Succession in riparian communities of the lower Yellowstone River, Montana
by Keith Webster Boggs
A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in
Biological Sciences
Montana State University
© Copyright by Keith Webster Boggs (1984)
Abstract:
Riparian plant communities of the lower Yellowstone River between Glendive and Sidney, Montana,
were studied during the Summers of 1980 and 1981. These communities originally colonized sand and
gravel bars deposited along the River channel. New sand deposits were invaded by Salix spp. and
Populus deltoides Marsh. seedlings. This original community developed sequentially to willow
thickets, to young cottonwood forests, and to mature cottonwood forests after about 3, 7, 34 and 92
years, respectively. While the mature cottonwood community is usually replaced by a grassland
community (principally Agropyron smithii Rydb.) via a shrub community (Rosa woodsii Lindl. and
Symphori-carpos occidental is Hook.), it may be replaced by a Fraxinus Pennsylvania Marsh. plant
community. Numbers of species with constancies of over 60% rose from 12 in the seedling community
to. 18 in the mature cottonwood community and declined to 13 in the grassland. Composition changes
are documented in releve tables. Populus deltoides density fell rapidly from 48/M2at 3 years to
0.02/M2 at 92 years; Salix spp. disappeared still more rapidly, with declines from 10/nr at 3 years to
near 0/m2 at 34 years. Aboveground biomass rose from 0.2 kg/m2 at 3 years to 30 kg/nr at 60 years
and declined to less than 0.5 kg/m2 in near-climax grasslands; most of the large mass observed at
mid-sere is living wood.. Belowground biomass rose from about 7 kg/m2 at 3 years to over 30 kg/nr at
90 years and declined to about 19 kg/m2 in grasslands; over half of this biomass was soil organic,
matter in every community. Root/shoot ratios declined from 3/1 in the seedling community to 1/2 in the
mature cottonwood community and rose again to 10/1 in the grassland. K, Na and N contents of the
communities and ecosystem rose and fell with the rise and fall in its biomass. In contrast, P rose more
rapidly in early succession and continued to rise slowly through the grassland stage. Management
implications of logging and altered Streamflow are discussed. ©
COPYRIGHT
by
Keith Webster Boggs
1984
Al I Rights Reserved
SUCCESSION IN RIPARIAN COMMUNITIES OF THE
LOWER YELLOWSTONE RIVER, MONTANA
by
Keith Webster Boggs
A thesis submitted in p a r tia l f u l f i ll m e n t
o f the requirements fo r the degree
of
Master of Science
in
Biological Sciences
MONTANA STATE UNIVERSITY
Bozeman, Montana
June 1984
APPROVAL
of a thesis submitted by
Keith Webster Boggs
This thesis has been read by each member of the thesis committee
and has been found to be sa tis fac to ry regarding content, English usage,
format, c it a t io n s , bibliographic s t y le , and consistency, and is ready
for submission to the College of Graduate Studies.
iT
Iw or n s j
Date
'
I'
Chairperson, Graduate Committee
Approved for the Major Department
Date
Head, Major Department
Approved for the College o f Graduate Studies
3
Date
Xa
Graduate Dean
STATEMENT OF PERMISSION TO'USE
In presenting this thesis in p a r tia l f u l f i ll m e n t o f the require­
ments fo r a master's degree a t Montana State U niversity, I agree that
the Library shall make i t availa b le to borrowers under rules of the
Library.
B rie f quotations from this thesis are allowable without spec­
ia l permission, provided that accurate acknowledgment of source is made.
Permission fo r extensive quotation from or reproduction of this
thesis may be granted by my major professor, or in his/her absence, by
the Director of Libraries when, in the opinion of e it h e r , the proposed
use o f the material is fo r scholarly purposes.
Any copying or use pf
the material in this thesis fo r fin an cial gain shall pot be allowed
without my w ritte n permission.
\
Signature
Date
iv
ACKNOWLEDGMENTS
I am grateful to my major professor. Dr. Theodore W. Weaver I I I ,
and to Dr. Richard J. Mackie for t h e ir advice and in te r e s t in this
study and t h e ir constructive c ritic is m o f the manuscript.
I am also
grateful to Dr. John H. Rumely fo r his help during the study and c r i t ­
ical review of the manuscript.
I thank Gary Dusek fo r his important
assistance and advice during my two f i e l d seasons o f c o lle c tin g data
and Dr. Jim Pickett fo r reviewing the manuscript.
I thank my w ife,
S a lly , fo r her assistance and encouragement throughout the course of
this study.
F in a lly , I thank the Montana State Department of Fish,
W ild lif e and Parks fp r providing funds and equipment needed to carry
out this project.
\
*.
V
TABLE OF CONTENTS
Page
LIST OF TABLES......................... ............................................................................
vi
LIST QF FIGURES....................................................................................
ix
. ABSTRACT:..............................................................................
x
INTRODUCTION...........................................................................................................
I
DESCRIPTION OF STUDY AREA...................................................................................
?
MATERIALS AND METHODS......................................................................
9
Species Composition...............................................
S ite Age....................................... ..................................................................
Stand Height........ ................
Plant Cover................................... ...................... ..........................................
Density o f Wopdy P lants.................................! .......................... .............
Aboveground Biomass----- '.................................................
Aboveground Dead Biomass..................................................; ......................
Belowground Biomass.....................................................................................
System Nutrient Mass...................................................................................
RESULTS AND DISCUSSION...........................................................
Grassland Sere Community Descriptions......................... .................., .
Green Ash Sere Community Descriptions....................
Peach-leaved Willow Community Descriptions................
Evidence for Successional R elations......................
Species Numbers....................................................................................
Density of Trees and Shrubs........................................... '.......................
Discussion of Serai Causes and Relations.........................................
Changes in Aboveground Biomass Through Succession....................
Changes in Belowground Biomass Through Succession.......................
Changes in Nutrient Content Through S u c c e s s io n ...,.....................
9
IO
10
10.
11
12
14
16
18
20
23
29
30
?1
33
.33
35
37
41
42
MANAGEMENT IMPLICATIONS............................................................................
49
LITERATURE CITED............................
5p
APPENDIX...................................................................... ’ .............................................
'5 5
vi
L I S T O f TABLES
Tabl e
Page
I.
Coverage {%) pf Common Sppcies in Ten Communities.
2.
Plpnt Coverage, Species Richness and Community
Height in Nine Community Types o f the Lowpr
Yellowstone River Floodplain___
. ..
3.
4.
V
Mean Age and Diameter of Popul us del toides Trees
Found in Four Community T y p e s ....
1
32
. Density (number/100 m2 ) of Woody Plant Specie? in
Nine Community T y p e s . . . , ...........
34
5.
Locations o f Stand? Studies.....................................
K
R
Vu
6.
Composite Vascular Plant Species1 L is t of the Lower
Yellowstone River Floodplain........ ............. ...........................
57
Ages of Representative Populus deltoides and S alix
f l u v i a t i l i s in Stands' S t u d i e s ! . : : : . . . ' ........ ..........
6?
Elevation (m), Relptiye tp the River Surface, pf Stands
S tudied!.......... ...............................................
1 r .
.............................r t . . f
63
Height (m) 1 o f the Upper Canppy Surface and Diameter
(cm)^ o f Ropulus deltoides in Stands Studied...................
64
7.
8.
9.
10.
Presence {% Coverage) 1 p f Spegie$ in the Seedling
Community Type. ...................................................... ;.
11.
Presence (% Coverage) of Species in the Thickett
Community Type......................................................
67
Presence (% Coverage) of Species in the Young
Cottonwood Community Type.................................
69
Presence (% Coverage) pf Species in the Mature
Cottonwood Community Type...............................
71
12.
13.
v ii
LIST
OF
TABLES— Continued
Table
14.
15.
16.
17.
18.
Page
Presence (% Coverage) o f Species in the Shrub
Community Type...............................................................................
73
Presence (% Coverage) of Species in the Grassland
Communi ty Type........................... .............................. ....................
75
Presence (% Coverage), of Species in the Green Ash
Community Type.................................................................................
77
Presence (% Coverage) of Species in the Willow-Shrub,
Peach-Leaved Willow and MarshCommunityTypes.....................
79
Density (number/10 m^) o f Trees, by Diameter Class1 , in
the Seedling and Thickett Community Types...........................
.81
19.
Density (number/1000 m2 ) of Trees, by Digmeter Class, in
the Yogng Cottonwood and Mature Cottonwood Community
Types........ ................ ........................................ . . ; ' ............ 82
20.
Density (number/10 m2) of Trees, by Diameter Class, in
the Shrub Community Type.......................................................
21.
22.
23.
24.
25.
83
Density (number/1000 m2) of Trees, by Diameter Class, in
the Green Ash, Willow-Shrub and Peach-Leaved Willow
Community Types..............................................................................
84
Density /ngmber/lOO m2 ) Qf Shrubs, by Canopy Diameter
C lass', in the Young Cottonwood and Mature
Cottonwood Community Types...............................................
86
Density (number/100 m2) o f
Class, in the Shrub and
p
Density (number/100 m ) of
Class, in the Green Ash
Leaved Willow Community
Shrubs, by Canopy Diameter
Grassland Community Types........
88
Shrubs, by Canopy Diameter
Willow-Shrub and PeachTypes......................
90
I
9
Constants and Estimates of Relative Error fo r
Regressions Relating Logarithm Plant Weights to Their
Logarithm D ia m e t e r s ...,.....................................................
92
viii
L I S T OF TABLES— C o n t in u e d
Table
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
Page
2
Bulk Density (g/m ) of Soils from Three Horizons
in Seven Community Types'..........................................................
93
Elemental Contents (%) of Riparian Plants of the
Lower Yellowstone R iv e r ! ............................................................
94
Organic Matter Contents (%) of Soils,from Three
Horizons in Seven Community Types .......................................
96
Kjeldahl Nitrogen Contents {%) of Soils from Three
Horizons in Seven Community Types ................................. ..
97
Total Phosphorus Content (%) of Soils in Three Horizons
in Seven Community Types'..........................................................
