Perspective

Yellowstone Wolves and the Forces That Structure

Natural Systems
Andy P. Dobson1,2*
1 Department of Ecology and Evolutionary Biology, Princeton University, Princeton, New Jersey, United States of America, 2 Santa Fe Institute, Santa Fe, New Mexico,
United States of America

ranchers who live outside the park and environmental example of nonlinear po-
Abstract: Since their introduction
in 1995 and 1996, wolves have had regard wolves as varmints, best used for litical expediency.
effects on Yellowstone that ripple target practice. The pack that I heard Once introduced in 1995 and 1996, the
across the entire structure of the howling outside the cabin every night was wolf population grew rapidly. At the time,
food web that defines biodiversity quickly dispatched by the local rancher the elk population was declining from an
in the Northern Rockies ecosystem. soon after I left; their pelts could be found all-time high and provided a large supply
Ecological interpretations of the for sale in one of the souvenir stores at the of prey to fuel wolf reproduction; the
wolves have generated a significant entrance to Yellowstone. population increased at close to the
amount of debate about the rela- Scientists initially appear as polarized in maximum rate ever recorded [1]. As the
tive strength of top-down versus their opinions of the role of wolves and wolf numbers increased, the elk numbers
bottom-up forces in determining large predators in ecosystems as the wolf decreased, but at a rate that was more
herbivore and vegetation abun- watchers and ranchers are about their parsimoniously explained by a prolonged
dance in Yellowstone. Debates value to the local economy. Wolves were drought and levels of human harvest, the
such as this are central to the introduced back into Yellowstone follow- decline in abundance far exceeding that
resolution of broader debates which could be accounted for purely in
ing the development of a huge environ-
about the role of natural enemies
mental impact assessment (EIA) that terms of elk consumed by wolves [2,3].
and climate as forces that structure
food webs and modify ecosystem attempted to predict the outcome of their Significant evidence does suggest that the
function. Ecologists need to signif- reintroduction. The EIA, a 4-ft-deep pile elk had changed their feeding habits in the
icantly raise the profile of these of documents, provided solid testimony to presence of wolves, avoiding areas where
discussions; understanding the the need for a deeper empirical and they could readily be ambushed [3–8].
forces that structure food webs theoretical understanding of how ecologi- This allowed vegetation in riparian areas
and determine species abundance cal food webs respond to species additions to recover; photographs taken at a variety
and the supply of ecosystem ser- and losses. At the time, even the suggestion of locations showed considerable recovery
vices is one of the central scientific of introducing wolves created huge discord of aspen in areas where it had become
questions for this century; its com- in the ranching community surrounding overgrazed in the years when elk were
plexity will require new minds, new Yellowstone; most ranchers (and some abundant [1,9]. Although these riparian
mathematics, and significant, con- ecologists) were convinced wolves would areas cover only a small area of the
sistent funding. feed exclusively on cattle and sheep; the ecosystem (,2%), the park was witnessing
ranching industry was dead set against the first significant growth of aspen for
reintroduction. A curious event then over half a century. More recent data
occurred: photographers started getting suggest that similar recoveries are being
In the half moonlight at dawn on a photographs of wolves that had naturally seen in cottonwoods and willows [1]; this
sharply cold January morning, they looked colonized the park. As any natural coloni- in turn has led to an increase in the
like small ponies galloping beside the old zation would provide the wolves with full abundance and diversity of riparian bird
railroad at the northern entrance of legal protection under the United States species [10]. All of this evidence suggests
Yellowstone National Park. They weren’t Endangered Species Act, the Ranchers that wolves have a strong top-down effect
ponies. This was the ‘‘Eight Mile’’ wolf Association hastily made a U-turn and on trophic structure of the ecosystem
pack, each member huge, healthy, and supported introduction on the grounds (Fig. 1).
vigorous, romping through the light snow that experimentally introduced wolves Alternatively, climate has been argued
on a morning quest for elk, bison, or were nonnative and could be shot if they to be the principal driver of ecosystem
anyone too slow to get out of their way. It left the park. I know of no better change, not wolves; changes in vegetation
was an incredible moment, one that
evoked feelings shared by the hundreds Citation: Dobson AP (2014) Yellowstone Wolves and the Forces That Structure Natural Systems. PLoS
of wolf watchers who come to Yellowstone Biol 12(12): e1002025. doi:10.1371/journal.pbio.1002025
every month of the year hoping to Published December 23, 2014
experience even a glimpse of the wolves. Copyright: ß 2014 Andy P. Dobson. This is an open-access article distributed under the terms of the Creative
The enthusiasm of the wolf watchers is Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium,
almost totally reversed by the many local provided the original author and source are credited.
Funding: A grant from the James McDonnell Foundation supports my work on food webs. The funders had no
The Perspective section provides experts with a role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
forum to comment on topical or controversial issues
Competing Interests: The author has declared that no competing interests exist.
of broad interest.
* Email: [email protected]

