It is a rule in ecology that big animals outcompete little animals. Sometimes the big animals kill the little animals, sometimes the big animals eat the little animals, and sometimes the big animals drive the little animals out of one territory and into another, safer one. That basic pattern – “interspecific competitive killing” – has pushed scientists to try to understand how large carnivores shape entire ecosystems. Continue reading

When Predators Vanish, So Does the Ecosystem

By Carl Zimmer

Mark D. Bertness, an ecologist at Brown University, began studying the salt marshes of New England in 1981. Twenty-six years later, in 2007, he started to watch them die. In one marsh after another, lush stretches of cordgrass disappeared, replaced by bare ground. The die-offs were wiping out salt marshes in just a few years.

“It’s unbelievable how quickly it’s moved in,” Dr. Bertness said.

Scientists have been witnessing a similar transformation in a number of plant species along coastlines in the United States and in other countries. And in many cases, it’s been hard to pinpoint the cause of the die-off, with fungal outbreaks, pollution, choking sediments stirred up by boats, and rising sea levels proposed as killers.

There is much at stake in the hunt for the culprit, because salt marshes are hugely important. They shield coasts from flooding, pull pollutants from water and are nurseries for many fish species.

Certain species of herbivore crab, such as this purple marsh crab, are proliferating in salt marshes where humans are trapping and fishing their main predators. CreditTyler Coverdale

In the journal Ecology Letters, Dr. Bertness and his colleagues have nowpublished an experiment that may help solve the mystery. The evidence, they say, points to recreational fishing and crabbing. A fisherman idly dangling a line off a dock may not appear to be an agent of ecological collapse. But fishing removes the top predators from salt marshes, and the effects may be devastating.

Once New England salt marshes started dying off, Dr. Bertness and his colleagues embarked on a broad survey. Quickly they noticed a difference between healthy marshes and sick ones. The dying marshes tended to be near docks, marinas or buoys where boats could anchor, or where there were other signs of fishing.

“It wasn’t a brilliant thing we thought of sitting around the lab,” Dr. Bertness said. “By the time we got to 10 marshes, we realized there was this huge disparity.”

Dr. Bertness and his colleagues wondered how fishing and crabbing were affecting the food webs of the salt marshes. If people pull out striped bass and blue crabs and other predators from a salt marsh, the animals’ prey species — including those that feed on plants, like marsh crabs — are left to thrive. A growing population of marsh crabs might wipe out the cordgrass in a marsh. Without the roots of the cordgrass to anchor the soil, the marsh would erode, making it harder for new plants to grow.

To test this idea, Dr. Bertness and his colleagues surveyed salt marshes in Narragansett Bay in Rhode Island, comparing marshes that were healthy with ones that were almost entirely dead. The scientists found that in dying marshes, the plants had more signs of being fed on by crabs. And when they looked for other proposed causes of marsh die-off, such as pollution, they didn’t find a correlation. They published their results in March in the journal PLOS One.

Next, the scientists took a step beyond simply observing the die-offs: They tried to cause them. If the predator hypothesis was right, then creating a predator-free salt marsh habitat should lead to the disappearance of cordgrass.

In May 2013, the scientists installed cages in a healthy salt marsh on Cape Cod. Each cage was three feet on a side, with mesh walls and an open bottom. Marsh crabs could feed on the cordgrass inside the cages by burrowing up through the mud, and the wire mesh walls protected them from predators like fish and blue crabs.

The experiment quickly yielded results. In a matter of weeks, the cages were crowded with marsh crabs, and much of the cordgrass inside the cages was dying off. “We were planning on it being a two- or three-year experiment,” Dr. Bertness said. “But by the beginning of July, I thought, ‘My God, this is really going fast.’”

William J. Ripple, an ecologist at Oregon State University who was not involved in the research, said, “This is an important new scientific discovery for salt-marsh systems, and more generally for ecology.” Scientists like Dr. Ripple have argued that predators are important to the ecological health of other ecosystems. But it’s been difficult to test the hypothesis directly the way Dr. Bertness has.

Merryl Alber of the University of Georgia agreed that the experiment showed that removing predators could decrease salt marsh grass. But she was reluctant to draw big lessons from the study. “It is still a leap to connect dieback to recreational overfishing,” she said.

Wade Elmer, a plant pathologist at the Connecticut Agricultural Experiment Station in New Haven, thinks that the full story of the salt marshes’ decline is more complex than just fishing. Dr. Elmer has identified a new species of fungus that attacks cordgrass in New England salt marshes. He has suggested that the fungus may weaken the plants in a way that prevents them from making chemical defenses to ward off the marsh crabs.