98
Bicarbonate Extractable Phosphorus Contents [%>) of
Soils in Three Horizons in SevenCommunity Types!..............
99
Ammonium Acetate Extractable Potassium Content (%) of
Soils from Three Horizons in Seven CommunityT y p e s !...
Ammonium Acetate Extractable Sodium Content (meq/lOOg) ,
of Soils from Three Horizons in Seven Community Types
O
Biomass (g/m ) Present in the Sandbar and Seedling
Community Types...............................................................................
0
Biomass (g/m ) Present in the Thickett and Young
Cottonwood Community Types........................................................
2
Biomass (g/m ) Present in the Mature Cottonwood and
Shrub Community Types..................................................................
P
Biomass (g/m ) Present in the Grassland Community Type...
100
101
102
103
105
107
ix
/
'
L I S T OF FIGURES
Figure
L
Page
Map o f the Study Area Showing Locations of
Specific Study S it e s ....................................................................
3
Mean, Maximum and Minimum Yellowstone River Flow
(cubic m/sec) a t Sidney, MT in the 1910-1980
Period (USGS 1910-1980)..............................................................
4
The Average Monthly Temperature and P recipitatio n
at Glendive, MT ( Ruffner 1 971)................................................
7
Flow Chart Summarizing Successional Changes Observed
in the Lower Yellowstone River Floodplain.........................
' 21
5.
Age^Range and Physiognomy o f Community Types..........................
22
6.
Th? Elevations of the Soil Surface Above the
Yellowstone Riyer Surface fo r Communities o f
Increasing Age.................................................................................
24
Height of the Upper Canopy Surface, Relative to the
Ground Surface, in Communities o f Increasing Age.............
26
Above and Belowground Biomasses o f Communities o f
Increasing Age, Grassland Sere.............................................
3?
Abovegrounfl Masses o f S alix s pp., Populus d e lto id e s ,
Shrubs and Herbs in Communities o f Increasing Age,
Grassland Sere......................................................................
40
Total Ecosystem Contents of Organic Matter, Phpsphorus,
Nitrpgen, Sodium and Potassium..............................................
451
Elemental Contents of Living Plants as Related to
System Age............................. '..........................................................
47
2.
3.
4.
7.
8.
9.
10.
11.
X
ABSTRACT
Riparian plant communities o f the lower Yellowstone River between
Glendive and Sidney, Montana, were studied during the Summers of 1980
and 1981. These communities o r ig in a lly colonized sand and gravel bars
deposited along the River channel. New sand deposits were invaded by
Salix spp. and Populus deltoides Marsh, seedlings. This orig inal
community developed sequentially to willow th ic k e ts , to young cotton­
wood fo re s ts , and to mature cottonwood forests a f t e r about 3 , 7 , 34 and
92 years, resp ectively. While the mature cottonwood community is
usually replaced by a grassland community (p r in c ip a lly Agropyron
sm ithii Rydb.) via a shrub community ( Rosa woodsi i L in d l. and Symphoricarpos occidental is Hook.), i t may be replaced by a Fraxinus pennsylvanica Marsh, plant community. Numbers of species with constancies of
over 60% rose from 12 in the seedling community to. 18 in the mature
cottonwood community and declined to 13 in the grassland. Composition
changes are documented,,!'n releve tables.
Populus deltoides density
f e l l rapidly from 48/nT a t 3 years to 0.02/m2 at 92 years; Salix spp.
disappeared s t i l l , more ra p id ly , with declines from 10/m2 a t 3 years to
near 0/m2 a t 34 y ears. Aboveground biomass rose from 0.2 kg/m2 at 3
years to 30 kg/m2 a t 60 years and declined to less than 0.5 kg/m2 in
near-climax grasslands; most of the large mass observed a t mid-sere is
liv in g wood.. Belowground biomass rose from about 7 kg/m2 a t '3 years to
over 30 kg/rrr a t 90 years and declined to about 19 kg/m2 in grasslands;
over hq lf of this biomass was soil organic, matter in every community.
Root/shoot ratios declined from 3/1 in the seedling community to 1/2 in
the mature cottonwood community and rose again to 10/1 in the grassland.
K, Na and N contents o f the communities and ecosystem rose and f e l l with
the ris e and f a l l in it s biomass. In contrast, P rose more rapidly in
early succession and continued to ris e slowly through the grassland
stage. Management implications of logging and altered StreamfTow are
discussed.
I
INTRODUCTION
Recent studies of w h ite -ta ile d deer ( Odocoileus v irg in ia n u s ) on
the lower Yellowstone River ( Dusek 1981) required information about the
rip a ria n communities.
A previous study of the area provided only qual­
i t a t i v e data concerning plant communities (Stevens et a l . 1978).
The
extent to which data from studies of s im ila r floodplain vegetation in
other lo c a l it ie s ( E v e ritt 1968, Hosner and Minckler 1963, Johnson et a l .
1976, Keammerer et a l . 1975, Ware 1949, Weaver 1960, Wikum and Wali
1974, Wilson 1970) could be applied to the lower Yellowstone River
floodplain required determination.
Thq objectives o f th is study were to id e n tify major rip a ria n plant
community types, q u a lit a t iv e ly and q u a n tita tiv e ly describe them, and
describe t h e ir successional relatio n s h ip s .
Community type descriptions
were to include stand age, species l i s t s with coverage estimates, woody
plant d ensities, and plant and soil biomass and nu trien t mass.
The re ­
sults of this study should be useful in the management o f r iv e r flow,
w i l d l i f e and a g ric u ltu re on the flo o d p la in .
Funding was provided by the Montana Department of Fish, W ild life
and Parks under Federal Aid in W ild lif e Restoration, Montana Project
W-12Q-R- I^ and 13 and to a lesser extent by a Montana State University
Faculty C re a tiv ity Grant (#2-6000-690).
2
DESCRIPTION OF STUDY AREA
The Yellowstone River originates, in northwestern Wyoming and flows
north into Montana.
I t turns east and crosses the southern portion of
Montana (Figure I ) , then joins the Missouri River in western North
Dakota.
All major t r ib u ta r ie s of the Yellowstone River have t h e ir
origins in Wyoming and flow north.
The t r ib u t a r ie s , in the order in
which they enter the Yellowstone, are the Clark Fork o f the Yellowstone
River, the Bighorn River, the Tongue River and the Powder River. The
P
to ta l drainage area o f approximately 179,000 km is nearly equally
divided between Montana and Wyoming, with less than 1% lying in North
Dakota (Stevens et a l . 1978).
The Y ellow tail dam on the Bighorn River
forms the only major reservoir in the Yellowstone drainage system.
Study sites were located on a 72 km stretch of floodplain between
Glendive and Sidney, Montana (Figure I ) .
are lis t e d in
Table 5.
Specific study s it e locations
Elevations w ithin the study area varied from
625 m above sea level a t Glendive to 577 m a t Sidney.
The Yellowstone River's main channel length in the study area is
101 km.
Streamflow measurements obtained from a gauging station near
Sidney (USGS Water Resources Data for Montana 1910 to 1980) show that
the mean discharge is highest during the months of May, June and July
and decreases dram atically between August and April
(Figure 2 ) .
age maximum discharge also peaks in May, June and July.
Ayer^
The highest
STUDY LOCATION
COUNTY LINE
RIVER
ROAD
TOWN
. IO MILES ,
DAWSON CO.
LENDIVE
Figure I .
Map of the study area showing locations of
s pecific study s ite s .
2400
///
Figure 2.
Mean, maximum and minimum Yellowstone River flow (nr/sec)
at Sidney, MT, in the 1910-1980 period (USGS, 1910-1980).
5
flow ever recorded was 4,502 m3/s on June 2, 1921 and lowest flow was
13 m3/s on April 17, 1961.
The lower Yellowstone Valley lie s along the western margin of the
W illis to n Basin, a "broad shelf-low r e l i e f '1 geologic feature (A lden
1932).
Marine sediments were deposited in the W illis to n Basin during
the Paleozoic, Mesozoic
and early T e r tia r y times.
The Fort Union
Formation, the la t e s t sediment la y e r, was formed during the Paleocene
Eppch (Veseth and Montagne 1980) and consists of soft nopmarine floodplain pediments derived from erosion o f western Montana during the
Laramide orogeny.
During the Pleistocene continental g la c ia tio n , a
lobe o f glac ial ice blocked the River v a lle y a t Intake, impounding the
River flow to form a glac ial lake with re su ltant deposition o f fin e
sediments upstream (A lden 1932).
The landscape of the W illis t e n Basin now consists p f gently r o l l r
ing h i l l s , wide v a lle y s , and f l a t divides with sandstone and c lin ker
beds forming ridges and buttes (Veseth and Montagne 1980).
Erosion by
the Yellowstone River has formed a broad f l a t floodplain with an aver­
age width of 4.4 km in the study a rea .
distance downstream.
The floodplain broadens with
The narrowest segment in the study area was 2.4
km at Glendive, the widest was 8.0 km a t Elk Is la n d .
Soils on the floodplain and low terraces are of the TremblesHavrelon-Lohler association.
This association is described by Pescado
and Brockmann (1980) as "deep, nearly level and gently sloping, well
drained and moderately well drained fin e sandy loams, s i l t loams, s i l t y
clay loams, and clays underlain by s t r a t i f i e d fin e sandy Ipam to s i l t y
6
ql^y alluvium". This soil association forms a substrate fo r the plant
communities id e n tifie d below.
'
In the Riyer, i n i t i a l sedimentation leading to the formation of
permanent land normally occurs with the formation of point bars.
These
o rig inate on convex curves within the confines of the r iv e r channel.
The opposing concave bank is c ut, providing sediment fo r deposition on
qonvex curves downstream ( Matthes 1941).
l a t e r a l l y across the flo o d p la in .
The channel thus meanders
The channel width remains constant as
vegetation colonizes the point bars and islands.
Surface heights of
pew deposits ris e as sediments are deposited on them by waters flowing
over theip a t high-water times.:
The lower Yellowstone River passes through a region which has a
semi-arid climate (Thornwai t e 1941).
