Documente Academic
Documente Profesional
Documente Cultură
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
University of Wyoming
Laramie, Wyoming
www.uwyo.edu/biodiversity
Knight, Dennis H.
pagescm
of the U.S. Copyright Law and except by reviewers for the public
I.Title.
QH105.W8K582014
577.09787dc23 2014015382
Contents
Preface, ix
Acknowledgments, xi
Mountains
1. Introduction, 3
2. Landscape History, 12
3. P resent-Day
Environments and
Climate Change, 27
5. Marshes, Playa
Wetlands, Wet
Meadows, and Fens, 66
Yellowstone
Ecosystem, 245
16. The Black Hills, Bear
Lodge Mountains, and
Devils Tower, 266
17. The Laramie Basin, 282
Epilogue, 319
Appendix A: Conversion
Table, 321
Appendix B:
Characteristic Soil
Types, 322
Notes, 325
References, 349
Index, 387
Preface
terms, but those few that remain are defined where they
in appendix A.
ix
Preface
sity of Wyoming.
Acknowledgments
Gilli
gan, Bonnie Heidel, and Joanna Lemly, and the
and Mary Pasti, along with Peter Strupp and Cyd West-
xi
xii
Acknowledgments
Wyoming Depart
ment of Agriculture, the Wyoming
Manage
ment, the University of WyomingNational
Chapter 1
Introduction
tracts of land that have never been plowed and are still
bison may have done the same. Grizzly bears, elk, and
to the extent they once did, but all are found in some
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
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
tion available.
Introduction
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
10
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
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
are summed for the time, labor, and energy required for
services.
11
Chapter 2
Landscape History
ERA
PERIOD
EPOCH
Holocene
Quaternary
2.6
Pliocene
CENOZOIC
(Age of Mammals)
Miocene
23
Oligocene
34
Paleocene
12
PALEOZOIC
(Age of Fishes)
Jurassic
Triassic
Pennsylvanian
Mississippian
Devonian
Silurian
Ordovician
Cambrian
PRECAMBRIAN
Cretaceous
Permian
Eocene
56
Neogene
Paleogene
0.01
Pleistocene
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
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
years.
13
14
continued uplift and erosion from the adjacent mountains during the Laramide orogeny, the accumulating
peat was overtopped with deep sediments here and there
est deposits. Some of the seams are more than 200 feet
Landscape History
chapter 15).
15
PETM but were less common until after the heat sub-
the biosphere.
evolution.
hydrologic cycle.20
15
Basin
Granite and
Related Rocks
Landscape History
With uplifting, the climate cooled, and some rivers began to flow more rapidly, increasing the rate of
the western slopes as the rising air cools and the mois-
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
Landscape History
mountain glaciers.
23
northern Montana.
19
20
thousands of years.
chapter 3).27
tory of this area over the past 30,000 years, during and
cypress, have not been found in the fossil record for that
time.
years ago.
29
Whitlocks
research
indicates
that
Engelmann
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
21
22
ago.40 Pinyon pine did not arrive in the state until about
the Green River flows into Utah.41 This small tree also
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
Landscape History
Big Pryor
East Pryor
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
2000
anthropological data, and old journals and photographs. What do such records reveal about the past few
3000
1000
centuries?
4000
5000
8001300
6000
ad
7000
8000
9000
10000
80
80
ad,
the area.49 We can infer that they used small trees for
23
Proportion of western
U.S. in drought (%)
24
0
10
20
30
Wetter
40
50
60
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).
industrialists.52
53
576,000 people. 57
Landscape History
the way for new towns and for mining gold, silver, and
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.
have occurred.
25
26
of 1870.
Photographs must be used cautiously in drawing conclusions, as they only provide snapshots in time of a
small area.65
the east.
Present-Day
Environments and Climate
Change
Chapter 3
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
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
0
0
50
50
100 km
100 miles
To the west of the Great Plains is a region of mountains and intermountain basins (see figs. 1.2 and 3.1).
the north and south. For this reason, the Pony Express,
feet, with the highest basins in the south (see figs. 1.1
and 3.1).
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)
29
30
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.
book.
timing of precipitation.
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
common
ly accumulates during winter. Some snow
31
32
wet
Wet
10361 ft
48 in
1
0
1
2
3
Dry
dry
900
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
13 in
C
0
10 in
mm
16 in
MONTH
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
Soil Characteristics
11
erals are derived. Some soils are shallow and not well
chapter 8).
growth.
soil depth or soil texture (see figs. 17.9, 17.11, and 17.17),
chapter 7).
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
100
70
36
90
80
90
sandy loam
70
60
silt
50
40
30
20
10
100
% sand
Periodic Disturbances
cannot survive.
and shrubs.
37
38
point out that unusually cold days will occur during the
has been that such changes are so slow that they hardly
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
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
comparable data. 24
glacial melting about 10,000 years ago, and also the cool
structed for the past >6000 years and are now similar
trend that has been under way for 30 years.32 More hay
41
Chapter 4
Riparian Landscapes
during the year for food, water, and shelter. Animals also
year. In general, all wetlands have soils that are wet long
floods. Also, old trees topple into the water along erod-
45
46
Wetlands
1
2
3
4
5
6
Great Basin
16 Bear
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
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
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)
Common
name
Latin name
AlderTall
Short
conifer
willow
willow
woodland shrubland shrubland
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
woodland shrubland shrubland
Cinquefoil
silver
Greasesagebrush
wood
shrubland shrubland
Coniferbroadleaf
woodland
Riparian
meadow
SEDGES
FORBS
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
High
CHANNEL STABILITY
meandering or incised.
Cut
bank
Floodplain
Bar
Point bar
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
Braided
Meandering
CHANNEL TYPE
Straight
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. 4.7. 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.
the stream for 300 or 400 yards are covered with a thick
that happens.
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.
(fig.4.11; see also fig. 4.8). Fertile soils and water avail-
silver
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
55
Wetlands
Bar deposit
Sandbar meadow
Without
willow or
cottonwood
Cottonwood woodlands
Young (1-29 yrs)
Meadow
Basin big
sagebrush
Suspended sediment
Fire
or
beaver
Fire
or
beaver
Saltcedar
willow
Annual
flooding
Saltcedar
Skunkbush sumac
Rabbitbrush
shrubland
Periodic
flooding
or fire
Bank erosion
56
Periodic
flooding
or fire
Skunkbush sumac
Rabbitbrush
Snowberry
Rose shrubland
Prolonged inundation
?
?
Greasewood
shrubland
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).
Riparian Landscapes
lar woodlands.
ecosystem services.
57
Riparian Landscapes
Engelmann spruce, subalpine fir, and thinleaf alder. Elevation 8,500 feet.
can build new dams. Once the tires are in place, the
available shrubs.25
59
60
Wetlands
of the country.28
to the same place for a year or more. Also, the upland forage then, away from the rivers, might have been attrac-
Livestock
mules are very fond of the bark which we strip from the
Riparian Landscapes
fed streams flow into all major drainages (see fig. 4.1).
61
62
Wetlands
Riparian Landscapes
ther. Some rivers were clogged with logs. After gold was
39
63
64
Wetlands
to the region in the 1930s (fig. 4.17; see also fig. 4.11).45
44
Riparian Landscapes
stands with more square feet of leaf area per square foot
life habitat.53
invasive
growth is desired.
plantssuch
as
saltcedarprobably
will
land managers.
65
Marshes, Playa
Wetlands, Wet Meadows,
and Fens
Chapter 5
(figs.5.15.3).
66
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
has more species (275) in the state than the sedge fam-
other way.
8species of shrews.
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
70
Wetlands
Marshes
the water.
slope quickly into deep water, and wave and ice action
that occupy the wetland during the dry period but are
ent group of plants (figs. 5.6 and 5.7; see also fig. 5.5).
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
Alopecurus aequalis
Tufted hairgrass
Deschampsia caespitosa
LM
LM
Triglochin palustris
Seaside arrowgrass
Triglochin maritima
Threesquare bulrush
Schoenoplectus pungens
Hardstem bulrush
Schoenoplectus acutus
Cottongrass
Eriophorum spp.
LM
Juncus arcticus
LM
LM
Analogue sedge
Carex simulata
LM
Blister sedge
Carex vesicaria
LM
Buxbaums sedge
Carex buxbaumii
LM
Carex praegracilis
LM
Mountain sedge
Carex scopulorum
Mud sedge
Carex limosa
Nebraska sedge
Carex nebrascensis
LM
LM
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
Purple marshlocks
Comarum palustre
LM
Common name
Latin name
Marsh
Wet meadow
Fen
Playa wetland
Resin birch
Betula glandulosa
Shrubby cinquefoil
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
Symphyotrichum foliaceum
American globeflower
Trollius laxus
Arumleaf arrowhead
Sagittaria cuneata
Balsam groundsel
Packera paupercula
Broadleaf cattail
Typha latifolia
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
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
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
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.
