Mountains and Plains-The Ecology of WyomingLandscapes PDF

Mountains
and Plains
Dennis H. Knight
George P. Jones
William A. Reiners
William H. Romme
Mountains
and
Plains
The Ecology
of Wyoming
Landscapes
S e c o n d Ed i t i o n
New Haven and London
University of Wyoming
Laramie, Wyoming
www.uwyo.edu/biodiversity
Published with assistance from the University of Wyoming
Library of Congress Cataloging-in-Publication Data
Biodiversity Institute (wyomingbiodiversity.org).
Knight, Dennis H.
Published with assistance from the Mary Cady Tew Memorial

Fund.
Mountains and plains: the ecology of Wyoming landscapes/

Dennis H. Knight, George P. Jones, William A. Reiners, William H.
Romme.Second edition.
First edition 1994. Second edition 2014. Copyright 1994, 2014
pagescm
by Yale University. All rights reserved. This book may not be
Includes bibliographical references and index.
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ISBN 978-0-300-18592-8 (pbk. with flaps : alk. paper)
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1.EcologyWyoming. 2.Landscape ecologyWyoming.
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I.Title.
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Frontispiece photo of Gannett Peak and Dinwoody Creek by Scott
Copeland (www.scottcopelandimages.com); see fig. 2.5.
To our students and colleagues,

from whom we have learned so much
Nature is an open book. . . . Each grass-covered hillside

is a page on which is written the history of the past,
conditions of the present, and predictions of the future.
. . . Let us look closely and understandingly, and act
wisely, and in time bring our methods of land use and
conservation activities into close harmony with the
dictates of nature.
John E. Weaver, 1954
Contents
Preface, ix
Part One. Wyoming and
Part Four. Foothills and
Acknowledgments, xi
the Rocky Mountain West
Mountains
1. Introduction, 3
10. Escarpments and the
2. Landscape History, 12
3. P resent-Day
Environments and
Climate Change, 27
Foothill Transition, 155

11. Mountain Forests, 173
12. The Forest Ecosystem,
200
13. M
ountain Meadows
and Snowglades, 221
Part Two. Wetlands

4. R iparian Landscapes, 45
14. Upper Treeline and

Alpine Tundra, 230
5. Marshes, Playa
Wetlands, Wet
Meadows, and Fens, 66
Part Five. Landscapes of

Special Interest
15. The Greater
Part Three. Plains and

Intermountain Basins
6. Grasslands, 83
7. Sagebrush, 109
8. Desert Shrublands and
Playas, 131
Yellowstone
Ecosystem, 245
16. The Black Hills, Bear
Lodge Mountains, and
Devils Tower, 266
17. The Laramie Basin, 282
9. Sand Dunes, Badlands,

Mud Springs, and
Mima Mounds, 142
Part Six. Sustainable Land

Management
18. Using Western
Landscapes, 305
Epilogue, 319
Appendix A: Conversion
Table, 321
Appendix B:
Characteristic Soil
Types, 322
Notes, 325
References, 349
Index, 387
This page intentionally left blank
Preface
Many changes have occurred in the Rocky Mountain
lands, forests, subalpine meadows, and alpine tundra.
region since 1994, when the first edition of this book
The ecology and management of croplands are beyond
was published. Wildlife habitat has been fragmented
the scope of this book, but we discuss the ecological
at alarming rates, the once abundant sage-grouse has
effects of irrigation, invasive species, livestock, feral
been proposed for protection by the Endangered Species
horses, elk, prairie dogs, and wolves. The book ends
Act, invasive plants have become more common,
with a chapter on current issues pertaining to land
wolves have been reintroduced, climate change is now
management and conservation.
well documented, epidemics of forest insects are more
Plant ecology is emphasized, because vegetation pro-
widespread, forest fires are more frequent, and new ap-
vides habitat and gives character to the landscape, but
proaches for conservation have been adopted. Our goal
we also discuss plant-animal interactions, hydrology,
for the second edition, 20 years later, is to provide a new
nutrient cycling, and the effects of disturbances, such
synthesis of the ecological research that is pertinent to
as fire, insect epidemics, and timber harvesting. Most
natural resource management, with a focus on the ecol-
chapters consider the implications of climate change.
ogy of Wyoming and adjacent parts of Colorado, Utah,
The chapters vary in length and structure, based on the
Idaho, Montana, South Dakota, and Nebraska.
information available and our perception of what most
The book has six parts. The first is an introduction

to ecology, conservation biology, and the concept of
readers will find interesting. We provide guidance to

further information in the endnotes.
ecosystem services. It also includes a chapter on the
Recognizing the diverse backgrounds of people inter-
geological development of the diverse landscapes of the
ested in western landscapes, we have written this book
region and changes during the past several thousand
for anyone intrigued by the natural history and wildlife
years, plus a chapter on present-day environments and
of the region. The text is essentially free of technical
climate change during the past few decades. The parts
terms, but those few that remain are defined where they
that follow focus on riparian and nonriparian wetlands,
first appear. Definitions can be located using the index.
semi-arid basins and plains, foothills and mountains,
We have used common names for plants and animals
and three landscapes of special interestthe Greater
throughout the book, but Latin names for most species
Yellowstone Ecosystem; the Black Hills, Bear Lodge
are provided in the tables. English units of measure are
Mountains, and Devils Tower; and the Laramie Basin.
used almost exclusively, with a metric conversion table
All the major kinds of ecosystems are discussed: flood-
in appendix A.
plains, marshes, fens, mixed-grass prairie, sagebrush
Online resources are now of great value for ecologists
steppe, desert shrublands, sand dunes, badlands, wood-
and land managers, providing quick access to additional
ix
Preface
information, photos, and distribution maps for the
about. We also have been involved with the application
plants, animals, and virtually every topic mentioned
of our research and that of other scientists to perplex-
in the text. Some of the pertinent website addresses
ing challenges faced by society today. Discussions with
are in the endnotes. Our book has a website as well. If
agency and private land managers have broadened our
you learn something that we should know about, go
perspectives. Our goal in writing this book is to present
to mountainsandplains.net. There you can make com-
information that is pertinent to ongoing debates and
ments, ask questions, read our responses to questions
that illustrates the long history of a place that has been
already submitted, and see additional photos. Other
our home. Two important questions are: How can we
useful links are at http://www.uwyo.edu/biodiversity/,
use ecosystems in ways that enable future generations
the website for the Biodiversity Institute at the Univer-
to benefit from them as well? And how can we antic-
sity of Wyoming.
ipate and adapt to changes associated with a warmer
We have worked together for more than 30 years.
and drier climate while conserving biological diversity?
Our careers have blended education at various levels
Answers require policies informed by science but cogni-
with research on many of the ecosystems that we write
zant of diverse values and traditions.
Acknowledgments
We have benefited greatly from the encouragement,
John Hauer, and Hollis Marriott. Helpful suggestions for
knowledge, and talents of many colleagues during the
the Laramie Basin chapter were provided by Don Boyd,
development of this book. Ken Driese provided numer-
Tony Hoch, and Larry Munn; Martin Curry, Jerry Han-
ous photographs and some of our maps, and he, Dan
sen, Bern Hinckley, Mindy Meade, Ruben Vasquez, Kim
Binkley, and Phil White offered detailed, helpful sug-
Viner, and Lindsay Wheat answered questions about
gestions on all of the chapters. Phil was extraordinarily
the Basin. The chapter on land management issues was
diligent in assisting with library work and last-minute
improved with the suggestions of Indy Burke, Brian
details. Chris Nicholson contributed the climate maps
Kuehl, Jay Lillegraven, and Chris Madson.
in chapter 3 and Ramesh Sivanpillai created three of the
Others have helped as well. Especially important
maps in chapter 17. Their work, along with that of vari-
has been the research of our students over the years.
ous photographers, is identified in the figure captions.
They have greatly influenced how we think about ecol-
Many colleagues offered helpful suggestions on early
ogy and natural resource management. Numerous col-
drafts of various chapters. Don Boyd, Steve Buskirk,
leagues and agency personnel provided information,
Steve Jackson, Jay Lillegraven, Larry Munn, Bryan Shu-
including Ken Anderson, Bob Dorn, Bonnie Heidel, Jeff
man, Art Snoke, and Cathy Whitlock assisted with our
Lockwood, Jim Lovvorn, Chris Madson, Jill Morrison,
discussion of geologic history, modern-day environ-
Larry Munn, Ginger Paige, Scott Schell, Pete Stahl, Alan
ments, human history, and climate change. The for-
Ver Ploeg, Danny Walker, and Lee Whittlesey. Ron Hart-
est chapters were improved by the comments of Craig
man, Bonnie Heidel, and Ernie Nelson assisted with
Benkman, Peter Brown, Brent Ewers, Tim Fahey, Dan
plant identification, and Steve Buskirk answered ques-
Tinker, and Tom Veblen, and the grassland and shrub
tions about Wyomings fauna. Allory Deiss drew essen-
land chapters by the comments of Jeffrey Beck, Angela
tially all the line figures, Judy Knight contributed four
Hild, Alan Knapp, William Lauenroth, Brian Mealor,
illustrations, and Joy Handley assisted with the tables
Rachel Mealor, and Elise Pendall. The wetland chapters
and plant nomenclature. Jean Thomson Black at Yale
benefited from the comments of David Cooper, Laurie
University Press was very supportive, as she was with
Gilli
gan, Bonnie Heidel, and Joanna Lemly, and the
the first edition. Her associates Samantha Ostrowski
alpine meadow and tundra discussions from suggestions
and Mary Pasti, along with Peter Strupp and Cyd West-
by Tim Seastedt and William K. Smith. The chapter on
moreland of Princeton Editorial Associates, drew our at-
the Greater Yellowstone Ecosystem was improved with
tention to details and problems that we had overlooked.
the help of Andrew Hansen, Matt Kaufmann, and Paul
Nancy Ovedovitz designed the book; Christy Spielman
Schullery, and the Black Hills chapter by Beth Burkhart,
and Bryce Tugwell developed our website.
xi
xii
Acknowledgments
Over the years our research has been made possible
Wyoming Depart
ment of Agriculture, the Wyoming
by grants from the National Science Foundation, the
Game and Fish Department, and the U.S. Forest Service.
U.S. Forest Service, the U.S. Department of Energy, the
For the second edition, we received a grant from the
U.S. Department of Agriculture, the Bureau of Land
Biodiversity Instit ute at the University of Wyoming, di-
Manage
ment, the University of WyomingNational
rected by Carlos Martinez del Rio. Thanks to his encour-
Park Service Research Center, the Wyoming Water Re-
agement and support, along with that of Indy Burke,
sources Research Institute, The Nature Conservancy,
director of the universitys Haub School of Environ-
and the Mellon Foundation. Grants to facilitate the
ment and Natural Resources, this book is much better
publication of the first edition were provided by the
than it would have been otherwise.
Part One Wyoming and the
Rocky Mountain West
Chapter 1
Introduction
Straddling the Continental Divide and with an aver-
grasslandsthe Thunder Basin National Grassland in
age elevation of 6,700 feet above sea level, the land-
northeastern Wyoming and the Pawnee National Grass-
scapes of Wyoming are similar to those encountered
land in northern Colorado. Only about 4 percent of
by explorers in the early 1800s. There are now roads,
Wyoming is currently under cultivation, leaving large
ranches, farms, cities, and industrial developments,
tracts of land that have never been plowed and are still
but outdoor enthusiasts, naturalists, and scientists
dominated by native plants.1 Overall, the most signifi-
are attracted by the wildlife and natural ecosystems
cant landscape changes have occurred in three set-
that still exist over large areas. The persistence of such
tings: (1) along rivers at low elevations, where irrigation
amenities can be attributed to a cool climate, rugged
water is available and people enjoy having their homes;
terrain, and extraordinary features that are highly
(2)in the foothills, where many homes have been con-
valued. Yellowstone National Park was established in
structed and fires have been suppressed; and (3) where
1872the worlds first national park. Nineteen years
the extraction of energy resources is feasible.
later, in 1891, a forest reserve was established nearby.
During the past 20 years, Wyoming and adjacent
It became the Shoshone National Forest. In 1906 the
states have experienced surprisingly rapid changes.
nations first national monument was established at
Still, there are many opportunities for learning about
Devils Tower in northeastern Wyoming, and Grand
the natural history of the Rocky Mountains and western
Teton National Park was established in 1929. Since
Great Plains. Bison have been replaced in most areas by
then fifteen wildlands in Wyoming have been desig-
cattle, but pronghorn, elk, and deer are still common.
nated Wilderness by Congress. National parks, wilder
Livestock have grazed some rangelands heavily, but
ness, and wilderness study areas account for about
bison may have done the same. Grizzly bears, elk, and
9percent of the states land (figs. 1.11.5).
wolves no longer wander across the basins and plains
In addition, native prairie and shrublands persist in
to the extent they once did, but all are found in some
some areas, primarily because 37 percent of the state is
areas.2 Roads, timber harvesting, and industrial devel-
above 7,000 feet elevation, where growing seasons are
opments have fragmented mountain forests and basin
too short and cool for cropland agriculture. At lower
shrublands, but not everywhere. The question is: How
elevations water is more likely the limiting factor, with
can the land be managed so that highly valued natu-
most agriculture requiring irrigation. Farmers tried vari-
ral resources are used wisely during a time of alarming
ous crops in the late 1800s, but their fields commonly
rates of habitat degradation and climate change? The
failed. The federal government purchased abandoned
sciences of ecology and conservation biology are perti-
homesteads in the late 1930s, and some became national
nent to answering this question.
4 Wyoming and the Rocky Mountain West
Fig. 1.1.Elevation map of Wyoming and neighboring parts of

Montana, South Dakota, Nebraska, Colorado, Utah, and Idaho.
The lowest elevations in the state are in the northeast. The
names of physical and cultural features are shown in figs. 1.2
and 1.4. Land cover is illustrated in fig. 1.5, and major rivers
are identified in fig. 4.1. The total area of Wyoming, including that portion of Yellowstone National Park in the state, is
97,814 square miles. The state is bounded by longitudes 1043
and 1113 and latitudes 41 and 45. Cartography by Ken
Driese.
Ecology
ing the interactions among plants, animals, micro-
Most ecologists would argue that it is important to

protect air and water quality, conserve biological
diversity, and maintain soil productivity. Yet advocacy
for env ironmental protection is not ecology per se.
Rather, ecology is a science dedicated to understand
organisms, humans, and their environment. The

interactions are studied to be understood, not to be
judged as good or bad. Ecologists strive to be objective
through systematic measurements, experimentation,
and analysis. Many also work to ensure that decision-
Introduction
LL
PL OW
AT S
EA TO
U NE
YE
O N MTS.
KIE
MT
S.
OW
LC
SP
CH
AR
RI
ER
D
IN
GRA
N
RAWLINS
UPLIFT
SHIRLEY
BASIN
HANNA
BASIN
LARAMIE
BASIN
DENVERJULESBURG
ME
BASIN
NORTH
PARK
BO
NE .
CI
DI MTS
TS.
UINTA M
.
MTS
BASIN
ROCK
SPRINGS
UPLIFT
RA
ER RE
SI AD
M
SS
MT
S.
DIVIDE
WASHAKIE
BASIN
FO
ITE
E
MI
RA
LA
OVERTHRUST BE
LT
TS.
GREAT
RIVER
IL SY
NCLINE
KM
S.
GREEN
BASIN
REE
WIND
RIVER
BASIN
MT
RIVER
BASIN
CA
ER
POWDER
FT
LI
UP
TE
T
TS
S
LL
HI
BASIN
K
AC
BL
BIGHORN
KA
VE GR
O
M NTR S
TS E
.
TS.
RN M
HO
IG
B
O
ABSA R
SH
WA
N
KSO
JAC OLE
H
WASATCH MTS.
PRYOR
MTS.
BEARTOO
TH
MT
S.
HAR
T
UPL VILLE
IFT
MADISONGALLATIN
MTS.
100 km
100 mi
Fig. 1.2. Mountains and basins illustrated in the previous

figure. See Love and Christensen (1985) for a detailed geologic
map of Wyoming. Adapted from Blackstone (1988) and Mears
(1993).
makers are provided with the best ecological informa-
to understand how organisms are adapted to their en
tion available.
vironment, whereas population ecologists seek to under-
The goal of modern ecological research, as in other
stand why population sizes fluctuate through time and
sciences, is to understand patterns and processes in
vary from place to place. Community ecologists focus
nature well enough to make predictions. This includes
on the interactions among coexisting species in a spe-
anticipating the effects of environmental changes on
cific areathe community. Others study ecosystems,
the distribution and population sizes of all kinds of life,
defined as areas where plants, animals, and microbial
and on the movement of water, nutrients, and energy.
organisms interact with one another and their environ-
Theoretical ecologists search for generalizations that
ment. Ecosystem ecologists commonly measure flows
apply everywhere. Others are empirical; that is, they
of energy, water, and nutrients through the soil, atmo-
focus on specific areas and address interactions asso-
sphere, detritus, and living organisms (fig. 1.6). Often
ciated with, for example, specific tracts of forests and
the innumerable species are grouped into green plants
rangelands. Each approach is beneficial for the other.
(the producers) that convert the energy of sunlight into
Modern ecology can be divided into numerous sub-
new plant material, herbivores that eat plants, carnivores
disciplines. To illustrate, physiological ecologists work
that consume other animals, omnivores that depend on
6 Wyoming and the Rocky Mountain West
Fig. 1.3. Land jurisdiction in Wyoming and adjacent parts of

neighboring states. Federal and private lands make up 48 and
43 percent of Wyoming, respectively. Most federal land is administered by three agencies: the U.S. Forest Service (national
forests and national grasslands), the National Park Service
(parks, monuments, and recreation areas), and the Bureau of
Land Management. Approximately 5 percent of the state has

been designated by Congress as Wilderness, all on federal
land. State and tribal lands cover 6 and 3 percent of the state,
respectively. State-owned lands are widely distributed in
small tracts and are not separated from private land on this
map. County lines are shown. Cartography by Ken Driese.
both food sources, and detritivores and decomposers that
how the abundance of different species and their influ-
derive their energy and nutrients from dead plants and
ences are affected by varying environmental conditions,
animals. As the populations of a community or ecosys-
and how they change with time following disturbances.3
tem change, other attributes change as well (for example,
Some ecologists specialize in the study of plants, whereas
rates of streamflow, photosynthesis, herbivory, and the
others concentrate on bacteria, fungi, insects, birds,
cycling of nutrients). Of interest to many ecologists is
mammals, or other organisms.
Introduction
Fig. 1.4. Highways, cities, and larger towns in Wyoming. The

two largest cities are Cheyenne and Casper, with populations
in 2012 of 62,000 and 58,000, respectively. Cartography by
Ken Driese.
Another branch of ecology is landscape ecology. Ecol-
objectives of a study or management challenge. The area
ogists working at this scale commonly focus on large het-
included may be as small as a pond or as large as several
erogeneous areas, for example, square miles of grasslands
counties or states, or even continents.
and shrublands. They are curious about how the land-
Ecology is a diverse science that fosters an improved
scape has changed in recent decades or centuries, and
understanding of all the various forms of life in an area
how it might change in the future. Another goal is to
and the ecosystems they inhabit. To facilitate ecologists
determine the causes and effects of different landscape
work, diagrams are drawn to illustrate the plethora of
patterns. Aerial photographs, satellite images, and maps
possible interactions, such as those illustrated in fig.1.6.
are important tools for their work. The terms landscape
Some of the drawings are similar to those prepared by
and ecosystem are sometimes used interchangeably. To
engineers, but there is a significant differencenatural
illustrate, the Yellowstone landscape made famous by the
ecosystems were not designed and constructed by
artist Thomas Moran in the 1870s is a mosaic of commu-
humankind. Rather, they emerged after millions of
nities and is now often referred to as the Greater Yellow-
years of evolution and centuries of ecosystem develop-
stone Ecosystem (see chapter 15). The boundaries of an
ment. Much remains to be learned about the species
ecosystem or landscape are established according to the
and processes that enable their long-term persistence.
Introduction
SOLAR RADIATION
Transpiration
Atmosphere
Photosynthesis
Heat
Herbivores
Leaves
Evaporation
to atmosphere
Carnivores
Fruit
Seeds
Omnivores
Rain
Snow
Surface
runoff
Nitrogen
fixation
Stems
Detritus
Roots
Mycorrhizae
Herbivores
Detritivores
Soil surface
Carnivores
Soil organic
matter
Soil
solution
Omnivores
Decomposers
Subsurface
runoff
Mineral
soil
Weathering
Fig. 1.6. Major components (indicated by boxes) and interactions (arrows) of a terrestrial ecosystem. Arrow width indicates
the relative amount of energy or water moving along a pathway. Temperature, water and nutrient availability, and growing season length determine the rates of transfer between
components. The irregular shapes indicate sources of water
and nutrients. Complex food webs exist above and below the
soil surface, both of which are linked by the organic matter
on the soil, known as detritus, litter, or mulch. Such simple
diagrams do not convey the complexity caused by the diverse
group of organisms represented by each box. Changes in one
component or process cause changes in others.
Conservation Biology
As for ecologists, conservation biologists can take several approaches to their research. In general, they are
experts on rarity and what, if anything, can be done
about it. Some species are rare because their habitat has
been degraded by human activity; others are naturally
rare and are found only in one or a few small areas. Ultimately, conservation biologists work to facilitate sound
management programs that lead to the maintenance or
recovery of threatened species, thereby reducing the need
for the strict mandates of the Endangered Species Act. In
Fig. 1.5. (left) Major vegetation types plus cultivated

land, lakes, reservoirs, and urban and industrial developments.Sagebrush-dominated shrubland is most widespread
(33 percent of the land area), followed by mixed-grass prairie
(18 percent) and lodgepole pine forest (7 percent). Some
categories are not shown, because they occur in patches too
small for the scale of this map. These include most riparian
woodlands, shrublands, and meadows; most subalpine meadows; most aspen groves; small woodlands of ponderosa pine
and limber pine; woody draws; playas with greasewood; small
cultivated fields; white spruce groves in the Black Hills; and
foothill grasslands and shrublands.Subheadings indicate associated species in the foothills (fh), the Greater Yellowstone
Ecosystem (gye), the Black Hills (bh), and the Sierra Madre
(sm).Adapted from the national land cover map (2011) of the
U.S. Geological Survey Gap Analysis Program. Land cover
percentages are from Driese et al. (1997). Cartography by Ken
Driese.
Wyoming, seven plants, six mammals, six birds, six fish,

and one amphibian are currently protected by federal law
or are under consideration for protection (figs. 1.71.9).
Others are of special interest because they are endemics, that is, they are found nowhere else.5 Such species
along with those that are common and widespread, and
all the varieties in each speciescompose the biological
diversity of an area.
The conservation of rare and endangered species
requires information gained by scientists in ecology as
well as other disciplines. More specifically, biologists
with expertise in evolution and species identification are
needed to verify that so-called rare species are verifiable
species and are indeed rare. Some of them are difficult to
10
Wyoming and the Rocky Mountain West
distinguish from related species that are common, which

leads to controversy about whether the restrictions of the
Endangered Species Act are needed. Do the advocates of
such action really know? The expertise of systematists,
taxonomists, and geneticists is required. Its clear, however, that scientists cannot accomplish the goal of conservation by themselves. Stemming the tide of habitat
destruction and extinction requires joining forces with
people from all walks of lifelandowners; policymakers;
nongovernmental organizations; and local, state, and
federal agencies. Collaborating with so many parties is
complicated but necessary as increased value is placed on
rare species and other limited resources. Notably, some
resources, though not traditional commodities, have the
potential of providing ecosystem services that can be
bought and sold.
Ecosystem Services
Earths ecosystems provide many benefits in addition
to habitat for wildlife and rare species. These benefits,
often referred to as ecosystem services, include
Fig. 1.7.The black-footed ferret (Mustela nigripes), a member of

the weasel family, was believed to be extinct until a few dozen
individuals were found in 1981 on a ranch near Meteetsee,
Wyoming. With the collaboration of private landowners,
biologists, and state and federal agencies, there are now about
1,000 ferrets living in twenty large prairie dog colonies in
Arizona, Colorado, Kansas, Montana, New Mexico, South
Dakota, Utah, Wyoming, Canada, and Mexico. Adults are
1824 inches long. The species is protected by the Endangered
Species Act. Photographed in Arizona by George Andrejko,
Arizona Game and Fish Department.
Fig. 1.8.The worldwide distribution of desert yellowhead

(Yermo xanthocephalus) is confined to several hundred acres
of desert shrubland in central Wyoming. A member of the
sunflower family, this plant was first discovered and named
by Robert Dorn in 1990 and is protected by the Endangered
Species Act. It grows to a height of about 12 inches. Photo by
Bonnie Heidel.
erosion control,
maintenance of soil fertility,
inhibition of invasive plants,
provision of clean water,
air filtration,
mitigation of droughts and floods,
provision of habitat for insects that pollinate crops,
detoxification and decomposition of waste materials,
sequestration of carbon dioxide,
control of agricultural pests, and
provision of meat from areas that are too dry to cultivate using standard agricultural practices.
Such benefits are provided by the diversity of native
plants, animals, and microbes that thrive in the harsh
environments of the region.
Introduction
fossil fuels, the value of ecosystem services is estimated

at trillions of dollars per year.6 The Farm Bill of 2008,
passed by Congress, called for measuring the environmental benefits gained from managing farm and ranch
lands more carefully.
Considering their value, it should not be surprising
that some ecological services now can be bought and
sold, thereby creating opportunities for landowners,
businesses, consumers, and governments to become
more directly involved with supporting sustainable
resource management. An example is the buying and
selling of carbon credits as a means of reducing carbon dioxide emissions to the atmosphere, a major
Fig. 1.9.The Wyoming toad (Anaxyrus baxteri), less than 2.5
inches long, is found nowhere else except in an area of about
2 square miles in the Laramie Basin. It has been raised successfully in captivity, but there are currently only thirty to
a hundred individuals living in the toads native habitat. It
is protected by the Endangered Species Act. Photo by Sarah
Armstrong.
contributor to global climate change. Similarly, the

Conservation Reserve Program of the U.S. Department
of Agriculture provides monetary incentives for farmers to establish perennial plant cover on erodible land,
thereby enabling carbon sequestration as well as soil
stabilization and water purificationpowered by solar
energy rather than fossil fuels. Other examples are discussed in chapter 18.
Ecosystem services generally have been undervalued

or overlooked, leading to inevitable and undesirable
In sum, whereas ecologists study adaptations, popu-
consequences. Their value becomes clear when the costs
lations, communities, and ecosystems, conservation
are summed for the time, labor, and energy required for
biologists strive to develop strategies for conserving our
water purification, herbicides, insecticides, fertilizer,
evolutionary heritage. And now, a newly emerging mul-
and the control of soil erosion. When economists cal-
tidisciplinary group of professionals seeks to identify
culate all the costs of replacing such services with mod-
and profit from investments in maintaining ecosystem
ern technology, much of which requires the burning of
services.
11
Chapter 2
Landscape History
Millions of Years Ago
ERA
PERIOD
EPOCH
Holocene
Rocks exposed in canyons, escarpments, and moun-
Quaternary
tains provide the data geologists need to interpret the
2.6
Pliocene
climate was tropical. The water receded and advanced

many times, depositing sand in some places and more
finely textured silt and clay elsewhere. Sandstones, silt-
CENOZOIC
(Age of Mammals)
changes have occurred. About 350 million years ago,

become Wyoming was covered by seawater and the
stones, claystones, and other kinds of sedimentary rock
Miocene
23
Oligocene
34
Paleocene
limestones. An example is the Madison Limestone that

MESOZOIC
(Age of
Reptiles)
forms many escarpments in the region today.
dragonflies with 30-inch wingspans. Primitive vascular

plants were abundant, including tree-sized clubmosses,
ferns, and horsetails. Fossilization was less frequent on
Fig. 2.1. Geologic time chart showing the ages (millions of

years ago) of eras, periods, and epochs. The Paleogene and
Neogene periods of the Cenozoic were previously known as
the Tertiary. Adapted from the International Commission on
Stratigraphy (http://www.stratigraphy.org).
12
PALEOZOIC
(Age of Fishes)
ginkgo, pine, spruce, fir, and sequoia. Mountain ranges
Jurassic
Triassic
Pennsylvanian
Mississippian
Devonian
Silurian
Ordovician
Cambrian
PRECAMBRIAN
primitive gymnosperms, such as the ancestors of cycads,
Cretaceous
Permian
the uplands, but there were woodlands that included
Eocene
56
marine organisms, rich in calcium carbonate, became
Also during the Paleozoic, shifting seas created enor-
Neogene
Paleogene
were formed. Skeletal fragments from coral and other
mous swamps and tidal flats that were frequented by
0.01
Pleistocene
development of western landscapes.1 Unimaginable

during the Paleozoic Era (fig. 2.1), the area that would
AGE
65
145
200
251
299
318
359
416
444
488
542
Landscape History
from the older rocks and washed into the newly defined
inter mountain basins.
The Mesozoic would have been especially exciting
WYOMING
IA
S
LAURA
for biologists, as dinosaurs were abundant in the stillequator
tropical climate. About forty different kinds are recognized in the fossil record from Wyoming alone.5 Some
were huge, suggesting that the vegetation was abundant
enough to provide large amounts of food. Others were
GO
NDW
ANA
LAN
small, less than a few feet long. These reptiles, along

with other nondinosaurian reptiles like the flying pterosaurs, were important residents of Earth for more than
Fig. 2.2. Approximate location near the equator, about 240

million years ago, of the land area now known as Wyoming.
Adapted from Lageson and Spearing (1988).
100 million years. Fossils of some of the most famous

dinosaurs have been found at Como Bluff, between the
towns of Medicine Bow and Rock River. They include
Diplodocus, Apatosaurus, Stegosaurus, Allosaurus, and
were uplifted and essentially leveled by erosion within
Dryosaurus.6 Bison, pronghorn, and other familiar
tens of millions of years.
mammals would not be present for another 200 million
By the early Mesozoic Era, about 240 million years
years.
ago, most of Wyoming was slightly above sea level
Coexisting with the dinosaurs were flowering plants
and still near the equator (fig. 2.2). Fluctuating coastal
(angiosperms), such as magnolia, palm, fig, breadfruit,
environments in the Triassic Period led to formation of
sassafras, cinnamon, sweetgum, and willow trees, along
the colorful sandstones and shales of the Chugwater
with tree ferns, giant horsetails, clubmosses, and vari-
and Spearfish formations, both rich in iron oxides and
ous kinds of gymnosperms.7 Each of these types of
known as redbeds. Subsequently, in the Jurassic Period,
plants had various species, some adapted to the low-
extensive sand dunes were consolidated into the Nug-
lands and others to the uplands. Some coal deposits
get Sandstone. Various episodes of volcanism occurred
began to form during this time, coincident with the
toward the end of the Mesozoic, during the Cretaceous
dinosaurs, but most coal in the Powder River Basin did
Period, leaving deposits of fine ash across the landscape
not begin forming until the Paleocene, about 60 million
and in the sea.3 Much of this ash became bentonite, a
years ago, long after the large reptiles became extinct.
sodium-rich clay of considerable economic and ecological importance.
The Mesozoic ended with a cataclysmic event that

many believe led to the worldwide extinction of dino-
Also during the Cretaceous, thrust faulting created
saurs and numerous other forms of life. Known as the
the Hoback, Salt River, Sublette, and Wyoming moun-
Cretaceous-Paleocene Extinction Event,8 its cause is
tain ranges, now located near the Wyoming-Idaho
still debated, but there is little doubt that it was dra-
border. These mountains, known collectively as the
matic, possibly lasting only a few hours. A widely
Overthrust Belt and now an important source of oil
accepted explanation associates the extinction with a
and gas, are composed of sedimentary rock. Another
large asteroid impact on what is now the north shore of
round of mountain building began about 75 million
the Yucatan Peninsula. This occurred about 65 million
years ago, during the late Cretaceous, with folding
years ago, spewing huge quantities of dust into space,
and faulting of Earths crust throughout the region.
greatly reducing the amount of sunlight reaching Earth,
For about 30 million years, Precambrian granite and
which created cold temperatures that many species
other igneous and metamorphic rocks, together with
could not tolerate.9
overlying sedimentary strata, were thrust upward. This
But recently, an alternative explanation associates
period of mountain building is known worldwide as
the extinction with heat rather than cold.10 According
the Laramide orogeny. The overlying Paleozoic and
to this hypothesis, Earths surface was heated by the air-
Mesozoic sedimentary strata gradually were eroded
borne material resulting from the asteroids impact. The
13
14
lofted dust and rocks fell to Earth through the atmo-
pine, redwood, spruce, sweet gum, sycamore, walnut,
sphere and were heated by friction to temperatures
and willow. Others, such as breadfruit and palm, are
higher than most unprotected animals could tolerate.
found today only in warmer climates. Some trees were
Fires would have burned across Earths surface wherever
large, up to 75 feet tall and 5 feet in diameter, as indi-
there was sufficient fuel. Supporting this hypothesis is
cated by fossil logs in Yellowstone National Park and the
the fossil record, which shows that, on land, only rela-
Shirley Basin south of Casper (see chapter 15).12
tively small animals survivedthose sheltered from
Coastal swamps were widely scattered during this
the heat in burrows or rock crevices. They included a
time, with cypress and other wetland treessimilar
variety of small mammals that had managed to coexist
perhaps to the cypress swamps of southeastern North
with the dinosaurs for millions of years. Also, protected
America today. Sphagnum bogs around the edges of lakes
by the water, aquatic life such as turtles and crocodiles
were also widespread. With a warm, subtropical environ-
survived. Could a comparable asteroid strike Earth
ment year-round, and an estimated annual precipitation
again? Over timespans of hundreds of millions of years,
of 100 inches of rain per year, plant growth was rapid.
the improbable becomes probable.
Moreover, the anaerobic conditions of the swamps and

bogs enabled preservation of dead plant material. With
The Age of Mammals

With the dinosaurs gone, mammals evolved to occupy
continued uplift and erosion from the adjacent mountains during the Laramide orogeny, the accumulating
peat was overtopped with deep sediments here and there
the vacated ecological niches. This marked the begin-
that compressed the plant material into coal. Today, most
ning of the Cenozoic. Early in this era, in the Paleocene
coal is too deep to mine, except where further uplifting
Epoch, the mammals included condylarths, creodonts,
and erosion occurred, leaving some of it near the sur-
tillodonts, pantodonts, and multituberculatesfor
face. That accounts for the abundant, easily accessible
which no common names can be found in the modern
coal in the Powder River Basinone of the worlds larg-
fauna. Most were small, described by paleontologists
est deposits. Some of the seams are more than 200 feet
as shrew-like, rodent-like, weasel-like, or beaver-sized.
thick. Because this coal developed largely in a freshwater/
Some of the herbivores had long, sharp front teeth,
estuarine environment, its sulfur content is low.13
possibly to assist with digging roots and tubers. They
Plant fossils in the Bighorn Basin have been studied
might have been omnivores. Though condylarths were
extensively. Scott Wing, a Smithsonian Institution paleo-
small in size, a branch of the group included the ances-
botanist, concluded in 1981 that 54 to 38 million years
tor of present-day hooved animals, such as horses, elk,
ago, during the Eocene, the basin had a subtropical cli-
and pronghorn. The pantodonts included the very first
mate with a dry season. The characteristic vegetation was
large mammals, which have been described as bear-
an evergreen broad-leaved forest with some conifers. For-
like or hippo-like herbivores. Early primates, rodents,
ests dominated by conifers most likely were characteristic
and coyote-sized carnivores also existed at this time in
of the mountains, but snowfall was probably a rare event,
the area that would become Wyoming. After another
and then only at high elevations. Many of the fossils are
10 million years of evolution, by the beginning of the
of plants that are common in the region today, whereas
Eocene Epoch, these forms, along with bats and oth-
others now occur naturally only in Southeast Asia, for
ers, had evolved into the verifiable early ancestors of
example, ginkgo and dawn redwood. Wing concluded
the mammals we recognize today.11
that the different kinds of plants were segregated into
Flowering plants also became increasingly common
different communities along gradients of temperature,
during the early Cenozoic. Some types of trees, shrubs,
water availability, and salinity. He estimated that the
and herbs found during this time, about 60 million
annual precipitation in the Bighorn Basin at that time
years ago, are still living in parts of North America and
was 60100 inches, much higher than the 424 inches
Eurasia. The trees included ancestors of alder, beech,
that falls there now. However, the shrub that became
birch, black locust, chestnut, cottonwood, cypress, dog-
known as greasewood was present, suggesting semi-arid,
wood, elm, fir, ginkgo, hickory, magnolia, maple, oak,
saline environments in some places.14
Landscape History
Even though the North American continent con-
Bighorn Mountains, Black Hills, and Uinta Mountains
tinued to drift poleward, the climate did not cool as
were formed. Another mountain range, known by geol-
would be expectedapparently because greenhouse
ogists as the Absaroka volcanic field, developed over a
gas concentrations in the atmosphere kept the region
shorter time, during the latter half of the Eocene. Dur-
about 18F warmer, on average, than today. Research
ing that time, the remains of entire forests became fos-
suggests that Earth warmed still further at the end
silized in what is now Yellowstone National Park, buried
of the Paleocene and beginning of the Eocene, about
by volcanic eruptions and subsequent flooding. Many
55million years ago. The warmest period, first reported
trees, now exposed by erosion, are still upright (see
in 1991 and now known as the Paleocene-Eocene
chapter 15).
Thermal Maximum (PETM), took about 10,000 years

to develop.
With erosion in the mountains and deposition in the
The PETM appears to have been associ-
adjacent basins, the topography was gradually flattened,
ated with an increase in the amount of carbon dioxide
broken here and there by hills of Precambrian granite
in the atmosphere, and it lasted for 100,000250,000
(fig. 2.3). Large rivers deposited prodigious amounts of
years. No mass extinctions of terrestrial organisms are
sediment during floods, and volcanism produced colos-
known to have occurred, though a large portion of lake
sal volumes of ash and volcanic debris. Both contributed
bottom protozoans disappeared. The climate became
to basin filling during the latter half of the Eocene and
significantly warmer and in some places more arid.
episodically throughout the Oligocene (encompassing
There would have been less plant growth. The fossil
45 to 23 million years ago). Also, huge freshwater lakes
record suggests that, with less food available, smaller
developed, covering 1025 percent of Wyoming at vari-
animals had a better chance of survival.
ous timesfirst Lake Lebo in the western Powder River
15
During the unusually warm PETM, the vegetation of
Basin (early Paleocene Epoch, 63 million years ago) and
present-day Wyoming shifted from dense forests, with
later Lake Tatman in the southern Bighorn Basin, Fossil
birch, sycamore, palm, magnolia, walnut, elm, laurel,
Lake near the eastern part of the Overthrust Belt, and
and dawn redwood, to more open dry forests, such as
Lake Gosiute in the Green River Basin and Red Desert
those found in parts of Central America today.16 Mem-
(early Eocene, 52 million years ago). Coal formed around
bers of the bean family, such as mimosa, became more
some of these lakes. Evidence for these lakes are fossilized
common, and the plants had smaller leavesan indica-
fish, now the primary attraction of Fossil Butte National
tion of a drier climate. Many tree species survived the
Monument near Kemmerer (fig. 2.4). Also found there are
PETM but were less common until after the heat sub-
fossils of stingrays, 13-foot-long crocodiles, banana trees,
sided, probably as carbon dioxide was reduced through
palm trees, water lilies, and numerous other plants and
storage in plant biomass, the oceans, and other parts of
animals.19 The climate was still warm and humid.
the biosphere.
Lake Gosiute is an example of how these now-extinct
The PETM is sometimes discussed in the context of
lakes affected todays vegetation patterns and land use.
current global warming, which is also associated with
This lakeshallow, saline, and without any mountains
increased concentrations of greenhouse gases. Much
nearbyfluctuated in depth as the climate changed
can be learned by studying the past, but Earth scien-
over 4 million years. It teemed with tropical life, in the
tists have calculated that the rate of warming today is
water and on shore. Eventually the Lake Gosiute eco
about fifteen times faster than at the end of the Paleo-
system led to the production of the worlds largest oil
cene.17 More rapid climate change provides less time for
shale and trona deposits, part of the Green River Forma-
organisms to adapt to their new environment through
tion. Trona is formed from sodium carbonate, sodium
evolution.
bicarbonate, water, and various impurities. As Craig
The Laramide orogeny, which began late in the Creta-
Thompson, professor at Western Wyoming College,
ceous, before the rise of mammals, continued well into
wrote, few passersby are aware that part of the steam
the Eocene. This mountain building produced modern
they see [from trona refineries near Green River] is water
configurations of the Wind River, Granite, Gros Ventre,
from Lake Gosiuteliberated and moving again in the
and Medicine Bow mountains (see fig. 1.2).18 Later, the
hydrologic cycle.20
15
Basin
Granite and
Related Rocks
Landscape History
As Lake Gosiute dried, continued erosion deposited

thick sediments over the salts and shales that had accumulated. The topography and climate continued to
change, until today the Chain Lakes wetland is located
in the bottom of the Great Divide Basin (see fig. 1.2),
surrounded by sagebrush steppe and desert shrubland
much different than the tropical life that existed there
in the past. The lakes occupy a small portion of the
depression where Lake Gosiute once existed.
Later, in the Miocene (about 10 million years ago),
regional uplifting became more pronounced, eventually bringing the landscape to near its present elevation
and initiating a new cycle of erosion. This uplifting was
accompanied by localized faulting and folding during
the Pliocene (5.0 to 2.6 million years ago) that led to the
formation of the Teton Rangethe youngest mountain
range in the region.21
Fig. 2.4. Fossil of a 16-inch fish (Diplomystus dentatus) that

lived about 50 million years ago in freshwater Fossil Lake, located near Kemmerer in present-day southwestern Wyoming.
Some fossil beds in this area are protected in Fossil Butte
National Monument. The fossils of palms and other tropical
plants and animals are found in the same area.
With uplifting, the climate cooled, and some rivers began to flow more rapidly, increasing the rate of
Pliocene, about 5 million years ago). 22 Drought-tolerant
erosion and eventually cutting canyons through the
plant species persisted or immigrated from nearby.
igneous rocks of buried mountain ranges (see fig. 2.3).
Some, such as sagebrush, needle-and-thread grass, and
Examples include the canyons of the Bighorn, Green,
bluegrass, migrated from the Old World across the Ber-
Laramie, Platte, Powder, Snake, Sweetwater, and Wind
ing Land Bridge that connected Siberia and Alaska at the
rivers. Most easily seen are the Snake River Canyon
time. Halophytic (salt-tolerant) plants found in modern
south of Jackson and the Wind River Canyon south of
inland salt marshes evolved from plants that are char-
Thermopolis. In addition to canyons, the accelerated
acteristic of marine coastal environments, such as salt-
erosion of claystones, siltstones, and other nonresistant
grass, alkaligrass, and cordgrass. Fires killed many of the
strata created badlands, a process that continues to this
trees that still grew on the upland, hastening the spread
day, carving escarpments in the elevated land.
of grassland and restricting woodlands to ravines, valley
As the mountains rose, the climate became more
bottoms, or ridges that burned less often. The grasslands
arid on the easternleewardside, caused by the rain-
created new ecological niches for mammals, and the pre-
shadow effect. This happens because the prevailing
decessors of all modern Wyoming animals appeared in
winds are from the west, and moisture is deposited on
the fossil record, including several kinds of pronghorn
the western slopes as the rising air cools and the mois-
that are known only from North America. Only one
ture condenses as it passes over the mountains. Along
species of pronghorn survives today. Also present were
with cooling, the drying climate caused the demise of
the camel, horse, mammoth, mastodon, rhinoceros,
some forests and favored the establishment or expan-
sabertooth cat, giant beaver, giant ground sloth, and
sion of shrublands and grasslands (beginning in the
short-faced bearall of which would become extinct in

North America about 10,000 years ago, a few thousand
years after the first humans arrived from Eurasia.
Fig. 2.3. (left) Intermountain basin development during the

past 70 million years. Ten to five million years ago the basins
were filled with sediments eroded from adjacent mountain
ranges. Since that time, rivers have cut canyons through
mountain ranges because of regional uplifting. Some basin
sediments have now eroded away, and the ancient mountain
ranges are again more fully exposed. See also chapter 17. From
Knight (1974, 1990). Drawings by Samuel H. Knight.
The Past 2 Million Years

Glaciers, warm interglacial periods, and erupting volcanoes characterized the Quaternary Periodthe past 2.6
million years. The first part of the Quaternary is known
as the Pleistocene Epoch or Ice Age. Snow accumulated in
17
Fig. 2.5.This landscape, with Gannett Peak in the background and Dinwoody Creek in the foreground, was buried
in ice about 21,000 to 16,000 years ago. Note the alpine
cirques and U-shaped valley, both formed by Dinwoody
Glacier. Woodlands of whitebark pine grow at treeline, and
forests of Engelmann spruce and subalpine fir occur on the

lower slopes. The riparian vegetation is dominated by various
species of willow (arctic, diamondleaf, Drummonds, grayleaf,
netleaf, and Tweedys ). At 13,809 feet, Gannett Peak is the
highest point in Wyoming. Photo by Scott Copeland.
Landscape History
Fig. 2.6 Jenny Lake in Grand Teton National Park is contained

by forested terminal and lateral glacial moraines at the base of
Cascade Canyon. The canyon is a U-shaped valley formed by
glaciers. The forests on the moraines are dominated by lodgepole pine, subalpine fir, and Engelmann spruce. Douglas-fir
and limber pine are common on the lower slopes of the
Tetons; subalpine fir and Engelmann spruce are abundant on

the north-facing slopes of the canyon, and whitebark pine is
common in the alpine zone. Shrublands dominated by mountain big sagebrush are found on the glacial outwash plains
below. The highest peak is Grand Teton (13,766 feet); the ele
vation of Jenny Lake is 6,779 feet. Photo by Michael Collier.
the mountains during cold periods, often forming drifts
about 7 times during the past 750,000 years (50 times
that persisted through the summer. More snow caused
since the mid-Pliocene). With each episode of melting,
higher reflectivity of solar radiation, lowering the tem-
widespread flooding shaped broad riparian floodplains,
perature further. Many snow masses were converted to
known as glacial outwash plains. Later these flood-
mountain glaciers.
plains became terraces and benches, such as along the
23
Far to the north, huge continental
glaciers moved southward, forcing the Missouri River
Snake River in Grand Teton National Park and along the
to flow into the Gulf of Mexico rather than into Hud-
Laramie River in the southeast (see chapter 4).
son Bay, as it had previously. Continental glaciers never
Huge freshwater lakes also formed, including Lake
reached Wyoming, but features such as terminal and
Bonneville in Utah, Lake Missoula in southwestern
lateral moraines, U-shaped mountain valleys, cirques,
Montana, and Lake Wamsutter in the Great Divide
glacial outwash plains, and kettle topography indicate
Basin of central Wyomingalong with the smaller Fre-
that extensive glaciers were present in most mountain
mont, Jackson, and Yellowstone lakes in northwestern
ranges and on the Yellowstone Plateau (figs. 2.5 and
Wyoming. 24 The grinding of rock by moving ice pro-
2.6). The nearest continental glacier was in present-day
duced fine glacial dust, which was washed down rivers
northern Montana.
and blown into the basins and plains. Deposition of this
During periods of glaciation, tremendous amounts
material, known as loess (pronounced lss), contributed
of water accumulated in the ice for thousands of years.
to the development of fertile soils in areas that now
Glaciers formed and then thawed during warm periods
support grasslands and shrublands. As the snow melted
19
20
each spring, mountain soils were subjected to flushing
Shoshoni, Sioux, and Ute.31 By that time the glaciers had
by large volumes of water, which eroded soil particles
receded into the mountains.
and slowed the accumulation of nutrients. Soil develop-
Learning about change during the Holocenethe
ment in the mountains was most rapid where soft rocks
past 10,000 yearshas been facilitated by the analysis
(such as shales) were exposed instead of harder rocks,
of fossil pollen and other plant fragments. In wetlands,
like granite, rhyolite, limestone, and some sandstones.
these materials settled and were preserved in anaerobic
Volcanic activity during this time included enormous
sediments for thousands of years. 32 Paleoecologists
caldera-forming eruptions followed by multiple lava
have learned to identify the kinds of plants from which
flows that created much of the topography seen in
the partially preserved pollen, cones, seeds, and leaves
Yellowstone National Park today (see chapter 15).25
came. Because sediments accumulated with time, the
Though commonly thought of as the Ice Age, the
deeper plant materials usually are older. Thus, changes
Pleistocene also had extended warm periods, even
in plant materials with depth suggest how the vegeta-
warmer than during much of the past 10,000 years. 26
tion of the surrounding land area has changed over
All or most of the ice melted at such times. Present-day
thousands of years.
glaciers formed about 400 years ago, during a period
Paleoecological studies in Wyoming and nearby pro-
known as the Little Ice Age (ca. 15501850). They are
vide evidence of continued change through the Pleis-
now small, confined to high mountain peaks, espe-
tocene to the present (fig. 2.7). 33 For example, plant
cially in the Wind River Mountains, and are now reced-
fragments collected from the northern end of the Teton
ing due to climate warming during recent decades (see
Range document the presence of ponderosa pine about
chapter 3).27
127,000 years ago, during the last interglacial period,
At the beginning of the Quaternary, 2.6 million
when the climate was warmer than it is today. This coni-
years ago, the flora of the region was similar to what it
fer is now absent from that area and the Yellowstone
is today.28 Forests dominated by familiar conifers were
region. Studies by Cathy Whitlock and her students at
common in the mountains, and shrublands and grass-
Montana State University describe the vegetation his-
lands were common in the basins and on the plains.
tory of this area over the past 30,000 years, during and
Cottonwood, aspen, bur oak, boxelder, paper birch, and
following the last glacial period (known as the Pine-
Rocky Mountain maple were present, but most other
dale). They concluded that ice covered the Yellowstone
species of broad-leaved trees, along with redwood and
region at times during the Pleistocene, 34 when the mean
cypress, have not been found in the fossil record for that
annual temperature was about 918F lower than it is
time.
today. Conifers were restricted to a narrow band in the

foothills, and tundra-like vegetation was widespread
Thousands of Years Ago
across the lowlands (fig. 2.8). Throughout Wyomings

basins, permafrost was widespread. Several of the preva-
The first people arrived in North America about 15,000
lent mammals at the time are now characteristic of the
years ago.
With their ingenuity, tools, and weapons,
Arcticcaribou, collared lemming, and barren-ground
they spread rapidly. Their hunting is commonly thought
muskox.35 Mammoth and mastodon roamed the inter-
to have been a primary cause of the extinction of large
mountain basins as well.
29
mammals, such as the mammoth, giant sloth, camel,
Whitlocks
research
indicates
that
Engelmann
horse, and various other species that existed at the time;
spruce was one of the first trees to colonize the areas
but contributing factors may have been climate warming
vacated by receding glaciers, but within a few centu-
and reduced food availability caused by droughts. The
ries it was joined by whitebark pine, subalpine fir, and
hunters became known as the Clovis people. They were
lodgepole pine. Such forests were widespread at middle
able to kill 10-ton mammoths with their stone-tipped
and high elevations throughout Wyoming until 11,000
weapons.30 Clovis descendants eventually organized into
years ago, when the climate became warmer and drier.
various tribes, including the Apache, Arapahoe, Black-
These conditions led to an upward shift in upper and
foot, Cheyenne, Comanche, Crow, Flathead, Kiowa,
lower treelines and the development of lodgepole pine
Landscape History
PINE
SAGEBRUSH
wetter
14
C yr
(ybp)
B.P.
3,520
SAGEBRUSH
PINE
drier
8 7 6 5 4 3 2 1 2 3 4 5 6 7 8 9 10 11 12
5,850
HOLOCENE
8,460
10,060
10,200
TRANSITION
25,540
22,300
20,880
21,890
PLEISTOCENE
21,190
26,640
Fig. 2.8. Fossil ice wedge exposed at the Rawlins landfill,
which suggests that a tundra-like environment existed during
the Pleistocene (Mears 1981, 1987). The rod is 3 feet long.
Elevation 6,900 feet. Photo courtesy of Brainerd Mears, Jr.
>29,000
TEPHRA
61,000 ybp
and Douglas-fir forest at middle elevations. Forest fires

became more common. In the past 6,000 years, the climate has generally cooled, allowing spruce, subalpine
fir, and whitebark pine to expand their range. This trend
was reversed during brief, warm, dry periods, such as
the Roman and Medieval warm periods, about 2,000
and 1,000 years ago, respectively. These intervals of past
Fig. 2.7.The ratio of pine and sagebrush pollen has changed

during the past 60,000 years. A higher peak on the left indicates that pine pollen was more abundant than sagebrush, an
indication that the climate was wetter; a higher peak on the
right indicates that sagebrush pollen was more abundant. Based
on data from Grays Lake, southeastern Idaho. 14C yr B.P. = years
before present as determined by carbon-14 dating. Adapted
from Beiswenger (1991).
drought provide important insights on how ecosystems

will respond to the warmer, drier conditions projected
for the future (see chapter 3).
A comparatively warm, dry period spanning 3,000
years in the early Holocene caused an expansion of
big sagebrush, greasewood, juniper, and grasses in
the lowlands. During this time, grasses covered some
north-facing slopes up to 7,900 feet elevation in the
mountains.36 Most mountain tree species, such as
Engelmann spruce, subalpine fir, and lodgepole pine,
survived at higher elevations, though they probably
21
22
also existed in relatively cool, moist valley bottoms at
the past 2 million years or so, now it is clear that Utah
lower elevations. Droughts during this time killed some
juniper arrived in southwestern Wyoming about 9,000
of the grasses that stabilized sand dunes in the region,
years ago, having migrated roughly 200 miles north-
resulting in more blowing sand and shifting dunes.
ward as the climate warmed following glaciation. 39 Even
There is evidence that the Platte and other large rivers
more surprising, ponderosa pine did not reach the Big-
were sometimes dry during this period. 37
horn Mountains of Wyoming until about 3,000 years
The work of paleoecologists also has been facilitated
ago.40 Pinyon pine did not arrive in the state until about
by the middens of bushy-tailed woodrats, often referred
700 years ago and is still found naturally only where
to as packrats, because they collect leaves, seeds, cones,
the Green River flows into Utah.41 This small tree also
bones, and other material and deposit them in their
occurs in northern Colorado, just south of Owl Canyon
nests (fig. 2.9). Not particularly tidy, the woodrats uri-
on the east side of U.S. highway 287.
nate and defecate on their middens time and time again,
University of Wyoming paleoecologist Stephen T.
creating hard masses that often persist for thousands of
Jackson and his colleagues studied middens in the
years. By identifying the plants represented in differ-
foothills of the Pryor Mountains, at an elevation of
ent layers of a midden, and studying middens of dif-
about 5,000 feet on the north side of the Bighorn Basin
ferent ages, scientists draw conclusions about how the
(fig.2.10).42 They observed the following changes. About
vegetation has changed where the woodrats have for-
10,000 years ago, when glaciers were retreating up the
aged. Radiocarbon dating provides a rather precise time
mountains, the climate in the foothills was cool and
frame, with one midden in northeastern Utah having
humid, as indicated by the presence of ground juni-
deposits that are 40,000 years old. 38 Studying vegeta-
per, Rocky Mountain juniper, chokecherry, wood rose,
tion change using pollen or middens found at treelines
and fernsall of which today are found primarily in
is especially instructive, because trees are sensitive to
relatively moist woodlands and shrublands. In more
water stress. Thus, climate change is easily detected as
shallow layers of the middens, deposited about 8,000
the tree fragments appear and disappear from top to
years ago, fragments of drought-tolerant plants were
bottom in the middens.
foundnamely, pricklypear cactus, saltbush, winterfat,
Middens have provided some surprising informa-
and later, Utah juniper. The plants represented in older
tion. First, although it is commonly thought that most
midden deposits were absent. From about 4,400 to 2,700
of the same plant species have been in Wyoming for
years ago, limber pine expanded into the area, for reasons that are not yet clear.43 About 2,700 years ago, more
arid conditions returned, as indicated by the absence of
Fig. 2.9. Bushy-tailed woodrats are 1118 inches long. About

half their length is tail. The rodents gather identifiable plant
materials for their nests, which may persist for more than
10,000 years. The nests are known as middens. Photo by Phil
Dotson/Science Source.
limber pine, presumably because it could not survive the

droughts. Utah juniper and curlleaf mountain-mahogany became more common, both of which are abundant
around the middens now.
This Bighorn Basin research hinges to a large degree
on the presence or absence of the various species of
shrubs and trees at lower elevations. Another study,
conducted in the Wind River Mountains by Patricia Fall
and her associates from Arizona State University, studied changes in alpine treeline using pollen data from
a lake at about 10,200 feet.44 They found that, prior to
about 11,300 years ago, with glaciers still nearby, the
climate was relatively cold, and the vegetation around
the lake was largely alpine tundra. However, by about
10,600 years before present, forests of whitebark pine,
Engelmann spruce, and subalpine fir surrounded the
Landscape History
Big Pryor
East Pryor
is often mentioned during discussions about causes of
U
ta
h
R
oc
ky
tn
.J
un
ip
Ju
er
ni
p
e
Ju
r
ni
pe
rp
ol
le
n
R
oc
ky
M
tn
.J
U
ta
un
h
ip
Ju
er
ni
p
e
Ju
r
ni
pe
rp
ol
le
n
present-day global warming (see chapters 3 and 18).46
landscape change by studying tree rings, soil profiles,
2000
anthropological data, and old journals and photographs. What do such records reveal about the past few
3000
Age (14C yr B.P.)
In addition to the study of pollen and plant fragments,

scientists from various disciplines have learned about
1000
centuries?
4000
First, tree-ring studies have documented ongoing

climate change (fig. 2.11), including droughts during
5000
8001300
6000
ad
that were far more severe than any since
that time. Tree-ring data collected by Steve Gray and his

associates at the University of Wyoming indicate that
7000
these extended droughts lasted 75 years or more and
8000
were long enough to cause changes in soil profiles.47 Soil

characteristics suggesting drought are associated with
9000
10000
Centuries and Decades Ago
fewer American Indian artifacts, perhaps because the

5
80
80
Relative abundance (percent)

Macrofossils (solid bars), pollen (open bars)
Fig. 2.10. Abundance of plant fragments from two species of
juniper at different times during the past 10,000 years, based
on data from woodrat middens at two foothill locations on
the north side of the Bighorn Basin. The length of the solid
horizontal bars indicates the proportion of all plant fragments
that originated from a single species, which reflects the abundance of each species in the surrounding area. The open bars
indicate the relative abundance (percent) of juniper pollen
(not distinguishable by species). 14C yr B.P. = years before present as determined by carbon-14 dating. Adapted from Lyford
et al. (2002).
number of bison was lower at such times. Since about

1300
ad,
Wyoming has experienced a relatively moist,
cool climate. Paleoecologists refer to the 300-year period

from about 1550 to 1850 as the Little Ice Agea period
when a 2F cooling of mean annual temperature was
apparently enough to cause further vegetation and soil
changes at the boundary between grasslands and big
sagebrushdominated shrublands.48 Grasslands were
more widespread during relatively warm, dry periods,
because sagebrush is less drought tolerant (see chapter7).
Compared to changes during the eras and epochs
of the more distant past, Wyoming landscapes have
changed relatively little during the past few centuries.
American Indians surely influenced the landscape,
lake. Moreover, the treeline was probably 300 feet
but the details are not well documented for most of
higher than it is today, suggesting that the climate was
the area.49 We can infer that they used small trees for
warmer at that time than it is today. This may have
building lodges, and that they started fires from time
persisted until about 3,000 years ago. Since then, the
to timesome burning over large areas. The adoption
alpine lake has been cooler and surrounded by a mosaic
of the horse in the 1700s greatly increased hunting
of subalpine meadow and tundra with scattered white-
efficiency. That and human-ignited fires probably had
bark pine, as is typical of the transition from subalpine
the most pronounced effect on western ecosystems.50
forest to tundra in this region. Dense forests persisted
Also, some 65,000 emigrants passed through Wyoming
only at mid-elevations, as they do today. Lodgepole
in 1850 alone, some bound for California to search for
pine became abundant during the past 3,000 years.45
gold, others headed to Oregon Territory for land and
Such studies indicate that the climate has changed
other opportunities.51 Military forts were built along
dramatically during the past 10,000 years, a fact that
the way. Some travelers decided to live in Wyoming
23
Proportion of western
U.S. in drought (%)
24
0
10
20
30
Wetter
40
50
60
Little Ice Age
Drier
70
800
1000
1200
1400
Year AD
1600
1800
2000
Black areas indicate drought. Note that the past 400 years
have been relatively moist. Long droughts of at least several
decades occurred before 1300 ad and were more severe than
the droughts of, for example, the 1930s. See also Wise (2012)
and fig. 3.5. Adapted from Gray et al. (2006).
Fig. 2.11. Drought occurrence and duration in the interior

western states during the past 1,200 years based on tree-ring
data collected from sites throughout the region. The horizontal line indicates the average proportion of the western
United States that experienced drought during this time.
Territory, becoming ranchers, farmers, miners, mer-
outstanding example and was used actively by Indi-
chants, railroad workers, telegraphers, educators, and
ans from about 500 to 200 years ago.55 Hunting pres-
industrialists.52
sure by Indians intensified after they acquired horses.
The first significant influence of EuroAmericans was
Even more bison were killed when the railroad arrived
53
in 18671868, with the tendency of early travelers to
Such trappers as Wilson Price Hunt, Jedediah Smith,
shoot large numbers of animalspart of the adven-
and Jim Bridger were lured to the region because beaver
ture. 56 Bison were essentially absent as a free-roaming
pelts were highly valued for the fabrication of felt used
animal by the 1880s, about the same time that domestic
through the trapping of beaver in the early 1800s.
for clothing, primarily mens hats. Their unregulated
livestock herds were increasing. By 1900 the population
trapping caused a sharp decline in beaver popula-
of all large mammals (bison, mule deer, whitetail deer,
tions. Soon, many beaver dams failedcausing soil
elk, pronghorn antelope, bighorn sheep, and black and
erosion and the development of gullies in some areas
grizzly bear combined) in Wyoming had been reduced
where ponds, meadows, and willow-dominated shrub-
to as few as 60,000. Only with the conservation move-
lands had existed previously (see chapter 4). Some early
ment fostered by President Theodore Roosevelt and oth-
explorers noted how the gullies could be a madden-
ers in the early 1900s did natural resource management
ing hindrance that would require traveling a circu-
improve so that big game populations in Wyoming
itous route of several miles just to make one mile in the
could recover. In 2013, the Wyoming Game and Fish
desired directioneven though the gullies were only
Department estimated that the state was providing hab-
a few yards wide. Dudley Gardner, author and arche-
itat for 528,000 pronghorn, 427,000 mule deer, 104,000
ologist, suggests that most gullies were the effect of
elk, 60,000 whitetail deer, 7,400 moose, 5,500 bighorn
trapping too many beaver, though he acknowledged
sheep, and about 4,000 bison (including Yellowstone
that other factors could have been involved, such as
National Park). For comparison, the state had about
an increase in wagon traffic and overgrazing by horses
576,000 people. 57
and cattle.54 Notably, the market for beaver pelts dimin-
A useful approach for determining landscape changes
ished rapidly in the late 1830s. Beaver are now common,
during the past 100200 years is to compare present-
though much less so than previously.
day conditions with those described in early journals.
Indians had subjected bison and other animals to
Robert Dorn, noted Wyoming botanist, ornithologist,
hunting for thousands of years, often using cliffs to
and natural historian, summarized salient observations
kill or trap their prey. The Vore Buffalo Jump, located
by trappers, explorers, and settlers in Wyoming during
along Interstate 90 in northeastern Wyoming, is an
18051878 and concluded the following:58
Landscape History
1. Bison were present in all parts of Wyoming, includ-
mer of 1886 and the devastating winter of 18861887,
ing subalpine meadows, but they were especially
which killed 4060 percent of the cattle in most of
common on the eastern plains. Evidence of heavy
Wyoming.62 Vegetation changes caused by livestock in
grazing by bison was common.
the late 1800s and early 1900s are difficult to assess,
2. Range fires occurred, but they were not mentioned

often; grazing by native animals may have mini-
but they surely were substantial and are still apparent

in some areas (see chapters 6 and 7).
mized fuel accumulation.
Forests also were subjected to new kinds of human
3. Forest fires were common.
influences in the 1800s. The Union Pacific Railroad, the
4. Big sagebrush was very common.
nations first transcontinental railroad, required rail-
5. Grizzly bears, black bears, wolves, and elk were com-
road ties, and lumber and timbers were needed along
mon in grasslands and shrublands of the basins and
the way for new towns and for mining gold, silver, and
plains but declined with increased hunting pressure.
copper. The remnants of slab piles can still be found,
6. Some streams were ephemeral and had steep-walled
indicating the locations of sawmills. Some forests were
gullies.
7. Grasshopper and Mormon cricket plagues occurred,
as they do now.
8. Vegetation patterns have changed very little since
the mid-1800s.
Human influences were minor when the early explorers were writing their journals in the 1800s. 59 Probably
fewer than 10,000 people resided in Wyoming Territory
at the time, less than 10 per square mile. However, more
than 350,000 immigrants traveled through Wyoming
along the Oregon Trail between 1841 and 1868, and
with the wagon trains came thousands of livestock.
These animals grazed along the way and sometimes were
concentrated in small areas. For example, one group of
cowboys worked unsuccessfully for five days to drive its
herd across the Platte River near Casper, and during this
time the grass was devoured for miles around.60 During the 1800s, the environmental impacts of people and
livestock were probably much greater along the Oregon
Trail than in the region as a whole.
essentially clearcut. Elsewhere, forests were thinned as

the tie-hackers cut only the trees they wanted. Railroad
ties typically were floated down creeks and rivers to railheads, especially during spring when water levels were
high. Such activity, along with placer mining for gold,
must have caused significant changes along some rivers
and creeks.
The first photographers provided another kind of historical record. Several studies have compared early and
modern photographs of the same areas. For example,
rangeland ecologist Kendall Johnson retraced the route
of William Henry Jackson, the famous landscape photographer who accompanied the 1870 expedition of
geologist Ferdinand Hayden across Wyoming, which
occurred prior to the cattle boom.63 Johnson located
and rephotographed 56 scenes in the 1980s, approximately one century later. The photo comparisons led
him to conclude:
1. Except in the relatively small area where the land had
been plowed, the grasslands of eastern Wyoming are
The effect of livestock became more widespread
very similar to those of 100 years ago. Overall, the
after the completion of the Union Pacific Railroad in
effects of livestock grazing appear not to have been
1869, which made western rangelands a source for
much different than the effects of native ungulates,
much of the nations beef, lamb, and wool. Large herds
although shifts in plant species composition may
of cattle were driven northward from Texas. Sheep were
have occurred.
imported by railroad, and some herds grew to 10,000
2. Big sagebrush has increased, decreased, or remained
animals. Investors and western stock growers became
about the same, depending on land use and site char-
wealthy. Livestock were forced to graze on marginal
acteristics, but it was abundant in the intermountain
rangelands, leading rangeland scientists John Mitchell
basins long before livestock arrived.
and Richard Hart to write, the land was filled entirely
3. Woodlands and forests on the uplands have become
with cattle.61 Fortunately for rangeland condition,
more dense, probably because of fire suppression,
the livestock boom ended in 1887, following a crash
but trees have not invaded adjacent vegetation types
in the beef market that coincided with the dry sum-
over large areas.
25
26
4. River bottoms have changed dramatically, caused
agricultural purposes, often accelerating erosion. Live-
by impoundments, irrigation, cultivation, livestock
stock have replaced bison almost everywhere, and many
grazing, and settlements; and some river margins
plant species have been introduced from other conti-
now have more trees than in 1870.64
nents, some becoming invasive weeds. Towns, cities,
5. Areas of sparse plant growth, such as desert shrub-
highways, coalmines, oil and gas fields, refineries, wind
lands, retain a nearly identical appearance to those
energy projects, power lines, dams, and reservoirs have
of 1870.
been built. Riparian landscapes and some grasslands
Photographs must be used cautiously in drawing conclusions, as they only provide snapshots in time of a
and forests have been altered most dramatically; trees

have been planted where there were none before. Some
small area.65
industrial impacts have accelerated to alarming rates in
Since Wyoming became a state in 1890, changes have
and western Great Plains. Still, many parts of Wyoming
continued to occur. Forest landscapes in many areas
and nearby states are little changed since American
have been fragmented by road building and timber
Indians first encountered white-skinned explorers from
harvesting, and some rangelands have been plowed for
the east.
the past 20 years, especially in the intermountain basins
Present-Day
Environments and Climate
Change
Chapter 3
Describing the environment in a way that is meaningful

for understanding the growth and abundance of plants,
animals, and other forms of life is one of the greatest
challenges faced by ecologists. Convenient numbers
are sometimes calculated, such as monthly or seasonal
averages of temperature, but organisms respond to the
full complex of interacting environmental factors
not to one individual factor or to statistical summaries. Moreover, extremes are usually more important
than averages, and young individuals are less tolerant
of extremes. Therefore, the distribution of a species
depends more on the environment prevailing when the
plants or animals were youngconditions that usually
are not known when the focus is on adult plants. Little
wonder, then, that correlations between environmental
variables and the presence or absence of a species are
sometimes hard to find. The challenge is to integrate
the diverse factors that determine whether an organism
will survive.
The environments of wetlands, grasslands, shrublands, forests, tundra, and other kinds of ecosystems are
discussed in subsequent chapters. Here, general patterns
are described, with emphasis on the effects of topography and elevation on precipitation, temperature, evaporation, growing season length, soil characteristics, and
the nature and frequency of periodic disturbances. All
of these variables interact in determining the nature of
the ecosystem in a specific areaand all are affected by
climate change, the topic of the last section.
Topography
Wyoming landscapes span an elevational range of 10,758
feet, from the lowest point on the plains at 3,125 feet,
where the Belle Fourche River flows into South Dakota,
to the top of Gannett Peak at 13,809 feet. More than
one-third of the state is above 7,000 feet, and 10 percent is above 9,000 feet (fig. 3.1). About fifty mountain
peaks are 13,000 feet or above, mostly in the Wind River
Mountains. Several major rivers originate in the state
and flow into the Missouri, Colorado, or Columbia river
drainages (see chapter 4).
Elevation affects all environmental variables. Temperature decreases as one climbs, whereas annual precipitation generally increases.1 These trends create a
cooler and relatively moist environment in the mountains. Some groups of closely related plants have different species adapted to lowland, foothill, montane, and
alpine environments (fig. 3.2). The effects of elevation,
however, are strongly moderated by the influences of
topography and the increased rates of evaporation that
occur at higher elevations because of lower atmospheric
pressure in the mountains.2 Thus, high south-facing
slopes that receive direct solar radiation throughout the
year sometimes are as dry as low-elevation deserts. In
contrast, north slopes at low elevationscooler because
of less direct sunlighthave vegetation normally found
at higher elevations. Such observations have led ecologists to conclude that topographic position and degree
of exposure to direct sunlight are more important than
27
28
Sheridan
Cody
Powder River Basin
Big Horn
Basin
Gillette
Worland
Newcastle
Wind River
Basin
Casper
Lander
Great
Divide
Basin
Green
River
Basin
Rawlins
Rock
Springs
Laramie
Cheyenne
9,000 ft
7,000 ft
5,000 ft
Fig. 3.1. General topographic map of Wyoming with the

names of selected towns and intermountain basins. Elevation
is given as number of feet above sea level. See fig. 1.2 for the
elevation in determining plant and animal distribution

patterns.
0
0
50
50
100 km
100 miles
names of mountain ranges and other basins. Adapted from

Martner (1986).
To the west of the Great Plains is a region of mountains and intermountain basins (see figs. 1.2 and 3.1).
Based on elevation, topography, geologic charac-
The Continental Divide passes through the middle
teristics, and location with regard to the Continental
of this region, splitting west of Rawlins to form the
Divide, several ecological regions can be identified. The
Great Divide Basin, which has no drainage to either
Great Plains characterize the eastern third of Wyoming,
the Pacific or Atlantic oceans. The Divide around this
with elevations ranging from 3,125 to 7,524 feet. Sum-
closed basin is relatively low, about 7,000 feet above sea
mer precipitation is more frequent on the plains than
level, and is characterized by sagebrush steppe rather
farther west. Grasslands are the most common vegeta-
than the coniferous forest or alpine tundra typical to
tion type, and there is evidence to suggest that bison
the north and south. For this reason, the Pony Express,
were more numerous there than in the Great Basin of
various wagon trails, and the first trans

continental
Utah and Nevada. With more bison, plants on the
railroad passed through the area in the 1800s. Most of
Great Plains evolved adaptations to withstand grazing
the basins range in elevation from about 3,960 to 7,260
from large herbivores; hence, they are more tolerant of
feet, with the highest basins in the south (see figs. 1.1
livestock grazing than are plants in the Great Basin.
and 3.1).
Present-Day Environments and Climate Change
ELEVATION (m)
1000
2000
3000
Fig. 3.2. Different species of the same plant
genus are often found at higher or lower
elevations, indicating different adaptations to
the varying environmental conditions along
mountain slopes. Note that some species have
broader environmental tolerances than others.
Pinus spp.
Whitebark pine
Lodgepole pine
Ponderosa pine
Pinyon pine
Limber pine
Artemisia spp.
Alpine sagewort
Mountain big sagebrush
Wyoming big sagebrush
Black sagebrush
Basin big sagebrush
Silver sagebrush
Salix spp.
Arctic willow
Planeleaf willow
Pacific willow
Coyote willow
Peachleaf willow
Elymus spp.
Scribner wildrye
Blue wildrye
Canada wildrye
Stipa spp.
Columbia needlegrass
Needle-and-thread
Green needlegrass
Poa spp.
Alpine bluegrass
Big bluegrass
Canby bluegrass
Sandberg bluegrass
3000
5000
7000
9000
11000
ELEVATION (feet)
Scattered throughout the Great Plains and inter-
Some of the more erodible sedimentary bedrocks,
mountain basins, and in the foothills of the mountains,
such as shales and siltstones, lead to the formation of
are topographic features best referred to as escarpments
classic badland topography (for example, Adobe Town
or ridges. They are usually composed of erosion-resistant
southeast of Rock Springs, Gooseberry Creek badlands
sandstones and limestones (fig. 3.3). The vegetation is
east of Meteetsee, Hells Half Acre west of Casper, the
distinctive, primarily because of a coarse, rocky sub-
Honeycomb Buttes northeast of Rock Springs, and the
strate with very little soil and because of drifting snow.
Wind River badlands near Dubois). Other topographic
Shrublands dominated by mountain-mahogany, big
features in the lowlands are sand dunes, arroyos (gul-
sagebrush, and skunkbush are common, as are wood-
lies), buttes, playas, deflation hollows, mima mounds,
lands characterized by juniper, limber pine, and pon-
glacial moraines, hogback ridges, bajadas (alluvial and
derosa pine (see fig. 1.5).
colluvial fans), canyons, nivation hollows, and braided
29
30
Fig. 3.3.On this sandstone escarpment, limber pine and

Rocky Mountain juniper grow only in cracks along fracture
planes where soil, water, and nutrients are available. This escarpment is part of the Frontier Formation and is located near
Muddy Gap, north of Rawlins and west of the Ferris Mountains. Wyoming big sagebrush and greasewood shrublands
are found on the adjacent fine-textured soils. Photo by Ken
Driese.
or meandering stream channels. The ecological sig-
sure of water availability.4 Where precipitation is higher,
nificance of such features is a recurring theme in this
such as in mountain forests, other factors may be more
book.
limiting (for example, temperature or the length of
Constituting another third of Wyoming, the moun-
the growing season). In all cases, water availability to
tains are characterized by lakes, streams, coniferous
plants is affected as much by the infiltration and water-
forest, aspen groves, subalpine meadows, and alpine
holding characteristics of the soil as by the amount and
tundra. Environmental factors vary greatly, with freez-
timing of precipitation.
ing temperatures possible every month of the year at
Throughout Wyoming and much of the region, the
high elevations. Geographers have classified the Lara-
mean annual precipitation varies by a factor of 10, from
mie, Medicine Bow, and Sierra Madre mountains in the
6 to 60 inches (about 15150 cm; fig. 3.4). The two dri-
southeast as part of the Southern Rocky Mountains and
est areas are the Great Divide Basin, near Wamsutter
most of the other mountain ranges as part of the Middle
on Interstate 80, and the northern part of the Bighorn
Rocky Mountains (see fig. 1.2). Only the Beartooth,
Basin, between Lovell and Powell. In general, the inter-
Madison, and Gallatin ranges of the Greater Yellow-
mountain basins in the western two-thirds of the state
stone Area are in the Northern Rockies.
are drier than the Great Plains to the east, with averages
of 612 and 1216 inches per year, respectively. The
Precipitation
As in other semi-arid environments, precipitation has
foothills and mountains receive 1660 inches, with the

highest annual precipitation in the southwestern corner
of Yellowstone National Park.
a significant effect on plants and animals, as well as
With prevailing westerly winds, the western slopes
on human activities. In a given temperature zone, the
of the mountains receive relatively more precipita-
amount of plant growth in grasslands and shrublands
tion, the result of condensation as air moves upward,
is typically correlated with rainfall or some other mea-
cooling as it crosses the mountains. The leeward

Fig. 3.4. Average annual precipitation, 19812010. From
Wyoming Water Resources
Data System, prepared with
PRISM Climate Group data,
Oregon State University
(http://www.prism.oregonstate.edu/terms.phtml).
Cartography by Christopher
Nicholson.
slopes and basins are drier than might be expected,
mountains varies greatly from one year to the next.
because descending air masses become warmer, which
This variation in Wyoming is not well correlated with
increases the potential for evaporation instead of pre-
El Nio and La Nia years, as seems to be true farther
cipitation. This is the well-known rainshadow effect.
to the south and north.6
Most Wyoming basins and plains are in a rainshadow,
The form, duration, and intensity of precipitation
with the relatively dry conditions ameliorated during
are as important as the amount. Rain that occurs dur-
years when more than average amounts of moist air
ing warm periods evaporates quickly, sometimes even
from the Gulf of Mexico reach the plains and moun-
before it wets the rooting zone of the soil. Heavier
tains. Similarly, a northerly flow of moist air from
rainstorms can cause infiltration down to 4 inches
the Gulf of California (the Arizona Monsoon) causes
or more, but plant water stress soon develops if sig-
added summer precipitation in southern Wyoming, a
nificant rainfall events are separated by more than
phenomenon that could account for the presence of
a week or two, as they usually are. In contrast, snow
Gambel oak and the extensive groves of aspen on the
common
ly accumulates during winter. Some snow
west slopes of the Sierra Madre (see chapter 10), where
will be sublimated, but there is the potential for
they grow much as they do in parts of Colorado and
deeper infiltration into the soil when the snow melts,
Utah. Some years have high amounts of precipitation,
at a time when less evaporation occurs because of
generally caused by flows of such air masses from the
lower temperature. Such deep percolation and storage
south and southeast. Other years are extremely dry,
enable plant growth well into the summer and seem
occurring when these air masses are diverted to the
important for the survival of big sagebrush in most
east. As with rain in the lowlands, snowfall in the
intermountain basins. Not surprisingly, big sagebrush
31
Palmer Drought Index
32
wet
Wet
Little Brooklyn Lake
Early 20th Century:

Wetness
Pre-1900: Multiple droughts >15 years

many of greater severity than 1950s
10361 ft
48 in
1
0
1
2
3
Dry
dry
900
Late 20th Century:

Relatively wet &
no major droughts
50+ year
megadroughts
1000
1100
1200
1300
1400
Year
1500
1600
1700
1800
1900
2000
Foxpark
9065 ft
Fig. 3.5. (above) Drought history for the past 1,100 years in the Green River Basin of southwest
Wyoming, as reconstructed from tree rings. The graph shows estimated values for the Palmer
Drought Severity Index, a measure of drought severity. Positive values of the index represent
relatively wet conditions; negative values indicate drought. Each point on the graph represents the mean over a 25-year period. See also fig. 2.11. Based on data from Cook et al. (2004);
adapted from Gray and Andersen (2009).
Fig. 3.6. (right) Climate diagrams for southeastern Wyoming that illustrate climate change with
elevation. The graphs show mean monthly precipitation (top line) and mean monthly temperatures from January to December. Black areas indicate periods of drought, when the precipitation
line drops below the temperature line. Each unit on the vertical axis is 20 mm of precipitation
(water equivalent) or 10C. The number at the top of each diagram on the left is elevation (feet);
on the right, mean annual precipitation (inches). A break in the horizontal bar in the summer
indicates the frost-free period; a black bar indicates that the mean daily minimum during a
month is below freezing, and a hatched bar indicates months when the lowest temperature is
below freezing. On the vertical axes, to convert millimeters of precipitation to inches and degrees Centigrade to degrees Fahrenheit, see appendix A. Based on data from Martner (1986) and
the Wyoming Water Resources Data System at the University of Wyoming.
Laramie
7267 ft
Cheyenne
6125 ft
occurs most abundantly in the basins of the western

two-thirds of the state, where the proportion of snow
is higher.7 Proportionately less snow and more spring
as the lowlands, with those of the 1930s, 1950s, and

the first decade of this century lasting up to 10 years.8
Much longer droughts have been identified using treering research in Wyomings Bighorn Basin, with some
lasting 50 years or more prior to 1700 (fig. 3.6).9 Could
multidecadal droughts occur again? Understanding the
effects of drought on ecosystems and local economies
is a logical step toward developing a strategy for adapting to the droughts of the future. Even today, water is a
limiting factor for economic growth.
13 in
C
0
(fig. 3.5), where big sagebrush often is restricted to

Periodic droughts characterize the mountains as well
10 in
mm
and summer rainfall occur in the eastern grasslands

places where snowdrifts develop.
16 in
MONTH
Fig. 3.7. Average maximum temperatures, 19812010. The

highest maximum temperatures correspond approximately
with the longest frost-free periods and the highest rates of
potential evapotranspiration. From Wyoming Water Resources
Temperature, Frost-Free Period, and Evaporation

The July mean daily high temperature ranges from 90F,
such as on the Great Plains and in the Bighorn Basin, to
less than 75F in the higher mountains.10 The July mean
annual low temperature is 55F on the plains and near
freezing in the alpine zone, where frost at night is common even in the summer (fig. 3.7). During a 24-hour
period, variation is most extreme in the alpine tundra,
where air temperature near the ground may be 80F or
more during the day and below freezing at night. Such
extreme variation is caused by rapid heating of the soil
during the day (stemming from reduced water vapor in
the atmosphere of the alpine zone), and, for the same
reason, rapid cooling at night. Although most plants
can acclimate to cold temperatures, many cannot adapt
to such extreme diurnal cycles. Notably, the whole
region is comparatively high and dry, with relatively
Data System, prepared with PRISM Climate Group data,

Oregon State University (http://www.prism.oregonstate.edu/
terms.phtml). Cartography by Christopher Nicholson.
rapid warming and cooling even in the lowlands, especially when there are few or no clouds.
Historically, the mean frost-free period varies from
125 days to fewer than 25 days, with the longest periods occurring in the Bighorn Basin, in a small area near
Riverton in the Wind River Basin, and on the eastern
plains (see fig. 3.7). Such areas have the largest amount
of cropland. Crop plants generally do not tolerate frost,
though many native plants do. Significantly, although
temperature has an important effect on the initiation of
plant growth in the spring, the termination of growth
is more often caused by drought, especially in the lowlands where precipitation is lower. Native plants can tolerate short growing seasons; most crop plants require
irrigation. Fallow dryland farming is an alternative,
which allows such crops as wheat to benefit from 2years
of soil moisture accumulation rather than 1 year.
33
34
Topography influences growing season length and
chapter 8). The greatest potential for wind-generated
mean temperature by causing the flow of heavier cold
electricity is in the south, especially in the southeastern
air from mountain slopes to lower elevations, often
quarter of the state.14
along waterways. This airflow can create temperature

inversions, that is, warmer temperatures on mountain
slopes than in the adjacent lowlandsthe reverse of what
Soil Characteristics
is normally expected. Thus, the cities of Jackson and
Plant growth, crop production, and the carrying capaci-
Cheyenne are at about the same elevation6,209 feet
ties for wildlife and livestock depend on the nature of
and 6,053 feet, respectivelybut Jackson is much cooler
the soila matrix of organic matter, clay, silt, sand,
because of its location in Jackson Hole, which traps cold
and stones, teeming with microorganisms, inverte-
air flowing downward from the nearby Tetons.
brates, and roots, and mixed periodically by burrowing
11
One of the most significant effects of temperature is
mammals, such as prairie dogs, ground squirrels, and
on the potential rate of evapotranspirationevaporation
gophers. Indeed, the amount of energy consumed by
from soil and open water plus the amount of water
belowground organisms in grassland ecosystems can
evaporated on the interior of plant leaves and lost as
exceed the amount used by animals aboveground (see
vapor through the stomata. Higher temperature and
chapter 6). Every spoonful of soil contains millions of
lower humidity cause higher evapotranspiration, which
organisms, including bacteria, fungi, yeasts, protozo-
leads to summer drought stress for plants and animals.
ans, nematodes, and insect larvae. These mostly minus-
The potential evapotranspiration, which can be calcu-
cule organisms contribute immensely to the structure
lated from temperature and solar radiation, generally is
and fertility of the soil. Food webs belowground are
greater than annual precipitation in the plains, basins,
equally as complex and interesting as those above
and foothillsindicating dry conditions, because more
ground, though much more difficult to study.
water can be evaporated than usually is available. In con-
A multitude of different soils exists in the region,
trast, potential evapotranspiration in the mountains is
caused by the interactions over thousands of years of
less than annual precipitation, creating relatively moist
varying climate, topography, rock substrate (parent
conditions and the potential for streamflow. In the
material), and plant and animal life (see appendix B).
mountains, where more precipitation usually falls than
Sometimes the soils change gradually from one place
evaporates, there is the potential for loss of nutrients as
to another, forming a gradient from comparatively dry
they are dissolved in water moving through the soil pro-
grasslands to relatively moist meadows. In other places,
file, a process known as leaching. Only about 10 percent
the soils change abruptly because of abrupt changes
of Wyoming has a precipitation-evaporation ratio greater
in topography or the rock type from which soil min-
than one.12 Climate change surely will alter the precipi-
erals are derived. Some soils are shallow and not well
tation-evaporation ratio, which will lead to subsequent
developed, largely because of recent glaciation in the
changes in all ecosystemsincluding croplands.
mountains, or an abundance of erodible slopes and a
Topography and wind play an important role in

controlling water availability. South slopes are warmer
comparatively dry, cool climate not conducive to soil

formation.
than north slopes, enabling more evapotranspiration,
The physical and chemical characteristics of soils are
and wind accelerates evapotranspiration and leads
heavily influenced by the nature of the parent material,
to snow redistribution and sublimation. Areas blown
whether bedrock or transported sediments. Igneous and
free of snow are drier during the growing season; areas
metamorphic rocks (such as granite, basalt, quartzite,
where snow accumulates are wetter. Strong winds are
and rhyolite) are found primarily in the mountains.
common in Wyoming, especially in the south, where
They weather slowly, and consequently the soils are
the prevailing westerly winds are funneled through
usually shallow and coarse. Glacial moraines also lead
the Rocky Mountains at a low point in the Continental
to the development of coarse soils. Sandstones and
Divide (generally along Interstate 80).13 This wind pat-
limestones are resistant to weathering, often forming
tern has contributed to the development of a narrow
escarpments and ridges with shallow soils. Where suf-
band of sand dunes that is 100 miles long or more (see
ficient weathering has occurred, sandstones contribute
to the formation of a coarse-textured soil with a high
aeration, and salinity. Infiltration is especially impor-
infiltration capacity for water. Sometimes the resulting
tant in semi-arid regions, where the potential for evapo-
sand is sorted by wind into extensive dune fields, such
ration is high due to relatively warm, dry air. Water that
as northeast of Rock Springs and between Shoshoni and
percolates rapidly to a depth of 2 inches or more evapo-
Casper (see figs. 1.5 and 9.1).
rates more slowly, remaining in the soil long enough
Fine-textured and deeper soils usually develop from
to be used by plants. Coarse-textured soils in semi-arid
more easily weathered shales, mudstones, and siltstones,
regions tend to have higher infiltration rates and con-
or from material transported by wind and water (such as
sequently higher rates of plant growtha principle
on much of the western Great Plains). Infiltration rates
referred to as the inverse texture effect. The name stems
are slower in fine-textured soils, but the water-holding
from the fact that, in eastern North Americawhere
capacity is higher. Such soils can be quite fertile,
the precipitation is generally higher than in the low-
though some are saline or high in toxic elements (such
lands of Rocky Mountain statescoarse-textured soils
as selenium). Saline playas develop where fine-textured
sustain less plant growth than those with fine texture
sediments accumulate in closed basins with no or little
(more silt and clay).15
outflow. Some shale-derived soils (for example, from the
The principle of the inverse texture effect can be
Mowry, Steele, Belle Fourche, Pierre, and Thermopolis
extended to rock outcrops, where the soil is thin or even
formations) are high in bentonitea clay that swells
nonexistent but where water is funneled into cracks.
and shrinks during wetting and drying cycles, which
Plant roots capable of growing through these fissures,
typically prevents the establishment of young plants.
such as the roots of mountain-mahogany and limber and
Similarly, soils high in soluble salts, gypsum (hydrated
ponderosa pine, probably have more water available to
calcium sulfate), and sodium may form crusts on the
them than would be the case for adjacent fine-textured,
surface that slow the rate of plant establishment (see
deeper soils. In fact, shrubs and trees throughout the
chapter 8).
region, which generally require more water than do
Predictably, soil characteristics change with eleva-
herbaceous plants, are often found on rock outcrops
tion. Organic matter content increases from arid sites in
or coarse soil, not on fine-textured soils, where water
the basins up to montane meadows, but then decreases
often evaporates before plants can use it (see fig. 3.3).
farther up into the alpine zone. Plant production is
Mountain-mahogany and other shrubs that grow in such
highest at mid-elevations, where more moisture is avail-
places are an important source of food for wildlife.
able but the climate is still not as cold as higher on the
Water-holding capacity is determined by soil tex-
mountain. Severe climates, such as in the alpine tun-
ture, soil depth, and the amount of organic matter.
dra, tend to restrict soil development as well as plant
Fine-textured soilswith a relatively high percentage of
growth.
silt, clay, and organic matterhold more water against
Salts and nutrients accumulate in the soil where
the flow of gravity than do coarse-textured soils. They
the precipitation-evaporation ratio is less than one, as
also enable more plant growth if the soil is saturated
would be expected during most years in intermountain
at least once during the year, such as often occurs near
basins and on the Great Plains. A layer of lime (calcium
snowdrifts. Abrupt transitions between grasslands and
carbonate) develops near the average depth of water
some shrublands are often associated with changes in
infiltration, forming a hardpan or caliche layer that
soil depth or soil texture (see figs. 17.9, 17.11, and 17.17),
may restrict root development. In contrast, a hardpan
with the more drought-tolerant grasses abundant on
does not develop in the mountains where the precipita-
shallower soils with a low water-holding capacity (see
tion-evaporation ratio is greater than one and the soils
chapter 7).
tend to be slightly to strongly acidic. With more water
Soil aeration is determined by texture and the
moving downward through the soil profile, the poten-
amount of water in the soil; it is important to plants
tial for nutrient leaching is higher.
because air spaces in the soil have the oxygen required
Five soil features seem especially critical in deter-
for root respiration. Oxygen is less readily available if
mining vegetation patterns in Wyoming and adjacent
the spaces are filled with water, which occurs more
states: infiltration rate, depth, water-holding capacity,
often in fine-textured soils or in wetlands. Greasewood,
35
Blue grama
Gardner saltbush
Big sagebrush
20
80
30
clay
40
60
sandy
clay
clay loam
sandy clay
loam
20
60
silty clay
loam
70
80
loam
silt loam
10
loamy
sand
sand
100
50
silt
silty
clay
cla
y
10
90
50
40
30
Fig. 3.8. Surface soil texture has a strong influence on plant

distribution, as shown here for a grassblue gramaand
two species of shrubs, Gardner saltbush and big sagebrush.
Adapted from Nichols (1964).
100
70
36
90
80
90
sandy loam
70
60
silt
50
40
30
20
10
100
% sand
a common shrub around the edges of alkali flats, is well
The weathering of some rock types leads directly to
adapted to saturated soils with low oxygen, whereas big
the formation of soils with high salt concentrations,
sagebrush cannot tolerate standing water within 10 cm
especially Cretaceous marine shales, such as those in
of the surface.16 Well-drained, well-aerated soils are nec-
the Steele, Niobrara, Cody, and Pierre formations. Saline
essary for the survival of big sagebrush and the growth
soils often develop in basins where considerable water
of crops (fig. 3.8).
accumulates on the surface during wet periods and
Salinity, and other chemical traits such as alkalinity
where subsequent evaporation leads to the accumula-
and sodium concentration, depend on the composition
tion of salts on the surface, often forming white crusts.
of the soils parent material, the amount of water that
Soils derived from bentonite tend to have high con-
has evaporated from the soil over the centuries, and
centrations of sodium (because bentonite is composed
little or no potential for the leaching of salts from the
largely of montmorillonitic clay, which develops in
soil (that is, where the precipitation-evaporation ratio is
high-sodium sedimentary environments).
less than one).17 Plants that tolerate saline, alkaline, and

high-sodium soils are known as halophytes, for example,
greasewood, saltgrass, and the various species of salt-
Periodic Disturbances
bush. Less salt-tolerant species, such as big sagebrush,
The environment is usually described in terms of tem-
are sometimes found growing with halophytes but usu-
perature, wind, water availability, and soil characteris-
ally in areas where salt concentrations are lower (along
tics, but just as important are the various disturbances
drainages where snow accumulates and some of the
that have affected ecosystems for millennia. Examples
salts are dissolved and washed below the rooting zone;
include fires; windstorms; floods; burrowing; epidemics
see chapter 7).
of certain insects; and, in the alpine tundra, frost heav-
ing of soils.18 Attempts to suppress disturbances may
re-established by sprouting. In the same way, grasslands
create an environment in which some native species
also recover in a short time, with little plant mortal-
cannot survive.
ity. In contrast, shrublands dominated by big sagebrush
Fire is a well-known natural disturbance that can
may not recover for 1020 years, simply because big
occur when there is sufficient fuel. Long before humans
sagebrush lacks the ability to produce new stems and
learned to use fire, lightning strikes provided the neces-
leaves (shoots) by sprouting from its root system.
sary ignition, even in grasslands. The number of light-
Windstorms constitute another periodic distur-
ning strikes to the ground is surprisingly large, often
bance. Individual trees are often toppled, creating small
several hundred during a single summer storm. Not all
canopy gaps important to the survival of some under-
of them cause fires, but once started, prior to the time of
story species. Occasionally, large numbers of trees may
active suppression, the fires probably burned for weeks
be blown down, as occurred in the Teton Wilderness in
or months over large areas. Native Americans frequently
August 1987 and the Routt National Forest in October
started fires, probably most often to facilitate their hunt-
1997. In a few minutes, the trees over large areas were
ing of big game. Whatever the source of ignition, early
broken or uprooted, creating a landscape reminiscent of
explorers often saw smoke on the horizon. Meriwether
that after the eruption of Mount St. Helens in Washing-
Lewis wrote about fire and the scarcity of timber along
ton; all other environmental factors are affected. Evi-
the Missouri River in 1805:
dence of windstorms is easy to find in mountain forests.
This want of timber is by no means attributable to

a deficiency in the soil to produce it, but owes its
origin to the ravages of the fires, which the natives
kindle in these plains at all seasons of the year. The
country on both sides of the river, except some of
its bottom lands . . . is one continued open plain,
in which no timber is to be seen except a few . . .
clumps of trees, which from their moist situation, or
the steep declivities of hills, are sheltered from the
effects of fire.19
Fire and windstorms are physical disturbances, but

biotic disturbances are just as important. Such burrowing animals as prairie dogs, pocket gophers, badgers,
and harvester ants create disturbances that often cause
the death of some organisms while providing a favorable environment for others (see chapter 6). The gaps
created in this way are individually small, but combined they modify the environment over large areas.
Furthermore, burrowing animals move about on the
landscape, subjecting a large proportion of the land
to burrowing over long periods. Such disturbances are
Fires could be annual events in some of the more pro-
sometimes a nuisance for livestock owners, but there
ductive grasslands, or they could be comparatively rare
are beneficial influences as wellbetter infiltration of
in some high-elevation forests. In the foothills, a fire
water in a semi-arid environment, better aeration, mix-
might be expected every 525 years. Modern-day sup-
ing of organic matter deeper into the soil, a higher level
pression of grassland fires creates a favorable environ-
of biological diversity, and generally more plant growth
ment for the establishment and growth of some trees
(see chapter 6). Though difficult to value monetarily, all
and shrubs.
are examples of ecosystem services.
All fires are conspicuous disturbances, but some are
Population explosions of certain insects also create
more severe than others. Crown fires in forests become
disturbances. The insects most commonly involved in
extremely hot and burn through the treetops, killing
the lowlands are grasshoppers and Mormon crickets,
most trees and abruptly changing the nature of the
which can reach densities of 525 per square foot in
ecosystem. In contrast, surface fires that burn slowly
grasslands and shrublands in some years, consuming
through the forest undergrowth often consume only
most of the herbaceous plant material in a short time.
the shoots of small plants and the organic material on
Similarly, several kinds of bark beetles have histori-
the forest floor; most mature trees survive, as do most
cally killed trees over large areas from time to time. In
shrubs, grasses, and forbs, because their roots and
the past 20 years the beetles have affected forests over
rhizomes remain alive. Small trees may be killed, but
a larger area than ever before observed, probably the
within a year or two, most of the herbaceous plants are
result of climate change. Aside from influencing spe-
37
38
cies composition, insect outbreaks accelerate nutrient
of an area over a period of several decades or more. They
cycling and may affect forest flammability (see chapter
point out that unusually cold days will occur during the
11). Notably, these insects are native species and have
warmest month on record and that above-average pre-
coexisted with their hosts for millennia.
cipitation can fall in the middle of a 10-year drought.
Overall, an environmental analysis for any area must
Climatologists also emphasize that climate trends
consider the interaction of many factors that change
will not be the same everywhere. Even with an over-
either gradually or abruptly across landscapes and over
all trend of warming on Earth as a whole, some areas
time. No two areas have exactly the same environment;
will be cooler or wetter than normal. For this reason,
generalizations must be made with caution. This com-
the term climate change is preferred, rather than climate
plexity makes the environment difficult to conceptual-
warming or global warming. The rate of change seems
ize, but it is often helpful to focus on a few variables.
slow to some, with an increase of only a fraction of a
Key variables in the region include soil characteristics
degree every decade when warming does occur, but
(especially infiltration rates, water-holding capacity,
such small increases can lead to a higher probability
and salinity of the soil), water availability in the soil,
of summer drought if there is no concomitant increase
and seasonal and elevational temperature differences.
in precipitation. The result is slower tree growth, more
Periodic disturbances by fire, wind, burrowing animals,
frequent forest fires, and less streamflow from the
and a few insect species further modify an already
mountains. Also, as will be discussed, diseases and
diverse landscape mosaic. Regionally, different climates
insect epidemics can spread in ways that have not been
have caused the formation of grasslands, shrublands,
observed before. And, with warmer temperatures, the
forests, and tundra. Locally, the nature of the vegeta-
atmosphere can hold more moisture, thereby providing
tion is dependent on soil characteristics; topographic
the energy for extreme weather events. Flooding and
differences; snow drifting; and such disturbances as
snowstorms could be more severe.
fire, flooding, and windstorms.
Computer models generate specific predictions about

climate change and until recently have focused on large
Climate Change in Western States
areas. Some observers are skeptical of this approach,

but such models are required to synthesize all the geo
Climate change is nothing new. As described in the pre-
physical, atmospheric, and ecological data and processes
vious chapter, tropical forests once grew in Wyomings
that are involved. Modeling is used with confidence
Red Desert; millions of years later, glaciers extended
for weather forecasting and launching space vehicles.
down to the foothills. However, the prevailing notion
Warming projections based on models suggest that the
has been that such changes are so slow that they hardly
mean annual temperature in Wyoming and adjacent
mattered at human time scales. There were admonitions
states will increase by about 6F by 2100, perhaps 7F in
to be prepared for occasional droughts and cold winters,
the southwestern part of the statewarmer than at any
but few considered long-term trends. Now, during the
time in the past 10,000 years.21
past 3040 years, worldwide records reveal increases in
Computer modeling has been evolving for decades
air temperatures, and, not surprisingly, satellite images
as a tool for understanding complex systems, but such
have documented a steady, dramatic decline in Arctic sea
models should be tested when possible with actual mea-
ice. Similarly, repeat photos show how mountain glaciers
surements from specific places over long periods. Bryan
are retreating wherever they still occur. Rocky Mountain
Shuman, paleoecologist and professor of geology at the
glaciers are no exception.20
University of Wyoming, analyzed such data for Wyo-
Questions about the validity of warming trends are
ming.22 He used records from 30 weather stations for
often asked when record low temperatures occur, or
the period 18952007 to determine changes in three
about a drought after a year of heavy snowfall. Clima-
variables: annual average temperature, the high tem-
tologists emphasize that there is a difference between the
perature on the coldest day of the year, and winter pre-
weather that most people think about on a daily basis,
cipitation (NovemberMarch). His results, summarized
and climate, which is defined by the average conditions
in fig. 3.9, show an increase since 1978 in the frequency
Mean Annual
Temperature
2012
45.6F
Average
41.7F
1895
40.9F
Mean March
Temperature
2012
40.2F
Average
30.1F
1895
27.8F
1895
0.97 in / m
November March
Precipitation
Average
0.72 in/ m
2012
0.65 in / m
1895 1905 1915 1925 1935 1945 1955 1965 1975 1985 1995 2005
Year
Fig. 3.9. Summary of historic climate data for Wyoming. The

trends suggest that, during the past two or three decades, the
frequency of unusually warm years and warm March temperatures has increased and the amount of winter prec ipitation
per month has generally declined. The shaded area indicates
1 standard deviation of the averages for 18952012. The
data come from thirty long-time weather stations distributed throughout the state, all with records starting in 1895,
and were obtained from the National Oceanographic and
Atmospheric Administration (http://www.ncdc.noaa.gov/
cag/). Adapted from Shuman (2012); supplemented with
recent data.
40
of unusually warm years and warm winters. This warm-
basins in Wyoming, there is no obvious trend in six of
ing has occurred throughout the state. Not surprisingly,
the basins, though a downward trend is suggested in
during this period some glaciers have disappeared and
four (see chapter 17). Changes in future rain and snow-
wildland fires and bark beetle epidemics have become
fall are likely to be variable across the state, but with
more widespread.23 Shuman confirmed that tempera-
warmer temperatures, evaporation will be greater. In a
ture trends were correlated with atmospheric green-
warmer climate, it is probable there will be less snow,
house gas concentrations. He could not find correlations
less irrigation water, less forage production, more acres
with solar irradiance, the Pacific Decadal Oscillation, El
burned in wildfires, and other related changes.
Nio/La Nia years, or changes in land use. Essentially
Although Earths climate has changed throughout
the same results have been found for Montana using
geologic time, the consensus of climate scientists is that
comparable data. 24
recent and ongoing climate change is occurring more
Could the temperature trends for Wyoming be
rapidly than before and that the best explanation for
part of a cycle when viewed over a longer period? To
these changes involves greenhouse gases produced by
address this question, Shuman examined changes in
human activities. 31 Also, even though most organisms
the 14,000-year-old pollen record preserved in lake
have adapted to previous episodes of climate change
sediments to infer temperature changes for Yellowstone
during the long history of life on Earth, through evolu-
National Park and the Bighorn Mountains (see chapter
tion or migrating to more favorable environments, biol-
2). These pollen records show the warming that caused
ogists wonder whether the current rate of change is too
glacial melting about 10,000 years ago, and also the cool
rapid for the evolution of many species to keep up. Local
and warm periods that have occurred since that time.
extinctions of species will accelerate if migration to
Shuman concluded that it is unlikely that warming dur-
areas having favorable climates is not possible. Almost
ing the past 40 or 50 years is part of a long cycle.25 He
everywhere, habitat has already been fragmented to the
also determined that recent temperatures in Yellow
point where conservation biologists are concerned that
stone National Park . . . have exceeded those recon-
some species will not have a pathway along which they
structed for the past >6000 years and are now similar
can move when migration is necessary (see chapter 18).
to the highest temperatures of the Holocene record.26
Also, potential destinations may already be fully occu-
Though less pronounced, his analysis of data for the
pied. Such issues are pertinent even if humans are not
Bighorn Mountains gave the same results.
the cause of the current episode of climate change.
In Wyoming, as elsewhere, changes in rain and
Slowing the rate of climate change will require sig-
snowfall are more variable and uncertain than tem-
nificant and controversial changes in the way industry
perature change. Shuman did find a decline in the
is powered, the way homes and workplaces are heated
frequency of wet winters. Moreover, based on climate
and cooled, and the modes of transportation that are
data since 1895, he found there were no winters for the
used. Progress has been made on using wind, solar
8-year period from 1999 to 2007 that were wetter than
power, and biofuels, and on the capture and sequestra-
the 83-year average from prior to 1978. Only 12 winters
tion of carbon dioxide, but some observers are skeptical
in the past 57 years have been wetter.27 Another study
about such alternatives. Proponents are convinced that
reported that the snowpack in the Rocky Mountains has
using energy more efficiently, combined with reducing
been dropping more in the past 30 years than in the
dependency on fossil fuels, is feasible and provides vari-
past few centuries, a decline reflected in data on stream-
ous benefits. These include opportunities for the eco-
flow and reservoir levels.28 According to the National
nomic growth associated with new industries that could
Weather Service, 2012 was the third-driest year in
help supply worldwide markets.
118 years.29 Still, the results are mixed. The National
Current rates of climate change are likely to adversely
Climate Assessment program projected that, by 2090,
affect some aspects of the economies of western states.
precipitation in Wyoming would be about the same or
For agriculture, the challenges will include finding
possibly 10 percent higher.30 When long-term precipi-
additional freshwater for more irrigation at a time when
tation records are examined for the 10 major drainage
water is even less available than it is now. This chal-
lenge is pertinent to the Rocky Mountain region, where
long as the beetles have susceptible hosts of sufficient
so much freshwater is provided by mountain snowfall.
size and number. The extent of mountain forests will be
Most likely, there will be less ice and snow, on aver-
greatly reduced, with shifts in the dominant trees.34 A
age, than today, and summers will be longer and drier,
recent report on the effects of climate change in west-
because the snowpack will melt earlier in the yeara
ern states concluded that big sagebrush will become less
trend that has been under way for 30 years.32 More hay
common, and invasive plants, such as cheatgrass, knap-
for livestock will have to be imported if plant produc-
weed, and leafy spurge, will become more widespread.35
tion rates are down, making meat more expensive.
In this same report, U.S. Forest Service ecologist Megan
Municipalities and other industries will be implement-
Friggens and her associates concluded, By the end of
ing water conservation programs more frequently (see
the century, 55 percent of future landscapes in the West
chapter 18). As Steven Gray and Chamois Andersen
likely will have climates that are incompatible with the
concluded in their prognosis for the Rocky Mountain
vegetation types that now occur on those landscapes.36
region, The concern now is that climate change may

increase the impact of droughts, just as population
In sum, a 2007 analysis of climate trends by paleoecolo-
growth and other factors have greatly increased the vul-
gists John W. Williams and Stephen T. Jackson sug-
nerability of the West to any type of drought.33
gested that the characteristics of the emerging climate
In the case of forests and rangelands, fires are likely
in the Rocky Mountain region and elsewhere are novel,
to be larger and more severe than before because of more
having never occurred before. 37 How will western eco-
frequent extended droughts that increase flammability.
systems change? How can municipalities, agriculture,
Also, the spread of bark beetle epidemics throughout
and industries adapt? Such questions are considered in
western North America during the past decade has been
subsequent chapters. Understanding environments of
to a large degree attributed to warming (see chapters 11
the present is important, but anticipating the effects of
and 12). Most likely, such epidemics will continue as
climate change in the future now seems urgent.
41
Part Two Wetlands
Chapter 4
Riparian Landscapes
Streamside landscapes are highly valued real estate
ing banks, and new sediments are deposited as bars or
ecologically and economically. They account for a mere 2
on floodplains. Such developments lead to a continu-
or 3 percent of the land area, but an estimated 75 percent
ously varying riparian mosaic of woodlands, shrub-
of native animal species depend on them at some time
lands, meadows, marshes, sand bars, and mudflats.
during the year for food, water, and shelter. Animals also
Physical factors that determine the nature of stream-
move along the riparian corridors in both directions,
side vegetation include elevation, climate, the steep-
often passing through otherwise hostile environments
ness of the terrain, soil type, channel sinuosity, and
(fig. 4.1). In addition, riparian zones filter sediments and
the width-to-depth ratio of the stream. Other factors
nutrients, thereby improving water quality, and during
are beaver dams, grazing by large mammals, and water
the spring they store water in alluvial sediments that sus-
diversion projects for agriculture, municipalities, and
tain streamflow late in the summer. Domesticated ani-
industry. Like the vegetation, riparian soils are highly
mals often congregate in riparian zones, as do people.
variable across a floodplain and are strongly influenced
Cities, towns, roads, and cultivated fields usually are
by hydrologic factors. Commonly, there are overlapping
located there, and, not surprisingly, most riparian zones
lenses of gravel, sand, and silt, indicating the locations
at low elevations are privately owned.
and velocity of former stream channels.
From the smallest rivulet at high elevations to the
Separating riparian habitats from other kinds of
largest river in the lowlands, the riparian landscape can
wetlandsnamely, marshes, fens, wet meadows, and
be viewed as a continuum. Creeks that begin with the
playas (see next chapter)is commonly done by con-
melting of alpine snow cross the full range of climatic
sidering water movement. Rivers and streams have
conditions, changing from incised streams flowing
free-flowing, oxygen-rich surface water most of the
over cobbles and gravel to meandering rivers underlain
year. Exceptions are ephemeral streams that usually
by sand or mud. The environment changes gradually
flow only in the spring and early summer. In contrast,
along the waterway, except at occasional beaver dams,
nonriparian wetlands are usually associated with still
waterfalls, and rapids, or at entrances to canyons. Some
or slow-moving water, and the bottom sediments are
streams are long, originating in the mountains; others
more likely to be anaerobic for a longer time during the
are short, such as those that begin as seeps or springs
year. In general, all wetlands have soils that are wet long
in the lowlands. Creeks and rivers with headwaters at
enough during the year to enable the growth of plants
high elevations have peak flows in the spring, at the
that require large amounts of water. Such species are
time of snowmelt. New channels may form during
known as hydrophytes and are used to define the nature
floods. Also, old trees topple into the water along erod-
and extent of wetlands for legal purposes. 2
45
46
Wetlands
1
2
3
4
5
6
Missouri River Basin

7 Little Missouri
Yellowstone
8 Belle Fourche
Clarks Fork
Big Horn
9 Cheyenne
Little Big Horn 10 Niobrara
Tongue
11 North Platte
Powder
12 South Platte
Great Divide Basin

13 Great Divide
Great Basin
16 Bear
Colorado River Basin Snake River Basin

14 Little Snake
17 Snake
15 Green
Fig. 4.1. A network of creeks and rivers connects the landscapes of the region. Many smaller creeks are ephemeral,
lacking water during the driest part of the summer. The locations of Yellowstone Lake and the larger reservoirs are shown.
Drainages 1417 lie west of the continental divide. Drainage
13 is the Great Divide Basin (see fig. 1.2), which is formed by

a split in the continental divide and has no outlet. About 72
percent of Wyoming contributes to the Missouri River drainage. Drawing by Linda Marston.
From Rivulet to River
aspen, and balsam poplar). Associated shrubs include
As rivulets merge, a first-order stream develops with

the adjacent vegetation dominated by several species
of sedges, grasses, and willows (table 4.1). Meadows and
short willow shrublands (see fig. 2.5) become more conspicuous as two first-order streams come together, forming a second-order stream, and as two second-order
streams merge to form a third-order stream. Lower on
the mountain, such shrubs as thinleaf alder, water birch,
and tall willows become more conspicuous (figs. 4.2 and
4.3). In the foothills, trees are more common, primarily narrowleaf cottonwood, but also Engelmann spruce
and blue spruce3 (and in some places, lodgepole pine,
red osier dogwood, silver buffaloberry, thinleaf alder,

water birch, and several species of willowoften in
dense thickets (table 4.1). At still lower elevations, such
as in the Green River, Wind River, and Bighorn basins,
and on the Great Plains, plains cottonwood becomes
the most conspicuous tree (fig. 4.4).
Several studies have examined streamside vegetation
in the mountains.4 Collectively, the results suggest that
the patchy riparian mosaic is affected by various environmental factors, primarily duration of soil saturation,
soil depth and texture, frequency of flooding, depth to
the water table, oxygen availability for roots, duration
of snow cover, growing season length and temperature,
Table 4.1 Some characteristic plants of riparian plant communities in Wyominga
Common
name
Latin name
Broadleaf
woodland
Coniferbroadleaf
woodland
AlderTall
Short
conifer
willow
willow
woodland shrubland shrubland
Cinquefoil
silver
Greasesagebrush
wood
shrubland shrubland
Riparian
meadow
TREES
Engelmann
spruce
Picea
engelmannii
Blue spruce
Picea pungens
Lodgepole pine
Pinus contorta
var. latifolia
Boxelder
Acer negundo
Green ash
Fraxinus
pennsylvanica
Narrowleaf
cottonwood
Populus
angustifolia
Plains
cottonwood
Populus
deltoides
Peachleaf
willow
Salix
amygdaloides
Russian olive
Elaeagnus
angustifolia
Basin big
sagebrush
Artemisia
tridentata ssp.
tridentata
Silver
sagebrush
(mountain)
Artemisia cana
ssp. viscidula
Silver sagebrush (plains)
Artemisia cana
ssp. cana
Common
snowberry
Symphoricarpos
albus
Greasewood
Sarcobatus
vermiculatus
Rubber
rabbitbrush
Ericameria
nauseosa
Saltcedar
Tamarix
chinensis
Silverberry
Elaeagnus
commutata
Skunkbush
sumac
Rhus trilobata
SHRUBS
(continued)
Table 4.1 (continued)
Common
name
Latin name
AlderTall
Short
conifer
willow
willow
Cinquefoil
silver
Greasesagebrush
wood
shrubland shrubland
Broadleaf
woodland
Coniferbroadleaf
woodland
Riparian
meadow
Water birch
Betula
occidentalis
Woods rose
Rosa woodsii
Red osier
dogwood
Cornus sericea
Shrubby
cinquefoil
Dasiphora
fruticosa ssp.
floribunda
Silver
buffaloberry
Shepherdia
argentea
Thinleaf alder
Alnus incana
Booths willow
Salix boothii
Diamondleaf
willow
Salix planifolia
Drummonds
willow
Salix
drummondiana
Geyers willow
Salix geyeriana
Grayleaf
willow
Salix glauca
Narrowleaf
willow
Salix exigua
Park willow
Salix monticola
Shortfruit
willow
Salix
brachycarpa
Wolfs willow
Salix wolfii
Alkali
cordgrass
Spartina gracilis
Alkali sacaton
Sporobolus
airoides
Basin wildrye
Leymus cinereus
Bluejoint
reedgrass
Calamagrostis
canadensis
Foxtail barley
Hordeum
jubatum
Meadow barley
Hordeum
brachyantherum
GRASSES
Latin name
Broadleaf
woodland
Kentucky
bluegrass
Poa pratensis
Nuttalls
alkaligrass
Puccinellia
nuttalliana
Orchard grass
Dactylis
glomerata
Saltgrass
Distichlis
spicata
Slender
wheatgrass
Elymus
trachycaulus
Western
wheatgrass
Pascopyrum
smithii
Smooth brome
Bromus inermis
Timothy
Phleum pratense
Tufted
hairgrass
Deschampsia
caespitosa
Nebraska sedge
Carex
nebrascensis
Northwest
Territory
(beaked) sedge
Carex utriculata
Water sedge
Carex aquatilis
Horsetail
Equisetum spp.
Mountain
bluebell
Mertensia
ciliata
Rocky
Mountain iris
Iris
missouriensis
Common
name
AlderTall
Short
conifer
willow
willow
Cinquefoil
silver
Greasesagebrush
wood
shrubland shrubland
Coniferbroadleaf
woodland
Riparian
meadow
SEDGES
FORBS
A dash indicates that the species is absent or uncommon.
Fig. 4.2. (above) Mid-elevation riparian shrublands on Jack

Creek in the Sierra Madre. Common shrubs and trees include
Geyers willow, Booths willow, diamondleaf willow, alder,
shrubby cinquefoil, Engelmann spruce, subalpine fir, and
lodgepole pine. Herbaceous plants include elephanthead
lousewort, red and yellow Indian paintbrush, and various wetland sedges. The upland forests are dominated by lodgepole
pine and aspen at lower elevations and Engelmann spruce
and subalpine fir at higher elevations. Mountain big sagebrush, Idaho fescue, lupine, and desert Indian paintbrush are
common on the drier, south-facing slope in the foreground.
Elevation 8,500 feet. See also fig. 2.5.
Fig. 4.3. (left) Bebb and Geyers willow occur along creeks
in the foothills, forming a riparian greenbelt in otherwise
semi-arid landscapes, such as here along Little Sage Creek on
the north end of the Sierra Madre. Riparian zones often are
subject to heavy grazing and browsing by large herbivores,
causing some shrubs to have a pruned appearance. Shrublands dominated by Wyoming big sagebrush occur on the
upland. Elevation 7,500 feet.
Riparian Landscapes
Fig. 4.4. Rivers and creeks with
dependable streamflow often have
woodlands dominated by two species
of cottonwood. Narrowleaf cottonwood
grows in the higher basins, such as here
along the Green River; plains cottonwood is more common at lower elevations on the eastern plains and in the
Bighorn Basin. Wyoming big sagebrush
is taller and more dense in ravines
where snow accumulates, as illustrated
in this photo. Elevation 6,200 feet.
big game browsing, and ice damage in the spring. All are
states, the material in the terraces was commonly
affected by features of the terrain, such as valley width
deposited during floods at the time mountain glaciers
and orientation, size of the drainage basin, and stream
were melting, more than 10,000 years ago. At that time
gradient and sinuosity. Abrupt changes in riparian veg-
the volume of water was much greater, and it was laden
etation are typically associated with different riparian
with sediments that were deposited as glacial outwash.
terraces, and islands (fig. 4.5). Beaver dams, debris jams,

and roads can also have dramatic effects. Often, within
High
landforms, namely, point bars, channel bars, cut banks,

Incised
channel
short distances, the channel can change from braided to

The flatter terrain of some mountain meadows and
the lowlands causes streams to flow more slowly, leading to the development of alluvial soils over sometimes
broad floodplains. Channel meandering is typical at
lower elevations, except where rivers have cut canyons or where steeper gradients lead to the formation
of braided streams (see fig. 4.5). Freshly deposited sand
and gravel form bars, which provide excellent sites for
the establishment of some species of willow and, at
CHANNEL STABILITY
meandering or incised.
Cut
bank
Floodplain
Bar
Point bar
lower elevations, cottonwoods and willows. Across from

the point bars are eroding cut banks of the floodplain,
Channel
boundary
the water (fig. 4.6). Cut bank erosion on meandering rivers produces some of the sediments that form new bars
downstream. This natural erosion, along with flooding
and the occasional formation of oxbows, creates a riparian landscape that is constantly changing.
Above the floodplain are terraces, former floodplains that now are well above the river. In western
Low
where older trees and other plants eventually fall into
Braided
Meandering
CHANNEL TYPE
Straight
Fig. 4.5. Braided, meandering, and straight channels have

different levels of resistance to changes in channel location,
providing different environments for riparian plants. Bars are
formed by the deposition of gravel, sand, and silt.
51
52
Wetlands
solved from calcareous rocks and deposited on the former floodplain. Now, with periodic wetting, the salts are
slowly moved upward when the soils are wet and then
deposited on the surface as the water evaporates. Consequently, halophytes (salt-tolerant plants) are sometimes
found on terraces. Less salt-tolerant species occur on the
adjacent floodplain, where floodwaters periodically dissolve the salts and transport them downstream.
The most successful cottonwood regeneration occurs
along sand and gravel bars after major floods that de
posit floating seeds high on the bank, where the seedlings that develop are not likely to be washed away by
subsequent floods.6 If groundwater is close enough to
the surface to provide the water needed by the saplings,
a curvilinear band of trees develops that provides a living record of flooding patterns and channel migration
(figs. 4.74.9). As the channel moves across the flood-
Fig. 4.6.The Snake River in Grand Teton National Park, with

young trees and shrubs on point bars (on the right) and a cut
bank (on the left) with some trees about to fall into the river.
Douglas-fir is the dominant tree on the escarpment in the
foreground, and mountain big sagebrush steppe occurs on
the terracea former glacial outwash plainjust beyond the
river. Lodgepole pine forests occur on glacial moraines in the
background. See fig. 15.3 for a photograph taken approximately 90 to the left. Elevation 6,700 feet.
Now, with the glaciers gone and with different climatic

conditions, the rivers are smaller. They also have less
of a sediment load and, consequently, are more likely
to incise streambeds. Along some rivers there are two
or three pronounced terraces, each the result of a different period of glaciation and downcutting (see fig.
15.3).
Such substrate characteristics as texture, salinity, and
depth to water table are not uniform, causing patchiness in the vegetation. In Jackson Hole, for example,
mid-channel bars of sand and gravel are often occupied
by sandbar willow, fine-textured soils on the floodplain
have Booths willow, and the terraces regularly have
Geyers willow.5 Terraces at low elevations are more
likely to become saline because of salts that were dis-
Fig. 4.7. Band of plains cottonwood seedlings along the Bighorn River east of Lovell. Most of the seedlings die, but a few
survive if they are high enough on the bank to avoid being
washed away by subsequent floods and if groundwater remains
accessible. A curvilinear band of trees often develops, such as
those apparent in fig. 4.9. See Cooper et al. (1999) and Scott
et al. (1997) for estimates of flood size required for successful
cottonwood seedling establishment. Elevation 3,700 feet.
Riparian Landscapes
Fig. 4.8. Aerial view of riparian
woodlands along the Bighorn River
east of Lovell. Former channel locations and bands of trees are visible.
Most of the area shown is dominated
by plains cottonwood. The tree-ring
record indicates that, historically, a new
band of trees became established every
1020 years. The older bands tend to be
farther from the river (see figs. 4.9 and
4.10). Desert shrublands are found on
the adjacent uplands. The red in this
infrared image indicates an abundance
of chlorophyll in the riparian zone
where water is available longer into the
summer, in contrast to the adjacent
desert shrublands. Similar patterns are
found along other meandering rivers,
for example, the Powder River south of
Arvada. Image from the U.S. Department of Agriculture.
plain, the older trees die if their roots cannot keep up
a mile or more in width, and the immediate banks of
with the deepening water table.7 Those that survive are
the stream for 300 or 400 yards are covered with a thick
likely to fall into the river when the channel is again at
growth of cottonwood. Similarly, D. B. Sacket wrote in
their baseassuming beavers do not cut them before
1877: All along the Bighorn River [north of Thermopo-
that happens.
lis] . . . much fine, large cottonwood timber grows.9 In
The riparian zone at lower elevations is often wooded,

dominated by plains cottonwood. Other trees include
both places, plains cottonwood would have been the

dominant species.
peachleaf willow, ash, boxelder, lanceleaf cottonwood,
Notably, portions of some lowland rivers were histor-
and, rarely, American elm.8 Along the Wind River near
ically devoid of trees (fig. 4.10), such as along the Sweet-
Riverton, W. F. Raynolds wrote in 1860: The valley is
water and Laramie rivers. While camped along Crow
53
54
Wetlands
Fig. 4.9.This row of aging plains
cottonwood trees is not likely to be
replaced as the trees die. The riparian
meadow in this area, along Walker
Creek in Converse County, has Baltic
rush and various sedges. Cattails grow
in standing water, and introduced
plants (such as timothy and smooth
brome) are found nearby. Photo by Ken
Driese.
Creek, near Cheyenne, P. G. Cooke wrote in 1845, for
viding water late in the summer. However, if ground
forty miles we have seen but one treefive miles off
water drops to a depth where the large trees cannot
and not a bush or shrub.10 The near absence of trees in
obtain the water they need, woodlands give way to
some areas could have been caused by various factors,
meadows and shrublands. Extended droughts can lead
including frequent prairie fires, large mammal brows-
to the death of cottonwoods, just as shifting channels
ing, insufficient flooding for tree seedlings to become
and altered irrigation practices can.
established on point bars, or stable channels dominated
The riparian mosaic at the confluence of the Big-
by grasses and sedges that reduced the chance of tree
horn and Shoshone rivers in northern Wyoming illus-
seedling establishment (as discussed below). It seems
trates landscape patterns that can be found elsewhere
unlikely that the lack of seed dispersal is the explana-
(fig.4.11; see also fig. 4.8). Fertile soils and water avail-
tion, as many riparian species, such as cottonwoods,
ability for irrigation, combined with a relatively long
have seeds that are both buoyant and easily spread by
growing season, sustain one of the most important agri-
water and wind.
cultural regions in the state. Much of the native ripar-
Riparian meadows on the eastern plains, with no or
ian vegetation has been lost, but remnants persist in
only a few trees and shrubs, are commonly dominated
the Bighorn Canyon National Recreation Area. Plains
by Nebraska sedge, Baltic rush, prairie cordgrass, redtop
cottonwood woodland is the most conspicuous vegeta-
bentgrass, and plains silver sagebrush, plus introduced
tion type along the rivers. Associated species include
plants, such as Kentucky bluegrass, smooth brome,
peachleaf willow, silver buffaloberry, and the intro-
sweetclover, Canada thistle, hounds tongue, and tim
duced Russian olive. Various shrubs, forbs, and grasses
othy. In areas where salts accumulate in the soil, such as
are found as well (see table 4.1). Shrub and herbaceous
on terraces that are flooded less frequently, greasewood,
plant cover is low where tree density is high because
inland saltgrass, and alkali sacaton are common. Flood-
of inadequate light beneath the woodland canopy and
plain marshes with standing water commonly have
competition from trees for the available water.
dense growths of cattail (see chapter 5).