Climatic records f o r Glendive
i l l u s t r a t e the p re c ip ita tio n and temperature fluctuations o f the region
(Figure 3, Ruffner 1971).
Average monthly p re c ip ita tio n is highest in
early summers with 8.1 cm f a l l i n g in June, and drops during winters to
0,8 cm in December.
The average annual p re c ip ita tio n is 30 to 35 cm.
Temperatures flu ctu ate about an average monthly high of 23.7 degrees C
in July to an average monthly low of minus 7.4 degrees C in February.
The present floodplain is occupied mainly by deciduous forests,
grasslands and cultiva te d land.
Immediately adjacent to the River,
g a lix spp. and Populus deltpides Marsh, (willow and cottonwood) seed­
lings and thickets are dominant.
Mature Populus deltpides stands,
grasslands (p r in c ip a lly Agropyron sm ithii Rydb.)
and shrublands ( Rosa
woodsii L in d l. and Symphoricarpos occidental is Hook.) are scattered
throughout the flo odplain.
Fraxinus pennsylvanica Marsh. (Green Ash)
AVERAGE PRECIPITATION CM
AVE. PPTN.
AVE • TEMP.
F M A M
S
O
N
MONTH
Figure 3.
The average monthly temperature and p re c ip ita tio n at Glendive, MT (Ruffner 1971).
The v e rtic a l scales are adjusted so evapotranspiration exceeds precipitation
whenever the temperature lin e rises above the p re c ip ita tio n lin e (Walter 1973).
8
stands are commonly found at the edge o f the flo odplain.
whiph also ocpur here, were not s tudied.
Marshes,
The semi-arid climate domin­
ates floodplain vegetation where deep soil deposits l i f t the soil sur­
face well above the water ta b le , but ground water and flooding have 3
s ig n ific a n t e ffe c t where soil deposits are shallow.
Man's impact on the floodplain vegetation stems mainly from a g r i­
c ultu ral use.
Many once-forested lands, shrublands
have been cleared fo r c u lt iv a t io n .
and grasslands
Most land not c u ltiva te d is used by
sheep or c a t t l e .
I
9
MATERIALS AND METHODS
Reconnaissance during the early Summer of 1980 id e n t if ie d 12 mejor
physiognomic plant community types.
Seventy sites representing the
pi^'Pr communities were selected fo r a n a ly s is .
These were: 19 in the
"Sandbar" type, 9 "seedling" type, 8 "thicket" type, 9 "young cotton­
wood" type, $ "mature cottonwood" type, 9 "shrub" type, 9 "grassland"
type, 7 "cireen ash" type, 3 "willow-shrub" type and I "peach-leaved
willow" type.
A species l i s t
"rparsh" community types.
(Table 6 )
was also made fo r three
Stand selection c r i t e r i a were th a t each s ite
represented one of the major community types indicated above, and that
the sites showed no evidence of disturbance by f i r e , logging, recent
gfdzing or other a g ric u ltu ra l use.
Measurements were taken to charac­
t e r iz e each community type with respect to species composition, age,
soil height, stand s t r a t i f i c a t i o n and height, community composition
(qover and/or d e n s ity ), biomass and n u trie n t masses.
Species Composition
rr-l
”
1------------------
Complete plant species l i s t s were kept for a l l s it e s .
Species ep-*
countered in sites not sampled, including marshes, were separately
recorded,
S c ie n tific names of plants follow Booth (1972) and Booth and
Wright (1966) o r, fo r species not lis t e d by Booth, Van Bruggen (1976).
I
10
Site Age
The age o f forested sites was estimated by counting the annual
rings o f the largest Populus deltoides present.
Growth rings were
observed by cutting seedlings and saplings, or coring older trees at
breast h e ig h t.
Four years were added to each core count to compensate
for the time the tree took to grow to breast height (Wilson 1970).
Stand Height
The height o f the soil surface above the River water level was
determined with a level and measuring tape.
Since spil elevations
were recorded a t d iffe r e n t times o f the summer and since the River
level varied throughout the summer, soil elevations are only approxi­
mate indicators o f true height p f the soil above the ^iyer le v e l.
Plant Cpver
Horizental cover was recprded fo r understory, shrub and overstory
plants op a transect crossing the community perpendicular to the River.
Graminpids, forbs, Artemisia ludeviciana N u t t ., Tpxicodendron rydbergii
(Small) Greene, and vines were sampled with s ix ty step-points and
presented as percent cover (Evans and Love 1957).
Shrub cover was
measured by c alculatin g canopy coverage ( p i . r 2 ) of each shrub present
in density plpfs (described below), summing and expressing the shrub
coverage as a percent of the tptal plpt area.
ages were based upon ocular estimates.
Tree and sapling cover­
11
Density of Moody Plants
Plots to sample the density of tree and willow species were located
a t three equally-spaced points on the cover transect.
The number of
trees and willows o f each species was recorded in each p lo t.
Seed­
lin g s , saplings and trees were defined by basal diameter as fo llo w s :
seedlings were less than or equal to 0.5 cm, saplings were less than or
equal to 6 cm, and trees were greater than 6 cm.
Basal diameter was
recorded fo r a l l saplings and the diameter a t breast height (dbh) was
recorded fo r a l l tree s .
Plot sizes used in various communities were
as follows; seedling community I x 0.5 m; th ic k e t community 2 % 3 m;
young cqttonwoqd community 5 x 5 m ; mature cottonwood community 20 x 20
m; shrub community I x 5 m; willow-shrub community 4 x 10 m; the green
ash and peach,leaved willow plots were 4 x 30 m.: T h irty meter ash,
willow plots were cut short i f 2Q trees were recorded before 4 x 30 m
was reached and the actual plot length was recorded.
Density plots fo r shrubs (except Artemisia ludoviciana, Toxico-
•
dendron rydbergij and vines) were also placed a t equidistant points .
along the transect.
The minimum size of each plot was I x 5 m an<i the
maximum size was 2 x 20 m, depending on which came f i r s t : 20 total
Shrubs or a maximum area of 40 square meters.
Each shrub's crown diam,
eter was recorded! in one-decimeter units as the average o f the widest
crgwn measure and the crown measure perpendicular to i t a t it s mid­
point.
Standing dead shrubs were not counted.
12
Aboveground Biomass
Biomass was measured only in communities representing the sandbar
through grassland sere.
Time lim ita tio n s prohibited measurements in
the willow-shrub, peach-leaved willpw and green ash communities.
Biomasses o f graminoids, forbs, Artemisia ludoviciana. Toxicoden­
dron rydbergii and vines were measured by harvest methods.
AU material
in fiv e I x 0.5 m plots equally spaced along the cover transect was
clipped a t ground le v e l, combined, dried to constant weight at 60 de­
grees C and weighed.
Biomasses of Populus deltoides and S alix spp. were
measured s im ila r ly in the seedling community.
Tree and shrub biomasses were estimated by summing the weights of
individual plants in each density p lo t.
Because only a lim ite d number
of shrubs and trees could be harvested, weights of trees were determined
from regressions r e la tin g weights o f plant parts to a plant dimension
(Whittaker and Woodwell 1968a).
The method is outlined in the followr
ing six paragraphs.
Large, medium and small specimens o f the shrubs Symphoricarpps
occidental i s , Rosa woodsii and Artemisia cana Nutt, were collected in
each o f three s it e s .
Crown diameter was measured and each shrub was
separated into leaves, wood (branches) less than 0.5 cm in diameter
and wood greater than 0.5 cm in diameter.
These components were dried
and weighed and t h e ir weights were regressed against crown diametgr
(Weaver 1977).
Nine specimens each o f Salix f l u v i a t i l i s Nutt, and Sa lix amygdaloides Anders, were collected from, three locations.
The basal diameter
13
of each Salix individual was recorded.
Sa lix spp. were separated into
leaves, wopql less than I cm in diameter and wood greater than I cm in
diameter.
Each component was then dried and weighed.
Subsamples o f .
the la rg e r specimens were used to determine the dry weight/wet weight
ratios needed to convert wet weights to dry weights.
S im ila rly , f iv e small Populus deltoides from the seedling and
th ic k e t communities were measured fo r basal diameter and the dbh of
f iv e Populus deltoides collected from the young cottonwood and mature
cottonwood communities were measured.
A ll specimens were separated
into leaves, wood less than I cm in diameter, wood greater than I cm
and less than 10 cm, wood greater than 10 cm in diameter, and weighed
while wet.
Subsamples were used to determine dry weight.
Dry weights
wpre regressed against diameters,
For each species, the logarithm o f dry weight wap regressed
against the logarithm o f diameter to create the graphs needed to e s t i ­
mate weights from an e a s ily measured diameter (Whittaker and Woodwell
1968a).
The regression parameters used to predict plant part masses
are presented in Table 25
according to the formula y=mx+b where
y=loglO o f the part weight in grams, b=the y in te rc e p t, m=the change
in mass with changing diameter and X=IoglO of the plant diameter in cm.
Plant diameter was canopy diameter for shrubs, basal trunk diameter fo r
Salix spp. and small (0-6 cm basal diameter) Populus d e lto id e s , and
diameter at breast height fo r Populus deltoides la rger than 6 cm basal
diameter.
To measure the closeness o f f i t between two variables along a
regression l i n e , the c orrelation c o e ffic ie n t ( r , Snedecor and Cochran
14
1980) and the estimate o f r e la t iv e error for a logarithmic regression
(E, Whittaker and Woodwell 1968a) were calculated (Table 25).
Total biomass estimates of trees and shrubs in communities older
than the th ic k e t community are underestimated s lig h t ly because in f r e ­
quently encountered species such as Juniperus scopulorum Sarg.,
Elaeagnus a n q u s tifo lia L . , Fraxinus pennsylvanica, Ribes spp. and
Cornus sto lo n ife ra Michx. were not included.
Aboveground Qead Biomass
L i t t e r mass was measured by harvesting three I x 0.5 m plots
equally spaced along the cover tran sect, drying and weighing.