Wet Meadows
76
Wetlands
survive.24
pugging.27
ness, from very wet sites that merge into marshes and
hydrology.
Fens
as well.
The differences between bogs and fens are caused
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.
running water.38
39
promise.
45
80
Wetlands
50
waters.
52
Intermountain Basins
Chapter 6
Grasslands
basins (see fig. 1.5). Some have been plowed for crop
high for a longer period of time each year, the soils have
appendix B).
plant diversity. 3
83
84
Grasslands
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).
favorable.
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.
87
88
Table 6.1. Some characteristic mammals, birds, and reptiles found in grasslands, shrublands, and escarpments at
low elevations
MAMMALS
Badger
Bobcat
Jackrabbit, black-tailedb
Chipmunk, least
Jackrabbit, white-tailedc
Cottontail, desert
Mouse, deer
Cottontail, Nuttalls
Coyote
Mouse, northern
grasshopper
Deer, mule
Fox, red
Fox, swift
Ferret, black-footeda
Pronghorn
Ground squirrel,
thirteen-lined
Woodrat, bushy-tailed
b
c
BIRDS
Bluebird, mountain
Meadowlark, western
Bunting, lark
Nighthawk, common
Eagle, golden
Owl, burrowing
Falcon, prairie
Sparrow, Brewers
Grouse, sage
Sparrow, lark
Harrier, northern
Sparrow, sage
Hawk, red-tailed
Sparrow, vesper
Jay, pinyon
Thrasher, sage
Kestrel, American
Towhee, green-tailed
Lark, horned
Vulture, turkey
Longspur, McCowns
REPTILES
Bullsnake
Rattlesnake, prairie
Lizard, spiny
Now rare.
neighbor
ing plants tap different parts of the soil for
10
Grasslands
from the plants that grew earlier and were eaten.13 With
drier.14
Elk consume more grass, and deer more twigs and leaves
Energy Flow
16
89
500
90
400
300
200
100
100
200
300
400
500
600
EVAPOTRANSPIRATION (mm/yr)
700
800
the plants use about 3040 percent for their own main-
than 1 percent.
or more.
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
Gross primary production (GPP), or total photosynthesis (g/m 2/yr), varies greatly from year to year, depending on water avail-
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
Aboveground carnivores include the coyote, fox, birds of prey, and snakes.
h
i
92
true that the only way humans derive food from semi-
Nutrient loss through leaching is an unlikely phenomenon in most western grasslands, because, unlike
Because
nutrients
accumulate,
grassland
soils
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
93
94
ity that the atom will again enter new herbage growth
sirable consequences.
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
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
95
96
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.
97
50
50
Lightning
NUMBER OF FIRES
98
30
20
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.
Fire
Prior to the advent of fire suppression, prairie fires
common. 55
53
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.
soil surface.58
Another effect of burning, especially in taller grasslands, is to remove the insulating mulch, allowing the
99
that remain.60
Drought
The continental climate of the western Great Plains
considerably.
Grasshoppers
drought-tolerant species.
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.
69
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
Grasslands
77
highly controversial.
the en
dangered black-footed ferret. Appropriately, the
croplands.
Invasive Plants
103
tor may be that western grasslands are too dry for most
Grasslands
82
first few years as they are later, after they have adapted
species can.
105
Careful manipulation
87
does not exist. Most of the native plants are there, and
some of the small native animals can be found if one
Grasslands
otherwise.
107
95
spheric deposition.
moderate warming.
96
Chapter 7
Sagebrush
109
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.
bly, big sagebrush is absent from very wet areas (fig. 7.3).
111
ELEVATION (feet)
10000
Mtn silver
sagebrush
Alpine sagewort
Mtn big
sagebrush
9000
Threetip
sagebrush
Wyoming big
sagebrush
8000
Black
sagebrush
or
Bud
sagewort
or
Birdfoot
sagewort
7000
Basin
big
6000 sagebrush
or
Silver
sagebrush
5000
Ravine
North
or
or
floodeast
plain
slope
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-
the region (figs. 7.5 and 7.6). Unlike most woody species
sagewort.10
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
40
Sagebrush
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.
shrubs canopy.
11
roots. If the deep roots are in moist soil and the shal-
115
season is longer.
Sagebrush
burning or herbicides.
117
deeper into the snowpack or possibly into the soil (if its
common occurrence.
haps by 15 percent,
29
30
runoff as streamflow.
Sagebrush
restoration.
well.44
Still, the benefits of improving habitat or forage availability for some species by burning must be balanced
highly valued.
Nutrient Availability
along with the potential for hydraulic lift and the tap-
119
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
Fire
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
ex
ample, horsebrush, rabbitbrush, and winterfat in
However, University of
60
121
root crowns, the depth of the latent buds below the soil
as sage-grouse.67
Cheatgrass
Sagebrush
300
Grasses
PERCENTAGE
200
Thickspike
wheatgrass and
Plains reedgrass
Bluebunch
wheatgrass
Bluegrass
100
300
Needle-and-thread
Shrubs
Horsebrush
Rabbitbrush
200
100
Sagebrush
1936
1942
1948
1954
YEAR
1960
1966
pepper
weed, Canada thistle, Dyers woad, halogeton,
tian toadflax.70
123
Fire suppression
Fire
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
accelerate.
management).77
Sagebrush
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
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
125
(former
ly known as blue-green algae), lichens, algae,
of these animals.87
Sagebrush
Fig. 7.16.The population of feral horses
commonly exceeds the carrying capacity of the rangelands they occupy. Photo
by Ken Driese.
continue.89
of purposes.90
127
91
ecosystem.93
chapter18).94
Sagebrush
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.
tors.
sagebrush.100
98
Un
fortunately, as discussed previously, fire manage-
129
101
Hundreds
times subtle and not obvious from the road, but aerial
Desert Shrublands
and Playas
Chapter 8
be saline (see figs. 1.2, 1.5, 3.4, and 8.1). The predomi-
in wetter zones.
131
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
Greasewood
Low
Pickleweed
Low
SALINITY
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).
a form of allelopathy.
the water and nutrients they need (see chapter 6). One
the two species help explain how they coexist. Other des-
133
11
the salts. The same might be true for many desert plants.
otherwise be expected.
In some areas the low shrubs are rare and the com-
18
135
Common name
Latin name
Saltbush
shrubland
Saltgrass
meadow
Desert
grassland
SHRUBS
Basin big sagebrush
ravines
ravines
Birdfoot sagebrush
Artemisia pedatifida
Bud sagebrush
Picrothamnus desertorum
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
Saltgrass
Distichlis spicata
Sandberg bluegrass
Poa secunda
Goosefoot
Chenopodium spp.
Green molly
Bassia americana
Hookers sandwort
Arenaria hookeri
Nuttalls povertyweed
Monolepis nuttalliana
Artemisia frigida
GRASSES
FORBS
Saltbush
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
30
25
20
15
Grassland
10
5
0
Desert shrubland
0 10
30 45 60
90
120
MINUTES
180
137
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
microbial activity?
139
arid environment.
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
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
Compared with surrounding grasslands and shrublands, some parts of the dune environment are more
the plant tissues grow and die, soil organic matter gradu-
lower elevations.
17
Badlands
for plant growth than the flats do. At one time, a slurry
conducted an experiment:
Mud Springs
sutter). These conical mounds are 315 feet high and are
the explorers.
Mima Mounds
than 2 feet high (figs. 9.9 and 9.10). They are common on
Washington.
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.
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
face. They also observed that the mound soil was lens-
like mounds.
Mountains
Escarpments and
the Foothill Transition
Chapter 10
(fig. 10.1).1
155
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
quently browsed by deer, especially in the winter. Notably, the shrubs respond by growing lateral branches.
Mountain-Mahogany Shrublands
Juniper Woodlands
157
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)
Wyoming big
sagebrush
Artemisia tridentata
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
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
Pseudoroegneria spicata
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
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
11
161
13
is important.
have been favorable for these tree species just as for juni-
17
24
than in Wyoming. 26
163
and Woods rose (fig. 10.11; see table 10.1). Great Basin
165
(see fig. 8.9), which keeps the stems and leaves densely
are different.38
Deciduous Woodlands
Four other vegetation types in the foothill-escarpment
mosaic are woodlands characterized by different combinations of aspen, chokecherry, Gambel oak, and bur
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.