Trees have invaded some meadows and shrublands,
Interspersed in the woodlands at low elevations are

shrublands dominated by basin big sagebrush, plains
possibly because dams and reservoirs reduced the fre-
silver
quency of tree-damaging floods.11 Also, return flows
sandbar willow, skunkbush sumac, western snowberry,
from irrigated land may have favored the trees by pro-
Woods rose, chokecherry, silver buffaloberry, saltcedar,
sagebrush,
greasewood,
rubber
rabbitbrush,
Riparian Landscapes
Fig. 4.10.The Sweetwater River, looking northeast from Independence Rock in 2013. Note the lack of trees and shrubs, just
as in 1870, when a similar photo was taken by William Henry
Jackson (U.S. Geological Survery photo284; see K. L. Johnson
1987). Common plants include alkali sacaton, fringed sagewort,

American licorice, plains silver sagebrush, and rubber rabbitbrush. Note the elongated, well-vegetated point bar. Elevation of
the river is approximately 5,800 feet.
and a variety of forbs and grasses. Abrupt transitions
ian zones grow rapidly during spring and early sum-
from woodland to shrubland, together with abundant
mer, providing an abundance of highly flammable fuel
evidence of fire-scarred trees and logs, suggest that
by late summer. Shrublands dominated by skunkbush
fire has played an important role in converting older
sumac often replace the woodlands that are burned.
woodlands to shrublands. Cattail marshes occur in
Another important factor causing rapid change is
flooded oxbows, and meadows are found nearby, such as
flood control. Along the Bighorn River, Akashi found
in recently abandoned channels.
evidence of much less channel shifting after the

enlargement of Boysen Reservoir in 1952. Consequently,
Shifting Riparian Mosaics
fewer and smaller point bars were formed, resulting

in fewer sites for the establishment of cottonwood
The mosaic of some riparian zones is relatively stable,
seedlings.13 At the same time, trees continued to die from
because the creek or river is at the bottom of a V-shaped
old age, fire, beaver cutting, agricultural clearing, and
valley incised in bedrock. In contrast, riparian mosaics
increasing depth to the water table. Thus, the land area
on the alluvial soils of broad floodplains change rapidly
in old-growth cottonwood woodlands declined, a trend
(see figs. 4.8 and 4.11). Comparing a series of aerial
observed in other areas as well.14 Just as fire suppression
photographs taken over the Bighorn River, University
has led to changes in some mountain forests, flood
of Wyoming graduate student Yoshiko Akashi found
suppression has changed the riparian zone.15
that some of the woodlands, shrublands, and meadows
Notably, the shifting mosaic along the meander-
changed locations during a 50-year period.12 Fire was
ing Bighorn River is different from that on the nearby
a major factor causing the change. Plants in the ripar-
Shoshone River, a more rapid, braided river. Cotton-
55
Wetlands
Bar deposit
Sandbar meadow
Without
willow or
cottonwood
Cottonwood woodlands
Young (1-29 yrs)
Meadow
Basin big
sagebrush
Suspended sediment
Middle-aged (30-79 yrs)

Old (>80 yrs)
Fire
or
beaver
Fire
or
beaver
Saltcedar
willow
Annual
flooding
Saltcedar
Skunkbush sumac
Rabbitbrush
shrubland
Periodic
flooding
or fire
Bank erosion
Increasing distance from channel or height above river level
56
Periodic
flooding
or fire
Skunkbush sumac
Rabbitbrush
Snowberry
Rose shrubland
Prolonged inundation
?
?
Greasewood
shrubland
Fig. 4.11. Probable riparian succession on new point bars

along the meandering channel of the Bighorn River east of
Lovell. As indicated, succession has changed considerably
in this area since the introduction of saltcedar (also known
as tamarisk). With flood control, many point bars are not
suitable for the establishment of cottonwood seedlings. Therefore, the proportion of the landscape dominated by cotton
wood decreases as older trees die; shrublands have become
more common. Elevation 3,700 feet. Adapted from Akashi
(1988).
wood seedlings are observed more frequently along the
cottonwood in western Wyoming and eastern Idaho,
Shoshone, perhaps because favorable flooding is more
sandbars by sandbar willow, and mudflats by Booths
frequent there. By whatever means, habitat for seedling
willow.17 Cottonwood often invades stream margins,
establishment is created more often. Similarly, young
but as they age and the channel moves farther away,
narrowleaf cottonwoods are common along the free-
the trees die and give way to shrubland or grassland.
flowing Gros Ventre River in Jackson Hole.16
As noted, tree mortality in such places could be caused
The riparian landscapes along meandering and
by a dropping water table as the channel moves across
braided streams change so rapidly that it is difficult to
the floodplain, which enables the invasion of plants
think about succession in the traditional sense. Before
that are more tolerant of water stress and also possibly
the biota has a chance to form a relatively stable com-
benefit from more oxygen in the soil, such as red osier
munity, the river channel typically shifts to a new loca-
dogwood, Rocky Mountain juniper, silver buffaloberry,
tion or a fire occurs (because of the abundant fuel).
skunkbush sumac, and rubber rabbitbrush. The sup-
Still, several successional trends have been observed.
pression of disturbances along some rivers allows the
Commonly, cottonwoods and some willows are the pio-
development of self-perpetuating shrublands, and also
neer species, with gravel bars colonized by narrowleaf
woodlands with trees other than cottonwood.18 Along
Riparian Landscapes
the Snake River south of Jackson, blue spruce and red
Many people living along creeks and small rivers ben-
osier dogwood have formed a relatively stable commu-
efit from soil laid down by generations of beaver work-
nity. In eastern Wyoming, ash and boxelder form simi-
ing for thousands of years. Some have suggested that the
lar woodlands.
beaver is a keystone species, affecting ecosystem structure

and function far beyond what might be expected, con-
Bank Storage, Stream Hydrology, and Beaver
sidering animal size and population numbers. Others

have referred to beaver as bioengineers because of the
The plant growth on floodplains slows the movement of
structures and services they provide. To illustrate, sedi-
floodwater and thereby increases the potential for sedi-
mentation behind their dams provides clearer water
ment deposition when floods do occur. The accumu-
downstream, and greatly enhanced bank storage leads
lated sediments increase the amount of water stored in
to sustained streamflow throughout the year (figs. 4.12
the bank and floodplain, which, along with other fac-
and 4.13). A study in Colorado found that a valley with
tors, increases the probability of streamflow through-
an active beaver colony had eighteen times more water
out the year. However, maintaining bank storage has
storage in the spring and higher streamflow in late sum-
its costs in the form of water consumption by phreato-
mer than a comparable drainage where the beavers had
phytestrees and shrubs that live with their roots in
been removed.19 Higher late-summer streamflow bene-
groundwater much of the year. Every pound of plant
fits fish and wildlife as well as landowners. Beavers also
material produced may require as much as 400 pounds
thin woodlands by cutting trees, thereby stimulating
of water, because up to 99 percent of the water entering
the formation of new sprouts and the growth of under-
a plant through the root system is lost to the atmosphere
story plants. When the beavers are removed, the habitat
by transpiration. This water is required to keep stomata
can change quite rapidly. 20
open for the uptake of carbon dioxide and to maintain
Somewhat similar to beaver dams are the debris jams
turgid leaves for better light interception. But water
that commonly occur along wooded creeks (fig. 4.14).21
transpired by plants is not available for downstream
Debris jams are initiated most often by a tree that falls
uses. Such losses, along with seepage to groundwater
across a stream, or by floating wood and other materi-
aquifers and evaporation directly from water surfaces,
als that become lodged in the channel, creating a dam.
are known as conveyance losses. Water consumed by
Both kinds of dams improve water quality and benefit
riparian vegetation is sometimes viewed as unfortunate
some wildlife.22 However, a common reaction has been
in semi-arid environments, but it can also be viewed as
to remove obstructions to streamflow. The dams some-
a small price to pay for the benefits of increased forage
times appear messy, and they can inundate land that
and shade for livestock, erosion control, sedimentation
owners prefer not be flooded. Indeed, beavers become
during floods, biological diversity, better wildlife habi-
a nuisance when they build dams at road culverts or
tat, and sustained late-summer flowsall examples of
when they interfere with irrigation projects.
ecosystem services.
Beavers and their dams are much less common now
Few data are available to estimate quantitatively the
than in the early 1800s. Alluvial sediments originally
amount of water used by riparian vegetation, but two
deposited behind beaver dams have washed down-
generalities seem probable: first, streamflow depletion
stream in many areas. With fewer dams to dissipate the
by phreatophyte transpiration (that is, the proportion
energy of spring floodwaters, new gullies are created in
of streamflow used by plants) is higher in small streams
a year or two, sometimes leaving fence posts hanging
than in rivers; and second, depletion is higher during
from barbed wire in midair. As the channel deepens, the
late summer, when streamflow is low and transpiration
water table is lowered, and the soils on either side of the
is more rapid. Preliminary estimates suggest that reduc-
gully become dry earlier in the summer. The riparian
tions in streamflow by phreatophyte transpiration are
vegetation changes from willows and moist meadows,
less than 2 percent in June when streamflows are high,
which are relatively rare in the landscape and valuable
and up to about 50 percent in late August when the vol-
for livestock and wildlife, to the drier, already wide-
ume of flow is low.
spread vegetation of the adjacent upland. The riparian
57
Fig. 4.12.Centuries of beaver

activity have created a relatively flat riparian ecosystem
in what once might have
been a steep-sloping valley.
Common plants in this
area are water sedge; beaked
sedge; resin birch; and several
species of willows, including
diamondleaf, Wolfs, Geyers,
and Booths. Photo taken
along the upper reaches of
Lake Creek in the Medicine
Bow Mountains, southwest
of Dry Park. Lodgepole pine
dominates the forested upland, though many trees have
succumbed to the mountain
pine beetle. Elevation 9,100
feet. Persico and Meyer (2009)
discuss the variable effects of
beavers and climate on riparian landforms.
Fig. 4.13. Prior to the construction of this new beaver dam,

a deep gully existed along Muddy Creek near the town of
Baggs. The alluvial soils might have been deposited originally
behind previous beaver dams, which could have failed after
the beaver population was reduced by trapping in the 1800s.
It is possible that the current beaver population will continue

building dams higher and higher, eventually filling the gully.
Willow shrublands and meadows could become re-established
where there are now shrublands with greasewood and big
sagebrush. Elevation 6,000 feet.
Riparian Landscapes
Fig. 4.14.Trees falling across streams create debris jams

that provide habitat for fish and other organisms. Here,
Mullen Creek in the Medicine Bow Mountains is flanked by
Engelmann spruce, subalpine fir, and thinleaf alder. Elevation 8,500 feet.
zone narrows, and gully erosion continues as long as
be more successful if other construction materials were
the flows are rapid and unobstructed. Such flows pro-
provided. He cut several aspen trees in the distant foot-
vide less opportunity for the benefits of bank storage.
hills and transported them to near the transplanted bea-
Ironically, erosion along lowland streams is some-
ver. Several days later a dam was being constructed, and
times of sufficient concern that landowners resort to
within 2 years much of the gully behind the dam was
constructing dams themselves, which is far more costly
filled. With the rising water level, sagebrush and grease-
than the problems caused by beaver. As one Wyoming
wood became less common, while meadow grasses and
rancher noted, Beavers can be a pain in the neck when
willows became dominant. This experiment involved
you get too many in the wrong place. On the other
working with beavers and has been repeated success-
hand, how are we going to get that erosion stopped if
fully in several locations. Initially it was natural to look
we dont use beaver? They work pretty cheap, and Ive
for aspen to assist the beavers, but riparian ecologists
never seen a lazy one yet.23
have learned that even old tires wired together can
The role of beavers in preventing erosion and creat-
serve as a stabilizing framework around which beavers
ing habitat has long been recognized. It was therefore
can build new dams. Once the tires are in place, the
natural for land managers to consider transplanting
beavers finish and maintain the project using readily
beaver to severely eroded gullies. In one case, in south-
available shrubs.25
western Wyoming, the transplanted beavers had diffi-
Significantly, gullies are not a new landscape feature
culty building and maintaining dams made from the
created by human activities. As noted in chapter 2, gul-
plant material that was available, namely, greasewood
lies are frequently mentioned in journals written in the
and sagebrush. 24 A Bureau of Land Management biolo-
1800s. Some could have resulted from excessive bison
gist, Bruce Smith, wondered whether the beaver would
grazing or beaver trapping by the first fur traders, or
59
60
Wetlands
by erosion caused by wagon trains. Yet even the best
limbs and give them every night as the buffaloe [sic]
beaver construction sometimes fails, especially with
have entirely destroyed the grass throughout this part
exceptionally high spring floods. Beaver populations
of the country.28
could have been diminished for other reasons as well,
However, the explorers did not provide information
including disease and periods of drought. Whatever the
on the amount of time the bison spent in a particular
cause, the evidence suggests that, though gullies were
area. Without fences, bison would have wandered at
present prior to EuroAmericans, they are probably more
will, possibly grazing an area heavily but not returning
common now (see chapter 2).26
to the same place for a year or more. Also, the upland forage then, away from the rivers, might have been attrac-
Livestock, Reservoirs, and Irrigation

Riparian habitats have long been important to people
living in the semi-arid West. Beaver trapping and placer
mining for gold took place along creeks and rivers, and
abundant forage and irrigation water facilitated agricultural development. Most farms, towns, and cities
are located nearby, partly because of the availability
of water but also because of the aesthetic appeal of
riparian zones. Floods continued to occur, often causing great damage to riverside settlements, but usually
the accepted solution for the flood problem was dam
construction or channelization. Floodplain amenities kept people from moving to higher ground. Dams
not only prevented spring floods, they also provided
opportunities for waterpower, irrigated agriculture,
and recreation.
Today, more people than ever want access to the
resources provided by riparian landscapes. The result
is high land prices and increased concern about riparian management. Three topics are especially relevant
in this regard: livestock grazing, reservoir construction
and management, and the effects of different irrigation
systems.
Livestock
tive to bison more than to livestock. Today water tanks

and salt blocks are used to draw cattle to the upland, at
least for a portion of the year. But in some areas it seems
as though the only solution to conserving the riparian
habitat and streamwater quality is additional fencing or
labor-intensive herding.
One of the undesirable consequences of heavy grazing along creeks and rivers is bank erosion, which leads
to a decline in bank storage, water quality, and habitat.29 Such erosion can occur at any elevation, reversing a natural tendency, namely, sedimentation near the
banks where the flow is slower and a gradual narrowing
of the channel as sediments are stabilized by plants. The
deeper, narrower streams often have overhanging banks
that provide excellent fish cover (fig. 4.15). Hooved
animals, however, whether livestock or big game, can
destroy the banks, thereby reducing water storage capacity, widening stream channels, and increasing water
temperatures. 30 Consequently, fish habitat is degraded.
Several studies have documented these trends and the
fact that better livestock control improves fish production in small streams.31
Poor livestock management is not the only potential
cause of stream degradation. Bank erosion often occurs
with road construction and other developments, and
nutrients can enter the water by seepage from feedlots,
fertilized cropland, and inadequate sewage treatment
Domestic livestock concentrate where water, food,
systems. Such factors may be as serious for lakes and
and shade are in close proximity, sometimes leading
reservoirs as for the streams. A case in point is Flam-
to excessive grazing. Large numbers of bison could
ing Gorge Reservoir near the Wyoming-Utah border,
have had the same effect on streams as cattle or sheep.
where the upper part of the reservoir has algal blooms
Indeed, travelers in the early 1800s observed thou-
that can interfere with recreational activitiespossibly
sands of bison grazing along rivers.27 One of the earliest
because of nutrient additions to the Green River that
explorers, Osborne Russell, wrote about southern Mon-
cause undesirable changes to the aquatic ecosystem.
tana in 1835: The bottoms along these rivers are heav-
In the Greater Yellowstone Area, various investiga-
ily timbered with sweet cottonwood and our horses and
tors have concluded that willow and cottonwood were
mules are very fond of the bark which we strip from the
once more common when concentrations of elk on win
Riparian Landscapes
tion and other uses later in the summer. 34 However,

about 85 percent of the streamflow in Wyoming still
flows into neighboring states. 35 Further reservoir construction has been stalled by economic constraints,
environ
mental concerns, and binding agreements
with downstream states. The most direct negative
effects of reservoir construction are the loss through
inundation of a free-flowing stream and its associated
riparian habitat. Sacrifices are always necessary and
must be weighed against the obvious benefits.
One reason reservoirs are controversial is that their
utility can be short-lived because of sedimentation,
which continues regardless of location. Sedimentation frequently is accelerated by human activities that
increase erosion. Dealing with large, sediment-laden
reservoirs will be a problem for future generations. Beaver also cause sedimentation, but their activities are
restricted to small areas; the breaking of a beaver dam
is not a big issue.
The various purposes of reservoirs include flood control, irrigation, power generation, and recreation. Typically, water levels fluctuate considerablyhigh at the
beginning of the growing season to maximize the water
available for agriculture later in the summer, but low by
the end of the growing season because of withdrawals
Fig. 4.15.Excessive grazing and trampling by large animals
can convert cool, narrow creeks with overhanging banks
to warm, shallow creeks that are less suitable for fish. Also,
the sediments (black) that contribute to bank storage and
late summer flows are lost. Streams in poor condition can be
restored.
for irrigation. Reservoirs with hydroelectric plants, such

as those on the North Platte River, may have water
level drawdowns at times of high demand for electricity. When water levels are down, nearly barren shores
are created that sometimes become saline mudflats and
sources of dust clouds. Most native plants cannot tol-
ter ranges were much lower, due to predation by wolves

and American Indians (see chapter 15).32 Browse lines
on willows, juniper, and aspen are now commonly
observed throughout the region, sometimes the result
of browsing by native ungulates as well as livestock.33
Whether such browse lines are an indicator of excess
grazing is debatable (see fig. 10.16).
Reservoir Construction and Management
erate the water-level fluctuations that occur, although

such sites can become dominated by weedy species capable of rapid growth, such as dock, foxtail barley, goosefoot, knotweed, saltcedar, sowthistle, summercypress,
sumpweed, and sweetclover. In general, sedimentation,
water-level fluctuations, and the streamflow changes
associated with reservoirs inevitably disrupt the riparian ecosystem. The seriousness of these changes must
be judged after considering what is lost, what is gained,
and how long the benefits are likely to last.
Because of the states high elevation, Wyoming is part
Flood control levees also disrupt riparian land-
of an area known as the headwaters of the West. Snow-
scapes. The most prominent example in Wyoming is
fed streams flow into all major drainages (see fig. 4.1).
in Jackson Hole, where in 1957 the U.S. Army Corps of
There are 116 reservoirs with storage of 1,000 acre-
Engineers and other entities began channelizing the
feet or more that retain some of this water for irriga-
Snake River to protect valuable private property from
61
62
Wetlands
flooding at the base of the Teton Range. 36 The levees
crops or retained in the soil. But trade-offs are involved.
now extend for a distance of about 20 miles. Between
First, some of the spring flood water is diverted into
the levees, the river is narrower and flows deeper and
reservoirs, reducing the magnitude of beneficial early
more rapidly than before, causing more erosion down-
season flows. Second, much of the irrigation water
stream. Spawning habitat has been greatly diminished;
is transpired or evaporated from croplands, thereby
and without periodic floods and with less groundwater,
reducing the amount of water returned to groundwater
the floodplain flora and fauna have changed. Although
or streams. In most western states, evaporation from
enabling residential and commercial developments on
reservoirs and cropland (evapotranspiration) account
the floodplain, the levees have reduced the values of
for more than 90 percent of total water consumption. 38
riparian ecosystem services. Attempts have been made
Losses through evapotranspiration constitute a high
to restore some spawning grounds in tributaries to the
proportion of the total irrigation water in drainages
Snake, but at considerable cost.
where the amount of water used for irrigation is low,
Like levees, the dikes built across riparian zones for
or where the amount of water required for percolation
highways and railroads modify patterns of stream-
back to the stream or groundwater is greater than the
flow, thereby altering the riparian ecosystem. Ponding
amount of water applied. In other words, if water added
of water usually occurs upstream, which can be ben-
to a meadow or crop is less than the soil profile can
eficial for waterfowl, but it also elevates the potential
retain against the pull of gravity, then all or most of the
for flooding if culverts and bridges are not well engi-
irrigated water will be lost to evapotranspiration and
neered. Downstream from a roadbed, the floodplain
very little, if any, will flow back to the stream. Many
can become stabilized, depending on land use practices
factors are involved, including climate, the hydraulic
in the area and distance to the next streamflow obstruc-
conductivity and water-holding capacity of the soil, and
tion. Of course, the most dramatic effects of levees and
crop water requirements.
dikes occur during periods of high water. Notably, dikes
Irrigation by flooding has disadvantages as well
increase the width of riparian zones in some areas, as
as the potential advantage of sustained streamflows
does irrigation; levees reduce the width.
in some watersheds. For example, nutrient leaching

occurs more frequently with flooding, which leads to
Irrigation and Streamflow
fertilizer losses and concomitant nutrient enrichment

(eutrophication) of streams and groundwater. Moreover,
By whatever means, and whether the water comes from
salts that have accumulated in the soil over long
reservoirs, pumps, or streamflow diversions, irrigation
periods are dissolved and transported to the stream
occurs when resources are available to construct a con-
or groundwater, sometimes degrading surface and
veyance system and the soils and growing season are
groundwater with salinity that is too high for livestock,
adequate for the crops desired. In Wyoming, the area
cropland, or human uses.
with potential for irrigated croplands is small, only
Many farmers now have sprinkler or drip irrigation
about 23 percent of the state. However, the effects are
systems that are more efficient than traditional
more far-reaching than the amount of land involved.
irrigation, because the water is spread more uniformly,
One effect is that some streams with formerly yearlong
and a smaller portion of the available irrigation water
flows are now dry late in the summer because of with-
is transported to a particular field. The same amount
drawals upstream, creating ephemeral streams that are
of water can thus be applied to a larger amount of land,
much less valuable for fish, wildlife, and livestock. In
thereby reducing costs. There also is less potential for
1986 the Wyoming state legislature recognized fishing
the loss of fertilizer by leaching, and consequently
and recreation as beneficial uses of streamwater.37
stream eutrophication occurs more slowly. Increased
Significantly, flood irrigation can lead to more
crop production and lower fertilizer requirements
streamflow lower on the watershed later in the summer.
help pay for the required equipment and pumping
This happens because of return flows, that is, water
costs. However, although using irrigation water more
applied by flood irrigation that was not used by the
efficiently has considerable appeal, there is a greater
Riparian Landscapes
potential for salts to accumulate on the irrigated soil
ther. Some rivers were clogged with logs. After gold was
surface. Eventually, this salinization may reduce crop
discovered, feverish digging began adjacent to stream
growth. One solution is to flood the soil periodically,
channels. About that time, herds of thirsty livestock
but this alternative simply washes the salts into other
also trampled the shorelines, here and there, possibly in
bodies of water. The losses of one ecosystem are the
a way that was not typical of bison herds.
gains of another. Also, sprinkler irrigation produces
Thus, many riparian zones were destroyed or clearly
much less return flow. Less water is applied to a field,
disrupted by the end of the nineteenth century, at a
leaving less water that percolates into wetlands or
time when numerous exotic plants had been introduced
back to the stream. Consequently, with more efficient
to western statesplants that were well adapted to sur-
irrigation, water levels in all kinds of wetlands can be
vive in the relatively moist, fertile riparian habitat.41
lower in the fall.
Some were invasive, that is, they gradually displaced
39
native species.42 A few were introduced intentionally,
Invasive Plants, Climate Change, and the Future
because it was thought they might be economically

or aesthetically valuable; others arrived on their own,
The very first large-scale disturbance in the region by
such as Canada thistle, leafy spurge, yellow sweetclover,
EuroAmericans, on any landscape, occurred in ripar-
Kentucky bluegrass, and smooth brome. In retrospect,
ian zones, namely, commercial beaver trapping in the
many land managers now recognize that introduced,
early 1800s (see chapter 2). Prior to that time, disease
invasive plantsother than cropscan bring more
may have caused fluctuations in the number of beavers,
problems than benefits. Moreover, controlling such
but intensive trapping greatly reduced their population
plants is costly, if even possible.
sizes throughout North America. Many beaver dams
Two noteworthy invasive plants in the region are
surely failed, and in some areas new gullies formed. A
Russian olive and saltcedar (tamarisk). Russian olive
few decades later, some of the same waterways were sub-
was widely planted for windbreaks on the semi-arid
jected to a second round of human-caused disturbances
uplands and as an ornamental tree in towns. However,
associated with the logging industry and gold mining.40
it has invaded many riparian zones to the point that
Stream banks were cleared in many places, and sluice
many people think of it as naturalized (fig. 4.16)the
dams were constructed to create flash floods that car-
next thing to a native. Unlike cottonwood, it does not
ried thousands of logs, disrupting the floodplain fur-
require periodic floods for the establishment of new
Fig. 4.16. Russian olive, with gray-green

leaves, is an introduced invasive tree in
riparian zones at low elevations, such as
here along the Bighorn River south of
Thermopolis. Elevation 4,345 feet.
63
64
Wetlands
seedlings, enabling it to reproduce more rapidly than
and also whether such nutrient enrichment favors
cottonwood on the many streams in western states
the invasion of other exotic plants. Unusually high
where streamflow is regulated and floods are suppressed.
proportions of the herbaceous plants in many riparian
The fruits are commonly eaten and dispersed by birds.
zones consist of introduced species.
Most reproduction is from seedling establishment,
The other invasive plant, saltcedar, was introduced
though root sprouts do occur. Invasion is hastened in
to the region in the 1930s (fig. 4.17; see also fig. 4.11).45
some areas because beaver preferentially cut and eat
It grows mostly at low elevations, such as in the Bighorn
cottonwood sprouts, thereby creating a more favorable
Basin; the Wind River Basin; the lower parts of the
environment for Russian olive. And once established,
Green River, Powder River, and North Platte basins; and
Russian olive better tolerates the aridity that develops
northward into Montana. It has also been found above
with drawdown of the water table. As is typical for
7,000 feet in the Laramie Basin. With warming of the
invasive plants, part of the exotic trees success may
climate, saltcedar could become more widespread, as it
be attributable to the frequent preference of most
tends be more drought tolerant than some native shrubs
herbivores for native species.43
and trees.46 Both saltcedar and Russian olive grow
The alluvial soils of riparian zones generally have

sufficient nitrogen for plant growth. Nevertheless,
together on floodplains where the groundwater level is

dropping and upland plants are invading.
Russian olive has a symbiotic association with a
The seedlings of saltcedar tolerate saline soils better
group of fungi known as Actinobacteria, which form
than some native riparian species do.47 This tolerance
actinorhizae (analogous to mycorrhizae) that are capable
can be attributed partially to salt-secreting glands on
of nitrogen fixation. This adaptation may facilitate the
the leaves. Salts released by the glands are deposited on
exotic trees survival on the uplands, even though it
the soil from time to time by rainwater and also when
may not be critical on the floodplains. Various studies
the leaves are shed. Consequently, established plants
have shown that the soil under Russian olives has
can elevate surface soil salinity through litterfall.48
44
higher nitrogen levels than under cottonwood trees.
Saltcedar often grows so densely and uniformly over
Some have wondered whether nitrogen enrichment of
large areas that few native species survive, and it does
the streamwater is higher because of the Russian olive,
not support wildlife as well as the native vegetation. Like
Fig. 4.17. Saltcedar is an introduced

invasive shrub in riparian landscapes
that is becoming more common at low
elevations in Wyoming and southern
Montana. Another invasive plant, hoary
cress, is visible in the foreground. Photo
by Brian Mealor.
Riparian Landscapes
cottonwood and willows, saltcedar is a phreatophyte.
and dikes have been constructed to prevent flooding.52
However, it generally transpires more water than the
Along with such developments, the ecosystem services
native species do, most likely because it often produces
provided by riparian ecosystems have been diminished,
stands with more square feet of leaf area per square foot
creating costs that often are not consideredcosts
of ground area (the leaf area index).49
related to water quality, sustained streamflow, and wild-
Controlling the spread of both Russian olive and
life habitat.53
saltcedar, and many other invasive herbaceous plants
And now an added concern is warming of the
in the riparian zone, has been difficult. Herbicides
climate. Higher temperatures could lead to continued
typically kill some native species as well as the
decline in cottonwood abundance if floodplains become
invasives, and the chemicals are not recommended for
drier, floods are insufficient to enable new seedling
use in close proximity to waterways. Cutting the shrubs
establishment, or fires occur more frequently. One
or trees followed by the application of herbicide to
effect of a warmer climate already has been observed:
the root crown has been effective, though costly, and
earlier snowmelt, which increases the probability of
there has been some success with biological control.50
drought-caused mortality of riparian plants during a
A fundamental approach is to manage the river so
longer, warmer summer.54 Some rivers and streams that
that cottonwood, boxelder, and other native trees are
formerly flowed all year may become ephemeral, causing
favored. As the native trees grow taller, the saltcedar dies
dramatic changes to the riparian flora and faunanot
because of insufficient light.51 Of course, this approach
to mention municipalities, industry, and interstate
may require allowing floods and reducing the beaver
compacts.55 With a warmer climate, frost-sensitive
population, especially if cottonwood establishment and
invasive
growth is desired.
expand their ranges, and Russian olive could become
plantssuch
as
saltcedarprobably
will
more common, because it tolerates drought better

There is one clear conclusion for riparian landscapes:
than cottonwood can.56 Water shortages will promote
streamflow regulation, agriculture, irrigation, livestock
controversial initiatives, such as cloud seeding and
grazing, invasive plants, fire suppression, and other
additional trans-basin diversions, and also will reduce
human activities have created riparian habitats that
the availability of water for ecologically beneficial
are quite different from those of the 1800s. To varying
instream flows or floods. Sweeping generalizations for
degrees, alterations are occurring in both the lowlands
all riparian ecosystems are not possible, but its clear
and mountains, especially where roads and summer
that benefiting from riparian resources in the future
homes have been constructed in valley bottoms, large
will require a coordinated effort by policymakers,
herds of livestock or big game congregate, land is culti-
riparian scientists, fisheries biologists, engineers, and
vated, dams have created reservoirs and regulated flows,
land managers.
65
Marshes, Playa
Wetlands, Wet Meadows,
and Fens
Chapter 5
Nonriparian wetlands are those commonly referred to
lost at alarming rates, the U.S. Fish and Wildlife Service
as marshes, playa wetlands, wet meadows, and fens.
was charged in 1974 with creating a National Wetlands
Such places, along with the riparian habitats described
Inventory.2 Based on that work, it is known that wet-
in the previous chapter, are recognized for the benefits
lands now cover some 950,000 acres (384,000 hectares)
they provide by filtering waterborne pollutants, regu-
in Wyoming1.5 percent of the state. Considering
lating water flows, storing carbon, and enriching the
this small area and their ecological importance, wet-
diversity of plants and animals.1 For these reasons, and
lands are appropriately thought of as keystone habitats
because wetlands nationwide were being degraded or
(figs.5.15.3).
Fig. 5.1. Swamp Lake wetland

northwest of Cody is a mosaic
of cattail marsh, sedge marsh,
and fen. The fen is the largest
known peatland in Wyoming
and is classified as nutrient
rich. It receives water and
nutrients from the limestone
and dolomite of the Cathedral
Cliffs in the background.
Elevation 6,600 feet.
66
Marshes, Playa Wetlands, Wet Meadows, and Fens 67
Fig. 5.2. (above) Marsh in Hutton Lake National Wildlife

Refuge in the Laramie Basin. Hardstem bulrush is the most
common plant in this area and becomes abundant when
water levels are favorable. This marsh is dependent on water
from creeks and irrigation canals. Elevation 7,150 feet. Photo
by Rhonda Foley/U.S. Fish and Wildlife Service.
Fig. 5.3. (left) The yellow-headed blackbird is a summer resident of lowland marshes. Photo by Ken Driese.
Only 5 percent of Wyomings wetlands are permanently flooded, and two-thirds of them are dry by midto late summer nearly every year.3 At low elevations,
about half of the wetlands are sustained in part by irrigation waters, whether by flood irrigation, leakage from
irrigation canals, or runoff from irrigated fields.4 It is
likely that natural wetlands have a higher biological
diversity than those maintained by irrigation waters.
However, that has not been documented, and many
have water during years when natural wetlands are dry.
Of the 2,692 vascular plant species found in Wyoming, 12 percent are obligate wetland indicators, that
is, plants that cannot tolerate upland environments.
This proportion is surprisingly high, considering that
wetlands occupy such a small area in the state. An additional 13 percent of the plants are ranked as facultative
wetland indicators, that is, plants that usually grow
68
Wetlands
in wetlands but can tolerate moist upland habitats as
in invertebrates, providing a critical food source for
well. In all kinds of wetlands, plants in the sedge fam-
migrating waterfowl and shorebirds. Also, during low-
ily (Cyperaceae) are very common, with 47 percent of
water seasons, some plants and animals are favored that
the 174 species of that family in Wyoming classified as
are less tolerant of high water. Fluctuating water levels
obligate wetland indicators. The grass family (Poaceae)
enable both groups of species to survive. Water diver-
has more species (275) in the state than the sedge fam-
sions intended to keep wetlands filled to a constant
ily, but far fewer are confined to wetlands (8 percent).
depth may not be advisable, though the impulse to do
Among the shrubs, willows are especially common in
so is understandable if most nearby wetlands have been
mountain wetlands and in riparian zones, with 11 of the
drained, or if they have been severely altered in some
states 41 species (27 percent) found only in wetlands.
other way.
Other wetland shrubs include greasewood, resin birch,

and shrubby cinquefoil. Numerous sedges, grasses, and
other graminoids and forbs are found in all kinds of
Surviving in the Wetland Environment
wetlands (fig. 5.4). Trees, though, are largely riparian or
An abundance of water would seem to be an unalloyed
found on lake shores.
benefit in semi-arid climates, but prolonged soil satu-
Mosses and algae also can be an important compo-
ration creates oxygen deficiencies that many species
nent of wetland vegetation. A study of 17 fens in the
cannot tolerate.12 Those that do survive have various
Medicine Bow Mountains documented 30 species of
adaptations. To illustrate, microbial organisms derive
mosses, compared to 105 species of vascular plants. In
their energy in ways that do not require oxygen, using
Yellowstone National Park, a survey of 166 fens found
nitrates, sulfates, iron, manganese, and even carbon
44 species of mosses and 254 species of vascular plants.
dioxide instead.13 By changing these elements from oxi-
Notably, mosses in fens often cover as much of the
dized to reduced forms, they also create the chemical
ground surface as vascular plants. Algae are less con-
environment characteristic of hydric soils. For example,
spicuous but are an important source of food for her-
reduced iron and manganese are water soluble and are
bivorous aquatic invertebrates.6
leached from the soil, leaving white or pale gray hori-
Among the vertebrates, amphibians have the most
zons (known among soil scientists as gleyed horizons).
obvious affinity for wetlands, because the adults need
Similarly, much of the sulfur is reduced to hydrogen sul-
water for breeding and their tadpoles are confined to
fide, the marsh gas with an odor of rotten eggs. Reduced
water. Of the 12 species of salamanders, frogs, and
nitrogen gases are commonly lost from wetland soils,
toads that live in Wyoming, 10 are closely associated
so nitrogen can be a limiting factor for plant growth.
with water. Birds are the most conspicuous wetland
Similarly, carbonwhich exists in many organic forms
vertebrates, with 82 of Wyomings 434 species.8 Over
in hydric soilscan be lost as methane gas. Even so, the
half of those (44) are summer residents only.9 Of the
rate of carbon accumulation in undisturbed wetlands
118 species of mammals in Wyoming, approximately
usually exceeds the rate of loss.14
35 (30percent) use wetlands for more than a source of
Woody plants generally avoid the problems of anaer-
water.10 They include the beaver, muskrat, moose, river
obic soils by growing in microenvironments where oxy-
otter, raccoon, meadow jumping mouse, meadow and
gen is more readily available, such as raised hummocks
water voles, 3 species of skunk, 16 species of bats, and
and wetland edges, or on slopes where groundwater
8species of shrews.
moving through the soil supplies sufficient oxygen.
As noted, wetlands sometimes dry out, or at least
Mosses use the same strategy by growing on the wet-
the water level fluctuates dramatically. This variation
land surface. Some herbaceous vascular plants that are
seems unfortunate for many wetland species, but there
rooted deeper in the saturated soil obtain their oxy-
are benefits. Drying excludes predatory fish and allows
gen via aerenchyma, a spongy tissue in their stems and
the development of food webs composed entirely of
roots through which oxygen diffuses from the atmo-
invertebrates, including mollusks, crustaceans, and
sphere above the water to the respiring root cells. This
insects.11 Saline wetlands in the region can be rich
exchange can be more than passive diffusion, as some
Fig. 5.4. Seven grass-like plantsgraminoidsthat are common in some wetlands. The two species of cattail can be
distinguished by leaf width and the presence of a gap between
the spikes of male and female flowers. The male flowers of

both species persist for only a short time in early summer.
Drawings by Judy Knight.
70
Wetlands
wetland plants develop a pressure gradient that acceler-
Among both plants and invertebrates, the mix of spe-
ates the transport of oxygen to the roots.15 Plants with
cies in a wetland at a given time depends on the length
aerenchyma include buckbean, common reed, horse-
and timing of the preceding wet and dry periods.
tails, pond lily, sedges, and white marsh marigold.
Similarly, wetland vertebrates have evolved to opti-
Another problem for wetland plants is the toxicity
mize their chances of survival in seasonal wetlands.
of reduced sulfur, iron, and manganese compounds
Wetland birds, for example, fledge their young in the
produced by the soil microbes. Adapting to this stress
spring while marshes are usually flooded and emergent
involves the diffusion of oxygen from fine roots into the
cattails and bulrushes provide nest sites and cover. Simi-
millimeter-thick environment immediately adjacent to
larly, tadpoles develop and metamorphose into adults
roots, known as the rhizosphere. Oxidation in the rhi-
quickly, usually before the ponds dry out. Animals with
zosphere essentially immobilizes the toxic compounds
limited mobility, such as many amphibians, must be
and renders them virtually harmless. The oxidized iron
adapted for dry periods and winter. Spadefoot toads
of the rhizosphere remains long after the roots have
burrow into moist soil or occupy burrows dug by small
died and decayed, appearing in the soil as red or orange
mammals on the nearby upland, where they hibernate
streakinga diagnostic sign of hydric soils.
until the next favorable period.16
Vertebrates and terrestrial invertebrates avoid the

oxygen deficiency problem by living at the well-aerated
wetland surface. Aquatic invertebrates have the same
Marshes
array of adaptations as aquatic organisms in lakes and
For many, the word wetland brings to mind blackbirds,
streams, such as gills capable of extracting oxygen from
marsh wrens, muskrats, and a variety of emergent plants
the water.
that includes cattailsall characteristic of marshes.
As noted, two-thirds of Wyoming wetlands usually
Of the various wetland types, marshes are flooded to
dry out every year. In such wetlands, the microbes,
a greater depth and for longer periods. The substrate
plants, and animals must be adapted for drought as
of silt, clay, and organic matter is soft and, when wet,
well as soil saturation. Perennial plants that survive as
is commonly referred to as mud or muck. Hard-bottom
roots or rhizomes actually may benefit from short dry
marshes have more sand. The frequency, duration, and
periods, because some of the soil organic matter decom-
depth of flooding exert strong influences on the kinds
poses when exposed to the air, increasing the supply of
of plants and animals that are present. Many lakes do
nutrients during the next wet period. Another group of
not have marshes, primarily because their banks often
plants persists as seeds that germinate when water again
slope quickly into deep water, and wave and ice action
floods the wetland. A third group consists of plants
prevent plant establishment, or because water levels fluc-
that occupy the wetland during the dry period but are
tuate excessively, such as on the shorelines of reservoirs.17
absent during the wet period. These plants grow from
In the mountains, marshes tend to occur in small
seeds that persist in the soil during the wet period or
areas, such as closed depressions in glaciated terrain,
that blow into the dry wetland from nearby shoreline
also known as potholes (fig. 5.5). Other mountain
plants. Considering that some organisms benefit from
marshes are often associated with old beaver ponds and
dry or low water conditions, it is not surprising that
abandoned stream channels.18 Two common sedges
the diversity of plants and animals in wetlands can be
Northwest Territory sedge and water sedgeare the
greater if water levels fluctuate.
common dominants in shallow and moderately deep
Invertebrates also have strategies for living in sea-
standing waters in high-elevation marshes. Forbs occur
sonal wetlands. Immobile or slow-moving species, such
less frequently, but one can often find twinleaf bed-
as clams, survive dry periods buried in moist sediments,
straw, elephanthead lousewort, white marsh marigold,
either as adults or larvae, or they survive as eggs that
and pond lily, among others (table 5.1). Willows along
hatch during the next wet period. In contrast, aquatic
the margin of some mountain wetlands are commonly
invertebrates that metamorphose into flying adults
Booths willow, diamondleaf willow, Drummonds wil-
leave the drying wetland and fly to nearby water bodies.
low, Geyers willow, and Wolfs willow.

Fig. 5.5. Pothole marsh on
the slope of Signal Mountain
in Grand Teton National
Park. Note the abundance
of pond lily with floating
leaves. Growing next to the
marsh is bluejoint reedgrass,
rough bentgrass (reddish), and
Northwest Territory sedge.
At low elevations, common marsh plants include
gated fields. Small marshes sometimes occur in surpris-
Nebraska sedge, cattail, common spikerush, threesquare
ing places, such as in swales between sand dunes, where
bulrush, and hardstem bulrush. Forbs often are com-
water percolating through the porous dune encounters
mon and may include arumleaf arrowhead and water
relatively impervious sediments (see chapter 9).
knotweed. Marshes in the lowlands are found in aban-
In many marshes and playa wetlands, the vegetation
doned stream channels and backwater channels, but
forms concentric zones, each dominated by a differ-
they are especially common in irrigated areas, more
ent group of plants (figs. 5.6 and 5.7; see also fig. 5.5).
specifically, in the slowly moving waters of ditches and
Plants that need or can tolerate longer dry periods and
in depressions that receive water from canals and irri-
shallower water grow adjacent to the upland, whereas
Table 5.1. Some characteristic vascular plants found in or adjacent to marshes, wet meadows, fens, and playa wetlands in
Wyominga
Common name
Latin name
Marsh
Wet meadow
Fen
Playa wetland
GRASSES
Alkaligrass
Puccinellia spp.
Alkali cordgrass
Spartina gracilis
Alkali sacaton
Sporobolus airoides
Bluejoint reedgrass
Calamagrostis canadensis
LM
LM
LM
Northern reedgrass
Calamagrostis stricta
LM
Common reed
Phragmites australis
Saltgrass
Distichlis spicata
Short-awn meadow foxtail
Alopecurus aequalis
Tufted hairgrass
Deschampsia caespitosa
LM
LM
RUSHES, SEDGES, AND ARROWGRASS

Marsh arrowgrass
Triglochin palustris
Seaside arrowgrass
Triglochin maritima
Threesquare bulrush
Schoenoplectus pungens
Hardstem bulrush
Schoenoplectus acutus
Cottongrass
Eriophorum spp.
LM
Mountain (Baltic) rush
Juncus arcticus
LM
LM
Analogue sedge
Carex simulata
LM
Blister sedge
Carex vesicaria
LM
Buxbaums sedge
Carex buxbaumii
LM
Clustered field sedge
Carex praegracilis
LM
Mountain sedge
Carex scopulorum
Mud sedge
Carex limosa
Nebraska sedge
Carex nebrascensis
LM
LM
Northwest Territory (beaked) sedge
Carex utriculata
Sheep sedge
Carex illota
Water sedge
Carex aquatilis
Common spikerush
Eleocharis palustris
LM
LM
Few-flowered spikerush
Eleocharis quinqueflora
LM
SHRUBS
Alpine laurel
Kalmia microphylla
Alpine wintergreen
Gaultheria humifusa
Bog blueberry
Vaccinium uliginosum
Greasewood
Sarcobatus vermiculatus
Silver sagebrush (mountain)
Artemisia cana ssp. viscidula
Silver sagebrush (plains)
Artemisia cana ssp. cana
Purple marshlocks
Comarum palustre
LM
Common name
Latin name
Marsh
Wet meadow
Fen
Playa wetland
Resin birch
Betula glandulosa
Shrubby cinquefoil
Dasiphora fruticosa ssp.

floribunda
Booths willow
Salix boothii
Diamondleaf willow
Salix planifolia
Drummonds willow
Salix drummondiana
Geyers willow
Salix geyeriana
Sageleaf willow
Salix candida
Wolfs willow
Salix wolfii
LM
LM
Alkali buttercup
Ranunculus cymbalaria
LM
LM
Alpine leafybract aster
Symphyotrichum foliaceum
American globeflower
Trollius laxus
Arumleaf arrowhead
Sagittaria cuneata
Balsam groundsel
Packera paupercula
Broadleaf cattail
Typha latifolia
OTHER HERBACEOUS PLANTS
Narrowleaf cattail
Typha angustifolia
Buckbean
Menyanthes trifoliata
Elephanthead lousewort
Pedicularis groenlandica
English sundew
Drosera anglica
Field mint
Mentha arvensis
LM
LM
Horsetail
Equisetum spp.
LM
LM
Knotweed
Polygonum spp.
LM
Largeleaf avens
Geum macrophyllum
Marsh grass-of-Parnassus
Parnassia palustris
Narrowleaf bur-reed
Sparganium angustifolium
Neckweed
Veronica perigrina
LM
LM
Red swampfire
Salicornia rubra
Redpod stonecrop
Sedum rhodanthum
Redwool plantain
Plantago eriopoda
Sea milkwort
Glaux maritima
Seepweed
Sueda spp.
Spiral ditchgrass
Ruppia cirrhosa
Subalpine fleabane
Erigeron peregrinus
White marsh marigold
Caltha leptosepala
Riparian species are listed in table 4.1. A dash indicates that the species is absent or uncommon. L = present mostly at low elevations;
LM = present at both low and high elevations; M = present mostly in the mountains.
74
Wetlands
Fig. 5.6. Vegetation around desert playa wetlands changes

with distance from open water, probably because soil salinity
decreases and depth to water table increases. Salicornia, also
known as red swampfire, is a succulent that tolerates high
salinity.
and figs. 5.6, 5.7, and 8.4).20 Red swampfire, also known
as salicornia or saltwort, is a short, annual forb with
succulent leaves that grows on many salt flats and is
conspicuous when large numbers of the plant form a
scarlet band in late summer along the edge of white salt
deposits. Many salt flats are virtually unvegetated much
hydrophytes (such as cattails and hardstem bulrush)
of the time. In the Laramie Basin, for example, mudflats
dominate the deeper water. Different plants and animals are associated with each of the zones, an illustration of how patchy environments contribute to the
biological diversity of the landscape as a whole.
Most large marshes are found in areas that have
received water from irrigation or diversions. For ex
ample, cattails occur along the shore of Ocean Lake
in the Wind River Basin, which receives much of its
water supply from irrigation drains. In Goshen Hole
of eastern Wyoming, marshes with cattails and bulrushes occur on the edges of the lakes and ponds
enlarged or constructed on the Table Mountain and
Springer
/
Bump-Sullivan Wildlife Habitat Management units. In the Laramie Basin, the water levels of
Hutton Lake National Wildlife Refuge are managed by
U.S. Fish and Wildlife Service biologists to maintain
large stands of hardstem bulrush.
Playa Wetlands
Wetland desiccation is more likely to occur at low elevations, which sometimes leads to the formation of playa
wetlands (see figs. 5.6 and 5.7).19 When the surface water
evaporates completelymore frequently in playa wetlands than in marshesthe salts precipitate and become
sufficiently concentrated to exclude most plants. Only a
few halophytes tolerate such environments, including
greasewood, red swampfire, and saltgrass (see table 5.1
Fig. 5.7. Playa wetland in the fall, when most of the water is
evaporated and red swampfire is conspicuous on the white,
salt-crusted soil surface. The tan vegetation on the left is
dominated by saltgrass. Photo taken in the Laramie Basin at
an elevation of 7,000 feet.

Fig. 5.8.Elongated wet
meadow south of Centennial,
with Sheep Mountain in the
background. This meadow
is defined by the presence
of shrubby cinquefoil, with
mountain big sagebrush and
scattered limber pine on the
upland. Elevation 8,100 feet.
with essentially no plants surround ponds with salinity
noids and forbs are the dominant plants; shrubby
greater than about 5 parts per 1,000. Ponds with less-
cinquefoil is common. (figs. 5.8 and 5.9).
saline waters support patches of marsh-like vegetation.
Sites that support wet meadows are diverse and occur
Playa wetlands are widespread on the plains and in
from the high mountains to the basins and plains.23
the basins of eastern and southern Wyoming. 21 Where
Commonly they are part of riparian landscapes. In the
water persists, the plants are mainly graminoids, such as
alpine zone, small meadows occur downslope from large
common spikerush, needle spikerush, threesquare bul-
snowbanks, where meltwater saturates the soil in spring
rush, mountain rush (also known as Baltic rush), and
and early summer. Broad glaciated valleys typically
meadow foxtail. Small forbs also are common, such as
support a mosaic of wet meadows and fens, with the
annual knotweeds. Around the depressions edge, where
meadows on the lower valley slopes and fens on wetter
facultative hydrophytes mix with upland plants, the
sites where groundwater discharges. Abandoned beaver
soil may lack the characteristic hydric features.
ponds are common sites for wet meadows, which replace

marshes as the ponds fill with sediments. In mountain
Wet Meadows
meadows, black alpine sedge and mountain sedge are

common at the highest elevations. Other species occur
Wet meadows develop where water tables are at or near
over a wide range of elevations, for example, tufted
the soil surface in the spring and early summer but
hairgrass, mountain rush, small-winged sedge, blister
where the water level drops below the surface as the
sedge, and water sedge. Common mountain forbs are
growing season progresses. Plant materials decompose
alpine leafybract aster, white marsh marigold, Ameri-
when water levels are low and oxygen is more readily
can globeflower, and largeleaf avens. Meadow shrubs
available, which prevents the accumulation of thick
include mountain silver sagebrush, shrubby cinquefoil,
layers of peat. Although wet meadows may not be rec-
and willowsespecially diamondleaf willow. Some
ognized as wetlands during part of the year, they have
meadows have substantial amounts of various mosses,
hydrophytes and the diagnostic hydric soils.22 Grami-
though they are more common in fens.
76
Wetlands
Fig. 5.9. Wet meadow in a bay on Upper Green River Lake,

in the foothills of the Wind River Mountains. The shrubs
are shrubby cinquefoil, and the single tree (foreground) is a
small lodgepole pine, both of which are growing where the
soils have more sand and are less anaerobic. The meadow is
dominated by graminoids, including blister sedge, bluejoint

reedgrass, fowl bluegrass, timber oatgrass, and tufted hairgrass. The forests are dominated by lodgepole pine, Engelmann spruce, and subalpine fir. Elevation 8,000 feet.
At low elevations, wet meadows commonly develop
and duration of high water. 25 In wetter areas the veg-
because of flood irrigation or leakage from irrigation
etation is composed of both obligate hydrophytes (like
ditches. Hayfields are an example, though they have
Nebraska sedge and water sedge) and facultative hydro-
mostly introduced species, such as Garrison creeping
phytes (such as bluejoint reedgrass and tufted hair-
foxtail, smooth brome, and orchardgrass. Native species
grass). The drier parts of meadows contain facultative
include tufted hairgrass, Nebraska sedge, clustered field
hydrophytes as well as upland plants.
sedge, common spikerush, and mountain rush. Wet
Because meadows remain moist during much of
meadows also occur on gently sloping valleys and broad
the growing season and are adjacent to rangelands on
flats without defined streams. Many low-elevation wet
the upland, plant growth is relatively high and they
meadows have areas with elevated salt concentrations
are often heavily grazed. 26 As in grasslands, prolonged
and halophytic species, such as alkali buttercup, alkali
grazing may cause changes in species composition,
cordgrass, alkali sacaton, Nevada bulrush, saltgrass, sea
from palatable to less palatable species. Also, excessive
milkwort, and seaside arrowgrass (see table 5.1). Wet
grazing can hasten drying and often causes the forma-
meadows can merge with salt flats, where few plants
tion of hummocks on the surface, a process known as
survive.24
pugging.27
Plants in wet meadows grow along a gradient in wet-
Wet meadows are desirable for real estate develop-
ness, from very wet sites that merge into marshes and
ments because of their proximity to surface water as well
fens, to drier sites that border upland vegetation. The
as to the upland, plus the availability of livestock forage.
important feature in this gradient may be the height
Federal laws restrict construction in wetlands because

Fig. 5.10. Fens are common south of
Mammoth Hot Springs in Yellowstone
National Park. Northwest Territory
sedge and water sedge are abundant on
the usually wet organic soils; diamond
leaf willow, Wolfs willow, resin birch,
and shrubby cinquefoil are found along
the margin; wet meadows and lodgepole
pine occur on mineral soils that are
somewhat drier.
of their widely recognized ecological and hydrological
by precipitation that has not passed through the soil or
values. Thus, considerable controversy can arise about
rock and contains only the nutrients that settle on the
whether a tract of land is in fact a wet meadow or a
wetland in dust, rain, or snow.28 Both kinds of peatland
grassland or shrubland. Such disputes are settled based
store far more water and nutrients than an equal volume
on the abundance of obligate hydrophytes, the pres-
of mineral soil, though many of the nutrients are not
ence of hydric soils, and an understanding of the sites
available to plants because they are chemically bound
hydrology.
in organic material. Mosses do well in this environment,

probably because the reliable supply of surface water
Fens
minimizes water stress late in the summer and they are

able to obtain the small amount of nutrients required
Fens are rich in graminoids and forbs, but shrubs are
by such small plants. Peat in fens is composed mostly
more commonespecially willows and a shrub known
of slowly decomposing sedges and mosses. The true peat
as resin birch (figs. 5.10 and 5.11). The peat substrate is
mosssphagnumis found in some of them.29
saturated all year. Many people think of fens as bogs,
Fens are most common in glaciated mountain land-
but wetland ecologists have determined that all peat-
scapes, such as where potholes have formed and in val-
lands in the Rocky Mountains are fens. The distinction
leys. They also occur on slopes where a reliable source of
is based on nutrient availability and acidity. Bogs have
groundwater comes to the surface in springs and seeps,
low nutrient availability and are highly acidic (pH <
sometimes developing under the canopy of trees. 30
5). In contrast, fens are fed by groundwater containing
Most mountain fens are small, between 1 and 10 acres.
relatively high nutrient content and are either basic or
Fens in the alpine tundra can be associated with palsas,
mildly acidic (pH 58). Some of the plants are different
which are vegetated domes caused by frost heaving.31
as well.
The differences between bogs and fens are caused
Fens also occur in sagebrush steppe in the Green

River Basin and on the Sweetwater Plateau in west-
by their sources of water: fens are supported by waters
central Wyoming. The latter site, known as Ice Slough
that have moved through the soil or underlying rock,
(see fig. 5.11), is the subject of a historical marker west
from which nutrients are dissolved; bogs are supported
of Jeffrey City. It became famous because immigrants
78
Wetlands
Fig. 5.11.In the mid-1800s,
summer travelers along the
Oregon Trail would dig for
ice preserved in the peat of
this fen, known locally as Ice
Slough. Photo taken along
Highway 287, 40 miles east of
Lander. Elevation 6,500 feet.
Photo by Ken Driese.
along the Oregon Trail were happy to find ice preserved
stone National Park, adjacent to Wyoming Highway 296
in the insulating peat there. Core samples show that Ice
(see fig. 5.10).34 This large wetland complex lies on gra-
Slough is underlain by 510 feet of peat.
nitic bedrock and glacial deposits, but it benefits from
Fens sometimes merge with marshes as water depth
waterborne nutrients coming from the calcium- and
increases. In some places the fen community extends
magnesium-rich limestone and dolomite of the nearby
over the deeper water, with the peat forming a floating
Cathedral Cliffs to the south.35
mat. Eventually, the peat may cover the entire pond or
Fens share many plant species with shallow marshes
pothole, with the only evidence of a former pond being
and wet meadows (see table 5.1). In the wetter parts
the undulation of the surface when walked on, or the
of fens, Northwest Territory sedge and water sedge are
surprise of ones foot slipping into the water below.
ubiquitous across a broad elevation range, along with
Deciding where the fen ends and a shallow marsh or
other sedges. Some unusual graminoids grow in many
wet meadow begins, or even if a wetland contains a fen
fens, such as cottongrass and grass-of-Parnassus. Com-
at all, can be a challenge. Peat thickness can be used,
mon forbs are elephanthead lousewort, redpod stone-
but fens at high elevations commonly have thinner peat
crop, and white marsh marigold. Floating peat mats
than those at low elevations.32
usually are composed of analogue sedge, mud sedge,
Wetland scientists customarily identify different
and slender sedge. Along the edge can be found shrubs,
kinds of fens based on acidity and concentrations of
such as diamondleaf willow, mountain willow, Wolfs
calcium, magnesium, and other minerals in the water.
willow, small-leaved laurel, and resin birch.
At the acidic, infertile end of the gradient are poor fens,
Sometimes mosses are as conspicuous as vascular
similar to bogs. With decreasing acidity and increasing
plants. 36 Growing on the well-aerated surface of the
mineral concentrations, fens are labeled inter mediate,
peat, they respond readily to differences in surface
rich, and extremely rich. In general, the number of plant
water chemistry across small distances. In contrast,
species increases with nutrient availability. Most fens
vascular plants rooted in the peat are influenced more
in the Rocky Mountains lie on granite or other rela-
strongly by differences in climate, topography, and
tively nutrient-poor rocks and are intermediate.33 The
elevation, along with differences in water depth, water
most nutrient-rich fen identified in Wyoming is at
flow, and peat thickness.37 Different groups of plants are
Swamp Lake, in the Clarks Fork valley east of Yellow-
associated with floating mats, areas of standing water,
hummocks, fen margins, and shallow channels with
as allelopathy (which has been difficult to demonstrate
running water.38
except in laboratory experiments).

Although both reed canarygrass and common reed
Management Issues and Future Challenges

Wetlands worldwide have long suffered degradation
are native to North America, they are also native to

Europe. Such a widespread distribution can be attributed to great genetic variation within the species. At
and outright destruction caused by myriad factors,
some point during the past century, seeds from Euro-
including the perceived need for drainage and water
pean populations of both species were introduced to
diversion, the accumulation of excess nutrients and
North America, where the resulting plants interbred
contaminants, and excessive grazing by livestock (see
with native populations. As it turns out, the hybrids
chapter 4). In Wyoming, 3540 percent of the original
appear to be more aggressive invaders than the native
39
wetlands have been lost, and elevated levels of ground-
populations. Thus, where reed canarygrass and com-
water contaminants are sometimes present.40 Generally,
mon reed appear to be invading wetlands, most likely
mountain wetlands have been less affected because
the plants involved are EuroAmerican hybrids. Slight
water is more readily available and the duration of graz-
genetic changes can greatly improve a plants ability to
ing is shorter.41 Also, pollutants are less widespread in
compete with its neighbors.
the mountains. Peat mining has severely disrupted
Invasive hybrids have also formed in cattails. Two
some fens in the mountains of Colorado,42 but that has
native species occur throughout much of North Amer-
not taken place in Wyoming.
icabroadleaf cattail and narrowleaf cattail. Until
Invasive plants have become a severe problem in
recently, the narrow-leaved species was so rare in mid-
many wetlands.43 Ecologists think of wetlands as land-
western and western states that some botanists, after
scape sinks, that is, low places on the landscape where
observing how its range was expanding, came to think
water, nutrients, sediments, and salts accumulate, and
of it as an introduced invasive plant.46 Indeed, narrow
where water levels fluctuateall conditions that favor
leaf cattail has expanded its range. More important,
weedy plants. Various Eurasian species are common
though, the two native speciesnow growing together
in Wyomings wet meadows, namely, Canada thistle,
more commonly than ever beforeare hybridizing. The
creeping bent grass, creeping foxtail, Kentucky blue-
sterile offspring, known as hybrid cattail, spread rapidly
grass, and timothy.
by rhizomes and are highly invasive.47 Very likely, Euro-
Some invasive wetland plants are now thought of
pean genes are a contributing factor.
as natives. Kentucky bluegrass is an example, as it has
An especially worrisome invasive plant in marshes
been in North America long enough to be viewed as
is purple loosestrife, a colorful forb that grows 3 or 4
naturalized. In contrast, some species that are veritable
feet tall. Though still rare in Wyoming, this European
natives behave like invasive species in altered wetlands.
species has overwhelmed native wetland plants in
For example, reed canarygrass expands rapidly in wet-
midwestern and eastern states. Seasonal wetlands are
lands created by irrigation. Such habitats could have
most vulnerable to invasion, because loosestrife seed-
been invaded by introduced species, but native plants
lings establish on bare sediments during dry periods.
often colonize the irrigated lands first. The species does
Preventing the spread of localized invasives like purple
provide good forage, but it tends to reduce the plant
loosestrife is generally more feasible than trying to
diversity of the areas where it becomes the dominant
reduce large populations once they are well established.
plant.44 Another invasive native, common reed (also
Also, early control is less damaging to the environment.
known as phragmites), grows to more than 6 feet tall
Biological control of purple loosestrife shows some
and is rapidly invading wetlands, especially at lower
promise.
This invader is thought
In addition to invasive species and the loss or degra-
to benefit from chemicals produced and released by
dation of wetlands, there are now concerns about the
the plants that suppress the growth of other species,
effects of climate change. Warmer, drier conditions
thereby facilitating its spreada phenomenon known
likely will result in shallower water, causing seasonal
elevations in adjacent states.
45
80
Wetlands
wetlands to dry up for longer periods and, for example,
ing through the soil from ditches and irrigated fields.53
converting marshes to playa wetlands.48 Sediments in
Surface inflows tend to be less saline, enabling different
Great Plains wetlands indicate that such changes have
wetland communities than where subsurface flows are
occurred in response to climate changes throughout pre-
the primary source.
history. During long dry periods, obligate hydrophytes

became less abundant and facultative hydrophytes
In sum, recognition of wetland values, their vulnerabil-
more abundant. In fens, increased decomposition rates
ity to degradation, and their poorly protected status has
resulting from warming and less flooding could cause
led state and federal agenciesalong with motivated
peat deposits to thin.49 Many kinds of wildlife and other
private organizations and landownersto promote
organisms will be affected adversely as the area in wet-
wetland conservation. One federal initiative is the Wet-
lands is reduced (see chapter 4). Ecologist Holly Cope-
land Reserve Program, administered by the U.S. Depart-
50
land and her associates conducted a recent assessment
ment of Agricultures Natural Resources Conservation
of Wyoming wetlands, concluding that climate change
Service. Landowners are paid to protect wetlands. Simi-
is most likely to affect wetlands at low elevationsthe
larly, the Wyoming Wetlands Conservation Strategy
same wetlands that are vulnerable to rural subdivisions
identifies wetlands where collaborative conservation
and oil and gas developments.51
management is needed and will work. 54 Unfortunately,
Agriculture is usually considered a cause of degrada-
wetlands often occur in places considered desirable for
tion in wetlands. However, as noted previously, many
new construction. When this happens, the prevailing
wetlands at lower elevations have been created or aug-
attitude is to construct new wetlands, to compensate
mented by irrigation projects. In Star Valley; Goshen
for those that are lost. Although commendable, this no-
Hole; and the Bighorn, Laramie, and Wind River basins,
net-loss policy sanctions the destruction of wetlands in
more than 50 percent of the wetlands receive irrigation
one place if a new wetland of comparable area is created
waters.
In the Laramie Basin of southeastern Wyo-
elsewhere. Many wetland scientists doubt that newly
ming, 30 percent of the inflows to wetlands were from
created wetlands provide the same benefits as those that
irrigation ditches, and 35 percent were waters percolat-
have been lost.55
52
Part Three Plains and
Intermountain Basins
Chapter 6
Grasslands
Grasslands are characteristic of the plains east of
scarlet globemallow, and various species of milkvetch
the Rocky Mountains and in several intermountain
and locoweed. Foothill grasslands at slightly higher
basins (see fig. 1.5). Some have been plowed for crop
elevations may have some of the same species, but
productionmostly winter wheatbut large areas of
others are more typical, such as bluebunch wheatgrass.
native vegetation can still be found (fig. 6.1). Prong-
Some plants are good indicators of soil characteristics.
horn are common. Diversity is added to the landscape
For example, sandy soils often have an abundance of
by scattered woodlands of ponderosa pine, limber pine,
Indian ricegrass, prairie sandreed, sand dropseed, sand
and juniper on outcrops of resistant bedrock, and by
sagebrush, and yucca, whereas saline soils have alkali
sand dunes and badlands (see chapters 8 and 9).
sacaton, fourwing saltbush, greasewood, and inland
Most grasslands in Wyoming have been classified
saltgrass. The most common soil types of upland
into two categories. One, known as shortgrass prairie,
grasslands, known as Aridisols, have accumulations of
is found only in the southeast corner of the state and
sand, silt, clay, and organic matter along with varying
has buffalo grass and blue grama as co-dominants. The
amounts of lime (calcium carbonate), gypsum (hydrated
other, mixed-grass prairie, covers a larger area (about
calcium sulfate), or various soluble salts. In swales and
17 percent), extending northward into Montana. Usu-
along drainages, where plant growth is comparatively
ally mixed-grass prairie does not have buffalo grass.
high for a longer period of time each year, the soils have
Sometimes the grasses in both types of grassland grow
more organic matter and are known as Mollisols (see
in small clumps and are known as bunch grasses, most
appendix B).
notably blue grama, junegrass, Sandberg bluegrass, and
Some grasslands mergegradually or abruptlyinto
Indian ricegrass. Others, such as western wheatgrass and
shrublands dominated by sagebrush or woodlands dom-
prairie sandreed, have rhizomes and are sod formers. 2
inated by juniper, limber pine, or ponderosa pine. Silver
Grasses provide most of the biomass in grasslands, but
sagebrush is common on sandy soils, in swales, and
forbs, succulents, and small shrubs provide most of the
along drainages; Wyoming big sagebrush is common
plant diversity. 3
in the intermountain basins (see chapter 7). Sand sage-
Mixed-grass prairie often has 50 or more differ-
brush is common on dunes, such as near Torrington,
ent kinds of plants in an area as small as 2 or 3 acres.
though it is less widespread in Wyoming than in east-
Common species include blue grama, Indian ricegrass,
ern Colorado. Around the world, shrubs are invading
junegrass, needle-and-thread grass, Sandberg bluegrass,
grasslands in a surprising number of places, a trend that
western wheatgrass, needleleaf sedge, threadleaf sedge,
has been attributed to various combinations of factors,
fringed sagewort, Hoods phlox, pricklypear cactus,
including less frequent fires, excessive grazing by large
83
84
Plains and Intermountain Basins
Fig. 6.1. Ponderosa pine grows on ridgetops adjacent to this

mixed-grass prairie on the western Great Plains; cottonwood
sometimes grows along creeks. The tan grass in the foreground is little bluestem. Other common plants are blue
grama, Sandberg bluegrass, western wheatgrass, junegrass,

needle-and-thread grass, and fringed sagewort. This grassland is near the Black Hills, at an elevation of about
4,500feet.
herbivores, nitrogen enrichment from atmospheric fall-
times causes the formation of patches that are one to
out in the form of dust and rain, and elevated levels of
several yards across (such as those for blue grama, prairie
carbon dioxide in the atmosphere.
sandreed, and yucca). Another cause of patchiness is the
Wherever grasslands occur, plant species composi-
burrowing of animals, such as badgers, ground squir-
tion varies with changes in topographic position, such
rels, harvester ants, pocket gophers, and prairie dogs.
as from hilltops to valley bottoms. Valley bottom soils
Their small disturbances favor some plants over others.
are deeper, finer-textured, moister, and more fertile than
Also, fairy rings may developan interesting but
soils on hilltops. Topography also affects snow drifting,
poorly understood phenomenon thought to be caused
with more snow accumulating in ravines and on leeward
by saprophytic fungi (fig. 6.2).5
slopes, causing them to be wetter than the surrounding

area. Depressions or playas with little or no drainage are
common, and typically they have saline or alkaline soils.
Surviving in the Grassland Environment
Differences in soil depth, salinity, and texture cause con-
Most of the grassland ecosystem is hidden in the
siderable variation in what sometimes appears as a uni-
soil, where 75 percent or more of the plant biomass
form vegetation covermonotonous during mid-day but
is found and, not surprisingly, most of the herbi-
inspiring when the sun is low in the sky.
vores also (fig. 6.3).6 Extensive root systems, required
Other patterns occur on a finer scale. For example,
to obtain water for the relatively small aboveground
cloning from root crowns, rhizomes, and stolons some-
shoots in this semi-arid environment, provide food
Grasslands
nial and herbaceous, having their perennating buds at

or just below the soil surface. This growth form is the
result of several factors: first, the semi-arid nature of
mixed-grass prairie limits the production of woody
stems aboveground. Second, and perhaps most important, is the frequency of fires in grasslands. Plants with
buds at or near the soil surface are less likely to be
killed, because the soil surface temperature during a
fire is relatively low. Most buds survive, and only the
herbaceous stems and leaves are burned. After the fire,
the buds sprout and grow rapidly. Third, plants with
their buds near the soil are well adapted to tolerate
grazing by large herbivores, which 10,000 years ago
included the camel and horse as well as bison, elk, and
pronghorn. Buds close to the ground, or buried in the
soil, are less likely to be eaten by the herbivores. Only
the easily regenerated stems and leaves are consumed
(fig. 6.4).
The ability to replace leaves and stems that have
been eaten or burned depends on the presence of special meristemstissues capable of cell division and
new growthand the capacity for energy storage in
the undamaged part of the plant belowground. The
meristems are contained in dormant buds on rhizomes
and root crowns, which are stimulated by herbivory to
Fig. 6.2. An aerial view of grasslands in the Laramie Basin
showing fairy rings that are about 30 feet in diameter, created by fungi that grow outward. A flush of nutrients is made
available as the fungi die and decompose, just inward from
the zone of major fungal activity, thereby stimulating plant
growth. Elevation 7,300 feet.
produce new stems and leaves. Another way that grass

leaves can be replaced is through intercalary meristems,
which are tissues capable of cell division that occur near
the bases of the leaves and some stems. As grass leaves
are eaten, new leaf tissue is produced by the leaf itself.
No buds are involved.7
Regrowth, whether from buds or intercalary meri-
for the largest group of herbivores in the ecosystem
stems, requires energy stored as carbohydrates in roots,
invertebrates. As the roots and other forms of life die,
rhizomes, bulbs, and corms. Carbohydrates are pro-
they become part of the soils organic matter. Some
duced by photosynthesis aboveground but are mostly
plants have deep roots for extracting moisture from
moved belowground soon thereafter. Considering that
throughout the soil profile, whereas others have shal-
75 percent or more of grassland plant biomass is below-
low roots to take advantage of light showers that wet
ground, there is a large amount of energy available to
only the surface soil. Generally, about 70 percent of
keep the plant alive and enable the production of new
the roots are within the top 4 inches.
leaves and stems.8 Of course, plants die from a lack of
Throughout the western Great Plains, grassland
energy if fire, herbivory, or drought are too frequent to
plants have experienced extended periods of drought,
allow for the replacement of the stored energy spent in
recurring fire, and grazing by herbivores. These envi-
previous episodes of regrowth.
ronmental factors led to a community composed
Most grassland plants have characteristics of xero-
largely of grasses but with a substantial number of
phytes, that is, plants adapted to dry environments.
sedges and forbs. Notably, most of the plants are peren-
Typically, they have small or narrow leaves, and some
85
86
Fig. 6.3. Most grassland biomass is in the soil. The enlargement in this drawing
shows a single root with root hairs and the hyphae of mycorrhizal fungi. A film
of water (stippled area) coats each soil particle and provides habitat for nematodes and numerous protozoans, including amoebae and ciliates. Air spaces
provide habitat for mites, insect larvae, and other invertebrates. Bacteria are
extremely abundant but are too small to illustrate at this scale. Magnification
about 15 times. Based on Weaver (1968) and Hunt et al. (1987).
species have fine, light-colored hairs (pubescence)
grazed heavily by thirsty herbivores were it not for spe-
on the leaves. Small size reduces the area exposed for
cial defense mechanisms, such as sharp spines.
heat absorption and transpiration on warm days, and
Animals, too, must be adapted for surviving in the
the light color reflects some solar radiation that might
grassland environment (table 6.1). With most of the
otherwise cause leaf warming and increased water loss.
plant biomass belowground, there are many subterra-
Most leaves of grassland plants are facultatively decidu-
nean herbivores, such as nematodes, mites, insect lar-
ous, that is, they fall or become senescent after a period
vae, and various burrowing mammals (such as pocket
when conditions for photosynthesis have not been
gophers).9 Aboveground, the primary herbivores include
favorable.
grasshoppers, ground squirrels, prairie dogs, jackrabbits,
Although grassland plants are adapted to tolerate low
and the larger mammals already mentioned. One of
water availability, some avoid drought stress by complet-
the major problems for large herbivores is obtaining
ing their growth early in the spring while water is readily
enough protein from coarse grassland plants with tough
available. Examples of drought-evading plants are spring
cell walls and high concentrations of lignin and silica.
wildflowers, such as sand lily and prairie smoke. Other
Chewing this kind of food causes rapid tooth abrasion.
grassland plants store water in tissues that remain suc-
Notably, large mammals common in grasslands have
culent throughout the dry summer, for example, prickly
teeth that continually grow from their crown. This
pear cactus and stonecrop. Succulent plants would be
adaptation began to appear in the fossil record at about
Grasslands
Fig. 6.4. Grassland plants are tolerant of grazing by large herbivores,
especially if most of the forage is consumed later in the growing season.
Nearly all the plants are perennials,
and most biomass is belowground.
This rangeland in the Laramie Basin
is in excellent condition. Problems
are inevitable near water sources and
where too many animals are confined by fences for too long in areas
that are too small.
the same time that grasslands became widespread, some

20 million years ago (during the Miocene).
Another adaptation of some large grassland mammals is the ruminant digestive system of bison, deer,
elk, and pronghorn (also shared by cattle and sheep).
The four chambers of their stomachs contain bacteria
and protozoans that facilitate digestion and the extraction of protein from coarse plant material. Regurgitation and additional chewing is an important feature of
this digestive system. Nonruminants, such as horses,
can also survive in grasslands, but they must consume
considerably more food, because it passes through their
digestive systems more quickly and with less energy and
nutrients extracted. To facilitate the digestion of plant
tissues in general, all mammalian herbivores, regardless
of their habitats or digestive systems, have longer intestines per unit of body weight than do carnivores.
Partitioning Grassland Resources

Many species of plants coexist in grasslands and occupy
nearly the same space, and many grassland animals
seem to share the same habitat. However, careful study
has demonstrated that there are subtle differences in
the ways that different species obtain what they need
to survive. Some plants, for example, complete their
growth at a different time of year than their neighbors.
Plants and animals that use different resources in the
same community or at different times of the year are
viewed as occupying different ecological niches. There
is a tendency during evolution for the niches of coexisting species to become less similar, reducing the intensity of competition.
In the 1950s, John Weaver at the University of
Nebraska conducted one of the first studies that suggested resource partitioning among grassland plants. He
87
88
Table 6.1. Some characteristic mammals, birds, and reptiles found in grasslands, shrublands, and escarpments at
low elevations
rooted plants can also benefit from water that percolates

to greater depths after snowmelt or heavy spring rains.11
While studying grassland root systems, Weaver would
sometimes dig by hand to a depth of 10 feet or more.
MAMMALS
Badger
Ground squirrel, Wyoming
Bobcat
Jackrabbit, black-tailedb
Chipmunk, least
Jackrabbit, white-tailedc
Cottontail, desert
Mouse, deer
Cottontail, Nuttalls
Coyote
Mouse, northern
grasshopper
Deer, mule
Pocket gopher, northern
Fox, red
Prairie dog, black-tailed
Fox, swift
Prairie dog, white-tailed
Ferret, black-footeda
Pronghorn
Ground squirrel,
thirteen-lined
Woodrat, bushy-tailed
On at least once occasion he decided it was prudent to

abandon the trench because of caving soil. In a sense,
the grassland is the reverse of a forest, where vertical
stratification exists aboveground, with trees above the
shrubs and herbaceous plants in the understory. Forest ecologists must watch for falling branches and even
b
c
whole trees, whereas grassland ecologists think more

about being buried in their trenches.
Grassland plants also partition resources seasonally, with cool-season plants completing their growth
by early summer or in the fall (when moisture is available), and warm-season plants growing in the summer
(until moisture becomes limiting). Junegrass, threadleaf
sedge, and western wheatgrass are examples of cool-
BIRDS
Bluebird, mountain
Meadowlark, western
season species; blue grama, buffalo grass, and little
Bunting, lark
Nighthawk, common
bluestem are examples of warm-season plants.12 This
Eagle, golden
Owl, burrowing
differentiation is based on their physiological charac-
Falcon, prairie
Sparrow, Brewers
teristics. Cool-season species commonly have lower
Grouse, sage
Sparrow, lark
optimal temperatures for photosynthesis and use C3
Harrier, northern
Sparrow, sage
metabolism. Such plants are usually less tolerant of high
Hawk, red-tailed
Sparrow, vesper
temperatures, thus growing early in the spring before
Jay, pinyon
Thrasher, sage
water stress develops. In contrast, warm-season species
Kestrel, American
Towhee, green-tailed
have a higher optimal temperature for photosynthesis,
Lark, horned
Vulture, turkey
can tolerate higher light intensities, and have a higher

water-use efficiency (that is, more photosynthesis per
Longspur, McCowns
unit of water uptake)all characteristics of C4 metabo-
REPTILES
lism. The separation of grasses into C3 and C4 species
Bullsnake
Lizard, northern sagebrush
has stimulated great interest among biologists and land
Rattlesnake, prairie
Lizard, spiny
managers, but it is only one of many ways that grass-
Now rare.
Eastern grasslands of Wyoming.
Western two-thirds of Wyoming.
land plants are adapted to their environment.

Such differences in physiology also appear to affect
responses to grazing and geographic distribution. With
regard to grazing, a critical time for ranchers and their
found that the root systems of some plants are mostly
livestock is in the spring, when hay supplies may be low
deep, whereas others are mostly shallow. In this way,
and there is little new forage production on rangelands.
neighbor
ing plants tap different parts of the soil for
The first plants to become green are the cool-season C 3
water and nutrients. To illustrate, 85 percent of blue
speciesthe plants most likely to be grazed first. The
grama roots are in the top 8 inches of soil. Neighbor-
warm-season species become important later, but by
ing plants with deep root systems, down to 3 feet or
that time there is more food available and less intense
more, include scarlet globemallow, skeleton plant, and
grazing pressure on any one group. Consequently,
slimflower scurfpea. All plants depend to some extent
repetitive early-spring grazing commonly leads to a
on surface soil derived from summer rain, but deeper-
decline in C3 species and an increase in some warm-
10
Grasslands
season species, probably because of less competition
The solar energy used for photosynthesis in grass-
from the plants that grew earlier and were eaten.13 With
lands, and all ecosystems for that matter, is typically
regard to geographic distribution, research has shown
less than 2 percent of the total impinging on the land-
that C3 species are more common at higher elevations,
scape. Nearly all remaining solar energy is reflected,
where one would expect cooler temperatures and less
used for the evaporation of water, or for heating the soil
water stress, and C4 species tend to be more common
and other ecosystem components. The low percentage
at lower elevations, where it is warmer and more often
of the solar radiation fixed by photosynthesis leads to
drier.14
an annual total net primary production (above- and
Coexisting animals also illustrate resource partition-
belowground) of 500900 g/m 2 in mixed-grass prairie,
ing. For example, bison, jackrabbits, and some grass
enough to maintain a diverse fauna of herbivores and
hoppers tend to eat more grasses, whereas pronghorn
carnivoreslarge and small.17
and some insects (including other grasshopper species)
As expected in semi-arid ecosystems, plant growth
eat more forbs.15 Thus, competition between representa-
in grasslands is strongly influenced by water availability
tives of these two groups is minimized. When the food
(fig. 6.5), though not in a linear manner. As the amount
habits of bison, cattle, pronghorn, and sheep are com-
of water increases, nitrogen can become limiting.18
pared, bison and pronghorn are most different, cattle
Notably, plant growth also is stimulated indirectly by
are similar to bison, and sheep are similar to pronghorn.
a heavy rainfall event that facilitates a pulse of organic
From an evolutionary perspective, bison and pronghorn
matter mineralization, thereby providing more nutri-
would be expected to have the most dis

similar food
ents. Various disturbances also can influence growth, as
preferences because they have coexisted for the longest
described later in this chapter.19
time. Similarly, elk and deer have coexisted for millions
Two other factors that influence plant growth in
of years, and their food preferences are quite different.
grasslands are the rate of water infiltration into the
Elk consume more grass, and deer more twigs and leaves
soil and whether precipitation comes as snow or rain.
of broad-leaved plants (called browse). Of course, the
Rain that does not infiltrate the soil evaporates quickly
diets of all coexisting animals are determined to some
and is unavailable to plants. Therefore, because coarse-
extent by what is available. Considerable overlap may
textured soils have higher infiltration rates, they often
occur in some seasons.
have higher rates of plant growth (the inverse texture

effect; see chapter 3). If more of the precipitation comes
as snow, the meltwater becomes available more slowly,
Grasslands from an Ecosystem Perspective
and there is more time for infiltration before the water

evaporates or is transpired. Generally, the mixed-grass
Energy Flow
prairie receives the majority of its annual precipitation
As in all terrestrial ecosystems, green plants make up
as rain, though snow can accumulate on the leeward
more of the grassland biomass than any other group of
sides of ridges and some taller plants (such as shrubs).
16
Plant growth and other aspects of grassland activity
The next largest component is not the large herbivores,
are often episodic, with pulses of growth, decomposi-
as is often assumed, but rather microscopic organisms,
tion, and nutrient transfers occurring after rain wets the
such as bacteria, fungi, mites, nematodes, and proto
soil. Moreover, because the rainfall is often patchy, the
zoans. Of these organisms, the weight of fungi has been
amount of plant growth and available biomass varies
estimated at four times that of bacteria. Earthworms are
from place to place. In African grasslands and savan-
rare in semi-arid grasslands, but there may be several
nas, patchy rainfall patterns determine the movements
million nematodes and a hundred thousand mites in
of large herbivores and the nomadic people associated
one cubic yard of grassland soil. Surprisingly, the total
with them. Patchy rainfall could also have affected the
weight of all mammals, birds, and insects combined is
movement of people, bison, and other animals in west-
less than 1 percent of the ecosystems living biomass,
ern North America, distributing the grazing pressure
even in grasslands grazed by domestic cattle.
over a large area.20
organisms96 percent, according to some estimates.
89

Fig. 6.5.The annual aboveground net
primary production (NPP) of grassland
ecosystems is correlated with evapotranspiration. NPP is the amount of new
plant biomass produced in a given area
over a known period of time; evapotranspiration is the sum of evaporation
and transpiration and integrates the
effects of temperature and moisture
availability. Each point on this graph is
for a different grassland. The response
of aboveground NPP to changes in
evapotranspiration at a single site would
be somewhat different from the response
illustrated in this graph. Adapted from
Webb et al. (1978).
500
ABOVEGROUND NPP (g/m2/yr)
90
400
300
200
100
100
200
300
400
500
600
EVAPOTRANSPIRATION (mm/yr)
700
800
By what pathways does the solar energy fixed by
zoans, mites, insects, nematodes, and small mammals
photosynthesis flow through the prairie ecosystem? For
that are mostly in the soil. Daniel Milchunas, William
green plants, 6585 percent of this energy is moved
Lauenroth, and their colleagues estimated that, of the
internally from the leaves to roots and rhizomes, and
plant growth that is consumed by herbivores annually,
the plants use about 3040 percent for their own main-
roughly one-third is eaten by cattle, one-third by insects
tenance. A large portion of the remainder is converted
(above- and belowground), and one-third by nematodes,
to stems and leaves that become what is known as stand-
with jackrabbits, rodents, and birds each consuming less
ing dead or detrital biomass at the end of the growing
than 1 percent.
season. Belowground detritus, such as dead roots and
It is interesting to compare the amount of energy
dead nematodes, adds organic matter to the soil. It also
available for human use in a plant-cow-human food
provides the base for an extensive microbial food web,
chain to a plant-human system. In general, people
enables increased infiltration rates, and provides the
obtain more food per acre if plants are consumed instead
nutrients required by plants as the detritis decomposes
of animals, because much of the plant-derived energy
(see fig. 1.6). Of course, fires can burn the aboveground
of the animals has been used for their maintenance or
biomass as well, converting a substantial portion of the
is converted to animal tissues or waste products that
energy to heat and having other significant effects in
people do not eat. Further, the second law of thermo-
some grasslands.21 During some years grasshoppers con-
dynamics dictates that every conversion of energy from
sume a large portion of the aboveground biomass.
one form to anotherfor example, from plant to animal
On average, herbivores consume only 1030 percent
tissue or from animal fat to animal movementleads
of the total annual plant growth (above- and below-
to the production of heat. This principle accounts for
ground), with more than twice as much energy going
the fact that energy flows rather than cycles through
to nematodes and arthropods in the soil than to large
ecosystems. All solar energy fixed by photosynthesis is
herbivores, such as cattle (table 6.2). Not surprisingly,
eventually converted to heat, though it may persist as
some ecologists have concluded that light to moderate
biomass, soil organic matter, or fossil fuels for centuries
levels of grazing have little or no effect on plant species
or more.
composition.22 More than plants and livestock, the food
Human food production is usually greater per unit
web of rangelands is composed of bacteria, fungi, proto
area when plants are eaten rather than meat, but it is also
Table 6.2.Estimated energy flow through a prairie ecosystem on the western Great Plainsa
Ecosystem component
Energy input
Lost by respiration
Tissue production
Production
consumption
SOLAR INPUT
Global radiation
4,155,000
Photosynthetically
active radiation
1,966,000
PRIMARY PRODUCTION
Grossb
21, 882
7,439
Netc
Aboveground
2,163
Belowground
12,280
Subtotal
14,443
HETEROTROPHS
Aboveground
Herbivores
Mammalsd
Macroarthropods
Carnivoresf
105
92
13
0.13
34
23
11
0.32
8.3
Subtotal
7.5
0.8
147
123
25
127
66
61
0.10
Underground
Herbivores
Macroarthropodsg
Nematodes
0.48
50
42
7.9
0.16
20
15
4.6
0.23
12,560
9,632
2,929
72
61
12
Carnivoresh
Detritivores
Microorganismsi
Nematodes
Others
Subtotal
TOTAL
9.2
6.7
2.5
12,838
9,822
3,016
12,986
9,945
3,041
0.23
0.16
0.27
Source: Adapted from Coupland and Van Dyne (1979).

The units are kilojoules per square meter.
Gross primary production (GPP), or total photosynthesis (g/m 2/yr), varies greatly from year to year, depending on water avail-
ability (see fig. 6.4).

c
Net primary production is calculated as GPP less the amount of GPP used by plants for their own maintenance (respiration).
Mammalian herbivores include cattle, pronghorn, rabbits, and ground squirrels; cattle grazing in the study area was light (one
yearling steer or heifer per 2530 acres for 180 days each year).
e
f
Mostly grasshoppers and other insects.
Aboveground carnivores include the coyote, fox, birds of prey, and snakes.
Mostly mites and insect larvae.
h
i
Belowground carnivores include mites.
Bacteria, fungi, and Actinobacteria.
92
true that the only way humans derive food from semi-
ing infiltration and the rate of plant growth. Fire, soil
arid rangelands is from livestock raised for meat and
disturbances, and grazing (above- and belowground) can
milkaside from hunting native animals and gather
stimulate plant growth as well, because they increase the
ing edible native plants. The alternatives are dryland or
rate of mineralization and improve nutrient availabil-
irrigated agriculture, which may lead to soil erosion and
ity.23 A simple mechanism by which mammals contrib-
other environmental problems.
ute to this process is through the deposition of urine and

feces, both of which are easily mineralized.
Nutrient Cycling in Grasslands
Nutrient loss through leaching is an unlikely phenomenon in most western grasslands, because, unlike
For their survival, organisms require nutrients as well as
in forests, rarely is there enough precipitation to cause
suitable temperatures and adequate energy and water.
water percolation below the rooting zone. Wind erosion
For example, calcium is required for plant cell walls;
during droughts or episodes of heavy grazing can lead
phosphorus is important for the storage and release
to some nutrient loss, but only a very small proportion
of energy during metabolism; potassium is needed for
of the nutrients in the soil are lost in this way. Nitro-
the regulation of cell water and manganese for the syn
gen can be lost during inevitable grassland fires, but
thesis of chlorophyll; and nitrogen is an important
again, most of the nitrogen remains unaltered in the
component of amino acids, proteins, and chlorophyll.
soil. Some volatilization of nitrogen has been reported
All nutrients cycle through grassland ecosystems, with
from animal urine and feces, but overall, nitrogen tends
losses occurring primarily during erosion events. Those
to accumulate in western grasslands (fig. 6.6). Nitrogen
losses that do occur are usually offset by nutrient inputs
loss attributable to denitrification bacteria is very small
from rock weathering, rain, snow, the deposition of dust
in mixed-grass prairie, if it occurs at all.
and finer aerosols, and such nutrient-specific processes
Because
nutrients
accumulate,
grassland
soils
as nitrogen fixation. Studies on cycling must consider
can become quite fertile and are excellent for crop
the rates at which nutrients are being added and lost,
productionassuming water is available (by irriga-
as well as the rates and pathways by which they move
tion or otherwise) and temperatures are suitable for
through the ecosystem. All aspects of cycling in grass-
the desired crop. Of course, the harvesting of crops is
lands cannot be considered here, but a few processes
itself a nutrient drain; fertilization is required after a
seem particularly interesting and relevant when think-
few years of cultivation.
ing about the western Great Plains.
Nitrogen fixation by bacteria in the nodules of
As noted, grassland plants typically have leaves and
legumes and other plants is an important long-term
stems high in lignin and cellulose, which are resistant to
nitrogen source for many ecosystems. In shortgrass
decomposition. For this reasonand because the warm,
and mixed-grass prairie, such legumes as locoweed,
moist conditions required for decomposition last for
lupine, milkvetch, scurfpea, and vetch are known to
only a short period each yeardead plant material tends
have nitrogen-fixing bacteria, and there is evidence
to accumulate on the soil surface as well as in the soil.
that a few nonleguminous plants may have the same.24
Among other effects, this surface litter (often referred
Lichens have been identified as nitrogen fixers in some
to as mulch or detritus) can improve infiltration rates.
desert shrublands,25 but so far there is no evidence that
In addition, soil organic matter increases the water and
its done by the most common lichen in Wyoming
nutrient storage capacity of the soil while providing a
grasslandsXanthoparmelia chlorochroa (fig. 6.7).
more erosion-resistant soil structure. Soil organic mat-
Symbiotic nitrogen fixation is commonly thought to
ter is resistant to decomposition, but nematodes, mites,
be the single most important mechanism that provides
and other microbes are able to use it as a food source,
nitrogen for plant growth, in the form of ammonium
directly or indirectly, and in the process nutrients from
and nitrate, but apparently this is not true in western
the organic matter are made available for plant growth.
grasslands, where legume density is low and suitable
Dead microbes also become part of soil organic matter.
environmental conditions for nitrogen fixation occur
In general, many processes create a friable soil, enhanc-
for only a short time each year. Nitrogen fixation by
Grasslands
Fig. 6.6.The amount of nitrogen, measured as grams per square meter, in the
major components (boxes) of a grassland
ecosystem, and the flows between the
components (arrows), measured in grams
per square meter per year. Based on data
in Woodmansee et al. (1981).
NITROGEN
Volatilization
Animals
0.01
Stems
Lvs
3
Roots
7
Soil
solution
2
2.3
24
Detritus
6
1.3
Animals
0.12
Atmospheric
deposition Fixation
0.5
<0.1
Soil surface
19
Soil organic
matter N
333
Microbes
5.2
Outflow (erosion)
0.1
bacteria accounts for only about 7 percent of the annual
This observation has led some managers to conclude
nitrogen inputs on a short-grass prairie south of Chey-
that fertilizing native rangelands might be profitable.
enne. 26 More important sources are rain, snow, dust,
Indeed, more forage of higher protein content could
and the microbial mineralization of soil organic mat-
result. However, several concerns have been identified:
ter, of which there is a large quantity. The sources are
(1) fertilizing only part of a rangeland could result in
diverse, but evidence suggests that nitrogen is a limiting
selective overgrazing as livestock seek out more nutri-
factor for plant growth in grasslands, especially during
tious forage; (2) certain undesirable weeds may be
wetter-than-average growing seasons.27
favored; and (3) despite increases in forage quality and

quantity, fertilization is not yet economical. Changes in
Fig. 6.7. Xanthoparmelia chlorochroa, a common lichen on the

soil surface in Wyoming grasslands and shrublands.
species composition following fertilization are typically

small unless the amounts applied are massive (>150
pounds of nitrogen/acre/year), but there is a chance that
cool-season species could be favored, because they grow
in the spring when nitrogen is more likely to be limiting
than water.28
Although annual nitrogen inputs may be low, this is
of little consequence if losses are also low and if there
is considerable nitrogen in the belowground biomass
and soil organic matter. Francis Clark, an agricultural
research scientist in Colorado, concluded that most
plant needs for nitrogen aboveground are met by the
transfer of stored nitrogen from the root system to the
leaves, with additional nitrogen being made available
by the mineralization of organic matter and microbial
93
94
tissue, plus atmospheric additions. In 1977 Clark wrote,
stored in roots is adequate for new growth (regrowth).
Once a given N atom makes its initial entry into the
In such cases, there is hardly any disturbance at all. 31
blue grama plant, there is a greatly increased probabil-
The effect of natural grassland disturbances is to
ity that the atom will again enter new herbage growth
augment an already high level of spatial variability. As
in each of several following years.29 More than half
described by ecologist Samuel McNaughton in 1985,
(5080 percent) of the nitrogen in the senescing leaves
while writing about African grasslands, there are con-
of western wheatgrass and blue grama is transferred to
tinual fluctuations of rainfall, grazing, nutrient availabil-
belowground perennial plant parts, thereby conserving
ity, and fire. These fluctuations generate pulses of rapid
the nutrient for use when growth resumes.
plant growth in different parts of the landscape, causing
As with plant growth, the accumulation and cycling
a nomadic way of life for large herbivores and humans
of nutrients is not uniform across the grassland land-
alike. Moreover, because of this nomadism, plants in
scape. Many years of natural erosion from ridgetops,
any particular place are not grazed continuously. Today,
with accumulation on lower slopes and in valley bot-
in North America, comparable animal movements over
tomscombined with pronounced differences in the
large areas are often impeded by exurban developments
microclimate and moisture availabilityhave led to
and various land management practices.
more productive soils where sediments are deposited.30

All aspects of nutrient cycling and the flow of energy
and water vary from one place to another. The effect
The Effects of Grazing
of this patchiness on plant and animal abundance is a
As noted, herbivores of any kind form a small portion
major topic in modern ecological research.
of the ecosystem biomass, but they play an important

role in determining the nature of grassland ecosystems.
Disturbance and Succession in Grasslands
Studying the effects of herbivory provides insights on

how coexisting plants and animals survive.
A common theme in ecological research is secondary suc-
On the surface it appears as though any grazing
cession, that is, the changes that occur as an ecosystem
would be disadvantageous to plants; aboveground, the
recovers after a disturbance. All ecosystems are sub-
leaf area for photosynthesis is reduced, and energy that
jected to various kinds of disturbances, whichthough
might be expended for the production of additional
often viewed as unfortunate by casual observershave
leaves is diverted to replacing stems and leaves that have
occurred for millennia and allow for the coexistence of
been eaten. However, some grassland plants seem well
more plant and animal species than otherwise would
adapted to tolerate considerable grazing pressure, and
happen. Also, nutrient cycling and plant growth are
ecologists have hypothesized that grasslands subjected
often more rapid after a disturbance. As described
to light to moderate grazing have higher plant cover,
later in this chapter, the suppression of disturbances to
more growth, and more species than do comparable
which some native species are adapted can lead to unde-
grasslands where herbivory is excluded (fig. 6.8). 32 With
sirable consequences.
a few exceptions, however, such studies have focused
Potential disturbances in grassland ecosystems,
on mammalian herbivores rather than the micro-
aside from plowing for crop production, are drought,
scopic belowground grazers that account for most
fire, periodic heavy grazing by native or domestic her-
energy flow. Scientists can construct fences to exclude
bivores, and the burrowing of small mammals and the
large mammals, but excluding nematodes, mites, and
western harvester ant. Each disturbance has the poten-
other belowground herbivores is necessary to test the
tial to modify the relative abundance of the different
hypothesisa formidable challenge. Not surprisingly,
species. Some disturbances, such as fire or drought, are
experiments on the effects of mammalian herbivory
largely physical phenomena, whereas herbivory and
alone have produced mixed results.
burrowing are biotic. Recovery to predisturbance condi-
How might mammalian herbivory lead to increased
tions often occurs in a few months or a year if damage is
plant growth in grasslands? There are several possible
restricted to aboveground plant parts and if the energy
mechanisms:
Grasslands
growth is increased to some optimal level by light to moderate grazing (see fig. 6.8), after which continued grazing
EFFECT ON NPP
pressure leads to a decline in growth because of an excessive drain on the energy stored in the root system and less
efficient water use (caused by soil compaction that slows
the rate of infiltration). Experiments on western wheat-
Grazing intensity
grass found that, after two days of clipping, the rate of
than for plants without clipped leaves.35 The increased
photosynthesis per unit leaf area was 510 percent higher

rates of photosynthesis did not completely compensate
for the photosynthesis that would have occurred had the
Fig. 6.8.Three hypothesized relationships between large

ungulate grazing intensity and plant growth (expressed as net
primary production; NPP). Curve A, the overcompensation
hypothesis, suggests that grazing stimulates NPP up to some
threshold level, after which the effect is negative; B illustrates
the partial compensation hypothesis, which states that plants
can maintain NPP despite some grazing on some plants; and
C suggests that any amount of grazing by large herbivores
causes a decline in NPP. Adapted from Heitschmidt (1990).
total leaf area remained on the plant, but partial compensation is suggested by the results.
Several investigators, most notably ecologists Elizabeth Painter and Joy Belsky, concluded in 1993 that few
data support the hypothesis that grazing benefits plants,
but the idea has appeal and will continue to guide future
experiments. Grassland ecologist Samuel McNaughton,
one of the early proponents of the compensatory growth
hypothesis, observed that compensatory growth did
not completely replace the vegetation consumed by herbivores and that it is improper to conclude that grazing
is strictly advantageous to the plants.36
1. If some older or senescent leaves and stems are

removed by the herbivore, more light is available for
photosynthesis in younger leaves. This mechanism
may be especially important on moist sites, where
the plants are comparatively tall.
2. If water is limiting at the time of grazing, less leaf
area for transpiration might increase water availability for the remaining leaves, possibly allowing for
more photosynthesis than had occurred before the
grazing started.
3. Herbivores improve nutrient availability for plants
by concentrating certain elements in feces and
urine, which are mineralized more readily than
plant detritus.
4. By eating the stems and leaves, herbivores inadvertently stimulate the plant to produce more cells and
initiate new growth. This process, known as the
removal of apical dominance, triggers the growth of
new leaves and stems that may be more active physiologically than the older plant material that was eaten.33
Grazing also can lead to changes in the abundance

of different kinds of plants. Some species are called in
creasers, because they increase in relative abundance
with grazing pressure, whereas others are classified
as decreasers, because they become less abundant. At
some point, grazing pressure, whether from native or
domestic herbivores, becomes so heavy that weedy
species (invaders) increase in abundance. Range managers have learned to use trends in the relative abundance
of increasers, decreasers, and invaders to assist with
evaluating range conditions. Today, however, managers
think more commonly about the ecosystem services
provided by a grassland, even if the species composition
is different than what might have been considered ideal
40 years ago.
Both bison and cattle undoubtedly grazed western
grasslands heavily at times. 37 Explorers passing through
Fort Laramie observed what they thought was overgrazed rangeland, and one noted that the area was
barren country abounding with prickly pearsa cactus still common today. Another traveler at about the
With such mechanisms in mind, the compensatory
same time reported that 125,000 head of cattle had
growth hypothesis was proposed.34 It states that plant
been driven through the same area, leaving the land
95
96
as devoid of grass as the streets of New York.38 Pre-
quickly throws the advantage in competition to the
sumably the cause was livestock abundance, perhaps in
side of the latter. Because of more water and light,
combination with drought. A hundred years later, the
their growth is greatly increased. They are enabled to
grasslands of this area appeared to have recovered. The
store more food in their propagative organs as well as
rate of recovery may be slow or rapid, depending on the
to produce more seed. The grazed species are corre-
plant species present at the time, weather conditions,
spondingly handicapped in all these respects by the
the history and nature of grazing pressure on the site,
increase of the less palatable species, and the grasses
and livestock management during the time of recovery.
are further weakened by trampling as stock wan-
Some plant ecologists attribute pricklypear cactus
der about in search of food. Soon bare spots appear
abundance to excessive livestock grazing, but Lewis and
that are colonized by weeds or weed-like species.
Clark observed in 1805long before the first domestic
The weeds reproduce vigorously and sooner or later
livestockthe prickly pears are so abundant that we
come to occupy most of the space between the frag-
could scarcely find room to lye [sic].39 Various investi-
ments of the original vegetation. Before this condi-
gators have observed how cactus increases with drought
tion is reached, usually the stock are forced to eat
and decreases following wetter than normal years, pos-
the less palatable species, and these begin to yield to
sibly because of greater susceptibility to insects. There
the competition of annuals. If grazing is sufficiently
is little or no evidence that cactus biomass increases
severe, these, too, may disappear unless they are
because of grazing pressure alone.40
woody, wholly unpalatable, or protected by spines.43
Travelers in Wyoming often wonder whether the

grasslands and shrublands are similar to those observed
prior to the arrival of Europeans. The photographic
record suggests that only a few changes have occurred,
but it is not known whether all the plant species are still
present. Were some plant species driven to extinction
by livestock grazing during the 1880s? Such questions
Lincoln Ellison, who in 1960 provided one of the

early comprehensive reviews of the effects of grazing on
grasslands, recognized that North American rangelands
have been subjected to grazing for millennia. He made
the following observation when comparing fires, plowing, and grazing:
are difficult to answer. Ecologists Richard Mack and
In arid lands, fires may be rare because the fuel is
John Thompson concluded in 1982 that Great Plains
eaten by grazing animals, denudation by plowing
plants are more tolerant of domestic livestock grazing
may be sporadic because of marginal returns and
than are plants in the Great Basin to the west. They
recurrent drought; but overgrazing, although caus-
attributed this difference to a much larger population
ing less complete denudation in any one season, is
of bison on the Great Plains for thousands of years, with
important because it is widespread year after year.44
the result that plants there evolved in the presence of

substantial grazing pressure.41 Long-term studies south
of Cheyenne suggest that changes in annual rainfall
affect plant cover and species composition far more
rapidly than livestock grazing does, and though heavy
grazing may cause declines in the abundance of some
palatable species, moderate grazing (defined as 50 percent removal of plant biomass) did not cause a decline
in range condition.42
The mechanism by which heavy grazing affects
grassland vegetation is generally well known and was
described, perhaps most succinctly, by Nebraska ecologists John Weaver and Frederick Clements in 1938:
In some areas, rangelands probably are still recovering

from the adverse effects of livestock in the late 1800s
and early 1900s, prior to the advent of modern management practices.
Notably, not all grasslands respond to grazing pressure in the same way. Daniel Milchunas, William Lauenroth, and their associates in 1988 compared grasslands
worldwide over a broad range of climatic conditions and
concluded that those in eastern Wyomingin a semiarid environment and with a long history of grazing by
such native ungulates as bisonare tolerant to grazing pressure. In contrast, the taller grasslands of more
humid climates, and grasslands in areas with no history
The more palatable species are eaten down, thus
of long-term grazing, change rather dramatically with
rendering the uneaten ones more conspicuous. This
the introduction of livestock.
Grasslands
Fig. 6.9. Mixed-grass prairie
in Wind Cave National Park,
with bison grazing in a
prairie dog town. Elevation
4,100 feet. Photo by James K.
Detling.
Of course, any grassland can be affected adversely
Serengeti appear to depend on wildebeest for their
if too many animals are fenced into small areas.45 In
survival, because the wildebeest eat the coarser for-
1991 Michael Coughenour described how the nomadic
age, thereby stimulating the growth of the new foliage
movements of large herbivores, triggered by the greater
required by gazelles.48 A month or so after the migra-
availability of preferred food nearby, constitute an im
tory wildebeest move through an area, consuming up
portant mechanism that enables some grasslands to tol-
to 85 percent of the green biomass, regrowth of the
erate grazing, simply because the animals seek food in
plants creates a grazing lawn on which the gazelles
new areas as forage becomes depleted where they are.46
feed. Samuel McNaughton, an expert on this eco
Various livestock management systems, such as short-
system, suggested that the magnificent carnivores in
duration grazing and rest-rotation grazing, are based
African savannas are more numerous because of the
on this principle, but there is still considerable debate
nutrient and energy flows facilitated by the herbivores
over their effectiveness, probably because generaliza-
and their interactions.
tions are difficult. Each ranch operation requires its
North American grasslands have had far fewer large
own management plan based on goals developed with
mammals since the great mammalian extinction about
an understanding of the environmental constraints of
12,000 years ago (see chapter 2), but interesting interac-
the available land.47 Rangeland ecologist David Briske
tions occur here also.49 Most notably, research by James
and his associates concluded that rangelands should be
Detling and his associates in Wind Cave National Park in
managed more to restore ecosystem services, and that
South Dakota have shown that bison, a nomadic herbi-
rotational grazing may not be the best approach when
vore, graze in prairie dog colonies (towns) more often
climate change, invasive species, fragmentation, and
than would be expected by chance (fig. 6.9). The forage
the conservation of biodiversity are concerns.
on the colonies is more nutritious than off, apparently
Research on African grasslands, where 25 or more
because more animal waste is deposited there. Nitrogen
species of large mammalian herbivores occur, has
uptake by the grazed plants is greater, and there are
revealed several interesting interactions. One is the
more young leaves (which tend to be relatively high in
way in which some African herbivores improve food
nitrogen content) in the available forage. In addition,
availability for others. To illustrate, gazelles on the
the regrowth of young shoots is stimulated by grazing.
97
Similarly, Krueger found that pronghorn used prairie
50
dog colonies preferentially, perhaps because of a higher

number of forbs there.
Cause of fire in 1960:

Human
50
Lightning
Such interactions among bison, pronghorn, and

40
burrowing animals surely must have been widespread

prior to the arrival of Europeans. Because of the prairie
dog and other burrowing animals, North American
bison may have been more abundant, along with their
predators (grizzly bears and wolves). One study found
more herbivorous insects on rangeland grazed by cattle
than on rangeland that had not been grazed for a long
period, and another study found more belowground
invertebrates in the soil of prairie dog towns than away
from the towns.51
NUMBER OF FIRES
98
30
20
The effects of large herbivores have attracted the

attention of most grassland scientists, yet the effects of
the abundant belowground herbivores, small as they
10
are, should not be ignored. For example, in an experimental study, a nematocide was applied to kill all the
nematodes in the study area.52 Plant growth declined.
Apparently, microbial grazers improve nutrient availability for vascular plants just as their much larger
aboveground counterparts do.
APR MAY JUN JUL AUG SEP OCT
Fig. 6.10.Number of recorded human- and lightning-caused

fires in the grasslands and shrublands of the Powder River
Basin north of Douglas, Wyoming, in 1960. Adapted from
Komarek (1964).
Fire
Prior to the advent of fire suppression, prairie fires
Most occur during July and August, when fuels are
occurred frequently because of the highly flammable
drier and thunderstorms more frequent. Thunder-
leaves and stems that rapidly accumulate. Without fire,
storms may produce enough rain to extinguish such
and if grazing intensity is low, the standing biomass
fires, but lightning strikes with no or little rain are
is gradually flattened to become part of the mulch on
common. 55
the soil surface. Once fires were ignited, plumes of
Native Americans also started fires, sometimes
smoke would have been visible. The fires would have
to facilitate their hunting.56 In 1985 George Gruell
burned for weeks or months until extinguished by wet
reviewed 145 historical accounts of fire by 44 observers
weather, or until an effective fire break was reached
in the Rocky Mountain region. He concluded that fires
(such as a sufficiently wide river or ridge, or an area of
set by Indians were most common in the lowlands and
inadequate fuel to sustain the fire). Fires might have
that they could have been annual events in some areas
been less common during extended droughts, largely
(although probably the same tract of grassland did not
because of less fuel.
burn in two consecutive years). For much of the Great
53
The source of ignition was usually lightning strikes,
Plains, mean fire intervals were from 2 to 25 years, with
which start fires in grasslands as they do in forests.
the longer intervals in areas of rough topography.57
An annual average of 1025 lightning-caused fires
Grassland fires in central Nebraska occurred at intervals
per 1,000 square miles has been reported for differ-
of 46 years from 1850 to 1900, and every 1530 years
ent parts of the northern Great Plains. 54 Lightning has
in western Nebraska. Wyoming grasslands probably
been documented as the major cause of prairie fires
burned less frequently owing to slower fuel accumula-
in the Powder River Basin north of Douglas (fig.6.10).
tion in a more arid climate (fig. 6.11).
Grasslands
Fig. 6.11. Late-summer fire in the
mixed-grass prairie of the Laramie
Basin. Fires in tallgrass prairies are
less easily extinguished. Photo by Ken
Driese.
Generalizations about the effects of fire on specific
soil to warm up and plant growth to begin earlier in
plants are difficult, but warm-season species seem to
the spring, when water is available. Very little nitrogen
have evolved a higher level of tolerance for burning
is lost to the atmosphere by burning, compared to the
than cool-season species, probably because they are
amount that remains in grassland soils. Furthermore,
more likely to be growing in the drier summer months
the amount lost is replaced in a few years by precipita-
when most fires occurred. Some species increase in
tion (see fig. 6.6).
abundance after fires, not because fires are directly
Because of these influences, total plant growth is
beneficial, but because additional water and nutrients
sometimes higher after a fire, at least in relatively moist
become available following the death of less-tolerant
prairies with tall grasses, such as in central and east-
species, or because a more favorable environment for
ern Nebraska and in eastern Kansas. This observation
their growth is created by the burning of mulch on the
has led some managers to encourage the burning of
soil surface.58
grasslands as a regular practice. However, mixed-grass
Fires also affect nutrient cycling and plant growth.
and shortgrass prairies in the West are more frequently
In some cases plant growth following fire is faster, with
water stressed than tallgrass prairie, a condition that
greener, more palatable nutrient-rich stems and leaves.
could be aggravated by fire because of increased evapo-
Burned grasslands initially appear black and totally dev-
ration from the warmer, blackened surface. 59
astated, but in a few weeks it is common to see herds of
The effects of grazing and fire on plant growth are
bison, pronghorn, and cattle grazing on the regrowth.
interactive. Grazing can be so intense that there is little
Indeed, Indians probably burned grasslands intention-
mulch left on the soil surface, which leads to lower flam-
ally to attract game to smaller areas, where they could
mability. If, however, a large amount of mulch accumu-
be more easily harvested. The animals would have been
lates due to fire suppression or very little grazing, then
lured by more nutritious forage, resulting from the sud-
much of the water in short summer showers can be
den availability of nutrients after accumulated mulch
intercepted by the mulch and evaporated without ben-
was burned. Combustion, like microbial decomposi-
efiting the plants at all. Such adverse impacts may be
tion, converts organic matter to the inorganic nutrients
offset to some extent by the increased trapping of snow
needed for plant growth.
by the accumulated biomass and less evaporation from
Another effect of burning, especially in taller grasslands, is to remove the insulating mulch, allowing the
the shaded soil surface, but this beneficial effect is not

known to happen regularly.
99
100 Plains and Intermountain Basins
Rather than consider the effects of grazing and
and shortgrass prairie could be used for crop produc-
burning separately, ecologists now sometimes recom-
tion in a manner similar to the tallgrass prairies to the
mend a patch-burn grazing system, where the grassland is
east. The consequences were staggering dust storms. A
exposed to both livestock grazing and relatively small,
subsequent drought occurred from 1952 to 1955, but
patchy burns in the same management unit, enhanc-
by that time land management practices had improved
ing the probability of higher biodiversity because of the
and blowing dust was less severe.
more heterogeneous landscape that results. Large herbi-
After the end of an extended drought, the recovery
vores are attracted to the recently burned area because
of mixed-grass prairie may require several years or more,
of more palatable and nutritious forage there, for rea-
with some species reinvading from the shelter of prickly
sons just described. Yet nearby, some birds and small
pear cactus.64 Livestock managers do not usually appreci-
mammals thrive on the tracts of unburned grassland
ate cacti, but spiny plants enable other plants to survive.
that remain.60
Forage availability increases dramatically after cactus are

burned, partly because of greater accessibility without
Drought
The continental climate of the western Great Plains
the cactus spines, but also because plant growth is higher

without competition from the cactus. Still, pricklypear
cactus can help maintain biological diversity.
has great fluctuations in annual precipitation, and its
With time, many of the original species become re
common to have several consecutive years with well-
established following extended drought. Seedling estab-
below-average amounts of rain and snow. Predictably,
lishment is more probable with increased precipitation,
plant growth and rangeland carrying capacity are much
but new shoots also sprout from surviving rhizomes.
lower during droughts. Just as important, plant cover is
Research suggests that some perennial natives survive
reduced and shifts in species composition may occur.
droughts in a dormant or near-dormant state, even for
Some research suggests that shifts in species com-
periods of 57 years. Above-average rainfall for 13years
position commonly attributed to heavy grazing could
changed a shortgrass prairie dominated by blue grama
be the result of drought.61 Common Wyoming plants
to a mixed-grass prairie with needle-and-thread grass,
that sometimes increase in abundance during dry peri-
prairie junegrass, and western wheatgrass as well as
ods include blue grama, Sandberg bluegrass, prickly
blue grama.65 Predictably, livestock forage increased
pear cactus, and threadleaf sedge. Those that decrease
considerably.
include junegrass, little bluestem, needle-and-thread

grass, red threeawn, silver sagebrush, and western
wheatgrass. One of the most successful species during
Grasshoppers
drought is Sandberg bluegrass, but even it declines with
Drought disturbances are frequently accentuated by
extended drought.62 Species that survive or increase do
outbreaks of grasshoppersarguably the most impor-
so partially because of reduced competition from less
tant insect in western grasslands (fig. 6.12).66 The causes
drought-tolerant species.
of outbreaks during drought are not well understood,
The changes associated with drought are most likely
but some research suggests that bacterial and fungal
reduced plant growth, biomass, and cover. Plant cover,
grasshopper diseases are more common under rela-
for example, decreased from 69 to 2 percent during a
tively moist conditions, limiting grasshopper survival.
drought in eastern Colorado, and from 28 to 2 percent
It would be of great practical significance to know that
in southeastern Montana.63 Such drastic changes might
population sizes are determined by the weather, but
be partially the result of grasshopper outbreaks, which
Gordon Watts and his associates cautioned that, even
tend to coincide with drought years.
if there is a climatic correlation with grasshopper abun-
Unfortunately, the drought years of the 1930s (1933
dance, There can be no assertion as to whether it is
1939, especially 1934) followed a 50-year period of grass-
a direct action of weather on the grasshoppers physi-
land plowing, spurred on in some areas by large markets
ology, an indirect action on the food plants, a differ-
for wheat and the erroneous notion that mixed-grass
ential effect on their predators, parasites, and diseases,
Grasslands
Fig. 6.12. More than 100 species of
grasshoppers are found in Wyoming.
Most are adapted for the grasslands and
shrublands in the lowlands, but some
species occur in the alpine tundra. This
one, known as the flambellate grasshopper (Melanoplus occidentalis), is common
in grasslands. It eats mostly forbs but
also grasses, seeds, and dying or dead
arthropods. Photo by Scott Schell.
or a measure of each.67 Favorable conditions for grass
by grasshoppers when their population sizes are high
hopper population outbreaks have also been attributed
over large areas. The costs of the various control mea-
to heavy grazing, whether by bison or livestock.68 Some
sures were justified, in part, by the expectation that they
of the most interesting and complex ecological ques-
would keep the population down for several years. That
tions pertain to the population dynamics of insects.
may be true in some areas, but a recent study concluded
The amount of energy that flows to grasshoppers can
there is little basis for prorating the benefits of con-
be small or large. In northern Colorado they consume
trol beyond the year of treatment and that managing
less than 0.5 percent of the plant growth during years
grasslands in ways that conserve the natural predators
with normal population sizes, but 63 percent of the
of grasshoppers is likely to be a more sound approach,
forage is eaten in outbreak years, with about 20 grass
ecologically and economically.72
hoppers per square yard.
Notably, the apparent veg-
All Wyoming grasshoppers are native and, like bark
etation damage is not a good measure of the amount
beetles in mountain forests, are integral components of
69
actually eaten by grasshoppers, as they cut up to 25
the grassland ecosystem. Curiously, the most notorious
times more forage than they eatleaving considerable
grasshopper in the region, the Rocky Mountain locust,
detritus on the soil surface.
became extinct in the early 1900s. From time to time
In 1941 B. W. Allred described the effects of grass
this locust swarmed over hundreds of square miles,
hoppers in the Powder River Basin during the 1930s
forming virtual clouds that consumed crops and native
drought, writing that there were 50100 per square foot
plants alike. Plagues of flightless Mormon crickets do
and that they destroyed all edible vegetation.70 Much
much the same, as do other kinds of grasshoppers. How-
earlier, in 1864, General Alfred Sully wrote, the only
ever, their swarms are not as large as those of the now-
thing spoken about here [Montana] is the grasshopper.
extinct Rocky Mountain locust.73
They are awful. They actually have eaten holes in my

wagon covers and in the tarpaulins that cover my stores.
Curiously, some areas are not affected even during the
Prairie Dogs and Other Burrowing Animals
worst outbreak years, which illustrates another way that
Burrowing animals can cause prominent changes
disturbances can cause patchiness in landscapes.71
in grasslands and mountain meadows. For example,
For many years, ranchers and scientists have worked
pocket gopher mounds sometimes cover 25 percent of
toward the goal of minimizing economic losses caused
the soil surface.74 Predatory skunks and badgers search-
101
ing for the gophers often amplify the effect with their
digging. This burrowing moves tons of soil every year,
helping to create a more friable soil structure that has
long-term benefits for plant growth. Pocket gopher
burrows provide habitat for the grasshopper mouse, as
well as the beetles and crickets on which the mice feed,
and though the gopher is nocturnal, some individuals
that make the mistake of coming to the surface during daylight provide food for raptors. Notably, pocket
gophers tend to occur near clumps of pricklypear cactus, possibly because the succulent cacti serve as a
source of water. The gophers are able to eat around the
spines.
The most studied burrowing mammal in grasslands
is the prairie dog (fig. 6.13; see also fig. 6.9). Two species
are found in Wyoming, the black-tailed prairie dog on
the Great Plains and the white-tailed prairie dog of the
intermountain basins. One of the most obvious effects
of prairie dogs is the extensive burrowing that characterizes their colonies (towns). As with pocket gophers, large
volumes of soil are moved, improving infiltration, hastening the incorporation of organic matter, facilitating
nutrient cycling, and increasing spatial heterogeneity.75
Generally, prairie dog burrows are dug to depths of 39
feet, deeper than pocket gophers. Along with the pocket
gopher and other burrowing mammals, prairie dogs
enable a higher level of biological diversity by creating a
variety of subterranean habitats, such as for burrowing
Fig. 6.13. Prairie dogs, along with other burrowing animals