L itte r
included a ll dead organic m atter, including leaves and dead wood less
than 10 cm in diameter lying on the ground or standing in the plot and
reasonably d is t in c t from the underlying s o i l ,
Weights o f dead Populus d e lto id e s , whether standing or lying on
the ground, were estimated by m ultiplying t h e ir pre-death weights,
estimated with the weight-diameter regressions determined as described
above, by a fa c to r correcting fo r loss due to decomposition and sub­
trac tin g weights of branches apparently lo s t to the l i t t e r component
described above.
This method is elaborated below.
To correct fo r decomposition, the dead trees were assigned to one
o f three categories: ro tte n , h a lf-r o tte n or s o lid .
The wood o f rotten
trees was weathered, rotted and could be kicked 9part; h a lf-r o tte n
trees exhibited a t le a s t some sign of ro t but the woqd could not be
kicked apart; solid trees showed no outward evidence o f r o t .
Three to
f iv e samples fo r the 1-10 cm and greater than 10 cm diameter size
15
classes were collected from the ro tte n , h a lf-r o tte n and so lid categpr?
ie s .
The samples were dried and weighed, t h e ir volumes were determined
and wood density (gm/cc) was calculated.
each category was calculated.
A mean for a ll the samples in.
The r a tio o f ro tte n , h a lf-r o tte n or
solid wopd cjensity to l i v e wood density provided a fa c to r used to
correct fo r decomposition.
Tp correct fo r loss of branches, dead trees were recorded as
branchless (only the bole o f the tree present), hal^branched (the bole
plus about h a lf of the branches present) or fully-branched (the bole
and most o f the crown present).
Weights o f trees in the branchless
category were calculated as wood greater than 10 cm in diameter from
the Populus deltoides regression lin e s .
Weights of trees in the h a lf-
branched category included the to ta l o f wood greater than 10 cm plus
one-half o f the 1-10 cm size class.
Weights of trees in the f u l l y r
branched category included a l l wood la rg e r than I cm in diameter.
If
the dbh o f the tree was less than I dm, f u l l y branched included a l l the
1-10 cm diameter wood, half-branched equaled three-fourths of the IrlO
cm wood and branchless equaled one h a lf o f the 1-10 cm wood.
The weights of individual dead Populus deltoides in the density
plots were then determined by f i r s t calculating the weight o f each dead
tree as i f i t were a liv e by using the Populus deltoides tree regres­
sions.
Depending on which category the tree was recorded in - b r a n c h ­
less, half-branched or fully-branched.— some weight o f the tree was
subtracted from the overall weight.
The weight of each tfe e was then
m u ltip lied by the appropriate dead w e ig h t/liv e weight r a tio to convert
16
these data to ro tte n ,, h a lf-r o tte n or s olid weights.
A ll dead tree
weights were totaled fo r each p lo t.
Non-woody vegetation samples were dried in v e n tila te d ovens (60 C)
fo r two wpeks, then weighed to the nearest gram.
All woody biomaps
collected was weighed wet in the f i e l d to the nearest gram, or poynd
fo r large samples, then dried in v e n tila te d ovens (60 C) fo r four weeks
and reweighed.
Dry weight/wet weight ratios were calculated to convert
f i e l d weights of wet wood to dry weights.
Belowground Biomass
Belowground biomass consisted of four categories: root crowns and
large roots greater than I cm, roots I cm to 0.1 cm in diameter, f ip e r
roots and soil organic matter.
Due to pre-analysis grinding, soil
organic matter included roots less than I cm in diameter.
Weights of large roots were measured by dimension analysis
(Whittaker and Woodwell 1968a).
Root systems o f six Populus deltpides
treps were p ither excavated or found pre-washed on gravel bars.
Roots
of each individual tree were separated into I to 10 cm and 10 cm or
greater diameter classes, dried and weighed.
Wherever systems were
too large to dry, wet weights were converted to dry weights by m u lti­
plying by dry weight/wet weight ratios calculated from subsamples.
logarithm weight-logarithm diameter regression
pared.
(Table 25)
A
was pre­
Weights of roots greater than I cm diameter of a l l trees in
each stand were estimated in a fashion p a ra lle lin g the calculation of
shoot biomasses.
17
Biomass o f roots less than I cm in diameter, soil organic matter
and soil n u trien t concentrations were estimated from soil cores in the
0-10, 10-30 and 30-150 cm horizons.
Five cores equally spaced along
the cover transect were pooled fo r each horizon sampled.
When the king
tube sampler could not be driven beyond 120 cm, comparable samples were
removed from the sides o f a soil p i t .
Each sample was dried a t 60
degrees C, mixed, weighed and subsampled (25% by weight) fo r laboratory
analysis of elements and organic material present,
Biomasses of small roots (less than I cm in diameter) in the 0-1Q,
10-30 and 30-150 cm soil horizons were estimated by weighing, on an
ash-fr^e basis, volumetric samples washed from soil cores and expanding
th a t value appropriately.
Roots la rg e r than I cm were discarded since
they were separately estimated by dimension a n a ly s is .
Roots and d e t r i ­
tus were washed from the samples following the methods o f Uackspn
(1956), except that s o i l s were not p re sifted with a 6 mm screen.
Rpots,
d e tritu s and soil remaining on the washing screen were placed on pre­
weighed Whatman 42 ashless f i l t e r paper.
Roots, s o il , d e tritu s and
f i l t e r paper were dried a t 60 degrees C fo r two days.
Roots and
d e tritu s were separated into size classes less than or equal to I mm
and greater than I mm in diameter.
Because roots and d e tritu s greater
than I mm in diameter were ra re , they were combined across a l l stands
o f a type to calculate average biomass.
remaining was ocularly estimated.
ashed and reweighed.
Percent root in the sample
The e n tire sample was then weighed,
Ash-free root biomass was calculated as (organic
matter + f i l t e r paper + soil traces - f i l t e r paper - ash) times the
percentage of root in the sample (Weaver 1982).
18
The organic matter content of soils acquired, in volumetric cores
was calculated as percent organic matter (gm/100 gm) times soil bulk
density times volume o f the soil layer considered.
Organic matter
contents were c o lo rim e tric a lly determined by the Montana State Univer­
s it y Soil Testing Laboratory a f t e r dichromgte oxidation (Sims and Haby
1970).
Soil organic matter was estimated as total organic matter minus
small root ( 0-1 cm diameter roots) organic matter.
System Nutrient Mass
Plant parts, l i t t e r and soils were analyzed for nitrogen, phos^
phorus, organic phosphorus o f s o i l , potassium and sodium by the Mpntana
State University Soil Testing Laboratory.
The analytical results are
summarized in Appendices K, M, N, 0, P and Q.
Only roots greater than
I cm in diameter were removed from the soil samples, thus soil analysis
also includes roots less than or equal to I cm in diameter.
Soil nu trien t masses in each horizon were estimated by m u ltip ly ­
ing nu trien t concentrations (gm/100 gm) by soil bulk density by the
volume o f soil in the horizon concerned.
data are summarized in
Table 26.
Stand-rby-stand bulk density
L i t t e r nutrient masses were deter­
mined by multiplying n u trien t concentrations by l i t t e r weight.
Nutrient masses o f plant parts were estimated by m ultiplying the
n u trien t concentration by plant part biomasses.
Neither roots nor dead
Populus deltoides were analyzed fo r n u trie n t contents (%).
To adjust
fo r t h is , fin e root contents ( 0-1 cm) were assumed to equal twig
(0-1 cm) contents of Populus deltoides and coarse root contents ( I cm+)
were assumed equal to those o f 1-10 cm wood.
S im ila rly , n u trien t
19
content {%) in l i v e Populus deltoides wood greater than 10 cm in diam­
eter was also used as an estimate o f the n u trien t content o f dead tr e e s .
Total nitrogen in both soils and plant material was measured by
the Kjeldahl method (Bremner 1965).
Potassium, sodium and phosphorus
contents o f plant material were measured by ashing samples and deter­
mining the quantities o f elements released by spectrophotometric (P) or
atomic absorption (K and Na) methods.
Potassium and sodium were ex­
tracted from soils with I M ammonium acetate and measured by atomic;
absorption (P ra tt 1965).
Inorganic phosphorus was extracted with a I M
s u lfu ric acid solution and measured spectrophotometrically (Olson and
Dean 1965).
Total soil phosphorus was determined by ashing, extracting
with I M HgSO^ and reading spectrophotometrically (01 son and Dean 1965).
Organic phosphorus was assumed to be to ta l phosphorus minus inorganic
phosphorus (Saunders and Williams 1955).
20
RESULTS AND DISCUSSION
Riparian communities o f the Yellowstone River floodplain were
re a d ily distinguishable.
These ranged from bare sandbars to willow
th ick e ts , to cottonwood forests and green ash forests, to grasslands.
Data from my studies are used below to characterize these communities
by composition, describe t h e ir serai re la tio n s h ip s , speculate on forces
driving the serai changes and measure changes in ecosystem contents of
organic m atter, nitrogen, phosphorus, potassium and sodium.
Figures 4 and 5 summarize the successional re lations o f the 12
'
plant communities which appear in the four seres id e n t if ie d , and which
w ill be described below.
The sere leading to the regional dry grass­
land climax dominates the flo odplain.
I t began with Populus deltoides
and Sglix spp. seedlings colonizing newly-formed River sandbars to ­
gether.
The Salix spp. i n i t i a l l y attained stature exceeding Populus
deltoides seedlings, but they died out a f t e r about 20 y e ars .. Populus
deltoides then assumed dominance but also disappeared a f t e r about IQO
years because dying trees were not replaced by seedlings.
Disappear­
ance of Populus deltoidgs allowed a shrub community to invade and,
eventually, a grassland community to assume dominance.
The green ash sere shared the sandbar through mature cottonwood
communities (Figure 4) with the grassland sere but apparently has a
green ash community climax.
Fraxinus pennsylvanica f i r s t appeared in
the young cottonwood community and s te a d ily became more common up
GREEN ASH
GRASSLAND
WILLOW SHRUB
PEACH LEAVED
WILLOW
SHRUB
MATURE COTTONWOOD
FRPE
A
YOUNG COTTONWOOD
FRPE
T H IC K E T
SEEDLING
SALIX >
SEEDLING
PODE SALIX
SANDBAR
Figure 4.