167
lead to an ex
pansion of oak woodlands if relatively
the roots are not usually killed and they sprout vigor-
vide grasses and forbs for big game during the winter,
169
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-
bedding.
explain why the groves are failing over large areas. How-
tem.48 More than one year of rest may result if the herd
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.
171
part of the new forest that develops. Its well known that
pine.51
Chapter 11
Mountain Forests
early July.1
systems and the ways that they affect the lowlands (for
mountains.
which allows relatively high rates of evapotranspiration.3 Still, plant water stress is less frequent in midelevation forests than in any other upland ecosystem.
173
their growing season into late fall and begin the next
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
Aspen
9000
Lodgepole pine
Willow
8000
Alder
Birch
2500
Aspen
Douglas-fir
or
Ponderosa pine
Foothill
shrublands
7000
Cottonwood
ELEVATION (m)
ELEVATION (feet)
3500
Alpine
6000
12000
9000
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
Mountain Forests
amphibians.14
years.
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 ponderosa
Pinus albicaulis
Abies lasiocarpa
Antelope bitterbrush
Purshia tridentata
Shinyleaf spiraea
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
Pseudoroegneria spicata
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
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
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.
objective.
all trees are cut, whether they are infected or not. This
Comandra
blister
rust
(Cronartium
comandrae),
infection.
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
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
REPTILES
Boa, rubber
Gartersnake, wanderingb
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
179
and fire.23
highis characterized by short, wind-pruned Engelmann spruce, subalpine fir, and limber pine in most of
Wyoming ranges, though whitebark pine is typical of
Mountain Forests
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
181
Mountain Forests
from the bark and fly toward other trees. They are most
183
inch long, about the size of a grain of rice. The first few
Curi-
36
The
40
38
stain fungus.41
Mountain Forests
trees often grow much faster than they did before the
44
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
tree seedlings.47
185
this migration.54
Mountain Forests
southern part.58
187
Douglas-fir Forest
Mountain Forests
Insects can also cause variation in forests of Douglas-fir, especially the Douglas-fir beetle and the western
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
limestone-derived soils.68
Wyoming.
67
Mountain Forests
closed and on the tree for many years until higher than
the forest floor if they are exposed to the sun for a long
69
pole pine trees have thin bark and are easily killed by
191
intervals are less than 200 years, canopy gaps are less
adaptation.
most trees were serotinous, and the fire scorched but did
Mountain Forests
Spruce-Fir Forest
of warmer temperatures.80
193
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
fir zone, however, lodgepole pine and aspen are not pres-
dant, but the trees are smaller and younger, rarely more
than 250 years old. Fir seedlings and saplings may also
turbance.
86
(which is not a host for the spruce beetle) and the small
195
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,
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).
96
seem that the small trees could tap the energy stored
197
ungulate numbers.
are found there all year long, such as the blue grouse,
age and better hiding cover. Elk, mule deer, and cou-
Some resource
food for birds that are agile enough to catch them. Even
102
forest.
Mountain Forests
199
Chapter 12
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:
200
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
0.9
Fixation
0.01
Soil
organic
matter
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).
Live trees
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
-1
-2
40
80
120
160
200
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
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).
scape mosaic.
STREAMFLOW (cfs)
8
7
6
5
4
3
2
1
1956-1971
1940-1955
APR
MAY
JUNE
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).
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
est, are the total amount of leaf area and leaf type (ever-
and the other a more open stand with about 800 trees
per acre. Both originated after the same fire in the late
the leaf area is low and the leaves are deciduous, such as
both stands had the same leaf area and about the same
transpiration rates.14
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
and fir use water at higher rates per unit of leaf area
are common.
is the amount of water that the soil can store, that is,
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
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).
tion is enhanced.
downstream.
occur.
tion are all subject to rapid change (see chapter 11), and
% ORIGINAL REMAINING
180
140
Nitrogen
100
Dry mass
60
20
0
12
24
36
48
60
TIME (months)
72
84
96
24
of disturbance.
26
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
ops following the fire, and whether the burn was patchy
longer time.
7
6
12
5
4
3
2
1
0
10
10
10
10
10
Pine saplings/hectare
16
10
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
With regard to insects, the extensive bark beetle outbreaks now sweeping across the West have important
reduce total tree growth for several years after the peak
often are not killed, and, with fewer large trees compet-
ing for water, nutrients, and light, they grow faster than
34
Total NEP
g C per m2 per yr
100
High density
50
0
-50
Low density
-100
0
50
100
150
200
250
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).
chapter 15).
42
ents from the soil, at least during the first few years after
43
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):
Fire
Coarse wood created
by disturbance
Pre-disturbance
Coarse wood
Initial Clearcut
Pre-disturbance
Coarse wood
Subsequent clearcuts
long after the fire.50 Also, over the long term, the addi-
and leaves over the past 10,000 years or more (fig. 12.13).
wood in the future as these trees die and fall. The dead
advisable.
immobilization.57
posed, given what has been learned thus far about forest
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
green stage early in the attack; the red stage, when the
the dead needles have fallen but most of the dead tree
boles are still standing; and the old stage, when the dead
and ladder fuels, which are small living trees that can
carry fire from the surface fuels into the canopy. Two
stage, but the dry, dead needles at this stage increase the
ate weather conditions, that is, when not too wet and
not too dry. When wet, nothing will burn and when
chapter 18.
flammable materials.
75
ous fuels and create places where fire spread and inten-
few sprouts.84
the dead trees have fallen and the surviving small trees
that exist throughout the current range, thereby creating a bioclimatic envelope. They then use climate models
Mountain Meadows
and Snowglades
Chapter 13
221
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
Sandberg bluegrass
Poa secunda
Timberline bluegrass
Alpine timothy
Phleum alpinum
Idaho fescue
Festuca idahoensis
Festuca saximontana
Oatgrass
Danthonia spp.
Prairie junegrass
Koeleria macrantha
Slender wheatgrass
Elymus trachycaulus
Thickspike wheatgrass
Elymus lanceolatus
Spike trisetum
Trisetum spicatum
Tufted hairgrass
Deschampsia caespitosa
SEDGES
Curly (rock) sedge
Carex rupestris
Dunhead sedge
Carex phaeocephala
Ebony sedge
Carex ebenea
Hoods sedge
Carex hoodii
Needleleaf sedge
Carex duriuscula
Carex utriculata
Obtuse sedge
Carex obtusata
Sheep sedge
Carex illota
Polygonum bistortoides
Alpine sagebrush
Artemisia scopulorum
Artemisia frigida
Arrowleaf balsamroot
Balsamorrhiza sagittata
Ballhead sandwort
Arenaria congesta
FORBS
Alpine bistort
Creeping sibbaldia
Sibbaldia procumbens
Elephanthead lousewort
Pedicularis groenlandica
Cerastium arvense
Fireweed
Chamerion angustifolium
Indian paintbrush
Castilleja spp.
Mules ears
Wyethia amplexicaulis
Pale agoseris
Agoseris glauca
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
Mertensia ciliata
Tufted fleabane
Erigeron caespitosus
Western yarrow
White locoweed
Oxytropis sericea
Caltha leptosepala
SHRUBS
Mountain big sagebrush
Silver sagebrush (mountain)
Shrubby cinquefoil
found where trees are excluded because the soils are not
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.
glades. The soil is covered, cold, and wet for too long and
disappear.
225
25
` 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
10
20
Distance (feet)
30
ting in the late 1800s and early 1900s. The best example
ponent of the summer range for elk and deer, and for a
227
roots.
cover.22
19
the winter.
229
Upper Treeline
and Alpine Tundra
Chapter 14
flowers. During the last Ice Age, the treeline was lower
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.
Elevation 10,825 feet.
230
and west slopes tend to have more snow later into the
snow cover, providing protection from abrasion and desiccation during winter. As important, they found that
Krummholz
alpine sunlight during the day but also from the cold
rates that are sufficient for plant growth and the for-
231
seedlings.
and meadows.
Avalanches
snow slides are common (fig. 14.4). Such areas are par-
233
threatened species.
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.
235
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-
SNOW COVER
Snowfree
Winter snow
cover
TOPOGRAPHIC
POSITION
Fellfield
Summit
or
ridge
Persistent
snow
Leeward
Upper
slope
source of water for most of the growing season. Consequently, the vegetation there has higher cover and
Turf
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).
otherwise.