“I think we all have our pet theories that explain what we see in our backyard,” said Dr. Elmer, “but these theories often fail as soon as we look elsewhere.”

Dr. Bertness doesn’t rule out the possibility that other factors are at play in the die-off of marshes. But he argues that fishing is having an enormous impact.

“The implications of these findings for the conservation of salt marshes are huge,” he said. “We need to maintain healthy predator populations.”


Wolves in a Tangled Bank

by Cristina Eisenberg

The wolves’ return to Yellowstone and the subsequent recovery of plants that elk had been eating to death in their absence has become one the most popularized and beloved ecological tales. By the 1920s humans had misguidedly wiped out most of the wolves in North America, thinking that the only good wolf was a dead one. Without wolves preying on them, elk and deer (also calledungulates) exploded in number. Burgeoning ungulate populations ravaged plant communities, including aspen forests. Decades later, the wolves we reintroduced in Yellowstone hit the ground running, rapidly sending their ecological effects rippling throughout the region, restoring this ecosystem from top to bottom. Yet today some scientists caution that this story is more myth than fact because nature isn’t so simple.

For decades scientists have been investigating the ecological role of wolves. In his 1940s game surveys, Aldo Leopold found ungulates wiping out vegetation wherever wolves had been removed. He concluded that by controlling ungulates, wolves could restore plant communities and create healthier habitat for other species, such as birds.

Since Leopold’s time, many scientists have studied food web relationships between top predators and their prey—called trophic cascades. In the 1960s and 1970s Robert Paine, working with sea stars, and James Estes, working with sea otters, showed that ecosystems without top predators begin to unravel. John Terborgh called the ensuing rampant species extinctions an “ecological meltdown.” Paine created the metaphorical termkeystone species to refer to top predators and noted that when you remove the keystone, arches and ecosystems collapse. Over the years ecologists found trophic cascades—also called top-down effects—ubiquitous from coral reefs to prairies to polar regions. However, William Murdoch and others have maintained that sunlight and moisture, which make plants grow, drive ecosystem processes from the bottom-up, making predators relatively unimportant. The Yellowstone wolf reintroduction provided the perfect setting to test these contrasting perspectives.

In the mid-1800s in his book The Origin of Species, Charles Darwin presciently described nature as a “tangled bank.” Nature’s complexity results from myriad species and their relationships with other species and all the things that can possibly affect them individually and collectively, such as disease, disturbance, and competition for food. Science works incrementally, taking us ever deeper into nature’s tangled bank as we investigate ecological questions. Each study answers some questions and begets new ones. Sometimes we find contradictory results. Learning how nature works requires what Leopold called “deep-digging research” in which we keep searching for answers amid the clues nature gives us, such as the bitten-off stem of an aspen next to a stream where there are no wolves.

waterton lakes

Trophic cascades science that focuses on wolf effects is still in its infancy, with huge knowledge gaps. For example, we’ve linked wolves to strong effects that cascade down through multiple food web levels. However, we’re just starting to parse how context can influence these effects. Some Yellowstone studies have found that wolves have powerful indirect effects on the plants that elk eat, such as aspens, due to fear of predation. With wolves around, elk have to keep moving to stay alive, which reduces browsing pressure. Conversely, a growing body of studies are finding no wolf effect—that aspens in places with wolves aren’t growing differently than those where predation risk is low. Other studies have found that wolf predation risk doesn’t affect elk feeding behavior. In my own research I’ve found that wolves need another keystone force—fire—to most effectively drive trophic cascades. With wolves and fire present, elk herbivory drops, aspens thrive, and biodiversity soars due to the healthy habitat created by young, vigorously growing aspen.

aspen, Glacier National Park

It’s human nature to try to find simple solutions. Today we are grappling with monumental environmental problems such as climate change and habitat fragmentation. Due to the wolf’s iconic status and our need to fix broken ecosystems, the environmental community and the media have run with the science that shows a strong wolf effect. This has inspired other scientists to prove that ecosystems are more complex than that. These dissenting studies demonstrate that the wolf dwells in a tangled bank, working alongside many other ecological forces.

Tangled banks seldom yield simple answers. However, arguing about what exactly carnivores do ecologically and why we need them is fiddling while Rome burns. Large, meat-eating animals improve the health of plant communities and provide food subsidies for the many species that scavenge on their kills. A system with wolves in it is far richer than one without and can support many more grizzly bears, coyotes, wolverines, and eagles. There are things we don’t know and disagreements about what we do know. But given the accelerated human-caused extinctions we are experiencing today, a precautionary approach to creating healthier ecosystems means conserving large carnivores.

