William J. Ripple

Trophic Downgrading of Planet Earth

Until recently, large apex consumers were ubiquitous across the globe and had been for millions of years. The loss of these animals may be humankind’s most pervasive influence on nature. Although such losses are widely viewed as an ethical and aesthetic problem, recent research reveals extensive cascading effects of their disappearance in marine, terrestrial, and freshwater ecosystems worldwide. This empirical work supports long-standing theory about the role of top-down forcing in ecosystems but also highlights the unanticipated impacts of trophic cascades on processes as diverse as the dynamics of disease, wildfire, carbon sequestration, invasive species, and biogeochemical cycles. These findings emphasize the urgent need for interdisciplinary research to forecast the effects of trophic downgrading on process, function, and resilience in global ecosystems.

Status and Ecological Effects of the World’s Largest Carnivores

Background The largest terrestrial species in the order Carnivora are wide-ranging and rare because of their positions at the top of food webs. They are some of the world’s most admired mammals and, ironically, some of the most imperiled. Most have experienced substantial population declines and range contractions throughout the world during the past two centuries. Because of the high metabolic demands that come with endothermy and large body size, these carnivores often require large prey and expansive habitats. These food requirements and wide-ranging behavior often bring them into conflict with humans and livestock. This, in addition to human intolerance, renders them vulnerable to extinction. Large carnivores face enormous threats that have caused massive declines in their populations and geographic ranges, including habitat loss and degradation,persecution, utilization, and depletion of prey. We highlight how these threats can affect theconservation status and ecological roles of this planet’s 31 largest carnivores. Ecologically important carnivores. Seven species of large carnivores with documented ecological effects involving (A) “tri-trophic cascades” from large carnivores to prey to plants, (B) “mesopredator cascades” from large carnivores to mesopredators to prey of mesopredators, and (C) both tri-trophic and mesopredator cascades. [Photo credits: sea otter (N. Smith), puma (W. Ripple), lion (K. Abley), leopard (A. Dey), Eurasianlynx (B. Elmhagen), dingo (A. McNab), gray wolf (D. Mclaughlin)] Advances Based on empirical studies, trophic cascades have been documented for 7 of the 31 largest mammalian carnivores (not including pinnipeds). For each of these species (see figure), human actions have both caused declines and contributed to recovery, providing “natural experiments” for quantifying their effects on food-web and community structure. Large carnivores deliver economic and ecosystem services via direct and indirect pathways that help maintain mammal, avian, invertebrate,and herpetofauna abundance or richness. Further, they affect other ecosystem processes and conditions, such as scavenger subsidies, disease dynamics, carbon storage, stream morphology, and crop production. The maintenance or recovery of ecologically effective densities of large carnivores is an important tool for maintaining the structure and function of diverse ecosystems. Outlook Current ecological knowledge indicates that large carnivores are necessary for the maintenanceof biodiversity and ecosystem function. Human actions cannot fully replace the role of large carnivores. Additionally, the future of increasing human resource demands and changing climate will affect biodiversity and ecosystem resiliency. These facts, combined with the importance of resiliente cosystems, indicate that large carnivores and their habitats should be maintained and restored wherever possible. Preventing the extinction of these species and the loss of their irreplaceable ecological function and importance will require novel, bold, and deliberate actions. We propose a Global Large Carnivore Initiative to coordinate local, national, and international research, conservation, and policy. Preserving Predators Large-bodied animals play essential roles in ecosystem structuring and stability through both indirect and direct trophic effects. In recent times, humans have disrupted this trophic structure through both habitat destruction and active extirpation of large predators, resulting in large declines in numbers and vast contractions in their geographic ranges. Ripple et al. (10.1126/science.1241484; see the Perspective by Roberts) review the status, threats, and ecological importance of the 31 largest mammalian carnivores globally. These species are responsible for a suite of direct and indirect stabilizing effects in ecosystems. Current levels of decline are likely to result in ecologically ineffective population densities and can lead to ecosystem instability. The preservation of large carnivores can be challenging because of their need for large ranges and their potential for human conflict. However, the authors demonstrate that the preservation of large carnivores is ecologically important and that the need for conservation action is immediate, given the severity of the threats they face. Large carnivores face serious threats and are experiencing massive declines in their populations and geographic ranges around the world. We highlight how these threats have affected the conservation status and ecological functioning of the 31 largest mammalian carnivores on Earth. Consistent with theory, empirical studies increasingly show that large carnivores have substantial effects on the structure and function of diverse ecosystems. Significant cascading trophic interactions, mediated by their prey or sympatric mesopredators, arise when some of these carnivores are extirpated from or repatriated to ecosystems. Unexpected effects of trophic cascades on various taxa and processes include changes to bird, mammal, invertebrate, and herpetofauna abundance or richness; subsidies to scavengers; altered disease dynamics; carbon sequestration; modified stream morphology; and crop damage. Promoting tolerance and coexistence with large carnivores is a crucial societal challenge that will ultimately determine the fate of Earth’s largest carnivores and all that depends upon them, including humans.