237
Common name
Latin name
Fellfield
Alpine turf
Wet
meadow
Willow
thicket
FORBS
Arctic alpine forget-me-not
Eritrichium nanum
Symphyotrichum foliaceum
Oxyria digyna
Alpine sagebrush
Artemisia scopulorum
American bistort
Polygonum bistortoides
Creeping sibbaldia
Sibbaldia procumbens
Dwarf clover
Trifolium nanum
Elephanthead lousewort
Pedicularis groenlandica
Tetraneuris grandiflora
Lesser spikemoss
Selaginella densa
Moss campion
Silene acaulis
Parrys clover
Trifolium parryi
Parrys lousewort
Pedicularis parryi
Redtop stonecrop
Rhodiola rhodantha
Paronychia pulvinata
Antennaria media
Ross avens
Geum rossii
Sticky polemonium
Polemonium viscosum
Tufted phlox
Phlox pulvinata
Varileaf cinquefoil
Potentilla diversifolia
Caltha leptosepala
Western yarrow
GRASSES
Alpine bluegrass
Poa alpina
Timberline bluegrass
Alpine timothy
Phleum alpinum
Purple reedgrass
Calamagrostis purpurascens
Festuca saximontana
Spreading wheatgrass
Elymus scribneri
Spike trisetum
Trisetum spicatum
Tufted hairgrass
Deschampsia caespitosa
Bellardi sedge
Kobresia myosuroides
Blackroot sedge
Carex elynoides
SEDGES
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
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
of terrestrial ecosystems associated with the same pollutants in northeastern North America.42
areas that are protected from wind scour, that is, places
43
But not all alpine treelines are the same. For exam-
down during the past 10,000 years (see chapter 2).44 But,
241
of surviving.
Special Interest
The Greater
Yellowstone Ecosystem
Chapter 15
along the north side of the Madison River are part of the
245
stone that now lies about 6 miles below the valley floor.
fault line on the east side, the west slopes are gentler and
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
deposits are mostly alluvium, often referred to as glacial outwash plains, deposited as the glaciers melted. Adapted from
Love et al. (2003).
some 25,000 years ago.7 At its peak, ice covered almost all
of what is now YNP and was about 4,000 feet thick above
spruce about 11,000 to 9,500 years ago, but the less fer-
along the Snake River were formed during this time (see
stone River leaves the park at 5,314 feet. More than 150
11
14
The accolades
rivers run clear. They also offer some of the finest trout
15
249
near the site of the old Buffalo Ranch in the Lamar Val-
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
251
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
7000
6000
5000
253
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
255
the water level was back to normal the next day. Kelly
plant distribution.
vious human activities. More research was also recommended, recognizing the many uncertainties about
Native Americans.40
tors were shot; and elk and bison were managed inten-
elk had occupied the winter range inside YNP until the
ent, but they became convinced that the park area was a
257
contended that YNP did not contain the key winter hab-
50
51
Fire Management
25
20
15
1988
Fires
10
5
1970
1975
1980
1985
1995
Wolf
reintroduction
1990
YEAR
1995
2000
2005
2010
it, often days later, the fire commonly had either gone
but could not stop the spread of fires across the land-
mid-September.64
259
the result that only small areas would have burned. The
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
20
40
60
80
1900
20
40
60
80
each year, and not every year would have had accept-
1988 fires.
Moreover, no aspen
71
261
263
those carcasses.86
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
90
93
in YNP).
region as a whole.
The rivers are clean, and the thermal features are fasci-
the gray wolf, the GYE again has all of the species and
265
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
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
Geomorphic Regions
was favorable.
gia for many forest plants and animals that are sepa-
Table 16.1. Some plants in the Black Hills representative of other floristic regions
Region/Common name
Latin name
Populus angustifolia
Ponderosa pine
Pinus ponderosa
Mahonia repens
Elkweed
Frasera speciosa
Heartleaf arnica
Arnica cordifolia
Grouse whortleberry
Vaccinium scoparium
Richardsons geranium
Geranium richardsonii
Cercocarpus montanus
Big sagebrush
Artemisia tridentata
Skunkbush sumac
Rhus trilobata
Blanketflower
Gaillardia aristata
Bottlebrush squirreltail
Elymus elymoides
Prairie junegrass
Koeleria macrantha
Threadleaf sedge
Carex filifolia
Ulmus americana
Boxelder
Acer negundo
Bur oak
Quercus macrocarpa
Common hackberry
Celtis occidentalis
Green ash
Fraxinus pennsylvanica
Hophornbeam
Ostrya virginiana
Bloodroot
Sanguinaria canadensis
Viola pubescens
Red columbine
Aquilegia canadensis
Virginia creeper
Parthenocissus quinquefolia
Betula papyrifera
White spruce
Picea glauca
Bunchberry dogwood
Cornus canadensis
Canada mayflower
Maianthemum canadense
Moneses uniflora
Twinflower
Ipomoea leptophylla
Crested pricklypoppy
Argemone polyanthemos
Sand sagebrush
Artemisia filifolia
Velvetweed
Gaura mollis
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
Spearfish
60
Lead
45
55 50
40
50
Rapid
City
Newcastle
45
Hot
Springs
10
10 mi
15
15 km
48
Custer
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).
Vegetation Mosaic
osa pine forests so extensive (fig. 16.5; see fig. 1.5). They
The Black Hills, Bear Lodge Mountains, and Devils Tower 271
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.
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.
The Black Hills, Bear Lodge Mountains, and Devils Tower 273
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
wood are left in place for the benefit of wildlife and the
road per square mile) in the Black Hills is still the high-
tion biologistss.
Foothill shrublands are common in the region, typically on the Hogback Rim and lower parts of the Lime-
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.
Jenney wrote:
The Black Hills, Bear Lodge Mountains, and Devils Tower 275
forage for livestock, deer, and elk (fig 16.9).37 After long
small aspens.30
Similarly, R. I. Dodge observed in 1876 that Throughout the Hills the number of trees which bear the marks
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
in the future.42
40
The Black Hills, Bear Lodge Mountains, and Devils Tower 277
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.
intensive crown fires, and still other areas did not burn
and other factors over a large area will promote the evo-
can be protected.
The Black Hills, Bear Lodge Mountains, and Devils Tower 279
wood. The older trees are now dying faster than they
vice versa.
The Black Hills, Bear Lodge Mountains, and Devils Tower 281
than todays.
52
After careful
the Black Hills and Bear Lodge Mountains that will per-
Chapter 17
was warming and the glacier that had reached the basin
282
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).
Albany County.
the basin. The general wrote in his journal that the val-
living in Laramie.
that the more resistant strata became ridges (see fig. 17.2).
ber pine and ponderosa pine are found on the rocky soils.
The raising of cattle and sheep has been the most viable
285
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).
mie. Distinctive for its mass and shape, this natural mon-
ument is more than 200 feet high and has been sculpted
sea that existed about 300 million years ago (fig. 17.4 ).
erosion.
287
over much of the area (see chapter 2). Such rocks have
elevation.
lows of 80F and 49F, respectively (fig. 17.6).16 Annually, most of the basin had only 75100 frost-free days
Green River basins to the west (see fig. 3.7). The mean
the basin.
66
2.4
43
1.6
32
19
-5
0.8
that moisture falls in the basin. The annual precipitaMonthly Precipitation (inches)
90
0.0
surface water.
Vegetation Patterns
west of the basin, the air rising over the Medicine Bow
289
291
basin floor.
the rock surfaces into cracks or into the soil under the
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.
293
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
phic rocks that form the main part of the ridge. Water
ing across the Airport Terrace and Big Hollow. The other
used the land (see chapter 2). Beaver were used hardly
295
Elevation, feet
Modern Little
Laramie River
floodplain
7,500
Little
Laramie
River
terrace
Pleistocene
floodplain
Airport Bench
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
Upland
grassland
Not
vegetated,
playa Upland
grassland
Irrigated Upland
Upland
hayland grassland
grassland
Riparian
today only on land that was too wet for domestic use,
a valuable commodity.
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
Meadow,
upland
grassland
Not
Riparian
vegetated, meadow
landfill
Upland
grassland,
sagebrush
Riparian
Sagebrush,
meadow
mtn.-mahogany
Foothill
Upland
shrubland
grassland,
sagebrush
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.
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
6.5
3.0
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.
