Animal Influences on Ecosystem Dynamics

Author(s): Robert J. Naiman

Source: BioScience, Vol. 38, No. 11, How Animals Shape Their Ecosystems (Dec., 1988), pp. 750-
752
Published by: American Institute of Biological Sciences
Stable URL: http://www.jstor.org/stable/1310783
Accessed: 09/04/2009 13:46

Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at
http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless
you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you
may use content in the JSTOR archive only for your personal, non-commercial use.

Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at
http://www.jstor.org/action/showPublisher?publisherCode=aibs.

Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed
page of such transmission.

JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the
scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that
promotes the discovery and use of these resources. For more information about JSTOR, please contact support@jstor.org.

American Institute of Biological Sciences is collaborating with JSTOR to digitize, preserve and extend access
to BioScience.

http://www.jstor.org
 Animal Influences
on
EcosystemDynamics
Large animals are more than passive components of
ecological systems
Robert J. Naiman

L ess than 200 years ago, vast munity resulting in increased produc-
numbers of mammals inhab- tion or population density, whereas
ited the grasslands and forests The implicationsfor other changes may be detrimental,
of North America, migratory birds wildlife management causing species replacement or lower
filled the skies, and seemingly unlim- production rates.
ited fishes and fur-bearing mammals are substantial A good example is provided by
populated the lakes and rivers. Early and long lasting Pastor et al. (page 770 this issue) for
explorations by Jacques Cartier, Sa- moose (Alces alces) in the boreal for-
muel de Champlain, Pierre Esprit est. Under certain conditions moose
Radisson, and Pierre Gaultier de Va- foraging on deciduous plants can shift
rennes revealed wildlife so abundant that herbivory or predation may alter the forest community towards coni-
as to be almost unbelievable today the behavior of the food supply, cause fers (e.g., spruce, Picea glauca, Picea
(Bakeless 1950). It is clear from their
changes in life history strategy and mariana), which produce a low-
descriptions, and those of Bates community composition, or induce quality litter. Over several decades,
(1863) for the Amazon River basin physical and chemical defenses. How- this shift affects soil formation and
and Bartram (1791), Hays (1871), ever, in only a few cases have the nitrogen cycling, which ultimately af-
and Morgan (1868) for North Amer- implications of changes at the popu- fect the productivity of the spruce,
ica, that animals had strong interac- lation and community level been ex- longer-term plant succession in the
tions with the ecosystems supporting amined at the ecosystem level. These forest, and the population dynamics
them. changes are important because they of the moose.
Although less widespread and eco- reverberate throughout all trophic As a general phenomenon, this
logically influential than in the past,
levels, causing alterations to the eco- process is complicated and difficult to
within our remaining natural systems system that cannot necessarily be pre- study because many animal popula-
animals continue to play significant dicted beforehand. Timely examples tion cycles occur over long periods
ecological roles that go far beyond are found in ecosystems invaded, and (i.e., decades); alterations to the eco-
their immediate requirements for disrupted, by exotic plants and ani- system are apparently subtle over
food or habitat. In many cases, they mals (Elton 1958, Vitousek 1986, Za- short periods (i.e., increased tree mor-
are responsible for biogeochemical, ret and Paine 1973). tality or altered soil formation); and
successional, and landscape alter- Large animals, due to their size, shifts in biogeochemical cycles or sed-
ations that may persist for centuries.longevity, and food and habitat re- iment and soil characteristics are not
Investigations over the past several
quirements, tend to have a substantial detectable over short periods (i.e.,
decades have clearly elucidated inter-impact on ecosystems. Both the ani- years). Nevertheless, these succes-
actions between animals and their mals' feeding strategies and their sional pathways often result in a het-
food supply, and the resultant effectsphysical alterations to the environ- erogeneous landscape that would not
on consumer population dynamics ment (Figure 1) affect plant and ani- occur under the dominating influence
and the community structure of the mal community composition, which of climate and geology alone; they
food supply. It is now well known in turn alters biogeochemical cycling require the intervention of animal ac-
of nutrients and ions in soils, sedi- tivity.
RobertJ. Naiman is the directorof the ments, and water. These alterations Even though animal population
Center for Streamside Studies, AR-10, have either positive or negative influ- sizes have severely diminished over
Universityof Washington,Seattle98195. ences upon the forage base and the the last two centuries, and remaining
? 1988 AmericanInstituteof Biological consumers themselves. That is, some populations are increasingly managed
Sciences. changes may benefit the forage com- to limit their effects, the physical his-