Beyond empiricism, scientists often operate based on instinct. Instinct led Darwin to dig more deeply into species adaptation and Leopold to doggedly delve into the effects of predator removal. For many of us who conduct trophic cascades science, our instincts are telling us that wolves should be conserved in as high a number in as many places as possible, due to the invaluable benefits they can bring to ecosystems. To do anything other than conserve wolves would be foolish, given all we’ve learned thus far.



Image © Kitch Bain | Shutterstock

In parts of Australia, people drop poisoned meat from airplanes or helicopters or leave it along dirt roads to keep dingo numbers under control. The justification is that dingoes attack livestock and need to be suppressed. But dingo poisoning has set off a cascade of ecosystem changes that affect other wildlife: according to a new study, small mammals are taking a hit too.

Figuring out what happens when you get rid of top predators is tough. It’s impractical to do a large-scale experiment in which predators are removed from an ecosystem, and such studies would often be illegal or ethically questionable anyway.

But the study authors had the chance to follow a “natural experiment”. In New South Wales, Australia, they identified seven pairs of sites with different levels of dingo control. Within each pair, dingoes had been regularly poisoned at one site for the last five years; at the other site, less than 50 kilometers away, people hadn’t made much of an effort to control dingoes.

The researchers monitored kangaroos, wallabies, foxes, cats, possums, and small mammals such as rodents at each site. They searched for the animals by identifying footprints, scanning the area while driving, or setting traps with peanut butter, oats, and honey.

At the sites with frequent poisoning, the authors found more kangaroos and wallabies and more signs of fox activity, presumably because fewer dingoes were hunting or harassing those animals. Since kangaroos and wallabies are herbivores, the density of understory plants went down. And the number of small mammals, which take cover among plants and are preyed on by foxes, also dropped.

“Dingo control programmes in conservation reserves may be counter-productive from a biodiversity conservation perspective,” the authors write in Proceedings of the Royal Society B. Managers will need to find a way to keep dingo numbers up and farm animals safe at the same time. — Roberta Kwok | 13 March 2014


Colman, N.J. et al. 2014. Lethal control of an apex predator has unintended cascading effects on forest mammal assemblages. Proceedings of the Royal Society B doi: 10.1098/rspb.2013.3094

Cougar Predation Key To Ecosystem Health

A new study by researchers from Oregon State University found that cougars in Zion National Park have a profound impact on other aspects of the ecosystem, primarily by controlling deer populations and the ecosystem alterations related to deer browsing.

The general disappearance of cougars from a portion of Zion National Park in the past 70 years has allowed deer populations to dramatically increase, leading to severe ecological damage, loss of cottonwood trees, eroding streambanks and declining biodiversity. Researchers are calling it a “trophic cascade” of environmental degradation.

This “trophic cascade” of environmental degradation, all linked to the decline of a major predator, has been shown in a new study to affect a broad range of terrestrial and aquatic species, according to scientists from Oregon State University.

The research was just published in the journal Biological Conservation — and, like recent studies outlining similar ecological ripple effects following the disappearance of wolves in the American West — may cause land managers to reconsider the importance of predatory species in how ecosystems function.

The findings are consistent, researchers say, with predictions made more than half a century ago by the famed naturalist Aldo Leopold, often considered the father of wildlife ecology.

“When park development caused cougar to begin leaving Zion Canyon in the 1930s, it allowed much higher levels of deer browsing,” said Robert Beschta, an OSU professor emeritus of forest hydrology. “That set in motion a long cascade of changes that resulted in the loss of most cottonwoods along the streambanks and heavy bank erosion.”

“But the end result isn’t just loss of trees,” he said. “It’s the decline or disappearance of shrubs, wetland plants, amphibians, lizards, wildflowers, and even butterflies.”

Until recently, ecologists had a poor understanding of how the loss of an important predator, such as wolves or cougar, could affect such a broad range of other plant and animal species. But the evidence is now accumulating that primary predators not only have direct effects in influencing the population sizes of native grazing animals such as deer and elk — they also have indirect effects in changing their foraging behavior, in what has been called “the ecology of fear.”

That phenomenon, the scientists say, has been shown as vividly in Zion National Park as any other location they have ever studied.

In Zion Canyon, which since the early 1900s has been a popular tourist attraction, cougars are virtually absent, mostly just scared off by the huge influx of human visitors. With their natural enemy gone, growing and ravenous deer populations ate young cottonwood trees almost as quickly as they sprouted, robbing streambanks of shade and erosion protection.

As a result, floodplains began to erode away. Other types of vegetation and the animal species dependent on them suffered. And unless something is done, cottonwoods in Zion Canyon may ultimately disappear in areas accessible to deer, the researchers said.