31
the mid-1980s (fig. 17.18; see also fig. 3.9). With regard
299
19
Laramie
45
40
15
35
13
30
11
17
25
9
20
7
19
45
40
15
35
13
30
11
17
25
9
20
7
1900
1910
1920
1930
1940
1950
1960
1970
1980
1990
2000
2010
ses have not yet been done for the Laramie Basin. Also,
winter precipitation can be a significant factor for grasslands if it results in considerable water in the soil at the
end of winter.
next chapter.
34
301
Management
Using Western
Landscapes
Chapter 18
lished homes along the way, but the first large wave of
trees used for fuel, shelters, sleds, and weapons. The great-
many people, but they have also led to a need for envi-
onto the land, but only at great cost. New reservoirs and
Water Development
of some rivers are now protected under the Wild and Sce-
are the headwaters of the Snake River and the Clarks Fork
307
ACRE-FEET (millions)
1930
1950
YEAR
1970
1990
2010
fossil fuels are still possible, but public lands are now val-
now cover a very small area and that the best use of such
Habitat Fragmentation
309
Roads
itat far more than the area required for the roadway.
measures is controversial.
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.
Ecosystem Management
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
needed.
313
trends are not good for the Wests economy. For exam-
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,
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.
315
identified.
agreement.21
science-driven approach.19 The process includes willing participants. The first application in Wyoming
habitat improvement.
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
Modern science, technology, and our economic system have greatly augmented the availability of many
319
320 Epilogue
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
2.590
MASS
Ounces to grams
28.4
Pounds to kilograms
0.454
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)
4.19
321
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
Soil order
Location/vegetation type
Entisols
Inceptisols
Aridisols
Mollisols
Alfisols
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
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
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
324
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
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).
western Wyoming.
3. The change that occurs after ecosystem disturbances is known
than those studying bears and humans. See Turner et al. (2001)
the Miocene, about 20 million years ago; bison did not appear
until late in the Pleistocene, less than 300,000 years ago. Bison,
20,000 years ago; see Dott and Batten 1981; Axelrod 1985; Hart
2008.
7. Tidwell 1975.
and federal and state agencies. The U.S. Fish and Wildlife Ser-
9. Schulte et al. 2010; one crater has been found in the present-
Species/Species_Endangered.html.
325
326
11. Gingerich and Clyde (2001) and Clyde (2001) list more than
29. Current estimates range from 16,000 to 13,500 years ago: http://
www.nytimes.com/2012/05/20/opinion/sunday/who-arrived
12. Tidwell 1975; Riedl (1959) describes the fossils of the Shirley
Great Plains and Rockies; see Janetski (1987) for human history
in northwest Wyoming.
and animals in the Bighorn Basin looked like during this time,
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
the bottom sediments and are able to identify the plants that
36. Reider 1983, 1990; Reider et al. (1988) describe grassland soils
15. Zachos et al. 2008; McInerney and Wing 2011; though a global
18. A portion of the initial Medicine Bow Mountains was later sepa-
ogy, and sand dune activity. See also Forman and Pierson
fig.1.2.
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.
45. Pine expansion is coincident with active sand dunes and low
water levels in lakes and riverstwo indicators of a dry or drying climate (B. Shuman, pers. comm.).
Chain Lake Flat, and Battle Springs Flat. About 10,000 years ago
48. Reider et al. (1988) conclude from soil profile data that fluctua-
25. Snoke (1993) and Good and Pierce (2002) describe the forma-
10. This quantity is known as the adiabatic lapse rate; Curtis and
Grimes (2004) estimated a drop of 5.5F per 1,000 feet for Wyo-
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
17. Branson et al. 1967, 1970, 1976. Common salts in the region
grazed the rangeland for only a short time. See also Wentworth
.wyohistory.org/essays/wyoming-cattle-boom.
62. In January 2011 about 1.3 million cattle and 360,000 sheep
1978.
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.
18. The grazing of large herbivores (such as bison, elk, cattle, and
too small.
327
328
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
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
with high snowfall and one with low snowfall; the trend con-
29. Laramie Boomerang, October 19, 2012; more than half (57 per-
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
32. Naftz et al. 2002; Woodhouse et al. 2006; Lundquist et al. 2009;
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.
14. Weynand et al. 1979; Miller 1979; Merritt and Cooper 2000;
37. Williams and Jackson 2007; Williams et al. 2007. For further
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
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.
many land managers are now using beaver for the benefits they
24. Beaver are able to obtain construction material for their dams
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.
49. Shafroth 2005; Owens and Moore 2007; Nagler et al. 2011;
37. Instream flow rights are also recognized on the Wind River
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.
39. D. Brosz, pers. comm.; see chapters 5 and 17 for the importance
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
329
330
30. Mattson 1984; Lemly 2007; Lemly and Cooper 2011; and reports
residents.
(http://www.uwyo.edu/wyndd/reports-and-publications/),
2004.
12. For reviews of wetland adaptations, see Rydin and Jeglum
(2006), Mitsch and Gosselink (2007), and Keddy (2010).
.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
Gage and Cooper (2012) state that 3540 percent of the wet-
19. Salt flats are a common feature in desert shrublands; see chapter
8.
20. The salt deposits are commonly referred to as alkali and the
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/).
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.
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.
16. Coupland and Van Dyne 1979; the weight of all microbial
Program at http://www.cnhp.colostate.edu/cwic/regulations.asp.
web: predatory mites, nematophagous mites, predatory nematodes, fungivorous nematodes, bacteriophagous nematodes,
Chapter 6: Grasslands
collem
bolas, mycophagous prostigmata, amoebae, flagellates,
and phytophagous nematodes.
17. Sims and Coupland 1979; Singh et al. 1983b; Lauenroth et al.
23. Ingham et al. 1985; Knapp and Seastedt 1986; Elliott and Cole-
man 1988; Stanton 1988; Holland and Detling 1990; Burke et al.
1997, 2008; Frank and Evans 1997; Oesterheld et al. 1999; Frank
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
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
buds remain dormant as long as the leaves and stems are intact,
beetles).
al. 1999; Stohlgren et al. 1999; Lauenroth and Burke 2008; Mil
331
332
57. Wright and Bailey 1980 (for the Great Plains); Bragg 1982 (for
Nebraska).
59. Redman (1978) found more water stress and lower plant growth
66. Reed and Peterson 1961; Rodel 1977; Watts et al. 1982; the
45. Rauzi and Smith (1973) and Gifford and Hawkins (1978) review
71. The quote is from Pfadt and Hardy (1987, p. 183); Crist (2008)
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.
ter than do others; see Detling and Painter 1983; Detling et al.
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
and sylvatic plague, for which the animals are now vaccinated
ity, ferrets have been released where large prairie dog colonies
can be found (with about 1,000 ferrets now living in the wild
Each ferret eats about 100 prairie dogs a year. Most present-day
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
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;
Zamora and Tueller 1973; Sabinske and Knight 1978; Baker and
of invasive plants.
13. Richards and Caldwell 1985; Caldwell et al. 1998; Ryel et al.
Hydraulic_redistribution.
18. Welch et al. 1991. Water extracts and volatile vapors from big
Preserve.
man and Hazlett 1977; Weaver and Klarich 1977; Ferreira and
birds are declining more than any other group of North Ameri-
can birds.
94. Burke et al. (1995) estimate 50 years for recovery of soil organic
matter.
333
334
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
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
57. Cowan 1929; Allred 1941; Furniss and Krebill 1972; Parmenter
27. Harniss and Murray 1973; West 1983; Singh et al. 1983b.
Baker (1972).
31. Grayson 2006; Mack and Thompson 1982; Van Vuren 1987;
60. Miller et al. 1993; Sturges 1993; Mensing et al. 2006; Morris et
63. Young and Evans 1978; Wright et al. 1979; Rennick 1981.
al. 1979.
66. Mihlbachler 1986.
67. Smith et al. 1985; Baker 2006.
sagebrush steppe.
72. Seefeldt et al. 2007; Davies et al. 2009b; Shinneman and Baker
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
Schupp (2009) for data on small mammals, which are less com-
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.
3. Smith and Nowak 1990; West (1983) reports that saltbushgreasewood desert shrublands have a mean net aboveground
(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-
86. Van Vuren (1987), Urness (1989), and Byers and Smith (2007)
1960 (p. 28). About William P. Hunt: Morris 2013 (p. 123);
7. White 1976.
8. West 1983.
Poore et al. 2009 (on climate change); see chapter 18 for discus-
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.
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-
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
335
336
38. West and Young 2000; Grayson (2006) concluded that there
would have been large grazers in the late Pleistocene (as discussed
loam, and clay loam. The soils tend to have low infiltration rates
2. Gaylord 1982.
see www.soilcrust.org.