(such as Richardsons ground squirrels, badgers, and harvester
ants), alter vegetation structure on the grasslands and shrublands of the intermountain basins and western Great Plains.
Their burrows can be 6 feet deep or more. The black-tailed
prairie dog, shown here, is found at lower elevations on the
western Great Plains; the white-tailed prairie dog is found
in the intermountain basins to the west. Photo by LuRay
Parker/Wyoming Game and Fish Department.
owls, rattlesnakes, grasshopper mice, desert cottontails,

reptiles and amphibians, which in turn encourages the
favorable. However, research suggests that large prairie
raptors that feed on such animals (most commonly the
dog populations are a symptom of heavy livestock graz-
ferruginous hawk). Large colonies also provide habitat
ing. They reason that prairie dog densities increase with
for the restoration of black-footed ferrets, which depend
heavy grazing because predator success is diminished,
on prairie dogs.76 Like the beaver in riparian zones (see
due to less plant cover around the town. Moreover,
chapter 4), prairie dogs influence grassland ecosystems
heavily grazed rangelands commonly have more forbs,
more than expected based on their low biomassthe
which are preferred by the rodents, and by pronghorn,
definition of a keystone species. Because beaver and bur-
too. The interaction of clipping, grazing, and predation
rowing mammals play such a deliberate role in creating
may be less likely in shortgrass prairie, where the veg-
their habitat, they are also known as ecosystem engineers.
etation is already short and visibility good.
In addition to extensive burrowing, the black-tailed
Although prairie dog towns add significantly to the
prairie dogs on the Great Plains keep the surrounding
biological diversity of the grasslands where they occur,
vegetation clipped close to the ground, even if they dont
they also appear to reduce the amount of livestock for-
eat itpresumably to improve their ability to detect
age. Moreover, some of the land is pockmarked with
stalking predators (see chapter 16). This clipping leaves
their burrows, creating a hazard for livestock and horse-
the impression of an overgrazed rangeland, which has
back riders alike. Consequently, like wolves, the rodents
led some to doubt that their effects on the rangeland are
are most often viewed as problem animals where live-
Grasslands
stock production is the primary management objective.
Ground squirrels are another group of small mam-
Often they are shot or poisoned even today, when their
mals that can affect vegetation through herbivory and
population sizes are a small fraction of what they were
burrowing. They do not form towns as prairie dogs do,
in the 1800s. Various studies have described how prairie
but their populations fluctuate between nearly zero and
dogs have little or no adverse effects on livestock weight
a hundred or more per acre. The causes of ground squir-
gain, and that long-term benefits for forage productivity
rel population fluctuations are still not well known.
outweigh what appear to be adverse effects. As noted,
Another common feature of the plains and basins
research has shown that the nutritive value of forage
are the mounds created by harvester ants (Pogono
on prairie dog towns is higher than in much of the rest
myrmex occidentalis).80 These seed-eating ants are some-
of the landscape, apparently because nutrient cycling
times viewed as destructive, because they denude areas
is facilitated by the activities of so many animals in a
around their dome-shaped mounds (fig. 6.14). An esti-
small area. There may be a need to control prairie dog
mated 6 million mounds existed in the Wind River
77
populations in some areas, but doing so has become
Basin in the 1970s. The common reaction to such num-
highly controversial.
bers is to lament the loss of forage for livestock. Yet, like
This issue recently came to a head on the Thun-
other burrowing animals, ants mix and aerate the soil,
der Basin National Grassland, where federal man
and the vegetation surrounding denuded areas is often
agers are trying to increase the size of a black-tailed
more vigorous, most likely because of improved water
prairie dog town so that it can sustain a population of
and nutrient availability caused by ant burrowing and
the en
dangered black-footed ferret. Appropriately, the
defecation. The absence of plant growth on the mound
managers are collaborating with adjacent landowners,
is probably compensated for by the increased growth
encouraging them to allow some prairie dog towns on
nearby (see fig. 6.14).
their property as wellat least in small colonies.78 Some
Disturbance by burrowing mammals and ants also
ranchers have consented to this plan, but only if they
adds patchiness to the grassland mosaicabove- and
can shoot or poison nearby dog towns where they are
belowground. The disturbances lead to continual
not wantedwhether on private or public land. Shoot-
change in the grassland community as species adjust
ing prairie dogs, however, does not make sense to those
to different environmental conditions. The native
who ask: How can poisoning black-tailed prairie dogs
flora and fauna are well adapted to such disturbances;
be justified when their benefits to the ecosystem are so
their diversity is high in large part because of them.
well known and they have been petitioned for protection
As in other ecosystems, however, the effect of people
under the Endangered Species Act?
has often been to modify the frequency of grassland
The dark cloud of sylvatic plaguea bacterial dis-
disturbances or to increase their intensity, resulting
ease carried by fleas spread by rodents and introduced
in a more homogeneous landscape with lower spe-
to North America with rats in the 1800scomplicates
cies diversity. In many areas, the natural disturbances
matters, because it spreads rapidly in prairie dog towns
have been replaced with new ones, like plowing over
and can cause local extinction of the rodents in a year
large areas, for which native species are not adapted.
or two. That has happened numerous times in the past
Thus, it is little wonder that introduced weedy species,
40 years.79 Insecticides have been used to kill the fleas,
originating in Eurasian landscapes with a much longer
with considerable successthough with the usual
history of agriculture, have done so well in American
expense and undesired effects of adding pesticides to
croplands.
the ecosystem. For two reasons, sylvatic plague could

foil the best-designed plan for restoring the ferret in
portions of its native habitat: the ferrets food supply
Invasive Plants
is diminished, and the ferret itself is susceptible to the
Native grasslands have been invaded in some places
disease. Vaccinating the ferrets against the disease has
by a variety of introduced plants that have predictable
shown some promise, but there are too many prairie
effects: crowding out native species, altering habitat
dogs for that to be practical.
and food availability for wildlife and livestock, reduc-
103
Fig. 6.14.The absence of plant growth on harvester ant

mounds probably is compensated for by the increased growth
around the mound. Plant growth on the perimeter would
be favored by better soil aeration caused by the ants, greater

availability of water and nutrients in the affected area, and
less competition from other plants.
ing biological diversity, affecting the amount of carbon
sometimes cattle. Herbicides should be avoided when
stored in the soil and nutrients available for growth
possible, as they kill some native species as well as the
(plant and microbial), increasing soil erosion in some
target plants and affect the soil biota in unknown ways.
areas, and creating the need for expensive weed con-
The challenge seems hopeless when weedy plants domi-
trol practices. Most introduced plants do not become a
nate a large areahence the admonition by weed con-
problem, but a few outcompete native species, because
trol specialists to learn how to identify noxious plants,
they are pre-adapted to their new environment.81
monitor rangelands closely for their appearance, and
Moreover, they are free of their native pathogens and
act promptly to eliminate the plants while their popula-
species-specific herbivores. Grassland invaders on Wyo-
tions are small and isolated.
ming watch-lists include Canada thistle, tumbleweed
Many Wyoming grasslands do not yet have invasive
(Russian thistle), cheatgrass, leafy spurge, and the vari-
weeds. Most likely this is the result of management
ous knapweeds (diffuse, Russian, and spotted). Cheat-
practices that maintain the native plants in good condi-
grass has received the most attention and is discussed
tion, especially during recent decades, when the seeds
in the next chapter.
of exotic plants have become so much more widespread.
Unlike other disturbances, the effects of invasive
Essentially, the native plants are fully occupying the
plants can be long-lasting and restoration is difficult.
spaces into which exotics might invade. Another fac-
Their removal from grasslandswhere that is judged
tor may be that western grasslands are too dry for most
to be importantmay require a combination of con-
invasive plants. To illustrate, Canada thistle and leafy
trol measures, including herbicides; biological con-
spurge tend to occur in the vicinity of riparian zones
trol; hand pulling; fire; changes in land management
or irrigated land rather than in semi-arid grasslands on
practices; and targeted grazing by goats, sheep, and
the upland. Realistically, though, land managers should
Grasslands
Fig. 6.15. Less-palatable plants are more likely to invade

grasslands when plants are weakened by excessive grazing,
such as to the left of this fence. Grass cover has declined and
the abundance of Douglas rabbitbrush, a native shrub, has

increased on the heavily grazed land.
anticipate that the invasive plants of the future could be
evolve to allocate less energy to species-specific chemi-
more drought tolerant.
cal defensive compounds and more energy to growth
82
Invasive species often gain a foothold when distur-
and reproductionan option that would not be adap-
bances reduce the vigor of native species. Some of these
tive for native species.83 Some observations suggest that
are human caused, such as excessive grazing (fig. 6.15),
invasive species are not nearly as abundant during the
plowing, and road construction; but others are natural,
first few years as they are later, after they have adapted
such as drought, fire, and the burrowing of small mam-
to their new environment in one way or another.84 Thus,
mals. A modern predicament is that even natural dis-
just because an introduced plant is localized does not
turbances can create habitat for undesired plants (see
mean that it will remain innocuous. Similarly and on a
chapter 16). Their seeds are widely dispersed, and some-
more positive note, some native species after a period of
times such plants become established before native
time may evolve physiological and morphological adap-
species can.
tations for competing with invasive species.85
An increased understanding of weed ecology and the
Another insight was proposed by Agricultural Re
characteristics that determine the susceptibility of some
search Service scientist Dana Blumenthal and his col-
ecosystems to invasion has led to several noteworthy
laborators.86 They observed that invasive species were
insights. First, once established, invasive plants (and
most successful when slow-growing native plants were
other invasive organisms) adapt further to their envi-
disturbed and when native enemies of the invasives did
ronment, enabling them to become even more aggres-
not exist. The first step to reduce the spread of invasive
sive. This could involve acclimatization as well as a
plants, they reasoned, was to enable the persistence of
genetic response. For example, once in an environment
the native plants, and if possible, find a herbivore that
without parasites and pests, the invasive species may
would feed on the invaders. It is tempting to infer that
105
irrigating or fertilizing the native plants would be good,
tion until a small population was found near Meeteetse
to increase their vigor, but Blumenthal and his associ-
in 1981. Now, through captive breeding, there are sev-
ates found that maintaining water and nutrients at low
eral thousand ferrets, and some are breeding success-
levels reduced the invasibility of the grassland they
fully in the wild.88
studied in northern Colorado.
Careful manipulation
The most conspicuous grazing in the twenty-first
of the time and nature of livestock grazing also would
century is by cattle. But, of the large herbivores, only
restore some level of herbivory, assuming the invasive
the pronghorn now completes its life cycle on the
species were eaten.
grasslandseach animal very likely dying in or near the
87
Where weeds are already well established and the
same county where it was born. In the early 1700s, Native
goal is to restore native species, Blumenthal and his
American hunters would have seen a greater diversity of
associates found that adding carbon to the soil, in the
large mammals, and very likely they would have heard
form of sawdust, reduced the availability of nitrogen.
the barking of prairie dogs almost every day during the
This causes a decline in soil fertility, which the native
summer. Raptors and other predators would have been
plants can tolerate better than the invasive species. In
more common with such a large prey base. There would
their experiment, adding nitrogen favored the intro-
have been many large animal carcasses, providing food
duced species. Thus, disturbances that make nutrients
for numerous kinds of scavengers. Nathaniel Wyeth
more available should be minimized while this kind
wrote in 1832 about an abundance of buffalo along the
of restoration work is in progress. Where feasible, her-
Sweetwater River valley, and in 1857long before the
bicides were integrated with this work in a way that
introduction of sheep and cattleWilliam A. Carter
favored the native species, helping them become estab-
wrote about seeing thousands of dead animals.89 Ravens,
lished. This research was motivated by the observation
vultures, coyotes, and other scavengers would have fed
that the primary expense of controlling invasives, by
on the carcasses. Now and then a grizzly bear or pack
whatever means, results from the fact that it has to
of wolves would have chased them away. Seeing a bad-
be done time and time again. If native species can be
ger would be common. The clickity-clack of grasshopper
restored and managed properly, they persist year after
wings would have been frequent in the summer, but with
year, preventing or slowing plant invasion well into the
no spotlights, the nocturnal swift fox and black-footed
futurean ecosystem service powered by solar energy.
ferret would have gone unnoticed most of the time.
Native plant growth and biomass may be less than that
Today, a grassland ecosystem like the one first
of the invasive plants, but the expense of maintain-
encountered by EuroAmericans in the early 1800s, and
ing the grassland is greatly diminished. Moreover, the
the first human immigrants some 15,000 years earlier,
adverse effects of the invading species are reduced.
does not exist. Most of the native plants are there, and
some of the small native animals can be found if one
Challenges for the Future
looks long enough. Grasshoppers still exist, some years

more than others, and occasionally a prairie dog town
Traveling across North America, its rare to find native
can be found. Coyotes will be heard. With luck, a swift
grasslands that extend to the horizon as they once did.
fox will be visible as it darts across the road at night.
Most have been plowed for cropland or converted to
However, off in the distance, the large animals are most
pastures of introduced plants. Prairie dog towns can
likely cattle rather than bison, deer, elk, pronghorn, or
be seen along the highways in the western states, but
large predatorsor the camels, mammoths, saber-tooth
the population of these animals is probably less than
tigers, and other extinct species of earlier times.
10percent of what it was in the 1800s. Many think that
Considering the relatively depauperate nature of
the black-tailed prairie dog should be listed federally
grasslands nowat least abovegroundthere is reason
as an endangered species, especially now that sylvatic
to promote the establishment of a grassland national
plague has been introduced to America and frequently
park in an area large enough to restore the diversity that
kills entire colonies. Because of the decline in prairie
once existed there. Neighbors would justifiably object
dogs, the black-footed ferret was on the brink of extinc-
to the re-establishment of wolves and grizzly bears on
Grasslands
the Great Plains, but all the other species could be re
Unfortunately, those circumstances are chang-
introduced. All dead animals would remain for scaven-
ing, leading to concerns that the values of grasslands
gers and nutrient cycling. The prairie dog towns could
will be lost. Plowing continues, often in concert with
grow to be large enough to sustain the black-footed fer-
shifts in crop prices and by landowners desiring to take
ret, assuming that the sylvatic plague can be controlled.
advantage of government subsidies. The cultivation of
Fires would be started or allowed to burn as judged
grassland soils is now more often sustainable than pre-
appropriate, and the area would be large enough that
viously, thanks to minimum till or no-till agriculture,
unintentional wildfires could burn without burning
but the grassland ecosystem is thoroughly disrupted in
the whole park. Protection for the neighbors would be
the process. Some grasslands have been restored, either
required. Some bison would be harvested to simulate the
by abandoning old fields and allowing secondary suc-
effects of the wolves and grizzly bears that are absent.
cession to occur, or through the planting of perennial
Traveling into this imagined park from any direction,
native species, such as farmers and ranchers are able to
the aboveground natural diversity of native grasslands
do with funding from the U.S. Department of Agricul-
would be immediately apparent.
tures Conservation Reserve Program. Wildlife invari-
Fortunately, some prairies in the semi-arid West
ably benefits, but decades if not centuries are required
have not been plowed, and they have been managed
before the original biological diversity is restoredif
for livestock production in a way that appears to be
thats possible at all.94 Such concerns cannot preclude
sustainable. The plants are mostly native species, the
farmers from making a living from their land, but the
all-important belowground food webs probably are still
best soils most likely have already been cultivated. With
intact, and the natural diversity aboveground is high if
so much of the grasslands already disrupted, there is
not complete. Prairie dogs and their associated species
reason to be concerned about further losses.
are present.90 Some intact grasslands are private; others
The intact grassland ecosystem requires an area that
are on public land, such as in Wind Cave National Park
is better thought of as a diverse landscape with ridges
and Badlands National Park in western South Dakota,
and riparian zones here and therenot a homogeneous
and a place known simply as Grassland National Park in
tract of native plants. Ranches in western states often
Saskatchewan.91 In Wyoming, the federal managers of
preserve such mosaics, along with the open space that
the Thunder Basin National Grassland are working with
is valued by many groups. But the challenges of keep-
adjacent landowners and others to develop a plan for
ing such large areas intact are becoming greater. The
profitable livestock grazing even while the prairie dog
primary threats are plowing, the widespread effects of
population is restored to a level that is sufficient for the
oil and gas development, new wind farms and trans
reintroduction of black-footed ferrets. More collabora-
mission lines, and the desire of many newcomers to
tions of this nature are needed.92
build their homes in the middle of that open space. The
Working with interested landowners, various land
services provided by grasslands are reduced still further
trusts are finding ways to conserve the native grasslands
with a diminished landscape and the introductions of
that remain in a way that provides economic benefits to
exotic plants and animals that invariably occursome
the landowners and maintains biological diversity. Such
of which could become as problematic as sylvatic plague
projects may not have all the components of the original
or cheatgrass (see chapter 7). Various conservation
ecosystem, but they provide a means of conserving some
organizations (such as The Nature Conservancy and
of them. This approach is highly valued by conservation
other nongovernmental groups) are helping to main-
biologists. Indeed, it is the most feasible one over much
tain intact grasslands where landowners and agencies
of the West. Traditional approaches to livestock manage-
have common incentives for doing soeconomic or
ment are being adapted to achieve the goals of sustain-
otherwise.
ability and conservation. Notably, the circumstances
Another challenge is presented by climate change.
that led to livestock production on large ranches in the
As described in chapter 3, this has been occurring for
West have also, inadvertently, conserved open spaces
millennia, but the socioeconomic impacts now could
and much of the biodiversitythough not all of it.93
be substantial. Crop production could decline because
107
of increased plant water stress and possibly less water
the season, such changes caused differences in nutrient
for irrigation. Forage could be reduced for large mam-
cycling and forage quality on the plots.97 For example,
mals of any kind. Further changes could occur if warm-
fringed sagewort, a small native shrub that is avoided
season grasses or shrubs become more common, which
by livestock, had much greater biomass than in the
have lower nutritive value than do cool-season grasses.
control plots; and there is evidence that less nutritious
For such reasons, the entire food web could change,
warm-season grasses will also become more common.98
belowground and above.
Research is under way to determine whether invasive
95
Notably, however, research in mixed-grass prairie
plants will be favored by elevated carbon dioxide as well
near Cheyenne indicates that increased atmospheric
as the nitrogen added by present-day levels of atmo-
carbon dioxide, while sometimes increasing the rate
spheric deposition.
of photosynthesis, has the additional effect of reduc-
Warming may also increase the rate of decomposi-
ing transpiration in some prairie plants. More water is
tion of soil organic matter, at least during times when
left in the soil for a longer time. A team of Colorado
the soil is moist, such as in the spring and on warm win-
and Wyoming scientists led by Agricultural Research
ter days. The overall effect could be a gradual decline
Service ecologist Jack Morgan conducted this research,
in the carbon stored in the soil with the concomitant
known as the Prairie Heating and Carbon Enhance-
release of more carbon dioxide to the atmosphere,
ment Experiment (PHACE). In replicated plots they
potentially aggravating global warming at a time when
increased levels of atmospheric carbon dioxide while
carbon sequestration is an important management
simultaneously warming the plant canopy and soil.
objective. Moreover, soils also could become more erod-
With higher amounts of carbon dioxide, the soil did
ible if plant cover declines; subtle changes could reduce
not dry as rapidly as in control plots, and plant water-
the capacity of grasslands to sustain grazing by live-
use efficiency was enhanced. This result is counter
stock. Animal scientists are already considering how the
intuitive, because if carbon emissions cause warming,
livestock industry might adapt its practices.
there should be higher rates of evapotranspiration,

causing more frequent periods of plant water stress.
Thus, the results of carbon dioxide enrichment and
However, as Morgan and his associates concluded, in
climate warming are more complicated than poten-
a warmer, CO2 -enriched world, both soil water content
tially causing higher rates of photosynthesis. If water
and productivity in semi-arid grasslands may be higher
stress develops quickly, or continues during extended
than previously expected. They added that, elevated
droughts, there will be no enhanced photosynthesis of
CO2 can completely reverse the desiccating effects of
the mixed-grass prairie, at least with the native plants
moderate warming.
that dominate the area now. The climate of the future
96
The PHACE study documented other changes as
may favor novel combinations of more drought-tolerant
well, including a rise in the importance of fungi, com-
species. Could this possibility be in the future for all
pared to bacteria, and an increase in the carbon-to-
kinds of ecosystems in the region? Will the ecosystems
nitrogen ratio of the soil. Along with more water later in
provide the services that many would like to have?
Chapter 7
Sagebrush
The iconic big sagebrush, Artemisia tridentata, dominates
Mention of sagebrush brings to mind wide-open
the most widespread ecosystem in Wyoming. Growing
spaces with few trees, few people, and scattered herds of
mostly west of the eastern grasslands, this shrub occurs
pronghorn and feral horses (fig. 7.1).2 Some would like
throughout the intermountain basins and covers about
to make the land more productive, whether by burn-
one-third of the state. Thats obvious to travelers. Not
ing to remove the shrubs or by cultivation. Indeed, the
so clear is that big sagebrush is but one of nearly a dozen
presence of knee-high big sagebrush is thought to be an
woody species of Artemisia that are commonly found
indicator of relatively deep soils that are suitable for crop
here. Others include alkali sagebrush, black sagebrush,
production. The potential for farming, however, is lim-
low sagebrush, silver sagebrush, and threetip sagebrush.
ited by scarce water and a cool, short growing season.
Fig. 7.1. Wyoming big sagebrush

steppe in central Wyoming,
southeast of Riverton. Associated
plants are junegrass, Sandberg
bluegrass, western wheatgrass,
blue grama, fringed sagewort,
phlox, and numerous other species. Scattered limber pine and
small groves of Douglas-fir occur
in the vicinity of Beaver Rim.
109

Fig. 7.2. Shrublands dominated by
Wyoming big sagebrush form a mosaic
with saltbush desert shrubland in this
area west of Rawlins. Big sagebrush is
restricted to ravines, where drifting
snow accumulates and provides more
water for plant growth. The adjacent
shrubland is dominated by birdfoot
sagewort, Gardners saltbush, western
wheatgrass, and winterfat. This area is
one of the driest in the state (see fig.
3.4). Elevation 6,750 feet.
The sagebrush-dominated ecosystem is far from uni-
ricegrass, pricklypear cactus, scarlet globemallow,
form. In some areas patches of sagebrush intermingle
horsebrush, and rabbitbrush. The common animals are
with patches of grassland, or with patches of small
often the same as well, including pronghorn, jackrabbit,
sagebrush plants intermingled with taller shrubs. This
badger, coyote, Wyoming ground squirrel, grasshopper
mosaic can sometimes be attributed to fires, which burn
and deer mice, several kinds of grasshoppers and spar-
unevenly across the landscape, creating patches of dif-
rows, and many others (see table 6.1).
ferent ages. The mosaic also is influenced by differences
Climate is an important factor in determining sage-
in soils and topography. Shallow soils and windswept
brush distribution and growth, as winter precipitation
ridges commonly have black sagebrush and small cush-
especially snowfallcontributes a larger proportion of
ion plants, whereas deeper, saline soils have birdsfoot
annual precipitation in the intermountain basins where
sage, Gardners saltbush, and greasewood. Soils along
big sagebrush is common (see chapter 3). 3 In contrast,
ephemeral creeks often support silver sagebrush. The
summer rains provide most of the water on the grass-
transitions are sometimes abrupt, due to abrupt changes
lands of the eastern plains. Unlike rainfall in the sum-
in moisture availability and soil conditions.
mer, which evaporates quickly, snow is more likely to
Many of the plants associated with sagebrush are
accumulate and when it melts, the water infiltrates
also found in grasslands, such as western wheatgrass,
to greater depths. This provides the more dependable
Sandberg bluegrass, needle-and-thread grass, Indian
water supply required by big sagebrush. In the Lara-
Sagebrush
Fig. 7.3. Mixed-grass prairie intermingles with Wyoming big sagebrush
steppe in the Powder River Basin. In this
area, the sagebrush cannot grow in the
ravines, probably because this habitat
is too wet for too long during the year.
Some of the shrubs on the edge of the
ravine are plains silver sagebrush, which
is more tolerant of wet soils. Elevation
4,800 feet.
mie and Shirley basins where there is less snowfall, or
comparatively moist ravines and valleys. It grows to a
toward the eastern fringes of its range, big sagebrush is
height of 6 feet or more, much taller than the other two
found almost entirely where snow drifting occurs, such
varieties. In Wyoming it is most often found in the Big-
as on the lee sides of ridges or in ravines (fig. 7.2). Nota-
horn Basin and the southwestern quarter of the state.
bly, big sagebrush is absent from very wet areas (fig. 7.3).
Veritable woodlands of head-high basin big sagebrush
Big sagebrush can be observed from the western
occur on stabilized dunes and other relatively moist
Dakotas all the way to the Cascade Mountains of Oregon
environments.5 Wyoming big sagebrush, much shorter
and Washington, and from Canada south to Arizona
than basin big sagebrush, is the most common shrub in
and New Mexico. It grows in habitats ranging from Utah
Wyoming. Normally less than 1.5 feet tall, it occupies
deserts at about 3,000 feet elevation to mountain shrub-
the drier uplands. The third subspecies, mountain big
lands at about 9,900 feet or higher. Surviving over such
sagebrush, is most common near woodlands of aspen
a broad range of environmental conditions requires
and low-elevation conifers but sometimes occurs in
considerable genetic variation. Three varieties with dif-
foothill ravines adjacent to shrublands with basin and
ferent adaptations to water stress are commonly recog-
Wyoming big sagebrush.6 Two of the subspecies can be
nized: basin big sagebrush, Wyoming big sagebrush, and
found growing together where environmental condi-
mountain big sagebrush.4 Basin big sagebrush is found
tions are intermediate. All three are known to hybridize,
at lower elevations (fig. 7.4) and is usually restricted to
which can lead to identification difficulties in the field.
111
ELEVATION (feet)
10000
Mtn silver
sagebrush
Alpine sagewort
Mtn big
sagebrush
9000
Rawlins, black sagebrush is found on windswept ridges

or slopes with shallow soils and very little snow. On
lower slopes with more snow accumulation and deeper
Threetip
sagebrush
soils (but minimal salt accumulation), Wyoming big

sagebrush occurs. Mountain big sagebrush grows in
moist ravines with deep snow. Desert shrublands (see
next chapter) occur in dry basins, where snow depth is
Wyoming big
sagebrush
8000
accumulation is exceptionally deep, only grasses and
Black
sagebrush
or
Bud
sagewort
or
Birdfoot
sagewort
7000
Basin
big
6000 sagebrush
or
Silver
sagebrush
5000
Ravine
North
or
or
floodeast
plain
slope
moderate and salt accumulation is high.9 Where snow
South
or
west
slope
Ridge
or
dry
basin
MOISTURE GRADIENT
Fig. 7.4. Approximate distribution of different species of
Artemisia (sagebrush and sagewort) in relation to elevation
and topographic position.
forbs can survive (fig. 7.8). Clearly, sagebrush distribution in the landscape depends on soil moisture, salinity, depth, and texture, all of which vary over short
distances because of wind and topography.
Wyoming big sagebrush sometimes occurs in ovalshaped patches 1050 feet across. Commonly referred
to as sagebrush islands (fig. 7.9), some are simply patches
of tall sagebrush surrounded by shorter sagebrush.
Most are patches of sagebrush in a matrix of grassland
or desert shrubland. One type of island forms where
small silt dunes or sand dunes develop on the lee side
of taller shrubs. These dunes could have been initiated
by a period of wind erosion, possibly after heavy grazing by bison or livestock during a dry period. Where
an occasional shrub managed to survive or a new one
became established, sand and silt accumulated, creat-
Silver sagebrush, black sagebrush, and low sagebrush
ing conditions favorable for the growth and establish-
are also common. Silver sagebrush has two subspecies,
ment of other shrubs. With time, a larger number of
plains silver sagebrushfound on relatively moist sites
taller shrubs increased the deposition of windblown
at low elevations, especially east of the Continental
particles, including snow, resulting in the develop-
Divideand mountain silver sagebrush, which is com-
ment of small dunes that are 6 feet deep or more. Such
mon in moist meadows in the mountains throughout
dunes are known as coppice dunes and typically have
the region (figs. 7.5 and 7.6). Unlike most woody species
spiny hopsage, rubber rabbitbrush, green rabbitbrush,
of Artemisia, silver sagebrush is capable of sprouting from
and sometimes basin wildrye in addition to Wyoming
the root crown following fire or other disturbances.7
big sagebrush. Spiny hopsage is an indicator of sandy,
Black sagebrush is usually less than 18 inches tall and is
relatively moist soils in the semi-arid intermountain
found on relatively dry soils associated with the slopes
basins (due to the inverse texture effect, explained in
of escarpments or where a shallow hardpan of calcium
chapter 3). A desert pavement is sometimes found sur-
carbonate (caliche) develops. Similarly, low sagebrush,
rounding the islands, often with scattered plants of bud
which is usually less than 10 inches tall, forms patches
sagewort.10
on shallow soils interspersed with patches of big sage-
Elsewhere, the sagebrush islands are not associated
brush, such as in the lowlands of Jackson Hole and on
with windblown material and are less easily explained.
the Sweetwater Plateau. In the Bighorn Basin and other
In such places, the islands typically have numerous
areas of low precipitation, shallow soils commonly have
small mammal burrows, which can disrupt shallow
grasslands (fig. 7.7).8
hardpans or create soil conditions in other ways that
Another factor influencing the shrubland mosaic is
favor infiltration of water and taller shrub growth. Per-
snow drifting. For example, in the foothills south of
haps the animals are attracted by the shade provided by
Fig. 7.5. Approximate distribution maps for shrublands dominated by eight different
species or subspecies of sagebrush (Artemisia) in Wyoming. Adapted from Beetle and
Johnson (1982).
ELEVATION (feet)
5500
5000
4500
4000
3500
Fig. 7.6. Big sagebrush and silver sagebrush are easily distinguished by examining leaf shape. The longer leaves on big
sagebrush are ephemeral, dropping off early in the summer,
whereas the shorter leaves remain on the shrub for a full year.
Both species are evergreen. Drawing by Judy Knight.
10
20
30
SOIL DEPTH (inches)
40
Fig. 7.7.The distribution of grasslands (1), black sagebrush

steppe (l), and big sagebrush steppe ( ) in relation to soil
depth and elevation. Note how big sagebrush occurs on
deeper soils. Data from the Bighorn Basin.
Fig. 7.8. Wyoming big sagebrush in

the lowlands of northern Yellowstone
National Park. Nivation hollows form
where drifted snow persists on leeward
slopes until late June, creating a
meadow, because big sagebrush cannot
survive where the soils are wet during
much of the early summer. Elevation
7,400 feet.
Sagebrush
Other sagebrush islands are associated with mima

mounds (see chapter 9), or with pockets of deeper soil
in areas dominated by threetip sagebrush. As is usually the case, a vegetation pattern can have several
explanations.
Sagebrush Adaptations
Plant distribution is determined more by the tolerances of delicate seedlings than by the characteristics
of adult plants. Seedlings of big sagebrush are commonly observed in some years, almost always when
the early summer has been relatively cool and wet.12
Lower temperatures minimize the rate of water loss
from the drought-sensitive seedlings, providing more
time for roots to grow down to a dependable water
supply. Though the roots may not grow more than
an inch or two in the first year, the presence of deep
soil water contributes to a relatively moist surface soil
environ
ment in three ways: (1) capillary movement
from below, (2) condensation in the surface soil of
water vapor that emanates from the deeper soil water,
Fig. 7.9. Wyoming big sagebrush sometimes grows in patches,
such as in this area east of Rock Springs. The patches can be
small or large, oval or irregular, and often are associated with
coppice dunes. Elevation 7,000 feet.
and (3) hydraulic redistribution from deep to shallow

soil via the root system of other plants (see below for
explanation). Once established, the adult plants tolerate the occasional years when little precipitation falls
during the winter. Surface roots use surface moisture
the shrubs. But how did the shrubs become established
early in the growing season, but other roots grow
in the first place? One explanation is that the patch
down to 6feet or more and make use of deep soil mois-
began with a single shrub that survived by chance,
ture aftersurface soils are dry. The root system extends
perhaps adjacent to a burrow. Its presence caused snow
laterally to a distance of about 5 feet, well beyond the
drifting on the lee side of the shrub, creating condi-
shrubs canopy.
tions favorable for the seedlings of additional shrubs
Significant soil water recharge at depth usually
perhaps from seed produced by the original plant. In
occurs only once a year, during the snowmelt period,
addition to snow, the taller shrubs could also lead to
but, as noted, surface soil dampening during a hot, dry
the accumulation of blowing organic matter and soil
summer can happen even without rainfall through
particles. This accumulation, combined with nutrients
the novel process of hydraulic redistribution.13 First
brought to the surface by the animals and litterfall from
described by ecologists James Richards and Martyn
the shrubs, creates islands of fertility. Such islands are
Caldwell in 1987, this process is now known to occur
found where water and other resources are inadequate
in numerous plants that have both deep and shallow
11
to support a uniform cover of sagebrush. Neil West, an
roots. If the deep roots are in moist soil and the shal-
ecologist at Utah State University, suggested in 1988
low ones in dry soil, as often happens in the summer
that, for sagebrush-dominated vegetation in the Great
where big sagebrush grows, then water is transported
Basin of Utah, any activity that destroys sagebrush
by the deep roots up to the shallow roots, where it is
islands reduces the productivity of the ecosystem as a
released into the soil. Initially the process was referred
whole. The same probably is true for Wyoming.
to as hydraulic lift (from deep to shallow soil), but sub-
115
sequent research found that water can be moved up or
The carbohydrates produced by photosynthesis are
down (always from wet to dry soil).14 This process pro-
distributed to all plant parts where energy is needed. In
vides more water in the summer for plants that have
big sagebrush, a large proportion of these energy-rich
only shallow roots, thereby extending the growing
compounds is stored in the twigs. Thus, both the twigs
season and increasing the overall productivity of the
and the leaves are a good source of food for herbivores.
sagebrush ecosystem. Hydraulic redistribution also can
Furthermore, the shrubs are tall enough to extend above
prolong microbial activity in the surface soil, which
the snow during most winters. Little wonder, then, that
facilitates decomposition and increases the supply of
historically the shrubs have been subjected to consid-
nutrients for the plants.
erable browsing pressure and that they have evolved
Several mechanisms allow the efficient use of water
a mechanism to minimize this herbivory, namely, the
by big sagebrush. The stomata close rapidly as water
terpenes that give sagebrush its characteristic aroma.
stress develops during the day, limiting transpiration
The terpenes apparently cause sufficient indigestion to
until the next morning, when they are again open.15
minimize the amount eaten by some animals. Without
The plant re-equilibrates at night, when water uptake
this adaptation, the shrub may not have survived the
exceeds losses. Stomatal closure limits photosynthesis
intense browsing that might have resulted from large
as well, but prolonging the growing season by con-
herbivore populations, especially those limited most
serving water seems to be adaptive. In addition, the
directly by winter food supply.18
plant is evergreen, retaining most of its leaves for one
Considering the genetic variation that exists in popu-
year. Moreover, acclimation to changing temperatures
lations of widespread plants, it should not be surprising
occurs readily, even down to freezing. Big sagebrush is
that different species of sagebrush, and even different
capable of photosynthesis in the early spring and even
individuals of the same species, vary in the amount of
during relatively warm winter days.16 In general, ever-
terpenes they produce. Herbivores have been observed
green plants fix carbon whenever water and tempera-
to preferentially browse on those plants with the low-
ture conditions are within tolerance ranges. With no
est terpene concentrations. Some animals tolerate the
time lost for the production of new leaves, the growing
terpenes. A good example is the pronghorn, which
season is longer.
eats sagebrush all year long (the shrub provides 7095
Water is most readily available in the spring, when
percent of its food in the winter). Other herbivores eat
big sagebrush has the largest leaf area for photosyn-
sagebrush as well, such as sage-grouseespecially when
thesis. Two leaf types are produced: ephemeral leaves
other foods are not available.19
in early spring, which fall as water stress develops
Big sagebrush, unlike most of its neighboring plants,
during the summer, and overwintering leaves that
lacks the capacity to sprout from roots or the root crown.
remain on the plant until the following spring (see fig.
Consequently, longevity and seed production are espe-
7.6).17 Essentially, the plant is evergreen, but the loss
cially important for its persistence. Some plants live for
of ephemeral leaves by early summer allows for some
100 years or more, though 4050 years is more com-
reduction in total leaf area as water becomes more
mon. 20 The shrub produces annual rings in the wood
limiting, providing another mechanism for conserv-
similar to temperate zone trees. With regard to sexual
ing water and extending the length of the growing
reproduction, big sagebrush produces thousands of tiny
season. Shadscale, another shrub in the basins, also
seeds each year, some of which remain viable for up to
produces both ephemeral and persistent leaves. The
4 years.21 Like many other plants, germination occurs
leaves of many species of sagebrush and other xero-
in late winter or early spring, when moisture is avail-
phytic plants often have a pale, silvery-green color,
able and the temperature is warm enough for growth.
attributable to dense, minute silvery hairs on the sur-
No cold period is required to break seed dormancy,
face. This color reflects light more readily than dark
although some investigators have found that this treat-
green does, which reduces water loss because the
ment causes higher germination rates in mountain big
leaves remain cooler.
sagebrush. Even very low salt concentrations can inhibit
Sagebrush
big sagebrush germination,22 an observation that may

explain why the plant is less common on saline soils.
A high percentage of big sagebrush seeds are viable
and germinate readily (80 percent or more). However,
seedlings survive only during favorable years and typically following a disturbance that reduces the competitive ability of neighboring plants.23 The age distribution
of a sagebrush population is therefore discontinuous; that
is, only a few age classes are represented.24 Conditions for
high growth rates early in the summer are essential, so
that the seedlings can develop a root system adequate to
cope with late-summer droughts. Jerry Schuman, mineland reclamation specialist, found that sowing too much
grass seed can lead to a level of competition from grasses
that reduces sagebrush seedling survival.25
The Sagebrush Ecosystem

Compared to grasslands, the distinguishing features
of sagebrush ecosystems are (1) the presence of an aromatic shrub with both shallow and deep roots and (2)a
large portion of the annual precipitation occurring in
the winter. Otherwise, grasslands and sagebrush steppe
are similar:
plant growth (net primary productivity) is limited
Fig. 7.10. Snow frequently accumulates on the lee side of

shrubs, which probably increases the amount of water that
infiltrates the soil.
by water availability and the length of the growing

season;
most of the biomass and half or more of the herbivory is belowground;
potential evapotranspiration usually is greater than
the annual precipitation;
nutrients are rarely leached beyond the rooting zone;
fire, drought, and burrowing animals are common
disturbances;
plant and animal species have evolved to minimize
competition by using different resources and by us
ing them at different times of the year; and
coexisting herbivores, above- and belowground,
interact in complex ways to affect nutrient cycling
and plant growth.
Hydrology and Plant Growth

Drifted snow on the lee side of shrubs (fig. 7.10) can be a
significant supplement to soil water recharge, especially
on comparatively level sites, where accumulation is not
affected by topography.26 Scientists and land managers
have learned that, without big sagebrush, relatively
more water and nutrients are available for the growth
of more palatable grasses and forbs, but overall, there
is probably less total water available and less total plant
growth than when the shrubs are present. Sagebrush
ecosystems have aboveground plant growth totaling
80250 g/m2/year, which is higher than that for many
mixed-grass prairies in Wyoming.27
Plant water availability may also be influenced by
This section focuses on differences caused by the pres-
the blackbody effect of shrubs that are above the snow
ence of an evergreen, deep-rooted shrub that is not able
in winter.28 Heat is absorbed by the shrub tops, caus-
to sproutand what happens when it dies because of
ing snowmelt around the shrub more often than would
burning or herbicides.
occur otherwise and creating a depression or well into
117
which more snow drifts during the next storm. In this
if a fire occurs. Big sagebrushdominated ecosystems
way, the shrubs facilitate the percolation of meltwater
burn from time to time, and because the sagebrush
deeper into the snowpack or possibly into the soil (if its
itself cannot sprout, nutrient and water resources are
not frozen), reducing the amount that is lost to subli-
made available to grasses and forbs for 10 years or
mation while creating the potential for additional snow
more. As herbaceous plants become more abundant,
accumulation as the snow melts around the shrub tops.
the carrying capacity for bison and cattle increases. Per-
The depressions around the shrubs can be created and
haps the optimal habitat for some animals is a mosaic
filled several times during a winter, augmenting mois-
of old-growth sagebrush intermingled with recently
ture input above that to be expected if the shrubs were
burned sagebrush. Before the arrival of EuroAmericans,
not there. This phenomenon could be especially impor-
the steppes dominated by sagebrush might have been
tant where the vegetation is more or less uniformly
more varied where periodic fires were possible, creat-
covered with snow except for the tops of shrubsa
ing patches of grassland and young sagebrush inter-
common occurrence.
mingled with old-growth sagebrush. Today, sagebrush
As noted, big sagebrush has roots that are deeper
cover might be more uniform and older in some places
than those of most grasses and forbs. As a result, more
than previously, the result of fire suppression and more
water is used by plants when the shrub is present. If the
extensive livestock grazing.32 However, elsewhere, sage-
shrubs are removed by fire or other means, total water
brush landscapes have been heavily fragmented, as dis-
consumption by the plants (transpiration) is less, per-
cussed later. Because of threatened species that depend
haps by 15 percent,
on sagebrush, the practice of prescribed burning in this
29
30
but total plant growth probably is
lower as wellas would be expected based on the cor-
ecosystem may not be advisable in some areas. 33
relation of plant growth with evapotranspiration in arid
Total annual plant growth is enhanced by the pres-
climates (see fig. 6.5). Closer to the mountains, where
ence of big sagebrush, in part because of its deeper root
snowfall is higher, less transpiration may allow for some
system but also because its green leaves persist through-
runoff as streamflow.
out the year, which lengthens the growing season. 34

Sagebrush growth tends to be greatest in the fall and
Energy, Carbon, and Soil Organic Matter
early spring, whereas the growth of associated grasses

and forbs is more rapid in late spring and early sum-
The food web of sagebrush-dominated shrublands is
mer. Fall-through-spring precipitation and temperature
similar to that of grasslands, with probably more than
are good predictors of total annual growth of sagebrush
half the herbivory and carnivory occurring in the soil.
and associated plants.35
However, less than half the annual precipitation comes
The flow of carbon through ecosystems parallels
during the summer, which limits the success of warm-
the flow of energy, because so much energy is stored
season C4 plants, such as blue grama. Much of the plant
as carbon-rich compoundsthat is, organic materials,
growth is concentrated in the less-palatable big sage-
such as cellulose and other carbohydrates. They are
brush, which competes for moisture that could be used
important sources of food for large and small herbivores,
by other forage plants, namely, cool-season grasses.
aboveground and below. Moreover, animal carcasses
Consequently, the forage available for large ungulates
along with plant remains that do not burn become part
might be less than on the grasslands, especially those
of the mulch and soil organic matterboth of which
on the Great Plains where the plants have evolved with
increase water infiltration and nutrient availability. Big
more bison herbivory (see chapter 6).31 Sagebrush is
sagebrush tends to concentrate such materials around
known to be an important food for pronghorn, mule
its base more than do herbaceous plants, creating a
deer, and elk, but primarily during the winter, when
small-scale island of fertility that can persist even after
herbaceous forage is scarce or buried by snow. Bison rely
sagebrush is burned or removed in some other way.36
on grasses and forbs more than shrubs, as do cattle.
This patchiness in the surface soil helps promote sage-
Just as in other ecosystems, the energy stored in
brush ecosystem recovery after disturbances. In general,
stems and leaves aboveground can be released as heat
mineland reclamation specialists have found that con-
Sagebrush
serving as much soil organic matter as possible, along
other plants, such as Indian ricegrass.39 Losses of nitro-
with its associated microorganisms, is fundamental for
gen may occur through erosion, denitrification, or the
restoration.
emission of nitrogen gases.40
Because of concerns about the effects of carbon di
Several investigators have studied the rates of litter
oxide on climate change, ecologists have been conduct-
fall and decomposition in shrublands dominated by
ing experiments to determine whether the sagebrush
big sagebrush, both important processes for nutrient
ecosystem is a source of atmospheric carbon or a sink:
cycling.41 Some evidence suggests that nutrients are
does an ecosystem with big sagebrush tend to store
reabsorbed by twigs before the leaves fall, thereby con-
more or less carbon than is released from the ecosystem
serving nutrients in the shrubs.42 Still, soil nitrogen
as carbon dioxide?37 As with most semi-arid shrublands
is enhanced under shrubs. There also is great varia-
around the world, sagebrush steppe tends to be a sink
tion in soil organic matter and nitrogen availability
during wet years but a source in dry years. Generally,
higher in ravines with mountain big sagebrush to less
the sagebrush ecosystem accumulates carbon as the
on ridgetops with black sagebrush.43 Burrowing mam-
shrubs age, in the form of organic matter above- and
mals and harvester ants affect nutrient dynamics as
belowground. When most of the shrubs die, the eco-
well.44
system could become a source if the unburned woody
The major effect on nutrient cycling of having big
sagebrush biomass decays rapidly. However, Wyoming
sagebrush or any shrub in the ecosystem is to have more
ecologists Meagan Cleary, Elise Pendall, and Brent
woody biomass aboveground. Nutrient accumulation is
Ewers did not detect an increase in carbon emissions
therefore greater aboveground than where grasses and
after firepossibly because the growth of grasses and
forbs predominate. When the shrub dies, or as twigs
forbs increases greatly soon after fires, and the sage-
and branches die, more of the nutrients are added to
brush roots and stems decompose slowly.38
the soil surface as wood rather than as herbaceous plant
Still, the benefits of improving habitat or forage availability for some species by burning must be balanced
material. Also, woody fuels accumulate that can create

conditions for a longer, hotter fire.
against the undesirable effects of additional atmo-
Another influence of shrubs is to create islands of
spheric carbon dioxide from the fire, not to mention
fertility, as discussed previously. Shrubs accumulate
the invasion of undesirable plants (such as cheatgrass)
windblown debris and associated nutrients, and the
and the loss of critical habitat for threatened species (see
root system of the shrubs extracts soil nutrients from
below). Federal and state guidelines often call for man-
several feet beyond the canopy while dropping them on
aging ecosystems on public lands so that they sequester
the surface as litter in a smaller areapossibly another
carbon, resist the invasion of exotic plants, and offer
mechanism creating patchiness in the vegetation. The
habitat for such species as the greater sage-grouse and
death of the sagebrush root system must be important
pygmy rabbitall ecosystem services that are now
as well, not only in reducing competition for water and
highly valued.
nutrients with neighboring grasses and forbs, but also

in providing a substantial pool of nutrients as the roots
Nutrient Availability
decompose. The decaying root system can also alter soil

structure, possibly leading to greater infiltration rates.
Nutrient distribution in sagebrush-dominated eco
Such effects may be lost at some point after big sage-
systems is similar to that of grasslands, with the largest
brush has been removed, whether by fire or herbicides.
amounts in the soil. Water is usually more limiting to
Deeper soil organic matter may gradually be lost as well,
plant growth than are nutrients, though nitrogen may
along with the potential for hydraulic lift and the tap-
be limiting during wet years. As in grasslands, the major
ping of deep soil nutrients, including phosphorus.45 Of
source of nitrogen is probably precipitation, but symbi-
course, big sagebrush typically becomes re-established,
otic nitrogen fixation occurs locally in biological crusts
as discussed in the next section, restoring the benefits
on the soil surface; in the nodules of lupine and other
of having this shrub in the ecosystemunless cheat-
legumes; and possibly adjacent to the roots of some
grass changes everything.
119
Recovery after Disturbances

Big sagebrush is easily killed because it cannot sprout,
but historically it re-established after disturbances.
The rate of recovery varies considerably in relation to
the amount of precipitation, the amount of litter on
the soil, the degree of competition from herbaceous
plants, the intensity of grazing, the number of sagebrush seeds in the soil, the number of live shrubs that
remain, and the kind of big sagebrush on the site.46
Mountain big sagebrush has been observed to reestablish after about 40 years, whereas Wyoming big
sagebrush takes much longer.47 Without big sagebrush,
the ecosystem appears and functions like a grassland,
with junegrass, needle-and-thread grass, western
wheatgrass, Sandberg bluegrass, and numerous other
species. In relatively moist environmentsand in the
absence of cheatgrasssagebrush cover becomes quite
high as the shrubs age, persisting until the next fire or
drought.48
Even without planting, managers have come to
expect that big sagebrush and other native plants will
eventually invade areas where the plants have grown
before. That happened, for example, on abandoned
farmland in the Powder River Basin.49 Interestingly, the
establishment of big sagebrush during mineland reclamation is often difficult. Some research suggests that
survival is higher when mycorrhizal fungi are available
in the topsoil and when the recommended seeding rate
western Wyoming and neighboring states.53 In contrast,

shrub dieback during the mid-1980s in the Great Basin
was attributed to unusually wet conditions that persisted
for 5 years.54 Shrubs that were affected included antelope
bitterbrush, big sagebrush, fourwing saltbush, rabbitbrush, shadscale, and winterfat.
Extreme, ecologically significant weather events are
likely to become more frequent with climate change.
Higher mean annual temperatures will lead to less snow
accumulation and earlier snowmelt, and also a higher
potential for evapotranspiration throughout the year.
Because of added warmth, drier conditions for longer
periods during the summer could result even if annual
precipitation stays the same. If plants begin growing earlier in the spring, late-spring frosts could kill
some of them. Predicting precipitation changes with
global warming is currently difficult, but if droughts
are more severe, some shrublands now dominated by
sagebrush could become dominated by more droughttolerant shrubs or possibly grasslands. 55 Silver sagebrush
might be affected most severely, but big sagebrush also
depends on higher levels of precipitation than do most
grassland species. Sagebrush reinvasion after disturbances probably would be slowed, if it occurs at all,
because the extended cool, moist conditions required
for seedling establishment may become less frequent.
Drought-tolerant invasive species, such as cheatgrass,
may have a better chance of becoming established.
of perennial grasses is not too high.50
Grasshoppers
Drought, Spring Frost, and Extended Wet Periods
Unusual weather can kill shrubs, though the degree of
disturbance depends on the species. In a study in eastern
Montana, the shoots of silver sagebrush were more susceptible to drought than were those of big sagebrush, but
the silver sagebrush recovered more rapidly because of its
ability to sprout from the root crown.51 Winter mortality
of sagebrush can occur as well, probably from water stress
created by frozen soils, low soil water availability, and
below-average snowfall.52 Another cause may be the premature breaking of dormancy during periods of aboveaverage air temperature, making the plants susceptible to
subsequent frosts. Extensive areas of mountain big sagebrush have been killed by spring frost damage in south-
Outbreaks of grasshoppers killed approximately 50percent of the big sagebrush in a portion of the Powder
River Basin during the drought years of the 1930s.
Allred wrote, the grasshoppers swarmed in such hordes
that they devoured all of the edible vegetation, ate the
leaves and bark from the twigs of the sagebrush, and
completely girdled the more tender stems.56 Other
insects known to kill big sagebrush include gall midges,
the aroga moth, some beetles, and Mormon crickets.
Small mammals (such as voles) may girdle the stems of
big sagebrush, causing a significant disturbance when
their populations are high.57
Grasshopper outbreaks tend to occur during drought
years that are relatively warm. Thus, continued warm-
Sagebrush
ing of the climate could favor grasshoppers, killing
elevations, where mountain big sagebrush is the domi-
many of the shrubs and reducing forage availability
nant shrub.61 Fire rotation can be thought of as the aver-
at a time when it already is in short supply because of
age time required for one or more fires to burn an area
drought. Historically, grasshopper densities fluctuate
equal to the entire landscape under consideration. Its
greatly from one year to the next, leading entomolo-
tempting to think that global warming and drought will
gists to conclude that control measures in most parts of
shorten the fire-return interval in sagebrush ecosystems.
Wyoming are beneficial only during the year that insec-
However, if droughts are severe enough, the interval
ticides are applied. Usually, little or no multi-year bene-
could be lengthened because of less fuel production.
fit occurs.58 It is still unknown whether global warming
Gaining confidence in fire rotation estimates is im

portant, because prescribed fire can be a good manage
will change that conclusion.
ment strategy in some places. However, at the present

time, how often should sagebrush be burned, if at all?
Fire
Has fire suppression really created a stand in need of res-
Once started, whether by humans or lightning, fire can
toration, or might the area be thought of as old-growth
burn large areas of sagebrush. In 1868 James Chisholm
sagebrush with values for wildlife? Answers to such
explored the Wind River Basin and the vicinity of
questions are required for the proper management of
South Pass City during the Wyoming gold rush. He
animals that are sagebrush obligates. As the sagebrush
observed the rapid spread of at least two human-caused
habitat is fragmented in one way or another, these
fires across what must have been sagebrush-dominated
species have less habitat available to them, potentially
uplands,59 writing in his journal:
reducing their populations to the point where they
The grass took fire and all our efforts could not extinguish it. This time the situation was really alarming,
for there was a prospect that the entire Wind River
Valley might go up in a flame. . . . The flame went over
the nearest hill with amazing velocity, Heaven knows
how far. . . . The mountains were black and bare over
which we travelled for the rest of that day, and we saw
the fire pursuing its way far ahead in several directions, but fortunately away from the valleys.
become locally rare or extinct. Significantly, the reestablishment of sagebrush can require several decades,
and if invasive plants (such as cheatgrass) already have
seeds in the soil, there is the potential that sagebrush
may not recover at all.
In general, the effects of fire on sagebrush eco
systems are more prominent than those following
grassland fires because many of the shrubs are killed
(figs. 7.11 and 7.12). Other native species also may
be reduced in abundance by burning, such as Idaho
The words mountains and away from the valleys
fescue, needle-and-thread grass, pricklypear cactus,
suggest that much of the burned area was in the foot-
and threadleaf sedgethough others can be favored,
hills or higher, where mountain big sagebrush probably
such as bottlebrush squirreltail, Sandberg bluegrass,
was the dominant shrub and fuel accumulation might
and western wheatgrass.62 Although the cover of big
have been greater.
sagebrush is greatly reduced by fire, several sprout-
The mean fire-return interval in sagebrush steppe
ing shrubs persist or increase in some locations, for
in the 1800s is thought to have ranged from 20 to 100
ex
ample, horsebrush, rabbitbrush, and winterfat in
years, depending on the potential for plant growth and
relatively dry basins; and antelope bitterbrush, prairie
fuel accumulation on the site.
However, University of
rose, silver sagebrush, skunkbush sumac, and western
Wyoming ecologist William Baker did a thorough review
snowberry in more moist environments.63 Sprouting
of the methods used for calculating such estimates and
species definitely are favored if fires occur more than
concluded that fire rotations probably were much longer:
once every 2025 years. As expected, perennial grass
325450 years in shrublands dominated by low sage-
production commonly increases when competition
brush; 100240 years in shrublands dominated by Wyo-
from shrubs is reduced.64 Unfortunately, undesired
ming big sagebrush; and 70200 years or more at higher
weedy species also can increase.65
60
121

Fig. 7.11.The growth of grasses and
forbs increases greatly after big sagebrush is reduced by burning, as in this
area northwest of Rock Springs. The
vegetation on the left provides more
forage for livestock but is less favorable
for sage-grouse. Burning sagebrush can
favor the establishment of cheatgrass
and other invasive plants (see fig. 7.13).
For herbaceous plants in the sagebrush ecosystem,
a critical resource, whether for livestock or big game.
susceptibility to fire-caused mortality seems to depend
Burning sagebrush may be useful, but it should be done
on the amount of fuel that has accumulated above the
only in the context of management plans and after
root crowns, the depth of the latent buds below the soil
careful consideration of the undesirable changes that
surface, and the stage of growth at the time of burn-
could result, including the spread of invasive plants and
ing. Some research suggests that western wheatgrass
undesirable effects on sagebrush obligate animals, such
vigor and production increase with spring burning but
as sage-grouse.67
decline with fall burning, and that burning at any time

reduces the productivity of needle-and-thread grass.66
Repeated burning in consecutive years can cause a
Cheatgrass
shift from a community dominated by both cool- and
Though problematic for some sagebrush-dependent
warm-season species to one dominated primarily by
animals, prescribed fire simulates a natural distur-
warm-season species, which are generally more fire
bance to which many native plant species are adapted.
tolerant. This shift in dominance can be significant to
Older shrubs are removed and herbaceous production
land managers if early-season production is viewed as
increases. Unfortunately, invasive plants also may ben-
Sagebrush
300
Grasses
PERCENTAGE
200
a winter annual that begins growth in the fall or early
Thickspike
wheatgrass and
Plains reedgrass
Bluebunch
wheatgrass
spring and gains a competitive edge over many native

species that are slower to initiate growth. Generally,
fire and heavy livestock grazing hasten the spread of
cheatgrass, though some research suggests that proper
Bluegrass
100
management of native perennial species can slow this

invasion.72 Native species are more likely to become
Idaho fescue
300
Needle-and-thread
established if actions are taken to favor such species

within a year or two of a fire, for example, by seeding
Shrubs
Horsebrush
desired native species and protecting the rangeland
Rabbitbrush
200
from excessive grazing. The maintenance of biological

soil crusts seems to be important as well (as discussed
later).
100
With cheatgrass as a component of the flora (or even

just in the seed bank), land managers are confronted
Sagebrush
1936
1942
with the conundrum of increasing desired native
1948
1954
YEAR
1960
1966
Fig. 7.12.Changes in the canopy cover of several grasses and

shrubs after the burning of big sagebrush steppe in eastern
Idaho. The burn occurred in 1937. An increase (or decrease)
in plant cover is indicated when the line for a species is above
(or below) the dashed line. The two shrubs that increased, rabbitbrush and horsebrush, are capable of sprouting from their
roots; big sagebrush declined, because it does not sprout, but
it eventually became re-established from seed. Adapted from
Harniss and Murray (1973).
grasses by burning big sagebrush without increasing the

abundance of an undesired, introduced weed.73 Moreover, cheatgrass can become so abundant that it greatly
increases the flammability of the vegetation. Fires then
occur more frequently, diminishing the chances of sagebrush re-establishment and causing a decline in some
perennial grass speciesfavoring cheatgrass expansion
still further (fig. 7.13).74 The economic impact of cheatgrass is moderated to some degree by the fact that it can
be good forage before it flowers, but converting semiarid steppes to vegetation dominated by an introduced
efit, some becoming so abundant that they reduce the
annual is a significant change that has undesirable
growth of native species and the potential for restor-
consequences. In addition to increasing flammability,
ing wildlife habitat. Cheatgrass (Bromus tectorum), also
cheatgrass is associated with an increase in soil temper-
known as downy brome, is the most notorious example.
ature and erosion, along with reductions in soil organic
This annual plant has greatly complicated the practice
matter and small mammal abundance and diversity.75
of prescribed burning as a management tool. For many
The cheatgrass problem is not restricted to land man-
years managers thought that cheatgrass would not grow
aged for livestock. For example, a fire burned a large stand
well at higher elevations, but it is now commonly found
of big sagebrush in Little Bighorn Battlefield National
throughout Wyoming, even at elevations above 7,000
Monument in southern Montana, where livestock have
feet.68 With climate warming, it very likely will become
not grazed for many years. Much of the sagebrush was
more common in some parts of the state.69 Other prob-
killed, and bluebunch wheatgrass, a co-dominant before
lematic species in shrublands include broad-leaved
the burn, became less abundant. Cheatgrass became
pepper
weed, Canada thistle, Dyers woad, halogeton,
more common on the burned area, as did yellow sal-
leafy spurge, Russian knapweed, spotted knapweed,
sify. Managing the vegetation of a national monu-
Russian thistle, western salsify, and yellow and Dalma-
ment so that it reflects conditions prior to the arrival of
tian toadflax.70
EuroAmericans is a goal that sometimes seems impos-
Cheatgrass was introduced to western North Amer-
sible once invasive species become established. Further
ica in the late 1800s from the steppes of Europe.71 It has
research is necessary to resolve perplexing ecological
become common in many areas, because the species is
problems such as this one.
123
Fire suppression
Fire
Native perennial grassland
Sagebrush steppe with perennial grasses
Heavy grazing,
particularly
during spring
Moderate
grazing
Possible recovery
if no fire and only
fall sheep grazing
Fire
Native grass cover with possible
shift toward less-palatable warmseason species; rabbitbrush and
sagebrush may increase slightly
Only annuals,
rabbitbrush, and
sagebrush
Fire again
Only annuals;
increased erosion
Possible recovery
with reduced grazing
if no significant loss
of soil and no fire
More
frequent fire
Near-permanent
deterioration of site
Cheatgrass and other annuals increase, as do sagebrush,

rabbitbrush, and less-palatable grasses and forbs
Fig. 7.13.The apparent effect of cheatgrass introduction,

heavy livestock grazing, and increased fire frequency in the
big sagebrush steppes of Utah and Nevada. A similar scenario
may apply to Wyoming if cheatgrass becomes more common.

The introduction of exotic species can cause undesirable ecosystem changes. Adapted from West (1988).
A confounding factor may be climate change, as
be the best approach, along with adjustments in live-
both warming and higher levels of carbon dioxide
stock management, but that may require highly dis-
would probably favor cheatgrass expansion. Even
ruptive weed control practices, and then resting the
without warming, annual plants often evolve rapidly
rangeland for many years while native species become
to tolerate new environmental conditions. In addition,
re-established.76 Restoring biological soil crusts may be
other disturbances caused by road building, plowing,
required, about which little is known.
and various industries have created new habitats for
One way to stop or slow the spread of cheatgrass
this invader and others. Of great concern, cheatgrass
is to suppress fires where cheatgrass invasion is likely
an annualpersists even with very little moisture.
to occur, at least until effective and affordable control
After a decade or more of cheatgrass occupancy and
strategies are found. Curiously, there is evidence for the
the concomitant reduction of native species, the land
intriguing idea that some native species may be evolv-
could be left with little plant cover. Soil erosion would
ing adaptations for competing with invasive species.
accelerate.
However, preventing the invasion of exotic plants and
Efforts to control cheatgrass have generally been
pathogens should be a high priority where that is still
futile. Experiments with herbicides, the introduction
an option and where there is a desire to maintain bio
of pathogens, and strategically timed fires and grazing
diversity and the services provided by native ecosystems
have had some success, but only at considerable cost.
(see chapter 6 for further discussion of invasive plant
Promoting the establishment of native species may
management).77
Sagebrush
Effects of Livestock and Feral Horses on

Sagebrush Ecosystems
As discussed for grasslands, grazing is not a disturbance
unless too many animals are confined in too small an
area for too long a time. Bison, pronghorn, elk, deer,
this plant. This climate has been found very favorable to the restoration of health, particularly in cases
of consumption; and possibly the respiration of air
so highly impregnated by aromatic plants may have
some influence.79
and other herbivores were integral parts of the sage-
Fremonts cartographer, Charles Preuss, wrote, When
brush ecosystem for thousands of years, and they still
the wind blows, it is as though one were in a pharmacy.
are in some places. Ungulate grazing does not generally
Ten years later, in 1852, Platt and Slater wrote, Along
kill the dominant big sagebrush, though excess herbiv-
the Sweet Water, most of the way, are narrow bottoms
ory is known to cause declines in plant cover and, in
of good grass. Adjacent to these bottoms are large, arid,
some places, an increase in undesired plants.
wild-sage plains, extending to the mountains.80
One of the most widely mentioned responses to live-
The same is true today (fig. 7.14). Big sagebrush clearly
stock grazing is an increase in big sagebrush cover. How-
has been a dominant feature of the Wyoming landscape
ever, sagebrush abundance should not be considered an
for hundreds of years. Its abundance might have been
artifact of livestock grazing pressure everywhere.78 Long
favored by grazing in some areas, because the removal
before the cattle drives from Texas, John Fremont wrote
of palatable species reduced competition for sagebrush
about his travels across Wyoming in 1842:
seedlings. Such trends might have been most common
One of the prominent characteristics in the face of

the country is the extraordinary abundance of the
artemisias. They grow everywhereon the hills,
and over the river bottoms, in tough, twisted, wiry
clumps; and, wherever the beaten track was left, they
rendered the progress of the carts rough and slow. As
the country increased in elevation on our advance to
the west, they increased in size; and the whole air is
strongly impregnated and saturated with the odor of
camphor and spirits of turpentine which belongs to
on the fringes of the Great Plainsfor example, in portions of the Powder River Basin, where summer rainfall
is favorable for grasses and forbs.81
Livestock grazing can have dramatic effects on the
grasses and forbs associated with big sagebrush, but
drought is another important factor. Several studies
have found that conditions appearing to have resulted
from poor livestock management were actually caused
by extended dry periods.82 Indeed, short-term shifts in
climate can cause great changes in plant growth and
Fig. 7.14.This photograph

along the Sweetwater River
east of Jeffrey City was taken
by William Henry Jackson
in 1870. The vegetation,
dominated by Wyoming big
sagebrush, is essentially the
same today (Johnson 1987).
The granitic Sweetwater
Rocks on the left have limber
pine and Rocky Mountain
juniper. Elevation 6,200 feet.
U.S. Geological Survey photo
JWH00292.
125
the best examples of soil crusts in Wyoming have been

found in the relatively warm, desert-like parts of the
Bighorn Basin (fig. 7.15).
Because of the dramatic changes in the sagebrush
ecosystem that can be brought about by poorly managed livestock grazing, some ecologists have concluded
that, in contrast to grasslands on the Great Plains to the
east, the plants of the sagebrush steppe did not evolve
with grazing by bison and other large mammals.85 The
rationale is that the environment was unfavorable for
large herds of bison because of summer drought and the
scarcity of warm-season C4 species at that time.86 Therefore, food to sustain lactating cows was less available
during the summer. Biotic crusts might have formed
under these conditions, where bison populations were
low. When cattle and sheep became abundant in the
late 1800s, trampling would have destroyed the crusts,
causing the soil to become more erodible and more suitable for the invasion of introduced weedy annuals, such
as cheatgrass.
But does this scenario apply to the higher elevation
landscapes of Wyoming, where the accounts of early
explorers suggest that bison were common in sageFig. 7.15. Microphytic soil crusts cover the soil between shrubs
in this grazing exclosure in the Bighorn Basin east of Greybull. The common shrub in the exclosure is winterfat, which
is characteristic of desert shrublands (see chapter 8). Soil crusts
tend to occur more often in drier areas and are less common
in Wyoming than in the Great Basin to the west. Photo by
Jennifer Muscha.
brush-dominated landscapes? During midsummer in

1832 and 1833, Nathaniel Wyeth observed buffalo by
the hundred thousands in the vicinity of South Pass
and Big Sandy Creekwhere sagebrush would have
been abundant. At one point he noted that the country was covered with buffalo. In the same area, Jason
Lee wrote in 1834 that The buffaloe [sic] have eaten
nearly all the grass. Near Crowheart in the Wind River
biomass in the semi-arid West.83 As with most ecological
Basin, fur trapper James Clyman wrote about Buffalo
phenomena, explaining the effects of grazing requires
plenty and how a thousand or more bison were killed
an understanding of the interaction of several factors.
on one November day in 1823. In the Green River
Grazing pressure can also cause a decline in biologi-
Valley, in 1811, Wilson Price Hunt wrote, We were
cal crusts on the soil surface of some intermountain
surrounded by mountains in which were disclosed
basins.84 The crusts are growths of cyanobacteria
beautiful green valleys where numerous herds of bison
(former
ly known as blue-green algae), lichens, algae,
graze; which made them much more interesting to us,
mosses, and fungi. They minimize erosion, facilitate
because, for several days, we had not seen a single one
nutrient cycling, and create favorable conditions for
of these animals.87
seedling establishment. Possibly, the crusts were less
As noted, biological soil crusts are easily destroyed
common in Wyoming than in the Great Basin to the
by hooved mammals, whether bison or livestock. Such
west, because of a higher level of herbaceous plant cover
crusts are found in Wyoming, but not as commonly as
that attracted more bison and pronghorn, or because
in the Great Basinpossibly because of livestock graz-
temperatures were too low at the higher elevations
ing since the late 1800s, or possibly because there were
when adequate soil moisture was available. Thus far,
more bison in Wyoming prior to the arrival of livestock.
Sagebrush
Fig. 7.16.The population of feral horses
commonly exceeds the carrying capacity of the rangelands they occupy. Photo
by Ken Driese.
If cheatgrass has not become the widespread problem in
condition in some places. Debate about the comparative
Wyoming that it is in the Great Basin, the explanation
effects of native ungulates and domestic livestock will
may be partially because Wyoming plants evolved with
continue.89
larger herds of bison. If true, native herbaceous plants
Several species of horses evolved in North America,
would have been more competitive and more resistant to
but all became extinct about 10,000 years ago, along
invasive species. As discussed for grasslands, maintain-
with the camel, mastodon, saber-tooth tiger, and
ing competitive stands of native species may be the best
numerous other large mammals (see chapter 2). Some
strategy for excluding undesired plants (see chapter 6).
species survived in Eurasia and were the ancestors of
The management of Wyoming shrublands is more
modern day Equus caballus. Domestication began about
refined today than it was a half-century ago, but the
3000 bc. Horses were reintroduced to North America in
impacts of excessive livestock grazing in those early
1539, when Hernando de Soto and his conquistadores
years probably exist to this day in some areas. Perhaps
brought a herd of 220 to present-day Florida. Mov-
the sagebrush-dominated rangelands most similar to
ing westward was difficult, even with horses, but the
those of the mid-1800s are located some distance from
Spaniards traveled a circuitous route to what are now
frequently used sources of watertoo far away for
Texas and New Mexico. Along the way, some of their
most cattle to useor where fires have not always been
horses wandered off, were traded, or were stolen. Indi-
suppressed, or where the seeds of cheatgrass or other
ans obtained the horse in the 1600s, and by 1750 horses
invasive plants are not yet in the soil. Do such places
were in Canada. The horse caused dramatic changes in
still exist? Debra Donahue, professor of law at the Uni-
the lifestyle of indigenous people, just as they have for
versity of Wyoming, argues that, because of past and
the many people who raise horses today for a multitude
ongoing environmental damage by livestock, there is a
of purposes.90
legal mandate for stopping livestock grazing on public
Feral horses are now common on the shrublands of
lands.88 Doing this even on public land only, she con-
the southwestern quarter of Wyoming and westward
tends, would facilitate the restoration of both the sage-
into the Great Basin states (fig. 7.16). Some, known as
brush ecosystem and nearby riparian zones. Restoration
mustangs, are similar genetically to the horses brought
of sagebrush-dominated shrublands to their prehistoric
to America in the 1500s. With few predators to keep
condition may be expecting too much, but it is plau-
their populations in check, these large herbivores
sible that such initiatives will lead to better rangeland
became, essentially, an invasive species, often forming
127
large herds that graze the rangeland excessively. In 2012
national incentives for energy independence that may
the Bureau of Land Management estimated there were
result in further loss and fragmentation of sagebrush
about 3,500 horses in Wyoming.
habitat. Also, fire commonly has been prescribed for
91
To maintain the feral horse population at appropriate
increasing forage availability for livestock. Indirectly,
levels, horses are rounded up periodically and offered
climate change could become a factor if precipitation
for adoption or for relocation to sanctuariesthe only
patterns change so that deep-soil recharge during spring
option, by federal law, for adjusting horse numbers to
snowmelt is not adequate to support big sagebrush. All
habitat conditions. This method of control has been
such factors interact in troublesome ways, to the point
questioned, because the number of people desiring
that the sagebrush steppe is arguably an endangered
the horses can be insufficient to keep the population
ecosystem.93
in check. Research is under way to develop a practical
Of all the declining sagebrush-dependent species,
method of birth control, but some observers have con-
the greater sage-grouse has received the most attention
cluded that more drastic measures are needed. Despite
(fig. 7.17)in part because it has a curious courtship
the damage they can cause to rangelands, many people
display in the spring. More importantly, the U.S. Fish
enjoy seeing a herd of feral horses loping across the
and Wildlife Service concluded in 2010 that this bird
plains, a vignette of the western frontier.
warrants protection under the Endangered Species Act.

Considering the adverse socioeconomic consequences
Disturbing Trends in Sagebrush-Dependent

Animals: Greater Sage-Grouse
of listing the sage-grouse (because the energy and

agricultural industries would be affected), conserving
the habitat for this species has become a priority for
Considering how much of the West is dominated by
many sectors of society. The Natural Resources Con-
sagebrush, it should not be a surprise that some ani-
servation Service, part of the U.S. Department of Agri
mals require this habitat, most notably, pygmy rabbit;
culture, now has a sage-grouse Initiative; and the state
sagebrush vole; and various birds, including Brewers
of Wyoming has identified core areas that are critical
sparrow, sage sparrow, sage thrasher, and greater sage-
for the maintenance of sage-grouse populations (see
grouse.92 Other species are common in sagebrush eco-
chapter18).94
systems, though they seem to survive in other habitats
There are several species of grouse in North America,
as well, for example, coyote, pronghorn, and mule deer.
each adapted to different ecosystemsan evolutionary
More than 200 species of vertebrates are associated with
process known as adaptive radiation that results in less
big sagebrush in one manner or another.
competition from closely related species. Not surpris-
Since the 1990s, the abundance of sagebrush-
ingly, the large land area occupied by sagebrush pro-
dependent species has been declining because of
vided a niche for one of them, the greater sage-grouse.95
habitat loss caused by various factors, including too-
Through evolution, this bird acquired adaptations that
frequent fires; inappropriate livestock grazing in some
enable it to eat sagebrush leaves, along with insects and
places; and habitat fragmentation stemming from road
forbs. For protection from predators, including ravens,
building, plowing, the construction of homes in rural
coyotes, and eagles, birds of all ages have color pat-
subdivisions, wind farms, and the extraction of oil, gas,
terns that provide camouflage as they hide under the
coal, and uranium. Anything causing a decline in both
shrubs and grasses. Moreover, the sage-grouse are large
shrub and herbaceous plant cover could lead to less
(the largest grouse on the continent), which is adaptive
food availability and less protection from predators for
for the often cold environment they occupy. The birds
sagebrush-dependent species. As the remaining tracts of
range over areas up to 30 miles across during the year,
sagebrush steppe become smaller, the impacts of distur-
but return faithfully to specific places known as leks for
bances have become significantly greater.
courtship and reproduction.96 Breeding adults do not
Reversing such trends is one of the most formid
readily adopt or develop new leks, and they avoid using
able challenges facing land managers today. There are
leks close to roads with considerable traffic at the time
Sagebrush
Fig. 7.17. Male (left) and female (right) greater sage-grouse.

The male is strutting on his lek in early spring. Because of
habitat fragmentation and other factors, the abundance of
of courtship. The birds also avoid leks that have been

compromised by the noise of traffic and drilling.97
Conservation biologists and some land man
agers
this bird has been declining throughout the West during the
past 20 years. In 2012 about 38 percent of the birds lived in
Wyoming. Photos by Mark Gocke.
To further improve sage-grouse habitat, new ap

proaches to livestock grazing are being considered,
with the goal of maintaining adequate grass and forb
have now taken steps where possible to conserve or
cover under the shrubs. This was less of a concern when
improve sage-grouse habitat. The leks are protected
sage-grouse were more abundant and large expanses of
from disturbances associated with new wells, wind
sagebrush steppe without industrial development still
farms, and other developments, and also by keep-
existed. Using small radiotransmitters attached to birds,
ing nearby traffic to a minimum during the breed-
habitat biologists have found that hens in relatively dry
ing season. Keeping tall structures to a minimum is
areas select sagebrush communities with more grass
important, as they provide perches for predatory rap-
cover and litter for nesting and raising their broods.
tors.
New roads are constructed along rights-of-way
Research is under way to determine whether thinning
that avoid critical habitat; the burning of sagebrush
the old shrubs mechanically leads to an increase in
is discouraged or done more judiciously.99 Fires did
nesting habitat as well as the establishment of young
occur historically, probably with minimal effects on
sagebrush.100
98
the sagebrush-dependent animals, but with current
But habitat degradation and road traffic are not the
levels of habitat loss and fragmentation, such events
only factors in play that cause population declines.
can have significant adverse effects in many areas.
Sage-grouse are susceptible to the exotic West Nile
Un
fortunately, as discussed previously, fire manage-
virus, which causes lethal encephalitis in the birds.
ment has been confounded by the invasion of cheat-
Transmitted by mosquitos, this disease was first iden-
grass, which increases fire frequency.
tified as a contributing cause of sage-grouse decline in
129
2003 and was associated with increased mosquito abun-
that could result from warming could cause an unstop-
dance, caused most likely by wastewater ponds created
pable decline in big sagebrush cover.102
during the extraction of coalbed methane.
101
Hundreds
of new ponds were constructed in sage-grouse habitat
Overall, traveling by car or foot though intermountain
in the Powder River Basin alone. Attempts to identify
basins offers views to the horizon of a seemingly nat-
the virus as the cause of population declines have been
ural sagebrush ecosystemmonotonous to some but
confounded by an extended drought that occurred at
inviting to others. The changes over the years are some-
the same time, which could have reduced food avail-
times subtle and not obvious from the road, but aerial
ability. Disease and drought together, along with habi-
views reveal miles of new pipelines, transm ission lines,
tat fragmentation, surely contributed to the alarming
and roads constructed for resource extraction. Recon-
80 percent decline in sage-grouse numbers in the Pow-
ciling the needs of threatened wildlife populations
der River Basin from 2001 to 2012.
with fossil fuel extraction is one of the greatest chal-
The control of road construction, traffic, prescribed
lenges facing western states.103 State governors have
fires, well location, and livestock grazing is feasible,
taken unusual steps in an attempt to reverse the trend
but now the less-controllable effects of climate change
for sage-grouse, as discussed further in chapter 18.
must be considered as well. Warming could benefit
Along with the confounding effects of climate change,
sage-grouse, possibly by favoring the production of the
the issues involved are complex and surely will affect
insects required by chicks. Unfortunately, warming also
the way people benefit from sagebrush-dominated eco-
favors the spread of West Nile virusand the droughts
systems in the future.
Desert Shrublands
and Playas
Chapter 8
Occupying the driest parts of the intermountain
shrubs are Gardners saltbush, winterfat, birdfoot sage-
basins, desert shrublands cover about 8 percent of
wort, shadscale, fourwing saltbush, and greasewood.1
Wyoming. The mean annual precipitation is less than
The dominant plants form a patchy mosaic associated
8 inches, and the generally fine-textured soils tend to
with the amount and timing of water availability, tem-
be saline (see figs. 1.2, 1.5, 3.4, and 8.1). The predomi-
perature at the time water is available, soil texture and
nant plants are different from those in grassland and
salinity, and depth to groundwater (fig. 8.2). Plants
sagebrush ecosystems, though big sagebrush can be
grow slowly and support fewer large herbivores than
found, especially where snow drifting occurs. Typical
in wetter zones.
Fig. 8.1. Desert shrublands

are found where the average annual precipitation is
8 inches or less. Common
plants include Gardners
saltbush, shadscale, fourwing
saltbush, birdfoot sagewort,
bud sagewort, pricklypear cactus, winterfat, and Wyoming
big sagebrush. Greasewood
grows adjacent to playas or
where groundwater seeps to
near the surface. Most of the
plants are shrubs. This area
is in the southern part of
the Great Divide Basin (also
known as the Red Desert),
between Rock Springs and
Rawlins (see fig. 1.2). Elevation about 6,800 feet.
131
High Big sagebrush

Winterfat
Gardners
Shadscale saltbush
are less than in grasslands and sagebrush steppe, primarily because of reduced water availability,3 and perennial
grasses and forbs are less abundant. Fires are infrequent.
Variation in plant distribution can be caused by slight
changes in topography. Playas occur in low depressions,
ARIDITY
where water sometimes accumulates before evaporating,

leaving a dry soil surface that is white with precipitated
salts.4 Plant life is sparse in such areas.
Salt accumulation in desert soils is a formidable problem for the survival of many plants, especially seedlings.
Most notable is that many plants cannot extract water
from soil if the salt (solute) concentration of the soil water
is higher than that of the root cells. Plants known as halo-
Greasewood
Low
phytes have evolved adaptations that solve this problem
Pickleweed
Low
SALINITY
by maintaining cellular concentrations at higher levels
High
Fig. 8.2. Distribution of six desert shrubs in relation to gradients of salinity and aridity. All are found in Wyoming except
pickleweed, the most salt-tolerant species, which is found in
Utah and Nevada. Adapted from West (1988).
than those found in the soil solution. In the leaves of

some halophytes, the salt concentrations promote water
uptake that causes leaf succulence. Greasewood and red
swampfire are examples.5 Other halophytes have evolved
special adaptations for excreting salts (fig. 8.3).6
Halophytes essentially function as pumps, collecting
salts from throughout the rooting zone and depositing
Except for soil salinity, desert shrublands have many
them in a smaller area as leaves fall. Some ecologists
features in common with sagebrush steppe: dominance
have hypothesized that salt deposition under shrubs
of drought-tolerant shrubs with small leaves; exten-
might prevent the establishment of other plants nearby,
sive, sometimes deep root systems for obtaining water
reducing competition for limited water and nutrients
and nutrients; sensitivity to various kinds of herbivores
a form of allelopathy.
and invasive plants; and processes that enhance the
As in grasslands and sagebrush steppe, competition
efficiency of water and nutrient use, such as hydraulic
for resources determines which desert plants and animals
redistribution in the soil and nutrient reabsorption from
manage to survive. This competition is often minimized
leaves prior to leaf fall. Plant cover and annual growth
because of seasonal differences in activity, or because
Fig. 8.3. Scanning electron micrograph

of salt-secreting glands on the leaves of
a salt-tolerant plant in the genus Cheno
podium. The bulbous glands, which are
modified leaf hairs, enlarge until they
burst, after which rainwater washes
the salts onto the soil. Magnification
about 33 times. Microphotograph by
HowardC. Stutz.
Desert Shrublands and Playas
different species tap different parts of the ecosystem for
shrubland, saltgrass meadow, desert grassland, saltbush
the water and nutrients they need (see chapter 6). One
shrubland, and mixed desert shrubland. Greasewood
study focused on how coexisting winterfat and shadscale
shrubland and saltgrass meadow are characteristic of
differed in their adaptations.7 The results showed that
comparatively wet depressions (fig. 8.4), as described
winterfat, a C3 species, is capable of rapid photosynthesis
in chapter 5, but they also are found on some slopes
in the spring and then becomes relatively inactive during
where water seeps to the surface, as sometimes occurs
the rest of the year, when moisture is limiting. In con-
in badlands. The desert grassland community occurs
trast, shadscale, a C4 species, has moderate rates of photo-
where soils are drier and less saline, where drainage is
synthesis throughout the growing season, about half that
not impeded by fine-textured soils, and on the lee sides
of winterfat in the spring but two times faster in the sum-
of slopes where snowdrifts occur. Saltbush shrubland
mer. Both species have a high tolerance for water stress,
is the community with the highest salinity and has an
and both are able to fix carbon at low temperaturesan
abundance of plants in the genus Atriplex. The fifth com-
important adaptation for the cool, short growing season
munity, mixed desert shrubland, is found on soils that
that characterizes northern deserts. Differences between
are intermediate in salinity and have a diversity of shrub
the two species help explain how they coexist. Other des-
species. Sagebrush steppe is sometimes thought of as
ert shrubs have similar adaptations.
desert shrubland, but it occurs at slightly higher eleva-
Sharp spines are another notable adaptation of some
tions or where annual precipitation is somewhat greater.
desert shrubs, especially those that are palatable. This
As described in chapter 7, the sagebrush community has
adaptation minimizes grazing in an environment where
a higher cover of perennial grasses and is likely to burn
the production of plant tissue is slow. Examples are
more frequently than desert shrub vegetation.
greasewood, spiny hopsage, bud sagewort, and shadscale.
Greasewood does not tolerate long periods of
Also, like big sagebrush, some desert shrubs produce both
drought. A deciduous shrub, it commonly grows to
ephemeral and overwintering leaves, improving the effi-
a height of 3 feet or more, and the roots have been
ciency of water use.8 A good example is shadscale.
observed at depths of 10 feet. Because of salt glands

on the leaves, soil salinity under the shrub is typically
The Desert Mosaic
higher than between shrubs.9 Notably, greasewood

seedlings can survive under the parent shrubs where
Five communities can be easily identified in the des-
the salinity is highest, though germination occurs
ert mosaic, based on the dominant plants: greasewood
early in the spring when the soil solution is less saline
Fig. 8.4. Greasewood shrublands in

the Great Divide Basin. Saltgrass and
Gardners saltbush are also common
in this area. The soil surface is often
white, because salts accumulate as water
evaporates. Elevation 6,550 feet. Photo
by Ken Driese.
133

Fig. 8.5. Greasewood is common on
terraces along many ephemeral streams,
with basin big sagebrush and plains silver sagebrush in ravine bottoms, where
salts are washed downstream more
frequently.
because of dilution by spring snowmelt and rains.10
tern is for saline meadows with inland saltgrass and its
Root sprouting also produces new shrubs. At least four
associates to be dotted with low mounds, where much
kinds of greasewood communities have been identi-
taller alkali sacaton grows. The mounds very likely are
fied: greasewood-shadscale, greasewoodbig sagebrush,
the result of silt deposition around the taller grass. Blue
11
greasewood-grass, and a greasewood monoculture.
grama and western wheatgrass can be found with the
Big sagebrush occurs with greasewood only sporadi-
saltgrass in some places, though these species are less
cally, where the soils have a higher infiltration rate and
tolerant of soil salinity.
are less saline.
Inland saltgrass has been the subject of several stud-
Normally greasewood grows at low points in the
ies because of its value as a forage plant. It normally
landscape, but an interesting anomaly sometimes occurs
grows at relatively low levels of salinity.14 Interestingly,
in desert riparian zones, where greasewood shrublands
saltgrass does not concentrate salts in its tissues. Seeds
are found on terraces and basin big sagebrush in ravine
of this grass germinate only when relatively high soil
bottoms, closer to the water table (fig. 8.5). Basin big
moisture coincides with optimal seedbed temperature,
sagebrush grows quite well in the ravine environment,
and when the available soil water is adequate to dilute
probably because of frequent flooding that washes away
the salts. The same might be true for many desert plants.
many of the salts that might otherwise accumulate and
Mixed desert shrubland occurs in dry upland habitats
inhibit the establishment of sagebrush seedlings. Also,
(fig. 8.6). Characteristic shrubs are shadscale, bud sage-
the ravine bottom soils tend to be more permeable than
wort, winterfat, spiny hopsage, and Gardners saltbush
terrace soils, which is favorable for sagebrush.12 Grease-
(table 8.1). Shadscale is more common on dry, some-
wood apparently receives adequate water on the terraces.
what sandier soils; spiny hopsage is common on sandy
Some saline depressions have no shrubs and are bet-
soils where snow accumulation is higher; and winterfat
ter referred to as saltgrass meadows. They are charac-
is more common on fine-textured soils (with more silt
terized by inland saltgrass, alkaligrass, alkali sacaton,
and clay). Big sagebrush can be found in ravines, where
and, in wetter areas, alkali cordgrass. Several halophytic
the soil is less saline and where water infiltration into
members of the goosefoot family are present here as
the soil in the spring is higher (fig 8.7).15 Although big
well. One, red swampfire (Salicornia rubra), is a small
sagebrush is thought to be intolerant of saline or alkaline
succulent plant that turns bright red in the fall and
soils, it sometimes occurs with greasewood.16 This might
seems well adapted to slickspot soilsa name derived
be explained by genetic variation,17 or by the fact that
from the slippery soil that develops after a rain because
plant species composition and growth are a function of
of the soil-particle-dispersing effect of high sodium
topography, aspect, snow accumulation, parent material,
concentrations. Red swampfire commonly forms a
and other factorsnot just soil pH or conductivity. No
band closest to the water, with concentric bands of
single factor explains why plants grow where they do.
saltgrass meadow, greasewood shrubland, and mixed
Desert pavement and coppice dunes are common in
desert shrubland occurring as distance from the pond
mixed desert shrublands. Silt and sand are easily eroded
increases (see chapter 5).13 A common vegetation pat-
by wind, leaving a surface of pebbles. The windblown

Fig. 8.6. Mixed desert
shrubland on the left, with
Wyoming big sagebrush in
ravines, near Green River.
particles accumulate around shrubs, forming small
products intended for human consumption.19 Any prac-
dunes (see fig. 7.9). Bud sagewort, galleta, greenmolly
tice that exposes selenium-bearing rock also increases
summercypress, and Gardners saltbush are common on
the density of plants that accumulate selenium. Even-
the drier desert pavement soils, whereas big sagebrush,
tually, as leaves fall and the plants die, this selenium
greasewood, Great Basin wildrye, Indian ricegrass,
is deposited on the soil surface, where it is subject to
spiny hopsage, and thickspike wheatgrass are common
redistribution by erosion. If this happens, toxic concen-
on the dunes. The coppice dunes apparently provide a
trations could develop in places where they would not
relatively moist and less saline environment. Excessive
otherwise be expected.
livestock grazing in deserts can promote the formation
Saltbush shrubland, sometimes called salt desert
of coppice dunes, as described for big sagebrush com-
shrubland, is dominated by Gardners saltbusha short
munities in the previous chapter.
shrub also known as saltsage or Nuttalls saltbush (fig.
Further variation in desert shrublands, and in some
8.8). Associated plants include birdfoot sagewort and
foothill shrublands as well, is caused by seleniuman
bud sagewort, both of which are species of Artemisia
element required for proper animal nutrition, but it can
(see table 8.1). This vegetation type occurs commonly
become lethal as concentrations increase. Some plants,
on fine-textured soils developed from shale or alluvium.
such as woody aster, princesplume, goldenweed, and
In some areas the low shrubs are rare and the com-
several species of milkvetch, accumulate selenium in
munity appears more like desert grassland, with blue
their tissues. The presence of these plants, known as
grama common on somewhat sandier soils and western
selenium indicators, has been used to map selenium-
wheatgrass on fine-textured clay soils. 20 On windswept,
rich bedrock, such as certain shales, siltstones, and
shallow soils with relatively low salinity, the dominant
claystones that are carbonaceous or have high concen-
species may be bluebunch wheatgrass, needle-and-
trations of iron. A concern in western states is that sur-
thread grass, and several species of cushion plants (fig.
face mining and irrigation could increase the amount of
8.9; see table 8.1).
18
land where selenium accumulates to toxic levels in both
Though its total biomass is low, Gardners saltbush is
water and livestock forage, and even meat and plant
valued for the forage it produces on winter ranges, and
135
Table 8.1. Some characteristic plants of desert shrublands in Wyominga
Common name
Latin name
Saltbush
shrubland
Mixed desert Greasewood

shrubland
shrubland
Saltgrass
meadow
Desert
grassland
SHRUBS
Basin big sagebrush
Artemisia tridentata ssp.

tridentata
ravines
ravines
Birdfoot sagebrush
Artemisia pedatifida
Bud sagebrush
Picrothamnus desertorum
Little (alkali) sagebrush
Artemisia arbuscula ssp.

longiloba
Wyoming big sagebrush
Artemisia tridentata ssp.

wyomingensis
Douglas rabbitbrush
Chrysothamnus viscidiflorus
Rubber rabbitbrush
Ericameria nauseosa
Fourwing saltbush
Atriplex canescens
Gardners saltbush
Atriplex gardneri
Shadscale saltbush
Atriplex confertifolia
Greasewood
Sarcobatus vermiculatus
Spineless horsebrush
Tetradymia canescens
Spiny hopsage
Grayia spinosa
Winterfat
Krascheninnikovia lanata
Alkali sacaton
Sporobolus airoides
Blue grama
Bouteloua gracilis
Bluebunch wheatgrass
Pseudoroegneria spicata
Thickspike wheatgrass
Elymus lanceolatus
Western wheatgrass
Pascopyrum smithii
Bottlebrush squirreltail
Elymus elymoides
Foxtail barley
Hordeum jubatum
Indian ricegrass
Achnatherum hymenoides
Needle-and-thread grass
Hesperostipa comata
Nuttalls alkaligrass
Puccinellia nuttalliana
Salina wildrye
Leymus salinus ssp. salinus
Saltgrass
Distichlis spicata
Sandberg bluegrass
Poa secunda
Goosefoot
Chenopodium spp.
Green molly
Bassia americana
Hookers sandwort
Arenaria hookeri
Nuttalls povertyweed
Monolepis nuttalliana
Prairie (fringed) sagewort
Artemisia frigida
GRASSES
FORBS
Saltbush
shrubland
Mixed desert Greasewood

shrubland
shrubland
Saltgrass
meadow
Desert
grassland
Suaeda spp.
Phlox hoodii
Common name
Latin name
Pepperweed
Lepidium densiflorum
Pricklypear cactus
Opuntia polyacantha
Red swampfire
Salicornia rubra
Saltlover (halogeton)
Halogeton glomeratus
Scarlet globemallow
Sphaeralcea coccinea
Seablight
Spiny (Hoods) phlox
Tanseyleaf tanseyaster
Machaeranthera tanacetifolia
Textile onion
Allium textile
Yellow spiderflower
(beeplant)
Cleome lutea
A dash indicates that the plant is absent or uncommon.
the seed is sometimes planted for reclamation purposes,
was found to be 810 inches in central Washington,
especially when the topsoil has high pH and salinity.
of which 1831 percent was transpired.24 As might be
Its roots spread laterally to a distance of 6 feet, with
expected, greasewood height, canopy cover, and total
taproots penetrating to 3 feet or more. Root sprouting
leaf area were highest when groundwater was near the
or layering is the major form of reproduction once the
surface. One study found that greasewood caused less
plants are established. These small shrubs are thought

to live more than 50 years.21
30
is winterfat, so named because some ranchers observed

that their livestock could gain weight by eating this plant
when it was dormant.22 In addition to sheep and cattle,
winterfat is eaten by pronghorn, elk, deer, and rabbits.
It also is one of the most useful shrubs for reseeding
disturbed rangelands where annual precipitation is less
than 10 inches. As with all desert plants, new seedlings
become established only when favorable moisture and
temperature conditions coincide for a long enough time.
The Desert Shrubland Ecosystem

With regard to total annual plant growth, cold-desert
ecosystems with different dominant species have been
ranked from high to low in this order: big sagebrush,
Gardners saltbush, greasewood, shadscale, spiny hopsage, and winterfat.23 The most productive shrublands
undoubtedly have the highest water consumption.
Annual evapotranspiration from a stand of greasewood
WATER INFILTRATION (cm)
Another highly valued forage for livestock and wildlife
Big sagebrush steppe
25
20
15
Grassland
10
5
0
Desert shrubland
0 10
30 45 60
90
120
MINUTES
180
Fig. 8.7. Desert shrublands often have finer-textured soils than

are found in grasslands or big sagebrushdominated shrublands, which accounts for much slower water infiltration (see
also fig. 3.8). In arid regions, less water is available for plant
growth in soils with slow infiltration rates. Adapted from
Nichols (1964).
137
Fig. 8.8. (above) Saltbush desert shrubland northwest of Rock

Springs. The dominant shrubs are Gardners saltbush, bud
sagewort, and birdfoot sagewort. The treeless escarpment is
formed from late Cretaceous sandstones and shales. Elevation
6,500 feet.
Fig. 8.9. (left) Windswept desert shrubland on a rim of the
Great Divide Basin, with cushion plants adapted for conserving water and heat. Common plants in this community are
Hookers sandwort, squarestem phlox, matted buckwheat,
stemless goldenweed, and stemless four-nerve daisy. Similar
growth forms but different species are found in alpine tundra.
soil water drawdown when leafless than did a stand of

evergreen big sagebrush.25 Generally, invasive cheatgrass grows more abundantly in relatively moist stands,
such as where greasewood occurs (see chapter 7 for a
discussion of cheatgrass). Research in Utah indicates
that cheatgrass can invade desert shrublands during
years of above-average rainfall.26
Other
than
greasewood-dominated
shrublands,
which occur where water is more readily available, desert ecosystems are characterized by potential evapotranspiration that greatly exceeds annual precipitation
and the water that might be gained by drainage from
surrounding slopes. Consequently, plants are frequently
under water stress. Infiltration rates vary considerably

and appear to influence the distribution of different
plants (see fig. 8.7). Nutrient leaching is rare, but low
levels of nitrogen can limit plant growth.27 Nitrogen fixation results from species of cyanobacteria, bacteria, or
lichens found in biological soil crusts, but atmospheric
inputs of nitrogen may be more important.28 Nutrient
losses are of little consequence, primarily because water
is usually the primary limiting factor, but also because
most nitrogen requirements for new plant growth probably are met by mineralization of soil organic matter
and reabsorption from senescing leaves. As in grasslands and other shrublands, herbivores and detritivores
increase the rate of nutrient cycling (see chapter 6), and
the presence of shrubs creates islands of fertility (see
chapter 7).
Except for the effects of livestock grazing, relatively
little is known about plant-animal interactions in desert shrublands, compared to grasslands and sagebrushdominated shrublands. Jackrabbits are thought to prefer
winterfat for food but use nearby patches of big sagebrush for hiding cover (fig. 8.10). In one study, winter-
Fig. 8.10. White-tailed jackrabbit in the Red Desert, near

Adobe Town. Photo by Ken Driese.
fat nearest the sagebrush was subject to heavier grazing

by the hares, which in turn facilitated the expansion
of big sagebrush. In this way the jackrabbits influenced
phytic plant biomass? In the case of greasewood shrub-
the vegetation mosaic, at least during years when their
lands, with finer-textured soils that are wet and possibly
numbers are high. Some investigators believe that jack-
anaerobic during much of the year, is most herbivory
rabbits in large numbers can graze rangelands more
still belowground, as in grasslands and some shrub-
heavily than livestock or big game.29
lands? How do high concentrations of selenium affect
In addition to jackrabbits, Townsends ground squir-
microbial activity?
rels can be abundant in desert shrublands during some

years, providing prey for raptors.30 In southern Idaho,
the rodent populations drop significantly when desert
Disturbances and Succession
shrublands are converted to communities dominated
The major disturbances in desert shrublands are
by annuals, such as cheatgrass.
drought and extraordinarily heavy herbivory, whether
Little research has been done on the hydrology,
from grasshoppers, Mormon crickets, bison, or live-
energy flow, and nutrient cycling of Wyoming desert
stock. Fires are less frequent in more arid environments
shrublands, but some interesting opportunities exist.
because of insufficient fuel, but they surely occur in
For example, just as it is helpful to consider the effects
greasewood and mixed desert shrublands during years
that the presence of big sagebrush has on ecosystem
when fuel accumulates, especially after periods of rela-
properties (see chapter 7), it also is interesting to con-
tively high rainfall that coincide with light grazing, or
sider the effects of salinity and selenium. Does nitrogen
where cheatgrass has invaded (see chapter 7). Harvester
become more or less available as salinity increases? Are
ants are abundant in some saltbush shrublands, creat-
soils high in sodium more susceptible to erosion? How
ing patchy disturbances,31 and sometimes two years or
do patterns of herbivory and decomposition change, if
more of drought will cause portions of a shrub to die,
at all, because of higher salt concentrations in the halo-
if not the entire plant.32 Shadscale is apparently very
139
sensitive to drought, with dramatic annual variations
In addition to the loss of soil crusts, introduced
in plant cover associated with spring precipitation and
annual plants have greatly affected some desert shrub-
the abundance of a scale insect that feeds on roots.33
lands, just as they have some shrublands dominated by
Drought is not the only explanation offered for des-
sagebrush (see fig. 7.13). Three common weeds are cheat-
ert shrub mortality. Between 1977 and 1986 the shrubs
grass, halogeton, and Russian thistle.40 Of the three,
over large areas in the Great Basin of Utah died, appar-
cheatgrass is by far the most invasive and has become
ently stemming from four factors triggered by several
a major problem in western intermountain basins (see
years of unusually high precipitation: (1) frequent
chapter 7). Some varieties of the species are adapted to
periods when the soil was anaerobic, (2) increased soil
tolerate saline soils and can invade even with little or no
salinity, (3) increased susceptibility of shrubs to soil-
livestock grazing.41 As in sagebrush, cheatgrass typically
borne diseases, and (4) the loss of shrub tolerance to
grows so densely in favorable years that fires burn in
the droughts that occurred after the wet period ended. 34
places where previously they could not because of in
Thus far, a similar dieback of desert shrubs has not been
sufficient fuel. And the fires destroy the patches of soil
reported for Wyoming.
crust that might still persist. Arguably, cheatgrass is the
Excessive livestock grazing has been a common
most severe problem facing desert shrubland managers
disturbance in most desert shrublands. 35 Cattle drives
in Utah, Nevada, and southern Idaho, and with climate
first started on the Great Plains, where the rangelands
change and ongoing evolution, it could become a severe
were adapted to the grazing of large mammals because
problem in Wyoming as well. Douglas Shinneman and
of large numbers of bison there. Moving westward, the
William Baker compared many sites in the Great Basin
cattle foraged in big sagebrush steppe, which also had
states,42 finding more rapid cheatgrass invasion where
considerable grass cover and a long history of grazing by
soil crust cover had declinedanother indication that
bison. Sometimes the concentrations of cows and sheep
managers must consider microbial organisms along
were much higher than livestock managers would allow
with large animals, fire, and the dominant plants as
today, and without careful herding, the same numbers
they strive to achieve sustainable land management.
of animals easily wandered into the adjacent desert
Though not yet reported for Wyoming, an unusual
shrublands, causing dramatic changes.36 Some of the
effect of cheatgrass invasion was to convert a shrubland
palatable plants declined with this added grazing pres-
in eastern Washington dominated by big sagebrush to
sure, while others increased. 37 Recovery after reducing
one dominated by greasewood. The following mecha-
the numbers of livestock was very slow because of the
nism was proposed by William H. Rickard: (1) a high
arid environment.
cover of cheatgrass led to the interception and use
One of the first adverse effects of livestock graz-
of more of the annual precipitation, resulting in in
ing was probably the degradation of delicate biologi-
adequate percolation of water to the depths required by
cal soil crusts, where they occurred, by trampling (as
big sagebrush; (2) fire frequency was increased by the
discussed in chapter 7). 38 When this happens, soil ero-
flammable cheatgrass, but greasewood (already present
sion increases, nitrogen fixation may be reduced in
in the community) persisted because it could sprout,
some years, and favorable sites for seed germination
unlike big sagebrush; and (3) greasewood caused an
and seedling establishment can be lost. Considerable
increase in surface soil salinity, which big sagebrush
research has been done on the importance of the crusts
seedlings could not tolerate.43 Similar changes could
for desert shrubland ecosystems in the Great Basin, but
occur in Wyoming if cheatgrass were to become abun-
their importance in Wyoming desert shrublands is still
dant where big sagebrush and greasewood coexist.
uncertain. Today they are most easily found in the Bighorn Basin, in places where grazing has been minimal.
Were the crusts less common in Wyoming to begin
Scenario for the Future
with, because of a higher cover of perennial grasses
Extrapolating from what is known about desert eco
between the shrubs and heavier grazing by bison? An
systems, a warmer climate would increase potential
answer is not possible at this time.39
evapotranspiration and rates of soil drying. There could
be occasional periods of high precipitation, but the dry
Whether disturbed by excessive livestock grazing or
periods are likely to be longer than during the past cen-
industrial developments, the restoration of desert eco-
tury, and more of the annual precipitation will come as
systems is difficult. Even under experimental conditions,
rain rather than snow. Big sagebrush in Wyoming and
new seedlings typically do not survive, and the devel-
the region will become less common if deep soil water
opment of new soil crusts takes 10 years or more. Soil
recharge is inadequate. Most likely, desert plants will
erosion occurs with less plant cover, leading to the for-
expand to occupy some land now dominated by sage-
mation of desert pavement and coppice dunes. Warmer
brush. Other changes in plant species composition will
temperatures will make reclamation even more difficult
occur as well. For example, already endangered species,
and most likely will cause a decline in the amount of
such as the greater sage-grouse, will lose some of their
soil organic matter, making the soils still more erodible
habitat as big sagebrush declines, and cheatgrass will
and a source of atmospheric carbon dioxidethereby
become more common and will occupy a larger area,
negating attempts to sequester carbon in terrestrial
very likely surviving the droughts (because it grows pri-
ecosystems. The desert shrublands of the twenty-first
marily in late fall and early spring, when sufficient mois-
centuryand the way they are usedare likely to be
ture probably will be available). The increased biomass
considerably different than during the twentieth cen-
of cheatgrass will probably increase fire frequency, caus-
tury, primarily because of mistaken assumptions about
ing further declines in big sagebrush. Carrying capacity
livestock management in the late 1800s, the invasion of
for big game and livestock in the intermountain basins
cheatgrass shortly thereafter, and the climate changes
will then decline.
now under way.
141
Sand Dunes,
Badlands, Mud Springs,
and Mima Mounds
Chapter 9
Sand Dunes
Stretching across the intermountain basins of central
Wyoming are elongated mosaics of active and stabilized
dunes (fig. 9.1). Sand dunes create a dramatically different environment in the semi-arid lowlands, primarily
because they provide a better water supply for plants,
due to the inverse texture effect (explained in chapter 3)
and because the shifting sand creates special problems
for plant establishment and growth. Consequently, the
flora of the dunes is considerably different from that of
the surrounding shrublands. Contributing to the biological diversity in some areas are small ponds that provide a source of water during the dry summer for plants
and animals that might not otherwise be able to survive
there.
Dune location depends on wind direction, a source
of sand, and barriers to sand movement. For example,
large deposits of sand are found on the westerly, windward foothills of the Ferris Mountains north of Rawlins,
the Laramie Mountains near Laramie, the Medicine Bow
Mountains in North Park, Colorado (east of Walden),
and the Sangre de Cristo Mountains (Great Sand Dunes
National Park) farther south in Colorado. Elongated
dunes are found in wind corridors where sand has been
funneled by the topography, such as the Killpecker
Dunes northeast of Rock Springs, and the dune fields
west and east of Casper.
The origin of such large volumes of sand is one of
the first questions asked. The answer lies in the physical
weathering of various kinds of rocks over long periods
142
and the power of water and wind to sort and move the
resulting particles. The sand grains (0.052.00 mm in
diameter) are gradually separated from coarser gravel
and much of the finer silt and clay. The sand then accumulates along creeks and rivers in the intermountain
basins. For example, most sand in the Killpecker Dunes
originated from the appropriately named Big Sandy
and Little Sandy creeks that flow from the Wind River
Mountains.1 These creeks were much larger when the
glaciers were melting, bringing large volumes of sand to
the area. Subsequently, westerly winds in the Holocene
blew much of this sand toward the east, across the Great
Divide Basin to the Ferris and Seminoe mountains.
Smaller silt and clay particles, known as loess, were
dispersed farther eastward onto the Great Plains. The
presence of different-aged sand grains in now-dormant
parts of the Killpecker Dunes suggests that sand movement occurred in this area at various times over the past
20,000 years, probably most often during dry periods,
when the sand was more easily blown out of the creek
beds.2
The kinds of plants growing on dunes depend on
the degree of sand stabilization as well as on temperature, moisture, and the amount of organic matter
in the sand. Only a few species can survive on shifting dunes in Wyoming. They include blowout grass,
sand lovegrass, prairie sandreed, sandhill muhly, rusty
lupine, blowout penstemon, alkali wildrye, Indian
ricegrass, and scurfpea. 3 The coarser soils of dune tops
typically have different species than the finer-textured
Sand Dunes, Badlands, Mud Springs, and Mima Mounds 143

Fig. 9.1. Location of the primary
sand dunes and badlands in
Wyoming. Some dunes shown
on this map are stabilized by
vegetation. Cartography by Ken
Driese.
soil between dunes.4 Alkali cordgrass is common where
Notably, the tallest, most vigorous stands of basin big
wetlands form between the dunes. Missing from Wyo-
sagebrush in the region occur on stabilized dunes, such
ming dunes is American beachgrass, commonly found
as some parts of the Killpecker and Ferris dunes where
on inland dunes to the east and along the Atlantic and
these shrubs grow to 6 feet tall or more. The tallest sage-
Pacific coasts, and yellow wildrye, a common colonizer
brush is typically on the leeward sides of dunes, where
in the dunes of southeastern Idaho.5
snow accumulates. Such sagebrush woodlands provide
Succession in dune ecosystems is primarily a process
important cover for the elk that live in the area.
of soil stabilization (figs. 9.29.4). The continued growth
However, stabilized dunes are subject to the same
of pioneer species gradually leads to more organic
kinds of periodic disturbances important elsewhere
matter in the sand and more soil binding by the root
drought, fire,7 burrowing, heavy grazing, and human
systems, which include the fungal filaments of mycor-
traffic. Such disturbances lead to a decline in plant
rhizae. The mere presence of the plants causes reduc-
vigor, subjecting small areas of the dune to rapid ero-
tions in wind velocity, diminishing the extent of sand
sion. The resulting blowout, created in a few days or
movement. This stabilizing process continues as plant
weeks, is eventually stabilized again through the per-
cover increases; eventually the entire dune is covered
sistence and expansion of the pioneer species. Signifi-
with plants. During stabilization, many other grasses,
cantly, the suppression of fires is thought to have greatly
forbs, and shrubs become common, including antelope
increased the proportion of some dune fields that are
bitterbrush, big sagebrush, rabbitbrush, silver sage-
now stabilized, such as in the Nebraska Sand Hills.8
brush, spiny hopsage, and others. Some plants create
Destabilization occurs with drought, heavy grazing, or
favorable microenvironments for others that could not
the recreational use of all-terrain vehicles.
grow on the dune initially. The pioneer species become

6
less common, but they persist in the community.
Compared with surrounding grasslands and shrublands, some parts of the dune environment are more
Fig. 9.2. (above) Killpecker Dunes,

located northeast of Rock Springsone
of the largest active dune fields in North
America. Dune colonizers in the foreground include Indian ricegrass, alkali
wildrye, slimflower scurfpea, veiny
dock, rusty lupine, skeletonplant, basin
big sagebrush, rabbitbrush, and spiny
hopsage. Elevation 7,000 feet. Photo by
Ken Driese.
Fig. 9.3. (left) Early stages of stabilization in the Killpecker Dunes, caused
here by veiny dock (Rumex venosus).
Aswith other dune-colonizing species,
such as slimflower scurfpea and alkali
wildrye, this plant spreads by rhizomes.
attracted to the dunes for this reason, including deer, elk,

waterfowl, and wild horses. The ponds also provide habitat for various aquatic organisms, including salamanders,
spadefoot toads, and freshwater shrimp. Because dunes
are an oasis in the semi-arid basins and plains, livestock
may concentrate there, creating management problems
similar to those encountered in riparian landscapes.
Indigenous people were also attracted to the dunes,
undoubtedly because of water and food availability.11
Artifacts, estimated at more than 10,000 years old, have
been found in the Killpecker Dunes and elsewhere in the
region, commonly in association with bison bones.
Although sand dunes are favorable for the growth of
established plants, shifting sands provide a poor substrate for seedling establishment. Moreover, established
plants are easily buried. Pioneer dune plants (such as
blowout grass and scurfpea) survive because their rhizomes grow up or down quickly enough to maintain
appropriate depths for the species. Aside from that, root
growth occurs in the direction of moisture availability.
As important, the rhizomes have considerable energy
stored in the form of carbohydrates, which enables the
growth of numerous shoots when the buds begin to
grow, all characteristics that diminish the probability
Fig. 9.4. Slimflower scurfpea. Note the rhizome and prominent taproots that have been exposed by wind erosion.
that the entire plant will be buried to excessive depths.

Roots also grow from the buried vertical stems of some
plants (adventitious roots), even on seedlings.12
Dune plants encounter other problems as well, such
favorable for the survival of many plants and animals.
as sandblasting of their leaves and stems. Not surpris-
The annual production of new plant biomass can be
ingly, most of them are herbaceous, with shoots that
more than double that of the adjacent sagebrush steppe.9
require maintenance for only a few months. Also, some
Even with limited precipitation, the rapid infiltration
species produce waxy coatings that reduce abrasion by
of water through the coarse sand provides good water
blowing sand. Unlike in the coppice dunes discussed
storage, and commonly snowdrifts on the lee sides of
in chapter 7, woody plants invade only after the dunes
dune crests are buried by drifting sand in the spring. It
have become partially stabilized and abrasion is less
is common to find buried snow in midsummer, which
severe. Sand abrasion is not all bad for the plants, as
provides a source of water for ponds and perennial
some of them have seeds that will not germinate until
streams as it melts.10 The landscape pattern is diversified
seed coats are abraded. Other adaptations contribute
further by the shrubs and small trees that occur along
to water-use efficiency in this environment, where the
such streams, for example, basin big sagebrush, com-
temperature at the soil surface can be high. Many dune
mon juniper, currant, Oregon-grape, Rocky Mountain
plants are water-efficient C 4 species, and a large propor-
juniper, water birch, Woods rose, and willow.
tion have both shallow and deep roots.13
Rapid infiltration reduces the time available for evap-
Extensive root systems with mycorrhizae are impor-
oration. This, along with buried snow that melts slowly
tant for obtaining nutrients, as sand is an infertile sub-
during the summer, contributes to the formation of small
strate. Also, two common dune colonizers in Wyoming
ponds between the dunes (fig. 9.5). Many animals are
are nitrogen-fixing legumes (scurfpea and lupine).14 As

Fig. 9.5. Small ponds sometimes form
between dunes, primarily because of
snowdrifts that melt slowly after being
buried by drifting sand. The colonizing plants in this photo, taken in the
Killpecker Dunes, are alkali wildrye and
slimflower scurfpea. Photo by Bonnie
Heidel.
the plant tissues grow and die, soil organic matter gradu-
turbance, however, the dunes eventually become sta-
ally accumulates. Some of it decomposes, providing
bilized and highly productivea classic example of
nutrients that, combined with water availability, create
natural succession. The dunes of central Nebraska have
one of the best environments for plant growth in the
been one of the most productive grasslands on the
intermountain basins. That may be short-lived, however,
Great Plains, providing forage even during extended
as the next blowoutwhether caused by an extended
droughts, such as occurred in the 1930s.
drought, fire, or other disturbancemay bring the
Most sand dunes in Wyoming occur at higher eleva-
organic matter to the surface, where it is widely dispersed
tions than in adjacent states, and consequently they
by the wind. Alternatively, the blowing sand may cover
have a relatively cold climate and short growing sea-
well-developed soils. Known as paleosols, such buried
son. With current predictions for increased warming
soils are useful for interpreting climate history.
and drought, reactivation of dunes over large areas
A newly activated dune generally has a deficiency
seems likely and may have already begun. Numerous
of nutrients on the surface. Without drought or dis
studies have shown that drought causes a rapid decline
in plant cover on stabilized dunes, which leads to the
ming, fossils of crocodiles, primates, and tapirs date
dune becoming reactivated.15 The effect of drought is
back to the Cretaceous and early Tertiary periods, indi-
hastened if fires, intensive grazing, disruption by off-
cating a subtropical climate. Some badlands are now
road vehicles, or other disturbances occur at about the
protected because of their value for paleontological
same time.16 Thus far, few invasive plants have been
studies and the number of people fascinated by such
observed in Wyoming dunes, but that is not true at
places. The most notable example is Badlands National
lower elevations.
Park in western South Dakota.
17
Plants are sparse in badland landscapes. The few
Badlands
that can be found also occur in adjacent grasslands

and shrublands, such as big sagebrush, bottlebrush
Some parts of the lowlands, known as badlands or breaks,
squirreltail, Gardners saltbush, juniper, shadscale, sil-
are nearly devoid of vegetation because of naturally high
ver sagebrush, tufted evening primrose, and western
rates of erosion associated with retreating escarpments
wheatgrass. Halophytes are common. In ravines or near
(figs. 9.6 and 9.7). Such areasfor example, Adobe Town,
seeps, basin big sagebrush, basin wildrye, greasewood,
Grizzly Buttes, Hells Half Acre, Honeycomb Buttes, the
narrowleaf cottonwood, and species of willow may be
Powder River Breaks, and Wind River Badlandsare some
present. The amount of plant cover varies considerably
of the most fascinating landscapes in the region. Badland
with elevation, substrate, precipitation, and topographic
topography, sometimes angular and sometimes rounded,
position. On steep slopes, erosion is a nearly continuous
occurs in arid climates on shales and mudstones. The
process, which prevents plant establishment.
clay-rich soils expand and contract with wetting and
Because of the abrupt, finely dissected topography,
dryingconditions that usually prevent plant establish-
the fauna of badlands can be more unusual than the
ment and facilitate erosion. Flash floods are common
flora. Hawks, eagles, swallows, and swifts find suitable
because of slow infiltration during thunderstorms.
nesting sites on exposed cliffs, and larger animals use
Because badlands are formed by rapid erosion, they
the badlands year-round, finding shelter on leeward
are favorite destinations for fossil hunters. Each year
slopes on cold, windy days. The vegetation mosaic is
new specimens are exposed on the surface. In Wyo-
unusual as well. In the Powder River Breaks of north-
Fig. 9.6. Honeycomb Buttes in

the Jack Morrow Hills northeast of Rock Springs, one of
numerous tracts of badlands
found in Wyoming. Here the
rocks are shales, claystones,
and sandstones of the Green
River and Wasatch formations
(Eocene). Elevation 6,888 feet.

Fig. 9.7. Skull Rim in a desert
wildland known as Adobe
Town, part of the Washakie
Basin southeast of Rock
Springs and southwest of
Rawlins. The badlands in this
area are carved from claystones and sandstones of the
Washakie Formation (Eocene).
ern Wyoming and southeast Montana, seven types of
surrounded by mudflats or playas with only a few plants
shrublands and woodlands have been identified, based
(fig. 9.8). The cones seem to provide better conditions
on their dominant plants: shadscalebig sagebrush,
for plant growth than the flats do. At one time, a slurry
big sagebrushshadscalewestern wheatgrass, big sage-
of mud oozed from their summits and then dried.19
brushwestern wheatgrass, greasewood, skunkbush
The geologist and explorer Ferdinand Hayden was
sumacwestern wheatgrass, Rocky Mountain juniper
the first scientist to discover these unusual landforms
western wheatgrass, and ponderosa pinejuniper. In
when he visited the area in 1877. According to his
addition, on mesa tops capped by erosion-resistant
report, the mounds at that time had pools of muddy
scoria, there are small patches of mixed-grass prairie.
water at the top. Observing that a bubble of gas periodi-
Greasewood is found near seeps on some slopes. Plant
cally rose to the surface of the pools, he and his men
cover is sometimes contoured in narrow bands that par-
conducted an experiment:
allel beds of gray clay, buff-colored silt, and lignitean

indication of strong edaphic control.18
Ponderosa and limber pine are found in badlands,
along with Utah and Rocky Mountain juniper, but only
on escarpments that are comparatively resistant to erosion. Elsewhere erosion occurs before mature trees
have time to develop. Landslides can cause significant
changes.
A rifle-ball shot down vertically into one of the

openings produced a sudden eruption of the whole
mass. Water and mud were thrown to a height of
about 10 feet, covering the luckless experimenter
from head to foot. From a safer distance the trial was
several times repeated and almost always followed
by the same result. . . . Crude as this test may be, it
shows the presence of gas at some depth, held there
under mechanical pressure. 20
Mud Springs
Today, the mud springs appear to be dormant. A
One of the most unusual features of the Great Divide
possible cause is the pumping of groundwater from the
Basin is the small area of mud springs found just east of
Great Divide Basin for livestock and industry, which
Circle Bar Lake (one of the Chain Lakes north of Wam-
could be relieving some of the water pressure that
sutter). These conical mounds are 315 feet high and are
was important for the creation of these springs. How-

Fig. 9.8. Mud springs in the Chain
Lakes area, located in the lowest part
of the Great Divide Basin, are currently
dormant. Plants include greasewood
and shadscale. A playa with little or
no vegetation surrounds the conical
mounds. Elevation 6,500 feet.
ever, it is possible that the mounds are rejuvenated by
Mima mounds raise two questions: What is the
periodic rather than continuous eruptions. Hayden
effect of the mounds on plant and animal distribu-
observed deep pools of muddy water in the area near
tion, and how did they originate? The effect on plants
the mounds, covered by a thick layer of crusty silt that
can be subtle, with no obvious differences between
gave the appearance of solid ground. He wrote, Innu-
mound and intermound vegetation. Elsewhere, the
merable bones of animals, who here sought to quench
vegetation on the mounds is taller or composed of
their thirst, provide the treacherous character of the
different species. For example, the mounds may have
soil. A nearby pond was christened Death Lake by
more big sagebrush and higher plant growth rates than
the explorers.
between mounds. 22 Some investigators have noted

that burrowing mammals, such as pocket gophers and
Mima Mounds
ground squirrels, are more common on the mounds.

Through their burrowing, the mammals create soils
A peculiar feature of some lowland grasslands and
that have a reduced bulk density (weight per unit vol-
shrublands in Wyoming are hundreds of closely spaced
ume), enhanced infiltration rate, and improved nutri-
mounds that are 1224 feet in diameter and usually less
ent availability. Vegetation differences can be expected
than 2 feet high (figs. 9.9 and 9.10). They are common on
when mound and intermound soils are different, but
the east slope of the Laramie Mountains (west of Chey-
whether the differences are due to the soil or associated
enne), in the Laramie Basin southwest of Laramie, and
animals is still unknown. Unfortunately, cheatgrass
in the Shirley Basin north of Medicine Bow (all at low
has been found on the tops of some mima mounds in
elevations). Equally puzzling mounds occur in numerous
the Laramie Basin.23
other places in North America and in Argentina, Kenya,
The more perplexing question about mima mounds
and South Africa.21 Known as prairie mounds, pimple
pertains to their origin. There are various hypotheses
mounds, and biscuit land, they are most commonly
that involve wind deposition (coppice dunes), erosion,
referred to as mima or mima-like mounds, because they
frost heaving, groundwater flows, burrowing animals,
were first described on the Mima Prairie in western
and even seismic activity. The erosion hypothesis sug-
Washington.
gests that the tops of mounds were once the location of
Fig. 9.9. (above) Mima mounds in the Laramie Basin. The vegetation is high-elevation mixed-grass prairie, dominated by
junegrass, Sandberg bluegrass, Indian ricegrass, needle-andthread grass, blue grama, fringed sagewort, and Hoods phlox.
Fig. 9.10. (left) Aerial view of mima mounds in the Laramie
Basin. In this area the mounds are partially on the broad
floodplain of the Big Laramie River (see chapter 17). Elevation
7,250 feet.
a stabilizing tree or tall shrub, with the soil in between

more susceptible to erosion. After the trees or shrubs
died, the mound persisted, though possibly with some
smoothing by wind. Indeed, the mound distribution
patterns are often similar to tree distribution patterns
in woodlands and savannas, but there is little or no supporting evidence for this explanation.
The frost hypothesis has received more attention than
the deposition and erosion hypotheses in Wyoming.
From the air, the mound pattern gives the impression of
sorted polygons that are common in the arctic tundra
(see fig. 9.10), an observation that suggests the mounds
were formed about 20,000 years ago when there were
glaciers in the mountains and the climate was colder.
Indeed, Wyoming intermountain basins at that time had
permafrost with relatively little snow cover.24 In such
environments, soil cracks 3 feet deep or more were cre-
ated by contraction caused by either freezing or desicca-
published in 1954, R. J. Arkley and H. C. Brown proposed
tion. These cracks filled with windborne soil particles,
the following explanation:
and today their approximate form and depth are apparent when the soil profile is exposed. Depending on the
cause of the cracks, the distinct patterns in the soil profile are referred to as fossil ice wedges (see fig. 2.8) or sand
wedge relics, the latter having been formed in desiccation cracks.
University of Wyoming soil scientists Lowell Spackman and Larry Munn dug trenches through mima
mounds in the Laramie Basin and determined that fossil ice wedges were a key factor in mound formation. 25
They concluded that the mounds could be attributed
to cryostatic pressure created from water entrapped
between a layer of permafrost or bedrock and a downward-freezing frost layer from the surface. Pressure was
ultimately released through planes of weakness created
The method by which the [pocket] gopher accumulates soil into a mound is explained by his tendency
to place his nest in a well-drained spot where the
soil is deepest; thus generation after generation of
gophers may keep building nests near the crest of
any high spot in the land surface, or over a window
in a hardpan. When a gopher is tunneling, he moves
the soil beneath his body, and forces it backward to
a surface opening already established. . . . Thus, over
a long period of time, the gopher, by digging outward from his nest, tends to move the soil toward it.
. . . The mound rises very gradually over a considerable period of time and perhaps many generations of
gopher occupation.28
by large fossil permafrost sand wedges. They found no
Others, primarily ecologist George Cox and his asso-
evidence for the burrowing rodent hypothesis. Previous
ciates, concluded that burrowing animals are the uni-
research had suggested that the mima mounds in Wash-
versal cause of mima mounds everywhere.29 Cox wrote,
ington were also formed in association with permafrost
Fossorial rodents are the builders of Mima mounds,
and ice wedges.26
the largest and most widespread landscape features pro-
Working in the same area, soil scientist Richard
duced by any mammal other than man. Cox reviewed
Reider and his associates proposed a mound-forming
studies indicating that, in addition to pocket gophers,
mechanism that involved low-pressure groundwater
mound building is caused by ground squirrels; badgers;
flow.27 They dug trenches through the mounds as well,
toads; fire ants; and, in Kenya, mole rats.
finding that all of them had (1) shallow impermeable

bedrock, (2) a thin layer of alluvial gravel on top of the
The rationale of Cox, suggesting that burrowing created
bedrock, and (3) well-developed soil at the ground sur-
all mounds worldwide, still needs to be reconciled with
face. They also observed that the mound soil was lens-
the evidence obtained supporting other hypotheses in
shaped in cross section and that it had concentrations
the Laramie Basin. A new dimension to the debate was
of sodium in patterns suggesting a swirling ground
added by geologist Andrew Berg, who concluded that
water flow. Reider and his associates concluded that the
most mima mounds are the result of seismic vibrations
explanation probably lies in blending their explanation
in Earths crust in areas with unconsolidated fine sedi-
with Spackman and Munns cryostatic hypothesis, but
ments on a relatively rigid, flat substratum. This led to
that no other explanation for mima-mound formation
a spirited exchange of viewpoints.30 Berg acknowledged
seemed applicable to their study area. They also advised
that the seismic hypothesis requires further work to
that this hypothesis should not be applied to all mima-
substantiate, but that it points the way to the solution
like mounds.
of a geologic enigma that has been with us for more
For biologists, the most intriguing hypothesis in
than 100 years. The most recent analysis, by Jennifer
volves burrowing animals, usually pocket gophers. In
Burnham and Donald Johnson in 2012, suggests there
one of the earliest papers on the origin of mima mounds,
could be more than one explanation.31
Part Four Foothills and
Mountains
Escarpments and
the Foothill Transition
Chapter 10
The lowlands of Wyoming are interrupted by a dozen
than the adjacent ecosystems. Escarpments appear as
or more mountain ranges (see figs. 1.11.2), all flanked
islands in a sea of sagebrush or grassland, though they
by foothills formed largely from glacial moraines and
can be narrow as well when associated with long ridges
eroding sedimentary rocks. Away from the mountains
(fig. 10.1).1
are numerous escarpments. Both provide distinctive
Foothills and escarpments are important habitats
habitats for plants and animals, adding immeasurably
for big game and other species because of their abrupt
to the aesthetic appeal of the region. The foothills are
topography. Windblown snow accumulates on leeward
comparable to the riparian zone, covering a small, nar-
slopes, which provides a more reliable source of water.
row, elongated portion of the landscape and providing
The lee slopes also serve as windbreaks for big game in
a more moderate environment at some times of the year
the winter, and windward and south-facing slopes are

Fig. 10.1.The coarse, rocky
soil of Cretaceous sandstone
escarpments in the Powder
River Basin supports little
bluestem, ponderosa pine,
Rocky Mountain juniper,
skunkbush sumac, and other
foothill plants. Wyoming big
sagebrush, blue grama, and
western wheatgrass dominate
the shrubland on deeper,
fine-textured soils in the foreground. Elevation 5,000 feet.
155
156 Foothills and Mountains
usually snow-free, providing easier access to forage. In
andin one or two localities in extreme south-central
such places, shelter and forage are available nearby
Wyomingpinyon pine or Gambel oak. Windswept
unlike the mountain forests, where there is abundant
ridges with shallow soils and little snow accumulation
shelter but little winter forage (owing to continuous
have bluebunch wheatgrass and a variety of cushion
snow cover), or the surrounding lowlands, where both
plants (see fig. 8.9).

In general, the foothill-escarpment mosaic is patchy,
shelter and forage may be hard to find.