By contrast, a nearby roadless watershed has similar native ecology but is sufficiently remote that it still has an intact cougar population and far fewer mule deer. In contrast to Zion Canyon, streambanks in this watershed have nearly 50 times more young cottonwood trees as well as thriving populations of flowers, lizards, butterflies, and several species of water-loving plants that help stabilize stream banks, provide food-web support, and protect floodplains for use by many other animal species.

“The documentation of species abundance that we have in this study is very compelling,” said William Ripple, a professor in the OSU Department of Forest Resources and lead author on the study. Researchers did a systematic survey of channel dimensions, streambank condition, vegetation and species presence along each study site.

“These two canyons, almost side by side, have a similar climate and their ecosystems should be quite similar,” Ripple said. “But instead they are very different, and we hypothesize that the long-term lack of cottonwood recruitment associated with stream-side areas in Zion Canyon indicates the effects of low cougar and high deer densities over many decades.

“It’s a great research setting and a great opportunity to assess the potential importance of a key predator,” he said. “We hope to conduct additional research in Zion National Park to further explore the findings of this initial study.”

It’s important to remember, the researchers said, that the ultimate driver behind all of these changes is humans — in the case of Zion Canyon, simply by their presence. That canyon receives nearly three million human visitors a year, the adjacent North Creek a stray handful of hikers. Cougars in Zion Canyon were not intentionally killed or removed, they just left due to the increased presence of humans.

As findings such as this — the way cougars affect deer and wolves affect elk — continue to mount, land managers may have to acknowledge the potentially enormous impact of these grazing animals on other ecosystem processes, scientists say. This could open the way to new management options once the role of herbivory by deer, elk, or other grazing animals is more fully understood.

In systems with wild ungulates, the sustainability of riparian habitats and biodiversity may require both predation on these herbivores as well as the fear of predation to further affect their behavior, the researchers concluded.

Ripple and Beschta considered other factors that may have played a role in loss of cottonwood trees in Zion Canyon, such as climate fluctuations or human interventions to stream channels, but concluded that those impacts could not have caused the enormous loss of trees and associated impacts to other biota that were found in the canyon.

The findings of this study may be relevant to other ecosystems in the U.S. and around the world where key predators have been removed, the researchers said, and high populations of native herbivores such as deer or elk — or domestic grazers such as cattle or sheep — affect native biodiversity.

This research was funded by the National Park Service.

 Story Source: The above story is based on materials provided by Oregon State UniversityNote: Materials may be edited for content and length.

John Terborgh: The Trophic Cascade Regulates Biodiversity

“The Trophic Cascade Regulates Biodiversity”

John Terborgh, Research Professor in the Nicholas School of the Environment and Earth Sciences at Duke University; Director of the Duke University Center for Tropical Conservation

In this presentation, Dr. Terborgh draws on his decades of ecological research in the Neotropics to explain how biological interactions intricately regulate biodiversity. Hypotheses on the maintenance of tropical forest diversity abound, but it is becoming increasingly recognized that interspecific interactions are vital to sustaining the rich diversity the tropics are famous for. Dr. Terborgh offers ecological insights on the regulation of biodiversity and describe how interactions between primary producers, herbivores, and their predators contribute to the richness of tropical forests.

PGE’s interdisciplinary Spring conference, “Conserving More Than Carbon: Valuing Biodiversity in a Changing World”, addressed the current state of knowledge of tropical forest diversity and outlooks for its protection.

Learn more about the conference and the participants at:

A behaviour-mediated trophic cascade involving Dugongs, Sea turtles and Tiger sharks.

‘Patterns of top-down control in a seagrass ecosystem: could a roving apex predator (Galeocerdo cuvier) induce a behaviour-mediated trophic cascade?’
By: Derek Burkholder, Michael Heithaus, James Fourqurean, Aaron Wirsing, Lawrence Dill

This manuscript presents data from a multi-year exclosure study to test a priori hypotheses regarding a behavior-mediated trophic cascade initiated by tiger sharks in a pristine seagrass ecosystem. We present evidence that seagrass communities are heavily influenced by large-bodied grazers, but only in areas where they can graze at lower risk from tiger shark predation. Although recent studies have suggested that roving predators, like tiger sharks, should be unlikely to trigger behavior-mediated cascades our work suggests that spatial heterogeneity can lead to such cascades. This study also suggests that the removal of large bodied predators could have wide-ranging consequeces for foundational species like seagrasses. Therefore, we believe that this manuscript should be of general interest to ecologists working in diverse marine, terrestrial, and freshwater ecosystems.

A behaviour-mediated trophic cascade from Journal of Animal Ecology on Vimeo.