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Fig. 1. Wolves in Yellowstone NP. Photo credit: Daniel Stahler/National Park Service (NPS) photo from http://www.yellowstonewolf.org/, a site at
which information and pictures of Yellowstone wolves can be found.
doi:10.1371/journal.pbio.1002025.g001

may have been driven by bottom-up and golden eagles, both of which have tion community to sanctify the wolves as
changes in water availability due to increased in abundance [2]. bringing only benefits to the ecosystem
changes in snow melt patterns [11]. Wolf Less well understood is the impact of [12], there is still a need for stronger data to
population expansion occurred at a time wolves on coyotes, the numbers of which support some of the beneficial claims made
when the Yellowstone region was entering may have declined since wolves were for wolf reintroduction. Some of this will
a prolonged drought that also reduced reintroduced [2]; carnivores are aggressive come from Yellowstone, but this needs to
forage available to elk; this combined with to other carnivores of similar but slightly be combined with studies of wolf reintro-
human harvest contributed significantly to smaller body size. As coyotes were the duction, or natural reestablishment, in
the declines in ungulate abundance. Fur- primary predators of sheep, you would other ecosystems. If the patterns observed
thermore, climate change has lengthened think that the sheep ranchers would in Yellowstone are repeated, as preliminary
the growing season for willows and aspen applaud wolves for the reduced loss of evidence suggests, then hard-core wolf
by around 27 days in the last couple of stock to predators; they have been notice- haters are going to need to reconsider the
decades [12], while the vegetation in many ably silent on this front. More subtly, the labelling of wolves as varmints.
areas of the park is dominated by conifer presence of wolves may help reduce the The research and debates surrounding
forest that has simultaneously been recov- threat posed by chronic wasting disease the role of wolves in modifying the
ering from the fires of 1989. Thus, it is not (CWD), an emerging prion pathogen that behavior and abundance of species on
straightforward to differentiate between is spreading from elk and deer to cattle multiple trophic levels in Yellowstone
postfire recovery and the indirect effects of and is arguably the biggest biological illustrates the complex interactions be-
carnivores on vegetation regeneration. threat to ranching in the region [15]. tween the forces that structure patterns
Concomitant to wolf introduction, the Unfortunately, the ranching community of abundance in natural ecosystems. The
grizzly bear population was increasing, does not recognize that the wolves may be debate gets to the heart of one of the
creating the potential for indirect compe- doing them a huge favor by removing sick central scientific challenge of ecology: how
tition between bears and wolves as the elk and mule deer infected with CWD can we understand the structure of food
latter selectively prey on old or injured elk (and elk and bison infected with brucello- webs? Central to any discussion of food-
in the winter. This predation reduces the sis) from the wild reservoir of infection. If web dynamics and ecosystem manage-
number of elk that would otherwise die CWD or Brucella enters cattle herds in the ment is the relative importance of top-
and become available for grizzlies emerg- states bordering Yellowstone, then federal down roles played by large predators and
ing from hibernation in the spring. This mandates will hugely restrict movement of pathogens and bottom-up forces driven by
absence of ‘‘frozen meals’’ caused grizzlies cattle in and out of these states. the climatological processes that determine
to switch to feeding on elk calves as an We may have to wait at least another ten plant growth. All of the work from
alternative spring food source when re- years before the impact of wolves on the Yellowstone cries out for the development
covering from their long winter fast [13]. Yellowstone ecosystem is fully quantified. of next-generation, population-based eco-
As elk numbers declined following the Although many strong patterns are ob- system models that focus on interactions
triple assault of drought, wolves, and served, several of these may be correlation between climate, vegetation, and the
bears, both grizzlies and some wolf packs without causation (for example, the in- dominant herbivore and carnivore species
switched their attention to bison [14], crease in beaver abundance is more likely in the park. In particular, food-web
which require larger packs to make an to be a consequence of beaver introductions ecologists need to more aggressively move
effective kill but ultimately provide a larger to the north of Yellowstone National Park beyond descriptions of the network geom-
meal. All of the extra carcasses have [NP] [12]). Furthermore, although there is etry of food webs and grasp the thistle of
provided a new bounty of food for ravens considerable pressure from the conserva- food-web dynamics. More generally, we