Wolves and the Ecology of Fear: Can Predation Risk Structure Ecosystems?

Abstract We investigated how large carnivores, herbivores, and plants may be linked to the maintenance of native species biodiversity through trophic cascades. The extirpation of wolves (Canis lupus) from Yellowstone National Park in the mid-1920s and their reintroduction in 1995 provided the opportunity to examine the cascading effects of carnivore–herbivore interactions on woody browse species, as well as ecological responses involving riparian functions, beaver (Castor canadensis) populations, and general food webs. Our results indicate that predation risk may have profound effects on the structure of ecosystems and is an important constituent of native biodiversity. Our conclusions are based on theory involving trophic cascades, predation risk, and optimal foraging; on the research literature; and on our own recent studies in Yellowstone National Park. Additional research is needed to understand how the lethal effects of predation interact with its nonlethal effects to structure ecosystems.

The Rise of the Mesopredator

Apex predators have experienced catastrophic declines throughout the world as a result of human persecution and habitat loss. These collapses in top predator populations are commonly associated with dramatic increases in the abundance of smaller predators. Known as “mesopredator release,” this trophic interaction has been recorded across a range of communities and ecosystems. Mesopredator outbreaks often lead to declining prey populations, sometimes destabilizing communities and driving local extinctions. We present an overview of mesopredator release and illustrate how its underlying concepts can be used to improve predator management in an increasingly fragmented world. We also examine shifts in North American carnivore ranges during the past 200 years and show that 60% of mesopredator ranges have expanded, whereas all apex predator ranges have contracted. The need to understand how best to predict and manage mesopredator release is urgent—mesopredator outbreaks are causing high ecological, economic, and social costs around the world.

Trophic cascades among wolves, elk and aspen on Yellowstone National Park's northern range

Quaking aspen (Populus tremuloides) biomass has declined in Yellowstone National Park (YNP) in the past century. We installed permanent belt transects (plots) for long-term monitoring of aspen stands both within and outside of established wolf pack territories on YNP’s northern range to determine if reintroduced wolves are influencing elk browsing patterns and aspen regeneration through a trophic cascades interaction. Wolves may have an indirect effect on aspen regeneration by altering elk movements, browsing patterns, and foraging behavior (predation risk effects). Elk pellet groups, aspen sucker heights, and the percentage of browsed suckers were the variables used to measure differences in aspen stands in high and low wolf-use areas of the northern range. The aspen stands in the high wolf-use areas had significantly lower counts of elk pellet groups in the mesic upland steppe and the combined mesic upland steppe and riparian/wet meadow habitat types. Based on our pellet group results, it appears that elk foraging behaviors may have been altered by the increased risk of predation due to the reintroduction of the wolf. In the riparian/ wet meadow habitat type, mean aspen sucker heights were significantly higher in the high wolf-use areas than in the low wolf-use areas. The percentage of browsed suckers in high and low wolf-use areas showed no significant differences in any of the habitat types. Considering the high browsing pressure in YNP aspen stands, it is uncertain whether the taller aspen suckers measured in the high wolf-use areas will eventually join the aspen overstory. These permanent plots represent a valuable baseline data set to assess any current and future aspen regeneration responses to the reintroduction of wolves in YNP. # 2001 Elsevier Science Ltd. All rights reserved.

Dynamics and Pattern of a Managed Coniferous Forest Landscape in Oregon

We examined the process of fragmentation in a managed forest landscape by comparing rates and patterns of disturbance (primarily clear-cutting) and regrowth between 1972 and 1988 using Landsat imagery. A 2589-km2 managed forest landscape in western Oregon was classified into two forest types, closed-canopy conifer forest (CF) (typically, >60% conifer cover) and other forest and nonforest types (OT) (typically, 914 m) maintained a greater percentage of CF than lower elevations (<914 m). The percentage of the area at the edge of the two cover types increased on all ownerships and in both elevational zones, whereas the amount of interior habitat (defined as CF at least 100 m from OT) decreased on all ownerships and elevational zones. By 1988 public lands contained - 45% interior habitat while private lands had 12% interior habitat. Mean interior patch area declined from 160 to 62 ha. The annual rate of disturbance (primarily clear-cutting) for the entire area including the wilderness was 1.19%, which corresponds to a cutting rotation of 84 yr. The forest landscape was not in a steady state or regulated condition which is not projected to occur for at least 40 yr under current forest plans. Variability in cutting rates within ownerships was higher on private land than on nonreserve public land. However, despite the use of dispersed cutting patterns on public land, spatial patterns of cutting and remnant forest patches were nonuniform across the entire public ownership. Large remaining patches (< 5000 ha) of contiguous interior forest were restricted to public lands designated for uses other than timber production such as wilderness areas and research natural areas.