750 BioScience Vol. 38 No. 11

toric evidence of animal influence on
ecosystem dynamics remains wide-
spread. Two common examples are
mima mounds and beaver meadows.
Mima mounds are earthmounds,
prevalent in western North America.
They may be more than 2 meters
high, up to 50 meters in diameter, and
of a density of more than 100 per
hectare. Until recently they were
thought to have originated from ei-
ther wind or water erosion, wind dep-
osition, periglacial freeze-thaw
dynamics, or soil translocation by
burrowing rodents such as gophers.
We now know that they were formed
by the activities of pocket gophers
(Thomomys spp.) over the past sev-
eral thousand years (Cox and Gakahu
1986). Likewise, North American
beaver (Castor canadensis) that build
dams have been shown to be instru-
mental in the formation of wetlands
and meadows, the geologic and vege-
tative features of which may persist
for centuries (Ives 1942, Rudemann
and Schoonmaker 1938; Figure 2).
Today people are the primary Herbivores,such as moose (Alces alces), can shift the forest plant communityfrom
agents of environmental change in deciduousplants to conifers.This effectcan ultimatelyinfluencesoil formation,plant
most areas of the world, and animal productivityand succession,and animalpopulationdynamics.Photo:PamelaMclnnes,
influences on ecosystems are not as Departmentof Fisheriesand Wildlife,Universityof Minnesota,St. Paul, MN.
easily recognized. Yet, animal influ-
ences remain important in natural
systems. In East Africa foraging by miferana), fall cankerworm (Also- environmenthave long-termimplica-
bush elephant (Loxodonta africana) phila pometaria), mountain pine bee- tions for ecosystem dynamics
is a major element of habitat and tle (Dendroctonus ponderosae), tent prompteda symposiumon this topic
landscape change (Laws 1970) and, caterpillar (Malacosoma disstria), for the 1987 annual meetings of the
in North America, insect herbivory and gypsy moth (Portheria dispar) set AmericanInstitute of Biological Sci-
has a substantial impact on forest in place a series of events that even- ences and the Ecological Society of
dynamics (Dyer 1986). Elephant feed- tually result in alterations to forest America held jointly in Columbus,
ing strategies involve physical distur- production, water yield, nutrient cy- Ohio. The articlesin this specialissue
bance of trees (especially Acacia) and cling, and streamwater chemistry of BioScience reflect animal-eco-
shrubs. This disturbance causes wide- (Dyer 1986). system interactionsoccurringover a
spread vegetation changes in wood- The activities of tree-cutting bea-
lands and savannas. These activities ver, defoliating insects, browsing ele-
alter the role of fire in the system and phants, and other conspicuous con-
severely affect the population dynam- sumers are not always detrimental, as
ics of other wildlife by changing the they might seem initially. Over the
food supply and the habitat. These longer term, the consumers and the
alterations are later reflected in soil forage base interact to influence criti-
formation, in the structure and cal community or ecosystem pro-
dynamics of riparian zones around cesses, thus developing a variety of
water courses, and in the biogeoche- system feedbacks that would not or-
mical cycling of important nutrients dinarily exist (Mattson and Addy Soils. Sediment. /
(Hatton and Smart 1984). 1975). An important management- and Water I.

Defoliating insects provide an in-
teresting contrast to elephants be-
cause, in forests, they cause consider-
able alteration to system function
without a major physical disturbance.
In North America, outbreaks of
spruce budworm (Choristoneura tu-
level question is how to plan for this
natural, long-term variability (or un-
certainty) in systems increasingly sub-
jected to short-term demands for sta-
ble resources.
The emerging realization that inter-
actions between animals and their
Figure 1. Animals influence ecosystem
dynamicsby their foragingstrategiesand
by physicalhabitat alterations.These im-
pacts are transmittedto the community,
resultingin long-termchangesto biogeo-
chemical cycles in soils, sediments, and
water.