9. Pearson 1965b.
10. High infiltration and storage in the Nebraska Sand Hills provide
15. Martin 1992; Clarke and Rendell 2003; Forman and Pierson
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
18. Brown 1971; the Powder River Breaks are found on weakly
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).
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).
31. Gruell 1983; Arno and Gruell 1986; see chapter 11.
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.
38. Welsh 1957; Baker and Kennedy 1985; Knight et al. 1987.
from seed.
8. Billings 1954.
10. Soils were shallow in all of Wight and Fissers (1968) 55 study
44. Krebill 1972; Beetle 1974a,b; Gruell and Loope 1974; DeByle
45. Elk damage mature trees through antler rubbing or eating bark,
13. Waugh 1986; similarly, Soul et al. (2004) concluded that graz-
14. Burkhardt and Tisdale 1976; Wright et al. 1979; Young and
Evans 1981.
15. Waugh 1986; Miller et al. 1993.
16. Romme et al. 2009.
17. Lyford et al. 2003; Romme et al. 2009.
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.
337
338
33. Lotan and Perry 1983; Christiansen et al. 1987; Amman 1989;
38. The blue-stain fungus does not affect wood strength and is
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.
42. Regniere and Bentz 2007; Chapman et al. 2012. Preisler et al.
46. Klutsch et al. 2009; Kayes and Tinker 2012; Nelson et al. 2014.
plant growth in aspen forests was not correlated with tree den-
47. Collins et al. 2011; Diskin et al. 2011; Kayes and Tinker 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.
21. Smith 1970, 1975; Benkman et al. 1984; Siepielski and Benkman
2007, 2008.
National Forests, see Meyer et al. (2005) and Dillon et al. (2005);
see also the websites of other national forests. Reed (1971, 1976)
57. Baker et al. 2007; Sherriff and Veblen 2007; Schoennagel et al.
2011.
58. Shinneman and Baker 1997.
59. Ponderosa pine re-establishment was extremely slow on some
and other land uses began to limit fire spread as early as the late
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
31. Veblen et al. 1991b,c; Brunelle et al. 2008; Raffa et al. 2008.
climate.
63. Arno and Gruell 1986, Goldblum and Veblen 1992; Baker 2009.
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
68. Moir 1969; Despain 1973, 1983, 1990; Whipple and Dix 1979;
Lotan and Perry 1983; Parker 1986; Peet 2000. Lodgepole pine
may be the only conifer species that can tolerate some infertile
sites.
69. Romme 1982.
ships with conifers and other woody species. Many early stud-
the slash open after a few weeks and release seed, which re
75. Benkman and Siepielski 2004; Benkman et al. 2012; see note 23.
103. Houston 1973; Loope and Gruell 1973; Beetle 1974b; Walters
et al. 1982; DeByle et al. 1987; Boyce 1989; Seager et al. 2013.
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;
alpine fir, white spruce is common in the Black Hills, and white
83. Romme and Knight 1981; Johnson and Fryer 1989; Veblen et al.
1991a.
87. Schmid and Hinds 1974; Alexander 1987a; Baker and Veblen
89. Oosting and Reed 1952; Shea 1985; Veblen 1986a,b; Alexander
or 100 years old. Sites having a higher site index are generally
4. Fogel and Trappe 1978; Maser et al. 1978, 1988; McIntire 1984.
339
340
7. Harder 1979.
9. For this unit of measure, the volume of water depends on the size
the volume of water that will cover a specified area 6 inches deep.
were initially elevated after the 1988 Yellowstone fires, but post-
1986) found that annual wood production per unit area returned
17. Gary and Troendle 1982; Troendle 1983, 1987; Knight et al. 1985;
19. Eighty percent of the roots are in the top 8 inches of the soil.
22. Fahey et al. 1985; Knight et al. 1985; Fahey and Knight 1986.
41. Pugh and Small 2012. Clow et al. (2011) observed higher soil
25. Gosz 1980; Fahey 1983; Prescott et al. 1989; Knight 1991.
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
48. Troendle 1987; Troendle and Nilles 1987; Prescott et al. 1989a;
and Franklin 1997; Perera et al. 2004; Dillon et al. 2005; Meyer
57. Knight et al. 1991; Parsons et al. 1994; killing 60 percent of the
extent possible.
types.
65. Bentz et al. 2010; little is known about the fir bark beetle.
66. Hicke et al. 2012; see also Klutsch et al. 2011; chapter 11.
8. Buckner 1977.
69. Lynch et al. 2006; the spatial pattern of burning in the 1988
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.
71. Westerling et al. 2006; Raffa et al. 2008; Logan et al. 2010.
ing http://www.firewise.org/resources/files/Around-Your-Home
.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.
341
342
.fs.fed.us/rm/highelevationwhitepines/Threats.
17. Resler and Tomback 2008; Keane and Schoettle 2011; Larson
2011.
19. Smith and Geller 1979; Tranquillini 1979; Hadley and Smith
1986.
20. Scott 1995.
21. Exceptions are the small annual plants studied by Reynolds
23. Scott and Billings 1964; Sakai and Otsuka 1970; Kimball et al.
1973; Wallace and Harrison 1978; Bell and Bliss 1979; Hamer-
lynck 1992.
24. Krner 2012.
25. Rochow 1970.
26. Spomer 1964; Campbell 1997 (plants); Molenda et al. 2012
(insects).
27. Knutson 1981.
30. Huntly and Inouye 1988; Armstrong et al. 2001; Seastedt 2001.
branches of spruce and fir produce roots while pressed to the soil
3). See Richmond (1949) for an early study in the Wind River
Mountains.
34. Billings and Mooney (1959) studied cryoturbation in the Medi-
14. Eversman 1968; Potter 1969; Mears 1975; Malanson and Butler
pine blister rust causes swelling on the branches and trunk, even-
National Park.
37. Bowman and Fisk 2001; Monson et al. 2001; as in forests, some
pine blister rust, the fungus requires plants in the genus Ribes
16. Janetski 1987 (chapters 2 and 3); Frison 1991; Kornfeld et al.
40. Initially the primary source for nitrogen was thought to be the
nitrous oxides coming from the Salt Lake City area, as reported
by Sievering (2001).
41. Fenn et al. 2003; Hood et al. 2003; Seastedt et al. 2004; Liptzin
and Seastedt 2010.
42. Fenn et al. 2003.
44. Carrera et al. (1991), who calculated that the upper treeline
Anderson 2006.
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,
al. 2012).
1. Eversman and Carr 1992; Hansen et al. 2002. Also see the Atlas
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.
3. Dorf 1964.
45. Ecologists Andrew Hansen and Jay Rotella (2002) found that the
13.
Haines 1996; http://www.cr.nps.gov/history/online_books/
haines1/iee2d.htm.
14. Meagher and Houston (1998) compared modern photographs
in YNP with those taken by early explorers.
343
344
Cross et al. 2010; Bienen and Tabor 2006; brucellosis and other
68. Shullery 1989; several other fires also were ignited by humans,
the controversy.
50. Elk are not the only wildlife species adversely affected by land
71. Kay 1993; Romme et al. 1997, 2005; Stevens et al. 1999.
75. For example, species that are now rare or absent in the GYE,
the warblers; the birds that are found in the park have come from
78. Tomback and Achuff 2010; the rust kills other five-needle pines
83. http://www.greateryellowstonescience.org/download_product/
538/0.
85. In 2012 more than 300,000 lake trout were removed from
86. Middleton et al. 2013d; see also Wilmers and Post 2006.
58. Patten 1968; Singer et al. 1994; Yellowstone National Park 1997.
across the GYE in 1956 and 2001; they found an average loss of
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
Bear Butte, Black Butte, Crow Peak, Custer Peak, Devils Tower,
and emphasize that general patterns and effects are not clear.
See also http://nyti.ms/1cHqikq.
types; see also Larson and Johnson 1999; Marriott et al. 1999;
history of the area and include road logs that identify geo-
23.
MacCracken et al. 1983; Hoffman and Alexander 1987;
tribes along the way, traveling most likely to the Bighorn Moun-
24. Boldt and Van Deusen 1974; Shepperd and Battaglia 2002.
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.
28. Fisher et al. 1987; Wienk et al. 2004; Brown 2006; Brown et al.
2008.
345
346
43. Brown (2006) found that fires were more frequent during La
45. Shepperd and Battaglia (2002) review the ecology of the vari-
Armillaria root rot, red rot, western gall rust, needle cast, and
Katabatic_wind.
16. The Wyoming Water Resources Data System (http://www
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.