MARSH
Flow chart summarizing successional changes observed in the
lower Yellowstone River flo odplain. Four l e t t e r s c ie n t if ic
name abbreviations are Fraxinus pennsylvanica (FRPE) and
Populus d e ltoides (PODET
-------------
RIVER
3I
H
|
YC1
40
60
MC
80 100
AGE (YRS)
Figure 5
V
w-s I ga
Iplw
23
through the shrub community.
When the Populus deltoides trees reached
senescence, Fraxinus pehnsylvanica trees invaded, apparently to persist
in d e f in it e ly on sites usually found near the flo odplain's edge.
The peach-leaved willow sere appeared to begin when S alix spp.
established without Populus deltoides on newly-formed sandbars (Figure
4).
Salix amygdaloides developed into stands o f trees which would
probably be replaced by a green ash community since Fraxinus Pennsyl­
v a n ia seedlings appear in the understory.
Although the marsh vegetation was not analyzed, a species l i s t was
compiled
(Table 17).
In the very long term, accumulating soil
would presumably allow a green ash or grassland community to establish
on these s ite s .
Grassland Sere Community Descriptions
The grassland sere began with the formation of sandbars along the
River.
The soil surface was a sand containing some gravel and debris
l e f t by high water (Appendix F . l ) \
Soil height averaged 0.8 m above
the River water level fo r new deposits (Figure 6 ).
Populus deltoides and Salix f l u v i a t i l i s ty p ic a lly were the f i r s t
plants to establish and the dominant species of the seedling community;
t h e ir principal associates are lis t e d in Table I .
this community averaged 3 dm (Figure 7 ) .
spp. cover averaged 27%.
The canopy height of
Populus deltoides and Salix
Forb and graminoid cpver averaged 8% and
formed only a minor component o f to ta l cover in any of the communities
to be described except grassland (Table 2 ) .
cover was 61% sand and 4% l i t t e r .
The remaining soil surface
Soil height averaged 1.3 m (Figure 6 ) .
24
H
X
LU
O
<
LL
CC
3
CO
20
Figure 6.
40
60 80
100
AGE (YRS)
The elevations of the soil surface above the Yellowstone
River surface for communities of increasing age. Bar
segments represent community types: sandbar (SA), seedling
(SE), th ick e t (TH), young cottonwood (YC), mature cotton­
wood (MG), shrub (SH) and grassland (GR).
25
TABLE I
E W F IP
Community types
TH
8
X±SE4 X±SE
2±1
SE2
Plant species
YC
9
X+SE
-
MC
6'
X+SE
-
SH
9
XtSE
-
GR
9■
XtSE
W-S
3
XtSE
GA
7
X+SE
Polygonum coccineum
(F )5
Echinochloa crusgalli
(G)
1±1
+
-
-
-
-
_
Rumex maritimus
(F)
1+0
+
-
-
-
-
_
_
_
Salix f lu v ia t ilis
(Se)
5±2
Salix f lu v ia t il is
(Tr)
-
Salix amyqdaIoides
(Se)
1+1
PLW
I
X+SE
_
-
-
-
-
-
_
30
-
-
-
-
_
.
-
-
_
_
5
+
+
-
-
20+3
_
-
-
-
-
-
-
40±4
+
-
-
+
1±1
+
+
1+0
+
_
Salixamygdaloides
(Tr)
-
Populus deltoides
(Se)
21±3
-
Populus deltoides
(Tr)
Melilotus o ffic in a lis
(F)
+
l±o
66±4
+
Aqropyron repens
(G)
-
3±1
5+2
1±1
+
1+0
3±3
Elymus canadensis
(G)
+
2±1
7+1
6±2
2±1
2±2 ,
3+1
5*2
-
Muhlenberqia racemosa
(G)
-
2±1
7±2
5±2
4±2
3±2
3±2
-
Glycyrrhiza lepidota
(F)
+
+
2+0
1±0
1±1
4+4
-
+
2±1
2±2
1±1
+
1+1
+■
2+2
12±8
12±10
Bromus Inermis
(G)-
30
. 4±2
60
_
-
2
Toxicodendron rydberqii
(S)
-
-
+
12+3
3+1
+
2+1
1±1
23
+
V itis rip a ria
(V)
-
. -
+
4+2
-
-
+
+
Ribes aureum
(S)
-
-
+
+
-
1±1
-
+
—
+
+
1+1
2±2
-
+
-
+
1±1
-
Parthenoclssus quinquefolia (V)
-
-
+
+
Ribes setosum
(S)
-
-
-
Rosa woods1i
(S)
-
-
+
12±3
Fraxinus pennsvlvanica
(Se)
-
-
+
+ '
Fraxinus Pennsylvania
(Tr)
1±0
-
14±2
+
+
-
11+5
3±3
+
+
9
+
-
-
-
+
+
Symphoricarpos occidental is (S)
-
-
+
6±2
8+1
Ulrnus americana
(Se)
-
-
-
-
-
-
-
+
_
Ulmus americana
(Tr)
-
-
-
-
-
_
-
' +
.
-
-
+
+
+
Medicaqo lupulina
(F)
-
Artemisia ludoviciana
(S)
-
-
+'
-
6±2
58±10
_
7+4
I
-
-
2+1
-
+
2±i
1+1
+
_
,Artemisia cana
(S)
-
-
-
-
-
1+0
-
_
_
Calamovil fa lo n q ifo lia
(G)
-
+
+
-
-
11+4
-
-
-
Aqropyron smithii
(G)
1±0
1+0
1+1
3±1
17+5
-
iA common species is one which occurred in over 6.056 o f the stands of any community type.
Community types are seedlings (SE), thicket (TH), young cottonwood (YC), mature cottonwood
(MC), shrub (SM), grassland (GR), willow-shrub (W-S), green ash (GA) and peach-leaved
,willow (PLW).
^The number_of stands of each community type sampled.
cThe mean (X) and standard error (SE) o f each species' coverage.
0Species habits are forb (F ), grass (G ), vine (V ), shrub (S ), tree seedling (Se) and tree ( T r ) .
26
hX
O
Z
<
CZD
6 0
AGE
Figure 7.
8 0
100
(YRS)
Height of the upper canopy surface, r e la t iv e to the ground
surface, in communities o f increasing age. Bar segments
represent community types: sandbar (SA), seedling (SE),
thicket (TH), young cottonwood (YC), mature cottonwood (MG),
shrub (SH) and grassland (GR).
TABLE 2.
Plant coverage, species richness and cummunity height in nine community types o f the
lower Yellowstone River f lo o d p la in . This table summarizes data found in Appendices
F . l through F.8 and E.
Community types1
XiSE2
COVER (%)
N3
Herb
9
8±2
TH
XiSE
8
28i6
YC
XiSE
9
MC
XiSE
6
SH
XiSE
9
GR
XiSE
9
W-S
XiSE
3
GA
XiSE
7
J 3-LW
XiSE
I
37i3
39i5
29+6
67+3
4 4 ill
41i7
57
23i2
2i0
15+5
23i3
32
+
-
20+3
5 8 il0
60
Shrub^
27+5
+
IiO
19i4
Canopy
-
625
66i4
40+4
9
8
9
6
9
9
3
7
I
T o ta l5
32
53
36
32
35
56
30
63
17
20%7
17
26
25
21
20
28
30
36
-
60%8
12
14
15
18
17
13
15
21
-
9
8
9
6
6
2
7
SPECIES RICHNESS
N
N
Community height (m)
0.3+0.2
2i0
1 9 il
25i2
IiO
-
-
2 il
13+2
^Community types are seedling (SE), th ic k e t (TH), young cottonwood (YC), mature cottonwood
(MG), shrub (SH), grassland (GR), willow-shrub (W-S), green ash (GA) and peach-leaved
2wi11ow ( PLW).
gThe mean (X) and standard error (SE) o f each species coverage.
^The number of stands o f each community type sampled.
gShrub (%) coverage includes seedlings o f a ll tree and Salix spp.
gThicket canopy coverage was estimated a f t e r the f a c t.
^Species richness includes the to ta l number o f plant species recorded in each stage.
LThe number o f species occurring in a t le a s t 20% of the stands sampled.
tiThe number of species occurring in a t le a s t 60% o f the stands sampled.
-
_
28
In t h e .th ic k e t community, Salix spp. and Pop,ulus deltoides con­
tinued to t?e the dominant species.
(Figure 7 ) .
Canopy height increased to 2.2 m
Populus deltoides and S alix spp. cover averaged 65% and
herb cover increased to 28% (Table 2 ) .
Only.24% of the soil surface
was covered by sand due to increased l i t t e r cover.
Soil height in ­
creased to an average o f 2.5 m (Figure 6 ) .
The young cottonwood community was dominated by Populus deltoides
tree s , since most S alix plants had died out (Table I ) .
The canopy '
height (Figure 7) t r ip le d over the thicket.community and canopy cover
averaged 66% (Table 2 ) .
The understory remained sparse, with a com­
bined herb and shrub cover o f 37% (Table 2 ) .
The soil surface was
mostly covered (66%) by l i t t e r and in a ll l a t e r communities i t covered
the m ajority o f the soil surface not covered by vegetation.
Soil
height increased s u b s ta n tia lly over the th ic k e t community to 3.1 m
(Figure 6 ) .
The mature cottonwood community was characterized by widely-spaced
towering Populus deltoides and a dense shrub-herb understory.
Canopy
height o f Populus deltoides increased to an average o f 24.6 m and the
canopy cover f e l l to an average o f 40% (Table 2 ).
Trees commonly con­
tained de^d branches and large dead trees were frequently encountered.
Shrub cover increased to 19% and herb cover was 39% (Table 2 ) .
Soil
height increased markedly, to 3.8 m (Figure 6 ).
In the shrub community, Rosa woodsi i and Symphoricarpos occident­
al is dominated,
No trees were present in the stands sampled although
seedlings and/or saplings of Fraxinus pennsylvanica were present at 5
o f the 9 sites
(Table 14).