The vegetation mosaic of escarpments and foothills
a reflection of abrupt changes in rock type, soil depth,
consists of shrublands, grasslands, woodlands, and
snow accumulation, microclimate, slope, and the in
windswept ridges (fig. 10.2). Shrublands dominated
verse texture effect. Vegetation banding is sometimes
by mountain-mahogany are abundant on many rock
observed, such as in the Centennial Valley, west of Lara-
outcrops with little or no soil. On deeper soils where
mie, where different kinds of vegetation are associated
snow accumulates, the common plants are antelope
with bands of sandstone, limestone, shale, siltstone,
bitterbrush, mountain big sagebrush, serviceberry,
and claystone. 2 Similarly, elongated aspen groves can
skunkbush sumac, and snowberry (table 10.1). Black
be found at the contact between impervious granite in
sagebrush or threetip sagebrush commonly occur on
the mountains and the overlying sedimentary strata of
windswept ridges or plateaus with shallow soils, and
the foothills, where runoff from the mountain encoun-
tufted hairgrass and other species sometimes are
ters deeper soils (fig. 10.3). Trees also grow on ridges,
found in leeward arroyos where unusually deep snow-
sometimes because the groundcover there is usually in
drifts develop, sometimes called nivation hollows (see
sufficient to fuel a fire. Thus, trees have a greater chance
fig. 7.8). Foothill grasslands support a sparse cover of
of becoming established than in the adjacent grassland
bluebunch wheatgrass along with other grasses and
or sagebrush steppe, where the competition for water
forbs. The woodlands have varying densities of juni-
from grasses and forbs is more intense and where the
per, limber pine, ponderosa pine, aspen, Douglas-fir,
time between fires is shorter.3
2800
Limber
pine
woodland
ELEVATION (feet)
8000
2600
2400
Juniper
woodland
7000
Basin or
Foothill
grassland
6000
5000
Lodgepole pine,
Douglas-fir, or
Ponderosa pine
forest
Saltbush
desert
shrubland
Sagebrush
desert
shrubland
Sagebrush
steppe
Mixedgrass
prairie
4000
Low
2200
Mixedfoothill
shrubland
or
Ponderosa
pine
savanna
Greasewood
shrubland
Woody
Riparian
draw
woodland
Moderate
High
2000
1800
ELEVATION (m)
9000
1600
1400
1200
WATER AVAILABILITY
Fig. 10.2. Distribution of various kinds of foothill grasslands,
shrublands, and woodlands in relation to gradients of water
availability and elevation in the Bighorn Basin, near the
Bighorn Mountains (Bighorn Canyon National Recreation

Area). Vegetation types that occur above and below the foothills are shown for comparison.
Escarpments and the Foothill Transition
Fig. 10.3. Aspen groves on the east slopes of Centennial

Ridge, in the Medicine Bow Mountains, occur in most ravines
and along the contour where the impermeable granite meets
the permeable soils that have developed on sedimentary
rocks. Sagebrush steppe, dominated by threetip sagebrush,
Plant Adaptations and Vegetation Dynamics
occurs on windswept ridges between the ravines. Douglas-fir

and limber pine occur just above the aspen. Forests at higher
elevations are dominated by lodgepole pine, Engelmann
spruce, and subalpine fir.
quently browsed by deer, especially in the winter. Notably, the shrubs respond by growing lateral branches.
Mountain-Mahogany Shrublands
The more heavily the shrubs are browsed, the more
One of the more conspicuous foothill shrublands is

dominated by mountain-mahogany (Cercocarpus spp.).
This shrub forms dense thickets on rocky or shallow
soils from low elevations on the western Great Plains
(~4,500 feet) up to elevations of about 8,000 feet in
the mountains (fig. 10.4; see table 10.1). Two species
of Cercocarpus occur in Wyoming: the deciduous true
mountain-mahogany in the southern Black Hills and
across the southern half of the state, and the evergreen curlleaf mountain-mahogany in the foothills
of the Bighorn Mountains and also to the west and
south (fig. 10.5). There is little overlap in the distribution of the two species. Both grow on a variety of
bedrock types, including sandstones, limestones, and
spiny the branches become, making further browsing

more difficult. In this way the plants develop their own
defense mechanism.
Some ecologists have observed expansion of curlleaf
mountain-mahogany into adjacent plant communities
during long periods without fire, suggesting that frequent fires historically restricted mountain-mahogany
to rocky sites.6 Curlleaf mountain-mahogany can live
150 years. The shrubs spread by layering or seedling
establishment and can sprout after fires.7 Large seed
crops are common, though often not in consecutive
years.
Juniper Woodlands
shales.4 The abundance of both species is undoubtedly
A characteristic feature of some foothills and escarp-
facilitated by their ability to fix nitrogen; and perhaps
ments are picturesque woodlands dominated by long-
because of their more nitrogen-rich foliage, they are fre-
lived Utah juniper or Rocky Mountain juniper (figs. 10.6
157
Table 10.1.Characteristic plants found in the different kinds of foothill vegetationa
Common name
Latin name
MountainMixed
mahogany
Juniper
foothill
Aspen
Conifer
Oak
shrubland woodland shrubland woodland woodland woodland
Woody
draws
TREES
Aspen
Populus tremuloides
Boxelder
Acer negundo
Bur oak
Quercus macrocarpa
Quercus gambelii
Gambel oakc
Green ash
Fraxinus pennsylvanica
Douglas-fir
Pseudotsuga menziesii
Limber pine
Pinus flexilis
Ponderosa pine
Pinus ponderosa
SHRUBS
American plum
Prunus americana
Antelope bitterbrush
Purshia tridentata
Black sagebrush
Artemisia nova
Mountain big
sagebrush
Artemisia tridentata
ssp. vaseyana
Silver sagebrush
(plains)
Artemisia cana ssp.

cana
Wyoming big
sagebrush
ssp. wyomingensis
Chokecherry
Prunus virginiana
Common juniper
Juniperus communis
Rocky Mountain
juniper
Juniperus scopulorum
Utah juniper
Juniperus osteosperma
Curlleaf
mountain-mahogany
Cercocarpus ledifolius
True (alderleaf )
mountain-mahogany
Cercocarpus montanus
Snowbrush
ceanothus
Ceanothus velutinus
Douglas hawthorn
Crataegus douglasii
Douglas rabbitbrush
Chrysothamnus
viscidiflorus
Rubber rabbitbrush
Ericameria nauseosa
Saskatoon
serviceberry
Amelanchier alnifolia
Utah serviceberry
Amelanchier utahensis
Skunkbush sumac
Rhus trilobata
Utah snowberry
Symphoricarpos oreophilus var. utahensis
MountainMixed
mahogany
Juniper
foothill
Aspen
Conifer
Oak
shrubland woodland shrubland woodland woodland woodland
Woody
draws
Common name
Latin name
Western snowberry
Symphoricarpos
occidentalis
Wax currant
Ribes cereum
Woods rose
Rosa woodsii
GRASSES
Basin wildrye
Leymus cinereus
Blue grama
Bouteloua gracilis
Bluebunch
wheatgrass
Idaho fescue
Festuca idahoensis
Spike fescue
Leucopoa kingii
Little bluestem
Schizachyrium
scoparium
Oatgrass
Danthonia spp.
Prairie junegrass
Koeleria macrantha
Sideoats grama
Bouteloua curtipendula
Arrowleaf
balsamroot
Balsamorrhiza
sagittata
Creeping barberry
(Oregon-grape)
Mahonia repens
Field (mouse-ear)
chickweed
Cerastium arvense
Hairy false
goldenaster
Heterotheca villosa
Hookers sandwort
Arenaria hookeri
Lupine
Lupinus spp.
Prairie (fringed)
sagewort
Artemisia frigida
Pricklypear cactus
Opuntia polyacantha
Purple locoweed
Oxytropis lambertii
Pussytoes
Antennaria spp.
Sulfur-flower
buckwheat
Eriogonum umbellatum
Western yarrow
Achillea millefolium
var. occidentalis
FORBS
b
c
Bur oak is found only in the Black Hills.
Gambel oak is found only in the Sierra Madre.
Fig. 10.4. (above) Curlleaf mountain-mahogany and a few

ponderosa pine grow on shallow, rocky soils in this area on
the east side of the Bighorn Mountains. Mixed-grass prairie
occurs on deeper soils. True mountain-mahogany is found in
similar habitats in the southern and eastern parts of Wyoming (see chapter 17).
Fig. 10.5. (right) True mountain-mahogany is easily distinguished from curlleaf mountain-mahogany. Drawing by Judy
Knight.
and 10.7). Depending on soil characteristics, associated

plants can include black sagebrush, big sagebrush, antelope bitterbrush, mountain-mahogany, limber pine,
and ponderosa pine (see table 10.1). Rocky Mountain
juniper is common in eastern Wyoming and the north-
tions. However, unlike mountain-mahogany, juniper is
ern Great Plains, where summer precipitation is higher
more capable of expanding into adjacent rangelands on
than farther west. Usually it occurs in ravines or on
deeper soils. Differences in the distribution of juniper
steep slopes, along with yucca, skunkbush sumac, and
and mountain-mahogany may be the result of differ-
ponderosa pine. Utah juniper grows on escarpments in
ences in microclimate, relative ability to spread into
the more arid basins of western Wyoming, for example,
adjacent habitats, or soil-bedrock features that have not
in the Wind River Canyon and Bighorn Basin.
yet been studied in sufficient detail.9 Juniper distribu-
The cause of juniper presence or absence is a puz-
tion in the Bighorn Basin ranges from about 3,600 to
zle, because both species are absent from many habi-
6,000 feet, overlapping with but generally lower than
tats that seem suitable. On the upper or northerly edge
mountain-mahogany shrublands. The lower limits in
of its range, such as in Wyoming, the junipers seem
specific areas seem to be where shallow rocky soils meet
to survive only in warmer thermal belts, where frosts
deeper alluvial or colluvial soils. The upper limit is prob-
are less frequent at night.8 Like mountain-mahogany,
ably controlled by a still-undetermined climatic factor.10
both species are found on a variety of geologic forma-
Most shrublands at low elevations in the Bighorn Basin
Fig. 10.6. (above) Juniper woodland occurs on escarpments

with coarse soils, such as in this area near Bates Creek in the
foothills of the Laramie Mountains, south of Casper. Rocky
Mountain juniper is common in the eastern half of Wyoming;
Utah juniper is common in the western half. Both species are
found near Lander.
Fig. 10.7. (right) The small, scale-like leaves of Rocky Mountain juniper are longer and narrower than the leaves of Utah
juniper. Drawing by Judy Knight.
are dominated by Wyoming big sagebrush, black sagebrush, and broom snakeweed, but curlleaf mountainmahogany commonly is found in the foothills of the
Bighorn Mountains, sometimes with juniper.
Juniper expansion into adjacent grasslands or shrublands, where juniper was previously absent, is one of
that have formed by juniper expansion into former
the most often-discussed ecological topics in the inter-
grasslands or shrublands during the twentieth century
mountain West. A related phenomenon is the estab-
tend to be found on relatively deep soils in regions hav-
lishment of new junipers in formerly sparse juniper
ing summer-dominated precipitation patterns. In con-
woodlandsknown as infilling. Expansion and infill-
trast, the older, persistent woodlands are most common
ing of juniper and pinyon during the twentieth cen-
on shallow, rocky soils and in regions where precipita-
tury have been documented in many areas throughout
tion comes mostly in the winter.12
11
the West. However, the process may be driven by more
What are the ecological processes that drive juni-
than one ecological mechanism, and many woodlands
per expansion and infill? Two mechanisms that have
are stable communities that have persisted with little
received much attention are the effects of livestock
change for hundreds or thousands of years. Woodlands
grazing and fire exclusion. Using tree-ring analysis,
161
rangeland ecologist W. J. Waugh found that juniper
establishment in many parts of the West.19 Increasing
in the Bighorn Basin had been restricted to limestone
atmospheric carbon dioxide may also be facilitating tree
outcrops until about 10 years after the introduction of
growth by increasing water-use efficiency.20 For what-
large herds of cattlea time that also marks the initia-
ever reason, woodland expansion and infill are occur-
tion of fire exclusion. Juniper has since expanded onto
ring in many parts of the West, though the causes may
adjacent sites. Waugh noted that more than 90 percent
not be the same everywhere.
of the juniper seedlings were becoming established
Regardless of the mechanism, herbaceous plant
under sagebrush, which apparently was functioning as
growth is greatly reduced with the establishment of
a nurse plant, and he hypothesized that an increase in
juniper in a grassland or shrubland, owing to more
sagebrush cover following livestock grazing had created
competition for water, light, and nutrients. For a time,
a more favorable environment for juniper invasion.
fires may become less frequent if the shrubs are widely
13
Another important indirect effect of livestock graz-
spaced, thereby perpetuating conditions favorable for
ing is to reduce the fine fuels available for fire. Fires
juniper infilling. However, eventually fuel continuity
at 30- to 60-year intervals formerly prevented juniper
increases and fires become more probable. Prescribed
expansion in parts of the Great Basin of Utah, and
fires are often used to reduce juniper density, but man-
observers have noted that burning can reduce the abun-
agers know that an undesired consequence could be
dance of sagebrush and juniper.14 Neither juniper nor
the invasion of introduced plants, such as cheatgrass
big sagebrush reproduce by sprouting, and woodlands
(see chapter 7). Another concern is that burning could
with old junipers are often restricted to rocky ridges,
reduce the amount of food and thermal cover avail-
where fires are less frequent. Heavy livestock grazing
able for deer and other wildlife. Animals stay warmer at
could also reduce the rate of evapotranspiration from
night under the juniper canopy because of less exposure
the rangeland ecosystem, which would reduce competi-
to the cold sky. Daytime shade during the summer also
tion for water from other plants. Moreover, in combi-
is important.
nation with an unusually wet period, livestock grazing
In northwestern Colorado, on the Colorado Plateau,
could enable the establishment of a large number of
extensive woodlands are dominated by pinyon pine
juniper seedlings. However, this scenario may not apply
and Utah juniper. 21 The two species coexist, but pin-
everywhere, as other observers have observed that
yon pine commonly is found at slightly higher eleva-
drought, in combination with grazing and fire exclu-
tions than Utah junipersuggesting that the juniper
sion, favors juniper invasion.15
is more drought tolerant than the pine. Pinyon pine is
Although fire exclusion and grazing are important
found in only one location in Wyoming, near Flaming
in some places, some young juniper stands represent
Gorge Reservoir in the foothills of the Uinta Mountains.
natural recovery by seedling establishment following
Rocky Mountain juniper is common, often occurring
past fires or other disturbances of mature woodlands.16
with limber pine and ponderosa pine in the foothills. 22
Near the edges of the current ranges of both species,

pinyon pine and the two junipers may still be migrating
into suitable habitats from their Pleistocene refugia. For
Ponderosa PineLimber PineDouglas-fir Woodlands
example, some juniper stands in Wyoming may repre-
Ponderosa pine, limber pine, and Douglas-fir occur com-
sent the first generation to colonize the sites since the
monly in the foothills and on escarpments (fig. 10.8).
last glacial advance.
Infill is also occurring in many
Growing in association with grasslands, juniper, and
stands that contain no evidence of past fire, that is, no
various kinds of shrubland, some of the tree-dominated
basal fire scars on living trees and no charred wood.18
woodlands have been increasing in density and expand-
Some of these woodlands have also been grazed only
ing their rangepossibly as a result of fire exclusion,
lightly or not at all. In such places, fire exclusion and
although climate conditions in the twentieth century
grazing cannot be the drivers of infill. Climate variation
have been favorable for these tree species just as for juni-
probably is more important. Several warm, moist peri-
per and pinyon.23 Increasing densities of ponderosa pine
ods during the 1900s were especially favorable for tree
can pose problems for ranchers, primarily because the
17
Fig. 10.8.Escarpments and foothills on the eastern plains

typically have ponderosa pine woodlands, such as along the
Hat Creek Breaks, north of Lusk (Oligocene and Miocene
sandstones and claystones). This ridge is part of Pine Ridge,
which extends westward to north of Casper and eastward into

southern South Dakota. Mixed-grass prairie is also common,
sometimes with silver sagebrush and Wyoming big sagebrush.
additional trees can reduce the amount of forage and
Although ponderosa pine is often common on escarp-
increase evapotranspiration (thereby reducing stream-
ments in eastern Wyoming, where deep water percola-
flow), and also because the pine needles can be toxic if
tion is presumably possible, some geologic substrata
their livestock are hungry enough to eat them.
provide better conditions for tree growth than others.
24
In Wyoming, ponderosa pine is found primarily in
Wyoming ecologist Jim States found that the growth
comparatively warm areas with higher summer pre-
of ponderosa pine was best on Mowry shale, where the
cipitation. Such environments are found in the eastern
trees had wider and less variable annual ring widths in
half of the state, primarily in the Black Hills and Bear
their wood.27 Low annual variability in ring width sug-
Lodge Mountains, at lower elevations on the east and
gests that this marine shale, despite the variable climatic
north slopes of the Bighorn Mountains and the Lara-
conditions from year to year, provided a consistent envi-
mie Mountains, in a few localities around the Medi-
ronment for growth. In contrast, trees on the Pine Ridge
cine Bow and Seminoe mountains, and on lowland
and Cloverly sandstones had narrower and more variable
escarpments scattered throughout the Great Plains. 25
annual rings, even though they grew near the Mowry
Associated plants include skunkbush sumac, Rocky
shale and surely had the same climatic conditions.28 The
Mountain juniper, and two grassessideoats grama
association of some foothill species with fracture lines in
and little bluestem (see table 10.1). Ponderosa pine is
the bedrock can be striking (see fig. 3.3).
essentially absent from western Wyoming, perhaps
Limber pine sometimes occurs with ponderosa
because of a growing season that is currently too short
pine, but it is more typical where the climate is drier
and dry. It does occur in the foothills of the Uinta
and colder. Unlike ponderosa, it extends upward to the
Mountains of Utah (such as in Dinosaur National
alpine treeline. The species tends to be restricted to rocky
Park), where the elevations are lower and the Arizona
soils and ridges, where the seedlings have relatively
Monsoon provides slightly more summer precipitation
little competition from other plants.29 It is commonly
than in Wyoming. 26
found on the lee side of boulders, where the seedlings of
163
tains just west of Yellowstone National Park, and in

other parts of the Greater Yellowstone Ecosystem (see
chapter 15). Compared to the eastern foothills, a larger
proportion of the annual precipitation occurs in the
winter, which apparently favors Douglas-fir. The soils
typically are derived from sedimentary rocks and are
more fertile. 30 Limber pine is sometimes associated with
Douglas-fir in western Wyoming, such as in Fossil Butte
National Monument.
The dynamics of Douglas-fir woodlands is much the
same as for ponderosa pine woodlands growing in similar situations, commonly with expansion into adjacent
grasslands and shrublands during long fire-free intervals.31 In parts of Montana, scars on trees have revealed
that fires occurred every few decades before 1890, confining the trees to rocky sites or the lee sides of slopes,
where the fires burned less frequently. Comparable
results were found for ponderosa pine woodlands in
Devils Tower National Monument, where, from 1770 to
1900, the average number of years between fires was 14.
Since 1900, fires have been much less frequent, about
once every 40 years, owing to fire suppression and the
Fig. 10.9. Limber pine occurs on rocky escarpments from low
elevations near Pine Bluffs in the southeast up to the alpine
treeline of some mountains in eastern Wyoming. In northwestern Wyoming, whitebark pine is found in the alpine
zone. This photo, taken in the Laramie Range, shows widely
scattered limper pine in a high-elevation shrubland dominated
by threetip sagebrush, fringed sagewort, junegrass, mountain
muhly, and Sandberg bluegrass. Elevation 8,000 feet.
great reduction in fires ignited by humans.32
Mixed Foothill Shrublands

Sagebrush steppe extends into the foothills in many
areas, often intermingled with mountain-mahogany
shrublands and woodlands dominated by pines,
Douglas-fir, and juniper. Species composition changes
as the environment becomes cooler and relatively more
the now-mature trees would have been protected from
moist. Mountain big sagebrush becomes the domi-
excessive wind damage. Once established, however,
nant variety instead of Wyoming big sagebrush, and
limber pine occurs on some of the windiest sites in the
other shrubs are common, namely, antelope bitter-
region (fig. 10.9). Indeed, the unusually limber branches
brush, common chokecherry, serviceberry, skunkbush
that give this tree its name probably are an adaptation
sumac, snowberry, snowbrush ceanothus, wax currant,
to minimize wind damage. Associated species include
and Woods rose (fig. 10.11; see table 10.1). Great Basin
various shrubs, such as common chokecherry, ground
wildrye and antelope bitterbrush grow in ravines and
juniper, mountain big sagebrush, Oregon-grape, wax
on sites where snow accumulates. Common grasses
currant, and western snowberry, along with forbs and
and forbs include bluebunch wheatgrass, Idaho fescue,
grasses (for example, spike fescue).
prairie junegrass, arrowleaf balsamroot, hairy golden
Foothill woodlands in the western half of Wyoming
aster, and lupine.
are dominated by Douglas-fir rather than ponderosa
Serviceberry is a conspicuous shrub in some mixed
pine (fig. 10.10). Many of the best examples are found
foothill shrublands. Two species are commonly rec-
in the Sunlight Basin, the Lamar Valley of Yellowstone,
ognized in Wyoming: Saskatoon serviceberry, which
Jackson Hole, the lower slopes of the Centennial Moun-
occurs throughout the state, and Utah serviceberry,
Fig. 10.10. (above) Douglas-fir is the most common tree on

north slopes at lower elevations in the Greater Yellowstone
Ecosystem. The south slopes are generally too dry for trees
but support a mixed foothill shrubland with bluebunch
wheatgrass, big sagebrush, Saskatoon serviceberry, and other
plants. Mountain big sagebrush dominates the shrublands
in the foreground. Photo taken looking west toward Hoback
Junction.
Fig. 10.11. (left) Saskatoon serviceberry, antelope bitterbrush,
rabbitbrush, and mountain big sagebrush dominate this mixed
foothill shrubland in the Sierra Madre. Aspen grows in the
ravine. Elevation 8,100 feet. Photo by Ken Driese.
which is found to the west and south.33 Both species

are deciduous, capable of root sprouting, and are important browse plants for deer and elk. At lower elevations,
where the frost-free period is longer, skunkbush sumac
is common, especially on the east side of the Bighorn
and Laramie mountains.
The relatively moist environment of the foothills allows for greater plant cover, which enhances
the probability of fires, whether caused by lightning
or humans. Periodic fires in some areas prevent the
growth of juniper and other conifers. Temporary shifts
165
in species composition may also occur following fire.
with succession or if grazing is confined to the period
Most of the shrub species have the ability to resprout
when the plant is dormant.37
from surviving roots and other belowground structures
Some foothill grasslands occur on extraordinarily
(big sagebrush and juniper are notable exceptions).
windswept plateaus or slopes with shallow soils. The
Fires commonly expose mineral soil, facilitating the
grasses are scattered and small forbs are common.
re-establishment or addition of some species through
Many of the forbs have the cushion plant growth form
seedling recruitment. Antelope bitter

brush, snow-
(see fig. 8.9), which keeps the stems and leaves densely
brush ceanothus, skunkbush sumac, and mountain-
aggregated at the soil surface, where the air is warmer.
mahogany are capable of nitrogen fixation because
Cushion plants are typical of windswept areas in alpine
of the actinorhizae formed by bacteria in the genus
tundra as well as in the lowlands, though the species
Frankia. This adaptation enables the establishment of
are different.38
these species on relatively infertile soils. Snowbrush

ceanothus is capable of layering, but in addition, seed
dormancy in this shrub is broken by the heat of a fire;
seedlings commonly are abundant in burns where the
species existed previously.
Greasewood occurs in a few places in the foothills,
Deciduous Woodlands
Four other vegetation types in the foothill-escarpment
mosaic are woodlands characterized by different combinations of aspen, chokecherry, Gambel oak, and bur
an interesting anomaly, as it is usually found where

groundwater is near the surface on saline soils in desert shrublands. The presence of greasewood on escarpments has been attributed to porous lignite seams that
cause groundwater to move to the surface on hillsides.34
Seeps or perched water tables, with concomitant salt
accumulation as some of the water evaporates, typically
develop on such sites.
Foothill Grasslands
Foothill grasslands can be found on windy slopes or plateaus, where snow does not accumulate in large quantities, soils are too shallow for most shrubs, or summer
rainfall is higher. Bluebunch wheatgrass is the characteristic plant, occurring most often on relatively warm,
dry sites. Idaho fescue is typical of the higher, more
mesic montane grasslands (above 6,900 feet).35 Threetip
sagebrush, a dwarf shrub, occurs with Idaho fescue on
windswept, shallow soils (see fig. 10.9). Little bluestem
is sometimes found on the slopes of escarpments at low
elevations.36
Bluebunch wheatgrass is a highly preferred forage
species for livestock and big game. At lower, warmer
elevations, it is usually found on north slopes or near
snowbanks, if it occurs at all. Various investigators have
observed that bluebunch wheatgrass becomes less common after hot fires or heavy grazing in spring or summer. Cheatgrass is a common invader following such
disturbances, but bluebunch wheatgrass increases again
Fig. 10.12. Ravines in the foothills of the Bear Lodge Mountains and Black Hills near Sundance typically have chokecherry, skunkbush sumac, Woods rose, and other shrubs.
Shrublands such as this are sometimes referred to as woody
draws. Mixed-grass prairie occurs on the upland, with ponderosa pine on some ridgetops. Elevation 3,900 feet.
Fig. 10.13. Aspen groves occur in some places on the lee

side of the Continental Divide south of Rawlins, where deep
snowdrifts develop during winter. The snow provides ade
quate water for shrubs and trees in an area that elsewhere is
semi-arid. This photo shows a portion of a doughnut-shaped
grove, known as an aspen atoll, where the center is dominated by shrubby cinquefoil, mountain silver sagebrush,
lupine, tufted hairgrass, and various other grasses and forbs.

Deep snow that persists into early summer prevents tree
establ ishment in the center. The shrubs in the foreground
and on the left are old aspen sprouts that are regularly
broken by the heavy snowpack. Aspen trees develop just
beyond the shrubby aspen, where snow accumulation is less.
oakall of which are deciduous and occur on mesic
Aspen-dominated woodlands tend to occur where
sites with deeper soils. The additional moisture appar-
the soils are deeper, such as on lower slopes where water
ently comes from a higher soil water-holding capacity,
seeps to the surface or on the lee sides of ridges where
more snow accumulation (from drifting), more summer
snow accumulates. Associated plants include species
precipitation, or runoff from the slopes above.
that occur in aspen woodlands at higher elevations (see
Chokecherry woodlands, also known as hardwood
chapter 11). Sometimes the aspen completely surround
draws or woody draws, are commonly part of a veg-
a snowdrift, forming a doughnut-shaped grove around
etation mosaic on the eastern plains that includes
a meadow that has been referred to as an aspen atoll
grassland and woodlands dominated by ponderosa
(fig.10.13). Such groves develop when trees first become
pine (fig. 10.12). In addition to chokecherry, common
established near ridgetops, which causes further accu-
shrubs include American plum, skunkbush sumac,
mulation of snow downwind. If the snow is deep
silver sagebrush, and western snowberry. 39 Trees com-
enough, it persists until midsummer and only herba-
monly found with the shrubs are boxelder and green
ceous plants capable of surviving with a short growing
ash (see table 10.1). Except for the lack of cottonwood,
season can persist. Gradually the aspen grow completely
chokecherry woodlands have the characteristics of
around the snowdrift and meadow by root sprouting.
some riparian woodlands. Specifically, they have more
The trees also expand inward, toward the edge of the
plant biomass than is found on adjacent shrublands,
central meadow. Here the average snow accumulation is
which provides good wildlife cover as well as forage
sufficiently shallow to allow the aspen to persist, but the
and shade for livestock.
weight of the snowpack causes breaking and bending,
167
resulting in trees that look more like shrubs. Only on

the outer fringes of the clone do aspen sprouts grow tall.
Bur oak woodlands are found in the northern and
eastern Black Hills (see fig. 16.6), and Gambel oak
woodlands can be found on the west side of the Sierra
Madre, on the Wyoming-Colorado border. Late spring
frosts and summer drought most likely inhibit Gambel
oak farther to the north by preventing establishment
of oak seedlings.40 Southward in Colorado and Utah
(see chapter 3), where Gambel oak becomes increasingly common, late-spring frosts are less frequent, and
summer precipitation is higher because of the Arizona
Monsoon. Gambel oak occurs in northern Utah primarily where temperature inversions regularly create
warmer thermal belts in the foothills, with an average
of 90 frost-free days a year.41 Here, Gambel oak is usually found on sloping upland sites between about 5,500
Plant-Animal Interactions in the Foothills

Escarpments and foothills provide habitat for a wide
diversity of wildlife. Of particular interest are elk and
other big game species, which tend to winter in these
landscapes, where snow cover is less and food more
available than higher in the mountains. Many winter
ranges have been greatly altered by agriculture, exurban
development, and other human activities. This loss of
habitat, coupled with the fact that winter ranges are less
extensive than the associated summer ranges, means
that animals are concentrated in small areas for long
periods each winter. Management of ungulate winter
ranges has long been controversial in Wyoming and
much of the intermountain West
Aspen, Elk, and Fire
and 7,500 feet, above the pinyon-juniper zone but
The management of foothill aspen groves has been
below ponderosa pine and aspen. Curiously, bur oak
one of the most interesting and highly debated natural
occurs only in the northeastern corner of the state
resource issues in the region.44 The controversy began
farther north, but the elevation is lower (and warmer)
in the mid-1950s, when some aspen groves in north
and annual precipitation is comparatively high (see
western Wyoming appeared to be dying (fig. 10.14). Such
fig.3.4). If it were not for late-spring frosts, oak would
groves were thought of as decadent. Aspen is unique,
probably be a co-dominant with ponderosa pine else-
because the numerous trees in a grove result from the
where in Wyoming, as is common in Colorado (for
root sprouting of only one or a few plants. Each shoot
example, near Colorado Springs). Oak might also occur
aboveground is better thought of as a branch. Seedling
with aspen and chokecherry. Climate change could
establishment is rare in the foothill environment, but
lead to an ex
pansion of oak woodlands if relatively
when successful, the roots sprout additional stems that
warm springs coincide with sufficient summer precipi-
grow into what most people think of as new treesa
tation for the seedlings to survive.42 Once established,
classical case of cloning that is more common in her-
oak tolerates cold, dry conditions.
baceous plants. Like other species of Populus, each stem
Oak woodlands burn readily during dry seasons, but
is comparatively short-lived, with few surviving past
the roots are not usually killed and they sprout vigor-
150 years. However, the root system lives indefinitely,
ously, often leading to an increase in density.43 Other
producing new stems around the edges of groves or
species invade during long fire-free intervals, such
where older stems are damaged.
as Rocky Mountain juniper and ponderosa pine (and
The apparent loss of foothill aspen groves is attrib-
pinyon pine, white fir, and canyon maple in north-
uted to two factors: fire suppression and the excessive
ern Utah). Both Gambel and bur oak woodlands pro-
browsing of sprouts by large mammals, particularly
vide grasses and forbs for big game during the winter,
elk.45 Browsing was the first factor to be debated.
and deer, elk, wild turkey, and squirrels consume the
Many of the so-called decadent groves in northwest-
energy-rich acorns. With time, forage production under
ern Wyoming occurred along known elk migration
oak canopies declines because of shading and competi-
routes, and many were located adjacent to winter
tion for water. Fire can be used to stimulate the growth
feeding groundsnotably the National Elk Refuge in
of more forage, but ranch buildings, summer homes,
Jackson Hole. Though elk are grazers in the summer,
and resorts located in the foothill environment can pre-
they browse on aspen sprouts in winter and spring.
vent this otherwise sound practice.
Too much browsing for too many consecutive years
reduces the number of sprouts. When aspen groves are

fenced to exclude large herbivores, the small sprouts
often grow into trees (fig. 10.15).
An interesting perspective on decadent aspen stands
was presented by U.S. Forest Service ecologist Norbert
DeByle.46 He suggested that one of the causes is political
pressure to maintain elk populations at the same high
level year after year, to satisfy hunters, guides, and game
and fish agencies that depend on license sales. DeByle
reasoned that in the 1800s the number of elk fluctuated considerably, largely stemming from higher mortality during some winters than others, and that aspen
sprouts had a better chance of developing into trees
during periods when the browsing pressure was low
Fig. 10.14. (right) An aspen grove in the Greater Yellowstone

Area that appears to be dying, most likely because of excessive
browsing by elk (see chapter 15). Mountain big sagebrush,
arrowleaf balsamroot, and other foothill plants occur in the
area; lodgepole pine and limber pine can be seen in the background. Elevation 7,100 feet.
Fig. 10.15. (below) Aspen groves usually recover quickly when
protected by a fence from browsing, such as in this area in
northwestern Wyoming. Aspen exists outside the fenced
exclosure, to the right, but only as small, heavily browsed root
sprouts. See chapter 15.
169
(anatural rest rotation). Elk populations still fluctuate,
and drier, as discussed in chapter 11. The magnitude
but probably to a lesser extent because of winter feed-
of the problem is not well known. What proportion of
ing programs. Notably, the number of elk has declined
the aspen groves in different mountain ranges is in a
during the past 12 years or so, almost certainly because
degraded condition? Could some aspen groves have
of the reintroduction of gray wolves in 1995 combined
been in poor condition long before the arrival of
with droughts. Furthermore, some evidence suggests
EuroAmericans in the 1800s?
that the wolves harass elk herds from time to time, causing them to move rather than stay in one aspen grove
for a long period. Aspen could be one of the beneficia-
Winter Rangelands for Deer and Elk
ries of restoring the regions top carnivore (but see chap-
The winter survival of pronghorn, deer, elk, and moose
ters 12 and 15 for further discussion of this topic and
depends largely on adequate food during the summer
sudden aspen decline).
and fall for the development of fat reserves sufficient to
Wildlife managers are sometimes sensitive about elk
sustain the animals during long, harsh winters. Deep
receiving so much of the blame for aspen groves that
snow or unusually cold winters cause a more rapid
appear to be in bad condition. They argue, correctly in
depletion of fat reserves, but fortunately, the snow com-
some cases, that livestock frequently congregate under
monly melts rather quickly on south slopes in the foot-
the trees because of the abundant forage that develops
hills. Also, the rugged topography makes it relatively
in the relatively moist environment where aspen is
easy for the animals to find windbreaks. The bark and
typically found. Cattle and sheep can cause consider-
twigs of trees and shrubs, though generally not pre-
able damage to aspen through browsing, trampling, and
ferred, often are consumed during this time, partly
bedding.
because they are visible above the snow. Browse lines
Fire is another factor that cannot be ignored in ex
develop on junipers, willows, and other tall shrubs but
plaining the apparent decline in some aspen groves.
are not necessarily an indication of excess numbers of
University of Colorado ecologists Thomas Veblen and
wintering animals at the present time (fig. 10.16).
Diane Lorenz, working in Colorado, suggested that
By the time spring arrives, winter ranges may seem
widespread burning in the late 1800s and early 1900s,
to be heavily grazed and browsed, yet the plants typi-
whether caused by lightning or humans, might have
cally recover quickly, because the herbivory occurs
triggered the initiation of new stands of aspen over
largely when they are dormant. Depending on manage-
large areas.47 Those stands are now aging, which could
ment practices and the extent of overlap between the
explain why the groves are failing over large areas. How-
summer plant preferences of livestock and the winter
ever, aging stands could also be the result of fire sup-
preferences of big game, winter range plants may have
pression during the past 60 or 70 years. Possibly more
a full summer of rest and become senescent before they
important, with fire suppression conifers have become
are again subjected to large ungulates. The seasonal
dominant in some groves, so that the fewer remain-
movements of the animals create a natural rotation sys-
ing aspen are subjected to more browsing than before.
tem.48 More than one year of rest may result if the herd
Large-scale fires could stimulate the development of
selects a different part of the foothills for wintering or if
new aspen forests, providing a larger food base and
high mortality during a particularly severe winter leads
removing some of the browsing pressure on any partic-
to fewer animals during the following winter.
ular grove. Notably, many new aspen seedlings became
A primary concern of wildlife managers during the
established in the Greater Yellowstone Area after the
past 10 years has been the decline in mule deer. Long
1988 fires (see chapter 15).
periods of droughtpossibly climate-change-induced
There is still much debate on the relative significance
droughtprobably have been a contributing factor,
of elk browsing, fire suppression, and wolf predation on
because inadequate moisture limits the summer forage
the condition of aspen woodlands in the foothills. Cli-
available to build up fat reserves. Another factor has
mate change could be another contributing factor, pos-
been winter habitat fragmentation by exurban devel-
sibly reducing aspen vigor if summers become longer
opments. All such factors reduce the amount of forage
transmission (such as brucellosis or chronic wasting disease), and the fostering of a public perception that protecting suitable winter habitat is not important.49 The
benefits are perceived to be fewer agricultural conflicts,
larger herds for hunters (and increased license sales for
the wildlife agencies), and relief from the angst associated with knowing that many animals are starving
during harsh winters because of industrial activity and
rural subdivisions.
Pine Seed Dispersal by Birds

One of the most interesting plant-animal interactions
in foothill and escarpment landscapes is that between
certain seed-eating birds and limber and pinyon pines.
Unlike most conifers, these pines have comparatively
large nutritious seeds, and the cone scales tend to hold
the seeds in place, even after the cones have opened.
Not surprisingly, the seeds are an important source of
food for various birds, squirrels, and even grizzly bears.
The pines benefit by having their seed dispersed and
sometimes planted by birds and squirrels.
The best-studied example is the Clarks nutcracker,
a common bird in the foothills and mountains (fig.
Fig. 10.16. Browse lines, such as on the taller, older Rocky
Mountain juniper in the background, are formed when
adequate food is not available for elk during the winter. Such
browse lines might have existed for centuries near elk and
deer winter ranges. The two junipers in the foreground are
younger and do not appear to have been heavily browsed,
most likely reflecting reduced populations of the wintering
elk and deer during their development. Photo taken between
Gardiner, Montana, and the border of Yellowstone National
Park. Elevation 6,000 feet.
availability directlyand indirectly as well, because

human activity forces the ungulates into areas where
the amount and quality of winter forage is either marginal or where another herd already resides.
Recognizing the problems associated with reduced
winter range, federal and state agencies initiated winter
feeding programs (see chapter 15). From the beginning,
the wisdom of feeding elk and deer in the winter has
sparked controversy, with Wyoming, Idaho, and Utah
having state-sponsored feed grounds, whereas Montana
and Colorado do not. Aside from rather high costs per
animal, the problems are excess herbivory of trees and
other plants in the vicinity of feeding grounds, disease
10.17).50 In Arizona, a single nutcracker was observed

carrying 95 pinyon seeds in a pouch behind its beak for
a distance of 13 miles. Usually the seeds are collected in
late summer and buried on south-facing slopes, which
Fig. 10.17.Clarks nutcrackers bury the peanut-sized seeds of

limber and whitebark pine in the soil, often on warm south
slopes. Seeds that are not recovered often develop into new
trees, some of which grow in clusters if a bird deposited more
than one seed in a single hole. Photo by LuRay Parker/
Wyoming Game and Fish Department.
171
are often snow-free during late winter. Many seeds are
sunny slopes that are frequently snow-free in winter,
relocated in the spring, a time coinciding with the feed-
conditions that might reflect an instinctive preference
ing of nestlings and when food for seed-eaters is in short
of the bird. Seeds cached on such sites would be more
supply. However, some seeds are left in the ground and
accessible in the winter, when food is scarce. Burned for-
develop into new treescommonly clumps of trees, one
ests are also selected by the birds, which could increase
from each of several seeds that were cached by the bird
the probability of limber and whitebark pine becoming
in the same hole. In a similar manner, the nutcrackers
part of the new forest that develops. Its well known that
also disperse the seeds of limber pine and whitebark
grizzly bears in the Greater Yellowstone Area consume
pine.51
large numbers of whitebark pine seed in the fall (see
As might be predicted, the distribution of nut
chapter 15). From the bears perspective, the more seeds
crackers overlaps with that of pines having large, wing-
planted by birds, the betterespecially at a time when
less seeds (in Europe as well as North America). The
many of the adult trees are dying from the depredations
plants benefit by having their seeds dispersed more
of mountain pine beetles and the invasive, non-native
widely and even planted. In fact, the large wingless
white pine blister rust. Conservation biologists are
seeds cannot be dispersed effectively over long dis-
attempting to maintain populations of whitebark pine
tances by any mechanism other than bird transport.
by planting seeds from trees that appear to be resistant
Seeds that simply fall to the ground are more likely
to the disease. (See chapter 14 for further discussion of
to be eaten by small mammals, and the seedlings are
white pine blister rust.)
less likely to become established on the litter layer of

the forest floor, a less favorable environment than that
Overall, the exposed bedrock of foothill and escarpment
provided by burial in mineral soil. Moreover, seedlings
landscapes creates sharp environmental boundaries and
that originate from fallen seed may be less likely to
a patchy vegetation mosaic. Such landscapes are highly
survive because of competition with the nearby and
valued real estate for people, providing the amenities of
already well-developed parent tree. This coadaptation
a moderate, low-elevation environment close to pictur-
between birds and some pines is further illustrated
esque escarpments that provide windbreaks, together
by the observation that the nutcracker distinguishes
with grand vistas of the plains below and the mountains
between viable and nonviable seeds. There is little to
above. Dispersed residential development has been the
be gained for either the bird or tree if less nutritious,
fastest growing land use in the United States since 1950
nonviable seeds are cached.
and has resulted in habitat fragmentation throughout
Bird dispersal appears to be one of the factors caus-
the intermountain Westoften in the foothills. As dis-
ing the patchy distribution pattern of limber and white-
cussed further in chapter 18, debates about how best to
bark pine. Typically, these trees are found on ridges or
use these lands surely will continue.
Chapter 11
Mountain Forests
Rising above the plains, basins, and foothills are moun-
uneven snow accumulation and melting, with some
tains that greatly diversify the landscapes of the region.
patches of ground becoming snow-free much earlier
Most are anticlinal or fault block in origin and have
in the spring than others. Some species, such as dwarf
cores of Precambrian granite, gneiss, and quartzite. In
huckleberry, cannot tolerate snow cover that lasts until
some areas the Precambrian rocks are still covered by
early July.1
sedimentary strata formed at the bottoms of ancient seas
The most noticeable environmental gradient in
that spread across Wyoming millions of years ago, before
mountains is a decrease in temperature with an increase
mountain building began. More than 50 peaks rise
in elevation (see fig. 3.6). The rate at which tempera-
above 13,000 feet, mostly in the Wind River Mountains.
ture cools can be 25F for every 1,000 feet of elevation
Some of these mountain ranges are massive (see fig. 1.1),
gain, depending on the time of year.2 The variability
whereas others are small islands in a sea of sagebrush.
also depends on differences in mountain extent, cold-
Glaciers have shaped portions of the higher mountains.
air drainage, snow cover duration, potential for tem-
Large or small, the mountain ranges are important

economically. Almost all of the forests are found there,
perature inversions, and other climatic and topographic

factors.
and most of the water used by agriculture, industry,
Other environmental factors also change with ele-
and municipalities originates there as snowfall. The
vation. The snow-free period and length of growing
mountains are also important for outdoor recreation.
season are typically shorter high on the mountains,
Ecologically, they are interesting because of their eco-
but precipitation generally increasesat least to mid-
systems and the ways that they affect the lowlands (for
elevations, where snow accumulation is greatest. Snow
example, through the development of alluvial fans and
accumulation combined with lower evaporation rates
rainshadows). Indeed, the landscape patterns observed
caused by cooler temperatures provides an environment
in the foothills and lowlands are caused by the nearby
that, in general, is considerably wetter than in the sur-
mountains.
rounding lowlands (see fig. 3.4). Countering the effect

of cooler temperature is lower atmospheric pressure,
Variation in Mountain Environments

Spatial variation in mountain forests can be attributed
which allows relatively high rates of evapotranspiration.3 Still, plant water stress is less frequent in midelevation forests than in any other upland ecosystem.
partially to environments that change with elevation,
The effects of topography are just as important as
slope steepness and aspect, and the influences of under-
those of elevation. South-facing slopes are warmer and
lying bedrock on soil development. Other factors are
drier and north-facing slopes are cooler and wetter
173
than might be expected based solely on elevation. Val-
all interact to determine the spatial distribution of any
ley bottoms are often cool because of cold-air drainage.
given plant species.4
Thus, species typically found high on the mountains

may extend to low elevations in ravines or on north
slopes with less direct sunlight, and species requiring a
Surviving in the Mountains
warmer environment may be found at unusually high
Perhaps the primary challenge for plant survival in the
elevations on south slopes (fig. 11.1).
Rocky Mountains is the short, cool, and sometimes
Soils also have a pronounced effect on the landscape
dry growing season. Several distinctive adaptations are
mosaic. For example, meadows occur in the Bighorn
involved. Most notable is that photosynthesis occurs
Mountains on comparatively dry, fine-textured soils
in montane plants at temperatures near freezing, or
even at high elevations where forests might be expected
sometimes even below,5 and many plants are evergreen
(see chapter 12). Forests of Douglas-fir, lodgepole pine,
or wintergreen (that is, they have chlorophyll in their
Engelmann spruce and subalpine fir grow on other
leaves or stems all year). The ability to tolerate low tem-
kinds of soils (Inceptisols and Alfisols), where moisture
peratures, combined with the presence of chlorophyll
stress is less severe in late summer (such as on north
throughout the year, enables some plants to extend
slopes). Lodgepole pine is typically the most common
their growing season into late fall and begin the next
tree in forests on coarse, less fertile soils derived from
one earlier in the spring. Evergreen conifers are excel-
granite or rhyolitic lava. Multiple factorssoils, local
lent examples, but other plants have similar adapta-
climate, influences of herbivores and pollinators, and
tions. For example, such deciduous plants as aspen
local history of disturbance and species migrations
and dwarf huckleberry have chlorophyll in their stems,

which permits photosynthesis even when they have no
leaves. The green stems of the huckleberry may account
Fig. 11.1. Approximate distribution of different forest types in

relation to elevation and moisture availability. Note that each
kind of forest typically occurs at the lowest elevation in cool,
moist ravines. Foothill shrublands occur just below woodlands and forests at lower treeline.
Some shrubs and herbaceous plants have leaves
Wet
meadow
Fellfield
Turf
Willow
enough to permit light penetration, when the temperature is at or near freezing.7 The capacity for photosyn-
Spruce-fir
Lodgepole pine
Aspen
9000
thesis at cold temperatures is an important adaptation,

3000
Aspen
9000
Lodgepole pine
Willow
8000
Alder
Birch
2500
Aspen
Douglas-fir
or
Ponderosa pine
Foothill
shrublands
but simply being able to survive the cold is another. A

comparison of low-temperature tolerances of numerous
tree species found that, unlike the trees from warmer
climates, those of the Rocky Mountains could tolerate
temperatures of 76F.8
Another problem for mountain plants is acquiring
nutrients from infertile, coarse-textured soils. All of the
tree species have mycorrhizae (fig. 11.2), which are con-
7000
Cottonwood
synthesis and growth as soon as the snow is shallow
ELEVATION (m)
ELEVATION (feet)
deciduous evergreen plants.
elk sedge and wintergreen. They are capable of photo-
3500
Alpine
6000
synthesis.6 Both aspen and huckleberry are examples of
that remain green even under the snowfor example,
12000
9000
for a substantial portion of the plants annual photo-
centrated in the top 12 inches of the soil, where limiting

2000
nutrients are most likely to be available. Once incorporated into plant tissues, the nutrients are retained for
Wet
Cool North
valley ravine slope
bottom
E-W
slope
South Ridge
slope
MOISTURE GRADIENT
extended periods simply by the longevity of leaves and

twigs. Lodgepole pine needles, for example, persist for
518 years, depending on environmental conditions.
Mountain Forests
Several studies have shown genetic differentiation over

short environmental gradients in Rocky Mountain
conifers, and aspen groves at high elevations are genetically different from those in the foothills.12 Considerable genetic variability, one aspect of biodiversity, can
be expected in landscapes with rapid spatial and temporal changes.
Vertebrates have an equally diverse range of adaptations for surviving in mountain landscapes. Some
migrate in the fall to warmer or less stressful environments at lower elevations or farther south, minimizing their exposure to cold temperatures, deep snow,
Fig. 11.2. Microphotograph showing swollen mycorrhizal root
tips on lodgepole pine. The filamentous hyphae of the fungus
extend from the bulbous tips into the soil, facilitating the
uptake of water and nutrients. Photo by Steven L. Miller.
and food shortages during the winter. Other species

survive because of remarkable insulation provided by
fur, fat, and feathers, and because they find or build
insulated shelters, such as in deep snow or under logs.
Energy stored during the preceding summer in fatty
tissue or food caches is especially critical for animals
When the leaves of evergreen plants are about to fall,
that are year-round residents. Many species have rel-
their limiting nutrients are first partially reabsorbed by
atively large feet that enable swift travel across deep
the twig.9 Thus, plants in nutrient-deficient environ-
snow, for example, red squirrels, martens, lynx, and of
ments have evolved mechanisms for conserving the
course, snowshoe hares.13 A few are able to enter a state
limiting nutrients they already have.
of torpor, or reduced physiological activity brought
Short, cool growing seasons also present problems for
about by decreased metabolism and body temperature.
seedling establishment, because plants that are only a few
Examples include bears, marmots, bats, reptiles, and
months old are the most susceptible to water, tempera-
amphibians.14
ture, and nutrient stresses. Seeds often germinate despite

cool temperatures in the spring, allowing the maximum
time possible for seedlings to develop the stored energy
The Forest Community
and cold-hardened tissues needed for the following win-
Forests are characterized by trees, but, with regard to
ter. Moreover, the formation of mycorrhizal roots typi-
the numbers of species, most plants in the commu-
cally occurs in the first few weeks after germination.10
nity are shrubs and herbaceous plants in the under-
Despite these adaptations, seedlings often succumb to
story. Common shrubs include buffaloberry, dwarf
summer drought, late-spring frosts, and other factors.11
huckleberry, and ground juniper; the herbs include
Because seedling establishment is so precarious, nearly
heartleaf arnica, lousewort, pyrola, and various grasses
all mountain plants are perennials that live for many
and sedges (table 11.1). Many animals and legions of
years.
microorganisms coexist with the plants. In one way or
Different species have evolved to tolerate differ-
another, all are adapted to living with the other spe-
ent levels of environmental stressa fundamental
cies, large and small. Indeed, some are dependent on
principle in considering the causes of patches in land-
others. A special example consists of plants known as
scapes anywhereand stress levels change rapidly
mycoheterotrophsthose that lack chlorophyll and
with changes in elevation, soil, and topographic posi-
obtain their energy from decaying tree roots by way
tion. Consequently, species composition often changes
of saprophytic fungi. A common example in Rocky
abruptly in the mountains. Variation occurs even with-
Mountain forests is pinedrops (Pterospora andromedea).
out changes in species composition, as many species
Clearly, understanding forest ecology is not possible by
have a wide range of genetically determined tolerances.
studying the trees alone.
175
Table 11.1 Some characteristic plants of mountain forests and woodlands in Wyominga
Common name
Latin name
Limber
Ponderosa
pine
pine
Douglas-fir
woodland
forest
forest
Aspen
forest
Lodgepole
Whitebark
pine
Spruce-fir
pine
forest
forest
woodland
TREES
Aspen
Populus tremuloides
Douglas-fir
Pseudotsuga menziesii
Engelmann spruce
Picea engelmannii
Limber pine
Pinus flexilis
Lodgepole pine
Pinus contorta var.

latifolia
Pinus ponderosa
Pinus albicaulis
Abies lasiocarpa
Antelope bitterbrush
Purshia tridentata
Shinyleaf spiraea
Spiraea betulifolia var.

lucida
Chokecherry
Prunus virginiana
Ponderosa pine
Whitebark pine
Subalpine fir
SHRUBS
Common juniper
Juniperus communis
Gooseberry currant
Ribes montigenum
Grouse whortleberry
Vaccinium scoparium
Ninebark
Physocarpus spp.
Oregon boxleaf
Paxistima myrsinites
Rocky Mountain
juniper
Juniperus scopulorum
Rose
Rosa spp.
Russet buffaloberry
Shepherdia canadensis
Saskatoon
serviceberry
Amelanchier alnifolia
Skunkbush sumac
Rhus trilobata
Utah snowberry
Symphoricarpos oreophilus
var. utahensis
Wax currant
Ribes cereum
Bluebunch
wheatgrass
Pinegrass
Calamagrostis rubescens
Idaho fescue
Festuca idahoensis
GRASSES
Limber
Ponderosa
pine
pine
Douglas-fir
woodland
forest
forest
Aspen
forest
Lodgepole
Whitebark
pine
Spruce-fir
pine
forest
forest
woodland
Common name
Latin name
Spike fescue
Leucopoa kingii
Mountain brome
Bromus marginatus
Wheelers bluegrass
Poa wheeleri
Geyers (elk) sedge
Carex geyeri
Ross sedge
Carex rossii
Arrowleaf
balsamroot
Balsamorrhiza sagittata
Bedstraw
Galium spp.
Fireweed
Chamerion angustifolium
Heartleaf arnica
Arnica cordifolia
Horsetail
Equisetum spp.
SEDGES
FORBS
Kinnikinnick
Arctostaphylos uva-ursi
Longstalk clover
Trifolium longipes
Meadow-rue
Thalictrum spp.
Nevada pea
Lathyrus lanszwertii
Creeping barberry
(Oregon-grape)
Mahonia repens
Wintergreen
Pyrola spp.
Sidebells
wintergreen
Orthilia secunda
Red baneberry
Actaea rubra
Sickletop lousewort
Pedicularis racemosa
Silky lupine
Lupinus sericeus
Sticky purple
geranium
Geranium viscosissimum
Timber milkvetch
Astragalus miser
Twinflower
Linnaea borealis
Western coneflower
Rudbeckia occidentalis

Whitebark pine occurs in northwestern Wyoming and northward in the Rocky Mountains.
The density of trees and understory plants in forests

varies considerably from place to place and also fluctuates through time.15 Where tree density is high, there
may be hardly any understory herbs or shrubs because
of too much competition from the trees for light, water,
and nutrients; in more open forests, the abundance
and diversity of smaller plants are often much greater.16
Lower tree density is found on sites that are too dry or
infertile to support a dense tree canopy, or in forests that
have been thinned by disturbances, as discussed below.
Snowdrifts persisting until early July restrict the growth
of some understory species, probably because the growing season is too short after the snow disappears.17
Plant parasites reduce tree growth, produce deformed
trees, or even kill some treesall problems in a setting
where wood production is a primary objective. Otherwise, native parasites add diversity to the forest community. Two that are especially important on Rocky
Mountain trees are dwarf mistletoe and comandra blister rust, both of which have coexisted with their hosts
for millennia.
Dwarf mistletoe (Arceuthobium americanum and other
species) is a flowering plant that is an obligate parasite
on pines (fig. 11.3). The pale green leaves and stems
are capable of some photosynthesis, but the plants
extend their roots into the hosts inner bark (phloem)
Fig. 11.3. Dwarf mistletoe is a parasitic flowering plant common on lodgepole and limber pine. The seed is spread to other
trees by explosive fruits, propelled by hydrostatic pressure.
and sapwood, where they obtain energy in the form

of carbohydrates, as well as water and nutrients.18 Tree
cides are avoided, because they generally have not been
growth is slowed, and sometimes the trees die.19 How-
effective, are expensive to apply over large areas, and
ever, bird species richness can be higher with mistletoe
have adverse secondary effects. Harvesting obviously
infection, suggesting that mistletoe eradication may be
infected trees is the most common approach, but often
undesirable where wood production is a less important
young mistletoe plants are difficult to see. Where foster-
objective.
ing the growth of a new generation of trees is a prior-
Mistletoe seed dispersal is accomplished by the ejec-
ity, clearcutting is the preferred option, whereby nearly
tion of seeds from the fruits by hydrostatic pressure,
all trees are cut, whether they are infected or not. This
with the seeds often traveling 2040 feet before landing
helps ensure that no infected trees are left behind. Most
on the leaves of another tree. The seeds have a slippery
likely, the prehistoric control of dwarf mistletoe was
coating and slide to the leaf base, where they germinate
periodic forest fires. Mistletoe now is probably more
in the spring. Infection occurs when the root success-
widespread than ever before because of fire exclusion,
fully penetrates the bark of 1- to 3-year-old twigs. The
and also because some early logging practices removed
mistletoe plant is hardly visible for several years, and
only the healthy trees.
new seeds are not produced for 56 years. Both healthy
Comandra
blister
rust
(Cronartium
comandrae),
and stressed pine trees are susceptible to mistletoe
another common native plant parasite, is a fascinat-
infection.
ing fungus that requires three very different plant spe-
Forest managers sometimes work to reduce the abun-
cies in close proximity to accomplish its life cyclein
dance of mistletoe to maximize tree growth. Herbi
this case, lodgepole pine, big sagebrush (or a few other
Mountain Forests
species), and a small herbaceous plant known as comandra or bastard toadflax. Comandra, the plant, is commonly found growing in dry mountain meadows and
is itself an obligate root parasite on sagebrush and various other plants. The disease therefore usually develops
only where sagebrush, comandra, and lodgepole pine
occur near one another. This arrangement is common
and the disease is widespread.20 Tree death cannot usually be attributed to the rust directly, but the treetops
die back and often a forked trunk develops. This is a
problem for forest managers focused on wood production, because it causes a reduction in the amount of harvestable wood. As with dwarf mistletoe, however, it is a
native species and is part of the ecosystem.
Animals also have significant effects on forest communities (table 11.2). For example, deer and elk can
reduce the rate of aspen regeneration by browsing the
small sprouts produced by the root system, and seedcaching birds (such as Clarks nutcracker) influence the
distribution of limber and whitebark pine in forests
as well as the foothills (see chapters 10 and 15). Pine
seeds are an important source of food for red squirrels
(fig. 11.4), with the cones being cached by the squirrels for winter consumption. However, not all seeds are
eaten, partly because the pine trees have adaptations
that deter squirrels; for example, the cones are resinous
and have thick scales. 21 All coniferous forests in Wyoming have red squirrels, but where they are not found,
to the north and west of Wyoming, the cones are less
massive and have twice as many seeds. Also, the seeds
are more easily removed. In such situations, the nutcrackers are more abundant. Where the nutcrackers
and squirrels do coexist, Wyoming ornithologist Craig
Benkman and his associates have found that the birds
bill is longer and stouter, presumably to give the bird a
better chance of prying open the scales of the heavyduty cones that have evolved in the presence of the
squirrels. 22
The serotinous cones produced by lodgepole pine
retain large numbers of seeds, which has made the
cones an especially rich food source for squirrels. As
Table 11.2. Some mammals, birds, amphibians, and

reptiles found primarily in mountain landscapes
MAMMALS
Bear, black
Bear, grizzlya
Bobcatb
Chipmunk, leastb
Cougar
Coyoteb
Deer, muleb
Elkb
Ground squirrel, Wyomingb
Hare, snowshoe
Marmot, yellow-bellied
Marten, American
BIRDS
Bluebird, mountainb
Chickadee, black-cappedb
Chickadee, mountain
Crossbill, red
Crossbill, white-winged
Dipper, American
Duck (various species)b
Finch, Cassins
Flicker, northernb
Flycatchers (various species)b
Goshawk, northern
Grosbeak, evening
Grosbeak, pine
Grouse, dusky
Grouse, ruffed
Hawk, red-tailedb
Jay, gray
Jay, pinyonb
Jay, Stellers
Junco, dark-eyedb
Kinglet, ruby-crowned
Nutcracker, Clarks
Nuthatch, red-breasted
Owl, boreal
Owl, great hornedb
Pipit, American
Raven, commonb
Robin, Americanb
Rosy-finch, gray-crowned
Rosy-finch, black
Rosy-finch, brown-capped
Sapsucker, red-naped
Sapsucker, Williamsons
Solitaire, Townsends
Sparrow (various species)b
Swallow, violet-green
Tanager, western
Thrush, hermit
Thrush, Swainsons
Veery
Warbler (various species)b
Woodpecker, American
three-toed
Woodpecker, black-backed
Woodpecker, downyb
Woodpecker, hairyb
AMPHIBIANS
Toad, boreal
Frog, boreal chorusb
Frog, Columbia spotted
noted, however, thicker cone scales have evolved to
REPTILES
minimize accessibility. This, in turn, has favored the
Boa, rubber
Gartersnake, wanderingb
evolution of larger jaw muscles in squirrels (compared
Moose
Mouse, deerb
Pika, American (alpine)
Pocket gopher, northernb
Porcupineb
Sheep, bighorn
Squirrel, red
Vole, heather
Vole, southern red-backed
Wolfa
Wolverine
Woodrat, bushy-tailedb
to those of squirrels in Douglas-fir forests, which do
not have serotinous cones). Craig Benkman and Matt
Frog, northern leopardb

Frog, wood
Salamander, tigerb
Snake, smooth green
Found only in the Greater Yellowstone Ecosystem.

Also found away from the mountains.
179

Fig. 11.4. Red squirrels have coexisted
with lodgepole pine for millennia.
Populations that depend on serotinous
lodgepole pine cones have extraordinarily strong jaw muscles for extracting the nutritious seeds from the
hard woody cones. The squirrels are
omnivores, as they also eat mushrooms,
bird eggs, and nestlings when available.
Photo by Fred Walsh.
Talluto concluded that seed predation by red squirrels
into forests dominated by ponderosa pine or Douglas-fir
reduces the occurrence of trees with serotinous cones.
at lower elevations, lodgepole pine at mid-elevations,
Thus, the characteristics of serotinous cones and the
and Engelmann spruce and subalpine fir at higher eleva-
proportion of serotinous trees in a specific forest is the
tions. 24 Considerable overlap occurs in the elevational
result of natural selection exerted by both the squirrels
distribution of these tree species, and in some areas the
and fire.23
foothill vegetation grades directly into lodgepole pine
Forest communities include hundreds or thousands
or Douglas-fir, as, for example, on the east slopes of the
of species of native insects. Most are inconspicuous
Medicine Bow Mountains and around Jackson Hole and
and spend their lives quietly pollinating flowers or
the Sunlight Basin. Ponderosa pine is rarely found in
decomposing dead organic matter. A few insects attract
western Wyoming, where Douglas-fir forests usually
attention, however, because they feed on living trees,
border the foothill vegetation.
sometimes killing their hosts. One groupthe bark
Intermingled with the coniferous forests are moun-
beetleshas gained prominence recently because these
tain meadows, also known as parks, along with rib-
insects are changing many forests in the Rocky Moun-
bon forests, snowglades, aspen groves, various kinds of
tains and westward. The causes and effects of bark
riparian communities, and shrublands or woodlands
beetle outbreaks are examined later in this chapter.
dominated by mountain big sagebrush and limber pine.

The upper treelinewhere the mountains extend that
The Forest Mosaic: Variation with Elevation

and Topography
highis characterized by short, wind-pruned Engelmann spruce, subalpine fir, and limber pine in most of
Wyoming ranges, though whitebark pine is typical of
The causes of vegetation changes with elevation have
the treeline in northwestern Wyoming and the North-
been of interest to scientists for more than two cen
ern Rockies. In central Colorado, bristlecone pine can
turies. In the Rocky Mountain region, foothill grass-
be found at upper treeline and also forms subalpine
lands, shrublands, and woodlands commonly grade
woodlands in some areas.
Mountain Forests
Disturbance and Change over Time: Another

Important Source of Variation in Forest
Communities
Forests are always changing, sometimes so slowly that
the changes are imperceptible. The ecological processes
that drive slow changes include competition among
trees for light and other resources, selective feeding
disturbances in forests are especially noticeable be

cause they commonly occur over large areas and their
impacts often last for decades (fig. 11.5). 26 In any mountain landscape, a shifting mosaic of forests responds
both to local environmental conditions and to the
effects of disturbances, some of which were recent and
others centuries old.
by herbivores, tree death, seed dispersal, and germination. Other changes are rapid and obvious, usually
resulting from discrete disturbances that kill or injure
many of the trees and other forest organisms. Examples include fire, an insect outbreak, a severe windstorm, and timber harvesting. Forests also change in
response to variation in the local climate. For example,
a severe but temporary drought may injure or kill susceptible plants; or a long-term directional change in
climate may lead to a gradual change in overall species
composition, with new species moving in while others
decline. In addition to the adaptations already discussed, surviving in Rocky Mountain forests requires
an ability to bounce back after the disturbances that
have occurred for millennia. 25
Landscape changes caused by disturbances and bi
ological processes are superimposed on the patterns
that develop in response to environmental gradients
in local climate and soils. Both kinds of pattern are
important in all vegetation types, but the effects of
Forest Fires
Two key aspects of all disturbances are their frequency
and severity. The natural frequency of fire varies from
decades to centuries among different forest types. Forest
fires tend to be most frequent in the foothills and lower
elevations of the mountains, where precipitation is sufficient to support flammable vegetation and where summers are often dry enough to permit fires to ignite and
spread over large areas. Late nineteenth- and twentiethcentury fire exclusion reduced fire frequency in many
lower-elevation landscapes.27 Fires became less frequent
in some areas because livestock grazing reduced the
abundance of fine-textured fuels, and roads, fields, and
towns broke up formerly continuous expanses of flammable vegetation.
At higher elevations, fires are notably less frequent
because of the typically moist conditions in those en
vironments: the snowpack melts later in the spring,
or not until midsummer. Also, summer rainstorms
often dampen the fuels, and relative humidity is often
Fig. 11.5.Tree-killing bark beetles of the genus Dendroctonus,

such as this mountain pine beetle, are native insects that have
coexisted with pine trees for thousands of years. The adults
are about 0.2 inch in length, though their wingspan is much
longer. Many species of bark beetles do not kill trees, subsisting instead on the inner bark of trees that have recently died
from other causes. Photo by Dion Manastyrski,from the collection of Lorraine Maclauchlan.
higher than at lower elevations. Lightning ignites fires

every year in high-elevation forests, but most fail to
spread beyond the ignition point because fuels are too
wet to burn. Only in unusually dry years are weather
conditions suitable for extensive fire spread. Significantly, its only during those historically infrequent
years that large fires occur in high-elevation forests.
Compared to climatic factors, fire suppression has
had a small impact on the frequency and size of highelevation fires.
Two related terms are fire intensity and fire severity. Fire intensity refers to the energy released in a fire,
expressed as BTUs per minute per unit area; fire severity refers to the effects of that energy release on organisms or ecosystems. Intensity must be measured while
the fire is in progress, which is difficult. Severity, however, can be estimated after a fire by characterizing tree
181
mortality and combustion of organic matter. Severity

ranges from surface fires that only consume a portion
of the forest floor and kill small trees, without harming most larger trees, to high-severity stand-replacing
crown fires that kill most of the trees and consume
essentially all of the forest floor. Forest fires typically
burn in a highly heterogeneous manner, responding
to wind direction, slope and aspect, and variation in
local weather and fuel conditions. The result is a mosaic
of severely burned patches, light or moderately burned
patches, and unburned forest.28
Each tree species has adaptations that enable it to persist in the face of periodic disturbance, and each major
forest type tends to be characterized by a different kind,
Bark Beetle Outbreaks

Rapid increases in the populations of some insect species
can lead to major disturbances that change forest structure and dynamics.31 The mountain pine beetle is the
insect most often mentioned as a disturbance agent in
Rocky Mountain forests, but other species of bark beetle
are also involved, each specializing in feeding on a particular tree species or group of species. These include the
Douglas-fir beetle, spruce beetle, and western balsam bark
beetle (which kills fir trees). All are natives. Defoliating
insects, notably the western spruce budworm and aspen
tent caterpillar, also modify forests. Because bark beetles
have had such dramatic effects over such extensive areas
frequency, and severity of disturbance. For example,

the thick bark of mature ponderosa pine and Douglasfir trees protects the critical inner bark tissues from heat
injury and enables these trees to survive low-intensity
surface fireswhich historically were the most common type of fire in many low-elevation ponderosa pine
and Douglas-fir forests. However, all conifers are killed
by fires that burn through the crowns of the trees, a
common type of fire in montane and subalpine forests.
Lodgepole pine is well adapted to such high-severity
fires, because even though the adult trees are usually
killed, each tree often has hundreds of cones that store
large numbers of viable seeds. At least some of the seeds
survive even an intense fire, as discussed later in this
chapter. These seeds germinate and grow best on the
bare soil exposed by a severe fire. Aspen also persist in
burned forests, despite fire killing all the aboveground
portions of the tree, because its roots usually survive
and produce abundant new sprouts that grow rapidly
into new trees. Soil is an excellent insulating material
for the roots.
Fires also affect animal abundance and distribution
patterns. For example, three-toed and black-backed
woodpeckers typically nest and reproduce in coniferous forests that have recently burned or been affected
by bark beetle outbreaks, because the insects inhabiting
the dead or dying trees provide an abundant and readily available food source. 29 Ungulates and bears benefit
from the greater abundance of herbaceous or shrubby
plants after such disturbances, and raptors are drawn
to disturbed forests because rodents and other prey are
more visible.30
Fig. 11.6.Numerous spots of resin and yellow sawdust on

the bark indicate that this lodgepole pine has been invaded
successfully by mountain pine beetles. Each spot represents
the entrance of one beetle. The beetles introduce the bluestain fungus that soon plugs the sapwood, causing tree death
within a few months. The leaves are reddish-brown by the
following summer (as illustrated in figs. 16.4 and 16.9 for
infested ponderosa pine). This pine also has a scar on the
trunk, possibly caused by a porcupine if not a fire.
Mountain Forests
Fig. 11.7. Many lodgepole pine on the mountain slope in the

background have been killed by mountain pine beetles. In
other areas, Engelmann spruce forests are affected similarly
by the spruce beetle. This area is near Green River Lakes in
the Wind River Mountains. Large numbers of elk are attracted
to this area in the winter because of a winter feeding program,
which may account for the poor condition of the aspen in

the foreground (see chapters 10 and 15). The soil on this
south slope is relatively moist, as indicated by the presence
of shrubby cinquefoil, mountain silver sagebrush, common
juniper, and western snowberry. The small conifer next to the
aspen is limber pine.
in recent decades, they are the focus of this section. Only
In late July and August the female beetles emerge
a few of the hundreds of different kinds of bark beetles in
from the bark and fly toward other trees. They are most
western North America actually kill trees; more than 99
often attracted to larger trees (those greater than 68
percent feed on trees that have already died from other
inches in diameter), which surely is adaptive because
causes, contributing to the decomposition of bark and
such trees provide the beetle larvae with more food,
woodan important ecosystem service.32
namely, the phloem, cambium, and sapwood of the
All tree-killing bark beetles have similar life histo-
inner bark. In addition to visual clues, the beetles can
ries and ecological effects, which can be illustrated by
be attracted by the odors emitted by stressed or weak-
the mountain pine beetle. This insect attacks all native
ened trees, whether caused by drought, age, scarring by
pines in Wyoming (lodgepole, ponderosa, whitebark,
disturbances, or other factors.34
and limber). 33 In most years the beetle population is
After landing, the female beetle bores a hole through
low, surviving in weakened or recently fallen trees,
the bark and is soon cutting egg-laying galleries in the
with populations that are referred to as endemic. Peri-
outer sapwood and phloem. However, if the tree is in
odically, however, the population increases rapidly, and
good health, resin is produced in such quantities that
vast numbers of trees are killedan event referred to as
the beetle is pitched out before any damage to the tree
an outbreak, epidemic, or eruption (figs. 11.6 and 11.7).
occurs. Such trees are easily recognized by small globs
An understanding of the life cycle of the beetle and of
of resin adhering to the external surface of the bark,
the causes of its population explosions is fundamental
in which a dead beetle can often be found. Healthy
to understanding forest ecosystem dynamics.
trees also produce chemicals that are directly toxic to
183
the beetles. Less vigorous trees are unable to produce
temperature threshold applies in all places and at all
enough resin or chemical defenses to repel the beetles.35
times. Great genetic and physiological variations occur
Following a successful attack, one often sees fine saw-
among different beetle speciesand among geographi-
dust around the holes made by the beetles and at the
cally segregated populations of the same speciesin
base of the tree.
the rate, timing, and effectiveness of cold hardening.
The mountain pine beetle is a tiny insect about 0.2
In general, however, bark beetle larvae are most vulner-
inch long, about the size of a grain of rice. The first few
able to a spell of cold weather in late fall and early win-
female beetles to successfully burrow through the bark
ter, before the cold hardening process is completed, as
promote mass attacks by emitting an aromatic chemi-
well as in late winter and early spring, when warmer
cal, known as an aggregating pheromone, which attracts
temperatures have stimulated the beginning of a break-
Curi-
down of the antifreeze compounds. In the late spring or
ously, once a sufficient number of beetles have been
summer, after further feeding and maturation, another
attracted to the tree, the successful beetles emit a dis-
generation of adult beetles emerges to attack other live
aggregating pheromone, which repels other beetlesan
trees, and the cycle continues.
other mountain pine beetles to the same tree.
36
amazing adaptation that helps ensure the survival of
This life cycle enables bark beetles to persist at low
the species. If too many beetles attack the same tree,
numbers, killing the occasional weak or stressed tree
fewer would reproduce successfully. Such specific
but having little impact on overall forest structure.
responses to specific stimuli by both the beetles and the
How, then, does a bark beetle population suddenly
tree are a reflection of the long history of co-evolution
explode into an outbreak in which the beetles kill so

many trees that everyone notices the effects? Two con-
of the insect and its host. 37

Following successful attack, the females lay eggs in
ditions are required: suitable climate conditions and
their galleries. At the same time, the spores of a fungus
susceptible trees. Warmer temperatures affect both, by
known as the blue-stain fungusare introduced.
The
enhancing survival of beetle larvae and contributing
fungal filaments grow into the sapwood, where they
to the development of tree water stress, thereby reduc-
restrict the flow of water and nutrients and kill the
ing the amount of resin produced and making the trees
tree in less than a year.39 The beetle eggs soon hatch,
more susceptible to beetle attack. Ecologist Teresa Chap-
and the larvae begin feeding on the inner bark, mov-
man and colleagues found that recent mountain pine
ing horizontally around the tree, cutting through the
beetle outbreaks in northern Colorado and southern
phloem and stopping the flow of carbohydrates from
Wyoming closely followed a period of warming tem-
the leaves to the roots. Although this girdling surely con-
peratures, which included several years of reduced pre-
tributes to the death of the tree, trees stressed only by
cipitation.42 Large numbers of trees were vulnerable to
girdling can live for 45 years.
beetle attack, because they were stressed by drought and
40
38
It seems clear that the
plugging of the sapwood by the blue-stain fungus is the
were of sizes preferred by beetles.
primary factor causing tree death. Notably, the larvae
As forests age, the trees experience increased compe-
also obtain important nutrients by feeding on the blue-
tition from their neighbors and many become more sus-
stain fungus.41
ceptible to beetle epidemics. Defensive chemicals, such
In the fall, the larvae begin to produce antifreeze-
as resin, are thought to have low priority in the alloca-
like chemicals in their bodies. They also eliminate
tion of limited carbon compounds by trees, compared
substances that could serve as ice-nucleating agents.
to the maintenance of leaf area and young roots, with
This process of cold hardening greatly increases winter
the result that less resin is produced and beetle inva-
survival with minimal protection from the coldonly
sion is more likely. Also, damaged trees, such as those
a thin layer of bark. Nevertheless, long periods of very
scarred by lightning strikes, fire, or logging activity,
cold temperatures at the right time of year can result in
are more susceptible. Some research suggests that the
high mortality of beetle larvae, and such cold-related
microclimate of closed, old-growth forests is a contrib-
mortality may be an important mechanism preventing
uting factor to initiating an outbreak. According to this
or terminating a bark beetle outbreak. No single lethal
hypothesis, closed canopies minimize air movement,
Mountain Forests
causing beetle-attracting odors to concentrate. Beetles
With less competition for light, water, and nutrients
are then attracted to such forests from a larger area, and
following the death of overstory trees, the understory
with high beetle densities, even the more vigorous trees
trees often grow much faster than they did before the
are susceptible to invasion.43 Under such conditions,
outbreak. For example, height growth rate of many sur-
beetle populations increase rapidly in one year. If the
viving lodgepole pine and subalpine fir trees doubled
forest is opened up by whatever means, the odor plume
after the mountain pine beetle outbreak in a northern
is dispersed and becomes less effective. Beetle parasites
Colorado study area, and radial growth rate increased
and beetle predation by birds may also be factors that
two- to threefold after a mountain pine beetle out-
reduce bark beetle populations.
break in the Greater Yellowstone Ecosystem.48 Modeling
Once an extensive outbreak is under way, the thou-
studies suggest that this rapid growth of the surviving
sands of adult beetles emerging from infested trees can
understory can restore pre-outbreak levels of basal area
overcome the defenses of healthy treeseven some rel-
within about 80 years.49 Recovery of basal area may be
44
atively small onesas well as those of unhealthy trees.
even faster under some conditions, as observed after a
The healthy hosts provide a superior food resource for
mountain pine beetle outbreak in Colorado in the 1980s.
the beetle larvae; hence, even more new adults emerge
Ninety percent of the basal area that had been present
the following year. When conditions are especially
prior to the outbreak had regrown after only 30 years in
favorable for the beetles, the outbreak may be so exten-
some lodgepole pine stands.50 In forests where lodgepole
sive that it becomes a regional or even continental phe-
pine dominated the pre-outbreak overstory but subal-
nomenon, as occurred in western North America during
pine fir and Engelmann spruce were more abundant in
the late 1990s and early 2000s.45 An outbreak usually
the understory, the composition of the post-beetle forest
ends only after a cold winter kills many larvae or when
shifts toward more fir and spruce; however, where lodge-
susceptible trees are no longer availableeither because
pole pine dominated both the overstory and the under-
the conditions causing host-tree stress have been ameli
story, lodgepole continues as the dominant species.51
orated in the area or because the beetles have simply

killed so many trees of suitable size. The last mountain
pine beetle outbreak in northern Colorado and south-
Major Forest Types
ern Wyoming apparently subsided because the beetles
Rocky Mountain forests are easily classified into sev-
could no longer find enough suitable trees; temperatures cold enough to terminate the outbreak were not
recorded.
An important feature of the disturbances caused
by most bark beetle species is that nearly all mortality
occurs in the larger trees, namely, those greater than
about 68 inches in diameter. Most smaller trees and
saplings usually survive, even where the beetles kill
most of the large trees in the overstory. For example,
severe mountain pine beetle outbreaks in lodgepole
pine forests of southeastern Wyoming and northern
Colorado during the 1990s and 2000s reduced the canopy by 10 to more than 70 percent in many stands.46
Nevertheless, even in the most severely affected stands,
the smaller trees that were not attacked were generally
eral major types (see table 11.1), each of which can be

further subdivided. The most thorough classification
was done using Rexford Daubenmires habitat-type
approach, where forests are classified according to two
factors: (1)the tree species that would dominate a stand
after a very long time without disturbance, when the
vegetation has presumably equilibrated with prevailing climate and soils, and (2) one or more understory
plant species that are indicators of other environmental
conditions. This classification assists in evaluating sites
for wood production, wildlife habitat, and other forest
values.52
Ponderosa Pine Forest
numerous enough to meet or exceed U.S. Forest Service
Ponderosa pine is the most widespread conifer in the
requirements for stand regeneration. Also, less competi-
western United States and is a major tree species at lower
tion from older trees enables the establishment of new
elevations throughout its range. In Wyoming it grows
tree seedlings.47
primarily in the eastern portions of the state, where
185
Fig. 11.8.On the left is an open ponderosa pine forest on

a moderately moist site in the Laramie Mountains south of
Douglas. The thick bark of mature ponderosa pine protects
many trees from easily controlled surface fires that kill
smaller trees, although some are scarred (see fig. 11.9). In the
same area, ponderosa pine forests develop extremely high tree

densities after long periods without fire or other disturbances
that coincide with climatic conditions favorable for seedling
establishment, as illustrated on the right. Such dense forests
are highly susceptible to severe wildfires. Elevation 6,700 feet.
summer precipitation is higher and the growing season
The importance of warmer temperatures was suggested
is warmer and longer (see figs. 1.5 and 3.4), especially
by a laboratory study that found that ponderosa pine
in the Black Hills and on the east slopes of the Bighorn
seedlings are more sensitive to cold temperatures than
and Laramie mountains. The importance of summer
were those of lodgepole pine, which is found at higher
precipitation is illustrated by the absence of ponderosa
elevations. 53 Probably for the same reason, ponderosa
pine on the southern end of the Bighorn Canyon, in the
pine is generally absent from the frost pockets in depres-
desert-like Bighorn Basin, and its presence at the same
sions, where cold air accumulates. Ponderosa pine is
elevation on the northern end of the canyon, where
a relative newcomer near the northeastern limit of its
summer rainfall is much higher (19 inches compared
distribution, such as in Wyoming, and probably is still
to 7 inches). Ponderosa pine is also abundant in por-
moving northward from Pleistocene refugia far to the
tions of the Uinta Mountains in Utah, the east slopes of
south (see chapter 2). Climate warming may accelerate
the Colorado Front Range, and throughout the moun-
this migration.54
tains of southern Colorado, New Mexico, and Arizona.
Tree density in ponderosa pine forests is a function
In all these areas, as on the Great Plains, a large por-
of local growing conditions and disturbance frequency
tion of the annual precipitation occurs in the summer.
and severity (fig. 11.8). On relatively wet sites, ponder-
Mountain Forests
osa pine can form a nearly closed canopy with a sparse

understory; on drier sites the forest is more open, or even
savanna-like, and often has a well-developed understory of shrubs, grasses, and other herbaceous plants.
Open ponderosa pine stands can also be maintained
by frequent surface fires that kill young seedlings and
saplings. The larger, older trees usually are not killed by
surface fires, owing to their thick bark and high crowns,
but they are often scarred by the fire. The scars can be
analyzed using dendrochronology to reconstruct local
fire history over the past several centuries (fig. 11.9).
Such fires burn the tops of understory shrubs and herbaceous plants, but the roots survive and soon produce
new sprouts, a type of vegetative reproduction that is
lacking in the conifers. Mountain pine beetle outbreaks
also thin ponderosa pine stands, although the resulting stand structure is different from that created by fire,
simply because the beetles tend to kill more of the large
trees and fewer of the saplings.
At the scale of an individual ponderosa pine stand,
fires commonly recurred at intervals of a few years to
a few decades prior to disruption of the historical fire
regime in the late 1800s.55 In some areas these fires were
predominantly low-severity surface burns that helped
maintain an open forest structure. Other ponderosa
pine forests were characterized by a variable-severity
fire regime that included a patchy mix of low- and
high-severity burning, both during individual fires and
in individual stands over time.56 High-severity (standreplacing) burning was an important component of the
fire regime primarily on wetter sitesduring dry years
Fig. 11.9. Ponderosa pine and Douglas-fir with thick bark

often survive surface fires, even though the base of the tree is
scarred. Fire scars develop when the heat scorches a portion
of the cambium, a layer of cells found on the inner side of
the bark that produces new bark and sapwood. Such scars can
be used to determine the years that fires occurred and the
number of years between successive fires. This photo of a ponderosa pine was taken in Devils Tower National Monument.
in the ponderosa pine zone, such as at higher elevations

and in mixed-conifer stands where other species, such
as lodgepole pine or Douglas-fir, were co-dominant. In
historically in the wetter northern part than in the drier
the Colorado Front Range, for example, fires at lower ele-
southern part.58
vations, especially near the prairie-forest ecotone, were
Severe burns were followed by either gradual or
predominantly low-severity surface burns that helped
rapid recovery of forest cover, depending on climatic
maintain low-density ponderosa pine forests. At mid-
conditions and seed availability. In some settings, a
elevations, ponderosa pine forests burned with variable
dense stand of ponderosa pine develops after a high-
severity, so that patches of stand-replacing fire were
severity fire; in other settings, such as on dry sites fol-
interspersed with areas of low-severity surface burn-
lowing severe burns, the post-fire stand may be sparse
ing. At the upper reaches of the ponderosa pine zone,
and slow to re-establish.59 Ponderosa pine seeds are rel-
where ponderosa transitions into lodgepole pine forest,
atively large and tend to disperse over relatively short
stand-replacing fires burned large areas of ponderosa
distances; consequently, the interior of a large patch of
and lodgepole pine forest.57 Similarly, in the Black Hills,
stand-replacing fire may become reforested very slowly,
dense stands and high-severity fires were more common
as has been seen after some recent fires in Colorado. The
187

Fig. 11.10. Douglas-fir forest
near Jenny Lake in Grand
Teton National Park. Elevation 6,850 feet.
establishment of new ponderosa pine seedlings tends to
Laramie Mountains near Casper and near Laramie Peak
be episodic, occurring primarily during periods of rela-
west of Wheatland, and a large fire of variable fire sever-
tively wet conditions, with or without fire.60 As a result,
ity burned near Laramie Peak in the summer of 2012.
a ponderosa pine forest often contains only three or
The same is true elsewhere in the region.
four easily identified age classes.

The earliest explorers surely saw a great diversity of
ponderosa pine forests as they traveled westward. How-
Douglas-fir Forest
ever, a combination of fire exclusion and heavy grazing,
Douglas-fir and ponderosa pine share several ecological
which enhances tree seedling establishment by reduc-
characteristics and sometimes they are found together.
ing competition from grasses and forbs, caused tree
Specifically, both species have a thick bark that often
density in many stands to increase during the twenti-
enables the trees to survive low-intensity surface fires.
eth century, especially on drier sites at low elevations
Also, the optimal environment for both is at lower eleva-
(see fig. 11.8). Wetter conditions also promoted seedling
tions, in the foothills and lower mountain slopes below
establishment and survival in areas that were logged in
about 8,500 feet (see fig. 11.1). Usually Douglas-fir for-
the early 1900s, when tree removal favored the growth
ests are found at slightly higher elevations than pon-
of young trees in many places.61 All these factors favored
derosa pine, or on somewhat wetter sites; for example,
the development of more extensive and uniform wood-
north-facing slopes often support relatively pure stands
lands than had occurred historically. Recent large fires,
of Douglas-fir, whereas on the opposite, south-facing
bark beetle outbreaks, and drought-caused tree mor-
slopes, stands may be dominated primarily by ponderosa
tality, as well as innovative new timber harvesting
pine. Limestone and other sedimentary substrata appear
methods, may be restoring some of the diversity that
to be especially favorable for Douglas-fir.62 Some research
characterized historical ponderosa pine forests. Nota-
suggests that Douglas-fir is more tolerant of competition
bly, since about 2000, extensive mountain pine beetle
for water, nutrients, and light in the understory than is
outbreaks have affected ponderosa pine forests in the
the pine, and that it can eventually replace the pine in
Mountain Forests
the absence of disturbance. In Wyoming, Douglas-fir is
matic conditions were very dry. For example, widespread
most common in the western half of the state, where it
fires in 1856 burned Douglas-fir forests in portions of
occurs just below the lodgepole pine forest zone. It is also
Jackson Hole, northern Yellowstone National Park, and
abundant on the east slopes of the Colorado Front Range,
the Centennial Valley just west of Yellowstone Park.64
and in adjacent Idaho and Montana, such as to the west

and north of Yellowstone National Park (fig. 11.10).
Insects can also cause variation in forests of Douglas-fir, especially the Douglas-fir beetle and the western
As with ponderosa pine forests, tree density and
spruce budworm (figs. 11.11 and 11.12). The bark beetles
understory cover in Douglas-fir forests vary with local
kill the larger trees, whereas the budworm kills mainly
environmental conditions and disturbance frequency
smaller individuals. Rarely do the insects kill all trees
and severity, ranging from sparse, savanna-like stands to
in a stand.65 Mixed-age Douglas-fir stands develop on
closed-canopy stands. Historical fire regimes were simi-
dry sites or where fires or insects have caused only par-
lar to those in ponderosa pine forests, with fire intervals
tial mortality of the canopy; even-aged stands (that is,
perhaps somewhat longer and high-severity fire perhaps
those made up of trees that are roughly the same age)
more common in Douglas-fir, especially on wetter sites.63
also develop, especially after high-severity fires or other
Large stand-replacing fires occurred in years when cli-
disturbances that kill all the trees.66
Fig. 11.11. Douglas-fir woodland in northern Yellowstone

National Park, 15 years after a severe outbreak of Douglas-fir
bark beetle. The beetles killed most of the larger trees in the
stand, which stimulated the growth of small trees, shrubs,
and herbaceous plants. Notably, in contrast to lodgepole pine,
Douglas-fir snags tend to break off rather than uproot. Photo
by Dan Donato.
Fig. 11.12. Larvae of western spruce budworm, a moth, commonly attack the buds and young needles of Douglas-fir, as seen
here in northern Yellowstone National Park in 2012. This budworm also attacks white fir in Colorado, Utah, and Idaho, and,
to a lesser extent, Engelmann spruce. It is said to be the most
widespread forest defoliator in western North America (Leatherman et al. 2009). Tree growth is slowed, but unlike bark beetles,
the western spruce budworm usually does not kill its host.
189
Fig. 11.13. An even-aged stand of 150-year-old lodgepole pine

above Centennial in the Medicine Bow Mountains. Tree-ring
data suggest that this forest was initiated following a fire. The
forest understory is dominated by dwarf huckleberry. Clusters
of small lodgepole pine are developing in openings. The blue

mark on the tree indicates that this tree will soon be cut,
probably for safety reasons. Elevation 8,400 feet.
As with ponderosa pine forests, tree density in many
Unlike the other conifers, lodgepole pine thrives on
Douglas-fir stands increased during the twentieth cen-
nutrient-poor soils (such as those that develop on gran-
tury as a consequence of fire suppression, grazing, and
ites and rhyolites) and is less common on more fertile
climatic conditions favorable for tree establishment. In
limestone-derived soils.68
some places, especially around meadows with big sage-
Lodgepole, as it is commonly called, has been viewed
brush, one can see the skeletons of sagebrush that died
as a pioneer species after fires or other severe distur-
for lack of sunlight as the Douglas-fir expanded into the
bances, because it generally does not tolerate the forest
shrubland. Recent fires and insect outbreaks also diver-
understory environment and is often the first tree to
sify stand structures.
become established after a disturbance, especially where

the soils are relatively dry and infertile. Under such con-
Lodgepole Pine Forest
ditions, lodgepole forms what is known as an even-aged

stand of trees. In fact, tree ages may differ by 20 years
The most common tree in the mountain forests of
or more, but the forest clearly became established after
northern Colorado, Wyoming, and much of the North-
a widespread disturbance. As the forests age and new
ern Rockies is lodgepole pine (fig. 11.13). It usually grows
gaps in the canopy are createdby localized windthrow
at mid-elevationsabout 6,00010,500 feet in north-
or other low-severity disturbanceslodgepole seedlings
ern Wyoming and about 7,00011,500 feet in southern
can become established and grow into the canopy.
Wyoming.
This pine is found at higher elevations on
In this way, tree establishment becomes episodic and
drier sites or in areas that have burned in the past cen-
patchy, resulting in a forest with multiple age classes.
tury or two. Engelmann spruce and subalpine fir may
The probability of lodgepole seedling establishment in
be co-dominant with lodgepole on relatively moist sites.
the forest understory appears to increase with moisture
67
Mountain Forests
Fig. 11.14. Lodgepole pine forests along the Rockefeller

Parkway between Yellowstone and Grand Teton national
parks. The stand on the left originated after the fires of 1988,
24 years before this photo was taken. The young trees are
very dense, because many of the pre-fire trees had serotinous cones, and also because the conditions for seedling
establishment during the two years after the fire were

favorable. The stand on the right has not burned for more
than 250 years. Many of the dead trees visible in the older
stand have been killed by mountain pine beetles during the
past 10 years. A riparian meadow and shrubland are in the
foreground.
availability, even without a canopy gap, but so does the
closed and on the tree for many years until higher than
chance of spruce and fir establishment.
usual heat causes the cone scales to open, such as dur-
Usually fires in lodgepole pine forests are infrequent,
ing a forest fire.70 The cones also open after falling to
separated by many decades or centuries in any given
the forest floor if they are exposed to the sun for a long
stand. This is primarily because of the cool, relatively
enough period.71 Numerous cone crops accumulate in
moist environments inhabited by lodgepole pine,
the treetops as the forest develops. Most of the seroti-
where snow persists until late spring and frequent sum-
nous cones open following a fire, and the seed is dis-
mer rain showers commonly keep fuels too wet to sup-
persed by wind at a time when a mineral soil seedbed
port extensive fire in most years. Only in exceptionally
and low competition for resources help to ensure seed-
dry years do large fires occur in lodgepole pine forests.
ling survival. At such times, more than a hundred thou-
Under these conditions, fires tend to be severe and
sand seeds may be dispersed per acre in a single year.72
stand-replacing, often burning over large areas.
69
Interestingly, not all lodgepole pine trees produce
In contrast to ponderosa pine and Douglas-fir, lodge-
serotinous cones. Individual trees tend to have either all
pole pine trees have thin bark and are easily killed by
serotinous or all nonserotinous cones, though a few trees
even low-intensity fires. However, lodgepole is well
may produce both types.73 The proportion of serotinous
adapted as a species, if not as individuals, to a high-
trees in stands of lodgepole ranges from 0 to nearly 100
severity fire regime because of its ability to produce
percent, with the highest proportions in young stands
serotinous cones (figs. 11.14 and 11.15). These cones are
at lower elevations, where the last major disturbance
produced more or less continuously, but they remain
was fire. The lowest proportions are in older stands at
191
Fig. 11.15.On the left, serotinous cones on a lodgepole pine

branch that has recently fallen to the ground. The diameter of
the branch indicates that the cones have remained closed for
many years, accumulating seed on the tree. One of the cones
is beginning to open, most likely because of higher temperatures at the soil surface than in the trees canopy. Cones that
remain on the tree typically open with the heat of a fire, as

shown in the center photo. The pine needles have burned,
but only the exterior of the cone scales was blackened. In
contrast, on the right, the green leaves around the open cone
indicate that the heat of a fire was not required for seed dispersal from the cones of this tree.
higher elevations, where the last major disturbance was
such as sagebrush-dominated shrublands. Some have
a windstorm, insect outbreak, or other nonfire distur-
suggested that forest management, in both wildlands
bance.74 To some degree, this pattern is genetically con-
and timber production areas, should allow for both fire
trolled and reflects long-term differences in selection
and nonfire disturbances, so that both genotypes can
pressures in different environments. Because fire inter-
be sustained, thereby maintaining biological diversity.
vals generally become longer with increasing elevation,
Seeds in both serotinous and nonserotinous cones
reaching 200300 or more years, trees growing at high
can be burned by intense fire, but rarely does fire kill
elevations are more likely to reproduce successfully from
all seeds of either type. Because the size of the can-
seeds dispersed into canopy gaps from nonserotinous
opy seed bank varies across the landscapeprimarily
cones. In contrast, at lower elevations, where typical fire
because of variation in the proportion of serotinous
intervals are less than 200 years, canopy gaps are less
trees and because of seed mortality during some fires
common and serotinous cones may be a more successful
the density of lodgepole pine seedlings after a fire varies
adaptation.
greatly. Seedling densities after the 1988 Yellowstone
As described previously, selection for serotinous
fires ranged from less than 5 per acre to greater than
and nonserotinous cones also may be influenced by
100,000 per acre. The fewest seedlings were found
seed predators, notably the red squirrel (see fig. 11.4).75
where the burned canopy was a mix of lodgepole pine
Considerably more seed is produced per year in non
and other species, essentially all lodgepole pine were
serotinous cones than in serotinous cones, a pattern
nonserotinous, and the fire burned through the canopy
that can be attributed to the need for allocating more
at high intensity. The most seedlings occurred where
energy to protective tissues in the serotinous cones
the burned canopy was essentially all lodgepole pine,
to minimize seed consumption by squirrels.76 Having
most trees were serotinous, and the fire scorched but did
both cone types in a stand seems adaptive for lodgepole
not consume all of the canopy.77
pine, as it ensures a seed source regardless of the type
As the seedlings grow into saplings and small trees,
of disturbance. Nonserotinous cones also enable the
the very dense post-fire stands become difficult to walk
establishment of lodgepole pine in nonforest habitats,
through. Such dense pine forests are commonly referred
Mountain Forests
to as doghair stands (fig. 11.16). The growth of individual

trees is slow in such places, but most survive and continue to produce seed for well beyond a century, though
remaining less than 4 inches in diameter. Not all lodgepole pine stands are so dense, however, as densities vary
greatly across the landscape. Historical reconstructions
in Yellowstone National Park revealed that, over time
spans of about 200 years, varying densities converge on
an average of about 500 trees per acre.78 Stands with different densities become more similar over time because
of self-thinning of initially dense stands and gradual
infilling of initially sparse stands (that is, by the establishment of young trees in older stands). Other disturbances affecting landscape patterns in lodgepole forests
include comandra blister rust, dwarf mistletoe, bark
beetles, root rot, and windstorms.
The effects of twentieth-century fire exclusion are
not nearly as significant in lodgepole pine forests as
in ponderosa pine and Douglas-fir forests, primarily because fires were infrequent and typically of
high severity in lodgepole pine forests even before
EuroAmericans ushered in fire suppression efforts.
Also, the fire regime is controlled more by weather and
climate than by ignition sources or variation in fuel
conditions.79 It is true that many fire ignitions have
been suppressed in the lodgepole pine zone, and that
some fires ignited in lower-elevation forests might
have burned up into the lodgepole had they not been
suppressed at the source. However, in most years such
fires would have been small, because the fuels at the
high elevations occupied by lodgepole pine historically were too wet for extensive fires. In dry years when
the weather is conducive to fire, the normal fuel struc-
Fig. 11.16.The trees in this dense stand of lodgepole pine

are small but are about 100 years oldprobably because the
previous forest that burned had a large number of trees with
serotinous cones. Also, the fire was not intense enough to
burn the cones and their seed, and the climatic conditions
after the fire were ideal for seedling establishment. With
about 6,000 trees per acre, there are few understory plants.
Unlike the dense ponderosa pine forest shown in fig. 11.8,
lodgepole pine forests such as this one are not caused by fire
suppression. Photo taken southwest of Dry Park in the Medicine Bow Mountains. Elevation 9,160 feet.
ture in lodgepole pine forests allows fires to spread

quickly and they are not readily suppressed. Had there
been no fire suppression during the twentieth century,
Spruce-Fir Forest
young lodgepole pine stands would be somewhat more
Engelmann spruce and subalpine fir often dominate
abundant today and old stands somewhat less sobut
forests at higher elevations and in cool ravines, most
todays distribution of stand ages does not differ greatly
likely because their seedlings tolerate the lower temper-
from what would have been seen in previous centuries.
atures and because more water is available longer into
Moreover, recent large fires in lodgepole pine forests
the summer than elsewhere (fig. 11.17).81 Both appear to
have created extensive stands of young forests (see
have lower water-use efficiencies than lodgepole, which
chapter 15). As discussed in the next chapter, climate
means that they require larger amounts of water for
change could increase the frequency and intensity of
every gram of new plant material they produce. Conse-
fires if the summers become drier, as predicted because
quently, the pine survives on drier sites at lower eleva-
of warmer temperatures.80
tions, where water stress is likely to develop. Ponderosa
193
Fires are less frequent in the spruce-fir zone than in

the lodgepole pine zone, because of even wetter conditions in the subalpine environment. Many spruce-fir
forests persist for centuries without fire. For example,
a tree-ring-based fire history study in a 75,000-acre
sub
alpine portion of Rocky Mountain National Park
recorded no significant fires during the previous 400
years in 27 percent of the forested area.82 This pattern
of few fires was true before the advent of efforts to put
them out, but those that did occur, and that occur
today, tend to be very hot and stand-replacing, just as
in lodgepole pine forests.
All four tree species in subalpine forestsspruce, fir,
aspen, and lodgepole pinemay become established
rapidly after a fire, depending on the availability of seed
Fig. 11.18.Occasionally subalpine fir and Engelmann spruce

develop clones, such as the two shown here. This occurs when
lower branches are pressed against the ground by snow and
new roots begin to grow, a process known as layering. A new
tree then develops from the rooted branch. Meadow plants at
this site in the Medicine Bow Mountains include alpine bistort, alpine timothy, and Idaho fescue. Elevation 10,400 feet.
Fig. 11.17. Subalpine forests dominated by Engelmann spruce
and subalpine fir are common at higher elevations, such as in
this area in the Medicine Bow Mountains. Note the presence
of downed wood, small and large trees, and the continuous
fuel ladder from the understory to the canopy. The spruce
beetle, a native insect, has recently killed many large trees in
this stand. The bright green plant in the understory is dwarf
huckleberry, an indicator of a relatively cool, moist environment. Elevation about 9,000 feet.
pine and Douglas-fir also are better adapted to the lowerelevation environment.
Unlike the pines and Douglas-fir, spruce and fir are
capable of vegetative reproduction when lower branches
are pressed to the ground by snow or a fallen log. The
branch then develops roots, known as adventitious
roots, after which the end of the branch begins to grow
upright into a new tree. Eventually the branchs connection to the parent tree is severed by death of the initiating branch and its decomposition. This process, known
as layering, produces clones of the original tree that are
commonly found locally at higher elevations. Clusters
of subalpine fir in meadows often can be attributed to
this cloning process (fig. 11.18).
Mountain Forests
and moisture.83 If lodgepole pine or aspen dominates the
spruce that the beetles generally do not kill. The growth
stand first, they will persist for a century or more. How-
of other plants is stimulated as well, for example, aspen,
ever, they are less tolerant of the understory environ
lodgepole pine, and numerous shrubs and forbs.
ment that typically develops at higher elevations and
Extensive areas of spruce-fir forest can also be affected
are gradually replaced by spruce and fir if no subsequent
by mortality of subalpine fir caused by the western bal-
stand-replacing disturbance occurs. One paleoecological
sam bark beetle (Dryocoetes) and its associated fungal
study, based on pollen analysis, found that dominance
pathogens. As with other bark beetles, warm, dry cli-
alternated over the centuries between lodgepole pine and
matic conditions favor this insect.88
spruce and firwhich may reflect post-fire dynamics or
The relative abundance of spruce and fir varies
shifting tree species distribution in response to climate
greatly from place to place. In stands older than about
changes.84 Spruce-fir forests also have been observed to
100 years, the largest and oldest trees typically are
develop on former meadows, sometimes following the
Engelmann spruce, as this species may live 500 years or
initial establishment of aspen.85 In much of the spruce-
longer.89 In contrast, subalpine fir is usually more abun-
fir zone, however, lodgepole pine and aspen are not pres-
dant, but the trees are smaller and younger, rarely more
ent. In such places, spruce and fir become re-established
than 250 years old. Fir seedlings and saplings may also
in the burned area without any competition from aspen
be 1020 times more numerous than those of spruce.
or lodgepole pine. Research in northern Colorado sug-
Apparently the roots of fir seedlings are better able to
gests that after 100200 years, on some sites, spruce no
penetrate the considerable litter that accumulates on the
longer reproduces in the forest understorya period
forest floor, whereas spruce seedlings are usually found
described as the spruce exclusion phase. After another
where mineral soil has been exposed, such as around
100 years, many of the dominant fir and spruce begin
the tipped root system of fallen trees. Some investiga-
to die, creating canopy gaps that initiate the spruce re-
tors have also observed spruce seedlings on rotting logs
initiation phase. The final phase is a second-generation
at sites where the decomposing wood remains wet well
spruce-fir forest which persists until the next severe dis-
into the summer, such as in moist ravines.90 Spruce pro-
turbance.
duces especially large numbers of seed every 25 years,
86
The likelihood of such a long interval with
no major disturbances caused by blowdown or insect

outbreak is low.
fir about every 3 years.

That subalpine fir reproduces more effectively than
Insect outbreaks also shape the structure and dy
spruce in the forest environment raises the question
namics of spruce-fir forests. Especially important is
of how spruce persists as a co-dominant. The answer
the spruce beetle, a bark beetle with a life cycle simi-
lies in their different life history strategies.91 Fir is
lar to the mountain pine beetle (described earlier in
shorter lived but produces many seedlings, whereas
this chapter).87 The spruce beetle attacks older stands
Engelmann spruce compensates for generally poor
with a high proportion of Engelmann spruce in the
reproductive success with increased longevity. Thus,
overstory. Susceptible stands usually display slow tree
the greater proportion of subalpine fir seedlings and
growth, suggesting that the trees are of low vigor and
saplings does not mean that fir will eventually become
are unable to produce sufficient resin to exclude the
the sole dominant.
beetles. Unlike mountain pine beetles, which reproduce
Engelmann spruce and subalpine fir are both com-
successfully only in live trees, spruce beetles can pro-
paratively tolerant of the understory environment; that
duce large numbers of offspring in trees that have died
is, young trees are tolerant of shade and competition for
recently from other causes. Moreover, an abundance of
water and nutrients. Therefore, the two species usually
dead trees, whether from logging or windthrow, may
develop uneven-aged forests, with both young and old
provide the energy base for the initial development of
trees. Because the overstory trees are the same species
a spruce beetle outbreak that spreads subsequently into
as in the understory, it is tempting to view spruce-fir
live trees. The infestation opens the canopy, greatly
forests as being in a state of equilibrium, with dying
accelerating the growth of the abundant subalpine fir
overstory trees being replaced with younger trees of the
(which is not a host for the spruce beetle) and the small
same species. However, all forests are subject to ongoing
195
changes caused by fire, insects, diseases, windthrow,

and other factors.
As with lodgepole pine forests, twentieth-century
fire exclusion had a relatively minor influence on the
structure and dynamics of spruce-fir forests. Timber
harvesting has been less widespread in subalpine forests; consequently, they appear today much as they
would have in the early 1800s.
Aspen Forest
Aspen forests extend from the foothills to the subalpine
zone, typically in depressions, ravines, valley bottoms,
and the lee sides of ridgeswhere water availability is a
little higher. Because of this distribution pattern, many
ecologists have assumed that aspen requires more water
than other trees do. In fact, adult aspen use considerably less water per unit leaf area than lodgepole pine,
subalpine fir, and Engelmann spruce.92 The general
restriction of aspen to wetter areas is probably due more
to the intolerance of aspen seedlings to drought than to
the intolerance of mature trees.
Aspen is uniqueit is the only deciduous tree in the
region that grows in an upland environment that seems
to favor evergreen plants. The loss of all leaves each fall
is not efficient with regard to nutrient conservation, and
this may partially explain why aspen seems restricted to
depressions, where nutrients are more readily available.
As with conifers, aspen twigs are probably capable of
reabsorbing nutrients from the leaves before they fall,
Fig. 11.19. Some aspen forests have an understory of suba lpine

fir or, in this area on the west slope of Wyomings Sierra
Madre, lodgepole pine. In the absence of another fire or harvesting of the conifers, the area dominated by aspen gradually
becomes smaller and conifer-dominated forests become more
extensive. Photo taken after leaf fall. Elevation 8,500 feet.
an important mechanism for retaining at least some

of the limiting nutrients in the leaves. Also, although
In Wyoming, aspen usually occurs in small groves
aspen loses its leaves in the fall, there is chlorophyll in
(figs. 11.19 and 11.20). However, extensive forests are
the barkeasily seen after scratching the bark. It was
found in a few areas, notably on the west slope of the
previously thought that bark photosynthesis was an
Sierra Madre south of Rawlins.94 Aspen abundance there
important component of the trees total annual pho-
may be enabled by the Arizona Monsoon, which brings
tosynthesis, especially during the leafless period. How-
more reliable summer precipitation to this part of the
ever, recent research indicates that little carbon dioxide
state (see chapter 3). Southward into Colorado and New
penetrates the transparent but corky bark outside the
Mexico, aspen becomes increasingly common, perhaps
layer of chlorophyll. Instead, the chlorophyll in the
because of more summer and fall rain. There, it tends
bark converts some of the carbon dioxide produced by
to occupy the same topographic positions as lodgepole
the trees own internal respirationa process known
pine does in the Central and Northern Rockies.95
as refixation. In this way, the bark contributes to the
The tendency for aspen to occur in small groves in
trees energy budget.93 The plant is effectively evergreen
Wyoming can be explained in part by its clonal nature,
despite having deciduous leaves. As noted previously,
with new trees developing from root sprouts, also
dwarf huckleberry also is a deciduous evergreen.
known as suckers. Seedling establishment is rare, not
Mountain Forests
Fig. 11.20. Aspen groves in
Wyoming tend to be small,
except on the southwestern
slopes of the Sierra Madre,
east of Baggs. Here the aspen
grow over large areas, similar
to the aspen farther south in
Colorado and New Mexico.
Patches of Gambel oak also
are found in this area. Note
that some of the aspen clones
are still green (see text for
explanation).
because there is a lack of seed but because the environ-
the ramets of some clones (genets) may intermingle.97
96
mental requirements for establishment are rarely met.
Genetic diversity undoubtedly explains why aspen is
Aspen seedlings are delicate and cannot tolerate even a
found over such wide environmental gradients in the
brief period of desiccation. They often die before their
Rocky Mountain region and, in fact, why it is the most
roots can penetrate through forest floor litter to the
widespread tree in North America.
mineral soil. Also, they may be unable to compete with
Because of aspens ability to sprout, it dominates
other plants for limited water. Whatever the mecha-
temporarily after severe disturbances in many conifer-
nisms that limit seedling establishment in undisturbed
ous forests, most likely because the sprouts have more
forests, aspen seedlings have been observed in recently
stored energy available for rapid growth than do conifer
burned forests where mineral soil is exposed and poten-
seedlings. More than 10,000 sprouts per acre may reach
tially competing plants have been temporarily reduced,
a height of 3 feet or more within a year following a fire.
such as in Yellowstone National Park after the 1988 fires
Conifers, especially subalpine fir, sometimes persist
(see chapter 15). Seedlings also have appeared on sedi-
in the understory and eventually overtop the aspen
ments exposed in abandoned beaver ponds.
because its seedlings become established on the aspen
With sprouts as the principal form of regeneration,
forest floor more readily than do those of other trees
nearly all groves develop as a genetically uniform clone
(see fig. 11.19). Lodgepole pine and Engelmann spruce
or a mix of several discrete clones. Essentially, what
also replace aspen under some conditions.98
nearly everyone refers to as a tree is more appropri-
Where other trees do not become established, aspen
ately termed a branch or shoot. Botanists use the term
continues to dominate stands indefinitely, with new
ramet. Although ramets typically die after about 100
sprouts replacing the senescing older stems. Generally,
years, the plant itselfthe genetlives for much longer,
most aspen sprouts do not survive in the understory
most likely for centuries. Thus, more genetic variabil-
shade of aspenan interesting anomaly, as it would
ity exists between aspen groves than within one grove,
seem that the small trees could tap the energy stored
as is apparent when, for example, two adjacent clones
by the larger trees (which have leaves fully exposed to
lose their leaves at different times in the fall. Of course,
sunlight). Plant hormones produced by the dominant
197
shoots apparently suppress the growth of the smaller
of the conifers, more prescribed fires, or reductions in
ones belowa case of apical dominance. Aspen stands
ungulate numbers.
burn periodically, at intervals of decades or centuries,

and the thin-barked adult stems are usually killed.99
However, abundant post-fire sprouting and rapid growth
Mountain Forests and Wildlife
result in the re-establishment of an aspen grove within
The forest-meadow-wetland mosaic provides essen-
a few decades. Chronic heavy browsing on the aspen
tial habitat for numerous birds and mammals. A few
sprouts by wild and domestic ungulates can inhibit this
are found there all year long, such as the blue grouse,
process of aspen regeneration (see chapters 10 and 15).100
boreal owl, Clarks nutcracker, gray jay, red crossbill,
Old landscape photos from the late 1800s and early
beaver, lynx, marten, red squirrel, snowshoe hare, and
1900s commonly show more aspen than is visible in the
voles. Others migrate to the foothills for the winter,
same scenes today, suggesting that aspen is declining
escaping deep snow and colder temperatures, returning
in at least some areas. This comparison may be mis-
to the mountains in the summer for more reliable for-
leading, simply because the late 1800s was a time of
age and better hiding cover. Elk, mule deer, and cou-
extensive burning throughout the West, and post-fire
gar are notable examples. Many birds, such as warblers,
aspen probably was exceptionally abundant around
thrushes, kinglets, and white-crowned sparrow, migrate
1900, compared with earlier in the nineteenth century
to warmer southern climates for the winter, along with
or in previous centuries. Thus, aspen abundance at the
the few ducks and mergansers that use mountain lakes
beginning of the twentieth century is not necessarily
and beaver ponds. As noted, some birds and small mam-
an appropriate historical benchmark for land manage-
mals rely on conifer seed for their food. Others feed on
ment.101 Moreover, photos from other areas show that
mushrooms or mature insects, or dig out insect larvae
aspen cover remained stable or even increased in some
from decomposing wood or the bark of living trees. The
places during the twentieth century.
Some resource
abundance of insects emerging from wetlands provides
managers are concerned that twentieth-century fire
food for birds that are agile enough to catch them. Even
exclusion and other factors have led to deterioration of
bears are known to eat moth larvae, which can be abun-
many aspen groves, as the adult aspen stems gradually
dant under the rocks on talus slopes.
102
die and are replaced by conifers or shrubs instead of
Commonly, mountain wildlife is secretive and not
new aspen sprouts. Aspen shoots can be killed by vari-
easily observed. As in other ecosystems, the best time
ous pathogens, including root rot and canker diseases,
to find them is in the spring, when the birds can be
especially if the bark is damaged. This trend of stand
located by their singing, or during the winter, when
deterioration is especially worrisome if the capacity for
many species congregate in flocks, or footprints can be
sprouting diminishes as stands age, or if the intensity of
observed in the snow. Aspen groves are often identified
ungulate browsing or beaver cutting is high (see chap-
as hotspots of biological diversity, perhaps because of
ters 10 and 15).103
the abundance and diversity of understory plants, the
The first decade of the twenty-first century saw exten-
frequent proximity of aspen to water, and the relative
sive death of aspen trees in Wyoming and across much
ease with which cavities can be found or created for
of the West, apparently triggered by the severe drought
nesting. In all kinds of forest, many animals depend on
of 20002003.104 The patterns and implications of this
dead trees, whether for perches or cavities.106 For many
mortality event, referred to as sudden aspen decline (SAD),
species, the juxtaposition of wetlands, meadows of vari-
are discussed in chapter 12. Despite such developments
ous kinds, and forests of different ages and types makes
in aspen forests, there seems little need to be concerned
for the perfect habitat. As always, the landscape mosaic
about aspen extinction.105 Still, the tree could become
must be considered, not just a few acres of meadow or
less abundant in some areas as aspen-dominated forests
forest.
gradually change to conifer-dominated forests or shrub-
Overall, forests of all kinds cover about 15 percent of
lands. To maintain the current abundance of aspen

forests in the landscape may require more harvesting
Wyoming and are found primarily in the mountains,

where temperature, moisture, and nutrient conditions
Mountain Forests
are sufficiently favorable to enable tree seedling estab-
amenities have led to intensive management in some
lishment and growth. Although only six tree species are
areas, and to concerns about whether current manage-
common in these forests, their adaptations vary consid-
ment practices will sustain wildlife populations as well
erably, and the influences of fires, insects, windstorms,
as wood production. Recent large fires, insect outbreaks,
and abrupt topographic changes create a mosaic that
and the specter of climate change raise questions about
includes a diversity of forest types intermingled with
the resilience of Rocky Mountain forest ecosystems, at
meadows and shrublands, along with mountain lakes
least as we know them todaya topic that is discussed
and alpine tundra. Demands for forest products and
further in the next chapter.
199
Chapter 12
The Forest Ecosystem
Thinking about forest ecosystems is less intuitive than
thousands of gallons of water and tons of chemicals
considering the adaptations and relative abundance of
streaming upward through tree trunks, photons of
different kinds of plants and animals, as in the previous
energy absorbed by leaves and put to work evaporat-
chapter. However, ecosystem science enables a better
ing water through leaves and fixing energy in organic
understanding of the factors determining tree growth,
compounds, food manufactured in leaves streaming
the development and maintenance of soils that allow
to growing points, insect predators quietly nibbling
that growth, the amount and quality of water flowing
away, rocks being broken down into useable nutri-
from watersheds, the effects of disturbances (such as
ents, microbes disassembling organic compounds
fires, insect outbreaks, and timber harvesting), and the
and freeing nutrients for reuse, all species playing
effects of climate change. Curiously, dead plants and
out their roles in reproduction, the forest ecosystem
animals are just as important as those still alive, if only
grudgingly restocking the forest stream with water,
because they provide microhabitats for various forms of
and a million other things.1
life. In some cases the dead plant materialthe O horizon of the soil (commonly referred to as the forest floor,
litter, or debris)functions as an important mechanism
for erosion control and for conserving nutrients that
may be limiting.
The intrigue of forest ecosystems, or any ecosystem
for that matter, is that the important processes are not
visible. To obtain data, scientists use various instruments to measure processes such as solar radiation (in
the wavelengths used for photosynthesis), the rates of
plant growth and litter decomposition, and the movement of nutrients and water. Ones perspective on a
forest changes when such information is revealed. The
late F. Herbert Bormann, eminent ecosystem scientist at
Yale University, wrote:
Energy Flow, Productivity, and Carbon Sequestration

Most of the suns energy reaching a forest, or any other
terrestrial ecosystem, is used for evaporating water and
heating the environment; a mere 13 percent is used for
photosynthesis. In mountain forests, because of high
elevation and a comparatively thin atmosphere with
low humidity, much of the heat gained during the day
is lost at night by radiation back into space. The result is
a relatively cool environment. Also, water is a limiting
factor in many places by mid- to late summer. Consequently, because of a cool, relatively dry environment,
tree growth rates are low in the Rocky Mountains compared to rates in other places in North America, such
Today when I visit a forest, there is . . . a sense of
as the Pacific Northwest and the Southeastern Coastal
being surrounded by an enormous dynamism:
Plain (fig. 12.1). Annual variation in wood produc-
200
The Forest Ecosystem 201
NPP (t/ha/yr)
Live tree
biomass
15
Scots pine
10
Atmospheric
deposition
0.3
Slash pine
5
Lodgepole pine
Fixation
0
25
50
0.01
0.6
Aboveground
litter
1.2
75
STAND AGE (years)

Fig. 12.1. Rate of forest plant growth, mostly tree growth
but also that of understory shrubs and herbaceous plants, is
known as net primary productivity (NPP)expressed here as
metric tons per hectare per year. NPP changes significantly as
forests age. Note the low growth rate for Wyoming lodgepole pine forests compared to slash pine in Florida and Scots
pine in northern Europe. One metric ton = 2,205 pounds;
1 hectare = 2.47 acres. Adapted from Knight (1991).
0.9
Fixation
0.01
Soil
organic
matter
tion can be observed in the annual rings of trees and

is caused mostly by annual variation in growing season temperature and precipitation. Also, younger forests tend to have higher growth rates than older forests.
Growth rate, also known as net primary productivity
(often abbreviated as NPP), is commonly expressed as
the average rate at which all new plant biomass is added
to the forest annually, whether in the form of trees,
shrubs, grasses, or forbs.2
Living trees, soil organic matter, and dead plant material on the forest floor are the largest components of
the forest biomass and change through time after a dis
turbance (figs. 12.2 and 12.3). The forest floor biomass is
composed almost entirely of leaves, twigs, branches, and
the boles of fallen trees. Some of itnot allis burned
during a forest fire, but the litter accumulates in total
weight for many years. This litter buildup occurs because
winters are long; summers are relatively dry; and most
plant tissues in Rocky Mountain environments have high
lignin content and high carbon-to-nitrogen ratios, both
of which slow the rate at which bacteria and fungithe
primary decomposerscan convert organic materials
to inorganic molecules. Microbial decomposition is also
limited by the relatively cool and sometimes dry conditions that prevail in the summer. Notably, considerable
Outflow
0.01
Fig. 12.2.Carbon and nitrogen distribution in a lodgepole
pine forest. The size of the boxes indicates the relative
amounts of organic matter in live tree biomass, aboveground
litter, and soil organic matter (which includes bacteria and
fungal biomass but not live root biomass). Roots are part
of the live tree biomass box. Shrub and herbaceous plant
biomass is not shown but is less than 1 percent of the tree
biomass. Annual nitrogen flows are shown by the arrows;
the numbers are grams per square meter per year. Much of
the nitrogen in the rooting zone is in organic matter that
is not readily decomposable. Note that nitrogen inputs are
larger than nitrogen losses, suggesting that nitrogen is accumulating in the tree biomass, litter, and/or soil organic
matter, probably because it is a limiting factor for plants and
microbial organisms. Also, note that the tree uptake estimate
is larger than the sum of the input estimates to the rooting
zone, which suggests that the amount of soil nitrogen is
gradually depleted in this forest as biomass accumulates. The
soil nitrogen pool is replenished as the forest ages further or
following such disturbances as fire. Based on data in Fahey et
al. (1985) and Fahey and Knight (1986).
Biomass Increment (t/ha/yr)
(such as the red squirrel and red-backed vole) commonly
Live trees
Forest floor litter

Dead trees and
downed wood
graze on mushrooms, truffles, and puffballs, the reproductive structures of fungi that decompose the detritus,
and birds commonly prey on these small mammals and
various invertebrate detritivores. Moreover, evidence
suggests that these animals are important in dispersing
the spores of fungi, which in turn are important not

only for decomposition butin the case of mycorrhizal
fungialso for the establishment and growth of new

plant seedlings, including those of all Rocky Mountain
trees.4
-1
The importance of mycorrhizal fungi is suggested

by estimates that up to 15 percent of the net primary
-2
40
80
120
Stand Age (years)
160
200
Fig. 12.3. Rates of biomass increment or decline in live trees,

forest floor litter, and dead trees and downed wood change
as forests grow to maturity and become old-growth forests.
Older forests have more dead trees and downed wood, but
tree growth slows as forests age (see also fig. 12.1). Dead trees
and downed wood, sometimes referred to as snags and coarse
woody debris, decline soon after fire, as decomposition rates
of this material are relatively high when water and nutrients
are more readily available for the decomposers. The forest
floor litter increases initially after a firewhich commonly
burns much of the pre-fire forest floorbut then remains
about the same, as litterfall becomes approximately equal to
the rate of litter decay. The patterns illustrated will be somewhat different for forests growing on different soils, at different elevations, or in different topographic positions. These
graphs are based on data collected from several lodgepole pine
forests at an elevation of about 9,500 feet in the Medicine Bow
Mountains. Rates of change are expressed as metric tons per
hectare per year; see fig. 12.1 for unit conversions. Adapted
from Pearson et al. (1987); see also Smith and Resh (1999) and
Kashian et al. (2013).
productivity in a coniferous forest goes to the maintenance of these fungi on roots.5 All studies thus far
indicate that, rather than being parasites, mycorrhizal
fungi develop a mutually beneficial association with
their hosts.6 The fungi derive energy from the plants in
the form of carbohydrates, while the fungal filaments,
known as hyphae, extend beyond the roots and enhance
water and nutrient uptake. The decaying biomass of the
fine roots and associated fungi may contribute more to
nutrient availability than do decaying leaves, twigs, and
branches.
On average, only 2 percent or less of the energy fixed
by plants during photosynthesis flows through animals
in terrestrial ecosystems, whether forests, shrublands, or
grasslands, yet animals often influence the ecosystem
in important ways. An example is the planting of whitebark and limber pine seeds by Clarks nutcracker (see
chapters 10 and 11). The populations of different herbivores and carnivores continually fluctuate, with the
result that their influences are greatly amplified when
they are abundant, such as during outbreaks of bark
decomposition occurs under snow during the winter, as
beetles. A less well-known example occurs when the
some of the decomposers are able to function at the soil/
populations of porcupines are high. Porcupines feed on
snow interface.3 Because detritus accumulates over time
tree bark, resulting in reduced flow of carbohydrates to
and is flammable when dry, fire is another important
the roots, which lowers plant growth in the forest as a
pathway for energy flowa decomposition process that,
whole.7 Ecosystems are dynamic, with all components
like bacteria and fungi, releases the inorganic nutrients
changing over time, as does the amount of energy flow-
required for new plant growth (fig. 12.4).
ing through the detrital, grazing, and fire pathways.
The combined biomass of forest floor litter, fungi,

and bacteria is far greater than the total biomass of
vertebrates and invertebrates. Not surprisingly, various
Hydrology of Forest Landscapes
animals have evolved to depend on detritus for a signifi-
Mountain landscapes are hydrologically distinctive
cant part of their energy. For example, small mammals
from the surrounding lowlands, because they receive
Fig. 12.4.Importance of nutrients and water for the growth

of lodgepole pine. In the photo on the left, an intense fire
has burned the forest floor and some of the downed wood
buried in the litter. The photo shows a pine root that had
grown horizontally in the top layers of the soil. When the
root encountered a decomposing log, now consumed by the
fire, it turned 90 degrees as it grew into the logmost likely
because the log provided a better source of nutrients or water.
In this case, the log was burned but not the root. In the photo
on the right, a dense stand of new pines is developing after
a clearcut that occurred about 12 years previously. However,
adjacent to the uncut forest is a band with very little vegetation around the entire perimeter of the clearcut, even with
full sunlight. This pattern is common and suggests that the
roots of the mature forest trees grow outward from the trunk,
to a distance of about 12 feet, and that the roots provide too
much competition for the establishment of tree seedlings and
other plants. Competition for water and nutrients may be
more important than competition for light in the understory
of lodgepole pine forests. The trees in the uncut forest on the
left are small because of their high density but are about 100
years old (see fig. 11.16).
more water than can evaporate during the year. In other
barely sufficient to wet the forest floor, to say nothing
words, the ratio between precipitation and evaporation,
of wetting the soil. In contrast, snowmelt normally
known as the P-E ratio, is greater than one. Conse-
saturates the soil, and then soil moisture declines
quently, the mountains are the regions primary source
more or less steadily during the summer and early fall,
of river water and groundwater, on which agriculture,
whether the vegetation is a forest, woodland, meadow,
industry, and municipalities depend as well as ripar-
or alpine tundra. 8 The actual amount of surface water
ian and wetland ecosystems. The mountains have been
draining from a watershed depends on at least seven
referred to as the water towers of the West.
factors: water content of the snowpack, potential for
Because snow historically has accounted for 5075
snow drifting, type of vegetation and the amount of
percent of the annual precipitation in the mountains,
water transpired, water-holding capacity of the soil,
streamflow has a pronounced flood peak during snow-
weather conditions, proportion of water that perco-
melt in the spring (fig. 12.5). Summer rains normally
lates into groundwater, and patchiness of the land-
contribute little to streamflow, because most rains are
scape mosaic.
STREAMFLOW (cfs)
In contrast, forests produce larger amounts of leaf
8
7
6
5
4
3
2
1
area and have higher rates of transpiration. Figure 12.6
1956-1971
compares the hydrologic budget of a typical lodgepole

pine forest in the Medicine Bow Mountains with that
of an old-growth Douglas-fir forest in Oregon (which
receives far more annual precipitation, though less
1940-1955
APR
MAY
JUNE
snow). In the Oregon forest, about 35 percent of the

JULY
AUG
SEPT
Fig. 12.5.Typical hydrograph (solid line) for a Rocky Mountain creek, showing how streamflow increases dramatically in
the spring when the snow is melting. More streamflow commonly occurs following timber harvesting in small patches
(dotted line). The units are cubic feet per second (cfs). Adapted
from Alexander (1987a).
annual precipitation evaporates and transpires during

a typical year, and 65 percent flows to the stream or
groundwater. Evaporation occurs primarily because
of interception. In both regions, interception varies
according to the amount of plant and litter surface area
exposed abovegroundwhether rainfall occurs as a
slow drizzle or quick thunderstormand wind velocity at the time of snowfall.10 In the Rocky Mountains,
Snow
The amount of water in snow cannot be predicted from
snow depth because snow can be comparatively wet or
dry. For example, a snowpack that is 3 feet deep may
have the equivalent of as little as 6 inches or as much
as 16 inches of liquid water.9 Snow drifting is typical,
and thus there can be considerable spatial variability in
depth as well as water content. Because of the importance of snow as a source of water for irrigation and
downstream reservoirs, the Natural Resources Conservation Service and other agencies routinely monitor the
water content of mountain snow. By combining such
data with information about the watershed, the amount
of runoff that will be available downstream during the
summer can be estimated.
Vegetation
Two characteristics of Rocky Mountain vegetation that
sublimation from snow may be more important than

interception.
Leaf area and other stand characteristics vary considerably from one forest to another.11 Not surprisingly,
the amount of leaf area tends to increase with water and
nutrient availability at specific locations.12 In a range of
lodgepole pine forest conditions in the Medicine Bow
Mountains, actual evapotranspiration for the period
from early spring to late fallwhen probably 95 percent
or more of evapotranspiration occursvaried from 10
to 20 inches.13 This represents 3395 percent (a mean of
75 percent) of the total annual precipitation. Outflow to
streams and groundwater accounted for 080 percent of
the snow water, depending on weather conditions during
the year and on forest characteristics, such as the leaf area
index and soil storage capacity. Almost no outflow occurs
during years of relatively low snow-water equivalent in
forest stands with substantial amounts of evergreen leaf
area capable of transpiration in the early spring.
strongly affect runoff, whether that of a meadow or a for-
The importance of leaf area is further illustrated by
est, are the total amount of leaf area and leaf type (ever-
another study done in the Medicine Bow Mountains,
green or deciduous). Leaves are important because water
where the water budgets of two adjacent but differ-
vapor is lost through their stomata (a process known
ent lodgepole pine forests were determined. One was
as transpiration), and also because water adheres to the
a dense doghair stand with about 6,000 trees per acre,
exterior of the leaves and evaporates without becoming
and the other a more open stand with about 800 trees
available to the plants (a process known as interception).
per acre. Both originated after the same fire in the late
The effect of vegetation on water outflow is least when
1800s. The two differed in tree density and biomass, but
the leaf area is low and the leaves are deciduous, such as
both stands had the same leaf area and about the same
in a meadow; probably 8090 percent of the snow water
transpiration rates.14
accumulating in meadows becomes part of streamflow

or groundwater.
Significantly, there are differences between the

hydrology of lodgepole pine forests and spruce-fir for-
Oregon
Douglas-fir Forest
Precipitation
237
Transpiration
Evaporation
Evaporation
5
Snow
2
43
218
37
Canopy
storage
Wyoming
Lodgepole Pine Forest
24
198
4
Canopy
storage
4
Litter
25
18
38
Rooting
zone
13
155
Outflow
Outflow
Fig. 12.6. Annual hydrologic budgets for stands of raindominated Douglas-fir in Oregon and drier, snow-dominated
lodgepole pine in Wyoming. The boxes represent major storage compartments; the arrows are processes affecting water
movement through the ecosystem. Wide arrows indicate
processes that affect relatively more water than processes
with narrow arrows. Numbers are centimeters of water (for
example, 25 cm is the volume of water required to cover any
22
40
Snow
181
Rooting
zone
Evaporation
Evaporation and
Sublimation
14
Litter
Precipitation
66
Transpiration
area 25 cm deep; to convert centimeters to inches, divide by

2.54). Canopy interception is higher in the Oregon forest,
because more rainfall and a higher leaf area index occur there
compared to Wyoming forests. Interception and subsequent
evaporation of summer rains is important in Wyoming, as is
sublimation of snow during winter. The figure for Douglas-fir
is adapted from Sollins et al. (1980); data for lodgepole pine
are from Knight et al. (1985) and Biederman et al. (2012).
ests.15 Forests dominated by spruce and fir in the Rocky
coniferous forests during long periods without distur-
Mountains have considerably more leaf area than do
bance (see chapter 11), with streamflow decreasing as
lodgepole pine forests, and furthermore, the spruce
this shift occurs.16
and fir use water at higher rates per unit of leaf area
Reductions in leaf area stemming from tree cutting,
than either pine or aspen. All things equal, streamflow
fire, and some other disturbances reduce evapotranspi-
would be less from watersheds where spruce-fir forests
ration and, therefore, potentially increase streamflow.17
are common.
Streamflow may continue at an elevated level for up to
Aspen forests are different because the leaves are
60 years, depending on forest type, intensity of distur-
deciduous and absent during most of the snowmelt
bance, and rate of vegetation regrowth, before declin-
period, precluding most early-spring transpiration.
ing to its predisturbance volume after the original leaf
Moreover, the leaves are shed in the fall, eliminating
area has been restored.18 Notably, for reasons described
the possibility of foliar transpiration during subsequent
below, the tree mortality caused by bark beetles does
warm days. Also, there is less potential for intercept-
not necessarily lead to an increase in streamflow.
ing snow on leaf surfaces, from which evaporation

and sublimation can occur. Some land managers have
noticed that the flow of mountain streams can increase
Soil Water-Holding Capacity
after aspen leaf fall. The net effect of forest domina-
Another variable affecting the amount of surface runoff
tion by deciduous trees is that less water is transpired
is the amount of water that the soil can store, that is,
and a larger portion of annual precipitation is available
the amount that is not lost to downward flow caused
for streamflow. Aspen forests may change to evergreen
by gravity. At some point, the soil becomes saturated
during most years, and additional water leaves the eco-
of less direct sunlight. In contrast, the snow on south-
system as surface or subsurface runoff. Thus, soil storage
facing slopes, especially at lower elevations, often melts
capacity at any particular time and place depends on
between snowstorms, with a considerable amount of
the amount of water already in the soil, with deeper and
the soil moisture evaporating on relatively warm days
finer-textured soils holding considerably more water
before additional snowfall occurs. Such periodic melt-
than shallow, coarse-textured soils. The uptake of water
ing makes the snowfall events analogous to summer
by plants leads to soil drying, enabling storage of addi-
showers, with each event producing too little moisture
tional water when it becomes available. The amount of
to percolate through the litter and rooting zone to cause
available storage generally increases during the sum-
substantial runoff. Thus, north-facing slopes contribute
mer, a function of the drying rate, rooting depth, soil
more to streamflow per unit area than do south slopes.
depth and texture, and the time since the last significant wetting eventusually snowmelt. By the end of
summer, the soil moisture of lodgepole pine forests may
be depleted to depths of 6 feet or more, as the pines have
taproots extending that deep.19
Percolation to Groundwater
Surface runoff in the form of streamflow is greatly
affected by the geologic substrate and by whether percolation into aquifers is possible. Granites and other igneous and metamorphic rocks are usually impervious to
Weather and Climate

Runoff from mountain watersheds is also affected by
weather. Gradual warming in the spring causes a prolonged snowmelt, which provides additional time
for early-spring transpiration, thereby reducing the
amount of water available for drainage to streamflow or
groundwater. In contrast, some springs are cold, becoming warm only when day lengths are long. Under such
conditions, the snow melts late and rapidly, and there is
water penetration, which enables more runoff. In contrast, watersheds underlain by sedimentary rock may
contribute significantly to the groundwater of intermountain basins, which is frequently tapped by wells.
Plants growing at the juncture of impervious bedrock
above and permeable sedimentary rock below benefit
from the higher amount of runoff (see fig. 10.3).
Patchiness of Landscape Mosaic
little time for spring transpiration. Moreover, the rate of
Heterogeneity or patchiness in landscapes, caused by
snowmelt occasionally exceeds the maximum infiltra-
disturbances and abrupt changes in various environ-
tion rate, causing overland flow and a higher potential
mental factors, has significant effects on watershed
for flooding, especially if the soil is already saturated
hydrology. For example, a landscape where forests are
or has become compactedas can happen when heavy
frequently interrupted by small meadows contributes
machines are used for timber harvesting. Water flowing
more water to streamflow than does an area of continu-
overland is not available for uptake by plants and can be
ous forest, primarily because wind causes snow drifting
a significant cause of soil erosion.
in such openings, where there is little leaf area and less
Fall weather may be as important as that of the
evapotranspiration.20 Furthermore, with more direct
spring. For example, heavy fall rains can saturate the
sunlight, snowmelt is more rapid in meadows, reduc-
soil just before the onset of winter, when evapotrans-
ing the time available for evapotranspiration. Con-
piration rates are low. Little storage capacity then exists
sequently, more snow water flows down beyond the
during the following spring, leading to the potential for
rooting zone than in areas uniformly covered by forest.
greater spring runoff. Also, if snow falls before the soil
Watershed managers sometimes recommend numer-
is frozen (and the snow is deep enough), the mountain
ous small patch cuts to increase the water yield from a
soils remain unfrozen for the entire winter and infiltra-
watershed. Based on research in northern Colorado, the
tion is enhanced.
largest increases in water yield are reached when 3040
Another factor causing variation in outflow is micro-
percent of a watershed is harvested in patches no larger
climate. For example, north-facing slopes in the North-
than 25 acres. Larger openings can have a different
ern Hemisphere are more likely to retain snow because
effect, because the stronger winds in the opening blow
more snow into the downwind forest, where the snow
potentially altering the chemical composition of water
melts more slowly and higher evapotranspiration may
downstream.
occur.
Inputs or additions of nutrients occur as the result of
In general, abrupt changes between forests of dif-
dryfall (dust), wetfall (rain and snow), rock weathering,
ferent ages, or between forests and other kinds of veg-
animal immigration, surface or subsurface run-on, and
etation (such as meadows), have significant effects on
nitrogen fixation (see fig. 12.2). Losses occur with soil
snow distribution and melting. Weather and vegeta-
erosion, leaching, animal emigration, and, for nitro-
tion are all subject to rapid change (see chapter 11), and
gen, loss to the atmosphere. The conversion of nitrate
these changes affect the amount of streamflow, evapo
back into atmospheric nitrogen is a microbial process,
transpiration, and sublimation.
known as denitrification. Research in lodgepole pine

forests reveals that nitrogen and sometimes potassium
Nutrient Cycling in Forest Landscapes
and phosphorus accumulate in undisturbed forests over

time, even during years of heavy snowpack and large
Unlike water and energy, nutrients cycle in an area for
volumes of water outflow.22 Uptake by plants probably
prolonged periods. Some are lost from the ecosystem,
is especially effective for nutrient retention when the
but they can also be replaced. The nutrient that has
plants are growing rapidly, incorporating additional
received the greatest attention in Rocky Mountain conif-
nutrients into new tissues. 23
erous forests is nitrogen, primarily because this element

often limits rates of plant growth and decomposition,
Dead wood is a common component of almost all

forests and plays a key role in many ecosystem processes,
Nitrogen is the most abundant element in the atmo-
including nutrient cycling and carbon sequestration. As
sphere (78 percent), where it occurs as N2 a form that
fig. 12.7 shows, the nitrogen content of freshly decay-
normally must be converted to inorganic ammonium
ing wood increases considerably over 5 years or more,
or nitrate before it can be taken up by plant roots. Cer-
probably because of nitrogen uptake by the fungi and
tain microbes living in the root nodules of some plant
bacteria that are using the carbohydrates in the wood
species, such as lupine, or microbes that are free-living
as their source of energy. The additional nitrogen most
in the soil, are capable of making this conversion
likely comes from two sources, namely, atmospheric
known as nitrogen fixation. Some fixation also occurs
deposition and transport by fungal filaments connect-
with lightning. Ammonium and nitrate are released
ing the log to the soil. Subsequently, nitrogen content in
during the process of decomposition as well. Notably,
the wood declines as decomposition continues, which
some plants take up nitrogen from the soil in organic

agricultural and industrial technologies have greatly
increased the amount of nitrogen available to eco
systems, which is considered a significant environmental problem on Earth as a whole (see chapter 14).
The potential for nutrient leaching from the soil is
of interest in mountain landscapes, because so much
of the water washes through the soil during the 3- to
6-week snowmelt period. That nutrients are retained
during this spring flush is remarkable, considering that
a large portion of litter decomposition occurs during
the winter just before the spring thaw. Essentially,
nitrogen-limited plants and microbes scavenge the
nutrients almost as soon as they become available,
thereby immobilizing them. Such immobilization
can be disrupted by disturbances, as discussed below,
Fig. 12.7.The percentage of original nitrogen remaining in

decomposing wood in lodgepole pine forests increases before
it begins to decline; the wood slowly releases this limiting
element for the growth of other plants and microorganisms.
Adapted from Fahey (1983).
% ORIGINAL REMAINING
forms, namely, amino acids.21 In one form or another,
180
140
Nitrogen
100
Dry mass
60
20
0
12
24
36
48
60
TIME (months)
72
84
96
frees the nutrient for use by trees, plants, and microbes.
canopies, with important consequences for the abun-
The microbes in decaying wood also immobilize phos-
dance and diversity of the organisms making up the
phorus and calcium. Considering that annual nitrogen
community. Disturbances also alter nutrient cycling
inputs from fixation and precipitation are low in Wyo-
and energy and water flowseffects that are both
ming coniferous forests, as are decomposition rates, the
immediate and long term. The following sections first
nutrient-enriched decaying logs appear to be an impor-
focus on the effects of fire and insects, disturbances
tant source of nitrogen for sustained site productivity.
24
that have affected Wyomings forest ecosystems for
Silvicultural practices that remove excessive amounts
millennia, and then compare such effects to those of
of wood may be detrimental to long-term site produc-
timber harvestinga more recent and different kind
tivity, as discussed later in this chapter. Notably, only
of disturbance.
about 4 percent of a forest ecosystems nitrogen is in

living biomass. Most of it, about 90 percent, is in soil
organic matter (see fig. 12.2), with about 6 percent in
Fire and Insects
aboveground litter (including decomposing wood). Soil
The effect of fire on ecosystem processes depends on
organic matter (including associated bacteria and fungi)
the amount of heat released and the amount of vegeta-
is mineralized very slowly; therefore, only a small pro-
tion burned, characteristics that are highly variable.
portion of the nitrogen in that compartment is made
At one end of the spectrum are low-intensity surface
available each year.
fires, which historically were most common in foothill
Limiting nutrients like nitrogen are retained tena-
woodlands dominated by ponderosa pine and Douglas-
ciously once they are in the coniferous forest biomass,
fir. Such fires burn primarily litter, small trees, and
both above- and belowground. 25 One of the most
understory plants. Most canopy trees survive the fire,
remarkable adaptations of many plants is the ability to
and most forbs, grasses, and shrubs quickly regrow by
withdraw some nutrients from senescing leaves back
sprouting, depending on rooting depth and fire inten-
into stems and twigs prior to leaf fall. This process,
sity. The leaf area of the forest as a whole remains about
known as nutrient resorption, conserves as much as
the same as before the fire, and only a portion of the
half the nitrogen that might have otherwise been lost.
litter is consumed. Consequently, low-intensity surface
In this way, the plants avoid the inevitable competition
fires bring about only small changes in plant growth,
with other organisms that would occur if the nutrients
streamflow, and nutrient cycling, though such fires are
were returned to the soil. More nutrient resorption was
important for maintaining an open stand structure in
found to occur in lodgepole pine than in Engelmann
ponderosa pine and Douglas-fir forests (see chapter 11).
spruce and subalpine fir, which surely contributes to the
At the other end of the spectrum are high-intensity
ability of the pine to tolerate less-fertile soils. In gen-
crown fires, which are typical in lodgepole pine and
eral, the nitrogen concentration of forest floor litter in
spruce-fir forests. These fires kill almost all above
coniferous forests is low (less than 1 percent), a factor
ground plant parts and greatly reduce the amount of
that limits the metabolism of bacteria and fungi and
leaf area (fig. 12.8). The immediate effects are dramatic.
thereby slows decomposition. Depending on site condi-
Plant growth stops until understory plants produce
tions, complete mineralization of the leaves of lodge-
new sprouts, and evapotranspiration is greatly reduced,
pole pine is a slow process, requiring 1222 years. Boles,
because most of the leaf area has been burned. Conse-
branches, and woody roots decompose slowly as well,
quently, a much larger portion of the annual precipita-
with a 12-inch diameter tree bole requiring about 100
tion leaves the watershed as streamflow. 27 Commonly,
years for complete mineralization.
flood peaks are higher than before the fire, depending
26
on such factors as weather conditions during the snow-
Effects of Disturbances on Ecosystem Processes
melt period, as discussed previously. Also, mineral soil is

exposed when the forest floor is burned, thereby increas-
As discussed in chapter 11, both natural and human-
ing the potential for erosion.28 The actual amount of
caused disturbances thin or temporarily remove forest
erosion depends on several factors, including slope, fire
Fig. 12.8. Grasses, sedges, and forbs sprouting from unburned roots and rhizomes 3 years after a high-severity
crown fire in a lodgepole pine forest on the northeast side
of Yellowstone Lake (the East Fire of 2003). Such plants
sequester limiting nutrients as they become available after
the fire, preventing their loss from the ecosystem. Small tree
seedlings were becoming established at the time this photo
was taken but are too small to be seen. One effect of such
fires is to mineralize the forest floor and most of the leaves,

twigs, and small branches. However, most roots, tree boles,
and large branches are not burned; and most of the organic
matter and nutrients in this unburned biomass is eventually
incorporated into the soil. In burned lodgepole forests such
as this one, about 80 percent of the large-diameter wood is
not burned, about 10 percent is converted to persistent charcoal, and the remainder is consumed.
intensity, amount of forest floor burned, water infiltra-
and total plant growth rates close to values measured
tion rate, the rate at which understory plant cover devel-
in comparable mature forests. 29 Most stands required a
ops following the fire, and whether the burn was patchy
longer time.
or uniform. The biggest erosion events usually follow
Using data from young lodgepole pine forests devel-
heavy summer rainstorms within the first year or two
oping after the 1988 fires, along with measurements
after the fire. Snowmelt is less likely to produce serious
from older forests that grew after earlier fires in Yellow
erosion than intense summer rain, even on a severely
stone National Park, Dan Kashian and colleagues pro-
burned site, because snow melts slowly and much of the
duced a model portraying the expected changes in
water percolates through the soil rather than flowing
total plant growth over a 250-year period following
over the surface.
fire or other intense disturbances. 30 Though initial
In burned lodgepole pine forests, leaf area and total
stand density makes a big difference in total productiv-
plant growth return to pre-fire levels within a few
ity, both dense and sparse stands converge on a simi-
decades. The rate of recovery depends mainly on the
lar low level of net productivity near zero by age 250
abundance of new lodgepole pine saplings, which can
years (fig.12.10). This level of net productivity does not
vary immensely, as discussed in chapter 11. In gen-
mean the trees are dead. Photosynthesis continues, but
eral, total plant growth in young stands increases with
accumulation of new plant tissues declines, for reasons
increasing tree density, although this productivity may
not yet understood.31 Lodgepole pine forests typically
be lower in stands having extreme densities (fig. 12.9).
recover more rapidly after an intense crown fire than do
Only 10 years after the 1988 Yellowstone fires, a few
any other forest types in the Rocky Mountains, except
stands of lodgepole pine already supported leaf areas
aspen forestswhich can be restored by sprouting from
7
6
12
5
4
3
2
1
0
10
10
10
10
10
Pine saplings/hectare
in the Rocky Mountains have a variety of impressive
Total LAI m2/m2
Total ANPP t/ha/yr
16
10
Fig. 12.9.Total aboveground plant growth rate, also known as

aboveground net primary productivity (ANPP), and leaf area
index (LAI), defined as square meters of leaf area per square
meter of soil surface, in relation to tree density (number of
saplings per hectare) in 10-year-old lodgepole pine forests
developing after the 1988 fires in Yellowstone National Park.
Both ANPP and LAI tend to increase with increasing sapling
density until extremely high densities of 40,000 saplings per
acre are reached. LAI in such places is almost as high as in
some mature forests. ANPP and LAI appear to level off or even
decrease in stands of even greater density (not shown here),
but the data do not permit this prediction. See fig. 12.1 for
unit conversion. Adapted from Turner et al. (2004).
mechanisms for conserving nutrients. As noted, microbial organisms, notably bacteria and fungi, sequester
them in their biomass. Also, some of the first plants to
grow back after fire absorb nitrogen and other nutrients
in excess of what they actually need for growtha phenomenon referred to as luxury consumption.35 As a result
of microbial immobilization and uptake by recovering
vascular plants, the amount of nutrients lost is less than
might be expected.36
Over a longer time frame, the nitrogen lost to a fire
in a lodgepole pine forest is typically replenished within
4070 years through local nitrogen fixation and inputs
from rain, snow, and dust.37 As the forest regrows, nitrogen and other nutrients are ever more tightly bound in
the living and dead biomass, both above- and belowground. The forest floor, including downed wood,
immobilizes the largest amounts of nitrogen, phosphorus, calcium, and magnesiumat least during the first
decade of post-fire stand development. Living biomass
appears to be the second most important factor in nutrient immobilization, especially after 6080 years. Rates
of nutrient sequestration remain positive even in very
old stands.38
the surviving aspen root system. Rocky Mountain coni-
With regard to insects, the extensive bark beetle outbreaks now sweeping across the West have important
fers lack that ability.