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cannot afford for this debate to become of the impact of these changes at ecosys- stone is a classic case study in this broader
polarized; that simply suggests to funding tem-level scales. However, the results of class of problems: it is about understand-
agencies and the general public that management experiments are rarely clear- ing how to measure forces and processes
ecologists do not know how ecological cut and often ambiguous. It is all too easy that act between operators at a variety of
systems function. Instead, we need to to be critical about lack of controls and different spatial and temporal rates within
frame the discussion as a major scientific absence of replication (which is nontrivi- a natural ecosystem that contains a
challenge that requires significant interna- ally a function of trivial budgets!), but diversity of natural heterogeneities (that
tional and national funding. understanding how food webs in national initially appear to confound the search for
There are curious and unexplored par- parks react to the addition and loss of broad patterns). If we re-pose ecology as
allels between work on food webs and species is as scientifically challenging as the science that examines the forces that
trophic interactions and that of physicists searching for tiny particles using very structure the central part of the universe in
who are trying to understand the forces that expensive particle accelerators. The cen- which we live, then more funding might be
determine the way the universe is struc- tral problem is ecological budgets are tiny available to address these complexities; we
tured at either the atomic or astronomical compared to those for ‘‘big science,’’ so we would also simultaneously attract more
level. At both scales, a series of nested forces need to use all sources of information that bright minds willing to grapple with
hold increasingly large particles together are available, including management ex- complexity.
using a mixture of centripetal and gravita- ercises, to interpret findings at the appro- From a much broader perspective, we
tional forces, which operate essentially as priate ecosystem-level scale. need many more ecosystem-level studies of
bottom-up forces (although this is almost a If we agree that physicists and ecologists how species interactions between preda-
metaphysical debating point!). Seen from are both trying to understand the forces tors, parasites, and prey change the
this perspective, the current controversy that determine the structure of the uni- patterns of spatial heterogeneity in vege-
about ecosystem-level effects of wolf rein- verse, what are the major scientific
tation that ultimately drive levels of
troduction to Yellowstone NP is every bit as differences between their approaches?
biodiversity at higher trophic levels. This
scientifically exciting as the recent discovery Ecologists are focusing on understanding
is an exercise that requires a new gener-
of the Higgs boson. Determining the these forces at the spatial and temporal
ation of spatial, multispecies, multitrophic
strength of the forces produced by the loss scales intermediate to that of physicists—
models and many more debates such as
or addition of particles or species to these less heroic, perhaps, but the scale that is
the current one about the role of wolves in
very different systems are key scientific directly relevant to humans. Less heroic or
Yellowstone. Resolving these discussions
questions for the 21st century. Although not, from the perspective of systems with
each discipline uses very different types of interacting components, ecosystems and will allow ecologists to present a much
equipment, budgets, and collaborations to their constituent species will always be as stronger case to funding agencies and the
undertake experiments that generate data complicated as those exhibited by atoms general public for ecology to be recognized
for subsequent analysis, they each seek the and bosons or galaxies and planets, as the central scientific discipline of the
answer to the same questions: ‘‘What are perhaps more so; food webs have many 21st century. Ecology’s mathematical
the fundamental forces that structure the different types of ‘‘particles’’ (species) that problems are as complex as anything in
universe in which we live, how do they interact, evolve, and behave nonlinearly in physics, and their solutions are required
operate, and how can we measure them?’’ a huge variety of time, and spatial, scales. with increasing urgency, particularly if we
The hunt for the Higgs boson—an Ultimately, we need to arrive at a want to test these assumptions and pre-
infinitely tiny particle whose energy is realization that the mathematics of food dictions against viable natural ecosystems.
required to hold the interior particles of webs and ecosystems is as complicated as
atoms in orbit—was an international that found in any of the problems of Acknowledgments
research collaboration with a budget that atomic and galactic structure studied in Time spent at the Santa Fe Institute constantly
exceeded all funding for ecology over the physics. Increasingly, we are realizing that stimulates me to think about the need to
last ten years, perhaps even over the last the quality of human life on the planet understand problems of ‘‘complexity.’’ The
century! In contrast, funding for work on depends on a deep functional understand- ideas described here were developed from many
natural ecological systems is usually cob- ing of the forces that structure the hours spent in Yellowstone in conversation with
bled together from a mixture of govern- dynamics of food webs and the ecosystem Peter Hudson, Emily Almberg, Tim Coulson,
ment and individual research funds; it is services they provide to the human Paul Cross, Mary Meagher, Doug Smith, Dan
rarely clear from year to year when, or if, economy. We may even need new math- McNulty, Dan Stahler, Bridgett van Holdt, and
Robert Wayne. Anieke van Leeuwen, Annarie
funds will appear for the next year’s ematics to deal with these levels and layers
Lyles, and Mercedes Pascual commented ex-
salaries. One benefit of working in nation- of complexity. tensively on an earlier draft. The opinions
al parks is that management occasionally The current controversy about the role expressed here are purely my own.
allows experimental introductions, or re- that wolves play in modifying the behavior
movals, of species that permit investigation and dynamics of other species in Yellow-

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