21. The Wyoming Game and Fish Department has a report titled
1. The one basin with no outlet is the Great Divide Basin, where
4. Shuman 2012.
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)
8. Fremont 1845; Stansbury 1852; Smith 1913 (p. 272); Dale 1918
5. http://www.rivers.gov/rivers/wyoming.php.
6. Ashworth 2006.
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).
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.
.wyo.gov/web2011/Departments/Hunting/pdfs/Regulations_
Ch62.pdf.
30. Ranchers prefer more grasses than sedges and rushes, which
15. Hansen et al. 2002 (p. 160); see also Power 2005.
17. Daily and Ellison 2002; Clark 2007; Esty 2007; Banerjee et al.
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.
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.
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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
aerenchyma, 6870
amphibians: in forests, 179t; in wetlands, 68, 70. See also specific types
Andersen, Chamois, 41
frost-free period and, 33; in grasslands, 83, 92, 93, 100, 1078;
antelope bitterbrush, 12021, 143, 156, 158t, 160, 16466, 165f, 176t,
254
Akashi, Yoshiko, 55
apomixis, 236
allelopathy, 79
arrowgrass, 72t
aspen forests, woodlands: 8f, 16770, 19698, 196f, 197f; in Black Hills,
272, 273, 273f; climate change in, 219220; cloning by, 168, 196
97; dominant plant species in, 15859t; 17677t; elk browsing on,
in, 236, 239; climate change in, 24142; dominant plant species
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)
badlands, 14748; animal species in, 147; dominant plant species in,
ture changes in, 40; formation of, 15; meadows in, 224f
14748; geologic history of, 17; topography of, 2930, 147
Index
bluestem, little, 83, 84f, 88, 100, 155, 159t, 163, 166, 268, 27374,
277
caldera, 245
Caldwell, M. M., 115, 331n8
cambium, 187f
bogs: vs. fens, 77; for geologic history, 14; lack of, in Wyoming,
canarygrass, reed, 79
329n1
Bonneville, Lake, 19
carbon credits, 11
Boysen Reservoir, 55
and, 34
Briske, David, 97
Casper Formation, 284f, 286, 287, 287f, 293, 293f, 295, 297f
broadleaf cattail, 79
brucellosis, 344n48
bryophytes, 330n29
334n73
bunch grasses, 83
tation by, 311, 311f; of mountain forests, 178. See also timber
harvesting
389
390 Index
Cleary, M., 119
coyote, 265
Clements, Frederick, 96
climate, 38; history of, 1223, 32f, 39f, 326n36; vs. weather, 38. See
of, 3840, 39f; current rate of, 15, 3841; in desert shrublands,
Crow Creek, 54
crown fires, 37; and bark beetle outbreaks, 217; in Black Hills,
7980
common reed, 79
Como Bluff, 13
compensatory growth hypothesis, 95
decreaser species, 95
by, 310
denitrification, 207
conveyance losses, 57
Cooke, P. G., 54
Copeland, H., 80
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-
328n13
soils of, 13134, 139, 335n4; water availability in, 131, 132, 132f,
13739; in Yellowstone National Park, 250
Coughenour, M., 97
Detling, J., 97
Index
Development by Design, 31516
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
Ellison, L., 96
El Nio, 31
endemic species, 9
entisols, 322t
drip irrigation, 62
erosion: in badlands, 147; beavers and, 59; in Black Hills and Bear
100, 332n62; history of, 24f, 32, 32f, 327n8; in sagebrush steppe,
eutrophication, 240
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
steppe, 116
Ewers, B., 119
extinction: climate change and, 40; in Cretaceous-Paleocene
mountain peaks, 27, 173; plant adaptations for, 27, 29f; and pre-
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,
of, 239
391
392 Index
fences: around national parks, 280; along roads, 311
fens, 7779, 77f; vs. bogs, 77; classification of, 78, 330n35; climate
change in, 80; dominant plant species in, 68, 7273t, 7779; soils
of, 6870
tion in, 31112; elevation gradient in, 173, 174f, 180; energy flow
228, 229; fires in, 18182, 20815, 21618, 216f; future chal-
on, 2526, 21115, 212f, 213f, 30812; hydrology of, 2027, 211,
fescue, Idaho, 50f, 121, 123, 159t, 164, 166, 176t, 194f, 221, 222t,
201, 201f, 203f, 2078, 210, 211, 21415; parasitic plants in,
in, 2034, 204f, 21516; slow vs. rapid changes to, 181; soils of,
fires: bark beetle outbreaks and, 21617, 341n64; in Black Hills, 187,
212f, 213f; tree growth rates in, 200201, 201f, 202f; understory
157, 162, 164, 16566, 170; frequency of, 121, 181, 216f; future
Fort Union Formation: badlands in, 336n18; coal deposits in, 326n13
Native American use of, 23, 37, 98, 99; as natural disturbance, 37,
Fossil Lake, 15
112, 118, 119, 12123, 122f, 123f, 334n32; in sand dunes, 143;
Extinction Event in, 14; Mesozoic, 13; Paleozoic, 1213; tree, 14,
first-order streams, 46
fracking, 3078
Friggens, M., 41
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;
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,
in, 16872; riparian zones compared to, 155; roads in, 311;
genets, 197
shrublands in, 156, 156f, 157, 16466, 165f, 290f; topography of,
girdling, 184
species in, 17980, 179t, 19899; area covered by, 8f, 215,
Index
glaciers: and climate change, 40; and floodplains, 19, 5152; forma-
of, 85, 87f, 94, 9697, 332n48; in prairie dog colonies, 9798,
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
Gondwanaland, 13f
239, 240f
Great Divide Basin, 28; desert shrublands in, 131f, 133f, 138f; lack of
Goshen Hole, 74
grama, blue, 36, 8384, 88, 94, 100, 109f, 118, 13435, 136t, 150,
248, 249; animal species in, 245; bark beetle outbreaks in, 185,
in, 248; elevation map for, 246f; elk in, 25658, 257f, 26465;
fires in, 216, 25862; foothills of, 165f, 169f; geologic history of,
of, 233f, 253; boundaries of, 252; dominant plants in, 25254,
253f; elevation of, 246, 247f, 253, 253f; establishment of, 3, 252;
228; riparian zones of, grazing in, 6061, 258; wolves in, 256,
83, 92, 93, 100, 1078; animals of, 8687, 88t; in Black Hills,
tion of, 1067, 333n94; desert, 133, 13637t; distribution of, 83;
dominant plants in, 8384; drought in, 86, 100, 332n62; effects
of climate change on, 1078; effects of grazing on, 95f, 105f, 298f
(food chain) in, 85, 8992, 91t; fairy rings in, 84, 85f; fertilizers
added to, 92, 93, 106; fires in, 37, 85, 98100, 98f, 107, 334n32;
grouse, species of, 128; dusky, 335n96; sharptailed, 335n96. See also
1036, 333n82; in Laramie Basin, 85f, 99f, 285, 289, 290f, 291f,
sage-grouse, greater
333n2; nutrient cycling in, 9294, 99; precipitation in, 32, 89;
Gruell, G., 98
117; sagebrush steppe mixed with, 110, 112, 114f; soils of, 34,
83, 8485, 86f, 9293, 108; succession in, 94; tallgrass prairie,
half-shrubs, 331n3
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
Hart, R., 25
intercalary meristems, 85
interception, 204
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,
27879; control of, 65, 79, 104, 1056, 124, 329n50; vs. crops,
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
horses, feral: human use of, 23, 24, 127; in sagebrush steppe, 12728,
hummocks, 76
hydrophytes, 45, 76
juniper, common, 145, 158t, 176t, 183f, 271, 272, 277, 295
hyphae, 202
juniper, Rocky Mountain, 23f, 157, 160, 161f, 162, 337n28; browsing
on, 171f
Kentucky bluegrass, 79
Index
keystone habitats, wetlands as, 66
levees, 62
keystone species, 57; beaver as, 57; prairie dog as, 102
Krueger, K., 98
livestock: arrival of, 25; current number of, 327n62; food preferences
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
La Nia, 31
loosestrife, purple, 79
Laramie Basin, 282301; climate change in, 299301, 299f, 300f; cli-
mate of, 28889, 289f; dominant plant species in, 28993; effect
of livestock grazing in, 298, elevation of, 283f, 288; fires in,
features of, 282, 284f, 28588, 288f; geologic history of, 282,
285, 287, 346n3; geologic map of, 286f; grasslands in, 85f, 99f,