All kinds of fire affect nutrient cycling, with more
effects on ecosystem processes (see chapter 11 for bark
severe burns causing greater effects.32 Some nitrogen is
beetle natural history and effects on forest structure).
volatilized by the heat, but most persists in the large
By selectively killing the larger trees in a stand, beetles
amount of unburned soil organic matter. A portion of
reduce total tree growth for several years after the peak
other nutrients (like phosphorus) remains in the ash,
of an outbreak. However, the smaller trees in the stand
which becomes incorporated into the soil within the
often are not killed, and, with fewer large trees compet-
first year after the fire if not carried away by wind or
ing for water, nutrients, and light, they grow faster than
water. With more water moving through the soil profile
before the outbreak. Understory shrubs and herbs also
as a result of reduced leaf area and transpiration, the
grow faster, resulting in recovery of total stand pro-
probability of nutrient losses from the soil is enhanced
ductivity and carbon uptake and storage within a few
with subsequent nutrient gains to streams.33 After the
decades or even a few years.39 Looking at the effects of
1988 Yellowstone fires, streams flowing from several
bark beetles and other native insects on ecosystem pro-
severely burned watersheds contained elevated levels of
cesses over long time frames, some ecologists have sug-
nitrogen and phosphorus for at least 5 years. Similarly,
gested that plant-feeding insects actually help maintain
nitrate concentrations were elevated for at least 5 years
high levels of total plant growth in forests by killing
in streams draining heavily burned watersheds after the
older, slow-growing trees and freeing resources for young
2002 Hayman fire in Colorado.
individuals. This intriguing idea requires testing, along
34
Although it would seem that nutrient losses asso-
with other possible effects of insects on ecosystems.
ciated with fire, especially losses of nitrogen, might
The death of dominant canopy trees during a beetle
impair post-fire recovery, conifer forest ecosystems
outbreak can result in a substantial reduction in total
Total NEP
g C per m2 per yr
100
High density
50
0
-50
Low density
-100
0
50
100
150
Years since fire
200
250
Fig. 12.10. Long-term trends in plant growth and carbon

dynamics in lodgepole pine forests developing after highseverity fire in Yellowstone National Park. Nearly all growth is
accounted for by trees, but the trends are based on data for all
plants in forests after increasing time since the last fire. Net
ecosystem production (NEP) is the difference between carbon
dioxide removed from the atmosphere by photosynthesis
and that returned to the atmosphere by the respiration of all
living organisms, including plants and decomposers. NEP
is initially negativethat is, respiration exceeds photosynthesisbecause of high rates of respiration by microbes and
other organisms at a time when, after the fire, water and nutrients are more readily available for decomposition and the
biomass of photosynthesizing plants is still low. Initial postfire tree density has a long-lasting influence on NEP, because
denser young forests have a higher rate of plant growth (see
fig. 12.9). Sparse young forests eventually become more dense,
reaching their maximum rates of NEP several decades later
than the forests that initially have a higher density. Both forest types eventually converge on similar low rates of NEP. NEP
units are grams of carbon per square meter per year. Adapted
from Kashian et al. (2006).
leaf area, which could, potentially, lead to changes in
the ecosystem is a short-term increase in useable nitro-
watershed hydrology.40 For example, following the
gen in the soil and increased concentrations of nitrogen
death of more than half the trees in a spruce-fir forest
in the leaves of surviving plants.44
in the Medicine Bow Mountains of southeastern Wyo-
Overall, bark beetle outbreaks lead to some inter-
ming, evapotranspiration declined and soil moisture
esting changes in hydrology and nutrient cycling, but
increased.41 That was predictable. However, a study by
changes in the quantity and quality of streamwater
Joel Biederman and his associates, working in southern
are still not well known.45 Additional research is also
Wyoming and northern Colorado, found no increase
needed to determine whether the overall effect of the
in outflow after beetle-caused mortality. The explana-
outbreaks is to cause forests to become a source or sink
tion appears to be that, although evapotranspiration
of carbonan important consideration in modern for-
was reduced by the beetle attack in the pine forest, the
est management.46 Beetle outbreaks may have cascad-
amount of sublimation from the snow increased under
ing effects on forest ecosystems,47 just as wolves appear
the more open forest canopy. Thus, no additional water
to have had in the Greater Yellowstone Ecosystem (see
was available for streamflow.
chapter 15).
42
Bark beetle outbreaks also affect nutrient cycling.

The magnitude of this effect is small compared with
the impacts of a high-intensity fire, but dead needles
Timber Harvesting Compared with Fires
falling from the canopy bring additional nutrients to
Changes in plant growth rates, hydrology, nutrient cy
the soil surface, and dead trees no longer take up nutri-
cling, and the abundance and distribution of different
ents from the soil, at least during the first few years after
plants and animals can result after timber harvesting,
an outbreak. Not surprisingly for an ecosystem where
just as they do after other disturbances. The biggest
nitrogen is a limiting factor, the surviving trees and
distinction is whether the forest was clearcut or selec-
understory plants sequester many of the nutrients that
tively harvested. Both clearcutting and crown fires kill
become available by decomposition. The net effect on
all or most trees, and it is tempting to conclude that
43
Fig. 12.12. After several generations of trees have grown

and died in a forest, every square foot of soil has had a tree
growing on it or a decomposing log laying over it. Thus, even
though evidence of decomposing logs may not be apparent on the surface (see fig. 11.17), the surface soil contains
many wood particles that contribute to soil development.
Decomposing woody roots contribute to soil development as
well. Reducing the amount of wood added to the soil by too
frequent timber harvesting could cause long-term changes in
soil structure and chemistry. Fires do not normally burn all
the wood (see fig. 12.8).
Fig. 12.11.In contrast to fire, timber harvesting removes much

of the larger wood. Leaves, twigs, and small branches are sometimes left behind as slash, but they decompose more rapidly
than the stem wood. In some places the slash is piled and
burned, reducing the amount of organic matter left on the site.
Clearcutting leaves most slash on the site, that is,

the branches, twigs, and leaves of the harvested trees
along with the bole wood of small trees that are not
removed; fire consumes most of this small-diameter
material.
An intense fire consumes much of the forest floor,
their effects are the same. Indeed, some ecological im
whereas the floor remains intact after clearcutting
pacts are similar. Both dramatically reduce leaf area,
except where the litter is disturbed by roads, various
resulting in reduced transpiration, more water moving
slash treatments, or large equipment.
through the soil, and increased streamflow. Leaching of
The amount of shade cast by dead standing trees after
soil nutrients potentially can increase with both distur-
a fire may be considerable, which can be beneficial
bances.48 Increased light and reduced competition allow
for the establishment of plant seedlings; a clearcut
shrubs and herbs to grow more prolifically after both
provides little shade if most bole wood is removed.
clearcutting and fire, which can be beneficial for herbivores, including elk and deer.
Still, there are important differences that can be enumerated as follows (figs. 12.11 and 12.12):
The differences in ecosystem structure that result

from fire and timber harvesting also lead to differences
in ecosystem processes. The intact forest floor that
remains after clearcutting continues to protect the soil
During clearcutting, most large-bole wood is re
from erosion; in fact, except on steep slopes, most ero-
moved, because wood is the desired commodity.
sion from clearcuts usually comes from the roads built
In contrast, following a crown fire, most dead boles
to access the timber, not from the harvested forest. The
remain. Even an intense fire consumes only a small
bare soil created by an intense fire is more vulnerable to
proportion of the bole wood of living trees.
erosion until plant cover is re-established by sprouting
Fire
Coarse wood created
by disturbance
Pre-disturbance
Coarse wood
Coarse wood added

by developing stand
Fig. 12.13. Predicted long-term trends

in coarse wood after natural fire (top)
or clearcutting (bottom) in lodgepole
pine forests. Time progresses from left
to right in these diagrams. Natural
fires have occurred at centuries-long
intervals in these forests. Thus, only a
single disturbance is shown in the top
panel. The bottom panel depicts a 100year clearcutting rotation, illustrating
how repeated harvest leads to a gradual
depletion of large pieces of wood (more
than 3 inches in diameter) and possibly
to changes in the amount and characteristics of soil organic matter. Adapted
from Tinker and Knight (2004).
Initial Clearcut
Pre-disturbance
Coarse wood
Coarse wood created

by disturbance
Subsequent clearcuts
shrubs and forbs, which usually takes 14 years.49 After
the slash are released slowly during decomposition,
a fire, many nutrients formerly tied up in the living and
reducing the likelihood of nutrient losses to streamflow.
dead biomass exist in the form of ash, which can be car-
Some of the most important differences between
ried away by water and wind; some are lost in smoke as
clearcutting and fire have to do with the dead trees left
well. Warmer daytime soil temperatures resulting from
by firethe same trees that would have been harvested.
removal by fire of the cover of insulating litter also
Dead standing trees provide habitat for many organisms
stimulates soil microbial activity, including the bacteria
long after the fire.50 Also, over the long term, the addi-
and fungi that decompose dead roots and other organic
tion of wood fragments to the soil enhances nutrient
matter, thereby accelerating nutrient release. Thus, the
availability and water-holding capacity and creates an
potential exists for considerable nutrient loss after a fire.
important substrate for microbial organisms. As com-
In contrast, aside from nutrients lost by erosion along
monly noted, the character of forest soils today is par-
roads, the nutrient loss from harvesting is more likely
tially the result of organic matter additions from wood
to be less than that following an intense fire, simply
and leaves over the past 10,000 years or more (fig. 12.13).
because the harvested wood has surprisingly little nutri-
Some wood is lost to periodic forest fires, but between
ent content compared to the slash that is left behind.
fires wood-derived organic matter gradually accumu-
Unless the slash is piled and burned, the nutrients in
lates. Clearcutting interrupts these inputs, especially if
stands are repeatedly harvested at short intervals and if
vides perches for some birds, such as raptors, as well as
the slash is burned.51
a continuous supply of dead standing trees and downed
With increasing recognition of the ecological
wood in the future as these trees die and fall. The dead
importance of large wood on the forest floor and in the
trees, known as snags, also provide important habitat
soil, researchers and managers are exploring ways to
for cavity-nesting birds, such as woodpeckers, small
modify clearcutting operations to increase the amount
owls, and nuthatches.55
of wood that remains after harvest. For example, mod-
Selective cutting leaves many trees in place (fig.
eling studies suggest that simply doubling the usual
12.14) and can mimic the spatial patterns and variable
amount of slash and large wood left on a site after
intensities of insect outbreaks and low-severity fires.56
every harvest could bring long-term wood dynam-
Moderate levels of evapotranspiration are still possible,
ics much closer to the conditions associated with the
and the root systems of the remaining trees continue
natural history of these forests.52 Piling and burning
to take up nutrients and minimize erosion. One study
this material, with the intent of reducing flammabil-
in a lodgepole pine forest estimated that removing
ity or to reduce habitat for certain insects, may not be
about half the total leaf area resulted in nearly dou-
advisable.
bling the amount of water outflow, yet the outflow of
It is disconcerting to some observers that forest
nitrogen remained low. In contrast, clearcutting led
firesa natural phenomenonmight lead to more
to a nearly threefold increase in water outflow and a
erosion and nutrient loss than clearcutting, which is
sixfold increase in total nitrogen outflow. The marked
an artificial, human-caused disturbance. Such adverse

effects must be placed in the context of current conditions. Erosion from mountain forests before road
construction and similar kinds of construction was
probably less common than it is today, a time when
all erosion is viewed as a sign of poor management.
However, prior to this modern predicament, the episodic but infrequent addition of sediments and nutrients to streams could have been beneficial, in terms
of higher stream productivity and possibly higher biological diversity. Various studies have hypothesized
beneficial effects of erosion and nutrient additions to
aquatic ecosystems after fires, even to the point of suggesting that pulses in aquatic productivity might be
tied to periodic fires that burn a substantial portion
of the watershed.53 Such benefits are doubtful today
because of the numerous sources of eroded materials
in most landscapes. Exceptions are in some large roadless areas, including those designated Wilderness by
Congress.
Thus far, clearcutting has been the focus of this discussion, because it is the most intense kind of humancaused disturbance and is most comparable with crown
fire, the most intense disturbance that some consider
natural. However, less-intense selective harvesting is
often feasible. 54 It is now common to leave numerous
trees standing in what otherwise might be considered a
clearcut. Known as green tree retention, this practice pro-
Fig. 12.14. Selective harvesting, or thinning, preserves much

of the forest structure in this ponderosa pine forest in the
Black Hills. Comparable results may not be achieved in other
kinds of forests.
differences in nitrogen outflow between thinned
ics become more frequent because of warmer winters,
and clearcut stands suggest that the surviving trees
and will the number of costly fires increase because of
in thinned stands play an important role in nutrient
longer fire seasons? What are future forests likely to look
immobilization.57
like, and how should the forests be managed to conserve
A confounding factor when selecting a silvicultural
their biological diversity? Plausible scenarios can be pro-
system for lodgepole pine forest is whether the manage
posed, given what has been learned thus far about forest
ment goal is to reduce the amount of dwarf mistletoe
ecology, the effects of recent climate change, and the
(see chapter 11), which, though a native species, is
responses of Rocky Mountain forests to previous epi-
viewed as a problem for timber production. If that is the
sodes of climate change (see chapter 2).
goal, then managers strive to cut all trees infected with
Warming has been occurring for many years and
the mistletoe. That seems simple, but the mistletoe is
is well documented (see chapters 2 and 3). Less well
difficult to see on many trees, which invariably leads to
known is whether precipitation over the mountains
some infected trees being missed and allowing mistle-
will decline. Even if annual precipitation does not
toe to spread to pine seedlings. As noted in the previ-
change substantially, warmer temperatures will cause
ous chapter, dwarf mistletoe was probably controlled in
evapot ranspiration to increase, especially in the sum-
pre-European times by periodic fires. In some places, it
mer, leading to more frequent droughts. Also, warmer
is difficult to use timber harvesting to accomplish the
winters will cause more precipitation to fall as rain
same objectiveexcept by clearcutting. Another con-
rather than snow, and a shallow snowpack combined
cern with partial cutting of lodgepole pine, Engelmann
with warm spring temperatures will lead to earlier
spruce, and subalpine fir forests is that the residual trees
snowmelt, earlier drying of fuels, a longer fire season,
are sometimes vulnerable to windthrow.
and a longer period for trees to be water stressed. Very
Regardless of whether the forest is subjected to par-
likely, climate change will lead to more frequent insect
tial or total tree deathwhether by burning, cutting, or
outbreaks; more frequent large fires; and reduced water
insect epidemicsthe lost nutrients are a small portion
supplies for towns, industry, and agriculture. Forest
of the total nutrient capital. The leaching of nutrients is
regeneration will sometimes fail if extended droughts
elevated for perhaps 10 years, possibly more, but the dis-
become more common58 and if more frequent fires
turbances do not slow nutrient inputs. Most likely the
occur. Most forest types may shift upward in eleva-
lost nutrients are replaced in a few decades at the most.
tion, as the alpine climate becomes favorable for tree
Soil erosion is a more serious concern. Still, with proper
growthresulting in less total forest cover because
precautions it should be possible to ensure the continu-
there is less land area at higher elevations.59 Consider-
ation of soil development and retention. Though hav-
ing the current climate requirements of the prevalent
ing significant impacts at the time they occur, both
forest trees today, U.S. Forest Service scientist Gerald
crown fires and clearcuts are infrequent events. A big-
Rehfeldt and his colleagues concluded that, by the end
ger concern is the proportion of a watershed or forested
of the twenty-first century, forests are likely to cover
landscape affected during a period of time.
less area in the western United States than they do

now.60 Today, forests cover only about 15 percent of
The Future of Mountain Forests
Wyoming (see fig. 1.5)

What can be done about the prospect of reduced
Many forests in the region show the imprint of a century
water supplies? Knowing that transpiration is reduced
of intensive use, having been fragmented by roads, tim-
and water outflow enhanced in forests having less leaf
ber harvesting, homes, and other developments. Others
area, as discussed above, some planners have suggested
have been altered only slightly, such as in roadless areas.
that water supplies could be augmented by increasing
Everywhere, however, big changes loom because of cli-
the harvesting of mountain forests.61 Although possible,
mate change. Will warming affect precipitation patterns
substantial increases could be produced only from forests
in ways that reduce water availability for tree growth
that receive the largest amounts of precipitation, that is,
as well as downstream users? Will bark beetle epidem-
forests at the highest elevations, and such increases in
61
Spring-Summer
Temperature
100
59
Wildfire Frequency
57
0
1970
1975
1980
1985
1990
1995
2000
55
Fig. 12.15.Trends in wildfire frequency and average springsummer temperature (March through August) in the western
United States since 1970. Fire frequency is the number of fires
greater than 1,000 acres in size per year. Temperature and
fire frequency are correlated, and both increased beginning

around 1985. Climate models suggest these trends are likely
to continue during the twenty-first century. Adapted from
Westerling et al. (2006).
water would be seen only in years of average or above-
Have Past Fires Increased the Probability of

Beetle Outbreaks?
average precipitation, which may become infrequent.

Because the regrowth of high-elevation forests is slow
and such forests are highly valued for other reasons
wildlife, recreation, and aestheticsintensive harvest
may not be a socially acceptable strategy for increasing
water supplies. Other approaches, such as improving
water conservation measures and augmenting water
storage facilities, may be more effective.
With regard to fires, an upsurge of large fires during the past 25 years has already occurred across most
of western North America (fig. 12.15). In addition,
extensive bark beetle outbreaks are now a continental
phenomenon.62 Several factors may be responsible for
this recent upswing in forest disturbance. One is the
extensive cover of dense, mature conifer forests that
developed during the twentieth century, when fires and
other natural disturbances were relatively infrequent,
and when human-caused disturbances (such as timber
harvest) affected only a small portion of the forested
land. Dense, mature forests generally are more susceptible to fire and insect outbreaks than are younger
or more open forests. It is tempting to attribute such
problems to inadequate forest management, that is, too
much fire suppression or too little timber harvesting.
However, the increase in fire and bark beetle activity
For many years ecologists thought that fire-injured trees

provided loci for subsequent mountain pine beetle outbreaks into surrounding unburned forests. The results
of one study in the Greater Yellowstone Ecosystem re
vealed that the number of new adult beetles in the next
generation was greater in trees that sustained moderate
fire injury than in severely injured or uninjured trees.64
However, moderately injured trees are usually scarce
after fire in lodgepole pine forests, as most trees touched
by fire are severely injured and die. Few new adult
beetles emerge from the severely injured trees because
of various factors, including damage to the inner bark
tissues on which the beetles depend.
Drought or other nonfire stresses are more likely
initiators of mountain pine beetle outbreaks, simply
because they cause healthy trees to be more susceptible
to beetle attack without damaging the food resource
used by the beetle larvae. Also, in contrast to the mountain pine beetle, spruce beetle outbreaks are known
to begin in windthrown timber or large-diameter logging slashthough drought or other conditions that
stress living trees are necessary to sustain a widespread
outbreak.65
is almost ubiquitous, occurring in forests with a great

variety of management histories throughout most of
the western and northwestern states and up into Alberta
and British Columbia. Climate change is the most likely
explanation;63 current climate models suggest that conditions favorable for beetles and fire will become more
frequent in the future.
Do Beetle Outbreaks Increase the Probability of Fire?

After a beetle epidemic, forests are mostly reddishbrown or gray and they appear alarmingly flammable.
Research suggests that this association is not as simple
as it seems. Jeffrey Hicke and his colleagues from the
University of Idaho developed a model that character-
izes general fuel conditions and potential fire behavior
extremely dry, everything will.68 Although beetle-
in conifer forests during four stages of an outbreakthe
affected forests appear highly flammable for several
green stage early in the attack; the red stage, when the
years after infestation, green forests can burn just as
trees are covered by dead needles; the gray stage, when
intensely under hot, dry weather conditions.
the dead needles have fallen but most of the dead tree
Most forest fires occur during relatively extreme
boles are still standing; and the old stage, when the dead
weather conditions, especially in lodgepole pine and
trees have fallen and a new overstory is developing.66
spruce-fir forests. Consequently, it is not surprising that
Their model incorporates data on surface fuels, which
the influence of beetles on flammability has been small
support a surface fire of varying intensity; crown bulk
or undetectable. One study of the 1988 Yellowstone fires
density or total mass of needles and small twigs on the
found that forests were more likely to have burned in
trees, which can support the burning of tree crowns;
1988 where high beetle-caused tree mortality occurred
and ladder fuels, which are small living trees that can
15 years earlierthough only about 10 percent more
carry fire from the surface fuels into the canopy. Two
likely.69 An examination of all fires in northwestern
types of crown fire are possible: torching (the burning
Colorado and southern Wyoming during the past cen-
of some individual tree crowns) and active crown fire
tury found that forests where bark beetle outbreaks had
(in which the flames spread from tree crown to tree
occurred previously were no more likely to have burned
crown and through most of the forest canopy, killing
with high severity than forests with no previous his-
the majority of trees).
tory of beetle activity.70 In contrast, years with large
Hicke and colleagues found no substantial change
high-severity fires were all drier than years without
in surface fuels or canopy bulk density during the red
fires. Overall, drought and windnot beetlesare most
stage, but the dry, dead needles at this stage increase the
highly correlated with high-severity fires.
probability of torching and active crown fire. Crown
The implication of these recent studies is that large
bulk density is greatly reduced during the gray stage,
fires and extensive insect outbreaks tend to occur
which reduces the likelihood of active crown fires. How-
together, not because one causes the other but because
ever, surface fuels increase during the gray stage, lead-
both respond to similar climatic conditionswarmer
ing to a potentially more intense surface fire, which in
winter and summer temperatures and earlier melting
turn may produce torching of individual tree crowns.
snowpack. These conditions create a longer fire sea-
In the old stage, the fallen boles of the beetle-killed trees
son and greater drought stress for trees in late summer.
contribute to a potentially intense surface fire. More-
Stressed trees are less able to defend themselves against
over, the ladder fuels that have developed at this stage
bark beetles, and the beetles themselves are better able
increase the potential for crown fires.
to survive in the winter, because long cold periods are
Two primary caveats are necessary when consider-
less frequenta convergence of conditions that favor
ing this conceptual model. First, there is disagreement
both fire and beetle disturbance and that is likely to
about many of the models interpretations, partly
become more frequent in the future.71
because bark beetle effects are variable and therefore

workers in different settings will reach somewhat different conclusions.67 Second, although the conceptual
Some Management Options
model focuses on the effects of changes in fuel condi-
In large roadless areas, such as the backcountry of
tions, weather conditions at the time of a fire are as
Yellowstone National Park, managers often allow fires
importantor more importantthan subtle differ-
and insect outbreaks to spread without interference.
ences in fuels. When fuels are dry, temperatures are
Few resources are threatened and such disturbances
high, and winds are strong, any of the four stages
bring about ecological benefits. Native plants and ani-
are easily burned. Indeed, the patterns described in
mals are adapted for such events; the ecosystems are
the model may be relevant only during intermedi-
resilient.72 This is true even if there are more fires in
ate weather conditions, that is, when not too wet and
coming decades. Studies of charcoal and pollen pre-
not too dry. When wet, nothing will burn and when
served in lake sediments reveal that the Yellowstone
country burned more frequently at times in the past,
wind or oriented parallel to the direction of expected
notably about 10,000 years ago, when summers were
fire spread. Significantly, the protection afforded by fuel
warmer than today, yet lodgepole pine apparently was
breaks of this kind are often compromised by glowing
as abundant at that time as it is today.73 The results of
embers, which can be transported by the wind for sev-
this research suggest that forests of lodgepole probably
eral hundred yards and often ignite new fires. This is a
will persist in spite of increased burning and insect
common cause of fire spread on windy days.
activity, though a recent study suggested that the future
Another landscape-level approach is to prohibit
climate in some areas may be unsuitable for this pine,
or discourage home construction in areas of greatest
irrespective of such disturbances.74 Generalizations are
wildfire risk. This might be best accomplished through
difficult; most likely, the dominant factors bringing
disincentives, such as higher property tax rates and
about change will vary from place to place.
insurance premiums, or by county restrictions that
But not all Rocky Mountain forests are in roadless
reflect the relative fire risk at a given location and the
areas. The number of people building homes in moun-
costs to the community as a whole. Access for fire sup-
tain forests has increased greatly over the past two
pression is often presented as a justification for building
decades because of the appeal of living in such an envi-
more roads, but most areas in the foothills and adja-
ronment. Timber harvesting also is an important activ-
cent mountains already have extensive road networks.79
ity in much of the West. Large forest fires are of great
The negative effects of additional roads are discussed in
concern in these settings. Two general approaches can
chapter 18.
be used to reduce the threat of fire. The first is local, that
How should managers deal with bark beetle out-
is, reduce the density of trees, shrubs, and other flam-
breaks? If an outbreak is not already in progress, man
mable vegetation in the immediate vicinity of a house or
agers can reduce the likelihood of one occurring locally
other structure and build the structure itself with non-
by using standard techniques of good forestry, that is,
flammable materials.
Many homes that implemented
by thinning and spacing trees for optimal growth and
the Firewise recommendations of the U.S. Forest Ser-
the maintenance of healthy trees that are capable of de
vice have escaped serious damage in recent fires.76 Fuel
fending themselves against bark beetles. 80 Making forest
reduction also can be accomplished in forests further
stands more resistant to bark beetles in this way is prob-
removed from vulnerable structures. The 2000 National
ably most appropriate on public and private lands where
Fire Plan directs federal agencies to conduct thinning
timber production is an important management goal
and prescribed fire treatments in this wildland-urban
or in campgrounds and communities where the death
interface; however, this mandate has proved challeng-
of many large trees would impair local aesthetics and
ing to accomplish, in part because most of the interface
create the hazard of falling trees. Such intensive, active
is on private, not public, land.77
management would be highly controversial in roadless
75
The second approach for reducing fire risks is applied
areas and in the backcountry portions of national parks.
on a broader scale. The objective is to break up continu-
Once a beetle outbreak is well under way, stopping
ous fuels and create places where fire spread and inten-
it does not appear to be an option at the present time.
sity can be reduced and where suppression crews can
Regardless of management, the outbreak will continue
attack fires effectively when that seems necessary. Forest
until bitter cold temperatures at the right time kill
fire scientists have identified optimal sizes and position-
the beetle larvae wintering under the tree bark, or the
ing of fuel reduction sites to achieve the greatest pro-
beetles simply run out of suitable host trees.81 Individual
tection of vulnerable resources for a given investment
high-value treessuch as those in campgrounds, town
in fuel reduction.78 For example, large fires are often
parks, or yardscan be protected from beetles even
driven by high winds coming from more-or-less pre-
during an outbreak by repeatedly applying appropriate
dictable directions. Therefore, rectangular fuel breaks
insecticides, but, over large areas, that solution has been
of sufficient size placed upwind of a community and
prohibitively expensive and problematic to date.
oriented perpendicular to the prevailing winds could be
Alternatives are available for handling the trees killed
more effective than similar fuel breaks located down-
by the beetles, such as removing them from around
homes, campgrounds, and along roads and trails to
growing; other clones appear to be dead or have very
prevent damage or injury when the dead trees fall. Un
few sprouts.84
fortunately, cutting and burning beetle-killed trees after
At the same time that aspen appeared to be declin-
the needles have turned red will do nothing to slow a
ing at low elevations, new aspen seedlings that became
mountain pine beetle outbreak, because the new cohort
established in burned forests after the 1988 Yellow-
of beetles has already emerged from the tree by the time
stone fires were growing best at higher elevationsin
the needles dry and begin to fall. Removing beetle-killed
some places higher than the pre-1988 range of aspen
trees is also questionable as a fire mitigation method,
in Yellowstone.85 The same pattern has been seen after
because, as discussed, beetle-caused mortality does not
recent fires in the Canadian Rockies. In Colorados San
appear to increase fire hazard.82 After 2050 years, when
Juan Mountains, a comparison of historic photos from
the dead trees have fallen and the surviving small trees
the early 1900s with recent photos of the same land-
are again forming a forest canopy, the heavy fuel load
scape revealed new aspen clones at the upper treeline,
will have the potential to support an intense fire. How-
apparently the result of seedling establishment even
ever, it is not known whether future fires in forests with
without any major disturbance by fire.86
many beetle-killed trees will actually be more intense
The pattern of aspen mortality at lower elevations
than fires in a green forest where that has not happened.
and new recruitment at higher ones suggests that aspen
Hence, at this time no clear guidelines exist for whether
is already responding to the climate changes of the past
trees killed by mountain pine beetle should be removed
quarter-century. Such trends are likely to continue,
from locations that are distant from homes, roadsides,
with some low-elevation clones dying as climate con-
and other obviously vulnerable resources. Doing so may
ditions become less favorable there. Forest ecologists
be more expensive than can be justified, considering that
and climate modelers have worked together to project
the next fire or beetle outbreak will probably be influ-
the future distribution of suitable habitat for aspen and
enced more by weather patterns than by fuel conditions.
other plants in western North America. Their approach
Again, however, the spruce beetle presents a different
is to map the current range of aspen and to then
problem, and the removal of dead trees may be advisable
describe the extent of variation in climatic conditions
when they are a threat.
that exist throughout the current range, thereby creating a bioclimatic envelope. They then use climate models
Aspen Forests: Bellwether of Future Changes?
to identify specific locations where suitable conditions

for aspen will be located in the future, assuming vari-
Quaking aspen is one of the most highly prized trees in
ous scenarios for climate change. For aspen, two major
western North America, primarily for the biological and
expectations emerge from these initial efforts: (1) the
aesthetic diversity it adds to the landscape. Thus, it was
total extent of suitable habitat will likely decrease dur-
especially alarming when large aspen trees began dying
ing the twenty-first century, although aspen will persist
in 2003 and continued to do so through 2007, mostly
in small pockets throughout most of its current range;
in Colorado and Utah but also in Wyoming and other
and (2) new groves will most likely become established
western states (fig. 12.16).83 This widespread phenom-
at higher elevations (up to 1,000 feet higher than where
enon has been called sudden aspen decline (or SAD).
aspen grows now).
For a time the cause of decline was unknown, but now
Similar studies have come to about the same conclu-
it appears that the severe drought from 2000 to 2002
sion for most other Rocky Mountain trees.87 Notably,
coupled with unusually high temperatures was the trig-
because there is less land area at higher elevations, and
gering factor. The greatest aspen decline has taken place
much of it lacks soil (such as on cliffs and talus slopes),
on south-facing slopes at lower elevations and on soil
the area occupied by each species will be reduced. In
types with low water-holding capacity, two observa-
Wyoming, the amount of forested land could decline
tions that are consistent with the drought hypothesis.
from about 15 percent at the present time to 78 per-
The root systems and some small trees of many clones
cent. There could be exceptions to this trend for spe-
survived and are now producing new sprouts that are
cific species. For example, Douglas-fir may be able to

Fig. 12.16. Sudden aspen decline affected aspen
stands in various places throughout the Rocky
Mountain region during 20012005. In some
stands, most large stems diedthough the root
systems did not. New sprouts are expected to
produce a new forest if they are not subjected
to another severe drought or to chronic excess
browsing by ungulates (see chapter 15). This
photo was taken on the Uncompahgre Plateau in
western Colorado. Elevation 9,500 feet. Photo by
Dan Binkley.
move up and across the expansive Yellowstone Plateau
mixes of species. Forests are likely to be more frag-
from the surrounding foothills where it now grows, as
mented in the future (a topic that is considered further
the cool climate on the Plateau becomes warmer. That
in chapter 18), and extensive tracts of interior forest
would happen, however, only if Douglas-fir can tolerate
may be less common than they are now. Meadows and
the relatively infertile rhyolitic soils found there.
parklands may become more common, just as they did

during the warm and dry intervals of the past 10,000
In sum, a cadre of scientists and managers is develop-
years. It seems prudent for managers to conserve native
ing procedures for predicting, detecting, mitigating,
species where possible, take actions to minimize the
and adapting to the ecological changes that can be
number of invasive species, and consider how the eco-
expected.88 The forests that future generations enjoy
system services provided by forests in the past can be
may cover less total land area and be composed of novel
extended into the future.
Mountain Meadows
and Snowglades
Chapter 13
Sometimes referred to as parks, meadows add diversity to
Higher on the mountain, Idaho fescue and tufted hair-
the mountain landscape (fig. 13.1). The plants are highly
grass are more common (see chapter 10). Wet meadows
variable due to differences in elevation, wind exposure,
are found at all elevations along streams and in depres-
water availability, and soil characteristics, but often they
sions, where melting snow provides a source of water
include such colorful species as arrowleaf balsamroot,
sufficient to support shrubby cinquefoil and mountain
Indian paintbrush, and lupine (table 13.1). Such plants
silver sagebrush in addition to various sedges and grasses
attract numerous butterflies, along with other animals
(see chapter 5).2 Meadows with intermediate moisture
that benefit from herbaceous plants. At lower elevations,
availability support the greatest diversity of plants.
the plants are similar to those in dry foothill grasslands,
A frequently asked question about mountain mead-
with bluebunch wheatgrass a characteristic species.
ows is why they have no trees. The answer depends on

Fig. 13.1. Mountain meadow
in the Wyoming Range west
of Big Piney. Mountain big
sagebrush and Idaho fescue
are common, with willowdominated shrublands in
the riparian zone. Arrowleaf
balsamroot and lupine are
abundant in the foreground.
The forests in this area are
dominated by Engelmann
spruce, lodgepole pine, and
subalpine fir, with whitebark
pine near the alpine treeline.
221
Table 13.1. Some characteristic plants of subalpine meadows in Wyominga
Common name
Latin name
Sagebrush
meadows
Dry subalpine
meadows
Mesic subalpine
meadows
GRASSES
Alpine bluegrass
Poa alpina
Cusicks bluegrass
Poa cusickii
Muttongrass
Poa fendleriana
Nodding bluegrass
Poa reflexa
Pattersons bluegrass
Poa abbreviata ssp. pattersonii
Sandberg bluegrass
Poa secunda
Timberline bluegrass
Poa glauca ssp. rupicola
Alpine timothy
Phleum alpinum
Idaho fescue
Festuca idahoensis
Rocky Mountain fescue
Festuca saximontana
Oatgrass
Danthonia spp.
Prairie junegrass
Koeleria macrantha
Slender wheatgrass
Elymus trachycaulus
Thickspike wheatgrass
Elymus lanceolatus
Spike trisetum
Trisetum spicatum
Tufted hairgrass
SEDGES
Curly (rock) sedge
Carex rupestris
Dunhead sedge
Carex phaeocephala
Ebony sedge
Carex ebenea
Hoods sedge
Carex hoodii
Needleleaf sedge
Carex duriuscula
Northwest Territory (beaked) sedge
Carex utriculata
Obtuse sedge
Carex obtusata
Sheep sedge
Carex illota
Polygonum bistortoides
Alpine sagebrush
Artemisia scopulorum
Prairie (fringed) sagewort
Artemisia frigida
Arrowleaf balsamroot
Balsamorrhiza sagittata
Ballhead sandwort
Arenaria congesta
FORBS
Alpine bistort
Creeping sibbaldia
Sibbaldia procumbens
Field (mouse-ear) chickweed
Cerastium arvense
Fireweed
Chamerion angustifolium
Indian paintbrush
Castilleja spp.
Mules ears
Wyethia amplexicaulis
Pale agoseris
Agoseris glauca
Mountain Meadows and Snowglades
Sagebrush
meadows
Dry subalpine
meadows
Mesic subalpine
meadows
Common name
Latin name
Purple locoweed
Oxytropis lambertii
Ross avens
Geum rossii
Silvery lupine
Lupinus argenteus
Sulphur-flower buckwheat
Eriogonum umbellatum
Tall fringed bluebells
Mertensia ciliata
Tufted fleabane
Erigeron caespitosus
Western yarrow
Achillea millefolium var.

occidentalis
White locoweed
Oxytropis sericea
Caltha leptosepala
Artemisia tridentata ssp. vaseyana
SHRUBS
Silver sagebrush (mountain)
Artemisia cana ssp. viscidula
Shrubby cinquefoil
Dasiphora fruticosa ssp. floribunda
local conditions. Some meadows occur on fine-textured
common, surface-soil texture is coarser. The presence
soils along rivers and streams, where the absence of trees
of a distinct stone line under the fine-textured surface
has been attributed to the soils remaining too wet for
soil in the meadow and its absence in the forest led
too long in the summer.3 Elsewhere, competition from
Doering and Reider to conclude that the meadow origi-
grasses and forbs may be too great. Other meadows are
nated after a severe fire or windstorm, perhaps several
found where trees are excluded because the soils are not
thousand years ago. Subsequent wind erosion removed
favorable for tree seedling establishment (fig. 13.2), or
much of the exposed fine material but left a layer of
where fires have occurred at high elevations and the re-
stones (fig.13.4). Gradually, new deposits of fine, wind-
establishment of new trees is slow. Transitions between
blown material covered the park. The dryness of the
meadows and forests in upland settings can be difficult
park probably is caused by two factors: (1) the diffi-
to explain, as soil differences often are not obvious.4
culty of water percolation through the shallow stone
Perhaps the most-studied meadow in the Rocky
line, with most water held in the shallow, fine-textured
Mountain region is Cinnabar Park in the Medicine Bow
surface soil, where it is readily evaporated; and (2) the
Mountains (fig. 13.3). Early reports suggested that this
strong winter winds that greatly reduce snow accumu-
dry meadow is slowly moving downwind, with trees
lation. Blowing ice and snow also damage the leaves
invading on the upwind side (where more snow accu-
and stems of young tree seedlings, leading to further
mulates) and older trees dying on the downwind edge.
mortality.6 Plant water stress in the meadows is sug-
However, soil scientists William Doering and Richard
gested by the obvious wilting of many plants during
Reider concluded in 1992 that the park was stable.
clear summer days.
They suggested, after detailed analysis of soil profiles,
Subalpine meadows also persist where summer frosts
that trees were absent in the meadow because of a shal-
are frequent. This seems counterintuitive, because trees
low, 6-inch accumulation of fine-textured surface soil
are found nearby. However, because of direct exposure
that provided a better environment for grasses and
of the meadows to cold skies at night, frosts at the soil
forbs. In the adjacent forest, where tree seedlings are
surface can occur on a third or more of summer nights.
223
Fig. 13.2. Large mountain meadow in the Bighorn Mountains, located where shales in the Gros Ventre Formation are
exposed. Shales lead to the development of fine-textured
soils that are not favorable for tree seedling establishment.
Common plants in the meadow include fringed sagewort,
hoary balsamroot, Idaho fescue, junegrass, prairie smoke,
silvery lupine, starry cerastium, wild geranium, and yarrow.
Aspen groves occur in the ravines. Forests here are dominated

by lodgepole pine and limber pine and are found on the
adjacent, coarse-textured soils derived from sandstones in
the Flathead Formation. Similar meadows exist where finetextured sedimentary rocks are exposed on the surface, such
as on the east slopes of the Wind River Mountains. Elevation
7,400 feet.
Fig. 13.3.Cinnabar Park, a

dry meadow in the Medicine
Bow Mountains, is located
on a high plateau. Strong
winds blow the snow off the
meadow, where it forms drifts
on the leeward side of the ribbon of lodgepole pine on the
left. Often the snow persists
until August, preventing
small trees from becoming
established. Canby bluegrass,
Idaho fescue, junegrass, sheep
fescue, and starry cerastium
are common in the dry
meadow; tufted hairgrass and
other wet meadow species
are found in the snowglade.
Fig. 13.4. Soil characteristics across Cinnabar Park. The stone

line probably formed during a period when fine soil particles
were eroded from the high plateau. New fine material gradually
accumulated to a depth of approximately 6 inches (15 cm), but

the establishment of tree seedlings in the dry meadow is still
difficult. See fig. 13.3. Adapted from Doering and Reider (1992).
Tree seedlings are easily killed by a combination of frost
topography, and the chance establishment of a small
damage and reduced rates of photosynthesis the next
tree or shrubwhich initiates snow deposition on its
day. In contrast, summer freezing is rare under the for-
leeward sideone or more bands or ribbons of trees are
est canopy, if it occurs at all. Two other factors limiting
sometimes found in the vicinity of snowglades.
tree seedling establishment could be excess competition
The ribbon forestsnowglade pattern takes at least
for tree seedlings with established herbaceous plants
two forms in the region.10 Commonly, there is just one
and the kind of vertical discontinuity in soil texture
band of trees and one band of meadow on the lee side
observed at Cinnabar Park.7
of a more extensive meadow, such as at Cinnabar Park

where the late-spring snowdrift may be 30 feet deep or
Snowglades and Ribbon Forests
more (see fig. 13.3). The initial origin of such meadows

is not easily explained. Dwight Billings, Duke Univer-
Meadows created by late-lying snow are known as snow-
sity ecologist, suggested in 1969 that forest fire near
glades. The soil is covered, cold, and wet for too long and
timberline . . . changes the snowdrift pattern enough
the growing season is usually too short for tree seed-
that trees in an unburned area to the lee of the burn
lings to become established.8 Also, deep snow that per-
are killed by late-lying snow during the summer. The
sists until midsummer favors the growth of molds that
dead trees are replaced by a wet type of snowglade
can kill buried branches and young trees, for example,
meadow.11 Support for this hypothesis comes from the
the blackfelt snowmold (Herpotrichia nigra) (fig. 13.5).
observation that huge snowdrifts form on the lee side of
In addition, pocket gophers that feed on tree roots and
some clearcuts, usually 1020 feet into the uncut forest,
bark are sometimes common under the snow.
and the effect of the drift is to reduce tree vigor. Dead
Tree seedlings, however, also have difficulty becom-
standing trees can be found even though the upwind
ing established where snow accumulation is low, in part
clearcut is less than 20 years old. Over a period of 500
because of insufficient moisture. As important may be
years or more, all remnants of the original forest could
insufficient protection from abrasion by blowing ice
disappear.
and snow. Working in the Medicine Bow Mountains,
In other situations, usually near upper treeline, sev-
Stephan Httenschwiler and William Smith found that
eral bands of forest and meadow may occur in sequence
tree seedlings were favored in places where snow depth
(fig. 13.7).12 Again, the role of wind and snow drifting
was at least 18 inches but no more than 56 inches (fig.
is obvious, because the bands are perpendicular to the
13.6).9 Due to the interaction of drifting snow, micro-
prevailing westerly winds and the deepest snow occurs
225
in the glades.13 The establishment of the first trees that

lead to the ribbon forest pattern is still a puzzle to be
solved (but see chapter 14).
A similar phenomenon has been observed where
aspen occur in doughnut-shaped or U-shaped groves
that are sometimes referred to as aspen atolls (see
fig.10.13). Commonly surrounded by sagebrush or subalpine meadows, the groves probably developed because
of the snow-fence effect of a few trees that by chance
became established. The original trees created the
only sites where new seedlings or sprouts could obtain
enough water to survive. Eventually, the clone expands
around all or most of the snowbank, such as south of
Rawlins on the north end of the Sierra Madre. Engelmann spruce and subalpine fir develop similar groves
in northwestern Wyoming, but at higher elevations.14
Trees cannot survive in the center, because the snow
persists too long, but the snowdrift provides a dependable supply of moisture for the trees on the perimeter.
More often seen are ridgetop ribbon forests. In this case,
deep snow accumulates on the lee side of ridges, maintaining a linear snowglade just above trees that benefit
from the meltwater and, consequently, are able to grow
where they might not otherwise occur.
25
The forest atoll or ribbon forestsnowglade pattern

may characterize only a small portion of the mountain
landscape, but the pattern is of considerable significance
economically, as they are a naturally developed high-
` ribbon _
20
15
W- and SW-winds
10
5
SW
NE
snow
20
15
30
20
10
Distance (feet)
none
none
none
10
none
Seedlings / ft2
Height above ground (feet)
Fig. 13.5. Blackfelt snowmolds can kill branches and small

trees that remain covered with snow late into summer, such
as in some snowglades. This photo shows an infected lodgepole pine sapling. The lower needles are compressed by the
black or brown fungal filaments (hyphae).
10
20
Distance (feet)
30
Fig. 13.6.Common pattern

in a subalpine meadow near
treeline, near 11,000 feet,
with ribbons of mature
Engelmann spruce and
subalpine fir (see fig. 13.7).
Note how the trees affect
snow drifting and how
moderate snow depths appear to favor tree seedling
establishment. The seedlings observed in this area
were 110 years old and
may not grow to maturity.
Snow accumulation depths
and spatial distribution
vary according to tree
height and the width of the
ribbon of trees. Adapted
from Httenschwiler and
Smith (1999).
higher in the mountain meadows because of the variety

of plant communities that occur up and down mountain slopes.15 Essentially, there is a wave of new plant
growth that the animals can follow, from the lowlands
in the spring to the high mountain meadows later in
the summer. Because of topographic diversity and different amounts of snow accumulation, a supply of green
forage almost always exists nearby.
Livestock managers have learned that the initiation
and duration of grazing is important to consider when
developing management plans. Ranchers prefer to rest
their low-elevation rangelands in the spring, when
plants there are actively growing and are most susceptible to damage from grazing. Consequently, they want
to move livestock onto the mountain meadows as soon
as possible. The trick is to move the animals at a time
when the soils are not so wet that damage from trampling occurs. Moreover, just as in the lowlands, the
mountain plants are subject to stress if grazed excessively in the early stages of their growth.16
Today, the grazing of mountain meadows is restricted
in many areas to a shorter summer period than in the
early 1900s. Fewer animals are permitted on federal
Fig. 13.7. At high elevations with strong winter winds, such
as in this area north of Medicine Bow Peak, parallel ribbons
of trees and snowglades sometimes develop. The trees are
predominantly Engelmann spruce and subalpine fir. See fig.
10.13 for a photo of an aspen atoll, caused by the effects of
blowing snow at lower elevations. Photo by William K. Smith.
land, and often a shepherd must be employed to keep

the animalsespecially sheepfrom grazing specific
areas excessively. As a result, some ranchers now find
it uneconomical to move their livestock to the mountains for such a short time, often midsummer to early
fall. Also, the amount of land area in meadows has
declined over the years in some areas, owing to fire sup-
water-yield ecosystem, in which most of the snow accu-
pression and the subsequent expansion of trees. This
mulates where there is comparatively little use of that
trend might be especially apparent in mountain ranges
water by plants. Not surprisingly, a common practice of
that were subject to considerable burning or tree cut-
watershed hydrologists desiring to increase streamflow
ting in the late 1800s and early 1900s. The best example
is to create small openings in the forest, where snow
in Wyoming may be the Sierra Madre, where extensive
can accumulate in much the same way (see chapter 12).
timber harvesting in conjunction with copper mining

created large areas of open rangeland. Now much of this
Livestock Grazing on Mountain Meadows
area is again forested.

Overall, mountain meadows are widely appreciated
Mountain meadows have long been an important com-
for the benefits they provide in terms of livestock for-
ponent of the summer range for elk and deer, and for a
age and water yield. As important, the edge between the
century or more many ranchers have come to depend
forest and meadow provides habitat for many birds and
on such meadows for supplemental livestock forage.
mammals. Trees have invaded some meadows, but only
Using the meadows in this way during the summer pro-
rarely are forests converted to meadows after fire or tim-
vides a period of rest for rangelands in the valleys below.
ber harvesting. Shifts in the location of forest-meadow
The carrying capacity for both wildlife and livestock is
boundaries may be an indication of climate change.
227
Forest Expansion into Meadows
Response to Climate Change
Although some meadows appear to be stable features
Predicting the effects of warming (or cooling) on plant
of the landscape, others are being invaded by trees.
growth in meadows is currently hampered by un
As noted, the origin of such meadows is often difficult
certainties about how water availability will change
to explain, but the presence of young trees suggests a
under future climates. If the annual precipitation
change of one or more environmental factors over time.
increases or remains about the same, then concomitant
The factors mentioned most often are less frequent fires,
higher temperatures will create drier conditions, because
which enables trees to grow where they could not sur-
the water will evaporate more rapidlyoften before it
vive when fires were more frequent; grazing by domestic
can enhance plant growth. The annual growth of many
livestock, which can favor trees by reducing competi-
plants would be slower if summer drought stress were
tion from herbaceous plants; and climate variation.
to begin earlier. If summer precipitation increases to a
Meadows that are otherwise apparently stable could
level that more than compensates for increased evapo-
also be undergoing very slow reforestation following a
transpiration, then a warmer climate could result in a
severe disturbance that occurred long ago. An example
longer growing season, depending on whether the addi-
of slow invasion is at high elevations on parts of Libby
tional water comes as snow or summer rain. If summer
Flats in the Medicine Bow Mountains (see chapter 14).17
rainfall were to increase sufficiently, then the growing
In considering the effects of climate variation, sev-
season for many mountain plants could be lengthened,
eral scenarios are plausible. First, consider what might
and the annual production of new plant biomass would
happen if the climate were to become cooler. Evidence
increase. Alternatively, more snow may not have much
suggests that tree establishment after such disturbances
effect on plant growth, because even low snowfall years
as fire would be slowed. For example, pollen data from
most likely would produce enough meltwater to saturate
the Wind River Mountains indicate that forests gave
the soils. The additional water could augment stream-
way to meadows about 3,000 years ago during a period
flow and reservoir levels but often would not increase
of cooling.18 Nocturnal summer frosts at this time
plant growth in the meadows.
probably became frequent enough to restrict tree seed-
Of course, not all meadows and plant growth forms
ling establishment as the older conifers died, thereby
would respond in the same way. Ecologist Diane
enabling herbaceous plants to dominate instead after
Debinski and her colleagues studied dry, moist, and
fires or windstorms. The long-term persistence of pine,
wet meadows in the Yellowstone area during a transi-
spruce, and fir depends on seedling establishment, as
tion from very wet conditions (19971999) to very dry
these trees lack the ability to sprout from stumps or
ones (20002007).21 The amount of plant cover gener-
roots.
ally declined, especially in dry meadows, but changes
In contrast, if warming occurs, then summer frosts
in forb and shrub cover varied considerably. Shrub
at night become less frequent, and tree seedlings may
cover actually increased in several of their dry, moist,
be able to survive more frequently (see chapter 14). A
and wet study areas. In addition to the direct effects
contributing factor would be that warmer air holds
of drought, various studies have documented variable
more water vapor, which itself is a greenhouse gas that
plant responses to changes in the duration of snow
enables warmer temperatures at night. Expansion of
cover.22
lodgepole pine into dry meadows since about 1870 in
To learn more about the mechanisms whereby sub
Yellowstone National Park apparently is driven by the
alpine meadows might respond to climate warming,
regional warming trend that began at the end of the
small-scale experiments have been conducted using elec-
Little Ice Age.
Similarly, trees became established in
tric heaters. One study in the Colorado Rockies found
some meadows in the Medicine Bow Mountains during
warming increased the water-use efficiency of mountain
this time, apparently for the same reason.20 Such obser-
big sagebrush, enabling higher rates of photosynthesis
vations suggest that tree expansion is likely to continue
early and late in the growing season.23 The results suggest
as the global climate warms over the coming century.
that mountain big sagebrush could become more abun-
19
dant with warming, partly because it has roots that tap
and biological diversity, are important sources of forage
deep soil water late in the growing season.24 In contrast,
and water, and are sensitive indicators of environmental
the rate of photosynthesis in shrubby cinquefoil did not
change.26 With ongoing climate warming, the expan-
increase with warming. Another heating experiment
sion of trees into some meadows is likely to continue
conducted nearby found no effect of warming on plant
where (1) desiccation does not preclude tree survival,
species composition after four growing seasons, possibly
(2) night-time temperatures enable tree seedling estab-
because the earlier drying after snowmelt canceled the
lishment, (3) competition from herbaceous plants is not
benefits of warming.25 Also, nutrients might have been
excessive, and (4) scouring by winter winds is not an
limiting to plant growth, rather than temperature or
important factor. The temptation to maintain some of
water, as observed in alpine tundra (see chapter 14).
these meadows by tree cutting or burning may increase.
If nutrients are a limiting factor for plant growth,
Snow may melt sooner because of warming trends, but
then the effects of warming may be indirect. For exam-
even low snowfall years are likely to produce enough
ple, warming may stimulate growth only after sufficient
water to saturate meadow soils at the beginning of the
time has passed for warming to promote the decomposi-
growing season. Thus, mountain meadows and tundra
tion of soil organic matter, thereby making more nutri-
will continue to be green for much of the summer, even
ents available at a time when water also is available and
during drought yearsin contrast to lowland grass-
temperature conditions are favorable. Invariably, mul-
lands. Consequently, the demand for livestock access
tiple factors are involved. The effects of environmental
to this forage may increase. Ribbon forests and snow-
change are not always immediately obvious; long-term
glades are likely to shift in their locations with changes
experiments are required.
in the timing of snowmelt, but they will continue to

be a feature of the subalpine landscape as long as suf-
In summary, mountain meadows are a highly valued
ficient snowfall and wind-caused drifting occur during
component of mountain landscapes. They add aesthetic
the winter.
229
Upper Treeline
and Alpine Tundra
Chapter 14
Found at the highest elevations, with views of the val-
many of its inhabitants will be unable to survive in the
leys below, alpine tundra is the land of the pika, rosy-
region? To make predictions, conservation biologists are
finch, wind-swept trees, and small plants with big
thinking broadly about the tundra environment and
flowers. During the last Ice Age, the treeline was lower
the ecology of the plants and animals that live there.
and tundra extended over a larger area (see chapter 2).
Subalpine forests give way to alpine tundra at eleva-
Alpine animals could move easily from one mountain-
tions ranging from about 11,480 feet in the Medicine
top to another. Since then, as the climate has warmed,
Bow Mountains, in southern Wyoming, to about 9,840
the tundra has been invaded by trees. Will the tun-
feet in the Beartooth and Bighorn mountains on the
dra habitat be further diminished to the point where
border with Montana (figs. 14.1 and 14.2).1 Slope expoFig. 14.1. Alpine tundra in the
Snowy Range of the Medicine
Bow Mountains. The bedrock
of this glaciated valley is
Medicine Bow quartzite.
230
Upper Treeline and Alpine Tundra

Fig. 14.2. Upper treeline in the Bighorn
Mountains. The trees are Engelmann
spruce and subalpine fir. Common
plants in the alpine turf are alpine
avens, alpine bistort, alpine forget-menot, alpine sagewort, blackroot sedge,
selaginella, stonecrop, and spike tri
setum. Elevation 9,600 feet.
sure also is important in determining the limits of tree
itous circumstances. William K. Smith and his associ-
growth, with treelines often at higher elevations on
ates, working in the Medicine Bow Mountains, found
the warmer south slopes with less snow cover. North
that seedling establishment was facilitated when the
and west slopes tend to have more snow later into the
area around the seedling favored a moderate amount of
spring, which causes the treeline to be lower.2
snow cover, providing protection from abrasion and desiccation during winter. As important, they found that
Krummholz
seedlings required at least a minimum of plant cover to

provide shade during the summer, not just from intense
The trees commonly found at treeline are Engelmann
alpine sunlight during the day but also from the cold
spruce, subalpine fir, limber pine, and, in the Greater
sky at night.4 Because of the cold-sky effect at night,
Yellowstone Area, whitebark pine.3 All are short at this
unshaded leaves could be colder than the minimum air
elevation. On exposed sites the trees have a windswept,
temperatures on near-freezing summer nights, which
shrubby growth form known as krummholz, a German
was found to inhibit photosynthesis the following day.
word meaning twisted wood. Some have vertical trunks
Summertime freezing is detrimental to plants and is
flagged with branches on the leeward side, caused by the
much more likely to occur when they are exposed to
abrasion of blowing ice and snow that prevent branch
the night sky. Moreover, unshaded leaves were consid-
growth upwind (fig. 14.3). Branches and the needle-
erably warmer than the maximum air temperature on
shaped leaves near the ground are denser, forming a mat
a summer day, which greatly increased rates of water
that accumulates a protective snowdrift in winter. The
loss and the likelihood of wilting. With adequate but
shrubby trees are widely spaced in a matrix of alpine
not excessive shading, seedlings have photosynthetic
tundra and subalpine meadows. In some wind-exposed
rates that are sufficient for plant growth and the for-
places they occur in ribbons because of the interactive
mation of mycorrhizaewhich in turn lessens water
effect of wind, snow deposition, and slight changes in
stress. The seedlings future is secured when the young
topography (see chapter 13).
tree is large enough to cause snow accumulation over
The stunted trees can be very old. Most likely they
its own branches for a sufficiently long time during the
became established when a seedling managed to survive
winter, or if the neighboring plants or microtopography
for several years because of a rare combination of fortu-
cause the needed snow drifting. This research suggests
231
Fig. 14.3.Trees at upper treeline often have the krummholz

growth form, with dense lower branches that are protected
from the wind by snow during the winter. Upper branches
develop only on the leeward side of the trunk because of
abrasion from blowing ice and snow. Most krummholz trees
are Engelmann spruce and subalpine fir, though whitebark
pine can have a similar growth form in northwestern

Wyoming. Common plants in the adjacent alpine fellfield
include alpine avens, alpine bistort, alpine sagewort, and spike
trisetum. Photo taken on Libby Flats in the Medicine Bow
Mountains. Elevation 10,800 feet.
that understanding the elevation of treeline depends as
matter and nitrogen under the trees. For a time, some
much on where tree seedlings are able to become estab-
alpine plants may do better on soils left behind after
lished as on climatic factors that limit growth.5
the trees have passed. However, wind scouring of the
Often the shrubby tree islands are initiated in slight
organic-rich soil most likely resumes after the protec-
depressions, where water is more readily available and
tion of windward branches is lost, leaving the tundra
seedling establishment is more likely to occur. Eventu-
with a soil similar to that prior to the establishment of
ally, after many years, the lower branches of the saplings
the first tree seedling.
are pressed to the ground by heavy snow and begin to
The islands of krummholz influence plant distribu-
produce roots. This process, known as layering, contrib-
tion in other ways, primarily through drifting snow
utes to the horizontal dimensions of the krummholz
and snowmelt water. 8 The snow provides protection
mat, creating the impression many years later of sev-
during the winter, and the temperature under snow
eral trees growing together.6 In fact, most tree islands
is warmer than on the soil surface of snow-free tun-
are one or a few clones developed from one or a few
dra. Downslope from the drifted snow, water is more
seedlings.
readily available during what would often be a dry
Once established, the patches of krummholz have
growing season, contributing to the amount of plant
several interesting features. First, they move very slowly
growth that can occur there. Commonly found under
downwind, perhaps an inch or so each yearthe result
the shrubby trees are such plants as dwarf huckleberry,
of abrasion on the upwind side and layering on the
mountain gooseberry, and other species associated
downwind side.7 As they move, the tundra soil under
with the less severe environment of subalpine forests
the krummholz is changed, with more soil organic
and meadows.
Though wind surely contributes to treeline forma-
violet radiation and the excessive reflection of solar
tion through its effect on snow accumulation and new
radiation from snow onto leavesboth of which can
seedling establishment, there is compelling evidence
retard physiological processes.
that heat deficiency is another cause.9 By their nature,

trees have woody stems and branches that must be
maintained by carbon-rich compounds produced by
Avalanches
photosynthesisa temperature-dependent process. As
Snow avalanche tracks are a conspicuous feature of
elevation increases, temperatures become too low for too
treelines where large volumes of drifting snow accumu-
much of the growing season to allow the amount of pho-
late, such as on the lee side of high ridges and where
tosynthesis required for the year-round maintenance of
snow slides are common (fig. 14.4). Such areas are par-
large plants. Hence, the trees become shorter and even
ticularly common in Colorado, northwestern Wyo-
shrubby in nature, and at some point, small seedlings
ming, and parts of the Northern Rocky Mountains.
cannot survive either. Worldwide, trees are absent where
Usually the snow accumulates in the alpine zone and
the growing season is shorter than 3 months and the

growing season average temperature is less than about
43F.10 Perennial herbaceous plants and small shrubs
are clearly better adapted for the cold alpine environment, probably because less photosynthesis is required
to maintain small plants and the roots and rhizomes of
such plants store carbohydrates for the production of
new leaves early the following year. Small plants also are
more likely to be protected by snow cover during winter.
Predictably, some trees have adaptations that
enable them to tolerate the cold treeline environment.
For example, treeline conifers have clustered needleleaf arrangements that tend to maintain daytime leaf
temperatures that are considerably warmer than the
ambient air.11 This temperature difference facilitates
photo
synthesis in the cool alpine environment and
occurs because leaf clusters (1) present a more dense
mass for the absorption of solar radiation and (2) minimize wind movement through the needles, thereby
reducing cooling. Limber pine and whitebark pine are
adapted in another way, namely, flexible branches that
reduce breakage from strong winds.12
The challenge of understanding treelines is illustrated by the hypothesis that trees cannot survive in
alpine environments because the summer is too short,
cool, and dry to allow development of the protective
tissues required to tolerate abrasion the following winter. Although plausible, Julian Hadley and William K.
Smith concluded that this hypothesis does not apply
to the Medicine Bow Mountains.13 Their results suggest
that wind-caused abrasion during the winter was more
important than a short growing season. Still other factors affecting treeline elevation are more intense ultra-
Fig. 14.4. Avalanche track in Cascade Canyon, Grand Teton

National Park. Tree damage is less frequent at the lower end
of the track, enabling the trees to grow taller there. However,
as the trees grow larger, they offer more resistance to the avalanche and are more likely to break or be uprooted. The trees
are subalpine fir and Engelmann spruce. The more flexible
shrubs higher on the slope include bearberry honeysuckle,
mountain ash, Rocky Mountain maple, rusty menziesia, and
serviceberry. Elevation at the bottom is 7,216 feet.
233
periodically slides along a topographic depression that

extends into the subalpine forest, ending in the runout
zone. Depending on the amount of snow and other conditions, avalanches can be gentle slides over short distances or thunderous events that break large trees in the
valley bottom. Sometimes several avalanches occur on
the same track in a single winter.
The vegetation of avalanche tracks is different from
the forest on either side or the alpine tundra above. In
the tundra, the track typically has sparse plant cover,
because the catchment area commonly has snow that
persists until midsummer, thereby shortening the growing season. Damage to small alpine plants is minimal,
because snow movement occurs within the snowpack
rather than at the soil-snow interface. Most physical
damage to plants occurs below the treeline, where tall,
woody plants are common. Even there small trees and
flexible shrubs are not damaged. However, trees with
diameters greater than 4 inches usually break, because
Fig. 14.5. White pine blister rust on a whitebark pine branch.

This exotic fungal parasite has killed a large portion of the
whitebark and limber pines in the Rocky Mountains and
is continuing to spread. The infection is fatal when the
parasite grows into the bark of the tree trunk. Photo by Anna
Schoettle.
boles of that size provide too much resistance to the

sliding snow.14
cut cone-bearing branches. Grizzly bears can switch to
Thus, there is a vertical gradient in the plant growth
other food sources, but the loss of whitebark pine over
forms that is correlated with avalanche frequency. Small
such a large area reduces habitat quality for this already
plants and flexible shrubs dominate at the top, with trees
threatened species.
becoming more common lower on the track because of
Millions of trees have been killed wherever five-
longer periods of uninterrupted growth. The largest trees
needle pines occur in North America, but the disease
occur at the bottom, which is rarely affected. Predictably,
currently is most severe in the Rocky Mountains. Infec-
the most severe disturbance occurs when trees are bro-
tion levels vary considerably, with the highest occurring
ken during large avalanches that extend to the runout
in close proximity to the rusts alternate hosts, which
zone. Such disturbances occur every 50100 years.
include wild currants and gooseberries (Ribes spp.) plus

a few other species.16 Some populations of the trees
White Pine Blister Rust at Treeline
seem resistant to the disease, which gives conservation

biologists hope that future generations of the pines will
A disturbing development is the invasion of an exotic
be more resistant. That seems to have happened around
disease that kills whitebark pinethe common species
the Great Lakes and farther to the east, where eastern
at treeline in the Greater Yellowstone Area and North-
white pine is still common today, even though it was
ern Rocky Mountains.15 Known as white pine blister rust
often infected by the blister rust in the mid-1900s. A
(Cronartium ribicola), this fungal parasite also kills lim-
complicating factor in western states, though, is that
ber pine and bristlecone pine in the Southern Rockies
whitebark and limber pine, even if generally resistant
(fig. 14.5). All three of these pines tolerate severe envi-
to the rust, may nevertheless be sufficiently weakened
ronments, and their peanut-sized seeds historically have
by the disease to be killed by the native mountain pine
provided an important food supply for grizzly bears,
beetle (see chapter 11). Research on this perplexing prob-
red squirrels, Clarks nutcracker, and other animals (see
lem and the resilience of the ecosystem is ongoing.17 As
chapter 10). The bears are able to build up fat reserves
at lower elevations, the nature of the ecosystem may
as they gorge themselves on the pine nuts, which some-
change considerably because of inadvertently intro-
times are accessible on the ground because red squirrels
duced invasive species.
Surviving in the Alpine Tundra

Above treeline, many of the shrubs and herbs that survive are also found in the Arctic. Both kinds of tundra
have cool, short growing seasons. However, there are
prominent differences: the Arctic has long summer days
with little diurnal temperature fluctuation, low ultraviolet and visible light intensities, and permafrost over
much of the landscape. In sharp contrast, alpine zones
have high intensities of ultraviolet and visible light,
shorter summer photoperiods, and great diurnal temperature extremes in the summeroften reaching more
than 68F during the day and dropping below freezing
at night. Also, permafrost is rare in the alpine tundra.18
Wind and cold are clearly important aspects of the
alpine environment, but drought is another factor
19
affecting plant survival.
Evapotranspiration is more
rapid because of lower atmospheric pressures at high elevations, which increases the probability of plant water
stress. Also, water uptake by roots is slower when the
soil is cold. Various investigators have noted the similarity of desert and alpine plants in terms of leaf size and
other structural features. Nutrients can be limiting as
well, which is partially compensated for by the presence
of mycorrhizae and nutrient resorption prior to leaf fall.
Notably, most of the root system, which has two to ten
times more biomass than the stems and leaves above
ground, is in the top 4 inches of the soil, where nutrient
availability is highest and the temperature is relatively
warm during summer.
Arguably, the alpine tundra has the most severe
conditions for plant growth to be found anywhere in
the region: extreme temperature fluctuations during a
24-hour period (which is very difficult for most plants
to tolerate); frequent freezing and thawing; rapid rates
of drying (away from melting snowdrifts); nutrient limitations; and a cool, short growing season. Still, several
hundred plant species tolerate this environment in
Wyoming alone.20
The predominant growth forms of alpine plants are
perennial herbs (including forbs, grasses, and sedges)
and short woody or semi-woody shrubs. Most biomass
is belowground. Annuals are rare because seedling
establishment is difficult in the alpine environment.21
Lichens are common, but they are not a dominant feature of alpine vegetation, as they are in the arctic tundra.
Annual plant growth of the tundra has been estimated

at 40 g/m 2/year on dry, windswept sites and 300 g/m2/
year on more protected, relatively moist sites.22 Such
growth rates, although low compared to subalpine forests, are reasonably high, considering that the growing
season is only 3075 days long.
Predictably, some unusual adaptations have evolved
to allow alpine plant growth.23 For example, all alpine
plants are capable of photosynthesis and growth at cold
temperatures, and many are drought tolerant.24 Evergreen and wintergreen leaves are common, enabling
growth early and late in the season, and even while
still covered with up to 20 inches of snow. Root systems
tend to be shallow, probably to facilitate nutrient uptake
as decomposition in the surface soils occurs, and the
roots usually have mycorrhizae that greatly increase the
surface area available for nutrient acquisition. Alpine
plants also seem well adapted for mitigating the adverse
effects of a short growing season. For example, alpine
pennycressa diminutive herbgrows from upper
treeline down into the foothills, but the higher populations complete their growth and reproduction more
rapidly than those at lower elevations.25
Another plant adaptation is the cushion plant
growth form, with branches and leaves that grow in
dense mats close to the soil surface (for example, moss
campion, alpine forget-me-not, and alpine phlox). Soil
temperatures during the day can be surprisingly warm
because of the intense sunlight at high elevations. The
dense branches absorb heat from the soilwarmed by
the sunand they slow the movement of air through
the plant, thereby slowing the rate of cooling. Dark color
and epidermal hairs also help maintain warmer leaf and
bud temperatures in this otherwise cool or even cold
environment. Some insects depend on the more moderate microclimate of the cushion plants.26
Reproduction of alpine plants is largely vegetative,
that is, sprouting from roots and rhizomes, because of
the difficulties associated with seedling establishment
in such a rigorous environment. Nevertheless, many
plants have evolved special adaptations for seed production. Some plants have unusually large and showy
flowers, presumably to attract scarce insect pollinators
during the short warm periods when the insects are
active. Flowers with large petals serve as windbreaks
for insects and are thought to provide a warmer micro
235
most of their livesand under snow for much of the

yearthey avoid bone-chilling winds and the coldest
temperatures. 30
Tundra Mosaic, Frost, and Burrowing Mammals

The plant communities of alpine tundra can be easily
classified into one of the following categories: fellfields,
alpine turf, wet meadows, and willow thickets.31 There
are also snow-beds, talus slopes, and boulder fields,
all of which have little or no plant growth. Lakes and
streams are common, with associated wetland vegetation that includes sedge-dominated wetlands as well as
willow thickets (see chapter 5).
Fig. 14.6.The big flowers of this 6-inch-tall alpine tundra
plant, known as alpine sunflower or old-man-of-the-mountain, attract pollinating insects during the short cool summer.
Alpine tundra is commonly discussed at three spatial scales. At the scale of tens of miles, variation is
caused by differences in local climate. 32 The tundra of
the Southern Rockies is different from that of the Cen-
environment (fig. 14.6).27 Because pollination often
tral and Northern Rockies. In contrast, at the scale of a
does not occur during the short summer, a high per-
few feet, abrupt differences can be caused by the pres-
centage of alpine plants are capable of producing seeds
ence of boulders, which absorb and store considerable
without fertilization, a process known as apomixis.
heat from the intense sunlight at high elevations, and
Animals have equally interesting adaptations. Aside
then release the heat at night to the benefit of plants
from fish, all vertebrates in the alpine zone are warm-
growing nearby. Also, snow accumulates on the lee side
blooded endotherms, which is adaptive as long as they
of boulders, and water from rain and condensation
have the food required to maintain body temperature.
drains rapidly to the soil below the boulders, where it
No alpine amphibians or reptiles have been found.
is less likely to evaporate. Because of these influences,
Some animals migrate up and down the mountain to
boulders provide warmer, wetter microenvironments
find food and shelter. During the winter, a few species

of birds leave the region altogether, like the American
pipit. Other animals have the extraordinary ability
to stop eating while they hibernate, such as the marmot. Mice, voles, and pocket gophers consume roots
and other accessible plant materials in their burrows,
even under the snow, whereas the pika works all summer long to create a cache of food deep in the boulder
fields and talus slopes where it lives (fig. 14.7). Known
as haypiles and composed largely of alpine avens and
other forbs, these caches are quite large, weighing on
average about 60 pounds in the Colorado alpine. For
comparison, each pika weighs less than 7 ounces.28
There can be six to eight pikas per acre, each one feeding outward from its nest. 29 Through their burrowing,
all small mammals contribute to soil development and
have effects on the vegetation where they live. Tons
of soil are aerated. By living in soil or under rocks for
Fig. 14.7.The American pika, 68 inches long, lives in alpine

boulder fields and talus slopes, where it creates haypiles for
warmth and food during the winter. It does not hibernate.
Copyright Wendy Shattil/Bob Rozinski.
SNOW COVER
Snowfree
Winter snow
cover
TOPOGRAPHIC
POSITION
Fellfield
Summit
or
ridge
Persistent
snow
Leeward
or tree islands typically have drifted snowbanks that

persist until midsummer. Few if any plants can survive
where snow persists late into summer, but plants growing downslope from late-lying snow have a dependable
Upper
slope
source of water for most of the growing season. Consequently, the vegetation there has higher cover and
Turf
growth. Moreover, the dominant plants of these mead-
Tufted
hairgrass meadow
Snow
accumulation
tion during the winter (fig. 14.8). The lee sides of ridges
Sedge meadow
Lower
slope
Ravine
Willow or fen
Windward
WIND EXPOSURE
Fig. 14.8. Distribution of different kinds of alpine vegetation
in relation to topographic position, wind exposure, and snow
cover. Adapted from Johnson and Billings (1962).
ows are different (table 14.1).

Perhaps the most unique process that causes pattern in the alpine landscape is cryoturbationthe freezing and thawing of moist soils that occurs on time
scales of a single day or a year. Soil water expands as
it freezes, pushing comparatively large objects, such
as stones and boulders, along the path of least resistanceusually toward the surface. After thousands of
freeze-thaw cycles, the larger objects are sorted from
smaller ones, creating patterned ground, such as frost
boils, stone nets, and stone polygons (fig. 14.9). On a
that allow some plants to grow where they might not
24-hour cycle, needle-ice forms and melts, which can
otherwise.
lead to the formation of frost boils on the soil surface
At an intermediate scale, differences are caused by
that damages the roots of seedlings (fig. 14.10). 33
topography and the presence or absence of tree islands,
Another frost-related process is solifluction, in which
as discussed previously. On windswept ridges or flats
frost and gravity combine to move wet soil downslope.
are fellfields, dominated by cushion plants and other
Solifluction occurs most often on lee slopes, where
drought-tolerant plants that require little snow protec-
snow accumulates and the soil becomes wet, gradually
Fig. 14.9. Alternating freezing and

thawing over long periods causes large
stones and rocks to be separated from
fine-textured soil particles, forming
stone polygons or nets, such as in this
area in the Beartooth Mountains. The
alpine turf is dominated by various
alpine sedges along with alpine avens,
alpine bistort, alpine sagewort, snow
willow, tufted phlox, and other species.
237
Table 14.1. Some characteristic plants of alpine plant communities in Wyominga
Common name
Latin name
Fellfield
Alpine turf
Wet
meadow
Willow
thicket
FORBS
Arctic alpine forget-me-not
Eritrichium nanum
Alpine leafybract aster
Symphyotrichum foliaceum
Alpine mountain sorrel
Oxyria digyna
Alpine sagebrush
Artemisia scopulorum
American bistort
Polygonum bistortoides
Creeping sibbaldia
Sibbaldia procumbens
Dwarf clover
Trifolium nanum
Graylocks four-nerve daisy
Tetraneuris grandiflora
Hookers mountain avens
Dryas octopetala var. hookeriana
Lesser spikemoss
Selaginella densa
Moss campion
Silene acaulis
Parrys clover
Trifolium parryi
Parrys lousewort
Pedicularis parryi
Redtop stonecrop
Rhodiola rhodantha
Rocky Mountain nailwort
Paronychia pulvinata
Rocky Mountian pussytoes
Antennaria media
Ross avens
Geum rossii
Sticky polemonium
Polemonium viscosum
Tufted phlox
Phlox pulvinata
Varileaf cinquefoil
Potentilla diversifolia
Caltha leptosepala
Western yarrow
Achillea millefolium var. occidentalis
GRASSES
Alpine bluegrass
Poa alpina
Timberline bluegrass
Poa glauca ssp. rupicola
Alpine timothy
Phleum alpinum
Purple reedgrass
Calamagrostis purpurascens
Rocky Mountain fescue
Festuca saximontana
Spreading wheatgrass
Elymus scribneri
Spike trisetum
Trisetum spicatum
Tufted hairgrass
Bellardi sedge
Kobresia myosuroides
Blackroot sedge
Carex elynoides
SEDGES
Curly (rock) sedge
Carex rupestris
Drummonds sedge
Salix drummondiana
Common name
Latin name
Fellfield
Alpine turf
Wet
meadow
Willow
thicket
Ebony sedge
Carex ebenea
Obtuse sedge
Carex obtusata
Water sedge
Carex aquatilis
Alpine laurel
Kalmia microphylla
SHRUBS
Alpine willow
Salix petrophila
Cascade willow
Salix cascadensis
Diamondleaf willow
Salix planifolia
Grayleaf willow
Salix glauca
Snow willow
Salix reticulata
Resin birch
Betula glandulosa
slipping or creeping downward. Formed over long peri-
about whether a stable tundra ecosystem ever devel-
ods, solifluction lobes or terraces cause a distinctive pat-
ops. However, gradual changes occur over time scales
tern (fig. 14.11).
of centuries. To illustrate, with gradual weathering and
Cryoturbation and solifluction, combined with the
soil development, boulder fields develop into fellfields,
burrowing of pocket gophers (fig. 14.12), are thought
which can become dry meadows, commonly known as
to be continual sources of disturbance to alpine vegeta-
alpine turf. Similarly, wet meadows dominated by water
tion, creating a mosaic of patches and raising questions
sedge develop into somewhat drier meadows dominated

by tufted hairgrass. On a smaller scale, episodes of cryoturbation cause changes in the dominant plants. 34
Disturbances that occur in alpine tundra are distinctly different from those occurring in ecosystems at
lower elevations. However, numerous plants and animals have evolved adaptations for this environment
and are not found elsewhere. Some observers refer to
tundra as fragile, because recovery after disturbances is
slow. To be sure, excess livestock grazing and damage
caused by foot traffic and off-road vehicles can cause
undesirable changes that persist for many decades. 35
Nitrogen Deposition
Nitrogen fertilization experiments in alpine tundra
Fig. 14.10. Frost boils, such as this one on Libby Flats in the
Medicine Bow Mountains, create small patches in the tundra
vegetation. Burrowing mammals, like the Wyoming ground
squirrel, may be beneficiaries of the loose soil, or they may be
involved with forming such features. Elevation 10,500 feet.
commonly lead to an increase in plant growth, indicating that nutrients can be limiting along with the cool,
short growing season.36 Nitrogen-fixing plants do occur,
such as alpine clover, but they are not widespread.
239

Fig. 14.11. Solifluction lobes or terraces
develop over a fellfield on slopes in
alpine tundra, such as in this area in
the Beartooth Mountains. Solifluction
creates different environments for plant
growth over short distances. Water
seeping through the lobes provides an
ideal environment for barrenground
willow. Elevation about 10,600 feet. See
Johnson and Billings (1962).
Limiting nutrients tend to be conserved in plant and
nutrient source. One study concluded that the eutro-
microbial biomass and will not normally be leached or
phication of Rocky Mountain aquatic ecosystems from
lost from the soil except after disturbances.37
atmospheric pollutants is analogous to the acidification
It was unusual, then, when investigators observed

high concentrations of nitrogen in the creeks drain-
of terrestrial ecosystems associated with the same pollutants in northeastern North America.42
ing the alpine zone of Rocky Mountain National Park

and other parts of the Colorado Front Range. 38 No obvious disturbances were evident, but further research led
to the conclusion that the addition of nitrogen in air
pollutants, mainly nitrous oxides, was causing a condition known as nitrogen saturation, when the amount
of nitrogen added to the ecosystem is essentially the
same as the amount lost.39 Nutrient enrichment, known
as eutrophication, leads to changes on the upland and,
downstream, enables algal blooms that can reduce fish
productivity and recreational opportunities.40
Notably, the added nitrogen apparently is first
deposited at high elevations in the tundra.41 Referred
to as nitrogen hotspots, such areas have large amounts
of bare rock and a small amount of biomass for nitrogen uptake. Moreover, much of the nitrogen is added in
snowfall that is blown into drifts, thereby concentrating the nitrogen in places that have the least amount of
plant growth and storage capacity. In contrast, the subalpine forest and many subalpine meadows have higher
nitrogen-storage capacities and currently are less of a
Fig. 14.12. Pocket gophers, though rarely seen, are common

in alpine tundra and subalpine meadows and often burrow in
the soil under snowdrifts, where they eat roots. The soil from
their burrows is pushed into tunnels through the snow, which
become middens when the snow melts. These small mammals
aerate the soil and facilitate nutrient cycling. The plants in
this area are sparse, because snow cover persists until early
July.
The response of tundra plants to nitrogen enrich-
higher precipitation (see chapter 3). Tree establishment
ment is currently under investigation. University of
could be favored by both warmer temperatures and less
Colorado ecologist Timothy Seastedt and his associ-
drought stress, but apparently such conditions have
ates have observed that willows are expanding their
not yet developed in some areas, such as in northern
distribution in the alpine zone, possibly because of
Colorado.49 Moreover, with ongoing disturbances (such
the combination of added nitrogen and climate warm-
as frost heaving, burrowing by small mammals, wind
ing. This result might be expected, he suggests, in wet
abrasion, and herbivory), the chances of new tree seed-
areas that are protected from wind scour, that is, places
ling establishment in the tundra are still lowand prob-
where organic matter accumulates as well as nitrogen-
ably will be for many years to come.50 Alpine ecologist
enriched snow and water. The more exposed dry mead-
Christian Krner concluded in 2012 that the advance
ows and fellfields would receive less nitrogen and would
of treelines will always lag behind climate change by
be less likely to change. Thats the hypothesis.
at least 50100 years, considering how long it takes for
43
new seedlings to become established and grow to tree
Advancing Treelines and Climate Change
height in the alpine environmenteven if it is slightly

warmer.
It is well known that treelines have moved up and
But not all alpine treelines are the same. For exam-
down during the past 10,000 years (see chapter 2).44 But,
ple, new tree seedlings have been found above treeline
if snowmelt is occurring earlier in the spring because
on relatively moist sites in the Front Range of Colo-
of climate warming, and glaciers are retreating, is the
rado.51 Also, in the Uinta Mountains of Utah, photos
alpine treeline moving upward throughout the region?
taken of the same place in 1870 and 2000 indicate a rise
Answering this question has been the subject of con-
in treeline of 180550 feet.52 In the same area, plants
siderable research, especially since the mid-1990s.45 The
other than trees have been observed moving upward in
results are mixed and offer an interesting example of
elevation through seedling establishment. Such upward
the complexities of interpreting the effects of climate
migration might have occurred even without the cur-
change. In general, the same results should not be
rent warming trends, but vegetation response to warm-
expected for alpine treelines everywhere.
ing clearly varies considerably.
Fundamental to treeline advance is the ability for
As pertinent to the effect of warming on alpine plant
new tree seedlings to become established higher on the
distribution is the paradox that earlier snowmelt in the
mountain than their parent trees could, that is, higher
spring from warming increases the probability of frost-
than the krummholz that already exists at treeline.46 In
related plant death or damage. Some alpine plants are
some places, newly established tree seedlings in the tun-
sensitive to cold temperatures after they begin growth,
dra have been hard to find, suggesting that trees at this
which is delayed by snowcover. If the snow melts ear-
elevation are not a good indicator of climate change.47
lier, frost damage is more likely.53 Global warming that
How could this be? One study in Rocky Mountain
causes higher mortality could retard the rate at which
National Park concluded that the treelines are relictual
treeline plants respond. Also, earlier snow-free dates
they persist because adult trees tolerate the present-day
could lead to colder soils and more cryoturbationtwo
climate, even though the more delicate seedlings can-
factors that reduce the chance of seedling survival.
not.48 In other words, the tree seedlings that founded
Climate warming also will affect alpine animals that
the tree islands of today became established at a time
are restricted to mountain tops. On mountains with less
when the climate was more favorable. Numerous seed-
land in the alpine zone, the populations of alpine species
lings were found invading openings in the subalpine
are smaller; and if treelines advance upward because of
forest below, especially in moist areas, but hardly any
climate change, their populations will become smaller
were found in the drier openings at treeline or above.
still, increasing the chances of local extinction. Ani-
In addition to warmer temperatures, one of the
mals on one mountain may be able to travel to others
predictions for climate change in the alpine zone is
nearby if they can fly or if they are able to survive or
241
tolerate the subalpine forest ecosystem as they move
able to survive below treeline where subalpine meadows
from one alpine zone to another. Populations of the
and talus slopes are in close proximity.
same species farther north, where there is more tundra

because treelines are lower, may have a greater chance
Overall, sorting out the causes of shifting treelines
of surviving.
and nutrient enrichment is a challenge that requires
Such discussions invariably consider the plight of
an understanding of microclimate; wind; soil; cryo
the American pika, a small mammal in the rabbit fam-
turbation; the effect of cold night skies; the chemistry of
ily that essentially is restricted to alpine tundra (see
the atmosphere and soil; the various factors that influ-
fig. 14.7). It nests in the crevices of boulder fields and
ence streamwater chemistry; and the adaptations of the
frequently is seen scurrying from one rock to another,
plants, animals, and microorganisms found on land
often stopping to bark along the way. If the forest mar-
and in water. Treelines are likely to expand upward,
gin continues to move upward because of global warm-
slowly at first. The alpine tundra, though far removed
ing, pika habitat will be squeezed into smaller and
from urban-industrial developments at low elevations,
smaller patches. At present, the alpine treeline appears
is nevertheless another bellwether of changes under
to be moving upward very slowly, if at all, and pikas are
way throughout the region.
Part Five Landscapes of
Special Interest
The Greater
Yellowstone Ecosystem
Chapter 15
The Greater Yellowstone Ecosystem (GYE) covers some
depth of 3 inches as far away as present-day Iowa. This
40,000 square miles and encompasses the largest wild-
time, with the loss of so much material within a mat-
land in the contiguous United States. It includes two
ter of hours or days, the center of Yellowstone country
national parks, seven national forests, ten designated
collapsed on itself, forming a giant depression known
wilderness areas, three wildlife refuges, and a variety
by geologists as a caldera. The underlying magma was
of other public, tribal, and private lands (fig. 15.1).1
still hot, however, and a long series of lava flows began
The influences of modern industrial society have been
filling in the depression, with major flows occurring as
minorcompared to those on the surrounding area
recently as 70,000 years ago. The thermal features that
and with the reintroduction of the gray wolf in 1995,
so many visitors enjoy today are evidence that Yellow-
all of the GYEs original pre-Columbian mammals are
stone is still underlain by a hot spot. The Yellowstone
present. Reading a list of the names of Yellowstones
caldera occupies about a third of YNP. The high cliffs
largest mammalselk, bison, wolves, moose, and griz-
along the north side of the Madison River are part of the
zly bearsis akin to contemplating Tyranosaurus rex,
north rim, Mount Washburn (10,223 feet) lies on the
Apatosaurus, and Pterodactylus coexisting in a landscape
northeastern edge, Mount Sheridan (10,298 feet) is on
with mysterious plumes of steam and water emanating
the southern edge, and Canyon Village is in the bottom
from the earth. Little wonder that the early explorers
at 7,915 feet. West Thumb and the central and northern
referred to the area as a wonderland.
portions of Yellowstone Lake also lie in the caldera.
In the Eocene, 55 to 35 million years ago, a series of
The processes that formed the Teton landscape, just
volcanic eruptions literally buried the mountain ranges
south of YNP, are different but no less amazing.5 Some
in and around what is now Yellowstone National Park
40 million years after mountain formation to the north,
(YNP) (see chapter 2). New forests developed after each
east, and south, the TetonJackson Hole region was still
eruption, one on top of the other. Fossil tree trunks are
relatively flat. The ancient Precambrian rocks were bur-
still visible today (fig. 15.2).3 The Absaroka Range, on
ied under nearly 4 miles of younger sedimentary and vol-
the horizon for someone looking east across Yellow-
canic rocks. Then, about 9 million years ago, uplifting
stone Lake, is a remnant of Eocene volcanic activity.
began along the 50-mile-long Teton fault, creating the
Most of the Eocene deposits to the west of the Absaro-
youngest range in the Rocky Mountains. As the Tetons
kas are gone, literally blown away by some of the most
rose, the softer rocks eroded. Today, essentially all sedi-
dramatic geological events in Earths history.4 Volcanic
mentary strata have been stripped from the steep eastern
activity occurred again, about 650,000 years ago, with
slopes and the highest peaksexcept for a telltale sand-
the ash deposited across much of North Americato a
stone remnant on the top of Mount Moran. Geologists
245
246 Landscapes of Special Interest
Fig. 15.1.Elevation map for the Greater

Yellowstone Ecosystem. See fig 1.3 for a
map showing land under the jurisdiction
of different agencies, and fig. 1.5 for the distribution of different kinds of vegetation in
the area. The land area covered by this map
is about 50,000 square miles. Cartography
by Ken Driese.
know this remnant is part of the same Flathead sand-
the bottom of Jackson Hole to the top of Grand Teton (fig.
stone that now lies about 6 miles below the valley floor.
15.3). As expected for a mountain range created along a
Slippage along the fault continues at about 12 inches
fault line on the east side, the west slopes are gentler and
per century in some places, with the valley floor sink-
are still covered by sedimentary strata (fig. 15.4).
ing while the mountains rise. Seven peaks in the Tetons
With regional uplifting, continental cooling, and cli-
rise above 12,000 feet, with Grand Teton the highest
mate change, glaciers crept over the Yellowstone region
at 13,770 feet. Nowhere else in the Rocky Mountains is
and then retreated numerous times (see chapter2). The
there such an abrupt rise in elevation6,500 feet from
most recent glaciation, known as the Pinedale, began
Fig. 15.2. (left) Petrified tree trunks on a ridge south of the

Lamar River valley, the remnants of forests buried by volcanic
deposits in what is now Yellowstone National Park during
Eocene eruptions, about 50 million years ago. The fossil trees
in the region include specimens of chestnut, magnolia, maple,
oak, redwood, sycamore, and walnutnone of which grow in
the area now. At the present time, Douglas-fir is common on
this steep north-facing slope. Elevation 8,400 feet.
Fig. 15.3. (below) The Tetons rise abruptly above the Snake
River and Jackson Hole to a maximum elevation on Grand
Teton of 13,770 feet. Shrublands dominated by mountain
big sagebrush are characteristic of the flat glacial outwash
plains, at an elevation of 6,600 feet, including the lower
terrace carved during the Pleistocene Epoch. Blue spruce,
Engelmann spruce, narrowleaf cottonwood, river birch, silver
buffaloberry, and various willows are common in the riparian
woodlands. Douglas-fir occurs on the slopes above the river.
See also fig. 2.6.

Teton Mountains
WEST
EAST
Grand Teton
13,770 ft.
n
to t
Te aul
F
Precambrian
sea level
0
0
Jackson Hole
Blacktail Butte
Jackson
Quaternary
Mesozoic
Paleozoic
10 km
10 mi
Fig. 15.4. Geologic structure of the Teton Range and Jackson

Hole. Continuing displacement along the Teton fault has
created a mountain range without foothills. The Quaternary
deposits are mostly alluvium, often referred to as glacial outwash plains, deposited as the glaciers melted. Adapted from
Love et al. (2003).
about 80,000 years ago and reached its maximum extent
in pothole topography. Also known as kettles, these
some 25,000 years ago.7 At its peak, ice covered almost all
potholes were created by the melting of huge blocks
of what is now YNP and was about 4,000 feet thick above
of ice buried in the moraines. Water-filled depressions
the Yellowstone Lake basin. Ice buried and subsequently
remained after the ice disappeared (see chapter 5).
carved many mountains in the area, including Mount
Based on fossil pollen records from lake sediments,
Washburn and Mount Sheridan. The ice also dammed
paleoecologist Cathy Whitlock and colleagues deter-
the Yellowstone River, causing Yellowstone Lake to
mined that tundra was prevalent in much of the GYE
expand and cover the area that today is Hayden Valley.
about 14,000 years ago.9 By about 11,500 years ago,
Fine-textured sediments deposited during that time in
Engelmann spruce began to dot the tundra where a
the bottom of the lake now support extensive meadows.
more fertile substrate had developed from andesitic
Several thousand feet of ice also covered the north-
rocks. Whitebark pine and subalpine fir joined the
ern part of Jackson Hole at times,8 and subsequent gla-
spruce about 11,000 to 9,500 years ago, but the less fer-
ciers flowing down the canyons of the Tetons created
tile rhyolitic soils of the Yellowstone Plateau remained
terminal moraines at the foot of the mountains, form-
unforested during this period. The climate gradually
ing lakes known today by the names of early explor-
became warmer, though it was still cooler and more
ersJackson, Bradley, Jenny, Leigh, Phelps, and Taggart.
humid than it is today. Continued warming led to the
Glaciers also shaped the jagged peaks and U-shaped val-
expansion of lodgepole pine throughout the region.
leys of the Teton, Beartooth, and Wind River Mountains
Whitlocks data indicate that limber pine was present at
(see fig. 2.6).
low elevations and that Douglas-fir expanded to higher
The glaciers began to shrink as the climate warmed,
elevations about 9,500 to 4,500 years ago. Subsequently,
about 20,000 years ago, leading to the gradual elimina-
the climate became cooler and wetter at high eleva-
tion of the vast Yellowstone ice field. Evidence suggests
tions, enabling spruce and fir to become more common,
that a glacier persisted in Jackson Hole until as recently
along with lodgepole pine, as they are today. Douglas-fir
as 9,000 years ago. It would have been witnessed by
now grows at lower elevations.
some of the earliest Native Americans as they entered
Today, the GYE spreads over fourteen mountain
the valley. With rapid thawing of huge masses of ice,
ranges and is a major source of water for three major
rivers became swollen with water and sediment. Great
river systemsthe Green flowing into the Colorado, the
floods occurred as moraines and ice dams collapsed,
Snake into the Columbia, and the Yellowstone into the
depositing large outwash plains (such as Antelope Flats)
Missouri and Mississippi.10 About a third of the land is
and scouring stream bottoms. The extensive terraces
forested, with lodgepole pine, Engelmann spruce and
along the Snake River were formed during this time (see
subalpine fir, and aspen dominating an estimated 20,
fig. 15.3). Further evidence of the glaciers is apparent
12, and 3 percent of the land, respectively. About 10
The Greater Yellowstone Ecosystem
percent is alpine tundra. Shrublands dominated by big
toward Yellowstone and the Tetons.17 He was followed
sagebrush are characteristic of the lowlands, growing on
by various trappers, including Osborne Russell, who
about 25 percent of the GYE.
wrote about the environment, wildlife, and people of

the Yellowstone region in the early 1800s.18
Today YNP occupies an area of 3,468 square miles.
Yellowstone National Park
The highest mountain peaks occur in the Absaroka
Early explorers along the headwaters of the Missouri
Mountains on the eastern boundary, rising to slightly
and Columbia rivers returned with tales of abundant
more than 10,900 feet; the lowest elevation is at Gar-
fish and wildlife, boiling mud, steam rising from holes
diner in the northwestern corner, where the Yellow-
in the earth, hot springs, spectacular waterfalls, and a
stone River leaves the park at 5,314 feet. More than 150
deep canyon with walls of yellow stone. So incredible
lakes cover 5 percent of the park, all small except for
were their stories that few took them seriously. Some
Yellowstone, Shoshone, Lewis, and Heart lakes. Soils
newspapers hesitated to print their reports. In 1870,
have developed mostly on two kinds of volcanic rock
General Henry D. Washburn led an expedition to eval-
the infertile rhyolite and more fertile andesiteplus
uate some of the claims. Cornelius Hedges, a member
the gravel, sand, silt, and clay deposited by retreating
of the expedition, was probably trying to preserve his
glaciers. The highest mean annual precipitation, up to
reputation as a keen observer when he commented,
70 inches, occurs on the high Madison and Pitchstone
I think a more confirmed set of skeptics never went
plateaus in the southwestern corner of the park and in
out into the wilderness than those who composed our
the Gallatin Mountains in the northwest, with the inte-
party, and never was a party more completely surprised
rior and lower elevations of the park receiving much
and captivated with the wonders of nature.12 Hedges
less (only about 10 inches at Gardiner, Montana). Two
complained of the wind and cold in August but, after
distinct climate regimes are recognized in YNP and the
11
walking along the rim of the Grand Canyon of the Yel-
GYE. In the areas located generally south and west of
lowstone River, he was filled with too much and too
the Continental Divide, winters are wet and snowy, and
great a satisfaction to relate.13
summers are relatively dry; but in areas located gener-
A year later, in 1871, Ferdinand V. Hayden led a sci-
ally north and east of the Continental Divide, winters
entific expedition to the region, sponsored by the U.S.
are somewhat drier, and summers are relatively wet.
Congress. Haydens report, with stunning photographs
Some vegetation patterns reflect this climatic pattern;
by William H. Jackson and paintings by Thomas Moran,
for example, aspen is more abundant in the winter-wet
was instrumental in Yellowstone being established in
region in the southern part of the GYE than in the sum-
1872 as the worlds first national park.
mer-dry region to the north.19
14
The accolades
continued. In 1875 General W. E. Strong wrote, Grand,
Rivers are a popular feature of the Yellowstone land-
glorious, and magnificent was the scene as we looked
scape, because they are big and clean. Pulses of sedi-
upon it from [Mount] Washburns summit. No pen can
ment-rich water occur following natural erosion events,
write itno language describe it. Today the wonders
especially in the spring and after heavy thunderstorms
of Yellowstone still entice and excite several million
over recently burned forests, but most of the time the
visitors each year.
rivers run clear. They also offer some of the finest trout
15
Of course, the ancestors of modern tribes had been
fishing in the world, with the native cutthroat trout con-
present 8,00010,000 years before these explorers
sidered a prize. Canyons, point bars, cut banks, terraces,
arrived.16 Aside from hunting and gathering their food,
and broad floodplains are part of the riparian mosaic
they collected and traded obsidian, a hard volcanic rock
(see chapter 4). Rapids and waterfalls are common. The
highly valued for spear points, arrowheads, and cutting
Grand Canyon of the Yellowstone River is about 1,500
implements. When Europeans first arrived, the major
feet deep, passing through rhyolite softened by hydro-
tribes living in the area were the Blackfoot, Crow, and
thermal activity. The canyon begins at Lower Falls, with
Bannock-Shoshone. In 1806 John Colter left the Lewis
a drop of about 300 feet, and continues for about 18
and Clark expedition in Montana and traveled south
miles to the mouth of the Lamar River.
249
Vegetation of Yellowstone National Park

About 80 percent of YNP is covered by forest (see
fig. 1.5). The parks forests appear uniform to many
visitors, yet ecologists have identified more than
thirty different forest environments (habitat types
and phases). 20 Lodgepole pine predominates over large
areas, especially on the rhyolite plateaus (see chapter
10). Lodgepole is probably the only tree capable of
thriving on such infertile soils; indeed, without lodgepole pine, there might be little or no forest over much
of the park. Some twenty-four habitat types in which
lodgepole pine is a major constituent have been identified. Some of the distinguishing understory species
include globe huckleberry and bluejoint reedgrass on
relatively moist and fertile sites, with grouse whortleberry, pinegrass, and elk sedge on dry and infertile
sites. Engelmann spruce and subalpine fir also are common and in some places dominant, especially on more
fertile substrates derived from andesite or sedimentary
rocks. Whitebark pine is common in high-elevation
forests (fig. 15.5), and Douglas-fir is common in the
foothills (fig. 15.6). 21 Tree-dominated riparian woodlands are not common in the park, but where they do
occur, the principal trees are aspen, balsam poplar,
narrowleaf cottonwood, and sometimes Douglas-fir,
Engelmann spruce, and lodgepole pine. 22
Fig. 15.5. Whitebark pine, shown here at the top of Mount

Sheridan in Yellowstone National Park, is the predominant
treeline species in northwestern Wyoming and the Northern
Rocky Mountains. In many areas the trees have been killed
during the past two decades by native mountain pine beetles
and the non-native white pine blister rust. Elevation 10,308
feet at the summit.
Nonforested land occupies most of the remaining

20 percent of YNP.23 Foothill grasslands dominated
grass. Interspersed throughout the subalpine forests are
by bluebunch wheatgrass and Sandberg bluegrass are
meadows dominated by tufted hairgrass and a variety
found at low elevations near Gardiner, where annual
of sedges and forbs.
precipitation is only about 10 inches. Also found in this
Several species of sedge are especially common in
area are desert shrublands dominated by fringed sage,
fens and wet meadows (see chapters 4 and 5), includ-
Gardner saltbush, greasewood, pricklypear cactus, and
ing beaked sedge, Buxbaums sedge, livid sedge, mud
winterfat. Meadows at low elevations along the Lamar
sedge, Northwest Territory sedge, slender sedge, and
River have the introduced Kentucky bluegrass, such as
water sedge. Sphagnum and philontus mosses occur
near the site of the old Buffalo Ranch in the Lamar Val-
with sedges in some areas. Wetlands with standing water
ley, along with native species that include Wheeler blue-
have hardstem bulrush and few-flowered spikerush,
grass, sheep sedge, and bearded wheatgrass.
with diamondleaf willow and Wolfs willow along the
Big sagebrush is found in the Yellowstone and Lamar
margin.24 In the riparian zone both meadows and shrub-
river valleys, where large numbers of elk and bison
lands occur. Riparian meadows typically have bluejoint
spend the winter; silver sagebrush and shrubby cinque-
reedgrass; tufted hairgrass; and other grasses, sedges, and
foil are found in meadows where the soils stay wetter
forbs. Shrublands along the creeks and rivers have alder,
for a longer time, such as in Hayden and Pelican val-
water birch, shrubby cinquefoil, silver sagebrush, and
leys (fig. 15.7). Associated plants include Idaho fescue,
various species of willow (diamondleaf willow, Geyers
junegrass, thickspike wheatgrass, and bearded wheat-
willow, sage willow, tealeaf willow, Wolfs willow, and
Fig. 15.6. Mountain big sagebrush steppe and foothill grasslands are widespread in the valleys of the Lamar and Yellow
stone rivers, an area known as the northern winter range
of Yellowstone National Park. The forests and woodlands at
this elevation are dominated by Douglas-fir, many of which

have been killed by the Douglas-fir beetle. The grasslands are
dominated by bluebunch wheatgrass and Sandberg bluegrass.
others).25 Dwarf willows (less than 6 inches tall) can be
thermal springs develop periodically, sometimes kill-
found above treeline, along with other alpine plants
ing trees over large areas.
common in the region (see chapter 14).

Several plant species are found only around hot
springs and geyser basins, including Ross bentgrassa
Grand Teton National Park
plant that is endemic to Yellowstone. The mineral-rich
Ferdinand Hayden returned to Yellowstone country in
water of these thermal features creates a saline envi-
1872 and sent a detachment, including photographer
ronment, and consequently, the characteristic plants
William H. Jackson, to explore Jackson Hole. The party
include such halophytes as alkali cordgrass, Baltic rush,
collected information on natural resources, took some
meadow barley, and seaside arrowgrass. The chemical
photographs, named many of the lakes and mountain
composition of the water varies from one geyser basin
peaks in Jackson Hole and the Tetons, and claimed to
to another, with the result that both terrestrial and
have been the first EuroAmericans to climb the Grand
aquatic species composition varies also. 26 For example,
Teton, though that has been disputed.27
the rock formed at Mammoth Hot Springs is rich in cal-
After 1872 the lands in YNP were off limits to set-
cium carbonate (travertine), but the rock around other
tlers, but Jackson Hole was still available. Named after
springs is rich in silica (sinter, also known as geyser-
early fur trader David Jackson, the town of Jackson
ite). Much of the color variation among geyser basins
was established in 1897.28 Cattle ranching became an
is caused by the varying temperature tolerances of dif-
important occupation in the valley at the same time as
ferent species of bacteria and algae, some of which are
potato farming developed in nearby Idaho at lower ele-
able to survive at near-boiling temperatures. New geo-
vations. To provide a more reliable source of irrigation
251
Fig. 15.7. Meadows and wetlands cover much of Hayden

Valley in Yellowstone National Park. The high point in the
distance is Mount Washburn, on the northern rim of the Yellowstone caldera. The valley is in the caldera and is underlain
by lake-bottom sediments deposited early in the Pinedale
glacial period. Moist meadows support mountain silver
sagebrush and shrubby cinquefoil; on drier sites, Idaho fescue

and Wyoming big sagebrush are common. Where the soil is
flooded during much of the summer, water sedge and beaked
sedge are common. The trees in the foreground are lodgepole
pine. Elevation 7,400 feet.
water, primarily for Idaho farmers, a dam on the Snake
on the mission of enlarging the park by buying as much
River was completed in 1906, thereby enlarging Jackson
land in Jackson Hole as possible and then donating it
Lake.29 Perennial concerns about possible dam failure,
to the federal government. 30 This was accomplished in
triggered by the 1976 failure of the Teton Dam 30 miles
1950, though not without criticism by many local land-
west (in Idaho), led to reinforcements on the Jackson
owners. As a compromise to enable park enlargement,
Lake dam (completed in 1989).
provisions were made for continued livestock grazing
The abundant wildlife and spectacular scenery south
and hunting. In 1972, the land along the highway con-
of YNP increased in popularity, and some began to
necting YNP and Grand Teton National Park was trans-
think that a second national park should be established.
ferred to the National Park Service by the U.S. Forest
Pierce Cunningham, a rancher near Spread Creek, cir-
Service and named the John D. Rockefeller Jr. Memorial
culated a petition in 1925 asking the state of Wyoming
Parkway, thereby acknowledging his vision.
or the federal government to set aside the valley for

the education and enjoyment of the Nation as a whole.
Four years later, Congress created Grand Teton National
Vegetation of Jackson Hole and the Tetons
Park. Predictably, the park boundaries encompassed
Jackson Hole is surrounded by mountains that extend
most of the rugged and uninhabitable parts of the
above treeline. Fifty-eight percent of Grand Teton
Tetons but almost none of the valley lying east of the
National Park is nonforested (alpine tundra, boulder
mountain frontyet it was the valley that needed pro-
fields, meadows, grasslands, and shrublands), 28 per-
tection from development. John D. Rockefeller, Jr., took
cent is lodgepole pine forest, 7 percent Engelmann
tundra
Whitebark p.
Whitebark pine
Subalpine fir
Engelmann spruce
Lodgepole pine
Douglas-fir
Limber pine
Elevation
(feet)
Spruce-fir-pine
Lodgepole pine
Douglas-fir
Limber pine
sagebrush steppe
9000
8000
Fig. 15.8.Elevational distribution of

major forest types near Grand Teton
National Park. The solid vertical lines
show the range of elevations over which
each tree species is important as a forest
dominant; the vertical dotted lines indicate the range over which the species can
be found. Adapted from Baker (1986) and
Whitlock (1993).
7000
6000
5000
sprucesubalpine fir forest, 4 percent Douglas-fir wood-
common on the valley floor, especially on the glacial
land, and 1 percent aspen groves.31 In general, plant dis-
moraines where soil moisture is available late in sum-
tribution is determined by elevation, topography, and
mer (fig. 15.9). 33 Also, temperature inversions caused by
geologic substrate (fig. 15.8).
cold-air drainage may cause the valley bottoms to be
Douglas-fir and lodgepole pine are the most common
cool enough for lodgepole pine.
trees at low elevations in the Jackson Hole area, typically
Aspen groves are found in upland areas that are rel-
occurring together on Blacktail Butte and in the Gros
atively moist, such as at lower elevations where snow
Ventre Mountains to the east, where soils are deep and
accumulates or where there is groundwater seepage
fertile enough to support tree growth. Five Douglas-fir
to the surface. Frequently the groves are interspersed
habitat types have been identified, with the distinguish-
in sagebrush-dominated shrublands. Douglas-fir is
ing understory species being ninebark and Rocky Moun-
a common associate. 34 Mixed foothill shrublands or
tain maple on comparatively moist sites, and mountain
Douglas-fir grow on drier sites (see chapter 10). Sage-
snowberry, pine reedgrass, and spiraea on the drier sites.32
brush-dominated shrublands occupy the floor of Jack-
Lodgepole pine is common on some mountain slopes,
son Hole, except on glacial moraines, where conifer
such as Signal Mountain, as well as the west slopes of
forests are common (see fig. 15.9). Rivers and streams
the Tetons and the mountains to the north and east of
are fringed by riparian woodlands or shrublands (see
Jackson Hole, but it is infrequent on the east face of the
chapter 4). When overlooking the Snake River from
Tetons. At higher elevations on the Tetons, Douglas-fir
Antelope Flats, ancient river terraces are conspicuous
forests are replaced by subalpine forests dominated by
(see fig. 15.3), with coniferous woodlands on the steep
Engelmann spruce and subalpine fir. Whitebark pine is
slopes of cut banks and willow-dominated point bars
common above 8,000 feet, especially on dry windy sites,
on the opposite shore. Extensive willow shrublands also
and extends to upper treeline. Avalanche tracks on the
occur near Jackson Lake, such as the Willow Flats area
steep-walled canyons of the Tetons are obvious to hikers,
southwest of Jackson Lake Lodge.
extending from above treeline into the forested valleys
Much of the Teton range rises above treeline. The
and across streambeds, sometimes reaching a short dis-
steeper slopes appear barren, supporting only lichens
tance up on the opposite slope (see chapter 14).
and herbaceous plants in crevices. Alpine meadows and
In some areas Douglas-fir extends to higher eleva-
fellfields are found where soil has developed, such as
tions than does lodgepole pine, such as around Jackson
around Solitude and Amphitheater lakes. In parts of
Lake. Lodgepole usually occurs above Douglas-fir in
the Gros Ventre Mountains to the east of Jackson Hole,
the Rocky Mountains, but in Jackson Hole the pine is
sparse plant growth is seen at lower elevations, appar-
253
Fig. 15.9.This view of Jackson Hole from the top of Signal

Mountain shows forests of lodgepole pine and subalpine fir
on glacial moraines, and shrublands dominated by mountain
big sagebrush in the foreground and on the glacial outwash

plains. The Gros Ventre Mountains rise to the east beyond the
Snake River. See also fig. 2.6.
ently because of the shallow, infertile, often reddish
suggests relatively deep soils having considerable water-
soils that have developed there on Cretaceous shales.
holding capacity. The second type of community, found

farther south near the Jackson airport and Blacktail
Sagebrush Mosaic
Most of Jackson Hole is an unusually flat sagebrush
steppe, interrupted only by the Snake River valley and
a few buttes and moraines. Geologists refer to the area
as a glacial outwash plain, because it was deposited by
the meltwaters of the mountain glaciers that developed
80,000 to 12,000 years ago. One of the primary effects
of the meltwater was to wash away much of the silt and
clay that would have greatly enhanced the nutrient and
water storage capacity of the soil. Subsequently, however, this loss was compensated for by windblown loess
deposits from the west, such as from the Snake River
plains of present-day Idaho.
Close examination of the sagebrush steppe indicates
that at least three types of plant communities are represented. East of the Snake River, there is a dense cover
Butte, is characterized by an association of mountain

big sagebrush and bitterbrush. The abundant bitterbrush suggests a higher percentage of sand or gravel in
the soil. The third type of community can be seen west
of the Snake River in the southern part of the valley
and north of Moran. Here the sagebrush community
forms an intriguing mosaic, with patches of low sagebrush here and there where mountain big sagebrush
predominates (fig. 15.10). Low sagebrush is known to be
an indicator of shallow soils, sometimes with impeded
drainage or low fertility. Overall, the sagebrush mosaic
probably reflects different glacial outwash patterns as
well as differences in the rock types found in the mountains to the west and east of the valley. 35
Gros Ventre Landslide
of mountain big sagebrush, Idaho fescue, and other
The mountain ranges to the east of Jackson Hole have
species. The abundance of big sagebrush in this area
diverse originsthe volcanic Absarokas to the north-
The forests and substrate slipped about 1,000 feet,

with many trees coming to rest at an angle. Only the
smaller trees survived, usually those less than 40 years
oldprobably because many of them were flexible
enough and light enough to avoid significant breakage.
Seedlings of subalpine fir, limber pine, aspen, cottonwood, and other trees have since become established
among the surviving Douglas-fir, Engelmann spruce,
lodgepole pine, and aspen. In contrast, the scar left by
the slide remains largely unvegetated.
Fears that the newly formed dam would break discouraged tourism. Indeed, 2 years later the upper 60
feet of the dam did fail, creating a flood of water, rock,
and mud that claimed the lives of six people. The
Fig. 15.11.The Gros Ventre landslide created Lower Slide Lake

when it dammed the Gros Ventre River on June 23, 1925. The
slide is more than a mile long and half a mile wide. The forests are dominated by lodgepole pine at lower elevations and
have lodgepole pine, aspen, Douglas-fir, Engelmann spruce,
and subalpine fir at higher elevations.
Fig. 15.10.East of the Snake River and north of Moran,

patches of mountain big sagebrush are found dispersed in a
shrubland of low sagebrush. Lodgepole pine and subalpine fir
dominate the adjacent forests on glacial moraines. Elevation
6,600 feet.
east, the gravel conglomerates of the Mount Leidy

Highlands, the sedimentary strata of the Gros Ventre
Mountains to the east, and the granite and gneiss of
Jackson Peak to the southeast. All are much older than
the Tetons. In the Gros Ventre Mountains, fine-textured
soils have developed that are highly susceptible to slippage when saturated. Landslides and mudflows are
common in this range. In fact, the largest slide in U.S.
history occurred there on June 23, 1925 (fig. 15.11).36
Despite a dense cover of coniferous forest at the time, a
block of earth a mile long, half a mile wide, and up to
300 feet deep slipped down the mountainside, crossed
the Gros Ventre River, and created a dam 900 feet long
and more than 200 feet high. Lower Slide Lake was created and now is about 3 miles long; ranch buildings
were inundated. All this occurred in about 2 minutes.
255
town of Kelly was swept away, and a 20-foot wall of
author was A. Starker Leopold, son of Aldo Leopold
water hit Wilson 2 hours later, 25 miles downstream.
the inherent dynamics of natural ecosystems were
A new alluvial fan was created in the Kelly area, but
explicitly recognized. 39 Earlier assumptions about the
the water level was back to normal the next day. Kelly
desirability of stable population sizes and vegetation
has been rebuilt at the same location, and new shrub-
condition were challenged. Instead of suppressing dis-
lands and riparian vegetation have developed over the
turbances, the parks were advised to accept and even
past 85 years. A trip to the Gros Ventre Slide provides a
encourage change driven by herbivory, predation, and
reminder of what could happen again, and is another
fire. This new natural regulation policy called for let-
illustration of how the geology of an area influences
ting such ecological processes operate to the greatest
plant distribution.
degree possible, with human intervention allowed as

needed for safety or to correct problems caused by pre-
Management Issues in the GYE
vious human activities. More research was also recommended, recognizing the many uncertainties about
Management concerns and priorities in the GYE have
whether specific ecological changes in the parks were
changed dramatically since YNP was established in
driven by natural phenomena or human interference.
1872.37 Initially, the objectives for YNP seemed simple
There also were debates about the long-term effects of
and straightforward: protect the geothermal features
Native Americans.40
from vandalism, protect the wildlife from predators
Whether natural regulation can really be an effective
and poaching, and protect the vegetation from fire
management approach in any ecosystem is still a topic
and overgrazing. Do these things and the park would
of considerable discussion. Consider, for example, the
be preserved. All fires, whether ignited by lightning or
challenges associated with the Yellowstone elk prob-
humans, were actively suppressed when possible; preda-
lem, the 1988 fires, the decline of whitebark pine, and
tors were shot; and elk and bison were managed inten-
the reintroduction of wolves.
sively, as though they were domestic livestock.