The shrub height averaged 1.3 m and
29
shrub cover averaged 23% (Table 2 ).
While few dead shrubs were ob­
served in the mature cottonwood community, ocular estimates indicated
th a t 20% to 50% of the shrubs in the shrub community were dead (Table
T4).
Standing dead shrubs were not included in e ith e r density or
coverage c a lc u la tio n s , consequently density and percent cover do
not f u l l y characterize the prevailing dense shrub aspect.
Herbaceous
cover decreased to 29% (Table 2) and the soil height decreased s lig h t ly
to 3..4 m (Figure 6 ).
Agropyron sm ithii and Calmovilfa lo n g ifo lia (Hook) Scribn. domin­
ate the grassland community (Table I ) , where the average graminoid and
forb cover was 67% as compared with a range of 8% to 39% in the e a r lie r
communities (Table 2 ) .
Artemisia cana Nutt, had the highest shrub per­
cent cover a t 1% (Table I ) .
The average soil height dropped to 3.3 m.
The grassland community apparently represents the clim a tic climax
(Clements 1936).
That is , i f a ll disturbances were removed from the
floodplain ( e . g . , . erosion, flo odin g), vegetation s im ila r to th a t occur­
ring on adjacent p r a ir ie ( Kuchler 1964) would eventually develop.
Green Ash Sere Community Descriptions
In the green ash sere the sandbars
through young cottonwood com­
munities were sim ilar to th a t of the grassland sere.
Fraxinus pennsyl-
vanica seedlings and saplings appeared, however, in the young cotton­
wood community and small trees were occasionally recorded in the mature
cottonwood community.
Thus, I speculate that the Fraxinus pennsylvanica
seedlings would develop under the Pop.ulus deltoides fo re s t canopy and
u ltim ate ly replace dying Populus d e lto id e s , ; Stands■characterizing the
30
green ash community were dominated by Fraxinus pennsylvanica tr e e s ,
with an average height (13.3 m) h a lf that of a mature cottonwood forest
and canopy coverage (.58%) s im ila r to a vigorous cottonwood stand (Table
2).
They also contained tr e e s , saplings and seedlings o f Ulmus ameri-
cana L. and Acer negUndo L.
Regeneration o f a l l three species was
patchy, i . e . , where any seedlings occurred there were many.
Elymus
canadensis L . , Muhlenbergia racemosa (M ichx.) BSP., Rosa woodsi i and
Symphoricarpos occidental is were the most common understory associates
(Table I ) .
The soil height averaged 4.3 m, the highest elevation
recorded fo r any community.
The willow-shrub community may be an intermediate between the
cottonwood community and green ash community in ephemeral stream
channel? or fla t!a n d adjacent to the River.
Rosa woodsii, Symphori-
carpos occidental is and Sa lix amygdaloides are the dominant species
(Table I ) .
In the ephemeral stream channels, Salix amygdaloides nor-
mally occurs on the banks with Rosa woodsii and Symphoricarpos occident­
al is in the channel bed.
Where the community occurred on level land
near the River, these shrubs were intermixed.
willow-shrub sites averaged 2.1 m (Table 2 ) .
and herb cover 44% (Table 2 ) .
The canopy height of
Shrub cover averaged 15%
Soil height averaged 3.9 m.
Peach-leaved Willow Community Description
In the peach-leaved willow stand sampled, Rosa woodsii and Bromus
inermis both had high cover values and were the dominant understory
species (Table I ) .
I saw no Salix amygdaloides regeneration, but
31
Fraxinus pennsylvanica seedlings were common (Table 2 ) .
The stands
were t y p ic a lly long and narrow.
Evidence fo r Successional Relations
The serai scheme described above was supported by observations
Of Populus deltoides ages, soil heights above the River water and over­
laps Of community composition.
The most solid evidence fo r the serai
scheme was provided by the annual rings of Populus deltoides trees in
the various community types in which they occurred: average tree ages
increased from 3 to 7 to 34 to 92 years in the seedling, thicket,"
young cottonwood and mature cottonwood communities, respectively (Table
3).
Unfortunately, this chronology cannot be extended to other com­
munity type$.
The rapid increase and then le v e lin g Of soil height through suc­
cession provided supporting evidence (Figure 6 ).
Soil height would be
expected to increase rapidly in e arly succession due to accumulation of
sand and s i l t during flooding.
Conversely, flooding o f older communi­
tie s would occur less frequently as soil height increased and less
s i l t would be deposited.
My data indicated that soil height increased
s te a d ily in early succession from an average sandbar height of 0.8 m to
3.8 m ip the mature cottonwood community (Figure 6 ) , a to ta l change of
3.0 meters.
In older communities, average ground level varied from a
low of 3.3 m in the grassland community to a high o f 4.3 m in the
green ash community, a difference o f only 1.0 m, r e fle c tin g a r e la t iv e ­
ly stable soil l e v e l .
TABLE 3.
Mean age and diameter of J3Opulus deitoides trees found in four
community types. This table ,summarizes data found in Appendixes
C and E.
Community types
I
N1
Diameter (cm)
XiSE
XiSE
XiSE
9
3.2±0.2
3
Mature
Cottonwood
5
Thicket
XiSE2
Age (years)
8
Young
Cottonwood
6
Seedling
<1.0
7 . It O - 7
3 4 .1 i3 .3
92.2+5.1
4 .3 i0 .7
2 3 . 2 i l .4
6 4 .8 i4 .4
gThe numberjof stands of each community type sampled.
3The mean (X) and standard erro r (SE) of Populus deltoides age and diameter.
Basal diameter was recorded in the seedling and th ic k e t communities and
diameter a t breast height was recorded in .th e young cottonwood and mature
cottonwood communities.
33
Community description emphasized vegetational linkage o f success­
ive communities.
To r e i t e r a t e , Salix spp. and PopuTus deltoides seed­
lings established together, with S alix spp. decadence Populus deltoides
assumed dominance, and the decadence of Populus deltoides allowed
establishment of a shrub community, followed by e ith e r a grassland or
a green ash community.
The peach-leaved willow sere began when Salix
amygdaloides established on sandbars and eventually developed into a
stand of peach-leaved willow trees.
Species Numbers
A complete l i s t o f vascular plant species in each community
appears in
Table 6.
The to ta l number of species found (Table 2) in ­
creased from the seedling community (32) to the th ic k e t community (5 3 ),
decreased to the mature cottonwood community (32) and then rose again
in grassland or green ash communities (5 6 -6 3 ).
In contrast, numbers of
plants with high constancy (60%+, Table 2) increased from early serai
communities (12-14) to young and mature cottonwood communities (19-18)
and then e ith e r f e l l to the grassland community (13) or continued up­
ward to the green ash community (21) (Table 2 ) .
Density o f Trees and Shrubs
Populus deltoides and S alix spp. established in vast numbers on
moist sandbars and died out rapidly through time (Table 4 ) .
Salix
f l u v i a t i l i s , which established a t densities of about nine seedlings per
square meter, maintained it s numbers through the th ic k e t community but
was e n t ir e ly absent at the beginning o f the young cottonwood community.
TABLE 4.
Density {number/100 hi ) of woody plant s:pecies an nine community types.
summarizes data 'found in Appendices G.l through G, 4.
Community types
N2
Plant species
Salix f l u v i a t i l i s
Salix f l u v i a t i l i s
Salix amygdalaides
Salix amygdaloides
Salix amygdaloides
Populus deltoides
Populus deltoides
Populus deltoides
Cornus stolo hife ra
Ribes au.reum
Symphoricarpos
occidental is
Rosa woodsi i
Ribes setosum
Ulmus americana
Ulmus americana
Fraxinus
pennsyl vam'ca
Fraxinus
pennsylvanica
Fraxinus
pennsylvanica
Artemisia cana
-SE
-9 3
X+SE
(S e)*
(Sa)
(S e)
(S a)
(Ir)
(Se)
(S a)
(Ir)
( s)
( s)
( s)
( s)
( s)
(Se)
(Tr)
(S e)
879±279
167±71
—
4815±787
-
TH
8
X±SE
YC
9
X±SE
MC
6
X±SE
S-H
9
X+SE
W-S
3
XlSE
GA
7
XiSE
PLW
I
XiSE
_
995±218
130+106
519+125
-
+
+
22±3
12±7
12±11
+
+
2±0
+
49+31
_
_
-
-
73±36
8±5
793±155
311165
_
+
18+8
_
_
_
16141264
601168
4+3
40114
1218
-
-
-
-
-
-
-
-
-
-
-
-
-
-
_
(Sa)
-
_
_
_
-
30114
5651281
3121119
+
_
.7
+
3+3
3l3
615+214
3261206
40+16
20+18
+
1+1
38+3
132195
73143
5±5
+
1811
3 ll
4+2
-
+
-
(Tr)
-
GR
. 9
XlSE
This -data
+
-
_
_
174
40
176
_
7
1112
47+20
-
-
Community types are seedling (SE), th ic k e t (TH), young cottonwood (YC), mature cottonwood (MG),
2shrub (SH), grassland (GR), willow-shrub (W-S), green ash (GA) and peach-leaved willow (PLW)
nThe number_of stands of each community type sampled.
^The mean (X) and standard error (SE) of each species' density.
The tree species are divided into three size classes: seedlings less than 0.5 cm basal diameter
(S e ), saplings less than 6 cm basal diameter (Sa) and trees greater than 6 cm basal diameter (Tr)
35
Populus d e lto id e s , which was i n i t i a l l y more numerous (.48 plants per
square meter) declined s te a d ily in density through the end of the
mature cottonwood community.
Several shrub species invaded as the willow thickets thinned and'
disappeared when grasslands became established (Table 4 ) .
The sum of
t h e ir densities was zero in the th ic k e t community, rose to 1 .1 , 11.5
and 22.4 per square meter in the young cottonwood, mature cottonwood
and shrub communities, resp ectively, and f e l l to 0.6 per square meter
in the grassland community (Table 4 ) .