Historic aspen recruitment, elk, and wolves in northern Yellowstone National Park, USA

We conducted an analysis of aspen (Populus tremuloides) overstory recruitment on the northern range of Yellowstone National Park (YNP) using information provided in a monograph published by Warren (Warren, E.R., 1926. A study of beaver in the Yaney region of Yellowstone National Park, Roosevelt-Wildl. Ann. 1, 1‐191), increment cores collected from riparian aspen stands in 1998, and an extensive random sample of aspen increment cores collected over YNP’s entire northern range in 1997 and 1998. We summarized aspen size classes reported by Warren and estimated overstory origination dates of the stands he described using a linear regression based on our riparian aspen diameter/age relationship. Applying our regression results to Warren’s diameter measurements, we predicted that the stands measured by Warren contained aspen that originated between approximately 1751 and 1920. The random set of aspen increment cores were used to analyze the age distribution of the current aspen overstory on YNP’s northern range. These increment core data showed that approximately 10% of the current overstory aspen originated before 1871, 85% between 1871 and 1920, and 5% after 1921. Based upon our analysis of the Warren data and our aspen increment cores, we conclude that successful aspen overstory recruitment occurred on the northern range of YNP from the middle to late 1700’s until the 1920’s, after which it essentially ceased. Rocky Mountain Elk (Cervus elaphus) browsing has been identified as significantly impacting aspen overstory recruitment on YNP’s northern range. We hypothesized why elk browsing has a diAerent influence on aspen now than it did historically. We discussed several potential social and ecological factors and hypothesize that a main factor is YNP’s loss of significant predator/prey relationships in the early 1900’s, especially the influence of gray wolves (Canis lupus). We found that aspen overstory recruitment ceased during the same years that wolves, a significant source of elk predation, were removed from YNP. Wolves may positively influence aspen overstory recruitment through a trophic cascades eAect by reducing elk populations, modifying elk movement, and changing elk browsing patterns on aspen. # 2000 Elsevier Science Ltd. All rights reserved.

Measuring forest landscape patterns in the cascade range of Oregon, USA

Abstract This paper describes the use of a set of spatial statistics to quantify the landscape pattern caused by the patchwork of clearcuts made over a 15-year period in the western Cascades of Oregon. Fifteen areas were selected at random to represent a diversity of landscape fragmentation patterns. Managed forest stands (patches) were digitized and analysed to produce both tobular and mapped information describing patch size, shape, abundance and spacing, and matrix characteristics of a given area. In addition, a GIS fragmentation index was developed which was found to be sensitive to patch abundance and to the spatial distribution of patches. Use of the GIS-derived index provides an automated method of determining the level of forest fragmentation and can be used to facilitate spatial analysis of the landscape for later coordination with field and remotely sensed data. A comparison of the spatial statistics calculated for the two years indicates an increase in forest fragmentation as characterized by an increase in mean patch abundance and a decrease in interpatch distance, amount of interior natural forest habitat, and the GIS fragmentation index. Such statistics capable of quantifying patch shape and spatial distribution may prove important in the evaluation of the changing character of interior and edge habitats for wildlife.

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.

MODELING OF SITE OCCUPANCY DYNAMICS FOR NORTHERN SPOTTED OWLS, WITH EMPHASIS ON THE EFFECTS OF BARRED OWLS

Abstract Northern spotted owls (Strix occidentalis caurina) have been studied intensively since their listing as a threatened species by the U.S. Fish and Wildlife Service in 1990. Studies of spotted owl site occupancy have used various binary response measures, but most of these studies have made the assumption that detectability is perfect, or at least high and not variable. Further, previous studies did not consider temporal variation in site occupancy. We used relatively new methods for open population modeling of site occupancy that incorporated imperfect and variable detectability of spotted owls and allowed modeling of temporal variation in site occupancy, extinction, and colonization probabilities. We also examined the effects of barred owl (S. varia) presence on these parameters. We used spotted owl survey data from 1990 to 2002 for 3 study areas in Oregon, USA, and we used program MARK to develop and analyze site occupancy models. We found per visit detection probabilities averaged <0.70 and were highly variable among study years and study areas. Site occupancy probabilities for owl pairs declined greatly on 1 study area and slightly on the other 2 areas. For all owls, including singles and pairs, site occupancy was mostly stable through time. Barred owl presence had a negative effect on spotted owl detection probabilities, and it had either a positive effect on local-extinction probabilities or a negative effect on colonization probabilities. We conclude that further analyses of spotted owls must account for imperfect and variable detectability and barred owl presence to properly interpret results. Further, because barred owl presence is increasing within the range of northern spotted owls, we expect to see further declines in the proportion of sites occupied by spotted owls.