By the 1960s, however, it had become increasingly
clear that this approach to park preservation was overly
The Elk Problem
simplistic and ineffective. With much less predation,
By the late 1800s, unregulated hunting of elk and other
the elk population increased rapidly and appeared to
wild game drastically reduced herds throughout the
threaten the persistence of aspen, willows, and grass-
West.41 To counter this trend, animals were fed and
lands on their winter range. Fire-dependent species
predators were systematically eliminated, especially
were declining, and the iconic grizzly bear was on the
wolves. By 1912 the summer elk population in YNP
verge of local extinction. Throughout the parks his-
was crudely estimated at more than 30,000.42 Concern
tory some scientists had argued that the greatest value
shifted from fears of elk extirpation to worries that elk
of Yellowstone lay in preservation of its wild nature,
were damaging their winter range. In response, manag-
and had questioned programs of intensive control and
ers began trapping and shipping excess animals to other
manipulation of the parks ecosystem. For example,
places. When the demand for such animals declined,
Theodore Comstock celebrated the park as one of the
rangers reduced herd sizes by shooting the animals,
few remaining places where one could study the evo-
even inside the park.43 Still, elk numbers remained high,
lution and natural selection of animals in a state of
and concerns began to mount about overgrazing on the
nature, and George Wright and Aldo Leopold famously
northern winter range, located between Mammoth and
advocated protection and restoration of predators in
Cooke City. For example, ecologist Charles Kay com-
Yellowstone and other national parks. 38 Nevert heless,
pared historic and modern photographs, noting an
major changes in management philosophy were not
abundance of tall willows and a lack of browsing-caused
made until the 1960s, as proposed in a 1963 report
scars on aspen bark in the 1800s, but few willows and
titled Wildlife management in the national parks.
almost ubiquitous scars on aspen bark in modern views
Known as the Leopold reportbecause the senior
of the same locations (fig. 15.12).44
every spring and fall between summer and winter

ranges. The summer ranges usually are at higher elevations, where plant productivity and forage availability
are greater because of the cooler, wetter environment.
Winter ranges are in the low-elevation foothills and
valleys (see chapter 10), where snow accumulation is
less and animals can forage until spring. However, the
relatively low productivity of winter range vegetation,
coupled with concentrations of animals for periods of
several months, leads to heavy use of forage plants
including shrubs and trees. Browse lines developed on
junipers, willows, and other tall shrubs (see fig. 10.16),
mature aspen were scarred after elk ate some of the bark,
and most aspen sprouts were snipped off before they
could grow into new trees (see figs. 10.14 and 10.15).
Recognizing that conversion of land to agriculture
and towns was reducing the availability of critical winter range for elk and deer throughout much of the West,
federal and state agencies initiated winter feeding programs to help maintain ungulate populations. In 1912
the National Elk Refuge in Jackson Hole was established
on the fertile bottomlands along Flat Creek. Wintering
elk began to congregate there by the thousands every
winter, as they do today.47 The Wyoming Game and
Fig. 15.12.The scars on these aspen were caused by elk feeding on the bark, a common occurrence on trees near winter
ranges.
Fish Department developed similar feeding programs

elsewhere. From the beginning, there has been controversy over the wisdom of this practice. The benefits are
perceived to be fewer agricultural conflicts, larger herds
for hunters, and less discomfort about animals starving
With the loss of tall willows came problems for
during harsh winters. However, the problems include
associated wildlife, including loss of nesting habitat
cost, injury of trees and other forage plants in the vicin-
for neotropical migrant songbirds.45 Competition with
ity of feeding grounds, disease transmission to livestock
elk for food was thought to be affecting populations
(such as brucellosis), and fostering the public perception
of deer and beaver. Some critics argued that few or no
that suitable habitat is not important.48
elk had occupied the winter range inside YNP until the
Arrival of the Leopold report in the 1960s led to
late-1800s, when land use changes north of the park
a fundamental rethinking of YNPs management
boundary restricted the animals to the park. To test this
approach. Some scientists argued that YNPs winter
hypothesis, historians Paul Schullery and Lee Whittle-
range was not overgrazed, pointing out that native
sey reviewed all observations made by explorers before
rangelands remained basically intact and continued
1882. They concluded that the record is not sufficiently
to sustain impressively large numbers of elk and other
detailed to know in any given year whether the elk
wildlife. Beginning in 1968, YNP terminated culling
population equaled, exceeded, or was less than at pres-
elk herds in the park, andas an experimentbegan
ent, but they became convinced that the park area was a
observing whether the Yellowstone elk population was
winter range for thousands of elk before 1882.46
capable of self-regulation. The park also implemented a
These concerns were not unique to YNP. In the GYE

and throughout much of the West, elk and deer migrate
research program to better understand the dynamics of

elk and their habitat.
257
Critics immediately challenged the natural regula-
mer ranges, which are more extensive than their winter
tion approach to managing the Yellowstone elk.49 They
ranges. These seasonal movements minimize grazing in
contended that YNP did not contain the key winter hab-
any particular area.54 Park biologist Douglas Houston
itat necessary to sustain a self-regulating elk population.
concluded in 1982, after 10 years of research, that reduc-
In winters since time immemorial, they argued, the
tions in the size of the elk herd were not necessary. 55
Yellowstone elk had migrated out of the relatively high-
In contrast to the grasslands, aspen and woody ripar-
elevation YNP to reach snow-free grasslands at lower
ian plants have continued to be significantly affected
elevations outside the current park boundary. Much of
by herbivores. 56 Approximately 85 percent of tree-sized
this area was now private land, converted to agriculture,
aspen on the northern winter range became established
roads, and other developments that impaired the qual-
prior to 1920; these old trees are gradually dying and
ity of the habitat for elk.
Hunting pressure in these
generally are not being replaced by young aspen sprouts
areas also reduced elk numbers significantly in severe
and seedlings.57 Willows and associated woody riparian
winters, and forced many of the elk to remain in the
vegetation in many areas appear degraded, with low
park during mild winters. Critics argued that regulation
stature, many dead branches, and new growth confined
of the elk population could never be truly natural, and
to the base of the plant rather than the top; seedlings
that unnatural overuse of the vegetation inside the park
of willow and cottonwood are often observed, but few
was unavoidable. To address this issue of elk migrating
survive.58 The National Academy of Sciences committee
out of the park during severe winters, wildlife agencies
evaluated several alternative hypotheses to explain the
and conservation groups eventually began to purchase,
lack of widespread aspen and willow regeneration dur-
or protect in other ways, some of the key elk wintering
ing the twentieth century. The committee determined
areas along YNPs boundaries.
that neither twentieth-century climate conditions nor
50
Notably, the assumption that wintering elk were
fire suppression were sufficient explanations in them-
irreparably damaging the vegetation of YNPs northern
selves for the paucity of recent aspen and willow regen-
range has not stood up to research findings. In 1998 the
eration and concluded that ungulate browsing was
U.S. Congress directed the National Academy of Sciences
the primary mechanism.59 Nevertheless, even though
to review all available science related to the manage
browsing is suppressing aspen and willows, the panel
ment of ungulates and the effects of ungulates on the
of scientists concluded that none of these species was in
vegetation of YNPs northern winter range, to clarify
imminent danger of extirpation on the winter range.60
what is known and what is not known. A committee of
Following cessation of artificial regulation in 1968, the
twelve eminent scholars concluded that the condition
northern Yellowstone elk population increased rapidly,
of the northern range is indeed different today than
growing from fewer than 4,000 animals in 1968 to more
when Europeans first arrived in the region, but that the
than 19,000 by 1988 (fig. 15.13).61 The controversy over
51
ecosystem is not in imminent danger of losing major
elk management in YNP and elsewhere did not subside,
components or crossing any threshold that would lead
but the plot thickened. In 1988 the largest fires in more
to irreversible changes. The committee determined that
than a century burned approximately a third of YNP,
grasslands on the northern winter range generally show
including a large portion of the northern winter range
less dramatic impacts of ungulate use than do aspen for-
and surrounding lands. With ongoing climate change,
ests and willow communities.52 Despite the consump-
the habitat has changed because of drought, a widespread
tion of 45 percent of the forage biomass by elk, the plant
epidemic of bark beetles, and the decline of whitebark
growth in grazed areas of YNPs northern winter range
pine due to an exotic disease. And then, in 1995, the gray
was 3685 percent higher than in exclosures, suggest-
wolf was reintroduceda major predator of elk. The so-
ing compensatory growth53 (see chapter 6).
called elk problem now became part of the broader chal-
One reason heavy grazing has had such minimal

adverse effects on the herbaceous vegetation of the win-
lenge of managing fire, forests, and wildlife as a whole.
ter range is that most foraging occurs during fall and
Fire Management
winter, when the plants are dormant. In spring and
Systematic efforts to eliminate wildfires were initiated
summer, elk and bison move to higher-elevation sum-
in YNP with the arrival of the U.S. Army in 1886,62
NUMBER OF ELK (thousands)
25
Northern Yellowstone Elk Herd
20
15
1988
Fires
10
5
1970
1975
1980
1985
1995
Wolf
reintroduction
1990
YEAR
1995
2000
2005
2010
Fig. 15.13. Population trends in the northern Yellowstone

elk herd from 1971 to 2010. Beginning in 1968, the park
ended its program of artificially controlling elk numbers.
For that reasonand because of a series of moderate winters
and low levels of predationthe population grew. Large fires
burned approximately a third of the park in 1988, including
much of the northern winter range, and wolves were reintroduced to the park in 1995 after nearly a century of absence.
The recent decline in elk numbers is due to several factors, as

discussed in the text. Three sets of data are shown (black and
red points): Schullery and Whittlesey (1992) for 19711992
(black); Vucetich et al. (2005) for 19852004 (red), and the
Yellowstone Center for Resources (unpublished) for 2005
2010 (red). The data overlap in the period 19861992 and
differ slightly, reflecting the difficulty in censusing wildlife
populations.
but they were generally ineffective in remote forested
mented in surrounding wilderness areas and in Grand
areas. By the time a fire was reported and a crew found
Teton National Park. However, in 1988 researchers and
it, often days later, the fire commonly had either gone
managers learned that 15 years was not long enough to
out or had become so intense that it could not be extin-
fully understand the ecology of fire in the GYE.
guished with hand tools. Only with the availability of
The summer of 1988 turned out to be one of the
new fire-fighting equipment after World War II, notably
driest and windiest on record. Some 248 fires were
aircraft and smokejumpers, were managers able to sup-
ignited in the GYE by lightning or humans and burned
press most ignitions.
approximately 1.4 million acres (fig. 15.14). A massive
In the wake of the 1963 Leopold report, researchers
fire-fighting effort25,000 fire-fighters and $120 mil-
began reporting that fire was an important ecological
lionsucceeded in minimizing loss of life and property
process in most western forests and rangelands and
but could not stop the spread of fires across the land-
that fire exclusion was leading to undesirable ecologi-
scape until significant precipitation finally arrived in
cal changes. Therefore in 1972in the spirit of natural
mid-September.64
regulationpark managers initiated a cautious pro-
Park managers convened a panel of experts to eval-
gram of allowing lightning-caused fires to burn with-
uate the ecological impacts of the fires and to predict
out interference in a remote portion of the park during
likely ecosystem responses and management implica-
moderate weather conditions, when the risk of human
tions.65 The panel concluded that the size and severity
injury or serious resource damage was minimal. In 1976
of the fires were influenced more by drought and wind
this natural fire program was expanded to encompass
than by fuels or previous management practices. They
most of the parks backcountry area. During a period of
noted that large and severe fires also had occurred in
15 years, from 1972 through 1987, a total of 235 light-
the early 1700s (fig. 15.15).66 Subsequent research has
ning-ignited fires were allowed to burn, of which only
shown that large fires have recurred in Yellowstone at
27 burned more than an acre; the largest burned 7,400
intervals of decades or centuries throughout most of
acres.63 YNPs natural fire program seemed to be suc-
the Holocene.67 The 1988 fires were not confined to the
cessful, and similar natural fire programs were imple-
park but burned substantial portions of surrounding
259

Fig. 15.14.Extent and severity of the
1988 fires in Yellowstone National Park.
Burn severity was calculated using
data from pre- and post-fire Landsat
Thermal Mapper images and ground
observations. Especially striking is the
heterogeneity of burn severity. Similar
heterogeneity probably characterizes all
large forest fires. Cartography by Brian
J. Harvey. Reprinted with permission
from Romme et al. (2011, fig. 1) and
Springer Science and Business Media.
national forests and private lands, destroying some tim-
were too close and too flammable, they argued that
ber resources and threatening human life and property.
manager-ignited fires, during moderate weather con-
Even at 2.2 million acres, YNP was not large enough
ditions and especially near human habitations, could
to contain a natural fire regime. Moreover, the North
have minimized the anxiety of 1988. Analyses by For-
Fork Fire, one of the largest individual fires of 1988, was
est Service ecologist James Brown substantiated this
ignited by humans outside YNP and spread eastward
claim, suggesting that the cost of fuel reduction near
into the park, burning about 490,000 acres.68
developments would have been reasonable compared
Just as with elk management, the 1988 fires focused
to the cost of fighting the 1988 wildfires.69 However,
attention on the question of whether natural regula-
the same analysis led to the conclusion that the scale
tion can really be an effective management approach,
of the 1988 fires would not have been reduced signifi-
or whether a more active program is called for, even in
cantly had there been a program of prescribed burns in
the largest parks and wilderness areas. For some observ-
YNP. Human-caused prescribed fires would have been
ers, the need for manager-ignited prescribed fires was
set only under moderate fire-weather conditions, with
eminently clear. Noting that gateway communities and
the result that only small areas would have burned. The
visitor centers were threatened because adjacent forests
resources needed to contain such fires if weather condi-
Complete
Suppression
Policy
Natural
Fire
Policy
LP3
40
% Area
in Each 30
Cover 20
Type
10
LP2
LP1
LP0
% Area 40
Burned 20
1700 20
40
60
80
1800
Fig. 15.15. Percentage area burned per decade from the

1690s through 1980s in a 450-square-mile landscape in central Yellowstone National Park, and percentage area covered
by four post-fire stages of forest development during that
time. LP0 is recently burned forest (typically 040 years
old), LP1 is young lodgepole pine forest (40150 years old),
20
40
60
80
1900
20
40
60
80
LP2 is mature lodgepole pine forest (150250 years old),

and LP3 is old-growth lodgepole pine forest, mixed with
spruce and fir on more fertile sites (250 or more years old).
Drafted by Michael Turner. Reprinted with permission of
M. G. Turner et al. (2001, fig. 7.15) and Springer Science and
Business Media.
tions and fire intensity had changed quickly would not
Natural fire policies in YNP and other national parks
have been sufficient to deal with more than a few fires
and national forests across the West were suspended
each year, and not every year would have had accept-
after the 1988 fires, while another review team evalu-
able weather conditions. Thus, assuming that human-
ated the scientific foundation of fire management.
caused prescribed fires had been used after 1972, when
The team concluded that YNPs natural fire policy was
fire was officially accepted as a necessary process in
basically sound but needed more specific criteria for
YNP, too little of the forest away from human habita-
deciding whether to suppress or to only monitor spe-
tions would have been burned to prevent the large-scale
cific fire ignitions. Yellowstone managers incorporated
1988 fires.
these changes in a new fire management plan in 1992,
Even if the 1988 fires cannot be viewed as entirely
and Grand Teton National Park and national forest
natural in size or spatial pattern, the ecosystem has
managers in the GYE developed similar modified poli-
shown great resilience, with surprisingly rapid natural
cies. Lightning-ignited fires are again allowed to burn
recovery of almost all components.70 There were some
in the GYE backcountry if they do not threaten life,
exciting ecological surprises as well: a large cohort of
property, or significant resourcesand if weather and
aspen seedlings appeared in burned areas the first year
fire-fighting resources are deemed such that another
after the fire (fig. 15.16). Genetic analyses confirmed
1988-like event is highly unlikely.
that the plants were genuine seedlings, not root sprouts,
But the Yellowstone fire story may have only just
Moreover, no aspen
begun. The 1988 Yellowstone fires were among the first
seedlings were observed in unburned areas, suggesting
of what has proven to be a dramatic increase in large
that successful sexual reproduction of this charismatic
forest fires across the American West since the mid-
Rocky Mountain tree may occur only in conjunction
1980s.73 The increase has been driven primarily by cli-
with severe fires. Another surprise was that, though the
mate changein particular, the trend of earlier spring
media sometimes gave the impression that YNP had
snowmelt, warmer summers, and a longer period each
been destroyed, visitation increased in the years after
year in which wildfires can burn. Anthony Westerling
the 1988 fires.72
and colleagues found a statistical correlation between
as are commonly seen in aspen.
71
261
fire rotation probably could not occur. Nevertheless, the

important message of this study is that fire will almost
certainly become more frequent in the future. With a
substantially shorter fire rotation, the older forests that
now characterize much of the GYE could not persist in
their present form. Moreover, post-fire vegetation development could be very different from the smooth recovery that occurred after 1988 if the climate is no longer
suitable for some of the major plant species.75 Westerling and his associates emphasized that there are many
uncertainties, but substantial changes in vegetation patterns appear likely.
Beetles, Blister Rust, Whitebark Pine, and Grizzlies

Whitebark pine is common at the highest elevations in
the GYE and Northern Rockies, where it stabilizes soils
and provides abundant seeds that are an important
food for grizzly bears and other wildlife. Recent fires
have burned whitebark pine forests, but this species is
capable of recovering from fire via seeds transported
and planted by Clarks nutcrackers.76 What the pine
may not recover from are the combined effects of global
climate change, mountain pine beetles, and a nonFig. 15.16.The small aspen in the foreground of this photo,
taken near Old Faithful, became established from seed after
the 1988 Yellowstone fires. These young aspen have been
heavily browsed by ungulates, but some have established new
clones in the area. The lodgepole pine in this photo are about
20 years old. The downed wood is from the trees killed by the
1988 fires.
native invasive pathogenthe white pine blister rust

(see chapter 14). Warming has allowed mountain pine
beetles to expand upward into whitebark pine forests
(see chapter 11 for beetle biology). Only recently have
entomologists learned that this pine lacks most of the
chemical defenses against bark beetles that are found
in lodgepole pine, most likely because the cold climate
large fires and warm summers in the Rockies from 1972
in whitebark pines high-elevation habitat formerly lim-
to 1999; they then applied this relationship to predict
ited bark beetle development and survival. When out-
fires for the next century, using four different global
breaks did occur in the past, they were infrequent and
climate change models.74 The results were startling: by
brief. However, the warm temperatures of the past 20
about 2050, large fires could be occurring almost every
years have allowed mountain pine beetles to success-
year, and the fire rotation in most of the GYE could be
fully attack whitebark pine over large areas; mortality of
reduced from the historical rotation of 120300 years
mature trees has been high.77
to less than 20 years. Fire rotation is a standard measure
The beetles generally have not killed the smaller
of fire frequency and is defined as the number of years
whitebark pine trees because, instinctively, the insects
required for cumulative area burned to equal the size
fly toward the larger trees that have adequate phloem
of the entire area of interest. This modeling procedure
to sustain a brood of beetle larvae. Ordinarily one
did not include changes in fuels; it assumed that fuels
would expect whitebark pine forests to recover from
would always be sufficient to permit burning even at
the beetle outbreak through growth of the smaller sur-
very short intervals. However, fires would begin to be
vivors. However, the white pine blister rust, a fungus
limited by fuel availability as intervals between fires
native to Eurasia that was accidentally introduced to
became ever shorter; the researchers noted that a 20-year
North America around 1900, kills whitebark pine trees

Fig. 15.17.The grizzly bear, an omnivore once common across the Great
Plains and throughout the Rocky Mountains, is now found only in the Greater
Yellowstone Ecosystem and northward.
Photo by Mark Gocke.
of all sizes, large and small (see fig. 14.5).78 In some
ignated a threatened species in 1975. Whitebark pine
areas, blister rust infection appears to make the larger
seeds had been an important late summer and autumn
trees more vulnerable to mountain pine beetle attack,
food for grizzly bears, contributing to the accumulation
probably by reducing their ability to produce sufficient
of fat reserves necessary for winter survival. Because
amounts of resin.79 Thus, a warming climate not only
individual trees produce large amounts of seed only
leads to increased burning of high-elevation forests, but
every 35 years, it was important that the bears have
it also permits native bark beetles to move into these
large numbers of trees over large areas, so that some
previously unsuitable areas. Meanwhile, the exotic blis-
seed is available each year.
ter rust kills many of the small survivorsall leading
In addition to a declining number of whitebark pine
to a grim prognosis for whitebark pine in the northern
seeds, grizzly bears have also lost another food source,
Rockies.80 Efforts are under way to locate and propa-
the native cutthroat trout in Yellowstone Lake. The cut-
gate blister rustresistant individuals, with the intent
throat trout are threatened by two other non

-native
of restoring whitebark pine woodlands.81 However,
specieslake trout and a microscopic myxozoan that
given the spatial extent and rapidity of whitebark pine
causes whirling disease. Lake trout were illegally intro-
decline, many timberline forests of the GYE and other
duced to the lake, apparently in the mid-1980s, and
parts of the Northern Rockies may become what ecolo-
they have proven to be voracious predators of the native
gist Jesse Logan refers to as ghost forests.82
cutthroat. Whirling disease, first recorded in YNP in
Extensive loss of whitebark pine has had a cascad-
1998, leads to abnormal growth of a fishs bony struc-
ing impact on other ecosystem components, notably,
tures, causing an inability to feed.83 Cutthroat trout
grizzly bears (fig. 15.17). Now rare in the contiguous
populations have plummeted, and spawning runs up
United States and restricted to remote areas, the grizzly
tributary streamswhere the bears formerly harvested
was once common throughout the Rocky Mountains
great numbers of trout every summerhave dwindled.
and across the Great Plains. Its current range is prob-
Unlike the cutthroat, the lake trout do not spawn in
ably less than 1 percent of what it was in the 1800s.
rivers and, in deep water, they are inaccessible to bears,
With fewer than 1,000 animals in Montana, Wyoming,
ospreys, pelicans, and other native wildlife that had fed
Idaho, and Washington combined, the grizzly was des-
on the cutthroat.84 Park managers have implemented
263
an aggressive program of lake trout control, but at best,
matically from approximately 19,000 animals in 1994
the program manages merely to hold down the lake
to only about 6,000 in 2010 (see fig. 15.13). However,
trout population: eradication of this non-native invader
a recent assessment determined that wolf predation by
appears unlikely.85 Adding further to the bears troubles,
itself was not sufficient to account for the drop in elk
a third important food source for grizzlies has been
numbers. Contributing to the decline were increased
diminished since the reintroduction of wolves in 1995:
predation of elk calves by grizzly bears, as described
the bears formerly fed extensively on winter-killed elk
previously, along with drought and elk hunting outside
and bison when they emerged from their dens in the
the park.95 The wolves preyed mainly on elk, and occa-
spring, but now the wolves are consuming many of
sionally bison, deer, pronghorn, and moose, but hun-
those carcasses.86
dreds of cattle and sheep were taken as well (though
Surprisingly, despite the decline of three key food
less than 3 percent of livestock losses in the area can
resources, grizzly numbers in the GYE have increased
be attributed to predation). Compensation programs for
over the past three decades.87 The bears have proved
livestock losses were initiated by a nongovernmental
themselves to be genuine opportunists, shifting to
organization, Defenders of Wildlife, and then adopted
alternative resources when a preferred food source is
to varying degrees by state game and fish departments.
in short supply. The grizzlies are now killing more elk
The costs of conserving biological diversity can be sub-
calves than they did before the decline of cutthroat
stantial when large predators are involved, but the ben-
trout, thereby obtaining the protein and fat they need.88
efits are also obvious. For example, wolves and grizzlies
The bears also feed on berries, succulent plants, fleshy
are a major tourist attraction.96
roots, insects, and mushrooms when they are avail-
There also are thought to be ecological benefits. Fol-
able.89 Yellowstones grizzlies appear to be coping so far
lowing the sharp decline in elk numbers on Yellow-
with the effects of climate change and non-native spe-
stones northern range, ecologists William Ripple and
cies, although there surely are limits to what they can
Robert Beschta proposed that the associated reduction
endure.
in browsing pressure had allowed sprouts of aspen, willows, and other woody riparian plants to grow tall again
after nearly a century of ungulate-caused suppression.97
Wolves, Elk, and Aspen
Their suggested mechanism involved not just the reduc-
By the early twentieth century, the wolf was function-
tion in elk numbers but also a change in elk behavior.
ally extinct across most of the West, and it was one
With wolves, they postulated, the elk moved around
of the first species to be protected after passage of the
more often, avoiding habitats where they would be
Endangered Species Act in 1973. The wisdom of preda-
more vulnerable to wolf predation. According to Ripple
tor eradication in YNP had been questioned as early as
and Beschtas hypothesis, such movements might be
the 1930s, and the possible role of the wolf in control-
in response to detection of a wolf in the distance. In
ling excess numbers of elk was an issue in the 1960s.91
both cases, the result would be less time spent browsing
Wolf reintroduction gained considerable public support
in any particular place, allowing more sprouts to grow
at that time, but it was not until 1995 that 14 wolves
tall. Though an intriguing concept, other research has
were captured in Alberta and released into YNPs north-
not supported this behavioral hypothesis.98 Many aspen
ern winter range.92 In 9 years the population grew to 170
sprouts on the northern range still have not elongated
wolves in 15 packs distributed throughout the GYE. By
into new tree-sized stems, except within fenced exclo-
2012 there were several hundred in the area (about 100
sures where elk could not reach them.99
90
93
in YNP).
Unlike aspen, willows have become conspicuously
Critics worried that wolves would reduce elk herds to
taller since about 2000 in some well-watered places,
the point of impairing big-game hunting opportunities,
such as along the Lamar River near Soda Butte Creek,
and that they would prey heavily on livestock outside
perhaps due partially to the presence of wolves. How-
the park.94 The elk population did, in fact, decline. For
ever, increased willow growth has not been observed
example, the northern Yellowstone elk herd fell dra-
everywhere. Colorado State University scientists Kristen
Marshall, Thomas Hobbs, and David Cooper concluded
underscored the importance of conservation efforts on
that a lack of willow and aspen food sources in the early
private lands in and surrounding the GYE. For example,
twentieth century led to local extirpation of beaver along
ecologists Andrew Hansen and Jay Rotella found that
some small streams, where beaver dams formerly main-
hotspots for bird diversity and abundance tend to occur
tained high water tables. Subsequent down-cutting in the
not in the high-elevation wilderness areas, but on low-
absence of beaver caused a lowering of the water table,
elevation private lands that are increasingly vulnerable
resulting in habitats that were too dry for willows, and
to degradation.102 In fact, this seems to be true for the
now, they reasoned, the willows in such places do not
region as a whole.
grow well even when fenced to exclude elk browsing.100
Given the natural complexity of this ecosystem
As another example of cascading ecological effects,
coupled with issues of boundary effects, land use
wolf reintroduction has led to a decline in the number
changes, invasive species, and climate changeit is
of coyotes in Jackson Hole, which in turn has resulted
difficult to know what management actions should be
in greater survival of pronghorn fawns.101 The value of
adopted, and to what degree natural processes are suf-
large wildlands for understanding ecological processes
ficient for maintaining the GYE.103 Managers are still
is again eminently clear.
learning about the effects of their decisions, and often

the lessons are unexpected. Historian Paul Schullery
On some days, the GYE is tranquil with elk, bison,
wrote, one of the most important contributions science
moose, and pronghorn at rest in expansive meadows.
has made to management dialogues in the past 30 years
The rivers are clean, and the thermal features are fasci-
is to elevate the admission of uncertainty as a credible
nating curiosities for resting hikers on a warm summer
management stance.104 Despite difficulties and con-
day. However, such tranquility belies the long winters
troversies, the Yellowstone country remains a national
that plants and animals must endure, the forest fires
treasure, indeed a global treasure. With the return of
that periodically burn across the landscape, and the
the gray wolf, the GYE again has all of the species and
molten rock that exists close to the earths surface. The
more or less the same ecological processes that existed
challenges of protecting such unique places, over such
when YNP was established in 1872. Equally important,
a large area, are formidable. Much research has focused
away from the roads, the GYE elicits a powerful sense of
on the GYEs public lands, but recent analyses have
wildness and beauty.
265
The Black Hills,

Bear Lodge Mountains,
and Devils Tower
Chapter 16
The mountains of western South Dakota and northeast-
hands, while the head-gear of the horses was deco-
ern Wyoming, known as the Black Hills and Bear Lodge
rated with wreaths of flowers fit to crown a queen of
Mountains, originated at approximately the same time
May. Deeming it a most fitting appellation, I named
as the mountains to the west.1 The highest point is Har-
this Floral Valley.
ney Peak, at 7,242 feet above sea level. Fifty million years
of erosion have stripped thousands of feet of sedimentary material, exposing the more erosion-resistant granitic core. Erosion occurred largely toward the east, the
direction in which the two major rivers flow, the Belle
Fourche and the Cheyenne (fig. 16.1). Devils Tower, an
unusual geologic feature, is commonly thought to have
been the neck of a volcano, though there is debate about
its origin (fig. 16.2).
Native Americans lived in the Black Hills when the
Vrendrye brothers, two French explorers, traveled
through the region in the mid-1700s. Most likely they
were the first Europeans to cross the northern Great
Plains and see the Rocky Mountains.2 More than a hundred years later, in 1874, Lt. Col. George A. Custer came
to the Hills from Fort Abraham Lincoln, near Bismarck,
North Dakota. Traveling with 1,200 troopers, 1,000
horses, and 110 wagons pulled by mules, plus 300 head
of cattle, his mission was to reconnoiter a route to the
Black Hills and explore their interior. For his journal
entry on July 25, Custer wrote:
Another member of his expedition observed, Everybody was making bouquets. . . . Some said they would
give a hundred dollars just to have their wives see the
floral richness for even one hour.3
The colorful flowers that Custer and his men enjoyed
most likely included anemone, blanketflower, bluebells,
blue-eyed grass, geranium, nodding onion, shooting star,
wild iris, and yellow owls clover.4 Custer also had his picture taken in the Black Hills with a dead grizzly bear.5
Congress designated much of the area as a forest reserve
in 1897, and today, most of the Black Hills and Bear
Lodge Mountains are in the Black Hills National Forest.
The Black Hills have been an important source of
gold over the years, with North Americas deepest and
largest underground gold mine located in Lead, South
Dakota. It closed in 2002 after 126 years of operation.
Many tourists toured the mine each summer. Visitors
continue to be attracted to the Hills by Devils Tower
National Monument, Jewel Cave National Monument,
Mount Rushmore National Memorial, Wind Cave
National Park, the Mammoth Site in Hot Springs, and
Every step of our march that day was amid flowers of
a summer climate that is cooler than that on the sur-
the most exquisite colors and perfume. So luxuriant
rounding plains. The region also is an important source
in growth were they that men plucked them without
of wood, water, and livestock forage.
dismounting from the saddle. . . . It was a strange
Botanically, one of the most interesting features
sight . . . the men with beautiful bouquets in their
of the area is the assemblage of plants found there
266
The Black Hills, Bear Lodge Mountains, and Devils Tower 267
Fig. 16.1. Geomorphic features of the Black Hills. The elevation ranges from 3,140 feet at Rapid City to 7,242 feet at the
top of Harney Peak, the highest point in South Dakota. On
the west side, the elevationa at Sundance and Newcastle are

4,750 and 4,334 feet, respectively. Adapted from Strahler
(1969).
(table 16.1).6 As might be expected, many Rocky
widespread about 10,000 years ago, when the climate
Mountain species occur in the area, such as elkweed,
was cooler and more humid. Today the Hills provide
grouse whortleberry, heartleaf arnica, narrowleaf cot-
an environment where many of these plants can still
tonwood, Oregon-grape, and ponderosa pine. Their
survive. Annual precipitation is considerably higher
presence is best explained by the generally accepted
than on the surrounding plains, ranging from 18 to 30
conclusion that Rocky Mountain plants were more
inches (fig. 16.3).
Fig. 16.2. Devils Tower, flanked by ponderosa pine savanna

and mixed-grass prairie. The grassland can be highly flammable. Common grasses include blue grama, western wheatgrass, needle-and-thread grass, and little bluestem. Fisher et al.
(1987) estimated that the historic fire-return interval in this
area was about 14 years. Devils Tower is one of a dozen igneous
intrusions in the Black Hills and Bear Lodge Mountains. The

others include Inyan Kara Mountain, Little Missouri Buttes,
and Warren Peaks in Wyoming, and Bear Butte, Black Butte,
Crow Peak, Custer Peak, and Terry Peak in South Dakota. Elevation at the top of Devils Tower is 5,117 feet. Photo by Hollis
Marriott.
More puzzling is the presence of plants that typi-
the somewhat cooler, wetter, largely forested environ-
cally are found in the deciduous forests of eastern
ment of the Hills, the biodiversity of the region as a
North America, such as American elm, bloodroot, box-
whole would be lower. Notably, the enabling legisla-
elder, bur oak, hackberry, and hophornbeam, or species
tion passed by Congress for establishing Wind Cave
from the boreal forests that stretch across Canada, such
National Park was to protect bison, elk, and pronghorn
as bunchberry dogwood, Canada scurvyberry, paper
as well as the cave itself. 8
birch, and white spruce. At least some of the eastern

species could have migrated westward along the moist
tributaries of the Cheyenne River, where they still occur
Geomorphic Regions
in favorable habitats.7 The northern species may have
The uplift and subsequent erosion of the Black Hills
migrated southward during the cooler periods associ-
and Bear Lodge Mountains have led to the formation
ated with the advance of continental glaciers. The Black
of five roughly concentric geomorphic regions that
Hills were not glaciated, but the boreal plants persisted
strongly influence landscape patterns (see fig. 16.1).9
a short distance south of the ice front where the climate
On the perimeter is the Hogback Rim, composed of
was favorable.
Mowry shale, Lakota sandstone, Fall River sandstone,
The Black Hills and Bear Lodge Mountains are refu-
Minnewasta limestone, and other erosion-resistant sed-
gia for many forest plants and animals that are sepa-
imentary strata that were tilted upward as the moun-
rated from their main populations by long distances
tains developed. The rim is sharply defined to the east
across grasslands. Conservation biologists place great
and south but is more like a plateau on the west. Pon-
value on such places, because they help maintain
derosa pine savannas predominate on rocky soils, with
genetic diversity. Without the habitat provided by
grasslands on fine-textured soils.
Table 16.1. Some plants in the Black Hills representative of other floristic regions
Region/Common name
Latin name
ROCKY MOUNTAIN SPECIES

Narrowleaf cottonwood
Populus angustifolia
Ponderosa pine
Pinus ponderosa
Creeping barberry (Oregon-grape)
Mahonia repens
Elkweed
Frasera speciosa
Heartleaf arnica
Arnica cordifolia
Grouse whortleberry
Vaccinium scoparium
Richardsons geranium
Geranium richardsonii
GREAT BASIN SPECIES

Alderleaf (true) mountain-mahogany
Cercocarpus montanus
Big sagebrush
Skunkbush sumac
Rhus trilobata
Blanketflower
Gaillardia aristata
Bottlebrush squirreltail
Elymus elymoides
Prairie junegrass
Koeleria macrantha
Threadleaf sedge
Carex filifolia
EASTERN DECIDUOUS FOREST SPECIES

American elm
Ulmus americana
Boxelder
Acer negundo
Bur oak
Quercus macrocarpa
Common hackberry
Celtis occidentalis
Green ash
Fraxinus pennsylvanica
Hophornbeam
Ostrya virginiana
Bloodroot
Sanguinaria canadensis
Downy yellow violet
Viola pubescens
Red columbine
Aquilegia canadensis
Virginia creeper
Parthenocissus quinquefolia
BOREAL FOREST SPECIES

Paper birch
Betula papyrifera
White spruce
Picea glauca
Bunchberry dogwood
Cornus canadensis
Canada mayflower
Maianthemum canadense
Single delight (woodnymph)
Moneses uniflora
Twinflower
Linnaea borealis ssp. americana
SOUTHERN GREAT PLAINS SPECIES

Bush morning-glory
Ipomoea leptophylla
Crested pricklypoppy
Argemone polyanthemos
Fendler (purple) threeawn
Aristida purpurea var. longiseta
Sand sagebrush
Artemisia filifolia
Velvetweed
Gaura mollis
Sources: Based on Buttrick (1914), McIntosh (1931), and Wright (1970).
104
resistant sedimentary strata. The vegetation is predominantly forests and woodlands of ponderosa pine, with
35
grasslands at lower elevations. The hills north of Newcastle are part of this region.11
Belle Fourche
Wyoming
South Dakota
55
50
40
Sundance
35
40
45
In the western part of the Black Hills, the sedimentary

strata form a comparatively flat area known as the Lime-
Spearfish
60
stone Plateau, with an elevation of 6,200 to 7,200 feet.

The underlying Madison Limestone (see chapter 2) is
Lead
45
55 50
permeable to water, which has enabled the formation of
40
numerous caverns, including Wind Cave and Jewel Cave.

Some creeks disappear into the bedrock before emerging
50
Rapid
City
Newcastle
again, such as Spearfish and Boulder creeks in the north.

44
marily ponderosa pine forests and savannas, commonly

45
50
45
Hot
Springs
10
10 mi
15
15 km
sandstones, forming the deepest and most spectacular

canyon in the Hills. The vegetation of the plateau is pri-
48
Custer
Spearfish Creek has cut through 400 feet of Paleozoic
intermingled with bur oak woodlands in the north and

groves of white spruce in canyons or on north slopes.
Also found are grasslands and foothill shrublands. A few
of the grasslands still have native species, but most are
now dominated by smooth brome, timothy, and other
introduced plants.12
The fifth geomorphic region is the igneous, granitic
Ardmore
Fig. 16.3. Mean annual precipitation patterns (shown in centimeters) in the Black Hills and Bear Lodge Mountains. Divide
centimeters by 2.54 to obtain inches. Adapted from Boldt and
Van Deusen (1974).
core of the Hills, which has been exposed through the

erosion of sedimentary rocks. Known as the Central
Area, it has many pinnacles and steep slopes (fig. 16.4).
The elevation is mostly 5,0006,000 feet.13 Again, the
vegetation is predominantly ponderosa pine forest, with
occasional groves of white spruce and aspen in wetter
habitats. Several lakes occur in the Hills, such as Sylvan
On the interior side of the Hogback Rim is the Red
Lake, but dams on creeks have created all of them.
Valleya name that stems from the red shales of the
The Bear Lodge Mountains, located to the northwest
Spearfish Formation that give a characteristic red color
of the Black Hills, occupy a smaller area and have less-
to the fine-textured soils.10 Also known as the race-
well-defined geomorphic regions. The highest point is
track because of its oval shape, the valley was formed
Warren Peak at 6,655 feet, about 600 feet lower than
by the erosion of softer shales, siltstones, and sand-
Harney Peak. Except for the absence of white spruce and
stones underlying the more resistant rocks of the Hog-
a few other species, the vegetation of the Bear Lodge
back Rim. Grasslands predominated in the valley when
Mountains is similar to that of the Black Hills.
European immigrants arrived, as they do today where

the land has not been cultivated. The towns of Sundance, Spearfish, Hot Springs, and the western part of
Vegetation Mosaic
Rapid City are found in the Red Valley, which ranges in
Nowhere in Wyoming and adjacent states are ponder-
elevation from about 3,000 to 3,600 feet.
osa pine forests so extensive (fig. 16.5; see fig. 1.5). They
Further to the interior, above the Red Valley, are
occur on nearly all soil types and exposures, occupy
the Minnelusa Foothills, composed of harder, erosion-
more than 90 percent of the Black Hills and Bear Lodge
Fig. 16.4. (above) The granitic Central Area of the Black Hills
has widespread forests of ponderosa pine, with groves of
aspen and white spruce along ravines and in other relatively
moist habitats. The red ponderosa pine on the left was invaded by mountain pine beetles the previous summer. This
photo was taken in 2010. Many more trees have been killed
by the bark beetles since that time. Elevation 5,0006,000
feet. Photo by Carl Christensen.
Fig. 16.5. (left) Ponderosa pine forests are common in the
Black Hills, the Bear Lodge Mountains, the Laramie Mountains, and on east slopes of the Bighorn Mountains. Common
juniper is a conspicuous shrub at higher elevations. This
photo was taken in the Black Hills. Elevation 6,400 feet.
Mountains, and can be classified into at least seven

habitat types.14 Rocky Mountain juniper occurs with
ponderosa pine on drier locations, such as on the Hogback Rim; and two shrubssnowberry and bearberry
are widespread associates in the Central Area and on
the Limestone Plateau. Similar forests occur on escarpments in the Chadron National Forest in northwestern
Nebraska and in Custer National Forest in southeastern
Montana. Where annual precipitation is higher, such as
in the northern Hills, ponderosa pine and bur oak form
a distinctive conifer-hardwood association (fig. 16.6).
pole pine and limber pine, both very common in the

Rocky Mountains to the west, are found in only one
or two localities in the Black Hills. Engelmann spruce,
subalpine fir, and Douglas-fir are not found there at all.
The Black Hills region has four kinds of deciduous
woodland: riparian, woody draws, bur oak, and aspen.17
The first, found in ravines and along rivers at low elevations, is dominated by boxelder, green ash, hackberry,
bur oak, American elm, and plains cottonwood.18
Woody draws are found nearby and have thickets of
chokecherry, American plum, currant, Woods rose,
hawthorn, western snowberry, red osier dogwood, and
various willows (see fig. 10.12). Historical photographs
suggest that these woodlands were more dense before
livestock grazing and farming began.
The third type of deciduous woodland occurs on the
upland at lower elevations and is dominated by bur oak
(see fig. 16.6). It occurs most often in the northern and
Fig. 16.6. Bur oak is found at lower elevations in the Bear Lodge
Mountains and Black Hills, sometimes growing under an open
canopy of ponderosa pine. This photo was taken on the east
slope of the Bear Lodge Mountains. Elevation 4,250 feet.
Common plants associated with the pine and oaks are

aspen, chokecherry, common juniper, mountain ninebark, paper birch, and white spruce.
In all localities, fire suppression has led to an increase
in the density of ponderosa pine, with highly flammable forests now found in many areas.15 Most of the
forests are second growth, having been subjected to logging for many years. An outbreak of the mountain pine
beetle has killed many of the trees during the past 10
years, but dwarf mistletoe is rare or absent.16
White spruce groves are found in canyons and
ravines, on north-facing slopes, and in cooler, wetter habitats at higher elevations (fig. 16.7). In especially humid locations, such as on the north slopes of
ravines, the tree branches sometimes are draped with
a lichen known as Old Mans Beard (Usnea cavernosa
and U. sorediifera). Spruce occurs on both igneous and
limestone substrates, commonly in close proximity
with ponderosa pine, aspen, and paper birch. Lodge-
Fig. 16.7. White spruce is common in the Black Hills at high

elevations and along valley bottoms with cool-air drainage. It
is rare in the Bear Lodge Mountains.
Fig. 16.8. Lt. Col. George A. Custer and his troops camped in
this meadow in 1874 (near the confluence of Silver Creek with
Castle Creek about 1 mile north of Deerfield Reservoir on U.S.
Forest Service road 110). The meadow appears the same today
as it was then based on a comparison of this photo with photos taken by expedition photographer William H. Illingworth.
However, the ponderosa pine forests in the background had
become more dense by the time this photo was taken in 2013.
A higher tree density could be attributed to natural forest
development after a fire that occurred a short time before
Custer arrived, or, alternatively, more trees could be explained
by fire suppression during the twentieth century. Notably,

mountain pine beetles are now thinning the forests, as shown
in fig. 16.9. Aspen groves can be seen on the edge of the
forest; Castle Creek flows through the foreground with various kinds of willows along its bank, including Bebb willow.
The meadow is now dominated by introduced species (such
as timothy, smooth brome, and Kentucky bluegrass), along
with green needlegrass, stiff goldenrod, and numerous other
natives. Illingworths photos of this meadow and others can
be viewed in Progulske (1974) and Grafe and Horsted (2002).
Elevation about 6,000 feet.
eastern parts of the Hills, mostly north of French Creek.
on the fringes of meadows (fig. 16.8). Associated species
Common associates include hophornbeam, ponderosa
include hazelnut, bracken fern, and wild sarsaparilla.
pine, green ash, American elm, and western snowberry.
Aspen can invade meadows and grasslands if browsing
On Mowry shale, bur oak forms a savanna with few
or grazing is not heavy. Aspen woodlands sometimes
understory plants. Sometimes the oak occurs as a shrub
give way to white spruce or ponderosa pine through
under ponderosa pine. Because of its ability to sprout,
succession. As with oak, the aspen and birch are able
oak can become the dominant tree if the pines are
to produce root sprouts and regain dominance more
killed by fire or harvesting. In fact, the re-establishment
quickly than ponderosa pine after a fire or other distur-
of pine may be difficult in some areas because of intense
bance (see chapter 11).
competition from oak and other species.19
The grasslands of the Red Valley and the fringes of
Aspen is the characteristic tree of the fourth type of
the Black Hills and Bear Lodge Mountains are typical
deciduous woodland.20 It occurs throughout the Black
mixed-grass prairie, dominated by blue grama, green
Hills on relatively moist sites. Occasionally aspen grows
needlegrass, little bluestem, needle-and-thread grass,
with paper birch or bur oak. The aspen woodland often
sideoats grama, western wheatgrass, and a variety of
forms a narrow band between the pine or spruce forest
sedges and forbs. 21 Where the soils are sandier, other
species are common, such as big bluestem, Indian rice-
wood are left in place for the benefit of wildlife and the
grass, and prairie sandreed. Westward, big sagebrush
maintenance of biological diversity. Research has shown
becomes more common. Streamside meadows in the
that at least 23 birds and 10 mammals in the Black Hills
same area have prairie cordgrass, tufted hairgrass, wild
depend on dead trees for food, nest sites, roosts, perches,
iris, and a variety of sedges. Inland saltgrass is com-
and dens.25 Sometimes the pines are thinned to provide
mon where the soils are saline and frequently moist.
more light, water, and nutrients for the growth of other
The interesting interaction of prairie dogs and bison on
desired species, such as bur oak or aspen. As tree density
the mixed-grass prairie of Wind Cave National Park has
is reduced, the growth and diversity of grasses, forbs,
been discussed in chapter 6.
and shrubs usually increase.26
At higher elevations, grasslands are found on the
Some believe there are too many roads in the Black
Limestone Plateau and in the Central Area near Deer-
Hills and that wood harvesting has taken precedence
field Reservoir. Those on the Limestone Plateau have
over other resource values, such as outdoor recreation,
been classified as Black Hills montane grasslands and
watershed protection, and biological diversity. Limit-
are identifiable by the presence of three grassesprairie
ing vehicular access is always controversial, but, though
dropseed, Richardsons needlegrass, and timber oat-
some roads have been closed, road density (miles of
grass. 22 The montane grasslands also have an abun-
road per square mile) in the Black Hills is still the high-
dance of colorful forbs, as Custer described in 1874.
est of any national forest in the region.
Several very large grasslands occur in the area, such as
The Black Hills also provide an abundance of live-
Gillette and Reynolds prairies. Only a few of the grass-
stock forage. 27 Excessive grazing still occurs in some
lands are still dominated by native plants, as most have
places (see chapter 6), especially in riparian meadows
been partially or completely converted to hay grasses,
and shrublands. Stream valleys, where much of the
mainly timothy and smooth brome. Those still domi-
private land is located, also are a preferred location
nated by native species are valued highly by conserva-
for summer homes. With livestock grazing and homes
tion biologistss.
concentrated in riparian zones, the habitat for big
Foothill shrublands are common in the region, typically on the Hogback Rim and lower parts of the Lime-
game, sensitive species, and other kinds of wildlife is

diminished.
stone Plateau. Skunkbush sumac and Rocky Mountain

23
juniper are common throughout the area, mountainmahogany occurs in the southwestern part of the Hills,
and snowbrush and russet buffaloberry are common in
the northern foothills.
Fire Suppression, Tree Density, and Streamflow

Perhaps the most subtle and far-reaching effect of land
management during the past century has been fire suppression, which is known to increase ponderosa pine
density. As discussed in chapter 11, surface fires char-
Land Management Issues

Timber Harvesting and Livestock
acterized many ponderosa pine forests and woodlands,

occurring historically every 1025 years, depending
on topographic position, elevation, and slope exposure. Surface fires kill most young trees, but usually not
Tree growth in the ponderosa pine forests of the Black
the older trees because of their thick bark. Such fires
Hills and Bear Lodge Mountains is the highest in the
also maintain a more open forest with relatively low
region because of the longer, warmer growing season
amounts of fuel. 28 Suppression led to the accumulation
and comparatively high summer rainfall.24 Ponderosa
of such large amounts of fuel that normally easy-to-
makes up 95 percent of the harvested wood. Consider-
control fires became stand-replacing wildfires.
able attention has been given to timber harvesting pre-
Although surface fires are thought to have been char-
scriptions that minimize adverse effects on other forest
acteristic of pre-European ponderosa pine forests, the
values. For example, clearcutting is generally avoided,
journals of early explorers suggest that crown fires also
with selective or shelterwood harvesting preferred, and
occurred. 29 For example, in 1880 H. Newton and W. P.
some standing dead trees (snags) and coarse downed
Jenney wrote:
The Black Hills have been subjected in the past to
neous and have fewer large trees. Their results support
extensive forest fires, which have destroyed the
the work of William Baker and his associates.36
timber over considerable area. Around Custer Peak
The apparent effects of fire suppression are various
and along the limestone divide, in the central por-
and wide ranging. To illustrate, tree density has clearly
tion of the Hills, on the headwaters of the Box Elder
increased, with trees often invading adjacent grasslands
and Rapid Creeks, scarcely a living tree is to be seen
and causing a concomitant reduction in the amount of
for miles. . . . Some portions of the parks and val-
forage for livestock, deer, and elk (fig 16.9).37 After long
leys, now destitute of trees, show by the presence
periods with high tree density, many understory plants
of charred trees and decaying stumps that they
die. In contrast, and predictably, thinning by timber
were once covered by forest, but generally the pine
harvesting leads to an increase in understory plant
springs up again as soon as it is burnt off, though
growth. Notably, a large number of tree seedlings often
sometimes it is succeeded for a time by thickets of
become established if thinning or some other kind of
small aspens.30
disturbance is followed by relatively wet weather the
Similarly, R. I. Dodge observed in 1876 that Throughout the Hills the number of trees which bear the marks
following spring. Tree density increases quickly if there

is no surface fire to kill the young trees (fig. 16.10).38
of the thunderbolt is very remarkable. . . . The woods are

frequently set on fire. . . . There are many broad belts of
country covered with tall straight trunks of what was
only a short time before a splendid forest of trees. Early
photographs also suggest a more open forest with many
dead standing trees.31
Lightning strikes are still a major cause of grassland
and forest fires in the region.32 Some areas might have
experienced a comparatively long fire-free period in
the 1800s, which would have enabled fuel accumulation sufficient for the extensive crown fires observed
by early explorers. Douglas Shinneman and William
Baker concluded in 1997 that frequent surface fires
maintained open savannas on drier sites, such as in the
southern Hills, but that forests at higher elevations and
in the northern Hills had less frequent, more dramatic
stand-replacing crown fires over areas of 20,000 acres
or more. Thus, its likely that the Black Hills and Bear
Lodge Mountains as a whole have a mixed fire-intensity
regime, with both kinds of fires possible, depending on
climatic and fuel conditions at the time.33 Native Americans surely had an effect on the fire regime as well (see
chapter 11). 34
Fires have been suppressed for such a long time
and timber harvesting has removed so much of the treering evidencethat reconstructing the original forest
structure and fire history using dendrochronology has
been difficult. However, ecologist Peter Brown and his
associates found evidence for a diversity of forests, with
savannas interspersed with high-density forests.35 They
concluded that the forests of today are more homoge-
Fig. 16.9. Young ponderosa pine are invading this meadow,

located to the south of the Castle Creek meadow shown in
fig.16.8. Above the meadow, mountain pine beetles are killing
many of the ponderosa pines, opening up the forest after a
long period of increasing tree density.
toward fire continues, fostered for many years by onesided Smokey Bear ads and by fires sometimes spreading onto private land from national parks and national
forests. Yet the negative effects of fire suppression have
moderated those views, and prescribed fires are now
common on both private and public lands.41
However, fire management plans are not easily implemented. Some forests have become so dense that uncontrollable crown fires develop easily. Often the labor of
tree thinning and removal is preferred (fig.16.11). Prescribed burning in grasslands also poses significant
challenges. For example, fires are usually kept small so
that they can be extinguished easily if weather conditions change unexpectedly, but the new regrowth of
burned grasslands typically attracts bison and cattle. If
the burns are too small, excessive grazing of the new
plant growth results.
Another challenge, in the Black Hills and other parts
of the Rocky Mountain region (see chapter 12), is the
large number of homes and communities dispersed
through most of the Hills. Managers hesitate to ignite
prescribed fires where homes are nearby. Such fires
almost always burn within the agreed-to limits of the
managers, and almost always they can be put out at
Fig. 16.10. Ponderosa pine seedling density can be high after
a period of weather conditions that favor seedling establishment. If surface fires occur frequently enough, the young trees
are killed before a highly flammable forest develops.
will. However, there are still risks. Consequently, prescribed fires are used less often than some managers
would like. And when they are ignited, when weather
conditions are favorable, the slow-moving fires usually burn only a small portion of the area that would
Increased tree density and forest expansion are also
benefit from burning. The conundrum is that overly
thought to have caused a reduction in streamflow, as
cautious prescribed fire causes higher risks of wildfire
evapotranspiration is higher in forests and woodlands
in the future.42
than in grasslands.39 Watershed managers know that
Fuel loadings are clearly an important factor con-
streamflow increases after tree cutting or burning (see
tributing to flammability in low-elevation ponderosa
chapter 12). The journals of early explorers suggest that,
pine forests, but weather is important as well. Peter
in the 1800s, there were more beaver, greater water stor-
Brown examined fire scar data from sites distributed
age capacity in the stream banks, more plant growth
throughout the Black Hills and found that the years
in widespread riparian meadows, and possibly more
with the most fires were associated with droughts.43 He
streamflow (but fewer flash floods).
also found that new seedlings of ponderosa pine were
40
The value of periodic surface fires for maintaining
more likely to become established during wet periods,
grasslands, savannas, and more open forests is now
with the most extensive recruitment associated with an
widely recognized. Among the benefits are reduced
extended wet period from the late 1700s to early 1800s.
fuels, a lower probability of hard-to-control crown
From this he concluded that even-aged cohorts often
fires, and more streamflow and forage production. Also,
become established because of a sequence of favorable
native plants are maintained, thereby reducing the
climatic conditions, not catastrophic fires or the initia-
likelihood of invasive plants. Negative public opinion
tion of fire suppression.
Fig. 16.11.This stand of ponderosa pine
has been thinned and is now thought
to be less susceptible to crown fires than
before. Because the small trees had little
market value, most of them were piled
and burned.
Considering the Black Hills region as a whole, the
story plants increased in abundance, such as snowberry,
largest fire in recorded history occurred in and around
yarrow, sideoats grama, needle-and-thread grass, Cana-
Jewel Cave National Monument in August 2000, follow
dian horseweed, and cudweed sagewort; others became
ing an extended dry period. Known as the Jasper Fire,
less abundant, such as common juniper, little bluestem,
it was of mixed severity and burned about 84,000 acres.
and ninebark. Unfortunately but predictably, several
Some of the forest had surface fires, other parts had
introduced species became more common as well, most
intensive crown fires, and still other areas did not burn
notably, bull thistle, Canada thistle, leafy spurge, and
at all. A patchy mosaic was created, with tree mortali-
Russian and spotted knapweed.
ties of about 20, 50, and 100 percent in areas of low,

moderate, and high burn severity, respectively.44 New
tree seedlings are becoming establishedthough slowly
Mountain Pine Beetles and Flammability
where burn severity was high, probably because most
Land management issues associated with insects, dis-
of the ponderosa seed supply was burned. Some under-
eases, and invasive plants are the same as those encoun-
tered elsewhere in the region (see previous chapters).45
In fact, the combination of beetles, fire, climate change,
An epidemic of the mountain pine beetle has been of
and other factors over a large area will promote the evo-
great concern during the past 20 years, causing much
lution and development of a more sustainable condi-
speculation about increased fire risk and what land
tion, whether managers take an active role or not.
managers might have done wrong to bring about such

conditions (see fig. 16.9). As noted, for a century or more
fires have been suppressed when possible and the trees
Invasive Plants, Climate Change, and Sensitive Species
have become more dense. The Forest Service and timber
Periodically the managers of federal and state lands are
industry collaborated in harvesting much of the wood
required to evaluate the ecosystems under their juris-
that was produced, though often not at the rate they
diction. In the Black Hills and Bear Lodge Mountains,
would have liked. The industry commonly claimed
that includes the Black Hills National Forest, Devils
the side benefit of improving forest health. Others felt
Tower National Monument, Jewel Cave National Monu-
that such harvesting caused habitat degradation over
ment, Wind Cave National Park, and Custer State Park.46
an excessively large area, or diminished the appeal of
Such assessments have been completed recently and, as
the area to tourists. Once again, land managers found
expected, all units are concerned about the challenge of
themselves in the middle of debates that often involved
dealing with the consequences of a long history of fire
governors and congressmen.
suppression and mountain pine beetle epidemics. Tim-
Presently, the furor seems to have diminished. More
ber harvesting has been accelerated where that option
people are recognizing that, on one hand, managing
is reasonable, and prescribed fires are used when there
ponderosa pine forests with attempts to suppress all
has been time to do the required preparation in places
fires will lead to other problems. Moreover, more people
where homes and other structures are either absent or
understand that the insects and diseases prevalent in
can be protected.
the Hills are nativepart of the natural biodiversity. To
Federal and state managers also are increasingly vigi-
manage the forest so that insect epidemics did not hap-
lant about the threats associated with introduced inva-
pen, or so that large fires did not occur, would create a
sive plants. Numerous species are involved, including
manicured forest that never existed before and many
buckthorn, burdock, Canada thistle, cheatgrass, com-
would not like. Even more roads would be required.
mon mullein, field bindweed, houndstongue, Kentucky
How to resolve such issues calls for creative collabora-
bluegrass, leafy spurge, spotted and Russian knapweed,
tion and acceptance of inevitable losses.
smooth brome, tumble mustard, white horehound, yel-
Current approaches for attempting to solve such
low toadflax, and yellow sweetclover. Curiously, the
problems include forest thinning and slash removal,
managers of Wind Cave National Park have observed a
which reduces the fuel load and increases the vigor of
rapid increase in white horehound, an introduced, inva-
surviving trees. The original structure of some pon-
sive plant in the mint family that is found primarily
derosa pine forests is thereby restored, at least to some
in association with the diggings of black-tailed prairie
degree. Prescribed fires are sometimes used when
dogs. Native species are adapted to such habitats as well,
weather conditions suggest that the risk of a wildfire
such as scarlet globemallow and fringed sagewort, but
is minimal and there are no mountain homes or busi-
they often are at a competitive disadvantage with the
nesses nearby, unusual circumstances in the area. Live-
introduced horehound. Mowing and pulling by hand
stock grazing is encouraged for reducing the abundance
are commonly used to keep weed populations in check,
of fine fuels, which may have an effect if that biomass is
and prescribed burning has been used successfully to
high after a year or two of above-average precipitation.
control some species.47 Herbicides in national parks
In some places the spread of aspen is promoted by cut-
are used carefully and with monitoring to minimize
ting the pines, which adds diversity to the landscape
impacts on other natural resources.
and could slow the movement of pine beetles. Accom-
The interaction between white horehound and black-
plishing such management activities over an area large
tailed prairie dogs is especially interesting because,
enough to be effective in a timely manner is difficult.
unlike the native plants, the horehound is not palatable
susceptible to predation, simply because the uneaten

plants provide more cover for stalking predators. If this
happens, the colony could be reduced in size, whether
from predation or because some of the rodents move
to places where more of the vegetation is palatable and
easily clipped. Horehound is a robust plant; clipping it
over large areas by prairie dogs consumes a considerable
amount of their energy.
Wind Cave National Park is one of two locations in
the area with sufficiently large prairie dog colonies to
support black-footed ferrets; the other is about 50 miles
to the east, in Badlands National Park and the adjacent
Conata Basin in Buffalo Gap National Grassland. The
ferret population at Wind Cave has been fairly stable
in recent years, with about 50 animals in 2011.48 If the
prairie dog population drops because of insufficient food
or other factors (such as sylvatic plague or canine dis
temper), or because of increased predation by other carnivores, then the ferret population will decline as well.49
The number of prairie dogs is difficult to estimate,
but it is relatively easy to determine the aerial extent
of active colonies. Notably, the size of the colonies at
Wind Cave National Park has been observed to fluctuate
Fig. 16.12.The dark area in this prairie dog town, located in
Wind Cave National Park, was caused by an herbicide applied
the previous summer for the purpose of removing white horehound, an introduced invasive plant that prairie dogs do not
eat. This photo was taken June 14, 2013. Managers anticipate
that native species will become re-established where such
control measures are taken, though the success of this treatment remains to be seen. See text for discussion of how white
horehound is thought to affect the animals adversely. There
are numerous prairie dog colonies in the park that do not yet
have horehound, as illustrated in fig. 16.13.
with weather conditions, expanding during relatively

dry periods.50 Prairie dogs concentrating in smaller
areas during wet yearsso they can clip the taller vegetation sufficiently to reduce predationcould explain
this pattern. An alternative, untested hypothesis is that
the number of prairie dogs, rather than their extent,
declines during wet periods because of more effective
predation during years when the plant cover is higher
which facilitates stalking by predators (fig. 16.13).
Considering that climate warming could lead to more
frequent droughts, prairie dogs may expand into larger
(fig. 16.12). Thus, prairie dogsalong with bison, elk,
areas of short vegetationassuming sufficient food is
and pronghornfeed primarily on the native species,
available and predation pressure does not increase as
thereby favoring the spread of horehound. Unless the
well (see chapter 6).
prairie dogs develop a taste for horehound, the animals
Bison and pronghorn populations are prominent
will suffer as their native food supply is reduced. Indi-
grazers in Wind Cave National Park, spending much of
rectly, this could have detrimental effects on the endan-
their time in the Red Valley portion of the park, where
gered black-footed ferret that was reintroduced to Wind
mixed-grass prairie is common. Both species are often
Cave National Park in 2007, not to mention the other
seen grazing on prairie dog towns, most likely because
species that depend on prairie dogs and their burrows
they find more nutritious forage there (see chapter 6).51
(see chapter 6). Moreover, if the horehound is not eaten
Pronghorn and bison population sizes in the park cur-
or readily clipped by the prairie dogsor removed by
rently are thought to be appropriate for the available
park managersthe prairie dogs may become more
rangeland, but, with the threat of increased drought
Fig. 16.13. A mixed-grass prairie with black-tailed prairie

dogs in Wind Cave National Park. The grass and other plants
are rather tall, an indication that the prairie dog population
was not large enough to keep up with plant growth at the
time this photo was taken. Normally the rodents clip the
vegetation low, both for food and, presumably, to reduce the
amount of cover for predators. The pale green plant is fringed

sagewort, which, along with scarlet globemallow, is more
abundant around prairie dog mounds and in other disturbed
areas. Both are native species; there are no invasive plants
in this colony. Missing from grasslands such as this are the
carcasses of large mammals, as discussed in chapter 6.
stemming from climate warming, there is concern that
easily jumped by elk. At 7 feet, the new fences are taller.
forage and water for the animals may be insufficient.
There is still no hunting allowed in the park.
Management options are a reduction in herd size or

supplemental feeding and watering.
An entirely different concern was identified in the

condition report for Devils Tower National Monument,
Elk are another large herbivore in the park, though
namely, a decline in riparian woodlands because of the
they are seen less commonly and are not attracted to
dam upstream on the Belle Fourche River, forming Key-
prairie dog towns. In recent years their population has
hole Reservoir. Constructed in 1952, the dam reduced
exceeded the number that managers think the park can
the frequency and magnitude of flooding, which is
sustainat least under current climatic conditions
important for seedling establishment of plains cotton-
without adverse effects on their habitat. To facilitate
wood. The older trees are now dying faster than they
achieving the goal of 400 elk, several hundred animals
are being replaced (see chapter 4). Small trees have
were recently herded out of the park through gates in
been planted to replace them, but survival has been
what is now thought to be an elk-proof fence. Generally,
poor. Also, the dam transformed a clear, fast-moving
such fences should be avoided around national parks,
river to a warm, sluggish stream with high turbidity.
but they were constructed originally to prevent the
The sediment load is partially attributable to livestock
bison in Wind Cave National Park from interbreeding
grazing along the bank outside the monument. As
with cattle. The genetically pure bison population
often noted, management activities outside a natural
in the park is highly valued by conservation biologists.
area can have significant effects on the insideand
The original fences were effective for bison, but were
vice versa.
Appropriately, condition assessments for the federal
Hills is marginal habitat for this species and that it also
and state lands now consider the implications of cli-
could become less common.53 The magnitude of such
mate change. Reports indicate that mean temperatures
changes can be debated, but the trends are the same.
in the region have risen by about 2F during the twen-
In general, future forests are likely to be much different
tieth century, and models predict an increase of 512F
than todays.
during the twenty-first century. As usual, there is less
As trees die from various inevitable disturbances, it
certainty about precipitation, but even if it increases,
would be ideal if landscapes could be restored to a condi-
evapotranspiration will increase as well because of the
tion that maintains the populations of sensitive species,
higher temperatures and longer growing season, most
such as aspen. Will that be possible with a warmer and
likely resulting in a drier climate. All managers in the
drier climate? Can the changes that are occurring be
region wonder what the effects of a warmer, drier cli-
guided so that the evolution of the native species can
mate will be.
continue, as they have for millennia? In fact, some sensi-
The most pertinent study on the effects of climate
tive species may not be able to survive; others may immi-
change was done in 2006 by U.S. Forest Service scien-
grate if the new environment is more favorable for them.
tist Gerald Rehfeldt and his associates.
The forests of the Black Hills and Bear Lodge Mountains
52
After careful
analysis of the climate required by ponderosa pine in
have changed dramatically before (see chapter 2).
western North America, they applied several climate

projections for the twenty-first century and concluded
Overall, the challenges of resource management are
that, most likely, ponderosa pine forests will be much
formidable in a landscape that has been modified by a
less dense and widespread in the Black Hills and Bear
century of fire suppression, habitat fragmentation, fenc-
Lodge Mountains by 2060 and that the species will no
ing, and the adverse effects of invasive species. And now
longer be reproducing by 2090 in some places where it
managers are faced with the uncertain effects of rapid cli-
now seems to thriveassuming unmitigated warming
mate change. The prospect of such changes is worrisome
continues. Mature trees can tolerate considerable tem-
to many, but two guiding principles are fundamental:
perature and drought stress, but as they die from old
conserve the diversity of native plants and animals and
age, beetle epidemics, fire, wind, or timber harvesting,
maintain the productive capacity of soils. One aspect of
the establishment of new seedlings will become less fre-
the Black Hills and Bear Lodge Mountains that will per-
quent than now because of the changing climate. Simi-
sist is the value of such places as refugia for mountain
larly, although aspen is the most widely distributed tree
plants and animals in an area otherwise dominated by
in North America, some think that much of the Black
the species of lowland grasslands and shrublands.
Chapter 17
The Laramie Basin
Westward from the Great Plains, beyond the Black
in previous chapters. Here the focus is on the Laramie
Hills and the Laramie Range, are a dozen or more inter-
Basin, one of the highest and most studied basins in the
mountain basins (see fig. 1.2). Long before they formed,
region (figs. 17.1 and 17.2).
the region was the site of inland seas in which thick

sequences of sedimentary strata accumulated (see chapter 2). As mountain ranges formed, the uplifted sedi-
Early Human History
mentary rocks were susceptible to erosion by water, ice,
The earliest evidence of humans in the Laramie Basin
and wind. Gradually, stones, sand, and silt produced
dates to about 13,000 years ago, based on the presence
by erosion were transported into adjacent basins,
of Clovis projectile points.2 By that time the climate
where they formed new strata. Hard as it is to imag-
was warming and the glacier that had reached the basin
ine, the basinsnow uplifted to a higher elevation
floor near Centennial had melted back to the slopes of
were nearly filled with material derived from adjacent
Medicine Bow Peak.3 Caribou, mammoths, mastodons,
mountain ranges. Some of the eroded mountains were
muskoxen, saber-tooth cats, and other exotic mammals
buried with these sediments. Geologists reached this
roamed across the tundra-like basin floor (see chapter 2).
conclusion because most basins are drained by a river
Cooling occurred again from about 1600 to 1850the
flowing through a canyon in the resistant rock of the
Little Ice Agebut not enough for glaciers to reach the
bordering uplift. Such a topographic feature develops
foothills as they had during the Pleistocene.4 Despite
where, at one time, the rivers course was established
climate changes, numerous Paleo-Indian artifacts sug-
over the buried mountain range. Gradually, the canyon
gest a frequent if not continuous occupation of the land
was cut by the river. As it deepened, more of the basin
in the vicinity of the Big and Little Laramie rivers.5
sediments could be washed away, further exposing the
The nomadic Blackfoot, Cheyenne, and Arapaho
mountain.1 Also indicative of a basins geologic history
tribes occupied the basin when Jacques La Ramie
are buttes and mesas, the tops of which are remnants of
arrived, at the age of about 35. Not much is known
ancient floodplains (see chapter 2).
about this French-Canadian trapper, but he is thought
Aside from these geologic similarities, the basins vary
to have been the first Caucasian in the area.6 Laura
in terms of elevation, climate, predominant vegetation,
Burdick wrote that La Ramie, in 1819, traveled up the
water availability, predominant land uses, exposed bed-
river on which he was camped, later to be named for
rock, mineral resources, water availability, and wildlife
him, against the advice of others who warned him
abundance. Some of this variability has been discussed
of the hostile Indians. He never returned and an
282
Fig. 17.1. Laramie Basin, with the Medicine Bow Mountains

on the west side and the Laramie Mountains on the east.
Centennial Valley, south of the town of Centennial, is defined
by the Medicine Bow Mountains on the left and Sheep Mountain on the right. The elevations of Laramie, Centennial, and
Woods Landing are 7,165, 8,074, and 8,095 feet, respectively.
The elevation of Wheatland Reservoir at the north end of
the basin is 6,959 feet, 1,136 feet lower than the Laramie
River at Woods Landinga straight-line distance of about

60 miles. The highest point in the Snowy Range, that part of
the Medicine Bow Mountains from which glaciers originated
during the Pleistocene, is Medicine Bow Peak12,013 feet.
The red dashed lines show the location of two transects used
for the profiles of geological surface types and vegetation
types shown in figs. 17.15 and 17.16. Cartography by Ramesh
Sivanpillai, University of Wyoming.
Medicine Bow
Mountains
Feet above
sea level
Snowy Range
12,000
Laramie
Range
Laramie
Basin
7,000
Mesozoic
Paleozoic
West
30 miles
East
Scale
Qa
Laramie River
Qa
Feet above
sea level
Laramie Range
Laramie Basin
8,600
Qa
7,200
Qa
Ku
Pf
Ps
Jm
West
4 miles
East
Scale
EXPLANATION
Quaternary
Alluvial deposits
Mesozoic
Cretaceous formations, undivided
Morrison Formation (Jurassic)
Chugwater Formation and Jelm Formation (Triassic)
Paleozoic
Forelle Limestone (Permian)
Satanka Shale (Permian)
Casper Formation (Pennsylvanian-Permian)
Precambrian
Granite
Undivided metamorphic rocks
Fault, arrow indicates relative direction of movement
Shear zone
Fig. 17.2. Geologic structure of the Laramie Basin. The top diagram is a generalized cross section of the entire basin, showing
the underlying sedimentary strata and the fault that was important in the formation of the Medicine Bow Mountains. The
bottom diagram shows more detail in the vicinity of Laramie.
The abundance of Quaternary alluvial deposits on the surface
is indicated, as is the thickness of the Casper Formationan
important aquifer in the basin. The Forelle Limestone forms a
prominent ridge east of Laramie; the colorful Casper Formation is exposed south of Laramie in the vicinity of Red Buttes
and Chimney Rock. Figure 2.3 illustrates the geologic development of the basin; a geologic time chart is presented in fig. 2.1.
Adapted from Mears et al. (1986).
unfinished cabin was found near Sybille.7 It would be
herds of buffalo, antelope [pronghorn] and mountain
good if more could be learned about his life, consider-
sheep. He also wrote about the many small streams that
ing that so many places bear his nameLaramie Peak,
were bordered with a thin growth of small willows and
Laramie River, Laramie City, Fort Laramie along the
richly stocked with beaver. Narrowleaf cottonwood
North Platte River, and Laramie County to the east of
also was observed. Another explorer, E.Willard Smith,
Albany County.
entered the valley from the south in 1840, observing
In 1825, about 5 years after La Ramie arrived, the fur
a great many buffalo every day as we passed along.
baron General William Ashley led an expedition across
In 1843 Captain John C. Fremonts party was in the
the basin. The general wrote in his journal that the val-
basin with Kit Carson, killing a pronghorn near Tie
ley and mountains were enlivened by in

numerable
Siding and camping along the Laramie River. As they
The Laramie Basin
passed around the north end of the Medicine Bow
established, and just north of the Laramie Basin, near
Mountains, sagebrush became more common, and the
Rock River, small deposits of coal were mined. With
party continued to encounter bison. Francis Parkman,
the exception of limestone for Portland cement, Albany
author of The Oregon Trail, was in the basin in 1846
County and the Laramie Basin generally are considered
and observed wolves, beaver, antelope, sage-grouse,
mineral poor. The rivers, lakes, and nearby mountains
elk, bighorn sheep, mule deer, and a forest fire. Three
provide many opportunities for outdoor activities. In
years later, in 1849, Captain Howard Stansbury and Jim
2012, Albany County had 37,000 residents83 percent
Bridger crossed the basin with a party of 18 mounted
living in Laramie.
horsemen, experiencing altercations with indigenous

tribe members along the way. Grasslands covered most
of the basin floor, as they do now. Stansbury also wrote
about bison, and he commented on an isolated grove of
Glacial Outwash, Hogbacks, Red Buttes, Big Hollow,

and Other Geologic Features
cottonwoods along the Laramie River, near where the
During the Pleistocene, glaciers in the Medicine Bow
city of Laramie would be founded 19 years later.8
Mountains of Wyoming and Colorado loosened large
The pace of EuroAmerican immigration accelerated
volumes of rocks, stones, sand, and silt.12 When the gla-
with the establishment of the Overland Stage Line by
ciers melted, much of this material was flushed onto the
Ben Holladay in 1862.9 This trail entered the basin from
basin floor, creating deltas. The waterways transporting
the southeast, near present-day Tie Siding, and contin-
and depositing this sedimentthe ancestors of the Big
ued westward around the north end of the Medicine
Laramie River, the Little Laramie River, and their tribu-
Bow Mountains (often referred to as the Snowy Range).
tariesshifted back and forth across the basin. These
Travelers with carts and wagons followed this trail.
waterborne sedimentslabeled Quater

nary alluvium,
Indian attacks led to the establishment of Fort Sanders
because they have been deposited since the beginning
in 1866, the location of present-day Laramie. The cav-
of the Pleistocene, 2.6 million years agoare the parent
alry provided security for cattle ranchers and for the
material for soils over about 75 percent of the Laramie
construction of the first transcontinental railroadthe
Basin floor (fig. 17.3). As might be expected, more large
Union Pacific. The railroad arrived in 1868, the same
rocks occur near the foothills than farther to the east
year that the city of Laramie was founded. After a brief
and north, where the flow of water was less forceful.
period of lawlessness and citizen vigilantes, Laramie
Before and following glacial advances, there were long
was settled with churches, schools, banks, a hospital,
periods of erosion from the basinbut only as rapidly
a courthouse, and, in 1873, the Territorial Federal Peni-
as the Laramie River could cut a canyon through the
tentiary. With the proximity to the railroad and the
Laramie Mountains to the northeast, a process that con-
availability of water, land privatization was more com-
tinues to this day.
plete than in basins to the west, where much of the land
Hogback ridges are conspicuous topographic features
to this day is in the federal domain. In 1886, before
on the perimeter of intermountain basins where sloping,
statehood in 1890, the Territorial Legislature designated
layered sedimentary strata are exposed. Strata of rela-
Laramie as the site for Wyomings first and, still today,
tively soft shale and siltstone erode more rapidly than
only 4-year university.
those of harder sandstone and limestone, with the result
During the 1900s there was great optimism about
that the more resistant strata became ridges (see fig. 17.2).
the potential for various land uses in the Laramie
The ridges create a different environment for plant and
Basin.10 Irrigated croplands were envisioned, but the
animal life. Mountain-mahogany and occasionally lim-
growing season proved too short and cool; only about
ber pine and ponderosa pine are found on the rocky soils.
3 percent of the basin is cultivated, mostly for hay.11
The ridges also add aesthetic appeal to the landscape and
The raising of cattle and sheep has been the most viable
provide desirable habitats for wintering animals, espe-
form of agriculture. Limestone and gypsum deposits
cially on leeward or south-facing slopes that are warmer
supported a brick and plaster industry for a time, and
in the winter (see chapter 10). People also enjoy having
later a cement industry. Several small oil fields were
homes in such places.
285
Fig. 17.3. Surface geology for the Wyoming portion of the

Laramie Basin. Recent alluvium and alluvial fans, shown in
yellow and buff colors, cover about 50 percent of the basin
floor. Green and blue colors show terraces and benches, which
originated as alluvium and occur on about 24 percent of the
floor. The Airport and Harmony benches are conspicuous (see
also fig. 17.5). Much of the surface has remnants of the Quaternary alluvium and older deposits that once filled the basin.
Some surface types listed in the legend are not easily distinguished at the scale of this map, including eolian deposits.
Cartography by Ramesh Sivanpillai, University of Wyoming.
Adapted from Case et al. (1998).
Prominent on the east side of the Laramie Basin are
aquifer is recharged by rain and snow on the west slope
the hard rocks of the Casper Formation, strata deposited
of the range, from its upslope edge near Lincoln Monu-
during the Paleozoic Era and now easily visible in the
ment down to the Laramie city limits. Protecting water
canyon east of Laramie, through which Interstate 80
quality in this aquifer is viewed as very important.14
passes. It is the oldest sedimentary formation in much of
The erosion of Quaternary alluvium and underlying
the basin, lying directly on top of Precambrian igneous
strata has been most rapid in the southern half of the
and metamorphic rocks (see fig. 17.2). It includes strata
Laramie Basin, where the elevation is about 1,000 feet
that contain the aquifer on which most people in the
higher than on the north end. This erosion has gone on
basin depend. Wells can be drilled to access this water,
sufficiently long that some of the basins oldest sedimen-
but artesian flows reach the surface in numerous places
tary strata are now exposed, often creating landforms
as springs, such as on the east side of Laramie.13 The
that are aesthetically, economically, and ecologically
The Laramie Basin
basin within a basin, about 10 miles long and 2 miles

wide, is easily seen along Wyoming Highway 130 (see
fig. 17.5) and is thought to have been formed by wind
erosion, starting most likely when glaciers extended
into the foothills above the town of Centennial. Strong
winds, possibly of hurricane force, apparently developed because of steep temperature gradients from the
top of the ice mass down to the valley.15 Such winds
dried and scoured the soil over a long enough period to
break down the deposits of calcium carbonate that previously had cemented the surface. Fine materials might
Fig. 17.4.Cross-bedding in sandstones of the Casper Formation indicate that the sand in this area was deposited in a
coastal dune field that existed here about 300 million years
ago. This photo was taken near Chimney Rock in the higher,
southwestern part of the Laramie Basin.
have blown eastward over the Laramie Range, whereas

coarser particles would have been carried downstream.
The bottom of Big Hollow is 300 feet below the surface
of the Airport Bench and has a playa lake in the bottom.
The smaller Bamforth Basin to the north most likely was
formed in a similar manner and also holds a playa lake.
Except for just above the town of Centennial, glaciers
important. For example, colorful sandstones of the Cas
never extended onto the floor of the Laramie Basin. The
per Formation are easily seen south of Laramie in the
melting of the latest glacier in this area began roughly
vicinity of Red Buttes. The brick-red pillars and monu-
10,000 years ago and led to the formation of a large,
ments add aesthetic appeal to the terrain. An equally
lobe-shaped glacial outwash plain, around which the
picturesque feature, known as Chimney Rock, is a rem-
Little Laramie River flows, first southward and then east
nant of Casper Formation sandstone and the underlying
and northeast. This relatively recent alluvium is easily
Fountain Formation, about 25 miles southwest of Lara-
seen along Highway 130 from Centennial eastward for
mie. Distinctive for its mass and shape, this natural mon-
about 6 miles, to the second crossing of the Little Lara-
ument is more than 200 feet high and has been sculpted
mie River (see fig. 17.3). To the west of Centennial, the
by erosion along Sand Creek. Stabilized sand dunes exist
highway climbs onto a terminal moraine marking the
in the area, formed from sand first deposited hundreds
farthest advance of the glacier.
of millions of years ago. The distinctive cross-bedding in
Various minerals have been extracted from the rock
the Casper sandstone exposed in the vicinity of Chim-
formations underlying the Laramie Basin. For a time in
ney Rock is evidence that it formed in a windblown
the nineteenth century, gold and copper mining took
coastal dune environmentnear the edge of an inland
place around Centennial and a few other localities.
sea that existed about 300 million years ago (fig. 17.4 ).
The most important minerals in the past 50 years have
Other topographic features also illustrate the geo-
been gypsum from the Pennsylvanian strata exposed
logic history of the basin. For example, west of Laramie
at the south end of the basin and limestone from the
are two large, flat, elongated benchesHarmony Bench
Casper Formation, along the west flank of the Laramie
and Airport Bench (fig. 17.5). Like buttes and mesas in
Range. Much of the stone used in the construction of
the area, these benches are capped by remnants of for-
older buildings on the University of Wyoming campus
mer floodplains. Now, on both sides, they are under
is sandstone from the Casper Formation.
going slow erosion by wind and water (see chapter 2).
Fossil fuel extraction has been a minor part of the
The tops of the benches have an abundance of river-
Laramie Basins economy. Only small amounts of oil
bed cobble that armors the surface, slowing the rate of
have been pumped from several small well fields, and,
erosion.
as noted, coalbeds are essentially nonexistent. Most
In addition, there is Big Hollow, reputedly the sec-
of Wyomings coal comes from Paleogene (early Ter-
ond largest deflation basin in the world. This elongated
tiary) strata, formed when subtropical swamps existed
287
Fig. 17.5.Topographic features in the Laramie Basin west of

Laramie, with Sheep Mountain in the background. Highways
and the Big and Little Laramie rivers are shown. Note the
location of the Airport and Harmony benches, and the large
deflation basin known as Big Hollow, west of the Airport
Bench. Bamforth Lake is in a smaller deflation basin. Lake
Hattie was enlarged to store more water for the irrigation of

hayland in the Pahlow strath, a term used for the valley between the south rim of Big Hollow and the Harmony Bench.
Drawing by Brainerd Mears, Jr., courtesy of the Wyoming
Geological Survey, Laramie.
over much of the area (see chapter 2). Such rocks have
annual temperature from 1991 to 2010 at the Laramie
been eroded from the Laramie Basin because of its high
Regional Airport was 41F, with July average highs and
elevation.
lows of 80F and 49F, respectively (fig. 17.6).16 Annually, most of the basin had only 75100 frost-free days
Elevation, Climate, and a Double Rainshadow
during this time. The short growing season combined

with relatively cool summer temperatures limits tillage
With an average elevation of about 7,200 feet, the
agriculture to mostly hay production where irrigation
Laramie Basin has the coolest climate of all the basins
water is available, such as near the Big and Little Lara-
in Wyoming, though similar to the Washakie and
mie rivers and along Rock Creek just to the north of
Green River basins to the west (see fig. 3.7). The mean
the basin.
The Laramie Basin

4.0
3.2
66
2.4
43
1.6
32
19
-5
0.8
that moisture falls in the basin. The annual precipitaMonthly Precipitation (inches)
Monthly Temperature (F)
90
0.0
Fig. 17.6. Average monthly precipitation (bars) and average

high and low temperatures (curves) at the airport in Laramie
(19812010). Comparable graphs from other weather stations
can be seen at http://www.usclimatedata.com.
tion at Laramie is only about 11 inches, most of which

falls in April and May. The mountains and basins to the
west tend to receive a higher proportion of the annual
precipitation in the winter, usually more than half. As a
result, sagebrush steppe is more common in the basins
to the west (see chapter 7).
Even with low precipitation, the Laramie Basin
was hospitable to both indigenous tribes and Euro
Americans because of snowfall in the adjacent mountains, which fed the Big and Little Laramie rivers in
addition to numerous springs on the east side of the
basin. Low-lying areas had numerous lakes and ponds,
such as in the vicinity of Lake Hattie and in the Sand
Creek drainage (see fig. 17.5).17 Waterfowl were abundant. Notably, creeks flowing into the basin from the
drier Laramie Range are ephemeral and provide little
The Laramie Basin is also windy at times, with a
surface water.
mean annual velocity estimated at about 12 miles/

hour. For comparison, mean annual wind velocities
for Missoula and Denver are about 6 and 9 miles/hour,
Vegetation Patterns
respectively. Snow drifting commonly occurs in lee-
Forests and woodlands are conspicuous as one crosses
ward locations (see also chapter 3), which favors the
mountain passes and descends into the Laramie Basin
development of shrublands dominated by big sage-
from the east, south, and west, whereas shrublands are
brush. Grasslands are found on surfaces that usually
more likely to be seen when entering at a lower eleva-
are blown free of snow.
tion from the north. However, mixed-grass prairie cov-
The basin is semi-arid for two primary reasons.
ers about two-thirds of the basin floor (fig. 17.7). Plant
First, evapotranspiration is more rapid at high eleva-
cover is sparse on windswept knolls, consisting of little
tions, especially when the wind is blowing. Second,
more than a few cushion plants and arid-land sedges
annual precipitation is quite low because of mountain
(threadleaf sedge and needleleaf sedge). In more favor-
ranges to the east and west that cause rainshadow on
able environments, the vegetation is dense, dominated
both sides. Several times each season it is common
by blue grama, western wheatgrass, Sandberg bluegrass,
to see storm clouds on the horizons that never reach
junegrass, needle-and-thread grass, fringed sagewort,
the basin. Those on the east are typically produced by
sand lily, Hoods phlox, and many other species (figs.
upslope storms, created when a low-pressure cell passes
17.817.10). Big sagebrush, mostly Wyoming big sage-
through southern Colorado or northern New Mexico.
brush, usually grows where snow accumulates on the
At such times, moist air from the Gulf of Mexico cir-
leeward side of ridges. Less than 10 percent of the basin
culates counterclockwise onto the east slope of the
has sagebrush-dominated shrublands, and much of that
Rocky Mountains. As the air rises, the air cools and
is black sagebrush on shallow soils in the foothills.18
condensation occursthe upslope conditions men-
Mountain-mahogany grows on exposed bedrock, and
tioned by weather forecasters. The precipitation that
some mountain big sagebrush is found where the soil
falls is mostly on the east side of the mountains. To the
is deeper and more snow accumulates (see chapter 7).
west of the basin, the air rising over the Medicine Bow
In general, plant growth and species composition vary
Mountains generally flows from the Pacific Ocean and,
largely as a function of wind exposure; the potential
having already passed over several mountain ranges,
for snow drifting; and the depth, salinity, and water-
is comparatively dry. Thus, only a small proportion of
holding capacity of the soil.
289
Fig. 17.7. Land cover in the Wyoming portion of the Laramie

Basin. Grasslands occupy about 66 percent of the basin floor,
with shrublands dominated by big sagebrush and greasewood
occurring on about 10 and 5 percent of the land, respectively. About 9 percent of the basin is in riparian woodlands,
shrublands, and meadows; about 3 percent of the land is in
cropland agriculture, half of which is irrigated. Some cover
types listed in the legend are not easily distinguished at the
scale of this map, including foothill shrublands on the lower

slopes of the mountains, small patches of irrigated land, and
the different kinds of riparian vegetation. The thicker dashed
lines show the location of two transects used for the profiles
of geological surface types and vegetation types, which are
shown in figs. 17.15 and 17.16. Cartography by Ramesh Sivanpillai, University of Wyoming.
The Laramie Basin
Fig. 17.8. (above) Mixed-grass prairie after a favorable growing

season in the northern part of the Laramie Basin. Common
grasses are blue grama, Indian ricegrass, junegrass, needleand-thread grass, Sandberg bluegrass, and western wheatgrass.
Fig. 17.9. (left) True mountain-mahogany grows on the shallow soils of the ridges (Forelle Limestone), and black sagebrush dominates the shrubland in the foreground, underlain
by Satanka shale. The city of Laramie is in the background, at
an elevation of 7,165 feet.
Above the basin floor, where annual precipitation

is much greater, mountain forests are more common,
especially to the west, where dense forests of lodgepole pine, Engelmann spruce, subalpine fir, and occasional groves of aspen are widespread (see chapter 11).
Curiously, driving into the Medicine Bow Mountains,
one travels almost directly from the grasslands and
shrublands into lodgepole pine or aspen forest, unlike
elsewhere in the Rockies, where a widespread foothill
transition zone occurs, with juniper, limber pine, ponderosa pine, or Douglas-fir (see chapter 10). Much of the
west side of the basin floor seems too high for the usual
foothill transition. The Laramie Mountains on the
east are drier, and the typical foothill pattern is more
291

Fig. 17.10.Chimney Rock, also known
as Camel Rock, on the WyomingColorado border, southwest of Laramie.
The upper part is being carved from
Casper sandstone, the base from Fountain sandstone. Black sagebrush grows
on the sandy soil in the foreground.
This photo was taken in mid-November,
after the deciduous shrubs had lost
their leaves. The light gray shrub on
the gentle slope is true mountainmahogany, and the darker shrub on the
steep slope next to the base is skunkbush sumac. Elevation 7,700 feet.
apparent, with foothill shrublands of true mountain-
funnel moisture into places where roots concentrate.
mahogany and black sagebrush at lower elevations.
Both ponderosa pine and limber pine grow in such
At mid-elevations, on south slopes, scattered limber
microenvironments. Evidence indicates that the pines
pines and a few ponderosa pines grow with an under-
existed in such places about 6,000 years ago, when the
story of mountain-mahogany. On north-facing slopes
climate was warmer than it is today.19 However, then
at the same elevation, dense forests of limber pine and
as now, mixed-grass prairie grew over most of the
Douglas-fir are found, such as east of Laramie along
basin floor.
Interstate 80 passing through Telephone Canyon. Far-
Riparian woodlands, shrublands, meadows, and
ther up the canyon, forests or woodlands are found on
hayfields currently cover about 10 percent of the low-
both slopes. At the summit, such as near Lincoln Monu-
lands (figs. 17.12 and 17.13). Such a large proportion
ment, small stands of lodgepole pine, subalpine fir, and
of the landscape in riparian vegetation is unusual for
Engelmann spruce can be found on north slopes; limber
Wyoming and is attributable to flood irrigation (see
pine savannas occur elsewhere.
chapter 4).20 On the fringes of the riparian zone and
Trees rarely grow in the lowlands, except in riparian
in depressions, sometimes in association with saline
zones and in association with outcroppings of rock (fig.
shales, salts tend to accumulate on the soil surface,
17.11). Plausibly, the boulders and exposed soil provide
even to the point of forming a white crust (see chapters
favorable microenvironments for the establishment of
5 and 8). Only halophytes survive in such areas, includ-
tree seedlings, absorbing heat during the day that mod-
ing greasewood, saltbush, red swampfire, alkali saca-
erates cold temperatures at night. Also, there may be
ton, and saltgrass (about 4 percent of the basin floor)
less competition from other plants in such places, and
(fig. 17.14). Where water persists, small lakes develop,
the boulders and ridges may provide leeward environ-
with playa wetlands having bulrushes, spike

r ushes,
ments where snow drifting occurs, thereby providing
and in some places, cattails (see chapter 5). 21 Playa
more water. Moreover, when it rains, the water flows off
wetlands, marshes, and associated lakes and ponds
the rock surfaces into cracks or into the soil under the
are the core of three national wildlife refuges in the
rocks, where evaporation is minimal, which provides
basin, namely, Bamforth, Hutton Lake, and Mortenson
a longer-lasting source of water. Essentially, the rocks
Lake. Water diversions from Sand Creek to some of the
The Laramie Basin
lakes have been constructed to help maintain water

levels for the benefit of wildlife, especially migratory
shorebirds and waterfowl. However, the U.S. Fish and
Wildlife Service has junior water rights; thus, during
times of low streamflow, the refuges may not have
much water available. Mortenson Lake National Wildlife Refuge was established to protect the endangered
Wyoming toad (see fig. 1.9), known to exist only in the
Laramie Basin.
Elsewhere in the basin, the vegetation is more characteristic of sandy soils, such as south of Laramie in the
Red Buttes area and along Sand Creek. Small tracts of
dunes can be found, some stabilized by Indian ricegrass,
prairie sandreed, sandhill muhly, scurfpea, and silver
sagebrush (see chapter 9). Sand is supplied by the erosion of sandstones in the Casper Formation. Periodically, flash floods carry large amounts of sand from the
mountains.
Even more curious than sand dunes are the large
tracts of mima mounds in the basin, some on floodplains along the Big Laramie River and some in the
upland grasslands, such as southwest of Laramie (north
Fig. 17.11. Ponderosa pine and limber pine on an outcrop of
the Casper Formation on the east flank of Sheep Mountain.
Ponderosa pine occurs infrequently in the Laramie Basin, as
this species requires warmer summers with more rainfall than
has been characteristic for the basin.
of Harmony, and also near Hutton Lake National Wildlife Refuge, along the road to Chimney Rock; see figs.
9.9 and 9.10). The uncertain origin of these features is
discussed in chapter 9.
Fig. 17.12. Riparian woodlands east of Centennial

along the Little Laramie
River. The trees are narrowleaf
cottonwood. Grasslands and
shrublands extend into the
foothills of the Medicine Bow
Mountains. Limber pine is
common on the mountain
slope in the distance. Photo
by Ken Driese.
293
Fig. 17.13. (above) Riparian meadows and willow-dominated

shrublands along the Big Laramie River south of Woods Landing in Colorado. Note the lack of cottonwood in this area.
Also note the terraces and benches through which the river
flows. This topography was formed in the same way as along
the Snake River in Jackson Hole (see chapter 2).
Fig. 17.14. (left) Shrublands dominated by greasewood are
common in the Laramie Basin, occurring in low areas and
on the upland where the soils are saline and relatively moist.
Associated species include Gardners saltbush, pricklypear cactus, alkali sacaton, western wheatgrass, and Nuttalls alkali
grass. Greasewood often is found adjacent to playa wetlands,
as illustrated in fig. 5.6.
Geobotanical Relationships
Geobotany is the study of the relationships between
native plants and the underlying rocks on which they
grow. Such relationships have been considered in previous chapters, but several interesting patterns in the
Laramie Basin are noteworthy. One example is the
association of selenium-rich shales of the Niobrara Formation and a group of plants that only grow on soils
developed from this formation. Known as selenium
indicators, the group includes two-grooved milkvetch,
tineleaved milkvetch, woody aster, and princesplume. 22
The Laramie Basin
The Niobrara Formation in the Laramie Basin is exposed
by the availability of water next to the mountain, soil
on the west flank of Sheep Mountain and at scattered
chemistry over the Niobrara Shale and Forelle Limestone,
localities on the basin floor, including Big Hollow.
and a coarse soil texture (such as on the sandstones of
Though selenium is a requirement for animal health in
the Fountain, Casper, Muddy, and Cloverly formations).
low amounts, it is poisonous if too much is eaten.
Big sagebrush was found growing in relatively deep, well-
Another example is the distinctive way in which
drained soils derived from all rock types.
elongated aspen woodlands grow along the contour
Another informative approach to studying geo
on the east flank of Centennial Ridge, on the west side
botanical relationships is to examine the concomitant
of the basin (see fig. 10.3). In this case, the association
variation in topography, geologic surfaces, and land
is between aspen and surface exposure of the contact
cover along transects. Figures 17.15 and 17.16 illustrate
between permeable Paleozoic sedimentary rocks and
two profiles that were prepared for the Laramie Basin.
the impermeable Precambrian igneous and metamor-
One is 17 miles long and extends southeastward from
phic rocks that form the main part of the ridge. Water
the Little Laramie River to the Big Laramie River, pass-
flowing from above mostly stays near the surface, just
ing across the Airport Terrace and Big Hollow. The other
under the thin soils, until it meets permeable sedimen-
profile is 4 miles long, extending from the Big Laramie
tary strata of the Fountain Formation, where it infil-
River eastward to the Laramie Mountains, crossing a
trates the soil and bedrock. The whole mountainside
subdued ridge formed on the Chugwater Formation and
essentially funnels water to this contact zone, creating
a prominent ridge formed from the erosion-resistant
favorable conditions for aspen.
Forelle Limestone, across the erodible shales of the
In 1969, botanist William Myers studied the effects
Satanka Formation, and then up the lower outcrop of
of bedrock type on the abundance of native plants
the Casper Formation. The importance of shallow soils,
in the basin.23 With the assistance of noted geologist
salinity, and inundation from flooding can be seen. For
Brainerd Mears, he was able to identify nine distinctive
both transects, grassland plants are most common, sug-
sedimentary strata exposed on the west flank of Sheep
gesting that they tolerate a broad range of soil condi-
Mountain. Four of the strata were more resistant to ero-
tions and topography.
sion and were fully exposed as hogback ridges, whereas

others were more easily eroded and formed low, soilcovered troughs between the ridges. Some plants were
Reflecting on the Past, Anticipating the Future
found only on the Niobrara Formation, as might be
The Laramie Basin has experienced ecological changes
expected because of its high selenium content. They
throughout the Holocene and for millions of years be
included the selenium indicators mentioned previ-
fore that. Tectonic activity, fires, flooding, and climate
ously along with greasewood, saltbush, and winterfat.
change have been most influential. Recently, over
Other plants were found only on the resistant rocks of
shorter periods, the changes can be attributable to
the Fountain Formation, probably because they ben-
transitions in how native and EuroAmerican cultures
efited from a more favorable water supply (for example,
used the land (see chapter 2). Beaver were used hardly
aspen, shrubby cinquefoil, serviceberry, and common
at all by the Indians, but they were nearly extirpated
juniper). Another group of plants was found on the hog-
by EuroAmerican trappers (see chapter 2). The people
back ridges some distance from the mountain (currant,
who lived here for 10,000 years or more harvested only
mountain-mahogany, limber pine, and ponderosa pine).
a small number of trees, mostly small, in contrast to the
Myers found no plants that grew only in the troughs
immigrants arriving about 150 years ago. They needed
between the ridges, probably because these low areas
railroad ties and lumber for construction. Unfenced
had soils that were influenced as much by weathering
bison herds roamed the plains until the 1860s, when
of the adjacent ridge rocks as by the underlying shales.
cattle and sheep replaced them. Large numbers of
Myerss study illustrated that rather few species of
domestic livestock survived on the prairie vegetation.
plants are restricted to one geologic formation. In his
Sheet-wash erosion left some slopes with little or no
study area, plant distribution is determined primarily
soil, most likely because of heavy livestock grazing near
295
Elevation, feet
Modern Little
Laramie River
floodplain
7,500
Little
Laramie
River
terrace
Pleistocene
floodplain
Airport Bench
Eolian deflation Dissected Laramie River

basinBig Hollow bench
terrace
Modern
Laramie
River
floodplain
Dissected
bench with
Pleistocene
overwash
7,400
7,300
7,200
7,100
7,000
Northwest
0
10,000
Southeast
20,000
30,000
40,000
50,000
60,000
70,000
80,000
90,000
Distance along profile, feet

Riparian
Upland
grassland,
hayland
Upland
grassland
Fig. 17.15. Seventeen-mile profile across the Laramie Basin

with topographic features and land cover indicated. As
shown in figs. 17.1, 17.3, and 17.7, this long profile extends
southeastward from the Little Laramie River, where a mosaic
of riparian woodlands, shrublands, meadows, and irrigated
hayland can be found. Grasslands are found over most of the
upland, across the Airport Bench, and down the slopes to near
the bottom of Big Hollow, where a playa with an ephemeral
pond has formed. There is no drainage from Big Hollow, but
standing water is rare because of low precipitation and no
streamflow. The Airport Bench is a remnant of the valley
floor at an earlier phase in the excavation of the basin and
has a surface that has been armored by coarse river cobbles
Not
vegetated,
playa Upland
grassland
Irrigated Upland
Upland
hayland grassland
grassland
Riparian
deposited during the Pleistocene. From the Big Hollow playa

southeastward, grasslands are found on the slopes and top
of a dissected bench. Beyond that, the former floodplain of
the Big Laramie River, known as a terrace, has a mosaic of
grasslands, shrublands, and irrigated hayland. Several lakes
also are located in this area.Riparian vegetation is found in
the current floodplain of the Big Laramie River. The profile
terminates on the dissected margins of a third, more ancient,
erosion-deposition surface. The Harmony Bench, illustrated in
fig. 17.5, does not occur along this profile. Plant species composition is influenced by topography, varying soil characteristics, and exposure to wind.
sources of water and railroad stockyards (fig. 17.17).
Irrigation had two general effects. First, it disrupted
Wolves were eradicated, enabling coyotes to become
the streamflow patterns of the rivers and creeks, reduc-
more abundant. Prairie dog densities declined; down-
ing flood peaks in the spring and emptying many
cutting of stream channels accelerated.
streams by late summer, when irrigation water was in
Equally profound change resulted from the establish-
greatest demand. Second, it enlarged the area of land
ment of irrigation systems, beginning in the 1870s.24 The
that was flooded. Essentially all floodplains and some
broad floodplains of the Big Laramie and Little Laramie
adjacent uplands were altered by this practice. In gen-
rivers enabled flood irrigation (see figs. 17.1 and 17.7).
eral, native riparian woodlands and shrublands exist
Dams, canals, and ditches eventually were developed,
today only on land that was too wet for domestic use,
including the Wheatland reservoirs on the Laramie
or where the presence of native plants provides benefits
River. Some of this irrigated land, though a small pro-
appreciated by the landowners, such as shade for live-
portion of the basin floor, was originally tilled for crops,
stock, erosion control, shelter for homes and barns, or
eliminating the native vegetation, but this practice soon
habitat for fish and wildlife.
proved uneconomical because of the cool, short grow-
Today the glint of numerous lakes across the basin is
ing season. Livestock production prospered; irrigated
especially obvious at sunset from the crest of the Lara-
haylandnow with mostly introduced speciesbecame
mie Mountainsall more or less permanent because of
a valuable commodity.
their connection to irrigation systems and providing
The Laramie Basin
Big Laramie
River floodplain
Chugwater
Formation
Tertiary
alluvium
Chugwater
Formation
Forelle
Limestone
Satanka
Shale
297
Casper
Formation
Elevation, feet
7,600
7,400
7,200
7,000
West
East
5,000
10,000
15,000
20,000
Distance along profile, feet

Riparian
meadow,
shrubland
Meadow,
upland
grassland
Not
Riparian
vegetated, meadow
landfill
Upland
grassland,
sagebrush
Riparian
Sagebrush,
meadow
mtn.-mahogany
Foothill
Upland
shrubland
grassland,
sagebrush
Fig. 17.16. Four-mile profile just north of Laramie, from the

Big Laramie River into the foothills of the Laramie Mountains, illustrating topographic features and land cover (see
figs. 17.1, 17.3, and 17.7 for location). Toward the east, beyond
the riparian meadows and shrublands, the surface slopes up
onto red beds of the Chugwater Formation. This rock is more
resistant to erosion than most materials in the middle of the
Laramie Basin, but the sloping surface has been altered by
erosional processes. At about 8,000 feet along the profile,
alluvial materials from higher slopes cover the Chugwater.
Soil development throughout this area is sufficient to support
grasslands. The portion of the profile lacking vegetation is the
Laramie Landfill. A narrow riparian meadow is intercepted
just beyond the landfill, where the profile crosses the bed of
an ephemeral stream. The steeper Forelle Limestone then
rises to a more pronounced hogback ridge with shallow soils

supporting foothill shrublands. The ridge is sharply defined
because of erosion of the underlying Satanka Shale beds to
the east. The profile ends on the slope of the Casper Formation, which extends to the crest of the Laramie Rangethe
hydrologic boundary of the basin. The lowest outcrops of the
Casper Formation mark the boundary of the valley floor for
much of the eastern side of the basin. Grasslands are characteristic of deeper soils developed over the Chugwater and
Satanka formations; shrublands with mountain-mahogany,
Rocky Mountain juniper, black sagebrush, and wild currant
are found on the ridges of the Forelle and Casper formations.
The Casper Formation is a very important aquifer, as discussed in the text, and its limestone strata are an important
source of calcium carbonate for the local cement plant.
wetlands that are important for wildlife (see chapter
endangered speciesthe whooping crane, piping plo-
5).25 Overall, irrigation has retained some of the water
ver, least tern, and pallid sturgeon. The Nebraska species
that previously flowed out of the basin, spreading water
depend on adequate water in the Platte River, of which
across the lower, flatter parts of the landscape, thereby
the Laramie River is a tributary. Potentially, more irri-
producing better-watered ecosystems than had existed
gation upstream in Wyoming, or more severe droughts
previously. Water withdrawals modified the rivers, but
anywhere in the watershed, could reduce the amount
irrigation waters that contributed to the formation of
of water downstream. For whatever reason, if more river
wetlands appear to have increased biological diversity
water is required to maintain viable populations of the
in the basin as a whole. Introduced trout still thrive in
endangered species in Nebraska, the Platte River Com-
the rivers, as they do in the lakes that had been created
pact and other regulatory statutes could require more
or enlarged. As noted, Congress designated some associ-
efficient use of Wyoming and Colorado water.27 That
ated wetlands as national wildlife refuges. Some species
could reduce water availability for Wyoming wetlands.
clearly benefited from wetlands maintained with irriga-
Can irrigation systems be engineered to use water more
tion projects, including mosquitoes. 26
efficiently and so enable survival of the Nebraska endan-
A conundrum has now developed that connects Wyo-
gered species without leading to the decline of Wyo-
ming wetland habitatand the endangered Wyoming
mings endangered species? That challenge would be
toadwith wetlands in Nebraska, where there are four
especially difficult if climate change leads to extended
Fig. 17.17.Exposed limestone of the Forelle Formation on the

east edge of Laramie now supports a shrubland of mountainmahogany. Most likely, parts of this area were mixed-grass
prairie prior to the arrival of the Union Pacific Railroad in
1868, when large numbers of cattle and sheep congregated
in the area because of close proximity to the city springs and
a spur of the railroad, where the animals were loaded onto
stock cars. Erosion over most the area would have been an
ongoing process because of the slope of the underlying bedrock, but it probably was accelerated by heavy grazing. Flash
floods are known to occur in the area, which would have
contributed further to soil loss. A remnant of the prairie can
be seen in the central part of the photo, where the soil has
not yet eroded. Limestone was quarried in this area, first for
building construction and later for the manufacture of plaster
and cement.
droughts. The way current laws are written, water rights
the potential for more wind farms, transmission lines,
for wildlife benefits have low priority.28
and roads (see chapter 18).
Another kind of change is associated with rural sub-
As in ecosystems throughout the region, invasive
divisions, especially around Laramie, along rivers, and
plants are a growing problem. 29 The grasslands and
in the foothills and mountains. As people move to the
shrublands of the Laramie Basin are still relatively
country, former grazing land is converted to residen-
free of invasive plants, though cheatgrass is becoming
tial use, wildlife habitat is fragmented, open space is
more common in the foothills. The riparian zones are
diminished, and the potential for problems with inva-
another matter, where Canada thistle, Dalmatian toad-
sive plants increases (see chapter 18). Moreover, the
flax, perennial pepperweed, white top, and spotted and
potential pollution of the Casper aquifer by a few land-
Russian knapweed are common. Some of their effects
owners on the outskirts of Laramie is a concern for the
are discussed in previous chapters. Native meadow
entire community. Guidelines have been proposed that
grasses that originally were harvested for hay included
would protect the aquifer, but this necessarily involves
meadow brome, redtop, and tufted hairgrass, but with
the cooperation of landowners who have other visions
irrigation they have been largely displaced by intro-
for their land. Currently, rural subdivisions occupy
duced species, such as foxtail, redtop, smooth brome,
only about 8 percent of the basin floor, but much of
and timothy. 30 One variety of foxtail, Garrison creeping
that is concentrated in the foothills, and the land area
foxtail, was introduced unintentionally when hay bales
involved increases annually as more people seek alter-
were brought to the basin from elsewhere. Initially,
natives to living in town. Accentuating such trends is
some ranchers appreciated its forage value, but now it
The Laramie Basin
43
6.0
42
5.5
41
5.0
40
4.5
4.0
39
3.5
38
1890
1910
1930
1950
Year
1970
1990
2010
Mean Annual Temperature (C)
Mean Annual Temperature (F)
6.5
3.0
Fig. 17.18. Mean annual temperature for Laramie from 1895

to 2011. An increasing trend since about 1982 is suggested,
though the highs and lows are still within the range of historic
variability. The solid horizontal line is the average temperature for the period of record (40.3F); the dashed lines are 1
standard deviation. See also fig. 3.9. From the Wyoming Water
Resources Data System, prepared with PRISM Climate Group
data, Oregon State University (http://www.prism.oregonstate
.edu/terms.phtml), courtesy of Christopher Nicholson.
has displaced many other meadow species. The hay,
estimate of trends, meteorologists now combine infor-
though abundant, is generally considered to be inferior
mation about typical storm patterns with data from a
to that existing previously.
Restoration to more desir-
network of weather stations to calculate annual precip-
able species will be difficult, should that be considered
itation for tracts of ground roughly 2.5 miles square.
a practical and worthwhile goal.
When this is done for Laramie, a slight downward trend
31
Climate change is likely to influence basin eco
is apparent (fig. 17.19). The mean annual precipitation
systems even more than other human activities. Chap-
in the past 10 years is 9 percent less than in the previous
ter 3 includes a discussion of climate change in general,
decade (11.1 inches instead of 12.2 inches).
and the potential effects are considered throughout
Similarly, climatologists have integrated precipita-
this book. For the Laramie Basin specifically, even
tion data for large drainage basins. Specifically, scien-
though mean annual temperatures during the past 10
tists with the National Climate Data Center divided
years are still within the range of historical variabil-
the state into ten basins and produced historical mean
ity, the record suggests a gradual warming trend since
annual precipitation for each year going back to 1895. 32
the mid-1980s (fig. 17.18; see also fig. 3.9). With regard
A cursory examination of the ten graphs suggests that
to precipitation, the trends are more difficult to ascer-
six of the basins have had no obvious trend upward or
tain. However, unless annual precipitation increases
downward during a period of 116 years. Four, includ-
along with temperature, a warmer climate will lead to
ing the Upper Platte River Basinof which the Lara-
more rapid drying and more frequent water stress dur-
mie Basin is a parthave had slight downward trends
ing the growing season.
(see fig. 17.19). Significantly, none of the basins had an
Though inconclusive, a closer look at precipitation
upward trend over the entire period of record. With
records is warranted. The most intuitive approach is to
warming, climate scientists predict that there will
look at records from a single weather station. However,
be less snowfall and more rainfall during the winter
there is great variation from year to year. Also, precipi-
months, and that snow cover will disappear earlier
tation events tend to be patchy across the landscape;
in the spring. As described in chapter 3, this trend is
downpours occur in one area but not nearby. For a better
already happening. Even if annual precipitation stays
299
19
Laramie
Annual precipitation (Oct-Sept)

5 year average precipitation
1895-2012 average precipitation
45
40
15
35
13
30
11
Annual Precipitation (cm)
Annual Precipitation (inches)
17
25
9
20
7
19
Upper Platte River Basin
Annual precipitation (Oct-Sept)

5 year average precipitation
1895-2012 average precipitation
45
40
15
35
13
30
11
Annual Precipitation (cm)
Annual Precipitation (inches)
17
25
9
20
7
1900
1910
1920
1930
1940
1950
1960
1970
1980
1990
2000
2010
Year (October - September)

Fig. 17.19.The top graph shows annual precipitation from
1895 to 2012 for a 2.5-square-mile tract of land in the vicinity
of Laramie, based on a network of actual data and knowledge
about snow and rainfall events in the area. The bottom
graph shows the same for the Upper Platte River Basin, which
includes the Saratoga Valley and Laramie Basin. A downward trend is suggested. The dashed line in both graphs is 1
standard deviation from the average for the period of record
(11.8 inches). The red line is a 5-year running average. Average
annual precipitation for the period of record is 12.1 inches.
The average annual precipitation in the early 1900s, when

irrigation systems and Wyoming water law were developed,
was 34 inches higher than at present. The top graph is from
the Wyoming Water Resources Data System, prepared with
the PRISM Climate Group data, Oregon State University,
http://prism.oregonstate.edu. The bottom graph is from the
Wyoming Water Resources Data System, prepared in collaboration with the National Climatic Data Center (http://
www.ncdc.noaa.gov/oa/ncdc.html), courtesy of Christopher
Nicholson.
The Laramie Basin
about the same, drought stress is likely to be more fre-
increase in mean annual temperature of 25F, which
quenta trend that is apparent for the western states
would likely increase potential evapotranspiration by
as a whole (see fig. 2.11).
0.010.04 inch/day. For this exercise, the first scenario
Overall, the data suggest a slight decline in annual
proposes increased winter precipitation and decreased
precipitation in the Laramie Basin during the past
growing season precipitation. The net effect would
10 years, a trend further substantiated by declining
cause a shift from grassland to mixed desert shrubland.
streamflow in some rivers, as is illustrated for the Green
The second scenario projects the same increase in tem-
River where it flows into Flaming Gorge Reservoir.33
perature, but with increased summer precipitation
However, no similar decline occurred for the Laramie
rather than winter precipitation. This scenario would
River. There is the potential that summer precipitation
favor continuing dominance of grassland, but with an
might be more closely associated with plant growth in
increase in cheatgrass. Ponderosa pine may also become
grasslands than is annual precipitation, but such analy-
more widespread than it is today, because of its apparent
ses have not yet been done for the Laramie Basin. Also,
reliance on warmer, wetter summers. 35
winter precipitation can be a significant factor for grasslands if it results in considerable water in the soil at the
In general, much remains to be learned about the impli-
end of winter.
cations of climate change combined with habitat frag-
Two scenarios for climate change have been proposed
mentation, invasive plants, and the long-term effects
for the east-central Rocky Mountain region that are per-
of living on the landa topic discussed further in the
tinent for the Laramie Basin.
next chapter.
34
Both assume a plausible
301
Part Six Sustainable Land
Management
Using Western
Landscapes
Chapter 18
The first EuroAmerican immigrants to Wyoming Terri-
West Coast along the Oregon Trail. 2 A few of them estab-
tory began building their homes in the 1820s. The human
lished homes along the way, but the first large wave of
population at that time numbered fewer than 10,000, 99
immigrants arrived after the completion of the Union
percent of them tribe members.1 Most likely, bison, elk,
Pacific Railroad in 1869. Others arrived in the 1950s and
deer, and pronghorn outnumbered people by 150 to 1.
1970s, when Wyomings oil, coal, uranium, natural gas,
Then, and for the previous 10,000 years or more, the
and soda ash became more valuable to the nation as a
primary human uses of the land were hunting and gath-
whole. Upticks in the annual census occurred with each
ering. Forty years later the population of domesticated
new development and were mostly associated with the
livestock still numbered only a few thousandmostly
energy industry. Tourists, campers, hunters, and anglers
horsesand timber harvesting was restricted to small
come to Wyoming during the summer and fall.
trees used for fuel, shelters, sleds, and weapons. The great-
During the past 200 years, new episodes of rapid
est human influence was igniting fires that sometimes
change have occurred. The world of Native Americans
burned for weeks or months. Tribal council meetings to
was turned upside down by the slaughter of bison and
discuss air and water quality were probably not necessary.
the introduction of horses, cattle, and sheep. Nothing
Today there are about 575,000 people and somewhat
so severe has happened since then. Still, the rate of
more than a million elk, deer, bison, moose, and prong-
population growth in western states has been above the
horn in Wyoming. Livestock, mostly cattle and sheep,
national average. Wyomings population grew 14 per-
number almost 2 million. A small proportion of the
cent from 2000 to 2010, with some counties growing
land is cultivated; agriculture is concentrated in a few
much more rapidly. For example, Campbell Countys
areas at low elevations. Fossil fuels rather than wood
population in the Powder River Basin grew by 37 per-
are the primary source of energy. Agriculture, techno-
cent during that 10-year period, and Sublette County in
logical advances, and towns and cities have benefited
the upper Green River Basin grew by 73 percent.
many people, but they have also led to a need for envi-
New oil and natural gas developments were the pri-
ronmental quality councils at federal, state, and local
mary cause for this latest wave of immigration, with
levels. Conservation biologists and land managers are
thousands of new wells drilled in the Powder River Basin
faced with new challengesthe topic of this chapter.
alone. Such activity has threatened biological diversity

through habitat fragmentation. An added threat has been
The New West
new rural housing developments, which provide places

where people can live close to the natural amenities that
Hundreds of thousands of EuroAmericans passed
more and more people enjoygood hunting, fishing,
through Wyoming in the mid-1800s, destined for the
hiking, and biking. But traffic has increased along the

305
306 Sustainable Land Management
Fig. 18.1.Open spaces have persisted largely because of short,

cool growing seasons and the challenges associated with
obtaining water. However, new roads and homes are now
being constructed in such places on private land, a trend that

is characteristic of the New West. The implications can be
significant. Photo by Nick Fuzessery.
thousands of miles of new roads that are required, and
of the past 20 years have had noticeable adverse effects
invasive plants have expanded their ranges. Among the
on many highly valued resourcesattracting the atten-
consequences are dramatic declines in the abundance of
tion of state governors and members of Congress. To
some native species, especially various birds found pri-
illustrate, Governor Jim Geringer sponsored work-
marily in shrublands and grasslands.
ing groups on the protection of open space, Governor
Alarmed by such trends in what some people refer
Dave Freudenthal promoted the sage-grouse conserva-
to as the New West, Chris Madson, long-time editor of
tion initiative and the Wyoming Wildlife and Natural
Wyoming Wildlife, wrote in 2007:
Resources Trust (approved by the Legislature in 2005),
Its not that we havent been working on conservation

in the last thirty years. Weve cleaned up our air and
water; weve pulled species like the black-footed ferret
and whooping crane back from the brink, but overall,
the conservation effort of the last generation is clearly
not up to the challenge of offsetting the damage to
land and wildlife we continue to inflict. . . . What will
Wyomings streams look like in twenty years, when
the last of the high glaciers in the Wind River Range
fades away to nothing? Can we adequately mitigate
the impacts of the latest round of gas drilling if the
current ten-year drought in sagebrush country continues for thirty, fifty, even a hundred years?3
and Wyomings congressional delegation sponsored

the Wyoming Range Legacy Act, which withdrew some
lands from future oil and gas leasing. The current governor, Matt Mead, supports the sage-grouse conservation
initiative and is promoting policies that may mitigate
some of the adverse effects of energy development.
Thanks to such initiatives, some concerns have been
addressed by state laws. Its clear that many residents
place great value on, for example, land reclamation
after mining, the protection of air and water quality,
the proper location of industrial activities, and the wise
management of threatened but still-abundant fish and
wildlife.4 Moreover, support is growing for the improved
management of sensitive areas, such as riparian zones
In 1994, when Mountains and Plains was first pub-
and other habitats critical for threatened species, and
lished, the generally accepted sentiment was that over-
for the curtailment of the spread of invasive plants and
all, Wyoming landscapes were about as pristine as could
animals. Such problems are discussed in previous chap-
be found in the lower forty-eight states (figs. 18.1 and
ters. Here the focus is on water development, multiple
18.2). That may still be true. However, the rapid changes
use on public lands, habitat fragmentation, the concepts
Using Western Landscapes
Fig. 18.2.Cultivated land, usually restricted to lowlands

where water is accessible, provides open space as well. The impact on the native flora and fauna is significant, but the land
still provides habitat for some kinds of wildlife, and many
people appreciate pastoral vistas, such as this one. In the New

West, land trusts work with property owners interested in
conserving the habitat and ecosystem services provided by
such places.
of ecosystem management and ecosystem services, and
onto the land, but only at great cost. New reservoirs and
adaptation to climate change.
trans-basin diversions are on the planning boards, and

existing diversions are being upgraded. State laws and reg-
Water Development
ulations have placed constraints on such developments

reflecting the view of many that the few unregulated
Not surprisingly, water is a limiting factor for human
rivers that remain are a valuable resource. Also, though
activities throughout the region. The traditional solu-
still difficult to implement, state laws now recognize
tions have been to construct reservoirs, irrigation sys-
instream flows as a beneficial use of water, and portions
tems, and trans-basin diversions, thereby making water
of some rivers are now protected under the Wild and Sce-
available wherever economically feasible and politically
nic Rivers Act of 1968. In Wyoming, the protected rivers
desirable (fig. 18.3). Faced with interstate competition
are the headwaters of the Snake River and the Clarks Fork
for water and the rights associated with prior use, a
of the Yellowstone River. Combined, they represent less
common attitude has been to use it or lose it. Pro-
than 0.4 percent of the states river miles.5
moted historically by the U.S. Army Corps of Engineers,
Groundwater is another issue. The Ogallala (High
the U.S. Bureau of Reclamation, and local chambers of
Plains) aquifer on the western Great Plains, extending
commerce, and heavily subsidized by state and federal
from Nebraska and eastern Wyoming into New Mex-
grants and loans, such projects have indeed stimulated
ico and Texas, has supplied agriculture and munici-
the economy. Also, the projects have often created
palities for many years, but the rate of withdrawal
additional benefits, such as flood control, hydroelectric
exceeds the rate of recharge. Significant drawdowns
power, and new recreational opportunities.
of this aquifer have forced some farmers to revert to
Still, regulated rivers have replaced rivers that flowed
dryland farming.6 Furthermore, groundwater in some
freely long enough to cut canyons through mountain
areas has been degraded through the deep percolation
ranges, and many reservoirs will probably be silt laden
of polluted surface water. Clearly, humans have the
in less than a century. Reservoir silt can be pumped back
capacity to affect deep groundwater supplies. Recent
307
ACRE-FEET (millions)

15
14
13
12
11
10
9
8
7
6
5
4
3
2
1
0
1890
to develop more efficient irrigation systems. Indeed,

people from all walks of life are striving to reduce the
amount of water they use.
Multiple Use on Public Lands

For many years, little attention was given to land management activities on federal lands administered by
the Forest Service and the Bureau of Land Management. However, by the late 1950s, concerns were being
expressed about excessive timber harvesting and live1910
1930
1950
YEAR
1970
1990
2010
Fig. 18.3. Water storage capacity in Wyoming reservoirs has

increased very little during the past 50 years. The last major
project was the High Savery Reservoir on the Little Snake
River, completed in 2004 and located northwest of Baggs.
Though locally important, it added only 22,433 acre-feet of
storage capacity. One acre-foot is the volume of water required
to cover one acre of land with one foot of water. Based on data
from Ostresh et al. (1990) and recent public records.
stock grazing. Responding to public sentiment in 1960,

Congress passed the Multiple UseSustained Yield Act,
a law that emphasized the importance of recreation,
watershed, and fish and wildlife values on national
forest lands as well as that of timber, livestock forage,
and other commodities. A similar philosophy was eventually adopted by the Bureau of Land Management,
though Congress did not formalize it until passage of
the Federal Land Policy and Management Act in 1976.
Many are comfortable with the multiple use concept.
concerns have emphasized the potential impact of
Timber harvesting and the extraction of minerals and
toxic hydraulic fracturing fluids, used in a process
fossil fuels are still possible, but public lands are now val-
known as fracking to extract oil and gas that previ-
ued for more than these resources alone. At times, some
ously was inaccessible.
stakeholders believed that the mandates of the Multiple
The demand for water has led to the consideration
UseSustained Yield Act were being abandoned, such as
of cloud seeding with silver iodide and other nucleat-
when the Wilderness Act of 1964 was passed. However,
ing agents as a means of increasing precipitation. Some
as the 1960 law states, multiple use is not necessarily
observations suggested that seeding might increase rain
the combination of uses that will give the greatest dol-
and snowfall by 15 percent or more, at least during some
lar return or the greatest unit output. Multiple use does
years. If droughts caused by climate change become
not require resource extraction everywhere.
more common, augmenting water supplies in any way
Much of the current debate over public land manage
is attractive. To be effective, however, the seeding must
ment centers on the loss of wildlands, whether from oil
happen when moisture-laden clouds exist, which are
and gas development, mining, or timber harvesting, and
less common during drought. Thus, at the time extra
the roads associated with these activities (fig. 18.4). Tim-
water is most needed, the likelihood of producing more
ber harvesting was the primary concern 20 years ago,
rain or snowfall is low. A committee of scientists orga-
but now controversy over fossil fuel extraction is in the
nized in 2003 by the National Research Council, an arm
news more often. Oil and gas developments are promoted
of the National Academy of Sciences, concluded there is
because of their high economic value and the potential
no convincing evidence that cloud seeding is effective.7
for less dependency on other nations; timber harvest-
This committee also doubted that the cost-to-benefit
ing is promoted because it, too, contributes to the local
ratio would be low enough to justify such a herculean
economies, and sometimes tree cutting can be used to
effort to increase water supplies.
improve wildlife habitat (see chapters 11 and 12). How-
Desires for more water are easily rationalized, but
ever, others maintain that the remnants of wildlands
learning to live with the amount provided by natural
now cover a very small area and that the best use of such
weather and hydrologic patterns is likely more sus-
areas is for wildlife habitat, dispersed recreation, and
tainable. In this spirit, agriculturalists are working
solitudeamenities that attract many people to the area.
Fig. 18.4. Habitat fragmentation from timber harvesting is

clearly visible around the mostly forested Savage Run Wilderness, located in the Savage Run Creek watershed on the west
slope of the Medicine Bow Mountains. This is the only entire
watershed in the Wyoming portion of this mountain range
The role of the public in public land management has

been evolving over recent decades. In the early days of the
without roads. Red indicates forest meadows; areas of gray on

the left are foothill shrublands. The geometric shapes in the
forests are clearcuts done at various times prior to the 1980s.
Landsat Thematic Mapper image, June 24, 1991. Cartography
by Ken Driese.
public interactions. Multiple use is better understood; the

values of wildlands are more widely appreciated.
U.S. Forest Service, for example, management decisions

lay almost exclusively in the hands of the agencys foresters. After World War II the public took more interest in
Habitat Fragmentation
what was happening on their lands, and such legislation
Long before Europeans arrived in North America, grass-
as the 1969 National Environmental Policy Act mandated
lands and shrublands extended as far as the eye could
that the Forest Service inform people of significant man-
see. In one direction or another, however, there were
agement issues and proposed actions. During the past
ridges, buttes, riparian zones, sandhills, or mountain
20 years, partnerships have sprung up across the West,
rangesislands on the plains. With variety in available
as illustrated by the Uncompahgre Plateau Partnership
environments, a greater diversity of plants and animals
in western Colorado. Community members from diverse
could survive. Bison surely found shelter on the lee sides
backgrounds join Forest Service and Bureau of Land
of ridges, as did Native Americans.8 The landscape was
Management employees in evaluating forest conditions
patchy, but at a rather coarse scale.
and monitoring the effects of forest management activi-
Today, aerial views show a more intricate patchwork
ties. This collaboration greatly increases understanding of
of croplands, human shelters, groves of trees, oil and
the complexities of land management while building trust
gas fields, and wind energy projectsall connected by
between the agencies and the public. Participants include
roads and superimposed on the landscapes that were
representatives of timber, grazing, motorized recreation,
there before. Many populations of animals have found
and environmental interests. As a result, forest manage-
their habitats broken into patches too small for their
ment has moved forward with less of the conflict and
survival, or too isolated from other favorable patches to
polarization that had previously characterized agency-
enable migration, reproduction, and the procurement of
309