Symphoricarpos occidental is and
Rosa woodsii were the major shrubs in a ll but the grassland stage where
Artemisia cana became established a f t e r other shrub and tree species
began to disappear.
In the preen ash sere, Fraxinus Pennsylvania density increased
f r om 5 plants per square meter in the young cottonwood community to 88
plants per square meter in the green ash community (Table 4 ) .
The
highest densities o f Fraxinus pennsylvanica seedlings and saplings
(132 pqr square meter) occurred in the willow-shrub community, provid­
ing evidence fo r eventual succession to green ash (see Appendix G for
complete l i s t and size classes).
Discussion of Serai Causes and Relations
j
~
■1
The observed serai changes apparently r e f le c t changes in water
a v a ila b ility .
Establishment o f Populus deltoides seedlings is depend­
ent on a moist soil surface fo r t h e ir f i r s t week of l i f e
Hosner 1957).
(Moss 1938,
S alix f l u v i a t i l i s and S alix amygdaloides seedlings also
establish glopg with Populus d e lto id e s .
During Spring high water, the
36
previous year's seedlings are inundated and sometimes buried under a
fresh lay e r of s i l t , however, both Populus' deltoides and Sa lix spp.
seedlings can survive long periods o f inundation and send up fresh
stems and leaves (.Hosner 1958).
and s i l t accumulates.
The seedlings s t a b iliz e the sandbars
Consequently, soil height above the River level
rises ra p id ly (Figure 6 ) .
As the soil height rises above the River le v e l, species such as
Populus deltoides and S alix f l u v i a t i l i s , which are dependent on wet
substrates fo r establishment, f a i l to regenerate.
The o rig in a l colon­
izers th in (Table 4) because they gradually die and do not replace
themselves under t h e ir own canopies.
This may be due to old age and
competition fo r availa b le water, sunlight and nu trien ts.
The increase in shrub cover as the Populus deltoides stand thins
may have been due to lowered competition fo r av aila b le n u trie n t and
water supplies.
As the Populus deltoides canopy opens (Table 2 ) ,
increased Tight may also contribute to the increase in shrub growth.
A fte r the Populus deltoides canopy disappears, the shrubs eventually
decline, perhaps due to water stresses associated with increased in s o l­
ation and wind flow.
Only r e l i c shrub populations appear in the
grassland community.
Species such as Agropyron sm ithii and Fraxinus
pennsylvanica, which are not d ir e c t ly dependent on r i v e r water for
growth, then may establish and subsequently dominate the vegetation.
In thq green ash sere, FTaxinus pennsylvanica seed!ings f i r s t
appear in the young cottonwood community and increase in density
through the shrub stage.
These seedlings are found to germinate
37
and grow over a wide range o f soil nitrogen, phosphorus and potassium
concentrations and availa b le water content values (Johnson et a l .
1976), although seedling densities have been found to be higher in
n u tr ie n t -r ic h , moist areas.
te n t.
I did not measure available moisture con­
However, I suspect that lack o f a v ailable water is the factor
which prevents a more general Fraxinus pennsylvanica establishment on
the flo odplain.
That water which occurs is re s tric te d to areas near
the flo o d p la in 's edge where runoff stored in organic-rich s oils may
provide enough moisture fo r seedling growth.
These sites may also be
sheltered from drying winds.
The willow-shrub community may be an intermediate between cotton­
wood communities and the green ash community type.
These areas pos­
s ib ly are more mesic than sites occupied by other la t e successional
communities due to e ith e r flooding or Spring stream flow.
Fraxinus
pennsylvanica seedlings and saplings are present in a ll transects
( Table 17)
indicating possible; succession to a green ash community.
Changes in Aboveground Biomass Through Succession
Several studies have dealt with the temporal and spatial changes
o f above- and belowground biomass during succession.
T y p ic a lly , above­
ground biomass increased during forest succession ( Egunjobi 1979, Gosz
1980, Grier et a l . 1981, Long and Turner 1975).
I studied changes in
biomass in the grassland sere and also found a pattern o f increasing
forest biomass as the cottonwood community aged.
However, to ta l biomass
f e l l in the ensuing shrub and grassland communities (Figure 8 ) .
38
MA SS ( K G / M 2)
SHOOT
LEAF
ROOT.IMM
ROOT
AGE
Figure 8.
( YRS)
Above and belowground biomasses of communities of increasing
age, grassland sere. Upper lines indicate le a f mass, total
liv in g shoot mass (including leaves) and to ta l aboveground
organic m atter. The lower lines indicate masses of roots
with diameters less than I mm, to ta l root mass (including
small ones), and total belowground organic m atter. Bar seg­
ments represent community types: sandbar (SA), seedling
(SE), th ick e t (TH), young cottonwood (YC), mature cottonwood
(MG), shrub (SN) and grassland (GR).
39
Total aboveground biomass rose s te a d ily from,164 grams per square
meter in the seedling community to 20695 grams per square meter in the
mature cottonwood■community , and then dropped to 784 and 505 grams per
square meter in the shrub and grassland communities, respectively
(Figure 8 ) .
Total l i v e biomass (shoot plus le a f ) followed the same trend,
ris in g from 118 grams per square meter in the seedling community to
19348 in mature cottonwood community and then decreasing to 140 and
169 grams per square meter in the shrub and grassland communities,
respectively (Figure 8 ) .
The dramatic ris e and f a l l of aboveground biomass re fle c te d the
development and decline of the Populus deTtoides trees (Figure 9 ).
The
contribution of Populus deltoides to the to ta l aboveground l i v e bioroa^s
increased from 36% in the seedling community to 99% in the mature
cottonwood community, then declined due to the death and decomposition
o f Populus deltoides trees and the lack o f tree regeneration (Table 4 ).
Salix spp. were important only in the seedling and willow communities
where they constituted 32% and 57% of aboveground biomass, respectively
(Figure 9 ).
Shrubs such as Rosa woodsi i and Symphoricarpos occident­
al is made only, a minor contribution to to ta l biomass u n til the shrub
community, where they provided 87% o f the aboveground l i v e biomass.
Shrubs also comprised 20% of the aboveground li v e biomass in the grass­
land community.
Graminoid and forb biomass constituted 11% o f the to ta l
li v e aboveground biomass in the seedling community, dropped to 1% in
the young cpttonwood stage and increased to 80% during the grassland
40
PODE
( 2 W / 9 »)
IO-
SSVW
8 0
100
(YRS)
Figure 9.
Aboveground masses of Salix s pp., Populus delto ides, shrubs
and herbs in communities of increasing age, grassland sere.
Bar segments represent community types: sandbar (SA)1 seed­
ling (SE), th ic k e t (TH), young cottonwood (YC), mature
cottonwood (MG), shrub (SM) and grassland (GR).
41
community (Figure 9; see Table 34
fo r a complete l i s t o f biomasses
by stand and structural component).
O v e ra ll, Populus deltoides dominated the seedling through mature
cottonwood communities, except in the th ic k e t community where Salix
f l u v i a t i l i s b r i e f l y contributed a greater percentage o f the to ta l liv e
biomass.
Shrubs comprised the bulk of l i v e biomass in the shrub com­
munity and graminoids dominated the grassland community (Figure 9).
Biomass of wood greater than I cm in diameter and wood less than
I cm in diameter followed the ris e and f a l l o f to ta l aboveground bio­
mass ( Figure 8 ) .
This again was due prim a rily to the growth and
decline of Populus deltoides trees (Table 4 ) .
Wood greater than I cm
nearly disappeared as the Populus deltoides trees died, occurring only
in the main stems o f large shrubs.
Wood biomass less than I cm in diam
e ter persisted into the grasslands as the structural wood o f shrubs
(Figure 9 ).
Leaf mass varied considerably and peaked while the cotton­
wood communities were r e la t iv e ly young (Figure 8 ).
Changes in Belowground Biomass Through Succession
Studies of fo rest soil organic matter through time have shown
rapid increases during early succession followed by s t a b iliz a t io n in
l a t e r stages (Dickson and Crocker 1954c, Grier et a l . 1981, Johnson et
a l . 1976, Olson 1958, Switzer and Nelson 1979, Syers et a l . 1970).
results showed the same trend (Figure 8 ) .
My
Soil organic matter rose
from 3097 grams per square meter on sandbars to 29204 grams per square
meter in the mature cottonwood and grassland communities ( Figure 8 ).
This rapid early successional ris e in soil organic matter re fle c ts the
42
production and decomposition o f roots and, to a lesser extent, of le a f branch l i t t e r .
Total belowground organic matter was especially high
in the mature cottonwood community because o f biomass stored in large
Populus deltoides roots and buried trunk bases.
Roots alone comprised
less than 10% o f belowground organic matter in a ll communities except
the young cottonwood and mature cottonwood, where they constitute
approximately 30% of belowground organic matter (Figure 8 ) .
Masses of roots less, than I mm in diameter rose from an average of
0.22 kg per square meter in the seedling community to 1.22 kg per
square meter in the grassland community.
The biomass o f roots greater
than I mm in diameter increased from an average of 0.13 kg per square
meter in the seedling community to 7.55 kg per square meter in the
mature cottonwood community, then declined to 0.49 kg per square meter
in the grassland community (Figure 7 ) .
A major proportion o f root bio­
mass greater than I mm in diameter consisted of Populus deltoides root
crowns and buried trunk bases.
Buried trunk bases were pooled with
roots because of t h e ir location below the soil surface.
Burial occurred
during early succession when s i l t was deposited by flood waters in
developing cottonwood communities.
Changes in Nutrient Content Through Succession
The ecosystem content of several elements was measured to determine
t h e ir changes through succession.
Elements studied were chosen on the
basis o f ecosystem flow characteristics hypothesized below.
In ­
organic solutes (sodium, potassium and phosphorus) enter the ecosystem
in p re c ip ita tio n ( Duvigneud and Danaeyer-de Smet 1970), on sediments
43
(Jenny 1941) and in r iv e r solution.
River deposits should accumulate
rapidly in early succession when flooding is common ( Syers et a l ,
1970, Switzer and Nelson 1979). As the land surface rises above the
River water l e v e l , io n -ric h c a p illa ry and flood water become less
a v a ila b le .
