moreover, they indicate that, without intervention, anthropogenic activities will soon precipitate a double truncation of the size distribution of the world’s vertebrates, fundamentally reordering the structure of life on our planet.

Range contractions of the world's large carnivores

The majority of the worlds terrestrial large carnivores have undergone substantial range contractions and many of these species are currently threatened with extinction. However, there has been little effort to fully quantify the extent of large carnivore range contractions, which hinders our ability to understand the roles and relative drivers of such trends. Here we present and analyse a newly constructed and comprehensive set of large carnivore range contraction maps. We reveal the extent to which ranges have contracted since historical times and identify regions and biomes where range contractions have been particularly large. In summary, large carnivores that have experienced the greatest range contractions include the red wolf (Canis rufus) (greater than 99%), Ethiopian wolf (Canis simensis) (99%), tiger (Panthera tigris) (95%) and lion (Panthera leo) (94%). In general, the greatest range contractions occurred in Southeastern Asia and Africa. Motivated by the ecological importance of intact large carnivore guilds, we also examined the spatial extent of intact large carnivore guilds both for the entire world and regionally. We found that intact carnivore guilds occupy just 34% of the worlds land area. This compares to 96% in historic times. Spatial modelling of range contractions showed that contractions were significantly more likely in regions with high rural human population density, cattle density or cropland. Our results offer new insights into how best to prevent further range contractions for the worlds largest carnivores, which will assist efforts to conserve these species and their important ecological effects.

Bayesian characterization of uncertainty in species interaction strengths

Considerable effort has been devoted to the estimation of species interaction strengths. This effort has focused primarily on statistical significance testing and obtaining point estimates of parameters that contribute to interaction strength magnitudes, leaving the characterization of uncertainty associated with those estimates unconsidered. We consider a means of characterizing the uncertainty of a generalist predator’s interaction strengths by formulating an observational method for estimating a predator’s prey-specific per capita attack rates as a Bayesian statistical model. This formulation permits the explicit incorporation of multiple sources of uncertainty. A key insight is the informative nature of several so-called non-informative priors that have been used in modeling the sparse data typical of predator feeding surveys. We introduce to ecology a new neutral prior and provide evidence for its superior performance. We use a case study to consider the attack rates in a New Zealand intertidal whelk predator, and we illustrate not only that Bayesian point estimates can be made to correspond with those obtained by frequentist approaches, but also that estimation uncertainty as described by 95% intervals is more useful and biologically realistic using the Bayesian method. In particular, unlike in bootstrap confidence intervals, the lower bounds of the Bayesian posterior intervals for attack rates do not include zero when a predator–prey interaction is in fact observed. We conclude that the Bayesian framework provides a straightforward, probabilistic characterization of interaction strength uncertainty, enabling future considerations of both the deterministic and stochastic drivers of interaction strength and their impact on food webs.

Rewilding the world's large carnivores

Earths terrestrial large carnivores form a highly endangered group of species with unique conservation challenges. The majority of these species have experienced major geographical range contractions, which puts many of them at high risk of extinction or of becoming ecologically ineffective. As a result of these range contractions and the associated loss of intact predator guilds, the ecological effects of these species are now far less widespread and common, with inevitable consequences for ecosystem function. Rewilding—which includes reintroducing species into portions of their former ranges—is an important carnivore conservation tool and means for restoring top-down ecological regulation. We conducted a global analysis of potential reintroduction areas. We first considered protected areas where one or more large carnivore species have been extirpated, identifying a total of 130 protected areas that may be most suitable for carnivore reintroduction. These protected areas include sites in every major world region, and are most commonly found in Mongolia (n = 13), Canada (n = 11), Thailand (n = 9), Namibia (n = 6), Indonesia (n = 6) and Australia (n = 6). We considered the sizes of protected areas, their levels of protection, the extent of human impacts within and around the protected areas, and the status of prey species in the protected areas. Finally, we used the ‘last of the wild’ approach to identify contiguous low human footprint regions within the former ranges of each species, identifying an additional 150 areas which could be the focus of conservation efforts to create conditions conducive to reintroductions. These low footprint regions were most commonly found in the USA (n = 14), Russia (n = 14), Canada (n = 10), China (n = 9) and Mauritania (n = 8). Together, our results show the global-scale potential for carnivore rewilding projects to both conserve these species and provide critical ecological and social benefits.