Fig. 18.5. Most of Wyoming is accessible by roads, though many are not
easily traveled or are private. With or
without fences, they affect the movements of pronghorn, deer, elk, and
sage-grouse. Moreover, their impacts
extend beyond the roadbed, contributing further to habitat fragmentation.
food. Some native animals have become locally extinct,
To illustrate, elk and mule deer in shrublands tend
or their populations have declined. Examples in forested
to stay about 1.5 miles away from roads in summer,
landscapes include the lynx, wolverine, grizzly bear, great
when traffic is greatest, and about 0.75 miles away
gray owl, boreal owl, northern goshawk, and northern fly-
from roads in winter.9 Much of their time is spent in
ing squirrel. In grasslands and shrublands, declining spe-
roadless areas. Similarly, sage sparrows and Brewers
cies include the sage-grouse, sage sparrow, sage thrasher,
sparrows tend to avoid roads and oil wells by about
Brewers sparrow, dickcissel, burrowing owl, lark bunting,
300 feet, and sage-grouse by 23 miles. Roads also are
bobolink, ferruginous hawk, prairie falcon, upland sand-
pathways along which invasive plants become estab-
piper, McGowns longspur, chestnut-collared longspur,
lished and expand.
Columbian sharp-tailed grouse, long-billed curlew, swift

fox, prairie dog, black-footed ferret, and pygmy rabbit.
Other effects of roads include direct mortality from

collisions. The number of animals killed by vehicles is
The adverse effects of fragmentation are most severe
small compared to fatalities from inadequate food sup-
during the inevitable episodes that lead to other stresses,
ply and hunting, but large ungulates on the highway
such as drought, severe winters, high levels of predation,
are genuine traffic hazards for people. Also, crossing
and avoidance behavior when humans or predators are
busy highways is one added stress that diminishes the
present. Even fires, beneficial for maintaining biological
energy in body fat required for surviving the winter. If
diversity since prehistoric times, can be detrimental if
energy reserves are insufficient, the animals are unable
they further fragment an already marginal habitat. Habi-
to return to their summer range, where their reserves
tat fragmentation can push animals into places where
are normally restored (see chapter 15).
they encounter excessive competition from others already
The best-known example of road effects on ungu-
there, or to habitats that are marginal, which reduces the
lates is at Trappers Point, along U.S. Highway 191 west
likelihood of survival during inevitable difficult times.
of Pinedale. This highway intersects the fall migra-
Roads, rural subdivisions, timber harvesting, and
tion route of pronghorn from Jackson Hole and Grand
energy development are common causes of habitat frag-
Teton National Park up the Gros Ventre River valley,
mentation and warrant further discussion (fig. 18.5).
across the divide to the headwaters of the Green River,
Roads
and then southward along the Green River valley to

the sagebrush steppe south of Pinedale and east of Big
The very presence of roads reduces the amount of hab-
Pineythe longest known migration of a mammal
itat far more than the area required for the roadway.
in the contiguous United States. If they survive, the
Fig. 18.6. Forest fragmentation by clearcutting in the

Medicine Bow Mountains. This mountain range is relatively
flat and accessible, which facilitates harvesting over larger
areas than, for example, in the more rugged Wind River and
Bighorn mountains. This aerial photo was taken in the 1980s.
New trees (mostly lodgepole pine) are now growing in all the
openings. Current management prescriptions call for smaller,
irregularly shaped openings, and tree thinning has become

more common than clearcutting. Much of the harvesting in
recent years has been of beetle-killed trees, which provides a
source of wood while reducing hazardous snags along roads
and near campgrounds. Another motivation for harvesting
in some areas is to improve the habitat for specific kinds of
wildlife. See also fig. 18.4. Photo by William K. Smith.
pronghorn return to Jackson Hole in the spring along
servation easements, bear-proof containers, and habitat
the same route. Collision mortality and the costs of
enhancements elsewhere.11 The effectiveness of such
vehicle repair have been a serious concern for many
measures is controversial.
years. To solve this problem, in 2012 the Wyoming

Highway Department completed a system of 25 miles
of 8-foot-high fences that funnel the pronghorn and
Timber Harvesting and Energy Development
deer through six underpasses and across two over-
The discussion of habitat fragmentation began about
passes along a 13-mile stretch of road. The $10 million
30 years ago with evaluations of the silvicultural prac-
cost of the project was justified largely by less vehicle
tice known as clearcutting, whereby all trees are cut
damage. Placing a value on each animal killed is still
whether they are harvested or not (fig. 18.6; see chap-
difficult.10 Without accommodations for wildlife, such
ter 12). The rationale for clearcutting was to enhance
as at Trappers Point, widening and dividing highways
the growth and health of new lodgepole pine trees by
to enable faster, safer travel for motorists has the oppo-
providing them with the maximum amount of light,
site effect on animals.
water, and nutrients, thereby mimicking some effects
New roads are often constructed into the foothills,
of a crown fire. Predictably, logging roads fragmented
because homebuilders are attracted to scenic locations
the forest still further, with the impact extending well
adjacent to national forests and national parks. Un
beyond the roadway. Conservation biologists distin-
fortunately, such places also are important habitat for
guish interior from edge forests, and have identified
some kinds of wildlife, especially during the winter.
species, mostly animals, that require one or the other.
People who live there commonly appreciate the abun-
Timber harvesting and road building can reduce the
dance of these animals, and sometimes they attempt
amount of interior forest and increase the amount of
to mitigate their own adverse effects, such as with con-
edge forest.
311
Debates often centered on whether natural forest disturbances, such as fire and blowdowns, had the same
effect as tree cutting (see chapter 12). Commonly, the
landscape patterns caused by natural disturbances in
wildland landscapes are thought to enhance biological
diversity, not because all native organisms find them
beneficial, but rather because the population sizes of
some species drop while others rise after the disturbance. The diversity exists because of the changes in the
landscape over long periods of time. Some landscapes
are large enough to accommodate such disturbances
without the adverse effects of fragmentation. Others are
already so patchy that further timber harvesting would
most likely lead to the loss of some species.
Although timber harvesting and roads have surely
contributed to forest fragmentation in some areas (see
fig. 18.4), some forests are naturally divided into small
patches, that is, mostly edge forest, because of abrupt
differences in mountain topography and soils.12 Harvesting might further fragment the habitat in such
places, but there is reason to question whether interior
forest is broken up in the process. The loss of interior
forest may be most significant in places of relatively flat
topography and extensive forest, such as in some parts
of the Medicine Bow Mountains.
Fig. 18.7. Habitat and open space fragmentation by oil and

gas development is now common in parts of the Powder River,
Green River, and Washakie basins. Photo Dave Showalter,
Lighthawk Aerial Support.
With regard to energy projects, a few well pads and

mines scattered about the landscape would have relatively little effect on wildlife. However, in the Powder
tors that use the towers for perches, such as ferruginous
River Basin alone, the energy boom during the past
hawks and golden eagles. All kinds of energy develop-
20 years has led to more than 10,000 new wells for
ment fragment wildlife habitat and have impacts on an
oil and gasall serviced by several thousand miles of
area larger than the land they occupy.13
new roads (fig. 18.7) and having adverse effects on air

quality. Similar intensities of development are occurring in other parts of the state, such as in the Green
Ecosystem Management
River and Washakie basins. Moreover, sixteen wind
Many difficulties associated with modern land man-
energy projects have been constructed in the state since
agement are attributable to rising demands on limited
1999, primarily south of Casper and along the Inter-
resources. This is occurring at a time when climate
state 80 corridor (fig. 18.8). More are envisioned. The
is becoming less favorable than when many societal
Chokecherry and Sierra Madre Wind Energy Project,
expectations and cultural traditions were developed.
now on the planning boards, anticipates 1,000 turbines
Ecosystem management is an attempt to achieve sus-
spread over an area of about 350 square miles of sage-
tainable land management by integrating existing eco
brush-dominated shrubland near Rawlins. Aside from
logical information with the very real and difficult
fragmentation concerns, when vertical structures are
challenges faced by managers. Effective eco

system
involved, such as the towers required for transmission
management also transcends artificial boundaries.
lines, prairie chickens and sage-grouse instinctively shy
Much of the best wildlife habitat, for example, occurs
away from the area to avoid the risk of predation by rap-
on private lands, whose owners depend on public land
Fig. 18.8. Wind energy projects also contribute to the fragmentation of habitat and open space. Towers for the turbines and
associated power lines provide perches for raptors that prey on
birds and small mammals more often than they would without
the perches, thereby extending the effects of this kind of

industrial development. Power companies strive to locate power
lines along corridors to minimize their visual and ecological
impacts. Photo Dave Showalter, Lighthawk Aerial Support.
resources (such as ranchers who have grazing leases, or
trary, as animals, plants, water, nutrients, sediments,
landowners with property that is highly valued because
energy, and pollutants move or disperse across them
it is adjacent to public land).
(see chapter 1). The size of the area needed to imple-
The ecosystem management concept reinforces
ment an ecosystem management approach depends on
such commonly accepted practices as environmental
what ecological issues the management plan is designed
impact assessment, public participation in the develop-
to address. For example, the area included in the Greater
ment of management plans, and monitoring of resource
Yellowstone Ecosystem was determined to a large extent
values to ensure that management practices are work-
by considering the land used by the widely dispersed
ing as intended. Goals include managing for long-term
elk and grizzly bear populations (see chapter 15). In all
sustainability, which means maintaining productive
cases, various landowners are involved; collaboration is
capacity of the soil (as mandated in the Multiple Use
needed.
Sustained Yield Act) and preventing species extinction
Understanding that some landowners might feel
wherever possible (as mandated by the Endangered
threatened by this approach, proponents of ecosystem
Species Act). The nature of ecosystems in the region
management strive to achieve sustainability while avoid-
certainly changed dramatically during the Holocene
ing adverse impacts on people who depend on the land-
and before (see chapter 2). Now, however, with greater
scapes of the region and who, indeed, can be important
societal demands on natural resources, combined with
stewards of the land. Enabling them to prosper reduces
habitat fragmentation and accelerated climate change,
the likelihood of land sales that could lead to sub
precautions seem prudent. Ecosystem management is a
divisions or other uses that are not helpful. Indeed, mak-
step toward being more cautious.
ing a living in a region with a semi-arid, cool climate
How much land is necessary for effective ecosystem
can be difficult. The most appropriate livelihoods and
management? Human-defined ecosystem boundaries,
management practices may not have been achieved dur-
like political boundaries, are always somewhat arbi-
ing a mere century of trial and error. Though difficult
313

Fig. 18.9. Monitoring water
quality provides an indication
of how successfully ecosystem
services on the watershed are
being maintained. Photo of
the Sweetwater River by Tony
Ferlisi.
to contemplate, some changes could lead to an economy
by native plants and that provide various amenities
that is more stable than the region has experienced since
and much of the Wests characterare now being
the late 1800s. Ecosystem management is an approach
fragmented.14 Some residents are convinced that such
that motivates willing participants to explore their
trends are not good for the Wests economy. For exam-
options more broadly than may have been done before.
ple, Montana State University ecologist Andrew Hansen

and his associates wrote, local communities may best
be able to promote economic growth by maintaining
Amenities and Ecosystem Services

A dollar value easily can be placed on water, meat,
wood, oil, gas, coal, and other commodities on which
the livelihoods of many residents depend. In the New
West, however, both old-timers and newcomers are
placing considerable value on open space, clean air,
free-flowing streams with clear water, and abundant
wildlifeincluding nongame species (figs. 18.9 and
18.10). For many years there seemed to be little conflict. Roads and ranches here and there contributed to
the pastoral scene that many enjoy; industrial developments and rural subdivisions were viewed as progress.
the natural amenities that are so attractive to new residents and businesses. If this is true, policies that favor
timber harvesting over scenery, mining over water quality, or intensive livestock grazing over wildlife habitat
might actually inhibit rather than expand economic
growth.15
Ecosystem services are also becoming more highly
valued, as discussed in previous chapters. The costs of
replacing them with new technology, where that is possible, can be high. Herb Bormann, long-time ecologist
at Yale University, wrote in 1976 that, to the degree eco
system services are lost,
Now such developments are becoming progressively
they must be replaced. . . . We must find replace-
closer to each other. Opportunities to appreciate open
ments for wood products, build erosion control
space decline with too many wind energy projects, oil
works, enlarge reservoirs, upgrade air pollution con-
and gas wells, pumping stations, ranchettes, power
trol technology, install flood control works, improve
plants, and power lines extending to the horizon.
water purification plants, increase air conditioning,
Widespread protective landscapesthose dominated
and provide new recreational facilities. . . . These

Fig. 18.10. Pronghorn are
now abundant in Wyoming,
numbering about 500,000,
but they were threatened
with extinction in the early
1900s from overhunting. The
conservation of this species
and some others has been a
success. Photo by Ken Driese.
substitutes represent an enormous tax burden, a
funding for such payments, the conservation banks
drain on the worlds supply of natural resources, and
sell conservation credits, usually to corporations seek-
increased stress on the natural systems that remain.
ing ways to mitigate the potentially adverse impacts of
. . . Increased consumption of fossil energy means
other developments they sponsor. The motivation for
increased stress on natural systems, which in turn
mitigation may be driven by corporate philosophy
means still more consumption of fossil energy to
which appeals to many consumers and investorsor by
replace lost natural functions if the quality of life is
regulatory mechanisms.
to be maintained.16
Bormann recognized, more than 35 years ago, the
economic value of using solar-powered ecosystems to
provide numerous services, including erosion control,
clean water, recreational opportunities, and habitat for
wildlife. Today we can add other services, such as carbon
sequestration and resistance to invasive plants. Fossil
fuels have put Wyoming in the position of subsidizing
new developments through the Wyoming Permanent
Mineral Trust Fund, made possible by royalties and severance taxes on oil, natural gas, and coal. However, such
nonrenewable resources can be used only as the basis
The federal government has developed similar programs. A good example is the Conservation Reserve
Program, whereby farmers are compensated for planting perennial plants on erodible soils that had been cultivated for annual crops. Erosion is reduced, there is less
dust in the air, water draining from such lands is cleaner,
and wildlife habitat is improved. Similar programs also
provide economic incentives to discourage the draining
of wetlands and the plowing of still-intact grasslands.
In this way, consumers from across the nation pay for
highly valued ecosystem services provided by participating landowners.
for a transitional economy. Recognizing environmental

constraints is fundamental to achieving sustainability.
The challenge is to find ways by which people can
Development by Design and Climate Change
profit from selling ecosystem services. Progress is being
The perceived antagonism between economic develop-
made. Numerous corporations across the nation have
ment and conservation has led to various approaches for
created conservation banks that are now federally regu-
setting priorities that are complementary to ecosystem
lated and employ experts in conservation finance.17 If
management. One example, known as Development by
landowners commit to making significant ecological
Design, was first articulated by The Nature Conservancy
improvements to their land, and if an objective third
(a nongovernmental organization) and stems from the
party confirms such improvements, payments are made
environmental impact statement process mandated by
to the landowner, entirely from private funds. To obtain
the National Environmental Policy Act of 1969.18 This
315
approach, now used by various entities, calls for assem-
previously degraded site, the purchase of conservation
bling the best available biological and ecological infor-
credits from a conservation bank, or the purchase of
mation about sites where a development is proposed.
conservation easements that prevent habitat loss on
That information is then displayed on maps using Geo-
other sites that probably would be the location of future
graphic Information Systems. The severity of anticipated
developments and the associated consequences. The
environmental impacts are assessed in a similar man-
approach is not perfect; population sizes of some sensi-
ner. After careful analysis, the impacts and conflicts are
tive species will decline. However, the process is cur-
identified.
rently viewed as minimizing adverse effects and may
But Development by Design does not stop there.
prove to be generally acceptable to diverse stakeholders.
The next step is to formulate a strategy for avoiding,
The goal is to stop declines in biological diversity. Com-
minimizing, or mitigating the adverse effects. Actions
pensation from a conservation bank is available for
are then taken, and the result is monitored to measure
landowners who can assist with the mitigation pro-
whether the plan was successful. Based on the results,
cess, as described previously. Also, for landowners who
plans developed in the future can be improveda
collaborate with U.S. Fish and Wildlife biologists in
decisionmaking process known as adaptive manage-
developing a conservation plan for sensitive species on
ment. The best scientific information is used, and
their property, there is the potential for a safe harbor
the analyses are peer reviewed, as required with any
agreement.21
science-driven approach.19 The process includes willing participants. The first application in Wyoming
Over the years, land managers have learned how to
was on the Jonah gas field south of Pinedale. To start
restore degraded ecosystems by fostering the growth
with, several corporations contributed some $25 mil-
of desired plant species, closing some roads, reduc-
lion to a fund administered by the Jonah Interagency
ing recreational impacts, applying new approaches
Office, which is overseen by a committee composed of
for conserving threatened species, and a variety of
specialists with the Wyoming Game and Fish Depart-
other initiatives. Ecosystem services often have been
ment, Wyoming Department of Agriculture, Wyoming
restored. Success is always more likely when the climate
Department of Environmental Quality, and the Bureau
is favorable and familiar. Now, however, the climate is
of Land Management. Working in collaboration with
changing in ways that are affecting land management
the Wyoming Wildlife and Natural Resources Trust,
practices. Also, through the widespread application of
established by the Wyoming State Legislature, the
fertilizers and nitrogen-rich effluents into air and water,
fund is used for various kinds of mitigation, including
ecosystems are becoming enriched with nutrients. The
habitat improvement.
environment of the future is likely to be more fertile as
According to Development by Design protocol, some
well as warmer and drier.
sites will have very high conservation value and the
Native plants and animals have evolved tolerances to
best strategy is to avoid the impact. When this happens,
the slowly changing climatic conditions of the past tens
developers are asked whether they could find another
of thousands of years. Can they evolve rapidly enough
place or method for their investment. If that is clearly
to thrive under the changes expected during the next
not possible, a plan is agreed upon to minimize the on-
century? Will mitigation or restoration be feasible with
site impact and mitigate the adverse effects. Mitigation
combinations of precipitation, temperature, and sea-
involves the use of offsets, that is, the reclamation or res-
sonal variations that have most likely never existed
toration of nearby habitat agreed to be in relatively poor
before? Will exotic species be favored over the natives?
condition. Ideally, by minimizing adverse on-site effects
Such topics have been discussed throughout this book.
and providing for offsets, there is a net gain in habitat for
Warmer temperatures and longer summers are likely
the sensitive species in the area. Mitigation is most likely
to extend the inevitable droughts. Fires already have
to succeed if it is done near the affected site.20
become more frequent; water for municipalities, agricul-
The extent to which mitigation is possible is con-
ture, and industry has become more expensive, and the
troversial, but it might involve active reclamation of a
reallocation of limited water supplies is controversial.
The carrying capacity for some rangelands will decline,
for the past few centuries or more are likely to be sorted
and the cost of hay is likely to go up. Large herbivores,
into new ecosystems. 23 Locally, will such species adapt
such as elk and deer, may have difficulty surviving long
rapidly enough to the new climate? Will they become
winters if forage availability on their summer ranges is
rare or extinct because of insufficient habitat regionally?
insufficient due to drought, or if extraordinary stresses
Will traditional forms of land use become impractical?
during migration reduce their limited fat reserves.22
A prudent strategy is to conserve the full range of bio-
Native ecosystems that have attracted and helped sus-
logical diversity wherever possible. If that valuable but
tain westerners for nearly 200 years will change, some-
vulnerable resource can be saved, and water resources
times in ways that are uncomfortable.
can be used more efficiently, then future generations
Some earth scientists have projected that, with climate change, species that lived in the same ecosystems
are more likely to continue benefiting from the amenities and services provided by western ecosystems.
317
Epilogue
The changes that have taken place in western North
development tended to be downplayed; the prevailing
America over tens of millions of years are staggering to
disposition has been simply to use resources. To illus-
comprehendthe sudden extinction of dinosaurs,
trate, an older building on the University of Wyoming
the slow evolution of bison and pronghorn from now-
campus is engraved with the words Strive onthe
extinct ancestors, the gradual transition from humid
control of nature is won, not given. Another building,
tropical forests to sagebrush, and the expansion and
completed 24 years later, in 1950, greets students with
retreat of glaciers. Substantive changes also have taken
The foundation of agriculture is not rooted in soil but
place during the past 15,000 years, since the ancestors
rather in the vision and attainment of men.
of Native Americans arrived. And now, modern civiliza-
But adverse environmental impacts are now so per-
tion has accelerated the rate of change in ways that we,
vasive that they can no longer be ignored. The experi-
the authors of this book, find troubling.
ence of land managers and the research of scientists are
The mountains and plains of the West make up an
providing insights on how to proceed, and Congress has
area for which many residents and nonresidents feel
passed legislation to help ensure the nationwide appli-
responsible. Considering that roughly half of the land
cation of certain restrictions. These laws complicate the
in the region belongs to the American public at large,
lives of resource managers and landowners. Thats inev-
decisions are not made solely at the local level. More-
itable as human pressures increase and many resources
over, the various groups involved, often referred to as
become less readily available. Resource management
stakeholders, have differing perspectives and values.
and economic development will never be as simple as
Debates commonly focus on land management prac-
they seemed to be 50 or 100 years ago. Classroom dis-
tices, the effects of unavoidable disturbances, and the
cussions now probe this complexity, providing a more
reasons that ecosystems respond the way they do. Fre-
realistic perspective than conveyed by a few words
quently the discussions consider strategies for benefit-
etched in stone on buildings constructed when the
ing well into the future from the amenities, services,
worlds population was less than half what it is today.
and natural resources that native ecosystems provide.
Providing a high quality of life for so manyone that
Modern science, technology, and our economic system have greatly augmented the availability of many
can be sustained for the benefit of future generations

is a formidable challenge.
natural resources, often improving the enjoyment of
Wyoming and the Rocky Mountain region in general
life in ways that once seemed sustainable for many
must contribute to the fossil fuels, minerals, wood fiber,
generations. Until recently, the limits to growth and
food, and other resources required for the great human
319
320 Epilogue
endeavor now under way. While providing these com-
of the plants and animals existed a million years or more
modities, however, we should not lose sight of the non-
before Homo sapiens arrivedand we are still learning
material qualities in which this region is so richopen
how to make a living from rugged western landscapes. As
spaces, abundant wildlife, free-flowing rivers, and others.
Aldo Leopold wrote in 1938, the oldest task in human
It is critical that the land be used with care and vision,
history [is] to live on a piece of land without spoiling it.1
so that less tangible but equally valuable resources and
Learning to live gently and sustainably, to be good stew-
services are not lost needlessly. Humans have been a pres-
ards, requires an understanding of both human nature
ence in this part of the biosphere for a short timemost
and the nature of ecosystems.
Appendix A Conversion Table
To convert
Multiply by
LENGTH
Inches to centimeters
2.54
Feet to meters
0.304
Miles to kilometers
1.609
Millimeters to inches
0.0394
AREA
Square feet to square meters
0.0929
Acres to hectares
0.405
Square miles to square kilometers
2.590
MASS
Ounces to grams
28.4
Pounds to kilograms
0.454
Tons (2,000 lb) to metric tons
0.907
VOLUME
Cubic feet to cubic meters
Cubic feet to liters
0.0283
28.3
Quarts to liters
0.946
Gallons to liters
3.78
TEMPERATURE
Fahrenheit to Celsius
5/9 (F 32)
Celsius to Fahrenheit
9/5 (C + 32)
(A difference of 1F is 0.556C; a difference of 1C is 1.8F)

ENERGY
Calories to joules
4.19
321
Appendix B Characteristic Soil Types
Table B.1. Soil orders and subgroups characteristic of vegetation types in Wyoming and adjacent parts of
neighboringstatesa
Soil order
Location/vegetation type
Entisols
Inceptisols
Aridisols
Mollisols
Alfisols
UsHAI
UsNAI
ArAUO
Black sagebrush
TyCOI
TyHAI
Big sagebrush
TyHAI
BoHAI
TyTFE
TyTPE
LOWLANDS
Mixed-grass prairie
Sagebrush steppe
Silver sagebrush
Riparian vegetation
Lowland
TyHAO
Foothill
CuHCO
Montane
HiCAP
Alpine
HiCAP
Saltbush
TyHAI
TyNAI
Mixed
TyNAI
Greasewood
BoNAI
TySOI
TyGOI
Basin grassland
UsHAI
UsNAI
Desert shrublands
322
Characteristic Soil Types 323
Soil order
Location/vegetation type
Entisols
Inceptisols
Aridisols
Mollisols
Alfisols
Sand dunes (stabilized)
TyTPE
Badlands
TyTOE
TySOI
TyHDO
Playas
FOOTHILLS
Shrublands
Mixed
Mountain-mahogany
LiTOE
Threetip sagebrush
TyCOE
TyCCP
TyHCO
Foothill grassland
ArHUO
ArAUO
Oak woodland
TyUOE
Juniper woodland
TyTOE
Ponderosa pine
LiTOE
LiUOE
TyHUA
TyHDA
Woodlands
Douglas-fir
Aspen
TyHAO
TyHDO
MOUNTAINS
Forests
Ponderosa pine
TyHDA
Lodgepole pine
TyCCP
TyHCA
Engelmann sprucesubalpine fir
TyCCP
TyHCA
Douglas-fir
MoHCA
TyHDA
Aspen
TyHCO
AqHCA
Dry
TyHCO
Moist
AqHCO
Wet
HiCAP
TyCAO
Fellfield
HuDCP
Turf
HuDCP
TyDCP
Wet
TyCAE
Subalpine meadows
Alpine tundra
Source: Courtesy of Larry Munn.

a
Subgroup abbreviations are defined in table B.2. A dash indicates that the soil type is absent or not so common. For descriptions of soil
orders and subgroups, see http://soils.usda.gov/technical/classification/tax_keys/.
324
Characteristic Soil Types
Table B.2. Soil subgroup name abbreviations

AqHCA
Aquic Haplocryalfs
TyCAO
Typic Cryaquolls
TyHUA
Typic Haplustalfs
AqHCO
Aquic Haplocryolls
TyCOE
Typic Cryorthents
TyNAI
Typic Natrargids
ArAUO
Aridic Argiustolls
TyCOI
Typic Camborthids
TySOI
Typic Salorthids
ArHUO
Aridic Haplustolls
TyDCP
Typic Dystrocryepts
TyTFE
Typic Torrifluvents
CuHCO
Cumulic Haplocryolls
TyGOI
Typic Gypsorthids
TyTOE
Typic Torriorthents
HiCAP
Histic Cryaquepts
TyHAI
Typic Haplargids
TyTPE
Typic Torripsamments
HuDCP
Humic Dystrocryepts
TyHAO
Typic Haplaquolls
TyUOE
Typic Ustorthents
LiTOE
Lithic Torriorthents
TyHCA
Typic Haplocryalfs
UsHAI
Ustic Haplargids
UsNAI
Ustic Natrargids
LiUOE
Lithic Ustorthents
TyHCO
Typic Haplocryolls
MoHCA
Mollic Haplocryalfs
TyHDA
Typic Hapludalfs
TyCAE
Typic Cryaquents
TyHDO
Typic Hapludolls
Source: Courtesy of Larry Munn.
Notes
Chapter 1: Introduction
1. Hay, including alfalfa, is by far the most abundant crop in
Wyoming and is grown on about two-thirds of the cultivated
land. Also common are sugar beets, corn, beans, wheat, and
barley. Livestock grazing is widespread. Timber harvesting in
mountain forests is a much smaller industry than agriculture.
About 29 percent of the gross state product in 2011 was generated from natural gas, oil, coal, trona, and bentonite, compared
to about 6 and 3 percent, respectively, from tourism and agriculture. See Hamerlinck et al. 2013.
2. Elk are numerous in many mountain ranges, and a small herd
persists in the desert northeast of Rock Springs, near Steam-
6. For further discussions of ecosystem services and their marketable values, see Costanza et al. (1997), Daily and Ellison (2002),
Clark (2007), Burke et al. (2008), Perrings et al. (2011), and
Hansen et al. (2013). For the effects of climate change on biodiversity and ecosystem services, see Frontiers in Ecology and the
Environment (vol. 11, issue 9).
Chapter 2: Landscape History
1. Snoke (1993) provides a detailed synthesis of the geologic history of Wyoming; Johnson and Raynolds (2003) provide an
illustrated overview; for a detailed geology map, see Love and
Christensen (1985).
boat Mountain and the Killpecker Sand Dunes. Grizzly bears
2. Tidwell 1975; some fossils were subsequently lost by erosion.
and wolves now occupy much of their former habitat in north
3. Most Cretaceous volcanism occurred outside the area that
western Wyoming.
3. The change that occurs after ecosystem disturbances is known
would become Wyoming.

4. Lillegraven and Ostresh 1988; Snoke 1993.
as secondary succession. Communities and ecosystems are con-
5. B. Breithaupt, pers. comm.
stantly changing in one manner or another.
6. In 2006, south of Laramie, the fossils of five dinosaurs were dis-
4. Landscapes may be large or small but are spatially patchy. Ecol-
covered in a trench dug for a natural gas pipeline. Three genera
ogists interested in grasshoppers focus on smaller landscapes
were found (Allosaurus, Apatosaurus, and Comarasaurus) in the
than those studying bears and humans. See Turner et al. (2001)
Morrison Formation of the Jurassic (K. Trujillo, pers. comm.).
for an overview of landscape ecology, and Chapin et al. (2012)
The ancestors of pronghorn appeared in the fossil record of
for an overview of ecosystem ecology.
the Miocene, about 20 million years ago; bison did not appear
5. Various organizations now exist for studying and monitoring the
until late in the Pleistocene, less than 300,000 years ago. Bison,
status of rare and endangered species. They employ conservation
mammoths, mastodons, camels, and horses were numerous by
biologists and are found in every state. The Wyoming Natural
20,000 years ago; see Dott and Batten 1981; Axelrod 1985; Hart
Diversity Database, affiliated with the University of Wyoming
2008.
(http://www.uwyo.edu/wyndd/about/index.html), provides ser-
7. Tidwell 1975.
vices to industry, municipalities, conservation organizations,
8. Formerly known as the Cretaceous-Tertiary (K-T) or Cretaceous-
and federal and state agencies. The U.S. Fish and Wildlife Ser-
Paleogene (K-Pg) Extinction Event.
vice administers the Endangered Species Act and provides lists
9. Schulte et al. 2010; one crater has been found in the present-
of endangered, threatened, and candidate species by state. The
day Gulf of Mexico, just north of the Yucatan Peninsula. Other
Wyoming website is at http://www.fws.gov/wyominges/Pages/
explanations include extremely large volcanic eruptions at
Species/Species_Endangered.html.
about the same time in India.
325
326
Notes to Pages 1323

10. Robertson et al. 2004.
26. Petit et al. 1999.
11. Gingerich and Clyde (2001) and Clyde (2001) list more than
27. Mears 1997.
150 genera of mammals known from Paleocene-Eocene fossils
28. Tidwell 1975.
in the Bighorn Basin; Lofgren et al. (2004) list the genera of
29. Current estimates range from 16,000 to 13,500 years ago: http://
Paleocene mammals for North America, based on many fossils
www.nytimes.com/2012/05/20/opinion/sunday/who-arrived
collected in Wyoming and nearby.
-in-the-americas-first.html. Gardner (2008a) provides a sum-
12. Tidwell 1975; Riedl (1959) describes the fossils of the Shirley
mary of human history for the region, focusing on the Red
Basin; Dorf (1964) reports on the fossil trees in Yellowstone
Desert, and Kornfeld et al. (2010) provide an overview for the
National Park; Wing et al. (2005) describe rapid global warming
Great Plains and Rockies; see Janetski (1987) for human history
and vegetation change in the Bighorn Basin during the PETM;
in northwest Wyoming.
Kunzig (2011) includes an artistic rendition of what the plants
30. The Colby Mammoth Site is located in the Bighorn Basin,
and animals in the Bighorn Basin looked like during this time,
near Greybull; see http://www.wyohistory.org/encyclopedia/
with estimates of temperature changes; Eberle and Greenwood

(2012) review the Eocene flora of the Arctic.
13. Pocknall and Flores 1987; the Wyoming State Geological Survey
colby-mammoth-site.
31. Reher and Frison 1980; see http://www.aaanativearts.com/
tribes-by-states/wyoming_tribes.htm.
website. The coal of the Powder River Basin is in the Fort Union
32. Working from boats, paleoecologists take vertical cores from
and Wasatch formations, deposited during the Paleocene and
the bottom sediments and are able to identify the plants that
Eocene epochs, respectively. Coal is found in other parts of

Wyoming as well, but often is under deep sediments or exists
only in thin seams.
14. Wing 1981; Gemmill and Johnson (1997) report on fossils
found in the Great Divide Basin that date back about 58 million
produced the preserved pollen.

33. Baker 1986; Walker 1987; Whitlock 1993; Shuman et al. 2009;
Mumma et al. 2012; Krause and Whitlock 2013.
34. Baker 1986; Whitlock 1993; Licciardi and Pierce 2008; Shuman
et al. 2009; Mumma et al. 2012; Krause and Whitlock 2013.
years (the late Paleocene); Barnosky (1984) described similar
35. Walker 1982, 1987.
vegetation patterns for the Jackson Hole area in the Miocene.
36. Reider 1983, 1990; Reider et al. (1988) describe grassland soils
15. Zachos et al. 2008; McInerney and Wing 2011; though a global
where forests now grow in the mountains; Marcott et al. (2013)
phenomenon, much of the data for the PETM was obtained in
conclude that the present-day global temperature is lower than
the Bighorn Basin of Wyoming, as reviewed by Kunzig (2011).
during the early Holocene but is likely to be higher by the year
16. McInerney and Wing 2011.

17. Jansen et al. 2007.
2100 (see chapter 3).

37. Minckley et al. (2012) present evidence for an earlier warm
18. A portion of the initial Medicine Bow Mountains was later sepa-
and dry period, based on hydrologic fluctuations, archeol-
rated by faulting and is known today as the Sierra Madre; see
ogy, and sand dune activity. See also Forman and Pierson
fig.1.2.
2003; Mayer and Mahan 2004 (Killpecker Dunes); Forman et
19. Sedimentary strata formed at the bottom of these lakes are now
in the Green River Formation, which is surrounded and overtopped by the lake-bordering fluvial and swampy deposits of
the Bridger Formation; Grande 1998; Thompson 2008.
al. 2005; Hart 2008; Halfen et al. 2010 (Casper Dune Field);
Mason et al. 2011.
38. Jackson et al. 2005; see M. E. Buskirk (2009) for a description of
packrat behavior and their nests.
20. Thompson 2008 (p. 111).
39. Lyford et al. 2003.
21. Love et al. 2003.
40. Norris 2006; Lesser and Jackson 2012.
22. Axelrod 1985; Leopold and Denton 1987; Mears 1993.
41. Gray et al. 2006.
23. Two periods of extensive glaciation are widely recognized in
42. Lyford et al. 2002; Jackson et al. 2002.
Wyoming: the Pinedale (about 30,000 to 14,000 years ago)

and the Bull Lake (about 150,000 to 100,000 years ago). For
43. Drying in the early Holocene also is indicated by the pollen

record, as noted previously.
the history of glaciation, see Mears (1997, 2001), Pierce (2003),
44. Fall et al. 1995.
and Licciardi and Pierce (2008). The Pinedale glacial period is
45. Pine expansion is coincident with active sand dunes and low
known in midwestern states as the Wisconsin glacial period.

24. Grasso (1990) describes Lake Wamsutter, located mostly north
water levels in lakes and riverstwo indicators of a dry or drying climate (B. Shuman, pers. comm.).
of Wamsutter, Wyoming. At times it was more than 180 feet
46. Jansen et al. 2007; Gardner 2008b; Marcott et al. 2013.
deep and covered 700 square miles, including Separation Flat,
47. Gray et al. 2004, 2006.
Chain Lake Flat, and Battle Springs Flat. About 10,000 years ago
48. Reider et al. (1988) conclude from soil profile data that fluctua-
the Killpecker Sand Dunes developed on top of a portion of the

lakebed.
tions occurred from about 3,000 to 200 years ago.

49. Some scholars assume that Native Americans had almost no
25. Snoke (1993) and Good and Pierce (2002) describe the forma-
impact on western North American ecosystems, because their
tion of the Yellowstone Plateau, caused largely by volcanic erup-
populations were low and their technology was limited. Others,
tions at about 2.0, 1.3, and 0.6 million years ago.
such as Charles Mann (2005), argue that Native Americans were
Notes to Pages 2337

numerous and had significant effects. Mann concluded that
European diseases decimated Indian populations, which led to
the erroneous impression that their numbers were low. Thomas
Vale (2002) concluded that the greatest human impacts prior
to arrival of Europeans could have been near popular camping
locations on the plains. See also Kornfeld et al. 2010.
50. Gardner 2008b; Beauvais (2008b) indicates that horses were in
Wyoming in the 1600s; Proulx (2008) reviews the history of
EuroAmericans in the Red Desert; Janetski (1987) writes about
Native Americans and the horse in Yellowstone.
51. Proulx 2008.
52. The first authenticated EuroAmerican in Wyoming arrived in
the Powder River Basin in 1805Franois Antoine Larocque,
a French-Canadian fur trader. The La Vrendrye brothers may
have been in the northeast part of the state in 17421743, but
that is not documented.
53. Hart (2008) reviews the history of land use for the western
Great Plains; see also Blair 1987; Crowe 1990; Gardner 2008a.
54. Gardner 2008a; Persico and Meyer (2009) conclude that the significance of valley filling by beaver and gully formation after
dam failure may be exaggerated in some areas. Earlier journals
suggest that gulleys were common before beaver pelts became
highly prized.
55. Reher and Frison 1980; Vorebuffalojump.org.
56. Occasional bison herds of tens of thousands were seen as late as
the 1870s, but by 1882 many Native Americans were starving,
because the bison were so rare; by 1895 only about 800 bison
Chapter 3: Present-Day Environments and Climate Change

1. Curtis and Grimes 2004; the climate of the shortgrass prairie is
described by Pielke and Doesken (2008); the highest precipitation tends to occur at mid-elevations in the mountains.
2. Smith and Geller 1979.
3. Mack and Thompson 1982.
4. Webb et al. 1978; Joyce 1981; plant growth is also known as
net primary productivity and is commonly expressed as grams/
square meter/year.
5. Moist air is brought to eastern Wyoming when the atmospheric
pressure to the south is low. Air flows are counterclockwise at
such times, which brings moist air from the Gulf coast north
and west to Wyoming.
6. Shinker et al. 2010; Wise 2010.
7. Martner (1986) presents data that show how spring and early
summer precipitation is more characteristic of the eastern
plains, whereas the western two-thirds of the state often has
a more even distribution of precipitation during the year (and
a larger proportion that occurs in the winter as snow). See also
Driese et al. 1997; Bates et al. 2006; Schlaepfer et al. 2011.
8. Depending on how drought is defined. Martner (1986) identified droughts in the mid-1930s, mid-1950s, and the early 1960s.
9. Curtis and Grimes 2004 (p. 97); Gray and Andersen 2009;
Woodhouse et al. (2006) observed that Colorado River water
allocation compacts were approved during one of the wettest
periods in the past 500 years; see also Wise 2012.
remained in North America. The last wild bison in Wyoming,
10. This quantity is known as the adiabatic lapse rate; Curtis and
outside of Yellowstone National Park, died in the Red Desert in
Grimes (2004) estimated a drop of 5.5F per 1,000 feet for Wyo-
1889 (Beauvais 2008).

57. The bison of the western Great Plains once numbered in the
millions (Cushman and Jones 1988; Hart 2001).
mings mean maximum temperature in July.

11. Martner 1986.
12. Martner 1986; Ostresh et al. 1990.
58. Dorn 1986.
13. Martner 1986; Curtis and Grimes 2004.
59. Larson 1977; Schullery and Whittlesey 1992; Kornfeld et al.
14. http://www.windpoweringamerica.gov.
2010.
60. Larson 1977; the effects of grazing by domestic livestock along
15. Sala et al. 1988; semi-arid is defined as having an annual precipitation of less than 15 inches.
the Oregon Trail might have been minimal if the animals were
16. Ganskopp 1986.
moved westward during midsummer or fall and if the animals
17. Branson et al. 1967, 1970, 1976. Common salts in the region
grazed the rangeland for only a short time. See also Wentworth
include sodium and magnesium chloride; sodium, calcium, and
1948; Mitchell and Hart 1987; Miller et al. 1993; http://www
magnesium sulfate; and bicarbonates of calcium and magne-
.wyohistory.org/essays/wyoming-cattle-boom.
sium. Saline soils can be defined as any soil with concentra-
61. Mitchell and Hart 1987 (p. 5).
tions of soluble salts that produce an electrical conductivity
62. In January 2011 about 1.3 million cattle and 360,000 sheep
of 4 dS/m or more. Alkalinity refers to soils in which the less
grazed on Wyoming rangelands (USDA National Agricultural
soluble calcium carbonate is important (pH 7.88.2), and sodic-
Statistics ServicesWyoming). Sheep numbers peaked in 1908
ity implies that sodium occupies 15 percent or more of the soil
at about 6 million. See also Abbott and Smith 1955; Larson
adsorption sites. Plant survival on such soils is dependent on
1978.
an ability to maintain cells with a solute concentration higher
63. Johnson 1987.
than that of the soil solution. If this is not the case, water flows
64. No trees are seen in early landscape photos along Crow Creek
out of the plant into the soil, creating water stress even though
near Cheyenne.
the soil is moist.
65. Hart and Laycock (1996) provide a bibliography of repeat-
18. The grazing of large herbivores (such as bison, elk, cattle, and
photo studies in the Great Plains and central Rocky Mountains.
sheep) becomes a disturbance in some grasslands only if too
Meagher and Houston (1998) show repeat photos in Yellow-
many animals are concentrated for too long in an area that is
stone National Park. See also Amundson 1991.
too small.
327
328
Notes to Pages 3756

19. Reported in Thwaites 1905 (p. 311).
20. See Wagner 2009 and online reports of the Intergovernmental
Panel on Climate Change (http://www.ipcc.ch), the National
4. Patten 1968; Starr 1974; Phillips 1977; Chadde et al. 1988;

Baker 1989a,b; Kittel 1994; Girard et al. 1997; Walford et al.
2001.
Climate Assessment program (http://www.globalchange.gov),
5. Houston 1967; Miller 1979.
and The Nature Conservancy (http://www.climatewizard.org/).
6. Akashi 1988; Friedman et al. 1996; Scott et al. 1997; Cooper et
21. See previous note.

22. Shuman 2012.
23. See Shuman 2012; chapter 12.
24. Wagner 2009; Pederson et al. 2010.
al. 1999.
7. Everitt 1968; Irvine and West 1979; Akashi 1988; Cooper et al.
1999; Hultine et al. 2010a,b; Rood et al. 2010.
8. Allred 1941; American elm occurs primarily in the Black
25. Naftz et al. (2002) examined oxygen isotope data in ice cores
Hills and has been killed in most areas on the Great Plains
taken from glaciers in the Wind River Range and concluded
by the introduced Dutch elm disease. Lanceleaf cotton-
that air temperatures had increased about 6F from the mid-
wood is a hybrid between plains cottonwood and narrowleaf
1960s to the early 1990s (about 9F from the mid-1800s to the

early 1990s). See also Naftz et al. 2009.
26. Shuman 2012 (p. 443).
cottonwood.
9. Raynolds 1868 (p. 83); Sacket 1877 (pp. 67).
10. Sun (1986) and K. Johnson (1987) write about the Sweetwater
27. Since Shumans study was completed, there has been one year
River; Cooke (1857, p. 402) about Crow Creek. Dorn (1986)
with high snowfall and one with low snowfall; the trend con-
reviews the journals of early explorers in Wyoming. Widely
tinues (B. Shuman, pers. comm.).
scattered groups of narrowleaf cottonwood were noted along
28. Woodhouse et al. 2006; Lundquist et al. 2009; Mote 2009;

Pederson et al. 2010, 2011; Shinker et al. 2010; Leppi et al. 2011;
Rice et al. 2012.
the Laramie River, at higher elevations in the southern part of

the Laramie Basin.
11. Based on repeat photos in K. Johnson (1987), Baker (1990),
29. Laramie Boomerang, October 19, 2012; more than half (57 per-
Knopf and Scott (1990), Friedman et al. (1996), and others;
cent) of Wyoming had extreme or exceptional drought, and 98
the distribution of cottonwood depends on flooding in some
percent had at least moderate drought.

30. Long-term precipitation data are available from the National
places, but floods also can damage young trees.

12. Akashi 1988.
Climatic Data Center (http://www.ncdc.noaa.gov/oa/ncdc
13. Sedgwick and Knopf (1989) conclude that cottonwood seedling
.html); downward trends are suggested in the Green, Upper
mortality along the South Platte River was caused by insufficient
Platte, Wind, and Yellowstone river basins, but not else-
water due to depth of the water table and by competition from
where in Wyoming. See globalchange.gov for projections of
nearby herbaceous plants. They suggest that cottonwood could
the National Climate Assessment and Development Advisory
be replaced by ash, Russian olive, or eastern juniper. In contrast,
Committee.
31. Anderegg et al. 2010; Stocker 2013. Carbon dioxide concentra-
along the North Platte River in Nebraska, cottonwood regeneration is now abundant and apparently does not depend on
tion in Earths atmosphere is now at or near 400 ppm (Science
the formation of point bars. Woodlands are expanding (W. C.
News, Dec. 28, 2013); it was 280 ppm in pre-industrial times.
Johnson, pers. comm.), reducing the amount of habitat avail-
32. Naftz et al. 2002; Woodhouse et al. 2006; Lundquist et al. 2009;
able for the thousands of sandhill cranes migrating through
Mote 2009; Pederson et al. 2010, 2011; Leppi et al. 2011; Rice et
central Nebraska in late winter and early spring. See also Fried-
al. 2012.
man et al. 1996.
33. Gray and Andersen 2009 (p. 12).
14. Weynand et al. 1979; Miller 1979; Merritt and Cooper 2000;
34. Rehfeldt et al. 2006.
Miller et al. (1995) did a similar study of the Rawhide Wildlife
35. Finch 2012.
Management Area on the North Platte River; Coble and Kolb
36. Friggens et al. 2012 (p. 1).
(2013) studied the effects of regulated and unregulated flows on
37. Williams and Jackson 2007; Williams et al. 2007. For further
the Delores River in Colorado.
reading on the effects of climate change on biodiversity and
15. The importance of floods in riparian landscapes is the topic
ecosystem services, see volume 11 (issue 9) of Frontiers in Ecology
of special issues of BioScience (September 1998) and Ecological
and the Environment.
Applications (June 2001).

16. Houston 1967; narrowleaf cottonwood reproduction is rare or
Chapter 4: Riparian Landscapes

1. Olson and Gerhart 1982; Hubert 2004; Lohman 2004;
McK instry et al. 2004.
absent along the nearby Snake River below Jackson Lake dam,
probably because flood peaks are lower there.
17. Houston 1967; McBride and Strahan 1984; Youngblood et al.
1985; Cooper et al. 1999.
2. The Clean Water Act and Food Security Act place restrictions on
18. Boggs and Weaver (1994) observed this pattern in Montana,
the management of wetlands that are connected to navigable
with the trees sometimes dying after only 60 years; Akashi
waters, whether by surface or groundwater.
(1988) observed a similar pattern along the Bighorn River in
3. Blue spruce is common in the foothill riparian zone along the

Snake River, lower Hoback River, and upper Wind River.
north-central Wyoming; see Miller (1979) for a discussion of

woodlands on the eastern plains.
Notes to Pages 5766

19. Neff 1957.
Medicine Bow Mountains that had been used for floating rail-
20. Rutherford 1954; Call 1966; Skinner et al. 1984; Parker 1986;
road ties from 1868 to 1940. Altered stream channels are still
Hayes 2012.
apparent.
21. Bragg 2000.
41. Stohlgren et al. 1998.
22. Brown et al. 1996.
42. Crops are not considered to be invasive plants, because they
23. D. Randall (1983); McKinstry and Anderson (1999) found that
usually do not displace native plants without some kind of dis-
many land managers are now using beaver for the benefits they
turbance, such as plowing; weeds may or may not be invasive.
provide. See also Cooke and Zack (2008).
The English sparrow and Eurasian collared dove are invasive
24. Beaver are able to obtain construction material for their dams
birds. Except for cutthroat trout, all species of trout in Wyo-
and lodges over distances of 150300 feet (B. H. Smith, pers.
ming are introduced. Among invertebrates, the New Zealand
comm.); Call 1966.
mud snail and a large, stalk-forming diatom are invasive on
25. Skinner et al. 1984.
stream bottoms in northwestern Wyoming (R. Hall, pers.
26. Dorn 1986: Gardner 2008a.

27. Dorn 1986.
28. Haines 1965 (p. 51).
29. Platts 1981; Platts et al. 1983; Kauffman and Krueger 1984;
Schulz and Leininger 1990; Clary et al. 1992.
30. Sedgwick and Knopf 1991; as on the adjacent uplands, grazing
can be a minor factor in riparian landscapes if carefully managed and if it occurs late in the growing season.
31. Platts 1981, 1982; Kauffman et al. 1983; Platts et al. 1983; Kauff-
comm.).
43. Lesica and Miles 2001a,b, 2004a,b; Katz and Shafroth 2003;
Pearce and Smith 2007.
44. Simons and Seastedt 1999; DeCant 2008; Shah et al. 2010;
Mineau et al. 2011; Reynolds and Cooper 2011.
45. Carman and Brotherson 1982; Sher and Quigley 2013.
46. Sexton et al. 2006; Pearce and Smith 2007; DeWine and Cooper
2008, 2010; Merritt and Poff 2010; Nagler et al. 2011.
47. Young et al. 2004.
man and Krueger 1984; Stabler 1985; Brookshire et al. 2002;
48. Ladenburger et al. 2006; Glenn et al. 2012.
McKinstry et al. 2004; Anderson (2007) observes that reduc-
49. Shafroth 2005; Owens and Moore 2007; Nagler et al. 2011;
tions in willow cover near winter feedlots for elk caused a

decline in the willow flycatcher, MacGillivrays warbler, and fox
sparrow.
32. Patten 1968; Kay 1993; Singer et al. 1994; Beschta 2003; Wolf et
al. 2007; Beschta and Ripple 2010.
33. Sedgwick and Knopf (1991) found that livestock preferentially
grazed on recently fallen cottonwood leaves during late autumn
along the South Platte River in Colorado.
34. State Engineers Office, pers. comm.; the Jackson Lake dam on
the Snake River was finished in 1906, enlarging Jackson Lake
and creating the oldest large reservoir in Wyoming (Ostresh et
al. 1990).
35. Marston and Brosz 1990; 51 percent flows to the Missouri River,
32 percent to the Columbia River, 14 percent to the Colorado,
and 3 percent to the Great Basin (Ostresh et al. 1990).
36. A report on the Snake River Levee System is available at http://
www.jhalliance.org/Library/Reports/levee.pdf.
Hultine and Bush 2011.

50. Hultine et al. 2010a,b; the Mediterranean tamarisk beetle (Diorhabda elongata) and related species have been effective; Lesica
and Miles (2004b) found that 3 months of flooding kills salt
cedar in northeastern Montana. For water use by saltcedar compared to native species, see Sala et al. (1996), Smith et al. (1998),
and Sexton et al. (2006).
51. Lesica and Miles 2001a,b; DeWine and Cooper 2008, 2010;
Bunting et al. 2011.
52. Poff et al. 2011; invasive aquatic plants and animals have also
created ecological problems.
53. Allan and Flecker 1993; Banerjee et al. 2013.
54. Pederson et al. 2011; see chapter 3.
55. Shinker et al. (2010) identify periods when the North Platte
River was dry and discuss the implications for Wyoming water
rights agreements with Colorado and Nebraska.
56. Extensive mortality of cottonwood and other riparian trees was
37. Instream flow rights are also recognized on the Wind River
observed during the drought of the 1930s (Ellison and Wool-
Reservation; two rivers in Wyoming have been protected by the
folk 1937; Albertson and Weaver 1944; Friedman et al. 1996);
Wild and Scenic Rivers Act, namely, the headwaters of the Snake
narrow
leaf cottonwood may be more drought tolerant than
River and the Clarks Fork of the Yellowstone River. Less than
plains cottonwood.
1percent of Wyomings rivers and streams have been protected

by this Act and the states instream flow option combined. The
flows of designated instream flow segments are managed by the
Wyoming State Engineer. See Madson 2012; https://sites.google
.com/a/wyo.gov/seo/surface-water/instream-flow.
Chapter 5: Marshes, Playa Wetlands, Wet Meadows, and Fens

1. McKinstry et al. 2004; Mitsch and Gosselink 2007; Keddy 2010;
wetland values are acknowledged by the Clean Water Act and
38. Gribb and Brosz 1990.
provisions in farm bills passed by Congress (Ryan and Squillace
39. D. Brosz, pers. comm.; see chapters 5 and 17 for the importance
2004). See chapter 4 for a discussion of riparian wetlands. As
of irrigation to wetlands.
40. Wohl (2005) describes the history of human impacts on creeks
and rivers in the Rocky Mountains and the challenges of restoration; Young et al. (1994) identify 61 streams in or near the
defined by wetland ecologists, bogs and swamps are not found

in Wyoming.
2. U.S. Department of the Interior, Fish and Wildlife Service,
Washington, D.C. http://www.fws.gov/wetlands/.
329
330
Notes to Pages 6779

3. Copeland et al. 2009, 2010.
4. Peck and Lovvorn 2001; Lovvorn and Hart 2004; Sueltenfuss et
al. 2013.
5. A total of 434 plant species, subspecies, or varieties are considered by the Wyoming Natural Diversity Database as rare
enough in Wyoming to be vulnerable to extirpation (B. Heidel,
pers. comm.). Approximately 20 percent of these plants are
obligate wetland indicators; an additional 10 percent are facultative wetland indicators.
6. Lemly and Cooper (2011) study mosses; Hart and Lovvorn
(2000) report on the importance of algae.
7. Orabona et al. 2009; Lewis 2011.
8. Orabona et al. 2009.
24. Salt flats are a common feature in desert shrublands; see

chapter8.
25. Gage and Cooper 2012.
27. Youngblood et al. 1985; Hansen 1995; Girard et al. 1997; see
chapters 4 and 13.
28. Peatlands are described by Rydin and Jeglum (2006), Mitsch
and Gosselink (2007), and Keddy (2010).
29. More than 200 species of bryophytes have been documented in
the Beartooth Mountains, mostly in fens, including Sphagnum
russowii and S. warnstorfii; see reports available at the Wyoming
Natural Diversity Database website (http://www.uwyo.edu/
wyndd/reports-and-publications/).
9. Twenty-nine species of birds are migrants, mostly shore-
30. Mattson 1984; Lemly 2007; Lemly and Cooper 2011; and reports
birds. Only eight species are considered year-round Wyoming
available at the Wyoming Natural Diversity Database website
residents.
(http://www.uwyo.edu/wyndd/reports-and-publications/),
10. Orabona et al. 2009; S. W. Buskirk, pers. comm.

11. Aquatic beetles, black flies, caddisflies, damselflies, dragonflies,
authored by B. Heidel, M. Jankovsky-Jones, G. Jones, S. Laursen,

E. Rodemaker, and R. Thurston.
fingernail clams, midges, mosquitoes, scuds, snails, water fleas,
31. Collins et al. 1984.
and water skippers; Wollheim and Lovvorn 1995; Gammonley
32. A minimum of 16 inches of peat is a common indicator of fens
2004.
12. For reviews of wetland adaptations, see Rydin and Jeglum
(2006), Mitsch and Gosselink (2007), and Keddy (2010).
in the United States.

33. Cooper and Andrus 1994.
34. See the 1992 report by W. Fertig and G. Jones and the 2000
13. The initial steps of respiration in all species in all environments
report by B. Heidel and S. Laursen, both available at the Wyo-
do not require oxygen, but a smaller portion of the energy
ming Natural Diversity Database website (http://www.uwyo
contained in food is available than in aerobic environments.

Mitsch and Gosselink (2007) and Keddy (2010) review the biogeochemistry of hydric soils.
.edu/wyndd/reports-and-publications/).
35. Lemly 2007; Lemly and Cooper 2011; fens on geothermally
altered sites on the Yellowstone Plateau, and in other parts of
14. Bernal and Mitsch 2012.
North America, are not easily classified using the nutrient-
15. Mitsch and Gosselink 2007.
availability gradient; such fens are acidic and similar to bogs,
16. Lewis 2011.

18. Various studies have focused on wetlands at higher elevations.
They include Briggs and MacMahon (1983); Mattson (1984);
though nutrient concentrations tend to be higher than in bogs.

36. The mosses include Aulacomnium palustre, Drepanocladus aduncus, Philonotis montana, Plagiomnium cuspidatum, Ptychostomum
pseudotriquetrum, and several species of Sphagnum.
Youngblood et al. (1985); Chadde et al. (1988, 1998); Girard et
37. Lemly 2007; Lemly and Cooper 2011.
al. (1997); Marriott and Faber-Langendoen (2000); Walford et
38. Cooper and Andrus 1994.
al. (2001); and reports accessible at the Wyoming Natural Diver-
39. Mitsch and Gosselink 2007; Copeland 2010; Keddy 2010.
sity Database website (http://www.uwyo.edu/wyndd/reports
40. The American Society of Wetland Managers estimates a loss of
-and-publications/), authored by W. Fertig, B. Heidel, M.
38 percent of Wyomings original 2 million acres of wetlands.
Jankovsky-Jones, G. Jones, and others.
Gage and Cooper (2012) state that 3540 percent of the wet-
19. Salt flats are a common feature in desert shrublands; see chapter
8.
lands present in Wyoming, Nebraska, and South Dakota at the

end of the eighteenth century have been lost, but they give no
20. The salt deposits are commonly referred to as alkali and the
estimates for each state. Both estimates include nonriparian
salt flats as alkali flats. However, alkalinity is related to acid-
and riparian wetlands. Dahl (1990) estimates 38 percent wet-
ity, not to the concentration of salts (carbonates, bicarbonates,
land loss for Wyoming between 1780 and the mid-1980s.
sulfates, and chlorides of calcium, magnesium, potassium, and
41. Youngblood et al. 1985; Hansen 1995; Girard et al. 1997.
sodium); Hart and Lovvorn 2000; Gage and Cooper 2012.
21. Holpp 1977; Wollheim and Lovvorn 1995; Brough 1996; Bartz
1997; Bowman 1997; Sanderson et al. 2008; reports available
from the Wyoming Natural Diversity Database website (http://
www.uwyo.edu/wyndd/reports-and-publications/).
22. Keddy 2010.
23.
See reports available at the Wyoming Natural Diversity
Database website (http://www.uwyo.edu/wyndd/reports-and
-publications/).
43. Zedler and Kercher 2004; Zedler 2011.

44. Garrison creeping foxtail has similar effects, though it is introduced; see chapter 17.
45. Kulmatiski et al. 2010; Gage and Cooper 2012; Uddin et al.
2012.
46. Zedler and Kercher 2004; Jarchow and Cook 2009.
47. The Flora of North America (volume 22, 2000) indicates that
narrowleaf cattail has expanded its range and has become more
Notes to Pages 8094

abundant, especially in roadside ditches and other highly dis-
9. Nematodes are a highly diverse group of microscopic round-
turbed habitats. The hybrid (Typha glauca) tends to occur in
worms. Some grassland species feed on plant roots; others feed
such places and is more invasive than narrowleaf cattail.
on fungi, bacteria, or protozoans. Some fungi have evolved
48. Poiani and Johnson 1991, 1993; Johnson et al. 2010; Shinker et
al. 2010.
49. Dise 2009; the dynamics of peatlands are complicated, and the
consequences of climate change are difficult to predict.
adaptations for trapping and digesting nematodes. Stanton

1988; Moore et al. 2008.
10. Weaver 1958,1968; Risser 1985.
11. Nippert and Knapp 2007.
50. Shinker et al. 2010.
12. Singh et al. 1983b; Lauenroth et al. 2008.
51. Copeland et al. 2010.
13. Ode and Tieszen 1980; Risser 1985.
52. Copeland et al. 2010; Sueltenfuss et al. 2013.
14. Boutton et al. (1980) study an elevational gradient from the top
53. Peck and Lovvorn 2001; data were obtained during a relatively
wet year.
of the Laramie Mountains to near Cheyenne.

15. Peden 1976; Vavra et al. 1977; Schwartz and Ellis 1981.
54. Copeland et al. (2010) review threats to the integrity of wetlands
16. Coupland and Van Dyne 1979; the weight of all microbial
in Wyoming. For a summary of federal regulations pertaining to
organisms in the soil is about half that of aboveground plant
wetland protection, see the report by Claudia Copeland (2010).
biomass in a Canadian mixed-grass prairie (Paul et al. 1979).
Also, see the summary prepared by the Colorado Natural Heritage
Stanton (1988) provides data on the relative abundances of
Program at http://www.cnhp.colostate.edu/cwic/regulations.asp.
different kinds of microorganisms. Moore et al. (2008) list
55. National Academy of Sciences 2001; Mitsch and Gosselink 2007
the following functional groups in the belowground food
(chapter 12); Sueltenfuss et al. 2013.
web: predatory mites, nematophagous mites, predatory nematodes, fungivorous nematodes, bacteriophagous nematodes,
Chapter 6: Grasslands
collem
bolas, mycophagous prostigmata, amoebae, flagellates,
and phytophagous nematodes.
1. In Wyoming, buffalo grass is found most commonly in the
17. Sims and Coupland 1979; Singh et al. 1983b; Lauenroth et al.
shortgrass prairie south and east of Cheyenne and sporadically
2008; aboveground plant growth is between 100 and 300 g/m 2/
into Montana. For more information on grassland distribution,
year, but belowground estimates are problematic because of the
see Singh et al. (1983a,b). Lauenroth and Burke (2008) review

the ecology of shortgrass prairie and classify most Wyoming
grasslands as northern mixed prairie.
2. There are occasional small patches of tallgrass prairie on sandy
soils and in some riparian zones on the eastern plains; common plants include big bluestem, Indiangrass, prairie dropseed,
difficulties of measuring root growth.

18. Sala et al. 1988.
19. Sala et al. 1988; Oesterheld et al. 1999; Lauenroth et al. 2008.
20. Coughenour 1991a,b.
21. Knapp and Seastedt 1986; Hunt et al. 1987; Lauenroth and
Burke 2008.
sideoats grama, and switchgrass. Buffalo grass reproduces veg-
22. Milchunas et al. 1988, 2008; Stohlgren et al. 1999.
etatively by stolons, which is unusual in the grass family.
23. Ingham et al. 1985; Knapp and Seastedt 1986; Elliott and Cole-
3. Forbs are herbaceous plants other than grasses and sedges
man 1988; Stanton 1988; Holland and Detling 1990; Burke et al.
(graminoids). Shrubs produce woody stems above the soil sur-
1997, 2008; Frank and Evans 1997; Oesterheld et al. 1999; Frank
face. Very small shrubs, such as fringed sagewort, are sometimes
et al. 2000, 2011.
referred to as half-shrubs. Forbs have more species in grasslands
24. Johnson and Rumbaugh 1981.
than any other growth form.
25. West 1990.
4. Epstein et al. 2002; Morgan et al. 2007, 2011.
26. Woodmansee et al. 1978.
5.
Hudson 1986; see also http://news.sciencemag.org/plants
27. Lauenroth et al. 1978; Lauenroth and Dodd 1978, 1979; Risser
-animals/2013/03/fairy-circle-mystery-solved.
6. Lauenroth et al. (2008) review above- and belowground biomass and productivity in grasslands.
7. Bud dormancy can be broken by changes in the production of
and Parton 1982; Wight and Godfrey 1985; Lauenroth and

Burke 2008; Dijkstra et al. 2010; Carrillo et al. 2012.
28. Houston et al. 1973; Houston and Hyder 1975; Rauzi and Fairbourn 1983; Senft et al. 1985.
growth-stimulating hormones, which often occur when leaves
29. Clark 1977 (p. 1322); Clark et al. 1980; Polley and Detling 1988.
and stems are eaten or are damaged in some other way. The
30. Burke et al. 1987.
buds remain dormant as long as the leaves and stems are intact,
31. Succession following the abandonment of agricultural land is
a process known as apical dominance. Briske (1991) reviews grass
described by Lang (1973) and Reichardt (1982); Peters et al.
morphology and physiology as it pertains to herbivory.
(2008) provide an overview of disturbances in grasslands,
8. Richards and Caldwell (1985) found that most of the energy
including the effects of fecal deposits from large mammals
required for regrowth in two species of wheatgrass was derived
and mortality caused by white grubs (the larvae of June
from current photosynthate by remaining leaves rather than
beetles).
from soluble carbohydrates stored in roots. Lauenroth et al.
32. Painter and Belsky 1993; Lauenroth et al. 1994; Oesterheld et
(2008) report that belowground plant growth is about three
al. 1999; Stohlgren et al. 1999; Lauenroth and Burke 2008; Mil
times more rapid than aboveground in shortgrass prairie.
chunas et al. 2008.
331
332
Notes to Pages 95102

33. Clipping plants uniformly may not give the same results as
57. Wright and Bailey 1980 (for the Great Plains); Bragg 1982 (for
actual grazingwhich typically leaves some part of the plant
central Nebraska); Wendtland and Dodd 1992 (for western
uneaten or severed at different heights. The novel idea that
Nebraska).
plant growth-promoting substances might occur in the saliva
58. Knapp and Seastedt 1986.
of mammalian herbivores has not been supported by most
59. Redman (1978) found more water stress and lower plant growth
research (Detling and Dyer 1981; Detling et al. 1981), although
a year after burning mixed-grass prairie in Saskatchewan, and
sheep saliva may enhance short-term growth under certain
Engle and Bultsma (1984) suggest that increases in plant growth
conditions (Gullap et al. 2011).
should not be expected following burning during drought years
34. Owen and Wiegert 1976, 1981, 1982; Briske 1991.
in western South Dakota. See also Scheintaub et al. 2009.
35. Painter and Detling 1981; Detling and Painter 1983.
60. Fuhlendorf et al. 2012; McGranahan et al. 2012.
36. McNaughton 1985 (p. 286).
61. Branson and Miller 1981; Branson 1985.
37. Heavy grazing by bison has been discussed by Larson (1940),
62. The effects of drought can vary; several investigators reported
Roe (1951), Lauenroth et al. (1994), and Hart (2001). McGinnies
that blue grama and buffalo grass increase in abundance with
et al. (1991) report on the resiliency to drought of grasslands in
extended drought (Albertson and Weaver 1944; Newbauer et al.
eastern Colorado; Laycock (1991) and Cid et al. (1991) review
1980), even to the point of creating a shortgrass prairie. Others
the conditions that favor recovery.
have reported a decline of the same species during similar con-
38. Davis 1959 (p. 4); Sturgis 1884 (p. 6).

39. Quoted in Branson 1985 (p. 4); April 20, 1805, in the Meriwether Lewis journal, published in 1814.
40. Houston 1963; Lauenroth et al. 2009.
41. Van Vuren (1987) and Urness (1989) did not agree with Mack
ditions (in southeast Montana: Ellison and Woolfolk 1937; Reed

and Peterson 1961).
63. Reed and Peterson 1961; Branson and Miller 1981.
64. Turner and Costello 1942; Houston 1963.
65. Weaver 1943, 1954; Newbauer et al. 1980.
and Thompsons (1982) explanation, suggesting instead that
66. Reed and Peterson 1961; Rodel 1977; Watts et al. 1982; the
bison numbers were sometimes low in the Great Basin because
distribution and characteristics of different species of grass
of intensive hunting by Indians. Early journals suggest large
hoppers in Wyoming are summarized by Lockwood et al.
numbers of bison in western Wyoming (L. Whittlesley, pers.
(1993) and Crist (2008). There are approximately 100 species of
comm.; Hart 2001).

42. K. Johnson (1987) includes historical photos; the relative effect
of climate and grazing is discussed by Milchunas et al. (1989).
grasshoppers in the state.

67. Watts et al. 1982 (p. 288).
68. Ellison 1960; Holmes et al. 1979; Quinn and Walgenbach 1990.
43. Weaver and Clements 1938 (p. 470).
69. Rodell 1977; Hewitt and Onsager 1983.
44. Ellison 1960 (p. 52).
70. Allred 1941 (p. 387).
45. Rauzi and Smith (1973) and Gifford and Hawkins (1978) review
71. The quote is from Pfadt and Hardy (1987, p. 183); Crist (2008)
the effects of livestock grazing on water infiltration.

46. Coughenour 1991a,b.
47. Briske et al. 2005, 2006, 2011.
48. McNaughton 1976; several studies have found that the growth
forms of western wheatgrass on prairie dog towns were shorter
and more prostrate than those away from the towns, suggest-
states that the effects of livestock grazing on grasshopper abundance are still uncertain, and the number of insects may be
triggered more by weather and plant growth.
72. Lockwood et al. 2000, 2002; Zimmerman et al. 2004.
73. Dorn 1986; Lockwood 2004; locusts are a form of grasshopper;
Mormon crickets are katydids, not crickets.
ing that some varieties of a single species tolerate grazing bet-
74. Foster and Stubbendieck 1980; Laycock (1958) studies the re
ter than do others; see Detling and Painter 1983; Detling et al.
vegetation of pocket gopher mounds; Peters et al. (2008) and
1986; Jaramillo and Detling 1988; Whicker and Detling 1998.
Stapp et al. (2008) review the effects of small mammals and
49. Coppock and Detling 1986; Krueger 1986; Day and Detling
1990a,b; Fahnestock and Detling 2002; Johnson-Nistler et al.
2004.
insects.
75. Archer et al. 1987; Whicker and Detling 1998; Sharps and Uresk
1990. Peters et al. (2008) report that prairie dog burrows may
50. Mielke (1977) suggests a mutualistic relationship between bison
be 39 feet deep, with 50300 mounds or more per colony.
and pocket gophers, with the bison improving conditions for
Towns can be tens to a thousand acres or more in size. Stapp
the forbs preferred by the gophers. The pocket gophers, he sug-
et al. (2008) and Davidson et al. (2012) provide an overview of
gests, improve soil fertility by their burrowing and feeding,

thereby increasing forage production for the bison.
burrowing mammals in grasslands.

76. Densmore (2011) reviews the history of black-footed ferrets,
51. Kamm et al. 1978; Ingham and Detling 1984.
which were declared extinct in 1979. Two years later, a colony
52. Ingham et al. 1985.
of about 130 animals was found near Meeteetse in the Bighorn
53. Oesterheld et al. 1999.
Basin. That number was soon reduced to 18 by canine distemper
54. Higgins 1984.
and sylvatic plague, for which the animals are now vaccinated
55. Woody plants on the fringe of burning grasslands are some-
when possible. After capture and a period of breeding in captiv-
times ignited and could sustain a fire during a thunderstorm.

56. Moore 1972.
ity, ferrets have been released where large prairie dog colonies
can be found (with about 1,000 ferrets now living in the wild

and 300 still in captivity). One of the first reintroduction sites
95. Morgan et al. 2011.
was the Shirley Basin north of Medicine Bow, where a breed-
96. Morgan et al. 2011 (p. 202).
ing population of several hundred ferrets has been established.
97. Dijkstra et al 2010; Carrillo et al. 2012; Nie et al. 2012.
Each ferret eats about 100 prairie dogs a year. Most present-day
98. Epstein et al. 2002; Morgan et al. 2007.
prairie dog towns are too small to sustain the ferret (Davidson
et al. 2012).
77. Freilich et al. 2003; Miller et al. 2007; Davidson et al. 2012.
78. Stapp et al. (2008) suggest that cultivation has more adverse
effects than shooting or poisoning and that conservation efforts
Chapter 7: Sagebrush
1. Driese et al. 1997.
2. Except in the southeast corner, the grasslands of Wyoming have
could be more fruitful by protecting land where the prairie dogs
been classified as northern mixed-grass prairie; the term steppe
are likely to establish new colonies, such as abandoned prairie
implies large areas of grassland or shrubland, often extending
dog towns. It is important to protect large colonies, as they are

a source of dispersing animals for other towns, and also because
larger towns provide more benefits to the associated species.
79. Pauli et al. 2006.
80. Scott 1951; Kirkham and Fisser 1972; Sneva 1979; Rogers 1987;
Peters et al. 2008.
to the horizon.
3. Houston 1961; West 1983, 1988; Miller et al. 1993; Driese et al.
1997; Bates et al. 2006; Poore et al. 2009; Schlaepfer et al. 2011.
4. Beetle and Johnson 1982; Barker and McKell 1986; Kolb and
Sperry 1999a,b.
5. Tall plants of basin big sagebrush are found in the dunes north-
81. Allen and Knight 1984; Evans et al. 2001; Ehrenfeld 2003;
east of Rock Springs; in the Ferris Dunes north of Rawlins; and
Strayer et al. 2006; Bradley and Marvin 2011. The adaptations
at the base of impermeable granitic domes, where water runs off
of invasive plants can include the production of large volumes
into the adjacent soil.
of seed that germinate earlier in the spring than the seeds of
6. Winward 1970; Shumar and Anderson 1986.
native species, and chemical defense compounds that local her-
7. Sand sage (Artemisia filifolia) and threetip sagewort (A. tripartita)
bivores and parasites cannot tolerate (Pfister et al. 2002).
also are capable of sprouting.
82. Bradley and Marvin 2011; in addition to cheatgrass, drought-
8. Welsh 1957; Robertson et al. 1966; Smith 1966; Myers 1969;
tolerant invasives that could become more common in Wyo-
Zamora and Tueller 1973; Sabinske and Knight 1978; Baker and
ming grasslands include camelthorn, medusahead wildrye,
Kennedy 1985. West of Laramie, black sagebrush is found on
rush skeletonweed, and ventenata grass.
soils with a hardpan at 812 inches, sometimes intermingled
83. Blossey and Notzold (1995) propose the Evolution of Increased

Competitive Ability hypothesis.
84. Strayer et al. 2006.
85. In Nevada: Leger 2008; Goergen et al. 2011; Rowe and Leger
2011; in Wyoming: Mealor and Hild 2007.
86. Blumenthal et al. 2003, 2010; Perry et al. 2010.
87. Blumenthal et al. (2008) added snow water to one plot of
with patches of grassland on more shallow soils.

10. Lundberg 1977; a desert pavement develops when strong winds
remove fine soil particles from the surface, leaving a layer of
stones too heavy for the wind to move.
11. West and Klemmedson 1978; Allen and MacMahon 1985; Longland and Bateman 2002.
mixed-grass prairie and observed an increase in the abundance
12. West and Young 2000.
of invasive plants.
13. Richards and Caldwell 1985; Caldwell et al. 1998; Ryel et al.
88. Miller et al. 2007; see note 76.
2002, 2004; Leffler et al. 2004, 2005; Meinzer et al. 2004;
89. Wyeth 1899; Carter 1857, as reported in Carter 1939.
Scott et al. 2008; Aanderud and Richards 2009; Bleby et al.
90. Roach et al. (2001) discuss the genetic implications of small,
2010; Prieto et al 2011; see also http://en.wikipedia.org/wiki/
isolated prairie dog colonies, finding that in some areas disper-
Hydraulic_redistribution.
sal from one colony to another is good if there is a network
14. Naithani et al. 2012.
of swales and drainages along which dispersing animals can
15. Richards and Donovan 2005; Naithani et al. 2012.
travel; further isolation could lead to inbreeding and a reduc-
16. DePuit and Caldwell 1973.
tion in genetic diversity.
17. Beetle and Johnson 1982; Miller and Schultz 1987.
91. Grassland National Park in Saskatchewan was established in
18. Welch et al. 1991. Water extracts and volatile vapors from big
1981; in 2009 it was designated as the worlds largest Dark Sky
sagebrush leaves are thought to affect plant growth (Hoff-
Preserve.
man and Hazlett 1977; Weaver and Klarich 1977; Ferreira and
92. Freilich et al. (2003) describe how livestock production has
Janick 2004), but until field experiments are conducted, it
changed grassland ecosystems and some of the steps that might
is prudent to assume that the primary adaptation of volatile
be taken to restore some lost features.
oils is to reduce herbivory rather than retard the growth rates
93. Wiens and McIntyre (2008) note that populations of grassland
of neighboring plants. Volatile sagebrush compounds may
birds are declining more than any other group of North Ameri-
reduce the rate of nitrification (West 1991); some evidence
can birds.
suggests that a browsed shrub emits volatile cues that reduce
94. Burke et al. (1995) estimate 50 years for recovery of soil organic
matter.
herbivory (insect or ungulate) on adjacent plants (Karban et al.

2011).
333
334

19. Some herbivores have evolved behavioral and physiological
46. Lommasson 1948; Frischknecht and Plummer 1955; Mueggler
adaptations for minimizing the adverse effects of terpenes and
1956; Robertson et al. 1966; Johnson 1969.
other plant toxins (Launchbaugh et al. 2001); see also Pfister et
47. Lesica et al. 2007; Ewers and Pendall 2008.
al. 2002.
48. Cooper 1953; Anderson and Holte 1981; Uhlich 1982; Anderson
20. Passey and Hugie 1963; Ferguson 1964; Biondi et al. 2007.
21. Robertson et al. 1966; Choudhuri 1968; McDonough and
and Inouye 2001; Ewers and Pendall 2008.

49. Lang 1973; in the Great Basin, Morris et al. (2011) found that
Harniss 1974; Daubenmire 1975; West et al. 1979; Cawker
the effects of cultivation persisted for more than a century.
1980; Romo 1984; Eckert et al. 1986; Meyer and Monsen 1991;
50. Allen et al. 1987; Stahl et al. 1998; Frost et al. 2001; Schuman et
Schuman et al. 2005.
al. 2005; Hild et al. 2006; Boyd and Svejcar 2011.
22. Choudhuri 1968.
51. Ellison and Woolfolk 1937.
23. Lommasson 1948; Robertson et al. 1966; Schuman et al. 2005.
52. Nelson and Sturges 1986; Allen et al. 1987.
Late spring frosts and cold winter temperatures can be fatal to
53. Hanson et al. 1982; Nelson and Tiernan 1983.
seedlings; the development of mycorrhizae on sagebrush roots
54. West 1988; Wallace and Nelson 1990.
facilitates establishment (Allen et al. 1987; Stahl et al. 1998;
55. Neilson et al. 2005; Poore et al. 2009; Finch 2012.
Schuman et al. 2005).
56. Allred 1941 (p. 387).
24. Daubenmire 1975; West et al. 1979; Cawker 1980.
57. Cowan 1929; Allred 1941; Furniss and Krebill 1972; Parmenter
25. Schuman et al. 2005; Hild et al. 2006.
et al. 1987; Miller et al. 1993. Sagebrush is commonly infested
26. Sturges 1975.
with gall-forming flies, but apparently with little detrimental
27. Harniss and Murray 1973; West 1983; Singh et al. 1983b.
effect; the effects of voles are described by Frischknecht and
28. Robertson 1947; Sonder 1959.
Baker (1972).
29. Inouye 2006.
58. Zimmerman et al. 2004; see also chapter 6.
30. Sheets 1958; Tabler 1968; Sturges 1986.
59. Homsher 1960 (p. 192).
31. Grayson 2006; Mack and Thompson 1982; Van Vuren 1987;
60. Miller et al. 1993; Sturges 1993; Mensing et al. 2006; Morris et
Urness 1989; Miller et al. 1993; bison abundance in the inter-
al. 2010; Nelson and Pierce 2010.
mountain basins may have varied from place to place, depend-
61. Baker 2006, 2011.
ing on the amount of grassland available. Large herds of bison
62. Wright et al. 1979.
were noted frequently by explorers traveling through western
63. Young and Evans 1978; Wright et al. 1979; Rennick 1981.
Wyoming in the early 1800s (L. Whittlesley, pers. comm.); see
64. Ewers and Pendall 2008; Cleary et al. 2010.
the journals of James Clyman, Warren Ferris, Zenas Leonard,
65. Cheatgrass, common dandelion, and yellow salsify; Wright et
W. F. Raynolds, Osborne Russell, Robert Stuart, and Nathaniel

Wyeth, summarized in Dorn (1986).
32. Mehringer and Wigand (1990) found that, over the past 5,500
al. 1979.
66. Mihlbachler 1986.
67. Smith et al. 1985; Baker 2006.
years, the abundance of charcoal has been inversely related
68. Chambers et al. 2007; Mack 2011a,b; Rau et al. 2011.
to the abundance of sagebrush pollen and directly related to
69. Bradley 2010; Bradley and Marvin 2011.
the abundance of grass pollen in the Great Basin of Utah. This
70. Tyrer et al. 2007; rush skeletonweed is now found in Idaho.
observation suggests that grasslands burned more often than
71. Young and Allen 1997; West and Young 2000.
sagebrush steppe.
72. Seefeldt et al. 2007; Davies et al. 2009b; Shinneman and Baker
33. Baker 2011.

34. Pearson 1965a.
35. Joyce 1981.
36. Mummey et al. 2002; Bechtold and Inouye 2007; Davies et al.
2007a,b, 2009a,b; Kwon et al. 2008; Wick et al. 2009.
37. Huxman et al. 2004; Prater et al. 2006; Kwon et al. 2008; Cleary
et al. 2010.
38. Havstad et al. 2007; Cleary et al. 2010.
2009; Prevey et al. 2010a,b.

73. The soil may have sufficient cheatgrass seedthe soils seed
bankto produce more than 2,000 seedlings per square yard
(Young and Evans 1975; Hassan and West 1986).
74. Young et al. 1976; Whisenant 1990; forage availability from one
year to the next can be highly variable where annual plants
dominate the vegetation, which creates problems for livestock
and wildlife management.
39. Murray 1975; Carpenter and West 1987; for nitrogen limita-
75. Gill and Burke 1999; Norton et al. 2004; Prater et al. 2006; John-
tions, see Charley (1977) and Doescher et al. (1990); West and
son et al. 2011; Mack 2011a,b; Rau et al. 2011; see Ostoja and
Skujins (1977) observed nitrogen fixation in microphytic crusts.
Schupp (2009) for data on small mammals, which are less com-
40. Matson et al. 1991.

41. Branson et al. 1976; Parmenter et al. 1987.
mon in cheatgrass than in adjacent big sagebrush.

76. Shinneman and Baker 2009; Prevey et al. 2010a,b; Eiseroad and
42. Charley and West 1975; Charley 1977.
Rudd 2011; Owen et al. 2011; Meyer et al. (2007) describe a fun-
gal pathogen that may have possibilities for biocontrol; see also
44. Friese and Allen 1993; Soul and Knapp 1996; Eldridge 2004,
2009.
45. McCulley et al. 2004.
chapter 6.
77. Havstad et al. 2007.
78. Vale 1975; Dorn 1986; Johnson 1987; Miller et al. 1993.

79. Fremont 1845 (p. 49).
80. Preuss 1958 (p. 23); Platt and Slater 1852 (p. 12).
3. Smith and Nowak 1990; West (1983) reports that saltbushgreasewood desert shrublands have a mean net aboveground
81. Bradley (2009) identified the Powder River Basin as an area
primary productivity of 20100 g/m2/year and big sagebrush
where big sagebrush ecosystems are at relatively high risk due
steppe a mean of 80250 g/m2/year. Carbon fixation by biotic
to climate change and other factors.

82. Reed and Peterson 1961.
soil crusts can be considerable when the soil surface is wet.

4. The most common salts in Wyoming desert shrublands and
83. Great year-to-year variation in shadscale-dominated desert
playas are magnesium and sodium sulfates, sodium chloride,
shrubland has been documented in the Great Basin through
and bicarbonates of calcium and magnesium. Less common
photographs taken of the same area annually by Sharp et al.
are calcium and magnesium sulfite salts and calcium sulfate
(1990).
di
hydrate (gypsum). All soils with high salt concentrations
84. Kleiner and Harper 1977; West and Young 2000; Belnap and
can be referred to as saline, with the term alkaline used for soils
Lange 2001; Muscha and Hild (2006) study soil crusts in Wyo-
where the hydroxides cause a pH of 8.4 or above. The term
ming; Shinneman and Baker (2009) found more rapid cheat-
sodic is used for soils where 15 percent or more of the exchange
grass invasion on sites with low cover of soil crusts.
capacity is occupied by sodium. Sodic alkaline soils commonly
85. Mack and Thompson 1982.
have high pH (exceeding 9 in some cases) and low rates of water
86. Van Vuren (1987), Urness (1989), and Byers and Smith (2007)
infiltration and air permeability. The soils of greasewood tend
emphasize the importance of intensive hunting by Indians; see
to be alkaline with a pH of 8 or higher and a conductivity of
also Grayson 2006.

87. About James Clyman: Wyeth 1899; Lee (1916, p. 134); Camp
410 dS/m or greater.

5. The adaptations of halophytes and other desert plants are
1960 (p. 28). About William P. Hunt: Morris 2013 (p. 123);
described by Jefferies (1981), Smith and Nowak (1990), Gure-
Townsend 1839; see also Dorn 1986.
vitch et al. (2002), and Sperry and Hacke (2002).
88. Donahue 2010.
6. Goodin and Mozafar 1972; Detling and Klikoff 1973.
89. Manier and Hobbs 2007.
7. White 1976.
90. Beever 2003; Gardner 2008; De Steigure 2011; Philipps 2012.
8. West 1983.
91. See http://www.blm.gov/wy/st/en/programs/Wild_Horses.html.
9. Nichols 1964; Sharma and Tongway 1973; Romo 1984; Romo
92. See http://gf.state.wy.us/wildlife/nongame/LIP/Sagebrush/index

.asp; see also Welch 2005; Holmes 2007.
93. Baker 2006, 2011 (on fire); Neilson et al. 2005; Madson 2006a,b;
and Eddleman 1985; Ladenburger et al. 2006.

10. Khan et al. (2002) found that greasewood seed germination
declines with increasing salinity.
Meinke et al. 2009; Davies et al. 2011 (on energy development);
11. Hamner 1964; Nichols 1964.
Poore et al. 2009 (on climate change); see chapter 18 for discus-
12. Nichols 1964; Brown 1971.
sion of habitat fragmentation.
13. Flowers and Evans 1966; Steger 1970; Detling and Klikoff 1973;
94. Crawford et al. 2004; Gilbert and Chalfoun 2011; see also http://
sagegrouseinitiative.com/.
95. The sharptailed and dusky grouse also use sagebrush in some
places.
96. Fedy et al. 2011.
Skougard and Brotherson 1979.

14. McGinnies et al. 1976; Bowman et al. 1985; Blaisdell and Holm
gren 1984; Tiku (1976a,b) and Cluff et al. (1983) found that saltgrass tolerates soils with a conductivity ranging from 5 to 25
dS/m and a pH of 710.
97. Blickley et al. 2012.
15. Branson et al. 1976; Baker and Kennedy 1985.
98. Dinkins et al. 2012.
16. Gates et al. 1956; Hamner 1964; big sagebrush has been reported
99.Baker 2006, 2011; Beck et al. 2009; Coates and Delehanty 2010;
Doherty et al. 2011a,b; Hess and Beck 2012a,b.
100.Holloran et al. 2005; Beck et al. 2012; Hess and Beck 2012b;
Kirol et al. 2012.
101.Naugle et al. 2004, 2005; Walker et al. 2007; Knick and Con-
on soils with a pH of 8.4 and a relatively low conductivity of

3.4dS/m.
17. Blaisdell and Holmgren 1984.
18. Beath et al. 1941; Kolm 1975. There are two groups of plants
that accumulate selenium. One group, the selenium indica-
nelly 2011; Dinkens et al. 2012.
tors, requires selenium for growth and includes two-grooved
102.Neilson et al. 2005; Poore et al. 2009.
milkvetch, tineleaved milkvetch, woody aster, and princes
103.Madson 2006a; Doherty et al. 2011a,b; Bush et al. 2011; Dzialak et al. 2011; Knick et al. 2011; Reese and Connelly 2011.
plume. In contrast, the other comprises facultative accumulators, such as Gardners saltbush, gumweed, locoweed,
mentzelia, snakeweed, spiny hopsage, winterfat, and certain
Chapter 8: Desert Shrublands and Playas
grasses. Plants in the latter group do not require selenium

but often develop high concentrations of the element. Some
1. West and Young 2000.
plants do not accumulate selenium even though they are fre-
2. Drenovsky and Richards 2006; Drenovsky et al. 2010; some
quently found on seleniferous soils. Both selenium indicators
shrubs are clearly deciduous, such as shadscale and greasewood;
and facultative accumulators can concentrate large amounts
others are evergreen or facultatively deciduous, such as fourwing
of the element in leaves and stems without adverse effects on
saltbush, Gardners saltbush, spiny hopsage, and winterfat.
the plant itself. For example, two-grooved milkvetch is known
335
336

to accumulate 4,000 ppm or more from soils containing only
1.1 ppm selenium. Grasses and cereal grains with only 1030
Desert shrubs probably would have increased in density during

this time, reducing the growth of the preferred forage plants.
ppm can be lethal or crippling to cattle and horses. Both the
37. Chambers and Norton 1993; Young 1994.
indicators and the accumulators increase surface-soil selenium
38. West and Young 2000; Grayson (2006) concluded that there
concentrations as their shoots decompose in the fall, thereby
would have been large grazers in the late Pleistocene (as discussed
possibly improving conditions for their own growth.
in chapter 2), but for tens of thousands of years, the plants of
19. Harris 1992.

20. Hamner 1964; Hodgkinson 1987.
most intermountain basins probably underwent relatively light

grazing by large mammals, compared to the western Great Plains.
21. Russey 1967; Gibbens 1972; Blaisdell and Holmgren 1984;
39. Muscha and Hild (2006) studied soil crusts in Wyoming;
several studies have found that saltbush shrubland exists on
Shinneman and Baker (2009) report rapid cheatgrass invasion
various soil textures, including sandy loam, sandy clay loam,
where the cover of soil crusts is relatively low.
loam, and clay loam. The soils tend to have low infiltration rates
40. Meyer et al. 2001.
compared to where big sagebrush and blue grama are common,
41. Scott et al. 2010.
probably because of their relatively high clay content or the dis-
42. Shinneman and Baker 2009.
persing effect of sodium on surface-soil particles. A soil pH of 8
43. Rickard 1964.
or above is common. The soils are not always saline, sometimes

having a conductivity of 1.5 dS/m or less. Soil salinity is often
higher under shrubs than between them.
Chapter 9: Sand Dunes, Badlands, Mud Springs, and Mima Mounds
22. Stevens and Monsen 1988.
1. Ahlbrandt 1973; Mayer 2003.
23. Branson et al. 1976.
2. Gaylord 1982.
24. Harr and Price 1972.
3. Prairie sandreed and blowout grass are restricted to dunes in the
25. Rickard 1967.

26. Meyer et al. 2001.
27. West 1991; James et al. (2005) suggest that phosphorus and
magnesium also can be limiting for salt desert shrub growth.
28. West 1990, 1991; Miller et al. 1993; West and Young 2000; Belnap and Lange 2001; nitrogen fixation by soil crusts may be bal-
eastern half of Wyoming, perhaps because of their intolerance

of the drier dunes to the west.
4. Barnes and Harrison 1982.
5. Chadwick and Dalke 1965.
6. Shumway 2000.
7. Wolfe (1973) studied the effects of fire on stabilized dunes;
anced by nitrogen losses caused by denitrification when the soil
chapter 2 of this book reviews the effects of drought.
is wet and anaerobic; for a discussion of biological soil crusts,
8. Weaver and Albertson 1956; Bleed and Flowerday 1990.
see www.soilcrust.org.
9. Pearson 1965b.
29. Currie and Goodwin 1966; Plummer et al. 1968.
10. High infiltration and storage in the Nebraska Sand Hills provide
30. Sharpe and Van Horne 1998.
much of the groundwater contained in the Ogallala aquifer, on
31. Wight and Nichols 1966; Kirkham and Fisser 1972.
which many Great Plains municipalities and farmers depend
32. Allred 1941; Blaisdell and Holmgren 1984.
(Zwingle 1993); see Ashworth (2006) for a modern synthesis of
33. Sharp et al. 1990.
the geologic development and status of the Ogallala aquifer.
34. Wallace and Nelson 1990.
11. Bleed and Flowerday 1990.
35. Succession in desert shrublands subjected to livestock grazing
12. Koske and Polson 1984; Brown 1997.
has been studied in several locations. Predictably, the most pal-
13. Barnes and Harrison 1982; Potvin and Harrison 1984.
atable species tend to decrease in abundance, for example, four-
14. Nitrogen fixation also occurs in the rhizosheaths of some
wing saltbush, spiny hopsage, winterfat, and some grasses. Less
grasses, including Indian ricegrass, prairie sandreed, and
palatable species increase, such as rabbitbrush, shadscale, and
needle-and-thread grass (Bergmann et al. 2009).
weedy species (including Russian thistle). The season of graz-
15. Martin 1992; Clarke and Rendell 2003; Forman and Pierson
ing can be important in determining rangeland response, with
2003; Mayer and Mahan 2004; Kelly et al. 2008; Hanson et al.
early-winter sheep grazing generally having less impact on desirable species than late-winter or early-spring grazing. Ranchers
2009, 2010; Schmeisser et al. 2009; Halfen et al. 2010.

16. Yizhaq et al. 2009.
commonly put sheep on the desert in winter and then transport
17. Sahara mustard is a problem in the dunes of California.
them to mountain rangelands in the summer. With protec-
18. Brown 1971; the Powder River Breaks are found on weakly
tion from livestock grazing, palatable species (perennial grasses
consolidated shales interspersed with lignite seams in the Fort
and forbs) increase, and the abundance of introduced annuals
Union Formation, from which most Wyoming coal is mined.
may decrease or remain about the same. Rangeland recovery in
Vegetation banding caused by abrupt changes in bedrock is
exclosures is variable, which has led to the hypothesis that some
found in various places in Wyoming, such as near Lander and
ecosystems are permanently changed after a disturbance of sufficient intensity and duration (the state transition hypothesis).
36. West and Young (2000) present evidence that the grazing
capacity of desert shrublands had been exceeded by 1900.
Kemmerer.
19. Similar mounds are found in northern Albany County, north of
Laramie (L. Munn, pers. comm.).
20. Both quotes are from Hayden (1879, p. 130).

21. Cox 1984, 2012; Washburn 1988.
22. Betancourt et al. 1991; pinyon pine populations also occur in
22. McGinnies 1960; Hansen 1962.
Idaho (west of Kemmerer) and in northern Colorado (Owl Can-
23. L. Munn, pers. comm.
yon, north of Fort Collins, east of Livermore).
24. Mears 1981, 1987.

25. Spackman and Munn 1984.
26. McFaul 1979.
23. Progulske 1974; K. L. Johnson 1987; Veblen and Lorenz 1991;

see note 27 in chapter 11 for the subtle distinction between fire
exclusion and fire suppression.
27. Reider et al. 1996, 1999.
24. Orr 1975; Shepperd and Battaglia 2002.
28. Arkley and Brown 1954 (p. 198).
25. Tolstead 1947; Potter and Green 1964; Weaver 1965; Brown 1971;
29. Arkley and Brown 1954; Cox 1984 (p. 45) Cox and Allen 1987;
Cox et al. 1987; Washburn 1988; Cox and Hunt 1990; Gabet et
al. 2014.
30. Cox 1990.
31. Burnham and Johnson 2012. The quotation is from Berg 1990
(p. 284).
Hoffman and Alexander 1987; Hansen and Hoffman 1988.

26. Welsh 1957; possibly, ponderosa pine can survive in western
Wyoming, but it has not yet migrated to the suitable habitats
there (see chapter 2).
27. States 1968.
28. Ponderosa pine is found with Rocky Mountain juniper on outcrops of shale that are rich in bentonite, such as near Upton
Chapter 10: Escarpments and the Foothill Transition

1. Kornfeld and Osborn (2003) discuss the ecological and sociological effects of patchy landscapes.
and Moorcroft; elsewhere it is found on outcrops of sandstone

(for example, southwest of Casper and north of Lusk) or granite (for example, west of Cheyenne and Wheatland).
29. Lepper 1974.
2. Hanna 1934; Myers 1969; see chapter 17.
30. Despain 1973, 1990.
3. States 1968; Wells 1970.
31. Gruell 1983; Arno and Gruell 1986; see chapter 11.
4. Johnson 1950; Medin 1960; Miller 1964; Brotherson et al. 1984;
32. Fisher et al. 1987.
the two species of Cercocarpus occur together on the west side of

Flaming Gorge Reservoir.
5. Vlamis et al. 1964; Youngberg and Hu 1972; Lepper and Flesch
ner 1977.
6. Duncan 1975; Young and Bailey 1975; Gruell et al. 1985; Arno
33. Major and Rejmanek (1992) view the two species of serviceberry
as one, citing several taxonomic studies.
34. Brown 1971.
35. Ludwig 1969; Mueggler 1975; Despain 1990.
36. Whitman and Hanson 1939; Hansen and Hoffman 1988.
and Wilson 1986; Johnson (1950) and Miller (1964) report on
37. Sauer 1978.
sprouting and layering, but new stands also become established
38. Welsh 1957; Baker and Kennedy 1985; Knight et al. 1987.
from seed.
39. MacCracken et al. 1983; Uresk and Boldt 1986.
7. True mountain-mahogany is thought to sprout more readily

than curlleaf mountain-mahogany.
40. Neilson and Wullstein 1983.

41. Neilson and Wullstein 1983; Harper et al. 1985.
8. Billings 1954.
42. Bartlein et al. 1997.
9. Wight and Fisser 1968; Spaeth 1981; Waugh 1986.
43. Harper et al. 1985.
10. Soils were shallow in all of Wight and Fissers (1968) 55 study
44. Krebill 1972; Beetle 1974a,b; Gruell and Loope 1974; DeByle
areas, with 3087 percent sand and a pH ranging from 6.8 to
1979; Olmsted 1979; Boyce 1989; Coughenour and Singer 1991;
8.1. Soil conductivity was low (0.451.50 dS/m).
Hessl and Graumlich 2002.
11. Romme et al. 2009.
45. Elk damage mature trees through antler rubbing or eating bark,
which facilitates the spread of fungal diseases, such as aspen
13. Waugh 1986; similarly, Soul et al. (2004) concluded that graz-
canker; Parker and Parker 1983; Veblen and Lorenz 1986.
ing and an increase in nurse shrubs facilitated western juniper
46. DeByle 1979.
expansion in parts of the Great Basin. For further discussion of
47. Veblen and Lorenz 1986.
sagebrush and livestock grazing, see chapter 7.
48. Coughenour 1991a.
14. Burkhardt and Tisdale 1976; Wright et al. 1979; Young and
Evans 1981.
15. Waugh 1986; Miller et al. 1993.
17. Lyford et al. 2003; Romme et al. 2009.
49. Smith 2001, 2012.

50. Vander Wall and Balda 1977, 1983; Ligon 1978; Tranquillini
1979; Tomback 1983; Arno and Hammerly 1984.
51. Bamberg 1971; Hutchins and Lanner 1982; Arno and Hammerly
1984.
18. Baker and Shinneman 2004; Gray et al. 2006; Romme et al. 2009.
19. Gray et al. 2004, 2006; Romme et al. 2009; Breshears et al.
(2005) observed widespread pinyon pine mortality in the
southwest stemming from to global-change-type drought.
20. Knapp and Soul 2008.
21. Welsh 1957; Foster 1968; Everett 1987; Miller et al. 1993; Gray et
al. 2004.
Chapter 11: Mountain Forests

1. Daubenmire 1943b; Knight et al. 1977.
2. MacMahon and Anderson (1982) adopted a general adiabatic
lapse rate of 3F per 1,000 feet, but rates can vary with time of
year.
337
338

3. Smith and Geller 1979; Smith and Knapp 1990.
32. Bentz et al. 2010.
4. Weaver and Perry (1978) found lodgepole pine on both sand-
33. Lotan and Perry 1983; Christiansen et al. 1987; Amman 1989;
stones and shales in the Bridger Range of south-central Montana.

5. Freeland 1944; Parker 1953; Perry 1971; Smith and Knapp 1990.
6. Kyte 1975.
7. Mooney and Billings 1960; Kimball et al. 1973; Salisbury 1984;
Hamerlynck 1992.
Bartos and Amman 1989.

34. The characteristics of high- and low-vigor lodgepole pine trees
are described by Kaufmann and Watkins (1990).
35. Waring and Pitman 1985.
36. Raffa et al. 2008; Bentz et al. 2010.
8. Sakai and Weiser 1973.
9. Knight 1991; Schoettle 1991.
38. The blue-stain fungus does not affect wood strength and is
10. Harvey et al. 1987.

11. Cui 1990.
12. Rochow 1970; Linhart et al. 1981; Knowles and Grant 1983;
Mitton 1985; Rehfeldt 1985; Schuster et al. 1989.
named for the bluish color it gives to the sapwood; some prefer
the more varied grain pattern of beetle wood.
39. Edburg et al. 2012.
40. Knight et al. 1991.
13. Buskirk et al. 1999.
14. Hibernation is a commonly used term, but there is ongoing
42. Regniere and Bentz 2007; Chapman et al. 2012. Preisler et al.
debate about which animals actually have extended slow-wave
(2012) concluded that warmer winter temperatures and drought
sleep and temperature-independent cellular depression over

extended periods of weeksthe definition of hibernation.
15. Meyer et al. (2005) and Dillon et al. (2005) synthesize informa-
were the strongest predictors of a beetle outbreak.

43. Amman 1989; Bartos and Amman 1989.
44. Raffa et al. 2008; Powell et al. 2012; Simard et al. 2012.
tion on the historical range of variability for the forests of the
45. Raffa et al. 2008; Bentz et al. 2010.
Bighorn and Medicine Bow mountains.
46. Klutsch et al. 2009; Kayes and Tinker 2012; Nelson et al. 2014.
16. In contrast, Severson and Kranz (1976) found that understory
The term stand is commonly used for a specific tract of forest.
plant growth in aspen forests was not correlated with tree den-
47. Collins et al. 2011; Diskin et al. 2011; Kayes and Tinker 2012.
sity or basal area.
48. Romme et al. 1986 (Yellowstone); Collins et al. 2011 (Colorado).
17. Kyte 1975; Knight et al. 1977.
18. Hawksworth and Johnson 1989.
50. Pelz and Smith 2012.
19. In the first edition of this book, published in 1994, the mistle-
51. Klutsch et al. 2009; Diskin et al. 2011; Kayes and Tinker 2012.
toe was identified as the single most important problem in
52. Using Daubenmires approach, Wyoming forests have been
lodgepole pine forests. Today bark beetles have that distinc-
divided into 9 general categories (series) and 110 habitat types
tion. See also Hawksworth and Johnson 1989.
and community types, including 9 different kinds of ponderosa
20. Geils and Jacobi 1990; Jacobi et al. 1993.
pine forest, 16 kinds of lodgepole pine forest, and 28 kinds of
21. Smith 1970, 1975; Benkman et al. 1984; Siepielski and Benkman
subalpine forest. Considering such detail is beyond the scope of
2007, 2008.
this book. For descriptions of the Bighorn and Medicine Bow
22. Benkman et al. 2012; see also Smith 1970, 1975.
National Forests, see Meyer et al. (2005) and Dillon et al. (2005);
23. Parchman et al. 2012; Talluto and Benkman 2013.
see also the websites of other national forests. Reed (1971, 1976)
24. Langenheim 1962; Patten 1963; Despain 1973; Veblen and

Lorenz 1986; Green and Conner 1989; Peet 2000.
25. Minckley et al. 2012; Bartlein et al. (1997) discuss the potential
reports on the forests of the Wind River Mountains.

53. Cochran and Berntsen 1973.
54. Bartlein et al. 1997.
effects of climate change on tree species distribution in Yellow-
55. Baker 2009.
stone National Park.
56. Sherriff and Veblen 2006; Brown et al. 2008.
26. For example, Veblen and Lorenz (1986) describe human-caused

disturbances in relation to gold and silver mining in the Front
Range of Colorado.
27. Active suppression of fire did not become consistently effective
57. Baker et al. 2007; Sherriff and Veblen 2007; Schoennagel et al.
2011.
59. Ponderosa pine re-establishment was extremely slow on some
until the mid-1900s in most parts of the West, whereas grazing
steep, mid-elevation, south-facing slopes after a severe Colo-
and other land uses began to limit fire spread as early as the late
rado fire in 1851 that was followed by a relatively dry decade.
1800s. Therefore ecologists usually refer to fire exclusion during
Fire-killed snags were still conspicuous, and ponderosa pine
the past century to account for the effects of both suppression

and land use changes.
28. Heterogeneous fire severity was apparent, for example, in the
1988 Yellowstone fires (Turner et al. 1994) and in the 2002 Hayman fire in Colorado (Finney et al. 2003).
29. Hutto 2008; Saab et al. 2014.
trees were sparse 148 years after the fire (Brown et al. 1999).
60. Brown and Wu 2005; Brown 2006; League and Veblen 2006.
61. Nacify et al. 2010.
62. Patten 1963; Despain 1973; Loope and Gruell 1973; Lanier
1978; Weaver and Perry 1978; Veblen and Lorenz 1986; Goldblum and Veblen 1992; Wyomings Douglas-fir is a different
30. Taylor and Barmore 1980.
subspecies than the Douglas-fir of coastal California, Oregon,
31. Veblen et al. 1991b,c; Brunelle et al. 2008; Raffa et al. 2008.
and Washington. The inland variety does not grow as large as

its northwestern cousin and is more tolerant of a drier, colder
95. Peet 2000.
climate.
96. McDonough 1985; Kay 1993.
63. Arno and Gruell 1986, Goldblum and Veblen 1992; Baker 2009.
97. Mock et al. (2013) found greater genetic diversity in aspen
64. Loope and Gruell 1973; Littell 2002; Korb 2005.
stands than was previously thought to exist, but the patches
65. Amman and Ryan 1991; Donato et al. 2013. The western spruce
of aspen that all leaf out in the spring or turn color at the same
budworm is a moth. In the Rocky Mountains, the larvae feed

primarily on the buds and young leaves of Douglas-fir. Outbreaks appear to occur when many of the trees are in a state of
low vigor.
time in the fall are usually dominated by a single genet.

98. Reed 1971; Youngblood and Mueggler 1981; Brown and DeByle
1989; Bartos et al. 1991; Peet 2000.
99. Shinneman et al. (2013) emphasize the great variability that
66. Korb 2005.
exists among aspen fire regimes in the western United States
67. Green and Connor 1989.
and propose five general types of aspen fire regimes in relation
68. Moir 1969; Despain 1973, 1983, 1990; Whipple and Dix 1979;
to environment and associated species.
Lotan and Perry 1983; Parker 1986; Peet 2000. Lodgepole pine
100.Seager et al. (2013) review the literature on effects of herbi
may be the only conifer species that can tolerate some infertile
vory on aspen forests and emphasize that the relationships are
sites.
69. Romme 1982.
complex, highly variable, and not well understood.

101. Kulakowski et al. (2013) review the wide range of settings in
70. Johnson and Gutsell 1993.
which aspen grows and the equally wide variety of relation-
71. Lotan 1964. Logging slash is sometimes scattered across the
ships with conifers and other woody species. Many early stud-
ground after a lodgepole pine clearcut; the serotinous cones in
ies of aspen were conducted in specific stands over periods of a
the slash open after a few weeks and release seed, which re
few decades, often in places where aspen was replaced by other
generates the stand (Aoki et al. 2011).
species. Recent studies over larger areas often reveal strong
72. Lotan 1975.
aspen persistence. Wiens et al. (2012) review potential pitfalls
73. Tinker et al. 1994.
of using historical conditions as benchmarks for evaluating cur-
74. Muir and Lotan 1985a,b; Schoennagel et al. 2003.
rent ecological conditions.
75. Benkman and Siepielski 2004; Benkman et al. 2012; see note 23.
102. Manier and Laven 2002; Kulakowski et al. 2004, 2013.
76. Muir and Lotan 1985a.
103. Houston 1973; Loope and Gruell 1973; Beetle 1974b; Walters
77. Turner et al. 1997, 2004; Schoennagel et al. 2003.
et al. 1982; DeByle et al. 1987; Boyce 1989; Seager et al. 2013.
78. Kashian et al. 2005.
104. Worrall et al. 2010; Anderegg et al. 2013.
79. Kipfmueller and Baker 2000; Schoennagel et al. 2004.
105.Kashian et al. 2007; but see Worrall et al. (2010) and Forest
80. Westerling et al. 2006, 2011; but see Gavin et al. 2007.
81. Running 1980; Knapp and Smith 1981, 1982; Kaufmann 1985;
Ecology and Management (2013, vol. 299, pp. 1100).

106. Saab et al. 2014.
Smith and Knapp 1990. As discussed in chapter 3, seedlings are

more sensitive to environmental conditions than mature plants
are. In addition to the widespread Engelmann spruce and sub
Chapter 12: The Forest Ecosystem
alpine fir, white spruce is common in the Black Hills, and white
1. Bormann 2000 (pp. 45); see also Chapin et al. 2012.
fir is known from two localities in southwestern Wyoming (in
2. Net primary productivity often is reported for aboveground
the foothills of the Uinta Mountains near Lonetree and on
plant growth only because of the difficulty of estimating the
Little Mountain in Sweetwater County).
growth of roots. Pearson et al. (1987) estimated that total net
82. Sibold et al. 2006.
primary productivity in mature lodgepole pine forests of the
83. Romme and Knight 1981; Johnson and Fryer 1989; Veblen et al.
Medicine Bow Mountains, including roots, is only 2.53.2
1991a.
metric tons/hectare/year. Forests dominated by spruce and fir
84. Bartlein et al. 1997; Minckley et al. 2012.
probably are similar. Somewhat higher productivity would be
85. Schimpf et al. 1980; Pearson et al. 1987.
expected in younger forests (see fig. 12.10); forests on more fer-
86. Aplet et al. 1988.
tile soils, such as on floodplains; or those at lower elevations,
87. Schmid and Hinds 1974; Alexander 1987a; Baker and Veblen
where the growing season is longer. Site index is an indirect
1990; Veblen et al. 1991b,c.
measure of the productive potential of an area and is expressed
88. Ciesla 2011.
as the height of dominant trees at a specified age, typically 50
89. Oosting and Reed 1952; Shea 1985; Veblen 1986a,b; Alexander
or 100 years old. Sites having a higher site index are generally
1987a; Veblen et al. 1991a; Peet 2000.

90. Oosting and Reed 1952; Loope and Gruell 1973; Alexander
1987a; Cui 1990.
91. Veblen 1986a,b.
more productive than those with a lower site index.

3. Fahey (1983) estimated that the amount of decomposition
under snow is about equal to that during the rest of the year;
Brooks and Williams 1999.
92. Kaufmann 1985.
4. Fogel and Trappe 1978; Maser et al. 1978, 1988; McIntire 1984.
93. Aschan et al. 2001.
5. Vogt et al. 1982; many mycorrhizal fungi produce mushrooms,
94. Severson 1963.
the spore-producing part of a fungus.
339
340

6. Allen 1991; Miller and Allen 1992.
costs associated with accumulation of wood and other non-
7. Harder 1979.
photosynthetic tissues reduced the net addition of new bio-
mass. However, recent experiments by Ryan et al. (2004) do
9. For this unit of measure, the volume of water depends on the size
not support this explanation. Instead, multiple mechanisms
of the area being considered. Six inches of water, for example, is
may be involved, including reduced gross primary productiv-
the volume of water that will cover a specified area 6 inches deep.
ity, reduced canopy leaf area, and reduced growth efficiency
10. Knight et al. 1985; Fahey et al. 1988.

11. In lodgepole pine forests of the Medicine Bow Mountains,
(amount of wood produced per unit of resource used).

32. Smithwick et al. 2009a,b.
Knight et al. (1985) found that transpiration commonly
accounts for 5061 percent of total annual evapotranspiration,
34. Minshall et al. 1997; Rhoades et al. 2011.
944 percent of which occurs during the spring drainage period
35. Chapin et al. 2002.
while snow still covers the ground (vernal transpiration). Esti-
36. Minshall et al. (1997) report that nitrogen levels in streams
mated vernal transpiration and outflow varied considerably
draining steep terrain underlain by friable volcanic substrates
among the stands that were studied, with vernal transpiration
were initially elevated after the 1988 Yellowstone fires, but post-
accounting for 420 percent of the snow water.
fire levels in other streams draining gentler topography were
12. Grier and Running 1977; Binkley et al. 1995.
barely detectable; see also Turner et al. 2007.
37. Smithwick et al. 2009a,b.
38. Pearson et al. 1987; Knight et al. 1991.
15. Kaufmann 1985; Kaufmann et al. 1987; Pataki et al. 2000.
39. A study in the Greater Yellowstone Ecosystem (Romme et al.
16. Jaynes 1978; Knight et al. 1985.
1986) found that annual wood production per unit area returned
17. Gary and Troendle 1982; Troendle 1983, 1987; Knight et al. 1985;
to previous levels or was even higher 1015 years after a bark
Troendle and Kaufmann 1987; but see Biederman et al. (2012)
beetle outbreak. Also, Brown et al. (2010) concluded that some
and the subsequent discussion of bark beetles in this chapter.
stands in British Columbia were still carbon sinks even during
18. MacDonald and Stednick 2003.
the first several years after severe bark beetlecaused mortality;
19. Eighty percent of the roots are in the top 8 inches of the soil.
and Pfeifer et al. (2011) projected that the amount of ecosystem
20. Gary and Troendle 1982.
carbon in Idaho forests would recover in 25 years or less after a
21. Nsholm et al. 1998.
bark beetle outbreak. Litton et al. (2003) studied the importance
22. Fahey et al. 1985; Knight et al. 1985; Fahey and Knight 1986.
of soil microbes and organic matter in the carbon budget.
23. Gorham et al. 1979.
40. Edburg et al. 2012; Pugh and Gordon 2012.
24. Harmon et al. 1986.
41. Pugh and Small 2012. Clow et al. (2011) observed higher soil
25. Gosz 1980; Fahey 1983; Prescott et al. 1989; Knight 1991.
moisture beneath beetle-killed trees in northern Colorado,
26. Fahey 1983.
which they interpreted as reduced evapotranspiration; similar
27. Some changes caused by clearcutting are similar to those caused
changes were documented in lodgepole pine forests of British
by fire. Stottlemyer and Troendle (1999) studied the effects of
Columbia by Boon (2012).
clearcutting on snowpack, soil solution chemistry, and nutrient
42. Biederman et al. 2012; Pugh and Gordon 2012.
export on the Fraser Experimental Forest in Colorado.
28. The proportion of the surface soil that is sterilized by the heat
of fires in the Rocky Mountain region is small and is soon mitigated by natural processes that facilitate occupancy by new
organisms.
44. Morehouse et al. 2008; Clow et al. 2011; Griffin et al. 2011; Griffin and Turner 2012.
45. Mikkelson et al. (2013a,b) found that mountain pine beetles
caused some changes in streamwater chemistry.
29. Turner et al. (2004) found that, in young forests 10 years after
46. Ecosystems become a source of carbon dioxide when the carbon
the 1988 Yellowstone fires, approximately 68 percent of the
fixed by photosynthesis is less than the carbon released by the
burned landscape was characterized by an aboveground net
respiration of plants and all other organisms.
primary production of less than 2.0 tons/hectare/year; 22 per-
cent by 24 tons/hectare/year; and 10 percent by greater than
48. Troendle 1987; Troendle and Nilles 1987; Prescott et al. 1989a;
4 tons/hectare/year. The tree stratum accounted for most of the

leaf area and productivity in almost all stands, even in those
with low tree densities.
Knight et al. 1991.

49. Intense fires in some areas can lead to a water-repellent surface,
which reduces infiltration and increases surface runoff and
30. Kashian et al. 2005. In the Medicine Bow Mountains, Pearson
possibly erosion. In burned Rocky Mountain forests, water-
et al. (1987) estimated that maximum total net primary produc-
repellent soils tend to be patchy, cover a small area, and remain
tivity rates of 2.53.2 tons/hectare/year are reached in stand

ages of 4060 years.
31. It was long thought that the decline in net primary productiv-
repellent only for several months at most.

50. Harmon et al. 1986; Maser et al. 1988; Tinker and Knight 2004;
Krzyszowska-Waitkus et al. 2006.
ity as forests age was because, although photosynthesis contin-
51. Tinker and Knight 2004; Krzyszowska-Waitkus et al. 2006
ued to produce sugars at previous levels, increasing respiration
52. Tinker and Knight 2001, 2004.

53. Romme and Knight 1982; Minshall et al. 1989.
79. Dillon et al. 2005; Meyer 2005.
54. Alexander 1986, 1987a,b.
80. Fettig et al. 2007.
55. Harmon et al. 1986; Bull et al. 1997.
81. Raffa et al. 2008.
56. There is growing interest in the idea of emulating natural distur-
bance patterns and processes in silvicultural operations (Kohm
83. Worrall et al. 2010.
and Franklin 1997; Perera et al. 2004; Dillon et al. 2005; Meyer
84. Anderegg et al. 2013.
et al. 2005). Because the ecosystems are adapted to these kinds
of disturbances, they are likely to respond well to anthropo
86. Zier and Baker 2006.
genic disturbances that mimic the natural disturbances. Other
87. See http://forest.moscowfsl.wsu.edu/climate/species/index.php
kinds of anthropogenic disturbances that have no natural prec-
for a summary of research done by Gerald Rehfeldt and his
edent are more likely to result in undesirable responses.
associates; see also Bartlein et al. 1997.
57. Knight et al. 1991; Parsons et al. 1994; killing 60 percent of the
88. Millar et al. (2007) describe approaches for adapting to climate
trees by girdling reduced leaf area by 43 percent and led to an
change, such as by conserving ecosystems that are resilient to
estimated 92 percent increase in water outflow compared to
climate impacts or assisting the migration of threatened species
clearcutting, which increased water outflow by 277 percent; see
to new habitats. They also suggest ways to mitigate climate
also Biederman et al. 2012; Rhoades et al. 2013.
change, such as by increasing carbon storage in forests, to the
58. Coops and Waring 2011.
extent possible.
59. Romme and Turner 1991.

60. Rehfeldt et al. 2006; see also chapter 16.
61. MacDonald and Stednick 2003.
62. Westerling et al. (2006) and Wotton et al. (2010) write about
fire; see Raffa et al. (2008) for more on beetle outbreaks.
63. Simard et al. 2012; Mitton and Ferrenberg (2012) found that
mountain pine beetles in Colorado now produce two generations per year rather than one, due to climate warming. For a
specific example and a review of the literature, see Raffa et al.
2008.
64. Powell et al. 2012. A somewhat different conclusion might be
Chapter 13: Mountain Meadows and Snowglades

1. Debinski et al. 2000.
2. Ellison 1954; Starr 1974; Windell et al. 1986; Debinski et al.
2000.
3. Dunnewald 1930; Daubenmire 1943a; Jackson 1957; Hurd 1961;
Patten 1963; Despain 1973; Dunwiddie 1977; Peet 2000.
4. Dunnewald 1930; Behan 1957; Patten 1963; Cary 1966; Miles
and Singleton 1975.
5. Miles and Singleton 1975; Vale 1978.
reached for ponderosa pine or Douglas-fir forests, where greater
6. W. K. Smith et al. 2003; see also Sibul (1995) on Dry Park.
numbers of moderately injured trees often remain after the
7. Munroe (2012) found no clear differences in the soil par-
fires of variable severity that typically characterize these forest
ent material under the meadow and forest. He classified the
types.
meadow soils as Typic Humicryepts and the forest soils as
65. Bentz et al. 2010; little is known about the fir bark beetle.
Inceptic Haplocryalfs, and assumed the differences could have
66. Hicke et al. 2012; see also Klutsch et al. 2011; chapter 11.
developed since the forest disturbance that created the park.
67. Compare Simard et al. 2011 and Schoennagel et al. 2012.
See also Coop and Givnish 2008.
68. See Simard et al. 2011; Schoennagel et al. 2012.
8. Buckner 1977.
69. Lynch et al. 2006; the spatial pattern of burning in the 1988
9. Httenschwiler and Smith 1999; Moir et al. 1999.
Yellowstone fires was significantly correlated with the spatial pattern of a previous mountain pine beetle outbreak that occurred
10. Bekker and Malanson (2008) review the types and causes of ribbon forests.
during 19721975, with the odds of burning increasing by 11
11. Billings 1969 (p. 204).
percent in areas of previous beetle activity compared with areas
12. Arno and Hammerly 1984; Httenschwiler and Smith 1999;
without beetles. However, no correlation was found between

burning and more recent beetle activity during 19801983.
Bekker and Malanson 2008; Bekker et al. 2009.

13. Buckner 1977; Bekker et al. 2009.
70. Kulakowski and Jarvis 2011; Black et al. 2013.
14. Bekker and Malanson 2008.
71. Westerling et al. 2006; Raffa et al. 2008; Logan et al. 2010.
15. Coughenour 1991a,b; Coughenour and Singer 1991.
72. Wallace 2004; Romme et al. 2011.
16. Stohlgren et al. (1999) studied grazing exclosures in the Rocky
73. Millspaugh et al. 2004.
Mountain region, finding that plant species richness was just as
74. Coops and Waring 2011.
great inside exclosures as outside and that grazing in their study
75. Firewise guidelines are available from many sources, includ-
area had no or little effect on the presence of exotic species.
ing http://www.firewise.org/resources/files/Around-Your-Home
17. Billings 1969; Moir et al. 1999; W. K. Smith et al. 2003.
.pdf.
18. Lynch (1998) studied Fish Creek Park in the Wind River Moun-
76. Cohen and Stratton 2003; for a booklet titled Living with Wildfire in Wyoming, see www.uwyo.edu/barnbackyard.
tains; see chapter 2.

19. Jakubos and Romme (1993) found no charred stumps or other
77. Schoennagel and Nelson 2011.
wood in the meadows they studied; Munroe (2003) worked in
78. Finney 2001.
the Uinta Mountains.
341
342

20. Anderson and Baker (2005) could not rule out the influence of
(Indian paintbrush). Such plants are known as alternate hosts;
livestock grazing in the Medicine Bow Mountains, as Jakubos
see Zambino et al. 2005; McDonald et al. 2006; http://www
and Romme (1993) could in Yellowstone National Park.

22. Forrest et al. 2010.
.fs.fed.us/rm/highelevationwhitepines/Threats.
17. Resler and Tomback 2008; Keane and Schoettle 2011; Larson
2011.
23. Harte and Shaw 1995; Shaw et al. 2000.
18. Billings 1973; Pw 1983.
24. Price and Waser 2000.
19. Smith and Geller 1979; Tranquillini 1979; Hadley and Smith
25. Price and Waser 2000.

1986.
20. Scott 1995.
21. Exceptions are the small annual plants studied by Reynolds
Chapter 14: Upper Treeline and Alpine Tundra

1. Peet 2000; Krner 2012; Daubenmire suggested in 1954 that
alpine treeline drops approximately 350 feet per degree of latitude between 30 and 60 north latitude.
2. Daubenmire 1954; west slopes tend to have more snow, because
(1984) in the Beartooth Mountains: Koenigia islandica, Poly

gonum confertifolium, and P. douglasii. Also, see Forbis (2003),
who found abundant seedlings of some plants in the alpine
tundra on Niwot Ridge in Colorado.
22. Terjung et al. 1969; Tieszen and Detling 1983; Bowman and Fisk
2001.
most storms come from the west, with precipitation occurring
23. Scott and Billings 1964; Sakai and Otsuka 1970; Kimball et al.
as the air rises on the western slopes. Generally, annual precipi-
1973; Wallace and Harrison 1978; Bell and Bliss 1979; Hamer-
tation is less on the leeward, east-facing slopes, stemming from

the rainshadow effect (see chapter 3).
3. Whitebark pine is a close relative to limber pine, with both having five needles per fascicle and many of the same ecological
characteristics; Driese et al. (1997) calculated a mean elevation
of 9,725 feet for upper treeline in Wyoming.
lynck 1992.
24. Krner 2012.
25. Rochow 1970.
26. Spomer 1964; Campbell 1997 (plants); Molenda et al. 2012
(insects).
27. Knutson 1981.
4. W. K. Smith et al. 2003.
28. Armstrong et al. 2001; Dearing 2001.
5. Krner (2012) describes factors that limit growth at treeline,
29. Huntly 1987.
whereas W. K. Smith et al. (2003, 2009) and Wieser et al. (2014)
30. Huntly and Inouye 1988; Armstrong et al. 2001; Seastedt 2001.
focus on factors limiting seedling establishment.
31. On Niwot Ridge in Colorado, the plant communities are classi-
6. Layering is the process whereby a new tree develops when low
fied as fellfield, dry meadow, moist meadow, wet meadow, late-
branches of spruce and fir produce roots while pressed to the soil
melting snowbank, and shrub tundra (Walker et al. 2001); see
surface during the winter by heavy snow. This is an example of

vegetative reproduction that results in a clone. See fig. 11.18.
7. Marr 1977; Benedict 1984; Holtmeier and Broll 1992; Seastedt
Thilenius and Smith 1985; Moir et al. 1999.