December 1988 751


Figure 2. Dam building by beaver often creates extensive wetlands persisting for long
periods. This wetland in Voyageurs National Park, Minnesota, is maintained by a series
of dams, of which only a few are actively repaired at any one time. The age of the dam
is not known, but sediment accumulations in the stream valley suggest a long period of
beaver activity. Photo: Naiman.
variety of habitats and of spatial and
temporal scales.
My colleagues and I (page 753 this
issue) document the influence of bea-
ver on the structure and dynamics of
North American streams. Carpenter
and Kitchell (page 764 this issue) dis-
cuss the role of predacious fish in
structuring the forage community and
ultimately the productivity of lakes.
Pastor and his colleagues (page 770
this issue) discuss the importance of
moose in altering plant communities
and soil dynamics in the boreal forest.
Whicker and Detling (page 778 this
issue) describe the influence of prairie
dog (Cynomys), and Huntly and
Inouye (page 786 this issue) discuss
the role of pocket gopher (Geomys
bursarius) on the vegetative composi-
tion, productivity, and nutrient char-
acteristics of North American grass-
lands. Finally, McNaughton and his
colleagues (page 794 this issue) de-
752
scribe the role of grazing mammals on
the dynamics of the Serengeti ecosys-
tem in Africa.
In each article the fundamental
mechanisms for ecosystem alteration
are similar. Animals, through physical
disturbance or through their feeding
strategies, alter community structure
and biogeochemical cycles in such a
way that the reverberations are trans-
mitted to all trophic levels. The path-
ways are complex but the fundamental
mechanisms are similar-and the im-
plications for wildlife management are
more substantial and long lasting than
most of us have yet realized.
References cited
Bakeless, J. E. 1950. The Eyes of Discovery.
J. B. Lippincott, Philadelphia, PA.
Bartram, W. 1791. The Travels of William
Bartram (1928 ed.). Dover Publ., New York.
Bates, H. W. 1863. The Naturalist in the River
Amazon (1920 ed.). MacMillan, London,
UK.
Carpenter, S. R., and J. F. Kitchell. 1988.
Consumer control of lake productivity. Bio-
Science 38: 764-769.
Cox, G. W., and C. G. Gakahu. 1986. A
latitudinal test of the fossorial rodent hy-
pothesis of Mima mound origin. Ann. Geo-
morphology 30: 485-501.
Dyer, M. I. 1986. The role of herbivores in forest
ecosystems: the case for biosphere reserves.
Pages 97-102 in Y. Hanxi, W. Zhan, J. N. R.
Jeffers, and P. A. Ward, eds. The Temperate
Forest Ecosystem, ITE SymposiumNo. 20. The
Lavenham Press Ltd., Lavenham, Suffolk, UK.
Elton, C. S. 1958. The Ecology of Invasions by
Animals and Plants. J. Wiley & Sons, New
York.
Hatton, J. C., and N. O. E. Smart. 1984. The
effect of long-termexclusion of large herbivores
on soil nutrientstatusin MurchisonFallsNa-
tional Park, Uganda. Afr. J. Ecol. 22: 23-30.
Hays, W. J. 1871. Notes on the range of some
of the animals in America at the time of
arrival of the white men. Am. Nat. 5: 25-30.
Huntly, N., and R. Inouye. 1988. Pocket go-
phers in ecosystems: patterns and mecha-
nisms. BioScience 38: 786-793.
Ives, R. L. 1942. The beaver-meadow complex.
J. Geomorphology 5: 191-203.
Laws, R. M. 1970. Elephants as agents of
habitat and landscape change in East Africa.
Oikos 21: 1-15.
McNaughton, S. J., R. W. Ruess, and S. W.
Seagle. 1988. Large mammals and process
dynamics in African ecosystems. BioScience
38: 794-800.
Mattson, W. J., and N. D. Addy. 1975. Phyto-
phagous insects as regulators of forest pri-
mary production. Science 190: 515-522.
Morgan, L. H. 1868. The American Beaver and
PA.
His Works.J. B. Lippincott,Philadelphia,
Naiman, R. J., C. A. Johnston,and J. C. Kel-
ley. 1988. Alteration of North American
streams by beaver. BioScience 38:753-762.
Pastor, J., R. J. Naiman, B. Dewey, and P.
McInnes. 1988. Moose, microbes, and the
boreal forest. BioScience 38: 770-777.
Rudemann, R., and W. J. Schoonmaker. 1938.
Beaver dams as geologic agents. Science 88:
523-525.
Vitousek, P. M. 1986. Biological invasions and
ecosystem properties: Can species make a dif-
ference? Pages 163-176 in H. A. Mooney, ed.
Ecology of Biological Invasionsof North Amer-
ica and Hawaii. Springer-Verlag,New York.
Whicker,A. D., and J. K. Detling. 1988. Eco-
logical consequences of prairie dog distur-
bances. BioScience 38: 778-785.
Zaret, T. M., and R. T. Paine. 1973. Species
introduction in a tropical lake. Science
182:449-455.
BioScience Vol. 38 No. 11