285, 289, 290f, 291f, 29899; human history in, 28285, 29596;
in, 295301; maps of, 283f, 286f, 290f; mima mounds in, 149,
150f, 151, 293; playa wetlands of, 7475, 74f, 29293; rainshadow
effect in, 288; sagebrush in, 11011, 289, 292; vegetation patterns
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
Laurasia, 13f
marshes, 7074; climate change in, 80; dominant plant species in,
mastodons, 282
Lebo, Lake, 15
395
396 Index
Mears, Brainerd, Jr., 295
230, 230f, 231, 232f, 233, 239f, 342n34, 343n45; fens in, 68;
foothills of, 165f; forests in, 190f, 194f, 204, 211, 340n30;
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
meristems, 85
of, 223, 225f, 341n7; wet, 75; in Yellowstone National Park, 228,
250, 252f
mountain pine beetle, 181f, 18285; in Black Hills, 27778; climate
change and, 216, 341n63; effects of, 18285, 182f, 183f, 219,
beetles
mudflats, 7475
Creek, 59f
Missoula, Lake, 19
Mitchell, J., 25
mycoheterotrophs, 175
mixed-grass prairie, 83, 84f; adaptations to, 85; agriculture in, 100;
by, 339n5
Myers, W., 295
moose, 343n45
mountain(s), 5f; droughts in, 32; elevation of peaks of, 27, 173;
formation of, 173; geologic history of, 1317, 173; map of, 5f;
345nn12; fires used by, 23, 37, 98, 99; horses used by, 127; hunt-
ing by, 24, 332n41; impact on ecosystems by, 23, 24, 326n49; in
249, 343n16
naturalized species, 63, 79
Index
natural regulation policy (Yellowstone National Park), 256,
25758
Painter, E., 95
paleosols, 146
pantodonts, 14
336n14; in wetlands, 68
peat mining, 79
Nugget Sandstone, 13
phreatophytes, 57, 65
physiological ecology, 5
pickleweed, 132f
pika, American, 236, 236f, 242
obsidian, 249
Ocean Lake, 74
O horizon, 200
of, 8f, 174, 338n4; doghair stands of, 193, 193f; dominant plant
species in, 17677t; fires and, 182, 191, 19293, 20810, 209f,
210f, 211f, 213f, 216, 218; forests of, 19093, 190f, 191f; in Grand
old-man-of-the-mountain, 236f
Oligocene Epoch, 15
open spaces: persistence of, 306, 306f, 307f; value of, 314
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
Hills, 27072, 271f, 274, 275f, 276f, 281; climate change and,
compared to, 18889; fires and, 182, 186f, 187, 187f, 274, 275f,
private lands: area of, 6f; ecosystem management in, 31213, 31516;
treeline, 231, 233, 234; white pine blister rust and, 234, 234f,
pine cones, serotinous, 17980, 19192, 192f, 339n71. See also pine,
lodgepole
Pinedale glacial period, 20, 24648, 326n23
pinedrops, 175
Pine Ridge, 163f
puffballs, 202
pugging, 76
purple loosestrife, 79
playa wetlands, 7475, 74f; dominant plant species in, 7273t, 7475,
radiocarbon dating, 22
Pliocene Epoch, 17
ramets, 197
Raynolds, W. F., 53
redbeds, 13
porcupines, 202
Powder River Basin: badlands in, 14748, 336n18; coal deposits in,
14, 326n13; escarpments in, 155f; fires in, 98, 98f; geologic his-
tory of, 13; grasshoppers in, 101, 120; habitat fragmentation in,
reed, common, 79
130
reed canarygrass, 79
rest-rotation grazing, 97
range of, 30; ratio of evaporation to, 34, 35, 203; in sagebrush
steppe, 11011, 112, 112f; in sand dunes, 145; seasonality of, 31,
rhizosphere, 70
Index
Richards, J. H., 115, 331n8
Laramie Basin, 11011, 289, 290f, 292; longevity of, 116; old-
riparian landscapes, 4565; area covered by, 45; bank storage in, 57,
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-
cal role of, 45; foothills compared to, 155; future challenges in,
tion of, 111, 112; in foothills, 164; in Grand Teton National Park,
Basin, 29293, 290f, 293f, 294f; livestock in, 6061; map of, 46f;
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,
in, 109; animals species in, 110, 128; climate change in, 120,
species in, 11012; drought in, 12021; energy flow in, 116,
118; fires in, 37, 110, 112, 118, 119, 12123, 122f, 123f, 334n32;
Rocky Mountain National Park: alpine tundra and upper treeline in,
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
of, 110; topography of, 110, 112; water availability and growth
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
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
fossil pollen from, 20, 21f; in Grand Teton National Park, 254,
399
400 Index
salt-secreting glands, 132f
Shirley Basin: black-footed ferrets in, 333n76; fossil logs in, 14; mima
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
in, 145; in sagebrush steppe, 112, 115f; soils of, 14243; stabiliza-
Sierra Madre: forests in, 196, 196f, 197f; mountain meadows in, 227
Sioux, 345n2
saprophytic fungi, 84
skunkbush sumac, 47t, 5456, 121, 148, 15556, 158t, 16067, 166f,
Smith, B., 59
second-order streams, 46
Smith, Jedediah, 24
wetlands, 68, 69f, 7071, 72t, 77, 77f, 78, 250; in Yellowstone
Snake River: channelizing of, 6162; dam on, 252, 328n16; vegeta-
tion along, 52f, 253; Wild and Scenic Rivers Act protection of
headwaters, 307
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;
335n18, 346n22
327n62
Sheep Mountain, 75f, 293f, 295
Index
material of, 3435; in plant growth, 3436, 36f; of riparian
surface fires, 37; in Black Hills, 274, 275, 276; in Douglas-fir wood-
Tatman, Lake, 15
spiny hopsage, 112, 13335, 136t, 137, 143, 144f, 335n2, 335n18,
336n35, 346n22
springs: in Laramie Basin, 286; in Yellowstone National Park, 251
Laramie Basin, 288, 289f, 299, 299f, 301; map of, 33f; range of,
Teton Range: elevation of, 246, 247f, 248f; formation of, 17, 24546,
Thompson, C., 15
streams: in alpine tundra, 236; ephemeral, 45, 62, 65; map of, 46f;
water movement in, 45. See also riparian landscapes
Strong, W. E., 249
evapotranspiration
401
402 Index
Trappers Point, 31011, 347n10
treeline, upper, 23042; adaptations to, 233; advancing of, 24142,
342n1, 342n3; krummholz at, 23133, 232f; soils of, 232; white
trona deposits, 15
Uinta Mountains: foothills of, 162; formation of, 15; upper treeline
in, 241
Uncompahgre Plateau, 220f
area covered by, 45, 66; biological diversity of, 67; climate change
role of, 45, 66, 329n1; future challenges in, 7980; hydrophytes
used to define, 45; invasive plants in, 79; irrigation water and,
67, 74, 76, 80, 346n17; as landscape sinks, 79; of the Laramie
loss of, 66, 79, 330n40; seasonal dry periods of, 67, 68, 70; soils
of, 6870; types of, 45, 66; water movement in nonriparian vs.
Utah juniper, 22, 23f, 148, 157, 158t, 160, 161f, 162
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,
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-
231, 233, 234; white pine blister rust and, 234, 234f, 26263; in
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,
Index
Wight, J. R., 337n10
distribution in, 8f; elevation of, 3, 4f; human population of, 305;
public, private, tribal, and state lands in, 3, 6f; roads and cities
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
wind: in alpine tundra and upper treeline, 231, 23233, 237, 237f;
xerophytes, 8586
34
Wind Cave, 270
Wind Cave National Park: bison in, 97, 97f, 268; black-footed ferrets
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
Wind River, 53
Park
Yellowstone Lake, 248, 344n85
Yellowstone National Park (YNP), 24951; bark beetle outbreaks in,
217, 341n69; dominant plant species in, 25051; early explorers
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
foothills of, 250, 251f; forests in, 189f, 192, 193, 250; fossil trees
in, 14, 15, 245, 247f; geologic history of, 245; grasslands in, 250,
258, 344n53; lakes and rivers of, 249; livestock in, 342n20(ch13);
plants in, 68; mountain meadows in, 228, 250, 252f; precipita-
tion in, average annual, 30; sagebrush in, 114f, 250; temperature
changes in, average, 40; topography of, 20, 246f, 249; volcanoes
in, 20, 245, 326n25. See also Greater Yellowstone Ecosystem
Yellowstone Plateau, 21920
403
ecosystems.
mate change.
federal agencies.
stone Ecosystem.
404