32. Billings (2000) suggested three terms that now are widely used
when describing vegetation patterns in the alpine tundra:
2001 (on soils); Seastedt and Adams 2001.
macrog radients, microgradients, and mesogradients. See also
8. Seastedt et al. 2004; Hiemstra et al. 2006.
Johnson and Billings 1962; Oberbauer and Billings 1981; Litaor
9. Griggs 1946; Daubenmire 1954; Wardle 1968; Tranquillini

1979; Arno and Hammerly 1984; Krner 2012.
et al. 2002, 2008.

33. Though freeze-thaw cycles still affect alpine tundra, some
10. Krner 2012; previously treeline was thought to occur where
patterned ground may be remnants of the Pleistocene, as sug-
the July mean temperature is less than 50F (Tranquillini 1979;
gested by solifluction terraces found in mountain foothills
Arno and Hammerly 1984).
and soil polygons in some intermountain basins (see chapter
11. Smith and Carter 1988.

12. Bristlecone pine also has flexible branches and is found in Colorado, northern New Mexico, and westward.
3). See Richmond (1949) for an early study in the Wind River
Mountains.
34. Billings and Mooney (1959) studied cryoturbation in the Medi-
13. Hadley and Smith 1986, 1987, 1989.
cine Bow Mountains, where in a few square yards the tundra
14. Eversman 1968; Potter 1969; Mears 1975; Malanson and Butler
changed from a frost hummock to a sorted polygon and back
1984; Butler 1985; Johnson 1987; Patten and Knight 1994.

15. Hoff 1992; Tomback et al. 2001; Keane and Schoettle 2011; white
to another frost hummock. Phillips (1982) observed similar

patterns associated with stone polygons in Rocky Mountain
pine blister rust causes swelling on the branches and trunk, even-
National Park.
tually killing the tree. In general, the disease infects five-needle
35. Willard et al. 2007.
pines, known collectively as white pines; see chapter 15. Bristle-
36. Bowman et al. (2006) concluded that changes in nitrogen
cone pine is found in central Colorado but not in Wyoming.

16. Pathogenic fungi known as rusts often require two unrelated
uptake after fertilization occur partially because of shifts in

species composition.
species of host plants to complete their life cycle. For white
37. Bowman and Fisk 2001; Monson et al. 2001; as in forests, some
pine blister rust, the fungus requires plants in the genus Ribes
nitrogen is absorbed by the roots in organic forms, such as the
(wild currant or gooseberry), Pedicularis (lousewort), or Castilleja
amino acid known as glycine.

38. Baron et al. 1994; Baron and Campbell 1997; Stottlemeyer et al.
15. Strong 1968 (p. 51).
1997; Heuer et al. 1999; Burns 2003, 2004.
16. Janetski 1987 (chapters 2 and 3); Frison 1991; Kornfeld et al.
39. Fenn et al. 2003; Porter and Johnson 2007.
2010; artifacts found on the northern end of Jackson Lake doc-
40. Initially the primary source for nitrogen was thought to be the
ument hunting camps dating back about 2,500 years.
Denver metropolitan area and large agricultural operations to
17. Schreier 1982.
the east, including feedlots. However, further research sug-
18. Haines 1965.
gested that much of the atmospheric nitrogen is in the form of
19. Despain 1990.
nitrous oxides coming from the Salt Lake City area, as reported
20. Despain 1983, 1990.
by Sievering (2001).
21. Despain 1990.
41. Fenn et al. 2003; Hood et al. 2003; Seastedt et al. 2004; Liptzin
and Seastedt 2010.
42. Fenn et al. 2003.
22. Chadde et al. 1988.

23. Patten 1959; Despain 1990.
24. Mattson 1984; Lemly 2007.
43. Timothy Seastedt, pers. comm.
25. Chadde et al. 1988; Chadde and Kay 1991.
44. Carrera et al. (1991), who calculated that the upper treeline
26. Sheppard 1971.
could rise 260460 feet with less than a 2F increase in the
27. Fryxell 1978.
mean summer temperature; Lynch 1998; Elias 2001; Toney and
28. Righter 1982.
Anderson 2006.
29. Palmer 1991.
45. Billings (1969) found evidence of fire at treeline in the Medicine

Bow Mountains, which had the effect of lowering the treeline,
because new tree establishment was slow.
46. W. K. Smith et al. 2003, 2009.
47. Kupfer and Cairns 1996; Malanson et al. 2007; Loffler et al.
2011.
30. Schreier 1982; Righter 1982.

31. Greater Yellowstone Coordinating Committee report of 1987,
available at http://fedgycc.org/index.html; Youngblood and
Mueggler 1981.
32. Steele et al. 1983.
33. Oswald 1966; Love et al. 2003.
48. Weisberg and Baker 1995a,b.
34. Loope and Gruell 1973.
49. Baker and Weisburg 1995, 1997.
35. Sabinske and Knight 1978.
50. W. K. Smith et al. 2003.
36. Blackwelder 1912; Voight 1982; Lawrence and Lawrence 1984.
51. Forbis 2003; Elliott and Kipfmueller 2010, 2011.
37. Pritchard 1999; Schullery 2004.
52. Munroe 2003; Stinson 2005; Cannone et al. (2007) worked in
38. Schullery 2010. Comstock wrote about the value of wildlands
the Alps, finding rapid upward treeline advance.
for evolution in the 1870s, at a time when such ideas were new
53. Inouye 2008; Wipf et al. 2009; Loffler et al. 2011; in Alaska,
and controversial. Wright and Leopold championed predators
yellow cedar is dying in some places because shallow roots are
in the 1930s. George Bird Grinnell described Yellowstone in the
killed by frost, due to reduced insulation from snow (Hennon et
late 1800s and early 1900s as a reservoir of water and wildlife
al. 2012).
which could sustain surrounding lands.

39. Leopold et al. 1963.
Chapter 15: The Greater Yellowstone Ecosystem
40. Schullery 2004.

41. The decade of the 1870s probably was the period of greatest
1. Eversman and Carr 1992; Hansen et al. 2002. Also see the Atlas
slaughter in and around YNP (Schullery 2004). The first super-
of Yellowstone (Marcus 2012) for a collection of maps pertaining
intendent of YNP, Nathaniel Langford, had almost no budget
to geography, wildlife, and ecology.
or staff and visited the park only twice during his five years
2. Schreier 1983; Good and Pierce 2002; Mogk et al. (2012) provide
a geologic map for YNP.
as superintendent; meanwhile, tourists and local residents

exploited park resources essentially without constraint. In 1886
3. Dorf 1964.
a troop of U.S. Cavalry was sent to protect YNP; the Cavalry
4. Good and Pierce 2002.
managed the park until creation of the National Park Service in
5. Love et al. 2003.
1916 (Schullery 2004).
6. Love et al. 2003.
42. Houston 1982.
7. Good and Pierce 2002.
43. Pritchard 1999; Schullery 2004.
8. Good and Pierce 2002; Love et al. 2003.
44. Kay 1993.
9. Whitlock 1993; Love et al. 2003; Millspaugh et al. 2004.
45. Ecologists Andrew Hansen and Jay Rotella (2002) found that the
10. Marston and Anderson 1991.
greatest diversity and abundance of birds in the GYE are associ-
11. Haines 1996; Schullery 2004.
ated with aspen, cottonwood, and willow habitats. In another
12. Helena Daily Herald, November 9, 1870 (p. 1).
study, Berger et al. (2001) compared bird communities in Grand
13.
Haines 1996; http://www.cr.nps.gov/history/online_books/
Teton National Park, where moose were relatively abundant
haines1/iee2d.htm.
14. Meagher and Houston (1998) compared modern photographs
in YNP with those taken by early explorers.
and fed heavily on willows, to nearby national forests, where

moose numbers and browsing intensity were lower. Species
richness and abundance of nests were greater in the national
343
344

forests. Also, two riparian specialists, the gray catbird and Mac
Gillivrays warbler, were absent in the park, where moose densities were high, but were present in the national forests.
61. Yellowstone National Park 1997; National Academy of Sciences

2002.
62. Schullery 2004.
46. Schullery and Whittlesey 1992.
63. Despain 1990; Renkin and Despain 1992.
47. D. W. Smith et al. 2003.
64. Schullery 1989.
48. Smith 2001, D. W. Smith et al. 2003; Kilpatrick et al. 2009;
65. Christensen et al. 1989.
Cross et al. 2010; Bienen and Tabor 2006; brucellosis and other
66. Romme and Despain 1989.
diseases have not been a problem for elk and bison.
67. Millspaugh et al. 2000, 2004; Higuera et al. 2011.
49. Chase (1986), Bonnicksen (1989), and Wagner (2006) were
68. Shullery 1989; several other fires also were ignited by humans,
major critics of park policy; Pritchard (1999) reviews some of
or were outside YNP boundaries, or both, and contributed to
the controversy.
50. Elk are not the only wildlife species adversely affected by land
the total acreage burned in the GYE in 1988.

69. Brown 1991.
use changes outside park boundaries. Hansen and Rotella (2002)
70. Wallace 2004; Romme et al. 2011.
analyzed nesting success of yellow warblers, neotropical migrants
71. Kay 1993; Romme et al. 1997, 2005; Stevens et al. 1999.
that nest in deciduous woodlands of aspen and cottonwoods,
72. Schullery 2004.
both inside and outside YNP. Woodlands in the park commonly
73. Westerling et al. 2006.
contained nesting yellow warblers, yet those birds often failed
74. Westerling et al. 2011.
to produce adequate numbers of surviving offspring to permit
75. For example, species that are now rare or absent in the GYE,
population growth because of low summer temperatures at the
such as ponderosa pine, could become more common, as noted
relatively high elevations that characterize park habitats. They

concluded that the park by itself cannot sustain a population of
by Bartlein et al. (1997) and Westerling et al. (2011).

76. Tomback et al. 2001.
the warblers; the birds that are found in the park have come from
77. Logan et al. 2010; Bockino and Tinker 2012.
more productive habitats elsewhere. Yellow warblers nesting in
78. Tomback and Achuff 2010; the rust kills other five-needle pines
woodlands at lower elevations, on private lands outside the park,

could potentially produce positive numbers of offspring under
also, such as limber pine (see chapter 14).

79. Bockino and Tinker 2012.
the warmer summer conditions in these habitats. However,
80. Logan et al. 2010; Tomback and Achuff 2010.
many woodlands on private lands are fragmented by agriculture
81. Keane and Schoettle 2011.
and exurban development, and this landscape pattern facilitates
82. Logan and Powell 2001.
nest predation by brown-headed cowbirds. As a result, yellow
83. http://www.greateryellowstonescience.org/download_product/
warblers generally fail to produce sufficient surviving offspring
538/0.
to sustain the population in these habitats as well. Apparently
84. Gresswell and Liss 1995.
the yellow warbler populations throughout the areas studied by
85. In 2012 more than 300,000 lake trout were removed from
Hansen and Rotella (2002) are sustained by birds coming into
Yellowstone Lake, and the number of juvenile cutthroat trout
this area from more suitable habitats elsewhere rather than by
appeared to be higher than in recent years (Yellowstone Associa-
local reproduction. See also Hansen and DeFries 2007.
tion E-Newsletter, December 2012).
51. National Academy of Sciences 2002.
86. Middleton et al. 2013d; see also Wilmers and Post 2006.
87. Schwartz et al. 2006.
53. Frank and McNaughton 1992, 1993. Deposition of urine and
88. Fortin et al. 2013; Middleton et al. 2013d.
feces, as well as soil scarification by the action of hooves, also
89. Mattson et al. 2002.
may contribute to high plant growth in grazed grasslands.
90. National Academy of Sciences 2002; Schullery 2004.
54. Coughenour 1991a,b; Coughenour and Singer 1991.
91. Pritchard 1999.
55. Houston 1982 (p. 198).
56. Chadde and Kay 1991.
93. Vucetich et al. 2005; seventeen additional Canadian wolves
57. Romme et al. 1995; Ripple and Larsen 2000.
were released in Yellowstone in April 1996.
58. Patten 1968; Singer et al. 1994; Yellowstone National Park 1997.
95. Middleton et al. 2013b,c,d; some evidence indicates that the
60. Research suggests that aspen persistence is not threatened in

the GYE as a whole. Brown et al. (2006) determined aspen cover
wolf population on the northern range of YNP has declined as

the number of elk has dropped.
across the GYE in 1956 and 2001; they found an average loss of
96. Vucetich et al. 2005.
10 percent of aspen cover overallfar less than was suggested
97. Beschta 2003, 2005; Ripple and Beschta 2004; Beschta and
by fine-scale studies of aspen that commonly have been conducted where ungulate browsing is conspicuous. Brown et al.
(2006) also determined that aspen is rare in the GYE, occupying only 1.4 percent of the region, and is more common in the
southern than in the northern part of the GYE.
Ripple 2010.
98. Beyer et al. 2007; Creel and Christianson 2009; Kauffman et al.
2010; Middleton et al. 2013a.
99. Kauffman et al. 2010; Middleton et al. 2013a. Kimble et al.
(2011) did find places where the density and size of aspen

sprouts apparently had increased between 1991 and 2006, but
part of the Limestone Plateau. Tertiary igneous intrusions in
cautioned that these patterns cannot be extrapolated to the
the Hills were formed 55 to 33 million years ago and include
landscape or attributed solely to the effects of wolves. Eisenberg
Bear Butte, Black Butte, Crow Peak, Custer Peak, Devils Tower,
et al. (2013) review the literature on wolf-elk-aspen research
Inyan Kara Mountain, the Little Missouri Buttes, Sundance
and emphasize that general patterns and effects are not clear.
See also http://nyti.ms/1cHqikq.
Mountain, Terry Peak, and Warren Peaks (Lisenbee et al. 1981).

14. Shepperd and Battaglia (2002) provide a review of all forest
100.Marshall et al. 2013; where willows have recovered, Baril et
types; see also Larson and Johnson 1999; Marriott et al. 1999;
al. (2011) found an increase in some species of songbirds; Baker
Marriott and Faber-Langendoen 2000; Wienk et al. 2004;
et al. (2005) describe the effect of elk herbivory on willows and

beaver; Hebblewhite et al. (2005) describe how human activity
has mediated the trophic cascade caused by wolves.
101. Berger et al. 2008.
102. Hansen and Rotella 2002; Parmenter et al. 2003; Gude et al.
2007; Hansen and DeFries 2007; Bilyeu et al. 2008.
103.Olliff et al. 2010.
104. Schullery 2010 (p. 12).
Battaglia et al. 2008.

15. Progulske 1974; Bock and Bock 1984; see chapter 10 and the
condition reports produced by the various units of the National
Park Service (available online).
16. Boldt and Van Deusen 1974.
17. Marriott and Faber-Langendoen (2000) provide a more detailed
classification.
18. Marriott and Faber-Langendoen (2000) describe the riparian
and wetland plant communities of the Black Hills.
Chapter 16: The Black Hills, Bear Lodge Mountains, and

Devils Tower
1. The Lakota name for the Black Hills is Paha Sapa, which trans-
19. McIntosh 1931.

20. Severson and Thilenius 1976.
21. MacCracken et al. 1983; Marriott and Faber-Langendoen 2000;
see chapter 5.
lates to hills that are black, an impression created by the
22. The three characteristic grasses are Sporobolus heterolepis, Achna
appearance of ponderosa pine forests from a distance. Lageson
therum richardsonii, and Danthonia intermedia, respectively. Mar-
and Spearing (1988) and Gries (1996) summarize the geologic
riott (2012) describes the Black Hills montane grassland type,
history of the area and include road logs that identify geo-
which is considered unique to the Black Hills; http://www
logic features; Froiland (1990) and Raventon (1994) review the
.natureserve.org/explorer/servlet/NatureServe; the large grass-
natural history of the Black Hills. The geologic origin of Devils
lands in the Central Area near Deerfield Reservoir, known as
Tower is unclear, but it is one of several igneous intrusions in
Reynolds, Slate, and Gillette prairies, have been largely con-
the area (see endnote 13).
verted to hay meadows.
2. Froiland 1990. The Vrendrye brothers encountered several
23.
MacCracken et al. 1983; Hoffman and Alexander 1987;
tribes along the way, traveling most likely to the Bighorn Moun-
mountain-mahogany is found only in the Black Hills east of
tains on a route north of the Black Hills (http://en.wikipedia
Newcastle and is rare in the Bear Lodge Mountains.
.org /w i k i/ Ve re nd r ye _ Brot he r s _ jou r ney_ to _ t he _ Ro c k y_
24. Boldt and Van Deusen 1974; Shepperd and Battaglia 2002.
Mountains). The Sioux arrived later in the 1700s. A treaty was
25. Shepperd and Battaglia 2002; Spiering and Knight 2005; see
signed in 1868 that gave the Sioux rights to the Black Hills. However, ownership is still the subject of a legal dispute between the
Sioux and federal government (http://www.defendblackhills
.org).
3. Custer 1875 (p. 506); Donaldson 1875 (p. 564); Ludlow 1875;
Raventon 1994; Grafe and Horsted 2002.
also chapter 12.

26. Wienk et al. (2004) found that a combination of thinning and
prescribed burning stimulates understory plant growth.
27. Uresk and Paintner (1985) studied cattle and found that shrubs
and trees in the northern Black Hills provide 37 percent of
their diet during September; in summer the diet was 54 per-
4. H. Marriott, pers. comm.
cent grasses, 17 percent forbs, and 28 percent shrubs and trees.
5. Progulske 1974; Grafe and Horsted 2002.
Elsewhere, cattle consume recently fallen cottonwood leaves in
6. McIntosh 1931; Raventon 1994; Larson and Johnson 1999.
September and October, such as along the South Platte River in
7. Buttrick 1914; Wright 1970.
Colorado (Sedgwick and Knopf 1991).
8. King et al. 2013.

9. Larson and Johnson (1999) describe six geomorphic regions
28. Fisher et al. 1987; Wienk et al. 2004; Brown 2006; Brown et al.
2008.
rather than five, adding the Gray Shale Foothills common
around the Bear Lodge Mountains.
30. Newton and Jenney 1880 (p. 322).
10. The Spearfish Formation is known as the Chugwater Formation

in Wyoming; gypsum also is found in the formation.
11. Marriott et al. 1999.
12. Marriott 2012.
13. Harney Peak 7,244 feet, Bear Mountain 7,164 feet, Crooks
Tower 7,137 feet, Terry Peak 7,082 feet, and Crows Nest Peak
7,045 feet. Harney Peak is in the Central Area; the others are
31. Quotation is from Dodge (1876, p. 62); photos in Progulske 1974.

32. Lovaas 1976.
33. Baker et al. (2007) concluded that a variable-severity fire model is
often more realistic and is pertinent to the restoration of forest
structure; see also Shinneman and Baker 1997.
34. Fisher et al. (1987) wrote about the effects of Native Americans
near Devils Tower.
345
346

35. Brown and Cook 2006; Brown and Schoettle 2008; Brown et al.
2008.
36. Shinneman and Baker 1997; Baker et al. 2007.
9. Mason 1987 (p. 41).

10. Laramie Chamber of Commerce 1913.
11. In 2010 hay was grown on 59,000 acres, about 3 percent of the
37. Shepperd and Battaglia 2002.
Laramie Basin, and almost entirely consisted of native grasses
38. Brown 2006; Brown et al. 2008.
(no or little alfalfa). Most alfalfa in Wyoming is grown in
39. Orr 1975.
Goshen and Fremont counties. Data from U.S. Department of
40. Early journals reviewed by Shepperd and Battaglia (2002).

41. Lovaas 1976; Wind Cave National Park has used prescribed fires
since the early 1970s.
Agriculture National Agricultural Statistics Service.

12. Mears 1991, 2001.
13. The annual water supply for the city of Laramie is approxi-
42. Battaglia et al. 2008.
mately 60 percent from surface water draining from the Medi-
43. Brown (2006) found that fires were more frequent during La
cine Bow Mountains of Wyoming and Colorado, and 40 percent
Nia years, cool phases of the Pacific Decadal Oscillation, and

warm phases of the Atlantic Multidecadal Oscillation.
44. Lentile et al. 2006.
from wells into the Casper aquifer.

14. The Casper Aquifer Protection Plan is at http://www.ci.laramie
.wy.us/index.aspx?NID=226.
45. Shepperd and Battaglia (2002) review the ecology of the vari-
15. Mears 1991; katabatic winds are generated by air becoming
ous insects and fungal diseases common in the area, namely,
cooler and denser on glacier surfaces, then flowing down
mountain pine beetle, pine engraver, red turpentine beetle,
glacial and mountain slopes; they are well known in Ant-
Armillaria root rot, red rot, western gall rust, needle cast, and
arctica and New Zealand. See http://en.wikipedia.org/wiki/
diplodia tip blight; see also Negron et al. 2008; chapters 11

and 12.
Katabatic_wind.
16. The Wyoming Water Resources Data System (http://www
46. The most recent condition assessments are available online or
.wrds.uwyo.edu/) provided temperature and precipitation data
from the headquarters of the parks or Devils Tower National
for a 2.5 mile 2.5 mile area in the Laramie Basin (41.25N,
Monument.
47. Battaglia et al. (2008) report that fires have helped reduce some
exotic plants, and that 39 percent of the grasslands and 19 percent of the forests in the Black Hills had prescribed burns in the
previous 10 years.
48. Wind Cave National Parks ferret population was estimated to
be 4664 in 2013; see the parks website for current estimates.
49. Pauli et al. 2006.
50. B. Burkhart, pers. comm.
51. Pronghorn prefer forbs, which are more common in prairie dog
105.75W), using PRISM Climate Group data, Oregon State University (http://www.prism.oregonstate.edu/terms.phtml).
17. Irrigation developments have increased the size of wetlands in
the basins.
18. Black sagebrush is found just east of Laramie, where Grand
Avenue joins Interstate 80; exposed bedrock in the same area is
dominated by true mountain-mahogany; see Thatcher 1959.
19. Elliott-Fisk et al. 1983; Norris et al. 2006; S. T. Jackson, pers.
comm.
20. Peck and Lovvorn 2001; Sueltenfuss et al. 2013.
colonies; bison graze on the younger, less fibrous grass sprouts
21. The Wyoming Game and Fish Department has a report titled
that are easily accessible in the colonies because of the clipping
Laramie Plains wetland complex: Regional wetland conserva-
done by the rodents.

52. Rehfeldt et al. 2006; see also Bartlein et al. 1997.
53. Shepperd and Battaglia 2002; King et al. 2013.
tion plan (March 19, 2012).

22. Beath et al. 1941; Kolm 1975; there are two groups of plants
that accumulate selenium. One, the selenium indicators,
require selenium for their growth (two-grooved milkvetch,
Chapter 17: The Laramie Basin
tineleaved milkvetch, woody aster, and princesplume). In

contrast, the other comprises facultative accumulators, such
1. The one basin with no outlet is the Great Divide Basin, where
as Gardners saltbush, gumweed, locoweed, mentzelia, snake-
the Continental Divide splits for a distance of about 100 miles
weed, spiny hopsage, winterfat, and certain grasses. They do
(see fig. 1.2).
not require selenium but often develop high concentrations of
2. Prasciunas et al. 2008.
the element. Some plants do not accumulate selenium, even
3. During the Pleistocene, much of the Laramie Basin was a cold,
though they are frequently found on seleniferous soils. Both
semi-arid, tundra-like steppe, as indicated by the presence of
selenium indicators and facultative accumulators can concen-
fossil ice wedges; see chapter 2.
trate large amounts of the element in leaves and stems without
4. Shuman 2012.
adverse effects on the plant itself. For example, two-grooved
5. Kornfeld et al. 2010.
milkvetch is known to accumulate 4,000 ppm or more from
6. Homsher 1965.
soils containing only 1.1 ppm in the soil. Grasses and cereal
7. Burdick 1987 (pp. 23); also, Homsher (1965) and Mason (1987)
grains with only 1030 ppm can be lethal or crippling to cattle
provide overviews of human history in the Laramie Basin.
and horses. Both the indicators and the accumulators increase
8. Fremont 1845; Stansbury 1852; Smith 1913 (p. 272); Dale 1918
surface-soil selenium concentrations as their shoots decom-
(letter from Ashley to Atkinson Dec. 1, 1825); Parkman 1950;
pose in the fall, thereby possibly improving conditions for
see also Dorn 1986.
their own growth.

23. Myers 1969.
24. M. T. Richardson (unpublished) reports that construction of the
disruption of disturbance regimes; it can be viewed at http://

wgfd.wyo.gov/web2011/wildlife-1000407.aspx.
Pioneer Canal from the Big Laramie River to Sodergren Lake
5. http://www.rivers.gov/rivers/wyoming.php.
was begun in 1879. This canal was extended to Lake Hattie in
6. Ashworth 2006.
1903. Units 2 and 3 of the Wheatland Reservoirs were created
7. Curtis and Grimes 2004 (p. 106); www.wrds.uwyo.edu.
by damming the Laramie River in the north end of the Laramie
8. Kornfeld and Osborn 2003.
Basin (see fig. 17.1). Richardson presents anecdotal evidence
9. Baker and Knight 2000; Ingelfinger and Anderson (2004) found
suggesting that irrigation caused an increase in the abundance
that the densities of Brewers sparrow and sage sparrow were
of mosquitos. For a map of the Pioneer CanalLake Hattie Irri-
reduced by 3960 percent within 300 feet of roads having low
gation District, see http://waterplan.state.wy.us/plan/platte/
traffic. See also Sawyer et al. (2007, 2009) for studies on the
atlas/upper/upper_agricultural_irrigation_districts.htm.
25. Lovvorn et al. 1999; Peck and Lovvorn 2001.
26. Laramie Sentinel, June 7, 1879; Laramie Boomerang, 1892 (interview with W. O. Owen).
effects of roads and traffic on elk and mule deer.

10. Ostlund 2012; prior to the completion of the Trappers Point
project, about 700 pronghorn and mule deer were killed along
a 27-mile stretch of U.S. Highway 191 during a 5-year period.
27. Peck et al. 2004; Shinker et al. 2010.
11. Pocewicz et al. 2011.
28. For a summary of Wyoming water law, see http://library.wrds
12. Knight and Reiners 2000.
.uwyo.edu/wrp/90-17/90-17.html. Water rights are granted

according to the first in time, first in right doctrine.
29. For weeds and pests, see http://www.wyoweed.org/statelist.html.
13. Kuvlesky et al. 2007; Pruett et al. 2009; Knick and Connelly
2011; Dinkens et al. 2012; LeBeau et al. 2014; Hamerlinck et al
(2013) describe recent trends in land use.
For fish- and game-related nuisance species, see http://wgfd
14. Eminent ecologist Eugene Odum suggested that a region could
.wyo.gov/web2011/Departments/Hunting/pdfs/Regulations_
be divided into four fundamental kinds of landscapes: urban,
Ch62.pdf.
industrial, agricultural, and protective. Properly managed
30. Ranchers prefer more grasses than sedges and rushes, which
rangelands can be considered as protective (see chapter 6).
tend to increase with excess irrigation; slender wheatgrass,
15. Hansen et al. 2002 (p. 160); see also Power 2005.
thickspike wheatgrass, and western wheatgrass are sometimes
16. Bormann 1976 (p. 759).
common along the edges of irrigated meadows.
17. Daily and Ellison 2002; Clark 2007; Esty 2007; Banerjee et al.
31. M. Curry, pers. comm.

32. Data from the Wyoming State Climate Office.
33. See the following website for other comparisons: http://www
.wrds.uw yo.edu/sco/data/normals/1971-2000/delta-Precip
-Annual.html.
34. Reiners 2003.
35. Norris et al. 2006.
2013; Hansen et al. 2013; the 2008 Farm Bill called for measuring the environmental benefits of conservation.
18. Hansen et al. 2002; Copeland et al. 2007, 2009; Kiesecker et al.
2007, 2009a,b; Rissman et al. 2007.
19. Pocewicz et al. (2013) identified specific areas important for
migratory birds.
20. Naugle 2011; Jakle 2012a,b.
21. A safe harbor agreement assures landowners that the U.S.
Chapter 18: Using Western Landscapes

1. Larson 1977.
2. Unruh 1993.
3. Madson 2007 (p. 4); see also Madson 2006b. Worster (1994)
wrote about the New West as an unsettled country. Baron
et al. (2013) summarize the rapid, unexpected changes that
occurred from 2002 to 2012 in the Rocky Mountain region;
Hamerlinck et al. (2013) describe land use trends in Wyoming.
4. The Wyoming Game and Fish Department developed a 900page plan for the conservation of wildlife and the habitat
they depend on. The plan describes the status of individual
species and identifies five primary threats: rural subdivision,
energy development, invasive species, climate change, and the
Fish and Wildlife Service will not require different management activities without the landowners consent, and that, at
the end of the agreement period, participants may return the
property to the conditions that existed at the beginning of the
agreement.
22. Middleton et al. 2013b,c,d.
23. Williams and Jackson 2007; Williams et al. 2007; Breshears et
al. (2005) describe global-change-type drought that has greatly
reduced the abundance of pinyon pine in the southern Rocky
Mountains.
Epilogue
1. Leopold 1938 (p. 254).
347
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Index
Page numbers for entries occurring in figures are followed by an f; those for entries in notes, by an n; and those for entries in tables, by a t.
Bold page numbers indicate pages where a definition can be found.
aboveground net primary productivity (ANPP): in grasslands, 90f; in
lodgepole pine forest, 210f, 340n29
Absaroka Range, 15, 245, 249
Actinobacteria, 64
23739, 237f, 239f; in Greater Yellowstone Ecosystem, 248, 249;

krummholz in, 23133, 232f; major plant communities of, 236;
nitrogen deposition in, 23941, 342nn3637, 343n40; range of
temperatures in, 33; soils of, 232
adaptive radiation, 128
alpine turf, 236, 23839t, 239
adiabatic lapse rate, 33, 173, 327n10, 337n2(ch11)
American beachgrass, 143
Adobe Town, 148f
American elm, 53, 328n8
aeration of soils, 3536
American Indians. See Native Americans
aerenchyma, 6870
American pika, 236, 236f, 242
African grasslands, 89, 94, 97
amphibians: in forests, 179t; in wetlands, 68, 70. See also specific types
Age of Mammals. See Cenozoic Era
anaerobic soils, of wetlands, 6870
aggregating pheromones, 184
Andersen, Chamois, 41
agriculture: area covered by and location of cultivated land, 3, 8f;
Anderson, D. C., 337n2(ch11)
carbon sequestration in, 11; climate change and, 4041; crops
Anderson, E. M., 329n31
of, vs. invasive plants, 329n42; dominant crops in, 325n1(ch1);
Anderson, M. D., 342n20(ch13)
frost-free period and, 33; in grasslands, 83, 92, 93, 100, 1078;
Anderson, S., 347n9
in Laramie Basin, 285, 288, 296, 346n11; limiting factors on, 3;
ANPP. See aboveground net primary productivity
as percentage of state economy, 325n1(ch1); in sagebrush steppe,
antelope. See pronghorn
109; soil characteristics in, 34. See also livestock
antelope bitterbrush, 12021, 143, 156, 158t, 160, 16466, 165f, 176t,
Airport Bench, 287, 288f, 296f
254
Akashi, Yoshiko, 55
ants, harvester, 103, 104f, 139
alder-conifer woodlands, 4749t
apical dominance, 95, 198, 331n7
alfalfa, 325n1(ch1), 346n11
apomixis, 236
alfisols, 174, 322t
aquifers: in Laramie Basin, 286, 298, 346n13; Ogallala, 307, 336n10
alkali flats, 330n20
arctic tundra, 235
alkalinity of soils, 36, 327n17, 335n4
aridisols, 83, 322t
allelopathy, 79
Arizona Monsoon, 31, 196
Allred, B. W., 101, 120
Arkley, R. J., 151
alluvium, in Laramie Basin, 285, 286, 286f, 287
arrowgrass, 72t
alpine pennycress, 235
Ashley, William, 284
alpine sunflower, 236f
aspen forests, woodlands: 8f, 16770, 19698, 196f, 197f; in Black Hills,
alpine tundra, 8f, 23042; adaptations to, 23536, 239; animal
272, 273, 273f; climate change in, 219220; cloning by, 168, 196
species in, 236; avalanches in, 23334, 233f; burrowing animals
97; dominant plant species in, 15859t; 17677t; elk browsing on,
in, 236, 239; climate change in, 24142; dominant plant species
16870, 169f, 256, 257f, 258, 264, 344n60, 344n99; as evergreen
in, 235, 23637, 237f, 23839t; elevation of, 23031; frost in,
tree, 174, 196; fires in, 182, 198, 20910, 261, 262f, 339n99;
387
388 Index
aspen forests (continued)
Berger, J., 343n45
in foothills, 156, 157f, 16770, 167f; genetic diversity in, 197,
Beschta, Robert, 264
339n97; in Greater Yellowstone Ecosystem, 169f, 253, 256, 257f,
Biederman, Joel, 211
258, 26465, 344n60; hydrologic budget of, 205; in Laramie Basin,
big game: current populations of, 24; in foothills and escarpments,
290f, 295; in mountain meadows, 226; sudden aspen decline in,

198, 220, 220f; understory density in, 338n16
aspen atolls, 167, 167f, 226
aspen tent caterpillar, 182
asteroids, 1314
15556, 16871. See also specific types

Big Hollow, 287, 288f, 296f
Bighorn Basin: droughts in, 32; foothills of, 16061, 160f, 162;
frost-free period in, 33; plants in geologic history of, 14, 22, 23f;
sagebrush in, 111; soil crusts in, 126, 126f, 140
atmospheric circulation, 31, 327n5
Bighorn Canyon National Recreation Area, 54
avalanches, 23334, 233f, 253
Bighorn Mountains: alpine tundra in, 230, 231f; average tempera-
badlands, 14748; animal species in, 147; dominant plant species in,
Bighorn River, 52f, 5356, 53f, 56f, 63f
ture changes in, 40; formation of, 15; meadows in, 224f
14748; geologic history of, 17; topography of, 2930, 147
Big Laramie River, 283f, 285, 294f, 296, 347n24
Badlands National Park, 107, 147, 279
big sagebrush. See sagebrush, big
Baker, W. L., 121, 140, 275, 336n39, 342n20(ch13)
Billings, W. D., 225, 342n32, 342n34, 343n45
Bamforth Basin, 287, 288f
bioclimatic envelope, 219
bank storage, 57, 59, 60
bioengineers: beaver as, 57; prairie dogs as, 102
bar(s), in riparian landscapes, 46, 51, 52, 55, 56, 56f
biological control of invasive plants, 65, 79, 329n50

biomass: in alpine tundra, 235, 240; in grasslands, 8485, 86f,
87f, 89, 331n16; in mountain forests, 2012, 202f, 210; in sagebrush steppe, 119. See also net primary productivity
birdfoot sagewort, 135
birds: in alpine tundra, 236; in foothills, 17172; in grasslands, 88t,
333n93; in Greater Yellowstone Ecosystem, 257, 265, 343n45,
344n50; migratory, 68, 330n9; in mountain forests, 178, 179t,
182, 198; roads avoided by, 310, 347n9; seed dispersal by, 17172;
in wetlands, 68, 70. See also specific types
biscuit land. See mima mounds
bison: abundance in grasslands, 334n31; in Black Hills, 27980;
decline in population of, 24, 327n56; effects on grasslands,
9598, 97f, 332n41, 332n50; food preference of, 89; in geologic
history, 13, 325n6(ch2); in Great Plains, 28; hunting of, 24,
332n41; in Laramie Basin, 28485; peak population of, 327n57;
in prairie dog colonies, 97, 97f, 27980, 346n51; in riparian landscapes, 60; sagebrush steppe and, 118, 12627
bitterbrush, antelope, 164, 254
black-backed woodpecker, 182
blackbird, yellow-headed, 67f
blackbody effect of shrubs, 11718
blackfelt snowmold, 225, 226f
black-footed ferret, 10f, 103, 106, 107, 279, 332n76, 346n48
Black Hills, 26681; bark beetle outbreaks in, 271f, 27778; fires in,
187, 27478; foothills of, 166f; formation of, 15, 266; geomorphic
regions of, 267f, 26870, 345n9; grasslands in, 84f, 270, 27374;
invasive plants in, 27879, 279f; management issues in, 27481;
meadows in, 273, 273f; Native Americans in, 266, 345nn12;
plants characteristic of, 26668, 269t, 27074; precipitation in,
267, 270f; topography of, 266, 267f
Black Hills National Forest, 266
black sagebrush. See sagebrush, black
black-tailed prairie dog, 102, 102f, 106, 27879, 280f
BLM. See Bureau of Land Management
Blossey, B., 333n83
bluebunch wheatgrass. See wheatgrass, bluebunch
bluegrass, Kentucky, 79
bluegrass, Sandberg, 29f, 83, 84f, 100, 109f, 110, 12021, 136t, 150f,
164f, 222t, 250, 251f, 289, 291f
bluestem, big, 331n2
bark beetles, 180, 18285, 183; in Black Hills, 27778; climate

change and, 41, 216, 341n63; in Douglas-fir forests, 189; effects
of, 18285, 21011; fires and, 21617, 341n64; fungi in, 182f,
184, 338n38; in Greater Yellowstone Ecosystem, 185, 216,
26263, 340n39; hydrology and nutrient cycling affected by,
211, 340n41; life history of, 18385; in lodgepole pine forests,
185, 216, 338n19; management options for, 21819; as periodic
disturbance, 37; predictors of, 18485, 338n42; species of, 182; in
spruce-fir forests, 195, 211
basin big sagebrush, 111; distribution of, 111, 333n5; greasewood
and, 134; in sand dunes, 143
basins, 5f. See also intermountain basins; specific basins
bastard toadflax, 179
Battaglia, M. A., 346n47
beachgrass, American, 143
bear, grizzly, 26364, 263f; distribution of, 3, 263, 325n2(ch1); pine
seed consumption by, 172, 234, 263; threats to, 26364
Bear Lodge Mountains, 26681; dominant plant species in, 27074;
fires in, 275; foothills of, 166f; formation of, 266; geomorphic
regions of, 270; management issues in, 27481; precipitation in,
267, 270f
Beartooth Mountains: alpine tundra in, 230, 237f, 240f, 342n21;
bryophytes in, 330n29
beaver: impact on riparian landscapes, 5760, 58f, 59f, 329nn2324;
in Laramie Basin, 295; trapping of, 24, 63, 327n54; and willows,
265
Bebb willow, 50f
beetle, Douglas-fir, 182, 189, 189f
beetle, Mediterranean tamarisk, 329n50
beetle, mountain pine, 181f, 18285; in Black Hills, 27778; climate
change and, 216, 341n63; effects of, 18285, 182f, 183f, 219,
340n45; in Greater Yellowstone Ecosystem, 26263
beetle, spruce, 182, 195, 216, 219
beetle, western balsam bark, 182, 195
Belle Fourche River, 266, 280
Belsky, Joy, 95
benches, in Laramie Basin, 286f, 287, 288f, 296f
Benkman, C., 17980
bentonite, 13, 35, 36, 337n28
Berg, A., 151
Index
bluestem, little, 83, 84f, 88, 100, 155, 159t, 163, 166, 268, 27374,
277
caldera, 245
Caldwell, M. M., 115, 331n8
blue spruce, 46, 328n3
cambium, 187f
blue-stain fungus, 182f, 184, 338n38
Camel Rock (in Laramie Basin), 287, 292f
Blumenthal, Dana, 1056, 333n87
Canada thistle, 104
bogs: vs. fens, 77; for geologic history, 14; lack of, in Wyoming,
canarygrass, reed, 79
329n1
canine distemper, 332n76
Bonneville, Lake, 19
canyons, geologic history of, 17. See also specific canyons
Boon, S., 340n41
carbon credits, 11
Bormann, F. Herbert, 200, 31415
carbon dioxide: in Cenozoic Era, 15; current concentrations of,
Boulder Creek, 270

boulder fields, 236, 239
boulders, in alpine tundra, 23637, 239
Bowman, W. D., 342nn3637
328n31; and grasslands, 108; in sagebrush steppe, 119; in soils of

wetlands, 68. See also climate change
carbon flow: in mountain forests, 201, 201f, 210, 211, 340n39,
340n46; in sagebrush steppe, 11819
Boysen Reservoir, 55
carbon refixation, 196
Bradley, B. A., 335n81
carbon sinks, 210, 211, 340n39
braided channels, 46, 51, 51f, 56
Carrera, P. E., 343n44
Breshears, D. D., 337n19
carrying capacity: in mountain meadows, 227; soil characteristics
Brewers sparrow, 310, 347n9
and, 34
Bridger, Jim, 24, 285
Carson, Kit, 28485
Bridger Formation, 326n19
Carter, William A., 106
Bridger Range, 338n4
Cascade Canyon, 19f, 233f
Briske, David, 97
Casper aquifer. See Casper Formation
bristlecone pine, 234, 342n12, 342n15
Casper Formation, 284f, 286, 287, 287f, 293, 293f, 295, 297f
broadleaf cattail, 79
Castle Creek, 273f, 275f
broadleaf woodlands, 4749t
caterpillar, aspen tent, 182
Brown, H. C., 151
cattail, broadleaf, 69f, 70, 74, 79
Brown, J., 260
cattail, hybrid, as invasive species, 79, 331n47
Brown, K., 344n60
cattail, narrowleaf, 69f, 70, 74, 79, 330n47
Brown, M., 340n39
cattle: effects on grasslands, 9596; food preference of, 89; popula-
Brown, P., 275, 276, 346n43
tion of, 25, 327n62. See also livestock
browsing. See grazing and browsing
Cavalry, U.S., 343n41
brucellosis, 344n48
Cenozoic Era, 12f, 1423
bryophytes, 330n29
Centennial Ridge, aspen and Douglas-fir on, 157f
bud sagewort, 135
Centennial Valley, 156, 157f
budworm, western spruce, 182, 189, 189f, 339n65
Central Area of Black Hills, 270, 271f, 345n13
buffalo. See bison
Chain Lakes wetland, 17, 149f
Buffalo Gap National Grassland, 279
Chapman, T., 184
buffaloberry, russet, 175, 274
cheatgrass: in desert shrublands, 138, 140, 141, 336n39; in foothills,
buffaloberry, silver, 46, 48t, 54, 56, 247f
166; in grasslands, 104; introduction of, 123; in Laramie Basin,
buffalo grass, 83, 331nn12
298; in mima mounds, 149; in sagebrush steppe, 12224, 124f,
Bull Lake glacial period, 326n23
334n73
bulrushes, 67f, 70, 71, 72t, 7476, 250, 292
Chimney Rock (in Laramie Basin), 287, 292f
Bultsma, P. M., 332n59
Chisholm, James, 121
bunch grasses, 83
Chokecherry and Sierra Madre Wind Energy Project, 312
Burdick, Laura, 282
chokecherry woodlands, 167. See also woody draws
Bureau of Land Management (BLM): on feral horses, 128; map of
Chugwater Formation, 13, 295, 297f, 345n10
land jurisdiction of, 6f; multiple use on lands of, 3089
Cinnabar Park, 223, 224f, 225, 225f
Burke, I. C., 331n1, 333n94
cinquefoil, shrubby. See shrubby cinquefoil
Burnham, Jennifer, 151
Circle Bar Lake, 148
bur oak, 168, 27273, 272f, 273f
cities, map of, 7f
burrowing animals: in alpine tundra, 236, 239; disturbances caused
Clark, F., 9394
by, 37; in grasslands, 84, 1013; in mima mound formation, 149,
Clark, William, 96, 249
151. See also specific types
Clarks Fork, 307
bushy-tailed woodrat, 22, 22f, 23f
Clarks nutcracker, 17172, 171f, 179, 202

Clean Water Act, 328n2, 329n1
C3 plants: in desert shrublands, 133; in grasslands, 8889
clearcutting: compared to fire, 21115, 340n27; habitat fragmen-
C4 plants: in desert shrublands, 133; in grasslands, 8889
tation by, 311, 311f; of mountain forests, 178. See also timber
cactus, pricklypear, 96, 100, 102
harvesting
389
390 Index
Cleary, M., 119
coyote, 265
Clements, Frederick, 96
crane, sandhill, 328n13
climate, 38; history of, 1223, 32f, 39f, 326n36; vs. weather, 38. See
creeks, map of, 46f. See also specific creeks
also climate change

climate change, 3841; in alpine tundra, 24142; approaches to
Cretaceous-Paleocene Extinction Event, 1314, 325nn89

Cretaceous Period, 12f, 13, 325n3(ch2)
adapting to, 21920, 341n88; approaches to slowing, 40; average
cricket, Mormon, 37, 101, 332n73
temperatures in, 3840, 39f; in Black Hills, 281; computer models
Crist, T. O., 332n71
of, 3840, 39f; current rate of, 15, 3841; in desert shrublands,
cropland. See agriculture
14041; fires and, 41, 26162; in grasslands, 1078; and land
cross-bedding, 287, 287f
management, 31617; in Laramie Basin, 299301, 299f, 300f; in
Crow Creek, 54
mountain forests, 21516, 21920; in mountain meadows, 22829;
crown fires, 37; and bark beetle outbreaks, 217; in Black Hills,
Paleocene-Eocene Thermal Maximum and, 15; and precipitation,

40, 41, 215, 328nn2930; and precipitation-evaporation ratio,
27475, 276, 277f; vs. clearcutting, effects of, 21112; intensity

and severity of, 182, 208, 209f; types of, 217
34; in riparian landscapes, 65; in sagebrush steppe, 120, 121, 124;
cryoturbation, 151; 237, 239, 342n34. See also alpine tundra
tree-ring studies documenting, 23; use of term, 38; in wetlands,
cultivated land. See agriculture
7980
Cunningham, P., 252
climate diagrams, 32f, 289f
curlleaf mountain-mahogany. See mountain-mahogany
cloning: in aspen, 168, 19697; in krummholz, 232; in spruce and
Curtis, J., 327n10
fir, 194, 194f

cloud seeding, 308
cushion plants: in alpine tundra, 235; in desert shrublands, 135,

138f; in foothills, 166
Clovis people, 20, 282
Custer, George A., 266, 273f, 274
Clow, D. W., 340n41
cut banks, 46, 51, 52f
Cluff, G. J., 335n14
cutthroat trout, 26364, 344n85
Clyman, James, 126

coal, formation of, 13, 14, 15, 28788, 326n13. See also energy
extraction
coarse woody debris, 209f, 21115, 213f, 262f, 271f
Dahl, T. E., 330n40

dams: beaver, 5760, 58f, 265; debris, 57, 59f; human-made, 252,
280, 296, 328n16, 329n34
Colby Mammoth Site, 326n30
Daubenmire, R., 185, 338n52, 342n1
cold hardening, 184
Debinski, D., 228
Colorado River: streamflow from Wyoming in, 329n35; water alloca-
debris. See O horizon
tion compacts for, 327n9
debris jams, 57, 59f
Colter, John, 249
DeByle, N., 16970
comandra blister rust, 17879
deciduous woodlands, 16668; in Black Hills, 27273
common reed, 79
decomposition: in grasslands, 92; in mountain forests, 2012, 2078,
Como Bluff, 13
compensatory growth hypothesis, 95
212f; in sagebrush steppe, 119; under snow, 202, 339n3; in

wetlands, 75
Comstock, T., 256, 343n38
decreaser species, 95
cones, serotinous, 17980, 19192, 192f, 339n71
deer, mule: decline in population of, 17071; food preferences of,
conifer woodlands, 4749t, 15859t. See also forests
89; in foothills, 17071; in grasslands, 89; in Greater Yellowstone
conservation banks, 315, 316
Ecosystem, 257; mortality on roads, 310, 347n10; roads avoided
conservation biology, 910, 11
by, 310
conservation finance, 315
Defenders of Wildlife, 264
Conservation Reserve Program, 11, 107, 315
defoliating insects, 182
Continental Divide, 28, 34
denitrification, 207
conveyance losses, 57
desert grassland, 133, 13637t
Cooke, P. G., 54
desert pavement: formation of, 333n10; in mixed desert shrublands,
cool-season plants, 8889
13435; in sagebrush steppe, 112
Cooper, D. J., 265, 330n40
desert playas, 132
Copeland, H., 80
desert shrublands, 8f, 13141, 131f; 134f; adaptations to, 13233;
coppice dunes: in desert shrublands, 13435; in sagebrush steppe,

112, 115f
cottonwood: drought tolerance of, 329n56; elk browsing on, 6061,
animals species in, 139; area covered by, 131; disturbances in,
13940; dominant plant species in, 131, 13637t; ecosystem of,
13739; future challenges in, 14041; major plant communi-
329n33; regeneration after floods, 5253, 5556, 56f, 328n11,
ties of, 13337; precipitation in, 131; resource partitioning in,
328n13
13233; sagebrush steppe compared to, 131, 132, 133, 335n3;
cottonwood, lanceleaf, 53, 328n8

cottonwood, narrowleaf, 46, 51f, 56, 328n10, 328n16, 329n56
soils of, 13134, 139, 335n4; water availability in, 131, 132, 132f,
13739; in Yellowstone National Park, 250
cottonwood, plains, 46, 51f, 52f, 53, 54, 329n56
desert yellowhead, 10f
Coughenour, M., 97
Detling, J., 97
Cox, G., 151
detritus: in grasslands, 90, 92; in mountain forests, 2012
Index
Development by Design, 31516
Yellowstone Ecosystem, 25658, 257f, 259f, 26465; in riparian
Devils Tower National Monument, 268f; establishment of, 3; fires in,
landscapes, 6061; roads avoided by, 310; tree damage by, 168
187f; geologic history of, 266, 345n1; management issues in, 280
70, 169f, 256, 257f, 264, 337n45; winter rangelands and feeding
dikes, 62
of, 17071, 25758; wolves and, 170, 264, 344n99
dinosaurs, 13, 325n6(ch2)
Ellison, L., 96
Dinwoody glacier, 18f
elm, American, 53, 328n8
Diplomystus dentatus, 17f
El Nio, 31
disaggregating pheromones, 184
endangered species: as focus of conservation biology, 910; in Laramie
disturbances, 3638; effects of suppression of, 37, 94; examples of,

Dodge, R. I., 275
Doering, W., 223
Basin, 29798; organizations monitoring, 325n5(ch1); water rights

and, 293, 298, 347n28; of Wyoming, 9. See also specific species
Endangered Species Act: in conservation biology, 9; sage-grouse and,
128; wolves and, 264
doghair forests, 186f, 193, 193f
endemic species, 9
dominance, apical, 95, 198, 331n7
energy extraction: Development by Design and, 316; and ecosystem
Donahue, D., 127
services, 315; through fracking, 3078; and habitat fragmenta-
Dorn, R., 10f, 2425
tion, 31112, 312f, 313f; landscape changes caused by, 3; in
Douglas-fir woodlands and forests, 18890, 188f, 189f; changes
Laramie Basin, 28788; and multiple uses of public lands, 308;
in distribution of, 21920; common plant species in, 17677t;

fires and, 182, 189; in foothills, 16264, 165f; in Grand Teton
National Park, 253; ponderosa pine compared to, 18889;
subspecies of, 338n62; western spruce budworm on, 189f; in
Yellowstone National Park, 189f, 250, 251f
as percentage of state economy, 325n1(ch1); and population

growth, 305; by wind projects, 313f
energy flow, in grasslands, 85, 8992, 91t; in mountain forests,
200202, 201f, 202f, 203f; in sagebrush steppe, 116, 118
Engelmann spruce, 8f, 46, 19396, 194f
Douglas-fir beetle, 182, 189, 189f
Engle, D. M., 332n59
Driese, K. L., 342n3
entisols, 322t
drip irrigation, 62
Eocene Epoch, 14, 15, 245
drought(s): in alpine tundra, 235; bark beetle outbreaks and, 216;
ephemeral leaves, of sagebrush, 116
climate change and, 4041, 65, 328n29; in desert shrublands,
ephemeral streams, 45, 62, 65
13940; grasshoppers during, 100, 12021; in grasslands, 86,
erosion: in badlands, 147; beavers and, 59; in Black Hills and Bear
100, 332n62; history of, 24f, 32, 32f, 327n8; in sagebrush steppe,
Lodge Mountains, 266; in Laramie Basin, 285, 286, 287, 298f;
12021; in sand dunes, 14647; in sudden aspen decline, 219
livestock and, 60, 298f; in mountain forests, after fires, 2089,
drought tolerance: of grassland plants, 86; of invasive plants, 64, 65,

105, 329n56, 333n82
21213, 214, 340n49; in riparian landscapes, 51, 59, 60

escarpments, 15572; big game animals and, 15556, 16871; com-
dunes. See sand dunes
position of, 29; distinctive vegetation on, 29; dominant plant
dusky grouse, 335n96
species on, 156; human impacts on, 172; plant-animal interac-
dust storms, 100
tions on, 16872; woodlands on, 15764
Dutch elm disease, 328n8
EuroAmericans, arrival in Wyoming, 2324, 327n52
dwarf huckleberry, 17375, 190f
eutrophication, 240
dwarf mistletoe, 17879, 178f, 215, 338n19
evaporation: elevation and, 27; as percentage of water consumption,

62; ratio of precipitation to, 34, 35, 203; temperature and, 34. See
ecological regions of Wyoming, 8f; 2830

ecology, 47
ecosystem(s), 5; boundaries of, 7, 313; vs. landscape, use of terms, 7;
major components and interactions of, 56, 9f; map of, 8f
also evapotranspiration
evapotranspiration, 34; in mountain forests, 204; as percentage of
water consumption, 62; potential, 34; temperature in rate of, 34.
See also water availability
ecosystem ecology, 5
even-aged stands, 189, 190
ecosystem management, 31214
evergreen plants: in mountain forests, 174, 175, 196; in sagebrush
ecosystem productivity, net, 211f

ecosystem services, 1011; examples of, 10; in land management,
31415, 314f; value of, 11, 31415. See also specific ecosystem types
steppe, 116
Ewers, B., 119
extinction: climate change and, 40; in Cretaceous-Paleocene
edge forests, habitat fragmentation in, 31112
Extinction Event, 1314, 325n8; habitat fragmentation and, 309;
elevation: of alpine tundra, 230; of Grand Teton National Park, 246,
hunting as cause of, 20
247f, 253, 253f; of Greater Yellowstone Ecosystem, 246, 246f; of

Laramie Basin, 283f, 288; of mountain forests, 173, 174f, 180; of
Fahey, T. J., 339n3
mountain peaks, 27, 173; plant adaptations for, 27, 29f; and pre-
fairy rings, 84, 85f
cipitation, 27, 32f, 327n1; range of, 2728, 4f; and soil character-
Fall, P., 22
istics, 35; and temperature, 27, 33, 173, 327n10; of upper treeline,
farm bills, 329n1; of 2008, 11, 347n17
23031, 342n1, 342n3; Wyoming average, 3
feces deposition, in grasslands, 344n53
elevation map, 4f, 28f
Federal Land Policy and Management Act, 308
elk: in Black Hills, 280; distribution of, 3, 325n2(ch1); food prefer-
fellfields, 236; dominant plant species in, 237, 23839t; formation
ence of, 89; in foothills, 16871; in grasslands, 89; in Greater
of, 239
391
392 Index
fences: around national parks, 280; along roads, 311
21920; beetle outbreaks in (see bark beetle outbreaks); climate
fens, 7779, 77f; vs. bogs, 77; classification of, 78, 330n35; climate
change and, 41; distribution in mountains, 8f, 174f; dominant
change in, 80; dominant plant species in, 68, 7273t, 7779; soils
plants in, 17579, 17677t; edge vs. interior, habitat fragmenta-
of, 6870
tion in, 31112; elevation gradient in, 173, 174f, 180; energy flow
feral horses, in sagebrush steppe, 12728, 127f
in, 200202, 201f, 202f, 203f; expansion into meadows, 223,
Ferrenberg, S. M., 341n63
228, 229; fires in, 18182, 20815, 21618, 216f; future chal-
ferret, black-footed, 10f, 103, 106, 107, 279, 332n76, 346n48
lenges in, 21520; vs. grasslands, 88; historical human impacts
fertilizers, added to grasslands, 92, 93, 106
on, 2526, 21115, 212f, 213f, 30812; hydrology of, 2027, 211,
fescue, Idaho, 50f, 121, 123, 159t, 164, 166, 176t, 194f, 221, 222t,
204f; in Laramie Basin, 29192; major types of, 18598, 338n52;
224f, 250, 252f, 254
management options for, 21719; nutrient cycling in, 17475,
fir, Douglas-. See Douglas-fir
201, 201f, 203f, 2078, 210, 211, 21415; parasitic plants in,
fir, subalpine, 19396, 194f
17879; plant-animal interactions in, 17980, 202; precipitation
fir, white, 339n81
in, 2034, 204f, 21516; slow vs. rapid changes to, 181; soils of,
fire scars, 187, 187f
174, 200202, 2056, 210, 341n7; timber harvesting in, 21115,
fires: bark beetle outbreaks and, 21617, 341n64; in Black Hills, 187,
212f, 213f; tree growth rates in, 200201, 201f, 202f; understory
27478; climate change and, 41, 26162; in desert shrublands,

139, 140; exclusion vs. suppression of, 181, 338n27; in foothills,
density in, 178; in Yellowstone National Park, 189f, 192, 193,

250, 261f. See also specific types of forest
157, 162, 164, 16566, 170; frequency of, 121, 181, 216f; future
forest floor. See O horizon
of, 26162; in grasslands, 37, 85, 98100, 107, 334n32; in Greater
Forest Service, lands administered by, 6f
Yellowstone Ecosystem, 216, 25862, 260f; intensity of, 181, 182,
Fort Union Formation: badlands in, 336n18; coal deposits in, 326n13
208; invasive plants and, 278, 346n47; management options for,
Fossil Butte National Monument, 15, 17f
21718, 276; in mountain forests, 18182, 20815, 21618, 216f;
fossil fuels. See energy extraction
Native American use of, 23, 37, 98, 99; as natural disturbance, 37,
fossil ice wedges, 21f, 151
121, 262; in riparian landscapes, 55; in sagebrush steppe, 37, 110,
Fossil Lake, 15
112, 118, 119, 12123, 122f, 123f, 334n32; in sand dunes, 143;
fossils: in badlands, 147; Cenozoic, 1423; Cretaceous-Paleocene
severity of, 18182, 208, 338n28; vs. timber harvesting, effects
Extinction Event in, 14; Mesozoic, 13; Paleozoic, 1213; tree, 14,
of, 21115, 212f, 213f, 340n27; at upper treeline, 343n45
15, 245, 247f
Firewise recommendations, 218
Fountain Formation, 287, 295
first-order streams, 46
foxtail, Garrison creeping, 29899, 330n44
fish: fossils of, 15, 17f; grazing and, 60, 61f
fracking, 3078
Fish and Wildlife Service, U.S., and endangered species, 325n5(ch1);
fragmentation. See habitat fragmentation
in Laramie Basin, 293; National Wetlands Inventory of, 66; in
Fremont, John, 125, 28485
safe harbor agreements, 316, 347n21; on sage-grouse, 128
Freudenthal, Dave, 306
Fisk, M. C., 342n37
Friggens, M., 41
Fisser, H. G., 337n10
fringed sagewort, 108
flambellate grasshopper, 101f
Frontier Formation, 30f
Flaming Gorge Reservoir, 60, 162, 337n4
Front Range, 241
flood irrigation, 62, 296
frost boils, 237, 239f
floodplains: former, as terraces, 51; glacial outwash, 19, 5152; rapid
frost pockets, 186
changes of, 55; soils of, 45, 51. See also riparian landscapes
floods in riparian landscapes: changes caused by control of, 54, 55,
6162; tree growth after, 5253, 54, 5556, 328n11
food webs: belowground, in grasslands, 34, 90, 91t, 107, 331n16;
fuel breaks, 218

fungi: bark beetles and bluestain, 182f, 184, 338n38; in grasslands,
84, 85f, 89, 331n9; in riparian landscapes, 64; at upper treeline,
234
plant-cow-human vs. plant-human, 9092; in sagebrush steppe,

118; in wetlands, 68. See also energy flow
foothills, 15572; big game animals in, 15556, 16871; dominant
Gage, E., 330n40

Gambel oak, 168
plant species in, 156, 156f, 15859t; evapotranspiration in, 34;
Gannett Peak, 18f
fires in, 157, 162, 164, 16566, 170; formation of, 155; grasslands
Gardner, Dudley, 24
in, 156, 156f, 166; human impacts on, 3, 172; in Laramie Basin,
Gardner saltbush, 132f, 13537, 138f
290f, 29192; loss of habitat in, 168, 17071, 172; mountain-
Garrison creeping foxtail, 29899, 330n44
mahogany shrublands in, 156; plant-animal interactions
gas development. See energy extraction
in, 16872; riparian zones compared to, 155; roads in, 311;
genets, 197
shrublands in, 156, 156f, 157, 16466, 165f, 290f; topography of,
geobotany, 294; of Laramie Basin, 29495, 296f, 297f
15556; woodlands in, 156, 156f, 15764, 16668; of Yellowstone
geologic history, 1226, 12f. See also specific place names
National Park, 250, 251f
Geringer, Jim, 306
Forbis, T. A., 342n21
Geyers willow, 50f
Forelle Limestone, 295, 297f, 298f
geyser basins, of Yellowstone National Park, 251
forests, 173220; adaptations to, 17475, 17980, 181, 182; animal
girdling, 184
species in, 17980, 179t, 19899; area covered by, 8f, 215,
glacial outwash plains, 19, 52, 254, 285
Index
glaciers: and climate change, 40; and floodplains, 19, 5152; forma-
of, 85, 87f, 94, 9697, 332n48; in prairie dog colonies, 9798,
tion of, 19, 20; in Greater Yellowstone Ecosystem, 24648;
1023, 27980; in riparian landscapes, 6061, 61f, 329n30; on
history of, 1723, 326n23; impact on landscapes, 18f, 19, 19f; in
sagebrush, 116, 118, 12528, 333n18, 334n19; in wet meadows,
Laramie Basin, 282, 285, 287; oxygen isotope data from ice cores
of, 328n25; present-day, 20
76
greasewood, 56f, 58f, 74, 74f, 59, 68, 83, 110, 13235; adaptations
gleyed soil horizons, 68
of, 13133; in badlands, 14748, 149f; cheatgrass and, 140;
global warming, use of term, 38. See also climate change
disturbances in, shrublands, 13940; in foothills, 156f, 166; as
gold mining, 63, 266
halophyte, 132; in Laramie Basin, 290f, 292, 294f; seed germina-
Gondwanaland, 13f
tion in, 13334, 335n10; shrublands of, 4749t, 13334, 13637t;
gophers, pocket: bison and, 332n50; in grasslands, 1012; in mima
water availability in, 13739; in Yellowstone National Park, 250
mound formation, 151; in mountain meadows and tundra, 225,
Great Basin wildrye, 164
239, 240f
Great Divide Basin, 28; desert shrublands in, 131f, 133f, 138f; lack of
Goshen Hole, 74
outlet in, 346n1; mud springs in, 14849
Gosiute, Lake, formation of, 1517
greater sage-grouse. See sage-grouse, greater
grama, blue, 36, 8384, 88, 94, 100, 109f, 118, 13435, 136t, 150,
Greater Yellowstone Ecosystem (GYE), 7, 24565; alpine tundra in,
155f, 159t, 268, 273, 289, 291f, 336n21
248, 249; animal species in, 245; bark beetle outbreaks in, 185,
grama, sideoats, 163, 277, 331n2, 332n62
216, 26263, 340n39; boundaries of, 313; dominant plant species
Grand Teton (mountain), 19f, 246, 247f, 251
in, 248; elevation map for, 246f; elk in, 25658, 257f, 26465;
Grand Teton National Park, 25156; avalanches at upper treeline
fires in, 216, 25862; foothills of, 165f, 169f; geologic history of,
of, 233f, 253; boundaries of, 252; dominant plants in, 25254,
24548; management issues in, 25665; mountain meadows in,
253f; elevation of, 246, 247f, 253, 253f; establishment of, 3, 252;
228; riparian zones of, grazing in, 6061, 258; wolves in, 256,
glacial moraines in, 19f; livestock grazing in, 252; mountain

forests in, 19f, 188f
Granite Mountains, 15
26465, 344n93, 344n95. See also specific areas

greenhouse gases: as cause of climate change, 40; in Cenozoic Era,
15. See also carbon dioxide
grasshopper, flambellate, 101f
Green River, 51f
grasshoppers: during droughts, 100, 12021; in grasslands, 100101;
Green River Basin, 15, 32f, 77
number of species of, 332n66; outbreaks of, as disturbances, 37;
Green River Formation, 15, 326n19
in sagebrush steppe, 12021
Green River Lakes, 183f
Grassland National Park, 107, 333n91
green tree retention, 214
grasslands, 8f, 83108, 84f; adaptations to, 8487; agriculture in,
Grimes, K., 327n10
83, 92, 93, 100, 1078; animals of, 8687, 88t; in Black Hills,
Grinnell, George Bird, 343n38
84f, 270, 27374; burrowing animals in, 84, 1013; classification
grizzly bear, 26364, 263f; distribution of, 3, 263, 325n2(ch1); pine
of, 83; climate change and, 1068; conservation and restora-
seed consumption by, 172, 234, 263; threats to, 26364
tion of, 1067, 333n94; desert, 133, 13637t; distribution of, 83;
Gros Ventre landslide, 25556, 255f
dominant plants in, 8384; drought in, 86, 100, 332n62; effects
Gros Ventre Mountains, 25456; dominant plant species in, 25354;
of climate change on, 1078; effects of grazing on, 95f, 105f, 298f
formation of, 15, 25455; landslide in, 25556, 255f
(see also grazing and browsing; livestock grazing); energy flow
Gros Ventre River, 56, 255
(food chain) in, 85, 8992, 91t; fairy rings in, 84, 85f; fertilizers
ground squirrels: in alpine tundra 239; in desert shrublands, 139; in
added to, 92, 93, 106; fires in, 37, 85, 98100, 98f, 107, 334n32;
grasslands, 103; Townsends, 139; Wyoming, 239f
foothill, 156, 156f, 166; future challenges in, 1068; in geologic
groundwater, 3078; percolation to, 206, 3078. See also aquifers
history, 17; grasshoppers in, 100101; invasive plants in, 95,
grouse, species of, 128; dusky, 335n96; sharptailed, 335n96. See also
1036, 333n82; in Laramie Basin, 85f, 99f, 285, 289, 290f, 291f,
sage-grouse, greater
29899; national, 3, 1067; northern mixed prairie, 331n1,
grouse, sage-. See sage-grouse
333n2; nutrient cycling in, 9294, 99; precipitation in, 32, 89;
Gruell, G., 98
resource partitioning in, 8789; sagebrush steppe compared to,
gullies, development of, 24, 5960
117; sagebrush steppe mixed with, 110, 112, 114f; soils of, 34,
GYE. See Greater Yellowstone Ecosystem
83, 8485, 86f, 9293, 108; succession in, 94; tallgrass prairie,
gypsum, 285, 287
331n2; topography of, 84; in Yellowstone National Park, 250,

258, 344n53
Gray, S., 23, 41
Grayson, D. K., 336n38
gray wolf. See wolves
grazing and browsing: in Black Hills, 274, 27980, 345n27; in desert
habitat fragmentation, 30912; causes of, 31012, 313f; climate

change and, 40; effects on wildlife, 30910; in foothills, 172; in
sagebrush steppe, 128
habitat loss: in foothills, 168, 17071, 172; in wetlands, 66, 79,
330n40. See also habitat fragmentation
shrublands, 139, 140, 336nn3536; as disturbance, 327n18;
Hadley, J., 233
effects of, on grasslands, 8889, 9498, 95f, 332n33; and fire,
half-shrubs, 331n3
99100; in foothills, 157, 16162, 16871; and grasshoppers, 101,
halophytes, 36; in desert shrublands, 132; in Laramie Basin, 292,
332n71; in Greater Yellowstone Ecosystem, 6061, 252, 25658;

and invasive plants, 105, 105f; by livestock (see livestock grazing); in mountain meadows, 227, 342n20(ch13); plant tolerance
294f; in playa wetlands, 74; salt excretion by, 132, 132f; on terraces of riparian landscapes, 52
Hansen, A., 265, 314, 343n45, 344n50
393
394 Index
hardpan (caliche), 35
inland saltgrass, 134, 335n14
hardwood draws, 167. See also woody draws
insect(s): climate change and, 41; disturbances caused by, 3738;
Harmony Bench, 287, 288f

Harney Peak, 266
in mountain forests, 180. See also bark beetle outbreaks; specific

types
Hart, R., 25
instream flows, 307, 329n37
harvester ants, 103, 104f, 139
intercalary meristems, 85
Hat Creek Breaks, 163f
interception, 204
Httenschwiler, S., 225
interior forest, fragmentation of, 31112
Hattie, Lake, 288f, 289, 347n24
intermountain basins, 5f; elevation of, 28; evapotranspiration in, 34;
hay: as dominant crop, 325n1(ch1); in Laramie Basin, 285, 288, 296,

29899, 346n11
Hayden, Ferdinand, 25, 148, 149, 249, 251
Hayden Valley, 250, 252f
geologic history of, 13, 15, 16f, 282; map of, 5f; precipitation in,
average annual, 30, 31f; river outlets of, 282, 346n1; topography
of, 4f, 2829. See also specific basins
invasive plants: adaptations of, 104, 105, 333n81; in Black Hills,
Hayman fire (2002), 210, 338n28
27879; control of, 65, 79, 104, 1056, 124, 329n50; vs. crops,
Hedges, Cornelius, 249
329n42; in desert shrublands, 140; drought tolerance of, 64, 65,
herbicides, for invasive plants, 65, 104, 106
105, 329n56, 333n82; fires and, 278, 346n47; in grasslands, 95,
herbivores. See grazing; specific animals
1036, 333n82; in Laramie Basin, 29899; in riparian land-
hibernation, 236, 338n14
scapes, 6365; in sagebrush steppe, 121, 12224; in sand dunes,
Hicke, J., 21617
147, 336n17; in wetlands, 79. See also specific plants
High Savery Reservoir, 308f
inverse texture effect, 35, 142
highways. See roads
irrigation, 6263; area of land with, 62; in Laramie Basin, 80,
Hobbs, T., 265

hogback ridges, 285, 295
Hogback Rim, 26870
Holladay, B., 285
29697, 346n17, 347n24; wetlands supplied by, 67, 74, 76, 80,
346n17. See also agriculture
islands of fertility: in desert shrublands, 139; in sagebrush steppe,
115, 118, 119
Holocene Epoch, 2022, 326n36

Honeycomb Buttes, 147f
Jack Creek, 50f
hopsage, spiny, 112, 134
jackrabbits, 139; white-tailed, 139f
horehound, white, 27879, 279f
Jackson, David, 251
horses, feral: human use of, 23, 24, 127; in sagebrush steppe, 12728,
Jackson, Stephen T., 22, 41
127f; species of, 127

hot springs, in Yellowstone National Park, 251
Houston, D., 258
huckleberry, dwarf, 17375, 190f
humans: arrival in North America, 20, 326n29; early impact on
Jackson, William Henry, 2526, 125f, 249, 251

Jackson Hole, 25156, 254f; dominant plant species in, 25254;
geologic history of, 24546, 248, 248f; riparian landscapes of, 52;
temperature inversions in, 34
Jackson Lake, 252, 343n16
landscapes, 2326; livestock vs. plants in diet of, 9092; popula-
Jackson Lake Dam, 329n34
tion in Wyoming, 305
Jakubos, B., 342n20(ch13)
hummocks, 76
James, J. J., 336n27
Hunt, Wilson Price, 24, 126
Jasper Fire, 277
hunting: extinctions caused by, 20; in Greater Yellowstone
Jenney, W. P., 27475
Ecosystem, 256, 258; by Native Americans, 24, 332n41
Jenny Lake, 19f, 188f
Hutton Lake National Wildlife Refuge, 67f, 74
Jewel Cave National Monument, 270, 277
hybrid cattail, 79, 331n47
Johnson, D., 151
hydraulic fracturing, 3078
Johnson, J. R., 345n9
hydraulic redistribution (lift), 11516, 132
Johnson, K., 2526
hydric soils, 68, 70, 75
Jonah gas field, 316
hydrophytes, 45, 76
juniper, common, 145, 158t, 176t, 183f, 271, 272, 277, 295
hyphae, 202
juniper woodlands, 8f; browsing on, 171f; expansion and infilling
Ice Age. See Pleistocene Epoch
juniper, Rocky Mountain, 23f, 157, 160, 161f, 162, 337n28; browsing
of, 22, 23f, 15762, 15859t, 161f, 337n13

Ice Slough, 7778, 78f
on, 171f
Inceptic Haplocryalf soils, 341n7
juniper, Utah, 22, 23f, 157, 160, 161f, 162
inceptisols, 174, 322t
Jurassic Period, 12f, 13
incised channels, 46, 51f

increaser species, 95
Kashian, D., 209, 340n30
index, site, 339n2
Kay, C., 256
infilling, of juniper, 16162
Kentucky bluegrass, 79
infiltration rates of soils, 3132, 35
kettles, 70, 71f, 248
Ingelfinger, F., 347n9
Keyhole Reservoir, 280
Index
keystone habitats, wetlands as, 66
levees, 62
keystone species, 57; beaver as, 57; prairie dog as, 102
Lewis, Meriwether, 37, 96, 249
Khan, M. A., 335n10
Libby Flats, 228, 232f, 239f
Killpecker Sand Dunes, 142, 143, 144f, 145, 146f
lichens: in alpine tundra, 235; in grasslands, 92, 93f
Kimble, D. S., 344n99
lightning-caused fires, 37; in Black Hills, 275; in grasslands, 98, 98f,
Knopf, F. L., 328n13, 329n33
332n55; in Greater Yellowstone Ecosystem, 259
Krner, C., 241
limber pine. See pine, limber
Kranz, J. J., 338n16
limestone: in Laramie Basin, 285; origin of, 12
Krueger, K., 98
Limestone Plateau, 270, 274
krummholz, 23133, 232f
Lincoln Monument, 292

litter. See O horizon
LAI. See leaf area index
Little Bighorn Battlefield National Monument, 123
Lake Creek, 58f
Little Ice Age, 20, 23, 24f
lake trout, 26364, 344n85
Little Laramie River, 285, 287, 293f, 296
Lamar River valley, 247f, 250, 251f
Little Sage Creek, 50f
lanceleaf cottonwood, 53, 328n8
livestock: arrival of, 25; current number of, 327n62; food preferences
landscape(s): boundaries of, 7; changed vs. unchanged areas of, 3;

vs. ecosystem, use of term, 7; geologic history of, 1226, 12f;
glaciers impact on, 18f, 19, 19f; protective, 314, 347n14; rate of
recent changes to, 3056
of, 89; in human diet, 9092; rise and fall in number of, 25; wolf
predation on, 264
livestock grazing: climate change and, 41; as disturbance, 327n18;
erosion caused by, 60; in grasslands, effects of, 8889, 9498, 95f,
landscape ecology, 7
105f, 298f; historical impacts of, 25, 327n60; in Laramie Basin,
landslides, in Gros Ventre Mountains, 25556, 255f
285, 29596, 298f; on public lands, 127; in riparian landscapes,
Langford, Nathaniel, 343n41
6061, 60f; in sagebrush steppe, 127
La Nia, 31
locust, Rocky Mountain, 101
lapse rate, adiabatic, 33, 173, 327n10, 337n2(ch11)
lodgepole pine. See pine, lodgepole
Laramide orogeny, 13, 14, 15
loess, 19, 142
Laramie (city), 285, 346n13
Logan, J., 263
La Ramie, Jacques, 28283
loosestrife, purple, 79
Laramie Basin, 282301; climate change in, 299301, 299f, 300f; cli-
Lorenz, D., 170
mate of, 28889, 289f; dominant plant species in, 28993; effect
Lower Slide Lake, 255
of livestock grazing in, 298, elevation of, 283f, 288; fires in,
low sagebrush, 112
99f; geobotanical relationships in, 29495, 296f, 297f; geologic
luxury consumption, 210
features of, 282, 284f, 28588, 288f; geologic history of, 282,
285, 287, 346n3; geologic map of, 286f; grasslands in, 85f, 99f,
MacGillivrays warbler, 344n45
285, 289, 290f, 291f, 29899; human history in, 28285, 29596;
Mack, R., 96, 332n41
irrigated land in, 290f; management issues and future challenges
MacMahon, J. A., 337n2(ch11)
in, 295301; maps of, 283f, 286f, 290f; mima mounds in, 149,
Madison Limestone, 12, 270
150f, 151, 293; playa wetlands of, 7475, 74f, 29293; rainshadow
Madson, C., 306
effect in, 288; sagebrush in, 11011, 289, 292; vegetation patterns
mahogany, mountain-. See mountain-mahogany
in, 28993, 290f; wetlands supplied by irrigation in, 80, 29697,
mammals: in alpine tundra, 236; current population of, 24; early
346n17
Laramie Mountains: foothills of, 161f, 164f; forests of, 186f; mima
mounds in, 149
Laramie River, 53, 283f, 285, 296, 297, 328n10. See also Little
Laramie River
human impacts on, 20, 2326; in geologic history, 1317, 20; in

grasslands, 8687, 88t, 89; in Greater Yellowstone Ecosystem,
245; in mountain forests, 17980, 179t, 182, 198; in wetlands,
Mammoth Hot Springs, 251
Larocque, Franois A., 327n52
mammoth, 20, 282
Larson, G. E., 345n9
Mann, C., 326n49
Lauenroth, W. K., 90, 96, 331n1, 331n8
Marcott, S. A., 326n36
Laurasia, 13f
Marshall, K., 26465
La Vrendrye brothers, 327n52
marshes, 7074; climate change in, 80; dominant plant species in,
layering, 194, 194f, 232, 342n6
7074, 7273t; invasive plants of, 79; soils of, 6870
leaf area index (LAI), 65, 204, 205f, 210f
Martner, B. E., 327nn78
leafy spurge, 104
mastodons, 282
Lebo, Lake, 15
McIntyre, N. E., 333n93
leks, sage-grouse, 12829
McNaughton, S., 94, 95, 97
Leopold, Aldo, 256, 320, 343n38
Mead, Matt, 306
Leopold, A. Starker, 256
meadows: riparian, 4749t, 54; sagebrush, 22223t; saltgrass, 133,
Leopold report, 25659

Lesica, P., 329n50
134, 13637t. See also mountain meadows; wet meadows

meandering channels, 46, 51, 51f, 56. See also riparian landscapes
395
396 Index
Mears, Brainerd, Jr., 295
distribution of, 111, 112; in foothills, 164; in Grand Teton
Medicine Bow Mountains: alpine tundra and upper treeline in,
National Park, 254, 255f; recovery after disturbances in, 120; in
230, 230f, 231, 232f, 233, 239f, 342n34, 343n45; fens in, 68;
foothills of, 165f; forests in, 190f, 194f, 204, 211, 340n30;
Yellowstone National Park, 251f

mountain-mahogany: curleaf, 157, 160f, 337n4, 337n7, 345n23;
formation of, 15, 326n18; habitat fragmentation by clearcut-
shrublands of, 156, 157, 15859t, 160, 160f; true (alder-leaf),
ting in, 311f; meadows in, 223, 224f, 225, 225f, 227f, 228,
157, 160f, 337n4, 337n7; in Laramie Basin, 289, 291f, 292, 298f,
342n20(ch13)
Mediterranean tamarisk beetle, 329n50
345n23, 346n18
mountain meadows, 22129, 221f; in Black Hills, 273, 273f; climate
Mehringer, P. J., Jr., 334n32
change and, 22829; dominant plant species in, 22123,
meristems, 85
22223t; dry, 22223t; forest expansion into, 223, 228, 229;
Mesozoic Era, 12f, 13, 13f
forests intermingled with, 180; formation of, 223, 225; livestock
Meyer, G., 327n54
grazing on, 227, 341n16, 342n20(ch13); snowglade, 22527; soils
middens, 22, 22f, 23f
of, 223, 225f, 341n7; wet, 75; in Yellowstone National Park, 228,
Middle Rocky Mountains, 30

Mielke, H. W., 332n50
250, 252f
mountain pine beetle, 181f, 18285; in Black Hills, 27778; climate
migratory birds, 68, 330n9
change and, 216, 341n63; effects of, 18285, 182f, 183f, 219,
Mikkelson, K. M., 340n45
340n45; in Greater Yellowstone Ecosystem, 26263. See also bark
Milchunas, D., 90, 96
beetles
Miles, S., 329n50
mountain silver sagebrush. See silver sagebrush
milkvetch, two-grooved, 335n18, 346n22
Muddy Creek, 58f
Millar, C. I., 341n88
mudflats, 7475
mima mounds, 14951, 150f, 293
mud springs, 14849, 149f, 336n19
mining: gold, 63, 266; in Laramie Basin, 285, 287; peat, 79
mule deer: decline in population of, 17071; food preferences of,
Minnelusa Foothills, 270
89; in grasslands, 89; in Greater Yellowstone Ecosystem, 257;
Minshall, G. W., 340n36
mortality on roads, 310, 347n10; roads avoided by, 310Mullen
Miocene Epoch, 17, 325n6(ch2)
Creek, 59f
Missoula, Lake, 19
multiple use on public lands, 3089, 313
Missouri River, 329n35
Munn, L., 151
mistletoe, dwarf, 17879, 178f, 215, 338n19
Munroe, J. S., 341n7
Mitchell, J., 25
mushrooms, in forests, 202, 339n5
Mitton, J. B., 341n63
mustangs, 127. See also horses
mixed desert shrublands, 133, 13435, 135f, 13637t, 139
mustard, Sahara, 336n17
mixed foothill shrublands, 15859t, 16466, 165f
mycoheterotrophs, 175
mixed-grass prairie, 83, 84f; adaptations to, 85; agriculture in, 100;
mycorrhizae: in forests, 174, 175, 175f, 202; mushrooms produced
climate change in, 108; distribution of, 83; dominant plant

species in, 83; drought in, 100; fires in, 99, 99f, 332n59; in
by, 339n5
Myers, W., 295
Laramie Basin, 289, 291f; nutrient cycling in, 92; precipitation

in, 89. See also grasslands
Naftz, D. L., 328n25
Mock, K. E., 339n97
narrowleaf cattail, 79, 330n47
mollisols, 83, 322t
narrowleaf cottonwood, 46, 51f, 56, 328n10, 328n16, 329n56
Monsoon, Arizona, 31, 196
National Academy of Sciences, 258, 308
Mooney, H. A., 342n34
National Climate Assessment program, 40
Moore, J. C., 331n16
National Climate Data Center, 289f, 299
moose, 343n45
National Elk Refuge, 168, 257
Moran, Thomas, 7, 249
National Environmental Policy Act, 309, 315
Morgan, J., 108
National Fire Plan, 218
Mormon cricket, 37, 101, 332n73
national grasslands, 3, 1067
Morrison Formation, 325n6(ch2)
national parks, 6f. See also specific parks
Mortenson Lake National Wildlife Refuge, 293
National Park Service, lands administered by, 6f
mosses, in wetlands, 68, 77, 78, 330n36
National Research Council, 308
Mount Sheridan, 250f
National Weather Service, 40
Mount Washburn, 252f
National Wetlands Inventory, 66
mountain(s), 5f; droughts in, 32; elevation of peaks of, 27, 173;
national wildlife refuges, in Laramie Basin, 29293, 297
evapotranspiration in, 34; fens in, 77; foothills of (see foothills);
Native Americans: arrival of ancestors of, 20; in Black Hills, 266,
formation of, 173; geologic history of, 1317, 173; map of, 5f;
345nn12; fires used by, 23, 37, 98, 99; horses used by, 127; hunt-
marshes in, 70; meadows in (see mountain meadows); other eco-
ing by, 24, 332n41; impact on ecosystems by, 23, 24, 326n49; in
systems affected by, 173; precipitation in, 3032; riparian plants
Laramie Basin, 28285; use of sand dunes, 145; in Yellowstone,
in, 46; topography of, 2830. See also specific mountains

mountain big sagebrush, 111; climate change and, 22829;
249, 343n16
naturalized species, 63, 79
Index
natural regulation policy (Yellowstone National Park), 256,
25758
packrats. See bushy-tailed woodrat

Pahlow strath, 288f
Natural Resources Conservation Service, 128, 204
Painter, E., 95
Nature Conservancy, The, 315
Paintner, W. W., 345n27
Nebraska Sand Hills, 143, 336n10
Paleocene-Eocene Thermal Maximum (PETM), 15
nematodes, 331n9; in grasslands, 86, 89, 98, 331n9
Paleocene Epoch, 13, 1415
net ecosystem productivity (NEP), 211f
paleoecology, 2023, 326n32
net primary productivity (NPP), 201, 327n4; aboveground, 90f,
paleosols, 146
210f, 340n29; in desert shrublands, 335n3; in grasslands, 89, 90f,
Paleozoic Era, 1213, 12f
95f; in forests, 201, 201f, 209, 211f, 339n2, 340nn2931
Palmer Drought Severity Index, 32f
Newton, H., 27475
pantodonts, 14
New West, 306
parasitic fungi, at upper treeline, 234
Niobrara Formation, 29495
parasitic plants, in forests, 17879
nitrogen: in alpine tundra, 23941, 342nn3637, 343n40; in desert
Parkman, Francis, 285
shrublands, 139, 336n28; fixation, 207; in grasslands, 9294, 93f;
parks. See mountain meadows
in forests, 201, 201f, 2078, 207f, 210, 211, 340n36; in riparian
patch-burn grazing system, 100
landscapes, 64; in sagebrush steppe, 119; in sand dunes, 145,
patterned ground. see cryoturbation
336n14; in wetlands, 68
Pawnee National Grassland, 3
nitrogen hotspots, 240
Pearson, J. A., 339n2
nitrogen saturation, 240
peat, in fens, 7778, 330n32
nivation hollows, 114f, 156
peat mining, 79
Niwot Ridge, 342n21, 342n31
Pendall, E., 119
nonriparian wetlands. See wetlands
pennycress, alpine, 235
northern mixed prairie, 331n1, 333n2. See also grasslands
P-E ratio. See precipitation-evaporation ratio
Northern Rocky Mountains, 30
Perry, D., 338n4
northern winter range of Yellowstone National Park, 251f
Persico, L., 327n54
North Fork Fire, 260
Peters, D. P. C., 332n75
North Platte River, 61, 328n13
PETM. See Paleocene-Eocene Thermal Maximum
Northwest Territory sedge, 70, 77f, 78
petrified trees, 245, 247f
Notzold, R., 333n83
Pfeifer, E. M., 340n39
NPP. See net primary productivity
PHACE. See Prairie Heating and Carbon Enhancement Experiment
Nugget Sandstone, 13
pheromones, aggregating and disaggregating, 184
nutcracker, Clarks, 17172, 171f, 179, 202
Phillips, V. D., 342n34
nutrient cycling: in alpine tundra, 235, 23941; in desert shrub-
photographs, early: in Black Hills, 345n15; information on landscape
lands, 139, 336n27; in grasslands, 9294, 99; in forests, 17475,
change in, 2526
201, 201f, 2078, 210, 211, 21415; in sagebrush steppe, 119; in
phragmites (common reed), 79
sand dunes, 14546
phreatophytes, 57, 65
nutrient enrichment, in riparian landscapes, 60, 64
physiological ecology, 5
nutrient resorption, 208, 235
pickleweed, 132f
pika, American, 236, 236f, 242
oak, bur, 168, 27273, 273f
pimple mounds. See mima mounds
oak, Gambel, 168
pine, bristlecone, 234, 342n12, 342n15
oak woodlands, 15859t, 168
pine, limber: arrival of, 22; in foothills, 16264, 164f, 17172; in
obsidian, 249
Laramie Basin, 292, 293f; seed dispersal by birds, 17172; in
Ocean Lake, 74
upper treeline, 233; whitebark pine compared to, 342n3; white
Odum, Eugene, 347n14

offsets, conservation, 316
pine blister rust and, 234; woodlands of, 17677t

pine, lodgepole, 8f, 19093, 190f; adaptations of, 174, 175f; bark
Ogallala aquifer, 307, 336n10
beetle outbreaks and, 182f, 183f, 185, 216, 338n19; distribution
O horizon, 200
of, 8f, 174, 338n4; doghair stands of, 193, 193f; dominant plant
oil development. See energy extraction
species in, 17677t; fires and, 182, 191, 19293, 20810, 209f,
oil shale deposits, formation of, 15
210f, 211f, 213f, 216, 218; forests of, 19093, 190f, 191f; in Grand
old-man-of-the-mountain, 236f
Teton National Park, 253; hydrologic budget of, in forests, 2045,
Old Mans Beard, 272
205f, 206, 340n11; on infertile soils, 190, 250, 339n68; nutrient
Oligocene Epoch, 15
cycling in, in forests, 207; parasitic plants on, 17879; serotinous
olive, Russian, 6365, 63f
cones of, 17980, 19192, 192f, 339n71; timber harvesting in,
open spaces: persistence of, 306, 306f, 307f; value of, 314
213f, 21415; in Yellowstone National Park, 191f, 250. See also
Oregon Trail, 25, 78, 78f, 305, 327n60

Overland Stage Line, 285
Overthrust Belt, 13
forests
pine, pinyon: arrival of, 22; in foothills, 162, 17172, 337n19,
337n22; seed dispersal by birds, 17172
397
398 Index
pine, ponderosa, 8f, 182, 18588, 186f; arrival of, 22, 186; in Black
precipitation-evaporation (P-E) ratio, 34, 35, 203
Hills, 27072, 271f, 274, 275f, 276f, 281; climate change and,
Preisler, H. K., 338n42
281; dominant plant species in forests of, 17677t; Douglas-fir
Preuss, Charles, 125
compared to, 18889; fires and, 182, 186f, 187, 187f, 274, 275f,
pricklypear cactus, 96, 100, 102
276f, 277f, 338n59; in foothills, 155f, 156f, 16264, 163f, 337n26,
primary productivity, net. See net primary productivity
337n28; in Laramie Basin, 292, 293f; in forests, 182, 18588
private lands: area of, 6f; ecosystem management in, 31213, 31516;
pine, whitebark: bark beetle outbreaks and, 26263; limber pine

compared to, 342n3; seed dispersal by birds, 172; in upper
fires on, 218; in Greater Yellowstone Ecosystem, 265; safe harbor

agreements on, 316, 347n21
treeline, 231, 233, 234; white pine blister rust and, 234, 234f,
productivity, net primary. See net primary productivity
26263; in Yellowstone National Park, 250, 250f
pronghorn: in Black Hills, 27980; conservation of, 315f; food
pine beetle, mountain. See mountain pine beetle
preferences of, 89; in geologic history, 13, 17, 325n6(ch2); in
pine cones, serotinous, 17980, 19192, 192f, 339n71. See also pine,
grasslands, 83, 89, 106; mortality on roads, 31011, 347n10; in
lodgepole
Pinedale glacial period, 20, 24648, 326n23
pinedrops, 175
Pine Ridge, 163f
prairie dog colonies, 27980, 346n51; sagebrush consumption

by, 116; wolves and, 265
public lands: area of, 3, 6f, 319; grazing on, 127; multiple use on,
3089
pinyon pine. See pine, pinyon
puffballs, 202
Pioneer Canal, 347n24
pugging, 76
plains cottonwood, 46, 51f, 52f, 53, 54, 329n56
purple loosestrife, 79
Platt, P., 125

Platte River, 22, 297
quaking aspen. See aspen forests, woodlands
playas, desert, 132
Quaternary alluvium, 285
playa wetlands, 7475, 74f; dominant plant species in, 7273t, 7475,
Quaternary Period, 12f, 1721
74f; in Laramie Basin, 7475, 74f, 29293; soils of, 6870

Pleistocene Epoch, 1720, 21f, 325n6(ch2)
radiocarbon dating, 22
Pliocene Epoch, 17
railroads, 24, 25, 285, 305, 329n40
pocket gophers: in alpine and subalpine meadows, 225, 239, 240f;
rainfall. See precipitation
bison and, 332n50; in grasslands, 1012; in mima mound forma-
rainshadow effect, 17, 31, 288
tion, 151; in mountain meadows, 225
ramets, 197
point bars, 46, 51, 52f, 55, 55f, 56f
rangelands. See livestock grazing
pollen record, 20, 21f, 22, 23f, 40, 326n32
Raynolds, W. F., 53
ponderosa pine. See pine, ponderosa
redbeds, 13
ponds, in sand dunes, 145, 146f
Red Buttes, 285
porcupines, 202
Red Desert, 15, 131f, 139f
potholes, 70, 71f, 248
Redman, R. E., 332n59
Powder River Basin: badlands in, 14748, 336n18; coal deposits in,
red (pine) squirrel, 17980, 180f, 192, 234
14, 326n13; escarpments in, 155f; fires in, 98, 98f; geologic his-
red swampfire, 73t, 74, 74f, 132, 134, 137t, 292
tory of, 13; grasshoppers in, 101, 120; habitat fragmentation in,
Red Valley, 270
312; sagebrush steppe in, 111f, 120, 125, 335n81; sage-grouse,
reed, common, 79
130
reed canarygrass, 79
prairie. See grasslands
Rehfeldt, G., 215, 281
prairie dog, black-tailed, 102, 102f, 106, 27879, 280f
Reider, R. G., 151, 223, 326n48
prairie dog, white-tailed, 102
relictual treeline, 241
prairie dogs: in Black Hills, 27879, 279f, 280f; conservation efforts
reptiles: in forests, 179t; in geologic history, 13; in grasslands, 88t
for, 103, 106, 107, 333n78, 333n90; disturbance in grasslands by,

1023, 102f, 332n75; grazing in colonies of, 9798, 97f, 1023,
27980; invasive plants and, 27879, 279f
Prairie Heating and Carbon Enhancement Experiment (PHACE), 108
prairie mounds. See mima mounds
precipitation, 3032; average annual, 30, 31f, 40; in Black Hills and
Bear Lodge Mountains, 267, 270f; climate change and, 40, 41,
215, 328nn2930; cloud seeding and, 308; in desert shrublands,
131; elevation and, 27, 32f, 327n1; in grasslands, 32, 89; infiltration into soil, 3132; in Laramie Basin, 289, 289f, 299301,
reservoirs, 6162; Boysen, 55; development of, 307, 308f; effects of

livestock on, 60; Flaming Gorge, 60, 162, 337n4; functions of, 61;
High Savery, 308f; Keyhole, 280; negative effects of, 61
residential development: in Black Hills, 274, 276; fires in, 218, 276; in
foothills and escarpments, 172, 311; in forests, 218; in Laramie
Basin, 285, 298; in riparian landscapes, 62
resource partitioning: in desert shrublands, 13233; in grasslands,
8789
restoration: of desert shrublands, 141; of grasslands, 1067, 333n94;
of sagebrush steppe, 127
300f; map of, 31f; in forests, 2034, 204f, 21516; in mountain
rest-rotation grazing, 97
meadows, 228; in mountains, 3032; in plant growth, 30, 33;
return flows, 62, 63
range of, 30; ratio of evaporation to, 34, 35, 203; in sagebrush
Reynolds, D. N., 342n21
steppe, 11011, 112, 112f; in sand dunes, 145; seasonality of, 31,
rhizosphere, 70
327n7. See also snow
ribbon forests, 22527, 227f, 231
Index
Richards, J. H., 115, 331n8
Laramie Basin, 11011, 289, 290f, 292; longevity of, 116; old-
Richardson, M. T., 347n24
growth, 118; precipitation in distribution of, 3132, 11011, 112f;
Rickard, W. H., 140
recovery after disturbances in, 12021, 122f, 123f; saline toler-
ridges: hogback, 285, 295; windswept, 156. See also escarpments
ance of, 134, 335n16; varieties of, 11112, 113f; in Yellowstone
riparian landscapes, 4565; area covered by, 45; bank storage in, 57,
National Park, 114f, 250. See also sagebrush islands; sagebrush
59, 60; beavers in, 5760, 58f, 59f; in Black Hills, 272; channels
in, 46, 51, 51f, 5253, 56; climate change in, 65; development of,
46; dominant plant species in, 46, 4749t, 50f, 5257; ecologi-
steppe; other species of sagebrush

sagebrush, black, 110, 112, 114f, 119; distribution of, 112, 156; in
Laramie Basin, 289, 291f, 292, 346n18
cal role of, 45; foothills compared to, 155; future challenges in,
sagebrush, low, 112, 254, 255f
6365; in Greater Yellowstone Ecosystem, 6061, 258; human
sagebrush, mountain big, 111; climate change and, 22829; distribu-
impacts on, 3, 45, 6063, 329n40; invasive animals in, 329n42;
tion of, 111, 112; in foothills, 164; in Grand Teton National Park,
invasive plants in, 6365; landforms in, 4653; in Laramie
254, 255f; recovery after disturbances in, 120; in Yellowstone
Basin, 29293, 290f, 293f, 294f; livestock in, 6061; map of, 46f;
National Park, 251f
vs. nonriparian landscapes, water movement in, 45; percentage
sagebrush, sand, 83, 269t, 333n7
of species dependent on, 45; rapid changes to, 5557; reservoir
sagebrush, silver, 112, 114f, 134f; adaptations of, 116; in grasslands,
construction and, 6162; soils of, 45, 51, 52; succession of plants
in, 54, 56, 56f; water consumption by plants in, 57; wet meadows
in, 75; in Yellowstone National Park, 249, 25051
Ripple, W., 264
Roach, J. L., 333n90
roadless areas, 21718, 309
roads: animal avoidance of, 310, 347n9; animal mortality on,
83; recovery after disturbances in, 120; two varieties of (plains

and mountain), 112
sagebrush, threetip, 112, 115, 156, 157f, 164f, 166, 333n7
sagebrush, Wyoming big, 109f, 110f, 111; distribution of, 111, 112; in
foothills, 164; in grasslands, 83, 290f; recovery after disturbances
in, 120. See also sagebrush, big; sagebrush steppe
sagebrush islands, 11215, 115f
31011; in Black Hills, 274; habitat fragmentation by, 31012,
sagebrush meadows, 22223t
310f; map of, 7f; for timber harvesting, 31112
sagebrush steppe, 8f, 10930; adaptations in, 11517; agriculture
Rockefeller, John D., Jr., 252
in, 109; animals species in, 110, 128; climate change in, 120,
rocks: in alpine tundra, 23637, 239; effects on plant distribution,
121, 124; conservation and restoration of, 127, 12830; desert
29495; as parent material of soils, 3435
shrublands compared to, 131, 132, 133, 335n3; dominant plant
Rocky Mountain juniper, 157, 160, 161f, 162, 171f, 337n28
species in, 11012; drought in, 12021; energy flow in, 116,
Rocky Mountain locust, 101
118; fires in, 37, 110, 112, 118, 119, 12123, 122f, 123f, 334n32;
Rocky Mountain National Park: alpine tundra and upper treeline in,
in Grand Teton National Park, 254; grasshoppers and, 12021;
240, 241, 342n34; fires in, 194
grasslands compared to, 117; grasslands mixed with, 110, 112,
Roman warm period, 21
114f; grazing and, 116, 118, 12528; invasive plants in, 121,
Roosevelt, Theodore, 24
12224; islands of fertility in, 115, 118, 119; loss of, 128; nutrient
root systems: in alpine tundra, 235; in forests, 202, 203f, 206,
availability in, 119; precipitation in, 11011, 112, 112f; recovery
340n19; in grasslands, 8485, 86f, 88; in sagebrush steppe,
after disturbances in, 12024; sage-grouse in, 12830, 129f; soils
11516, 118, 119; in sand dunes, 145
of, 110; topography of, 110, 112; water availability and growth
Rotella, J., 265, 343n45, 344n50

ruminants. See grazing; specific animals
runout zone, 234
rushes, 69f, 72t
in, 11011, 11718; in Yellowstone National Park, 250, 251f

sage-grouse, greater, 12830, 129f; adaptations of, 12829; conservation efforts for, 306; sagebrush consumption by, 116; threats to,
12830
Russell, Osborne, 60, 249
Sage-grouse Initiative, 128
Russian olive, 6365, 63f
sage sparrow, 310, 347n9
rust, comandra blister, 17879
sagewort, birdfoot, 110f, 112f, 131, 131f, 135, 136t, 138f
rust, white pine blister, 17879, 234, 234f, 26263, 342nn1516,
sagewort, bud, 112, 112f, 131f, 133135, 138f
344n78
Ryan, M. G., 340n31
sagewort, fringed, 108, 222t, 224, 250, 278, 280, 289, 331n3
sagewort, threetip. See sagebrush, threetip
Sahara mustard, 336n17
Sacket, D. B., 53
salicornia. See red swampfire
SAD. See sudden aspen decline
salinity of soils, 35, 36, 327n17; vs. alkalinity, 327n17, 335n4; in
safe harbor agreements, 316, 347n21
desert shrublands, 13134, 132f, 139, 335n4; irrigation and,
sage, sand. See sagebrush, sand
6263; in Laramie Basin, 292; in playa wetlands, 7475; in ripar-
sagebrush, basin big, 111, 134f; distribution of, 111, 333n5; in sand
dunes, 143. See also sagebrush, big
sagebrush, big, 10930, 113f, 114f; adaptations of, 11517, 333n18;
ian landscapes, 52
saltbush, Gardner, 13537, 138f
saltbush shrubland, 133, 13537, 13637t, 138f, 139, 336n21
area covered by, 8f, 109, 290f; ecosystem of, 11719; in desert
saltcedar (tamarisk), 54, 56f, 61, 63, 6465, 64f, 329n50
shrublands, 134, 137f; in forests, parasitic plants on, 17879;
salt desert shrubland. See saltbush shrubland
fossil pollen from, 20, 21f; in Grand Teton National Park, 254,
salt flats, 7475, 76, 330n19
255f; grasshopper outbreaks and, 120; in grasslands, 83; grazing
saltgrass, inland, 134, 335n14
on, 125; greasewood and, 134; juniper expansion and, 162; in
saltgrass meadows, 133, 134, 13637t
399
400 Index
salt-secreting glands, 132f
Shinneman, D. J., 140, 275, 336n39, 339n99
salts in soils. See salinity of soils
Shirley Basin: black-footed ferrets in, 333n76; fossil logs in, 14; mima
salt-tolerant plants. See halophytes

sand bars, 52
Sandberg bluegrass. See bluegrass, Sandberg
Sand Creek, 29293
sand dunes, 8f, 14247; adaptations of plants for, 145; animals
mounds in, 149; sagebrush in, 111

shortgrass prairie, 83; agriculture in, 100; distribution of, 83; dominant plant species in, 83, 331n1, 331n8; fires in, 99; nutrient
cycling in, 92, 93. See also grasslands
Shoshone National Forest, 3
species in, 145; disturbances in, 143, 14647; dominant plant
Shoshone River, 54, 5556
species in, 14243, 336n3; factors in location of, 142; formation
shrubby cinquefoil, 48t, 50f, 68, 73t, 75, 75f, 76f, 77f, 167f, 183f,
of, 3435, 142; geologic history of, 13; invasive plants in, 147,
336n17; in Laramie Basin, 287, 293; in mixed desert shrublands,
13435; origin of sand in, 142; ponds in, 145, 146f; precipitation
221f, 223t, 229, 250, 252, 295

shrublands: desert (see desert shrublands); foothill, 156, 156f, 157,
16466; in Laramie Basin, 292, 294f; riparian, 4749t, 5455
in, 145; in sagebrush steppe, 112, 115f; soils of, 14243; stabiliza-
Shuman, B., 3840
tion process in, 143
Sierra Madre: forests in, 196, 196f, 197f; mountain meadows in, 227
sandhill crane, 328n13
Signal Mountain, 71f
sand sage (sagebrush), 83, 269t, 333n7
silver sagebrush. See sagebrush, silver
sandstones: geologic history of, 13; in Laramie Basin, 287, 287f; as

parent material of soils, 3435
silviculture. See timber harvesting

sinks: carbon, 210, 211, 340n39; wetlands as, 79
San Juan Mountains, 219
Sioux, 345n2
saprophytic fungi, 84
site index, 339n2
Saskatoon serviceberry, 16465
Skull Rim, 148f
Satanka Formation, 297f
skunkbush sumac, 47t, 5456, 121, 148, 15556, 158t, 16067, 166f,
Savage Run Wilderness, 309f
176t, 269t, 274, 292f
Schullery, P., 257, 265
slash, 21214, 212f
Schuman, J., 117
Slater, N., 125
scurfpea, slimflower, 145f
slickspot soils, 134
Seastedt, T., 241
slimflower scurfpea, 145f
secondary succession. See succession
Smith, B., 59
second-order streams, 46
Smith, E. W., 284
sedge(s): in alpine tundra, 23839t; in forests, 177t; in grasslands,
Smith, Jedediah, 24
85; in mountain meadows, 222t; in riparian landscapes, 48t; in
Smith, W. K., 225, 231, 233
wetlands, 68, 69f, 7071, 72t, 77, 77f, 78, 250; in Yellowstone
snags, 202f, 214
National Park, 250. See also specific types
Snake River: channelizing of, 6162; dam on, 252, 328n16; vegeta-
sedge, Northwest Territory, 70, 77f, 78

sedge, water, 70, 77f, 78
tion along, 52f, 253; Wild and Scenic Rivers Act protection of
headwaters, 307
Sedgwick, J. A., 328n13, 329n33
Snake River Canyon, 17
sediment deposition: in reservoirs, 61, 307; in riparian landscapes,
snow: in alpine tundra and upper treeline, 23234, 236, 237, 237f,
57
seed dispersal: in foothills, by birds, 17172; in forests, 178, 179; in
riparian landscapes, 54
seed predation: by birds, 171, 179, 202, 234, 262; in forests, 17980,
192; by grizzly bears, 172, 234, 263; at upper treeline, 234
selective timber harvesting. See timber harvesting
selenium: accumulation in plants, 135, 29495, 335n18, 346n22;
241; in avalanches, 23334, 233f; climate change and, 40, 41;

decomposition under, 202, 339n3; in in forests, 173, 2034,
204f, 206; grasslands, 32, 89; infiltration into soil from, 3132;
in mountain meadows, 22527, 226f, 228; in Pleistocene Epoch,
1720; sagebrush distribution affected by, 3132, 11011, 112; in
sagebrush steppe, 32, 111, 112, 114f, 115, 117, 117f; water content
of, 204. See also nivation hollows
in desert shrublands, 135, 139; indicator plants, 135, 29495,
snowglades, 22527, 226f
335n18, 346n22
snowmold, blackfelt, 225, 226f
serotinous cones, 17980, 19192, 192f, 339n71. See also pine,

lodgepole
serviceberry, in Saskatoon and Utah, 156, 158t, 16465, 165f, 176t,
233f, 295, 337n33
Severson, K. E., 338n16
shadscale, 116, 132f; adaptations of, 133; drought and, 13940; in
mixed desert shrublands, 134
Snowy Range, 230f. See also Medicine Bow Mountains

Sodergren Lake, 347n24
sodicity of soils, 327n17, 335n4
sodium in soils. See salinity of soils
soil(s), 3436; alkalinity of, 36, 327n17, 335n4; of alpine tundra and
upper treeline, 232; characteristics of, 3436; climate change and,
108; composition of, 34; of desert shrublands, 13134, 139, 335n4;
shales: geologic history of, 13; mountain meadows and, 224f
development of, 1920, 3435; fires and, 208, 210, 340n28; of
sharptailed grouse, 335n96
forests, 174, 200202, 2056, 210, 341n7; and geologic history,
sheep: in grasslands, food preferences of, 89; population of, 25,
1920; of grasslands, 34, 83, 8485, 86f, 9293, 108; infiltration
327n62
Sheep Mountain, 75f, 293f, 295
rates of, 3132, 35; of mountain meadows, 223, 225f, 341n7;

Ohorizon of, 200; orders of, by vegetation type, 32223t; parent
Index
material of, 3435; in plant growth, 3436, 36f; of riparian
Sully, Alfred, 101
landscapes, 45, 51, 52; of sagebrush steppe, 110; salinity of (see
sunflower, alpine, 236f
salinity); of saltbrush shrubland, 13537, 336n21; of sand dunes,
surface fires, 37; in Black Hills, 274, 275, 276; in Douglas-fir wood-
14243; subgroups of, 32223t, 324t; in temperatures ranges, 33;

transitions between types of, 34; of wetlands, 6870
soil crusts: in desert shrublands, 140, 336n39; in sagebrush steppe,
123, 124, 126, 126f
lands, 18890; intensity and severity of, 182, 208; in ponderosa

pine woodlands, 187. See also fires
sustainable land management. See land management
swampfire, red (salicornia). See red swampfire
solifluction, 23739, 240f, 342n33
Swamp Lake, 66f, 78
Soto, Hernando de, 127
swamps: in geologic history, 14; lack of, in Wyoming, 329n1
Soul, P. T., 337n13
Sweetwater Plateau, 7778
Southern Rocky Mountains, 30
Sweetwater River, 53, 55f, 125f
South Platte River, 328n13
sylvatic plague, 103, 106, 279, 332n76
Spackman, L., 151

sparrow, Brewers, 310, 347n9
tallgrass prairie, 331n2
sparrow, sage, 310, 347n9
Talluto, M., 17980
Spearfish Creek, 270
talus slopes, 236
Spearfish Formation, 13, 270, 345n10
tamarisk. See saltcedar
sphagnum, 14, 77, 250, 330n29, 330n36
Tatman, Lake, 15
spiny hopsage, 112, 13335, 136t, 137, 143, 144f, 335n2, 335n18,
tectonic uplift, 13, 14, 17
336n35, 346n22
springs: in Laramie Basin, 286; in Yellowstone National Park, 251
Telephone Canyon, 292

temperature(s), 3334; average high and low, 33, 33f, 3840,
sprinkler irrigation, 6263
39f, 328n25; elevation and, 27, 33, 173, 327n10; in evapo
spruce, blue, 46, 328n3
transpiration, 34; and fire frequency, 216f; inversions, 3334; in
spruce, Engelmann, 8f, 46, 19396, 194f
Laramie Basin, 288, 289f, 299, 299f, 301; map of, 33f; range of,
spruce, white, 272, 272f, 339n81
33; tolerance of mountain trees, 174; topography and, 3334; at
spruce beetle, 182, 195, 216, 219

spruce-fir forests, 19396, 194f; bark beetle outbreaks in, 195, 211;
dominant plant species in, 17677t; hydrologic budget of, 2045.
See also forests
upper treeline, 233, 342n10

terpenes, 116, 334n19
terraces: in Laramie Basin, 286f; in riparian landscapes, 46, 5152;
along the Snake River, 247f
spurge, leafy, 104
Teton Dam, 252
squirrel, red (pine), 17980, 180f, 192, 234
Teton Range: elevation of, 246, 247f, 248f; formation of, 17, 24546,
squirrels, ground: in alpine, 239f; in desert shrublands, 139; in

grasslands, 103
248f. See also Grand Teton National Park

thinning. See timber harvesting
stands (forest), 338n28; doghair, 193, 193f; even-aged, 189, 190
thistle, Canada, 104
Stansbury, Howard, 285
Thompson, C., 15
Stapp, P., 333n78
Thompson, J., 96, 332n41
state lands, area of, 6f
threetip sagebrush. See sagebrush, threetip
States, J., 163
threetip sagewort. See sagebrush, threetip
state transition hypothesis, 336n35
three-toed woodpecker, 182
steppe, 333n2. See also sagebrush steppe
Thunder Basin National Grassland, 3, 103, 107
Stohlgren, T. J., 341n16
thunderstorms. See lightning
stone nets, 237, 237f
Tiku, B. L., 335n14
stone polygons, 237, 237f, 342n34
timber harvesting: in Black Hills, 274, 278; compared to fire, 21115,
streams: in alpine tundra, 236; ephemeral, 45, 62, 65; map of, 46f;
water movement in, 45. See also riparian landscapes
Strong, W. E., 249
212f, 213f, 340n27; future of, 21516; habitat fragmentation by,

31112, 311f; in multiple use on public lands, 178, 308, 309f;
selective, 21415, 214f, 277f, 341nn5657.
subalpine forests. See spruce-fir forests
toad, Wyoming, 11f, 293, 297
subalpine fir, 8f, 19396, 194f
toadflax, bastard, 179
subalpine meadows. See mountain meadows
topography, 2730, 28f; ecological regions based on, 2830; and
sublimation, 34, 118; from forests, 2047, 205f, 211

succession, primary, in sand dunes, 143, 145, 146
succession, secondary, 94, 325n3(ch1); in aspen, 196-98; in desert
temperature inversions, 3334; and water availability, 34. See also

rainshadow; specific ecosystems and regions
torpor, 175
shrublands, 13940, 336n5; in grasslands, 94, 107; in riparian
tourism, as percentage of state economy, 325n1(ch1)
landscapes, 54, 56, 56f
Townsend, John, 126
succulents, in grasslands, 83, 86
Townsends ground squirrel, 139
suckers, 196. See also aspen forests
transpiration, 204; in forests, 204, 205f, 340n11; in grasslands, 108;
sudden aspen decline (SAD), 198, 219, 220f
in riparian landscapes, 57; in sagebrush, 116, 118, 136. See also
sulfur, in soils of wetlands, 68, 70
evapotranspiration
401
402 Index
Trappers Point, 31011, 347n10
treeline, upper, 23042; adaptations to, 233; advancing of, 24142,
water consumption: evaporation as percentage of, 62; by plants in

riparian landscapes, 57
343n44; avalanches in, 23334, 233f; average temperature at,
water development: cloud seeding, 308; reservoirs, 3078, 308f
233, 342n10; climate change at, 24142; elevation of, 23031,
water-holding capacity of soils, 35
342n1, 342n3; krummholz at, 23133, 232f; soils of, 232; white
water movement, in riparian vs. nonriparian wetlands, 45
pine blister at, 234, 234f. See also alpine tundra
water rights, endangered species and, 293, 298, 347n28
tree-ring studies, 23, 24f, 32
water sedge, 70, 77f, 78
Triassic Period, 12f, 13
Watts, G., 100101
tribal lands, location of, 6f
Waugh, W. J., 162
trona deposits, 15
weather vs. climate, 38
trout, cutthroat, 26364, 344n85
Weaver, John E., vi, 8788, 96
trout, introduced species of, 329n42
Weaver, T., 338n4
trout, lake, 26364, 344n85
weeds. See invasive species of plants
true mountain-mahogany. See mountain-mahogany
West, N. E., 115, 335n3, 336n36
tundra. See alpine tundra
Westerling, A., 26162
Turner, M. G., 340n29
western balsam bark beetle, 182, 195
two-grooved milkvetch, 335n18, 346n22
western spruce budworm, 182, 189, 189f, 339n65
Typic Humicryept soils, 341n7
western wheatgrass, 332n48

West Nile virus, 12930
Uinta Mountains: foothills of, 162; formation of, 15; upper treeline
in, 241
Uncompahgre Plateau, 220f
wetland indicators, 6768, 330n5

Wetland Reserve Program, 80
wetlands, 6680; adaptations to, 6870; animals species in, 68, 70;
Uncompahgre Plateau Partnership, 309
area covered by, 45, 66; biological diversity of, 67; climate change
Union Pacific Railroad, 25, 285, 305
in, 7980; conservation of, 80; construction in, 7677, 80;
uplift, tectonic, 13, 14, 17
dominant plant species in, 6770, 69f, 7273t, 330n5; ecological
Upper Green River Lake, 76f
role of, 45, 66, 329n1; future challenges in, 7980; hydrophytes
Upper Platte River Basin, 299, 300f
used to define, 45; invasive plants in, 79; irrigation water and,
upper treeline. See treeline, upper
67, 74, 76, 80, 346n17; as landscape sinks, 79; of the Laramie
Uresk, D. W., 345n27
Basin, 80, 29697, 346n17; legislation on management of, 328n2;
urine deposition in grasslands, 92, 95, 344n53
loss of, 66, 79, 330n40; seasonal dry periods of, 67, 68, 70; soils
Urness, P., 332n41
of, 6870; types of, 45, 66; water movement in nonriparian vs.
Utah juniper, 22, 23f, 148, 157, 158t, 160, 161f, 162
riparian, 45; in Yellowstone National Park, 250. See also riparian
Utah serviceberry, 16465
landscapes; specific types of wetlands

wet meadows, 7577, 75f, 76f; in alpine tundra, 236, 23839t, 239;
Vale, Thomas, 327n49

Van Vuren, D., 332n41
construction in, 7677; dominant plant species in, 7273t, 7577,

23839t; invasive plants in, 79; soils of, 6870
Veblen, Thomas, 170
wheatgrass, 331n8
vegetation, maps of: in Wyoming, 8f; in Laramie Basin, 290f. See also
wheatgrass, bluebunch, 83, 156, 159t, 164, 165f, 166, 176t, 221, 250,
specific types of vegetation

Vrendrye brothers, 266, 345n2
volcanoes: geologic history of, 13, 15, 17, 20, 245, 325n3(ch2); in
Yellowstone National Park, 20, 245, 326n25
Vore Buffalo Jump, 24
251f
wheatgrass, western, 49t, 83, 84f, 88, 94, 95, 100, 109f, 110, 120,
12123, 13435, 136t, 14748, 155f, 156, 268f, 273, 289, 291f,
294f, 331n8, 332n48, 347n30
whirling disease, 263
whitebark pine: bark beetle outbreaks and, 26263; limber pine com-
Walker Creek, 54f
pared to, 342n3; seed dispersal by birds, 172; at upper treeline,
Wamsutter, Lake, 19, 326n24
231, 233, 234; white pine blister rust and, 234, 234f, 26263; in
warbler, MacGillivrays, 344n45
Yellowstone National Park, 250, 250f
warbler, yellow, 344n50
whitebark pine woodlands, 17677t
warm-season plants, 8889
white fir, 339n81
Wasatch Formation, 326n13
white horehound, 27879, 279f
Washburn, Henry D., 249
white pine blister rust, 17879, 234, 234f, 26263, 342nn1516,
Washburn, Mount, 252f

water allocation: on Colorado River, 327n9; and endangered species,
29798
water availability: climate change and, 4041; contributing factors
344n78
white spruce, 272, 272f, 339n81
white-tailed jackrabbit, 139f
white-tailed prairie dog, 102
in, 30; conveyance losses in, 57; in desert shrublands, 131, 132,
Whitlock, C., 20, 248
132f, 13739; in forests, 2027, 211; in grasslands, 89, 90f; in
Whittlesey, L., 257
mountain meadows, 228; in sagebrush steppe, 11011, 11718;
Wiens, J. A., 333n93
topography in, 34; wind in, 34
Wigand, P. E., 334n32
Index
Wight, J. R., 337n10
Wyeth, Nathaniel, 106, 126
Wild and Scenic Rivers Act, 307, 329n37
Wyoming: area of, 4f; economy of, 31415, 325n1(ch1); ecosystem
wilderness: area of, 3; location of designated, 3, 6f
distribution in, 8f; elevation of, 3, 4f; human population of, 305;
wild horses. See horses, feral
public, private, tribal, and state lands in, 3, 6f; roads and cities
wildlife: in forests, 198, 216; refuges, in Laramie Basin, 29293, 297;

in Yellowstone, 245, 249, 252, 25658. See also specific species
wildrye, Great Basin, 164
wildrye, yellow, 143
of, 7f
Wyoming big sagebrush, 109f, 110f, 111; distribution of, 111, 112; in
foothills, 164; in grasslands, 83; recovery after disturbances in,
120. See also sagebrush, big
Williams, J. W., 41
Wyoming Game and Fish Department, 347n4
willow(s): in alpine tundra, 241; in Greater Yellowstone Ecosystem,
Wyoming Natural Diversity Database, 325n5(ch1), 330n5
25657, 258, 26465; in riparian landscapes, 4546, 50f, 56; in
Wyoming Permanent Mineral Trust Fund, 315
wetlands, 68, 70, 236, 23839t
Wyoming Range, 221f
willow, Bebb, 50f
Wyoming Range Legacy Act, 306
willow, Geyers, 50f
Wyoming toad, 11f, 293, 297
wind: in alpine tundra and upper treeline, 231, 23233, 237, 237f;
Wyoming Wetlands Conservation Strategy, 80
for electrical power, 313f; in foothills, 164; in forest fires, 218;
Wyoming Wildlife and Natural Resources Trust, 306
katabatic, 287, 346n15; in Laramie Basin, 287, 289; in mountain

meadows, 223, 22526; as periodic disturbance, 37; in sagebrush
Xanthoparmelia chlorochroa, 92, 93f
steppe, 112; in sand dune formation, 3435; in water availability,
xerophytes, 8586
34
Wind Cave, 270
yellow cedar, 343n53
Wind Cave National Park: bison in, 97, 97f, 268; black-footed ferrets
yellowhead, desert, 10f
in, 279, 346n48; climate change and other management issues
yellow-headed blackbird, 67f
in, 27481; elk management in, 280; establishment of, 268;
Yellowstone fires of 1988, 25962; aspen forests after, 219, 261, 262f;
fires in, 346n41; intact grasslands in, 97, 107; invasive plants in,
bark beetle outbreaks and, 217, 341n69; and elk, 258; end of, 259;
27879, 279f; prairie dogs in, 97, 97f, 27879, 279f, 280f
extent and severity of, 25960, 260f, 338n28; lessons learned
wind energy projects, and habitat fragmentation, 312, 313f
from, 25961; in lodgepole pine forests, 191f, 192, 209, 210f,
Wind River, 53
211f; net primary productivity after, 209, 210f, 211f, 340n29;
Wind River Basin, 103
nutrient cycling after, 210, 340n36. See also Yellowstone National
Wind River Canyon, 17

Wind River Mountains: bark beetle outbreaks in, 183f; formation of,
15; Gannett Peak, 18f; glaciers in, 20; location, 5f; meadows in,
228
Park
Yellowstone Lake, 248, 344n85
Yellowstone National Park (YNP), 24951; bark beetle outbreaks in,
217, 341n69; dominant plant species in, 25051; early explorers
Wind River Reservation, 329n37
in, 249; ecosystems of, 25051; elk in, 25658, 264; establish-
Wing, Scott, 14
ment of, 3, 249; fens in, 68, 77f; fire history in, 209f, 25859, 261f
winterfat, 132f, 133, 134, 137
(see also Yellowstone fires of 1988); fire management in, 25862;
winter feeding programs, 171, 257
foothills of, 250, 251f; forests in, 189f, 192, 193, 250; fossil trees
wintergreen plants, 174
in, 14, 15, 245, 247f; geologic history of, 245; grasslands in, 250,
winter rangelands: in foothills, 17071; in Greater Yellowstone
258, 344n53; lakes and rivers of, 249; livestock in, 342n20(ch13);
Ecosystem, 251f, 25758
management issues in, 25665, 343n41; mosses vs. vascular
Wisconsin (Pinedale) glacial period, 326n23
plants in, 68; mountain meadows in, 228, 250, 252f; precipita-
wolves, 26465; distribution of, 3, 325n2(ch1); and elk popula-
tion in, average annual, 30; sagebrush in, 114f, 250; temperature
tions, 170, 264, 344n99; in Greater Yellowstone Ecosystem,

256, 26465, 344n93, 344n95; reintroduction of, 170, 245, 258,
26465, 344n93
changes in, average, 40; topography of, 20, 246f, 249; volcanoes
in, 20, 245, 326n25. See also Greater Yellowstone Ecosystem
Yellowstone Plateau, 21920
wood decomposition, 2078, 212f
Yellowstone River, 248, 249, 307
Woodhouse, C. A., 327n9
Yellowstone River valley, 250, 251f
woodpecker, black-backed, 182
yellow warbler, 344n50
woodpecker, three-toed, 182
yellow wildrye, 143
woodrat, bushy-tailed, 22, 22f, 23f
YNP. See Yellowstone National Park
woody draws, 15859t, 166f, 167, 272
Young, J. A., 336n36
Wright, G., 256, 343n38
Young, M. K., 329n40
403
About the Authors
Dennis H. Knight. Professor emeritus at the Univer-
William A. Reiners. Professor emeritus at the Uni-
sity of Wyoming, where he taught ecology and forest
versity of Wyoming, he taught at the University of
management for 35 years. Author of the first edition,
Minnesota and Dartmouth College before moving to
his research focuses on the effects of natural and
Laramie in 1983. His special interests include biogeo-
human-caused disturbances on forest and rangeland
chemistry, ecosystem analysis, and the effects of cli-
ecosystems.
mate change.
George P. Jones. Ecologist and associate director of the
William H. Romme. Professor emeritus, Colorado
Wyoming Natural Diversity Database at the University
State University, he taught at Fort Lewis College for
of Wyoming. His work focuses on the analysis of plant
many years before moving to Fort Collins. Most of his
communities and the provision of ecological informa-
research has been on the effects of fire and other dis-
tion to industry, consulting firms, and local, state, and
turbances on forests, especially in the Greater Yellow-
federal agencies.
stone Ecosystem.
404

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Mountains

New Haven and London

Published with assistance from the University of Wyoming

Library of Congress Cataloging-in-Publication Data

Biodiversity Institute (wyomingbiodiversity.org).

Published with assistance from the Mary Cady Tew Memorial

Mountains and plains: the ecology of Wyoming landscapes/

First edition 1994. Second edition 2014. Copyright 1994, 2014

by Yale University. All rights reserved. This book may not be

Includes bibliographical references and index.

reproduced, in whole or in part, including illustrations, in any

ISBN 978-0-300-18592-8 (pbk. with flaps : alk. paper)

form (beyond that copying permitted by Sections 107 and 108

1.EcologyWyoming. 2.Landscape ecologyWyoming.

press), without written permission from the publishers.

Yale University Press books may be purchased in quantity for

A catalogue record for this book is available from the British

To our students and colleagues,

Nature is an open book. . . . Each grass-covered hillside

Part One. Wyoming and

Part Four. Foothills and

the Rocky Mountain West

10. Escarpments and the

Foothill Transition, 155

Part Two. Wetlands

14. Upper Treeline and

Part Five. Landscapes of

Part Three. Plains and

9. Sand Dunes, Badlands,

Part Six. Sustainable Land

This page intentionally left blank

Many changes have occurred in the Rocky Mountain

lands, forests, subalpine meadows, and alpine tundra.

region since 1994, when the first edition of this book

The ecology and management of croplands are beyond

was published. Wildlife habitat has been fragmented

the scope of this book, but we discuss the ecological

at alarming rates, the once abundant sage-grouse has

effects of irrigation, invasive species, livestock, feral

been proposed for protection by the Endangered Species

horses, elk, prairie dogs, and wolves. The book ends

Act, invasive plants have become more common,

with a chapter on current issues pertaining to land

wolves have been reintroduced, climate change is now

management and conservation.

well documented, epidemics of forest insects are more

Plant ecology is emphasized, because vegetation pro-

widespread, forest fires are more frequent, and new ap-

vides habitat and gives character to the landscape, but

proaches for conservation have been adopted. Our goal

we also discuss plant-animal interactions, hydrology,

for the second edition, 20 years later, is to provide a new

nutrient cycling, and the effects of disturbances, such

synthesis of the ecological research that is pertinent to

as fire, insect epidemics, and timber harvesting. Most

natural resource management, with a focus on the ecol-

chapters consider the implications of climate change.

ogy of Wyoming and adjacent parts of Colorado, Utah,

The chapters vary in length and structure, based on the

Idaho, Montana, South Dakota, and Nebraska.

information available and our perception of what most

The book has six parts. The first is an introduction

readers will find interesting. We provide guidance to

10. Escarpments and the

14. Upper Treeline and

9. Sand Dunes, Badlands,