S im ila r ly , with the passage o f time, these elements are
f i l t e r e d by the roots, organic matter and clays in increasing amounts
o f ho rizon ta lly-in te rv e n in g s o ils .
Consequently, sodium, potassium
and phosphorus inputs decrease through succession.
As the a v a i l a b i l i t y
of River and ground water decreases, stores of inorganic solutes should
s t a b iliz e or even decrease i f losses from leaching, fir e s and plant
harvesting exceed imputs.
The rate of f a l l is hypothesized to decline
from sodium to potassium to phosphorus since the binding c a p a b ility to
soil p a rtic le s increases over this series (Jenny 1941).
Carbon content,
measured as biomass, should ris e with biomass development (Wright et
a l . 1959, Switzer and Nelson 1979) in moist early successional commun­
i t i e s because carbon enters the system from the a i r and thus the source
is unlimited.
Actual uptake, however, is determined by water a v a i l­
a b i l i t y via stomate aperture.
Water is abundant in early serai commun­
i t i e s but, as the ground level rises and the River becomes more dista n t,
the community becomes more dependent on scarce r a in f a ll
(Figure 3).
This decrease in av aila b le water lim its carbon fix a tio n so t h a t , when
community re sp iration exceeds carbon f ix a t io n , the overall community
carbon content w ill f a l l toward that o f the regional climax.
content was hypothesized to p a ra lle l carbon content.
Nitrogen
Nitrogen can enter
the communities from unlimited supplies in the a i r through nitrogen f i x ­
ing prokaryotes.
Since i t is covalently bound to carbon in biomass,
44
leaching should be minimal so long as to ta l biomass is increasing,
However, l a t e r in succession when net organic matter decomposition
occurs, the nitrogen released is v o la t iliz e d or leached from the com­
munity.
These hypotheses were tested in the grassland sere.
Inorganic solutes, in f a c t , rose ra p id ly in early succession and
then e ith e r leached out of s t a b iliz e d .
Sodium rose from a mean total
mass o f 0.1/ kg per square meter in the sandbar community to 0.6 kg per
square meter in the mature cottonwood community, then dropped to 0.4 kg per square meter in the grassland community (Figure 10).
Potassium
rose s im ila r ly from a mean to ta l mass of 50 g per square meter in the
sandbar community to a high o f 348 g per square meter in the mature
cottonwood community, then dropped to 235 g per square meter in the
grassland community (Figure 10).
The loss o f sodium and potassium in
the la te serai communities was mainly from the 3-15 dm soil la y e r.
They remained r e l a t iv e ly stable in the 0-3 dm layer (Appendices Q and
P).
Both elements may be removed from the 3-15 dm layer by leaching
downward, as well as through upward wieking by plants tran s piring and/or
c a p illa r y r is e .
The upward wieking apparently prevents decline by re ­
plenishing the nu trien t mass in the 0-3 layer lo s t by leaching.
Late
in succession, loss in the 3-15 dm layer was aggravated by decomposition
of organic ion exchange sites which held both sodium and potassium
(Jenny 1941, Fortesque and Martin 1970).
In contrast, phosphorus rose
from a mean total mass o f 0.74 kg per square meter on sandbars to
approximately 1.00 kg per square meter in the young cottonwood community
but f a ile d to show the la t e succession decline (Figure 10) exhibited by
the more mobile ions.
In plant m a te ria l, a l l three inorganic solutes
45
OM/30
20
Figure 10.
40
60
AGE
80
100
( YRS)
Total ecosystem contents of organic matter (On), phosphorus
( P ) 1 nitrogen (N ) , sodium (Na) and potassium ( K ) . Totals
include contents of soil and plant materials both liv in g and
dead and above and below ground. Bar segments represent
community types: sandbar (SA)1 seedling (SE), thicket (TH),
young cottonwood (YC), mature cottonwood (MC), shrub (SH)
and grassland (GR).
46
followed the gain and loss o f biomass due to storage o f the elements
in woody vegetation and subsequent loss as the Populus deltoides mass
decomposed (Figure 11).
Ecosystem organic matter content increased r a p id ly , from a mean
to ta l o f 3.1 kg per square meter in the sandbar community to 51.7 kg
per square meter in the mature cottonwood community, but then decreased
to 22.3 kg per square meter in the grassland community (Figure 10).
If
we assume that most of the organic matter is cellulose (glucose), i t is
approximately 53% oxygen, 40% carbon and 7% hydrogen.
Total below­
ground mass increased from 3.1 kg per square meter in the sandbar com­
munity to 30.0 kg per square meter in the mature cottonwood community
(Figure 8 ) .
I t then dropped when the Populus deltoides died and th e ir
roots decomposed.
Aboveground to ta l organic matter rose sharply as the
Populus deltoides trees matured and dropped almost as sharply when the
Populus deltoides senesced and shrub or grasslands invaded (Figure 8 ).
Nitrogen content increased from 0.23 kg per square meter on sandbars to 0.88 kg per square meter in the mature cottonwood community
(Figure 10).
community.
I t then f e l l to 0.74 kg per square meter in the grassland
This loss in nitrogen content was p rin c ip a lly from the 3^15
dm soil layer (T a b le 29).
composition of organic matter.
I t was probably associated with the de­
As with the other elements, nitrogen
remained r e la t iv e ly stable in the 0-3 dm layer in la t e succession.
As
predicted, total plant nitrogen dynamics follow the changes in organic
matter (Figure 11).
47
20
40
60
AGE
Figure 11.
80
100
(YRS)
Elemental contents of liv in g plants as related to system
age. Bar segments represent community types: seedling (SE),
th ic k e t (TH), young cottonwood (YC), mature cottonwood (MG),
shrub (SH) and grassland (GR).
48
In summary, the ecosystem contents of potassium, sodium, carbon
and■nitrogen rose and f e l l through time.
Sodium and potassium contents
rose in early- succession and were then lo s t from the system as input
supplies and storage capacity decreased.
S im ila rly , carbon and n it r o ­
gen rose rapidly as biomass was accumulated and decreased with the
demise o f the Populus deltoides tre e s .
In contrast, phosphorus showed
a more rapid i n i t i a l ris e and no sign o f eventual decline.
I a ttrib u te
the ris e in phosphorus to deposition with s i l t and the f a ilu r e to de­
c lin e to the continued t ig h t binding by the system.
49
MANAGEMENT IMPLICATIONS
Logging or burning of cottonwood communities may speed succession
to s e ra iIy advanced communities such as. grasslands or green ash.
This
w ill occur since, i f the land surface has risen well above the River
le v e l, Populus deltoides and Salix spp. seedlings cannot establish
while grasses and Fraxinus pennsylvanica can.
I f flooding is minimized by water impoundment or diversion, bank
cutting w i ll be reduced and consequently s e ra iIy advanced communities
may become more common at the expense o f early serai communities.
Channelization by concrete embankments or rip -ra p would create the same
r e s u lt.
Under these conditions, destruction o f older communities by
erosion w i ll occur less frequently and new sandy deposits needed for
colonization by early serai communities w i ll be minimized.
I f over-bank flooding is reduced by water impoundment or diversion,
the land surface would be elevated less due to reduced s i l t deposition
during high water.
This may prolong the presence of plants dependent
on shallow ground water.
Also, River channels that are now flooded in
the Spring but r e l a t iv e ly dry in the Summer may remain permanently dry,
allowing permanent plant colonization.
LITERATURE CITED
LITERATURE CITED
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_______ , and J. C. Wright. 1966. Flora o f Montana.
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Part I I .
Montana
Bremner, J. 1965. Inorganic forms of nitrogen. Ini Methods of soil
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52
Fortescue, J . A. C., and G. G. Marten'. 1970. M icronu trien ts: forest
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_______ . 1958. The e ffects of complete inundation upon seedlings of
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29-41.
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York. 281 pp.
Univer­
McGraw-Hill Bopk Co., New
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472 pp.
1967.
Vegetation mapping.
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53
Moss, E. H. 1938. Longevity of seed and establishment o f seedlings
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Madison., Wisconsin, pp. 1035-1049. ■
Olson, J. S. 1958. Rates of succession and soil changes on southern
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Edited by C. Black et a l . Amer. Soc. o f Agronomy, Madison, Wis­
consin. pp. 1022-1030.
_____ . 1965. Sodium. L% Methods o f soil a n alysis. Part 2. Edited
by C. Black et a l . Airier. Soc. o f Agronomy, Madison, Wisconsin,
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V ol. I .
Gale Research
Saunders, W., and E. W illiam s. 1955. Observations on the determination
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w i l d l i f e b io lo g is ts .
1980. Checklist o f North American plants fo r
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soil organic matter. Soil S c i. 112: 137-141.
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7th
Stevens, M. A . , R. L. Kerr, and E. Olgeirson. 1978. Yellowstone River
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54
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633-654.
USGS.
1931 i
The climates of North America.
Geogr. Rev. 21:
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Hudson Bay Basin, Missouri
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VanBruggen,
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H 7 pp.
Walker, H. 1973.
237 pp.
Vegetation of the e arth .
Springer-Verlag, New York
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---------- 1982.
s o ils .
D is tribution of root biomass in well-drained surface
Amer. Midland N a tu ra lis t 107: 393-395.
Whittaker, R. H ., and G. M. WoodwelI .
1968a. Dimension and production
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Ecol . Monogr. 44: 441-464.
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soils on a llu v ia l deposits in the Northwest T e r r it o r ie s . Can. J.
Soil S c i. 39: 32-43.
55
APPENDIX
56
TABLE 5
•
Locations o f Stands Studied
Location
Number
Biomass
Stand
Local Name
I
M, (N)
Glendive
2
M
Glendive
3
Gentry's
land
4
5
6
Above intake
N
7
Intake
Downstream
o f intake
8
N
9
M
Mary's
Island
10
" "
11
12
0
(N ), 0
13
14
Elk Island
Crittendon
Island
K
15
16
Elk Island
Seven Sis­
ters Island
T16N, R56E, S18, SW
corner o f NW 1/4 sec.
T17N, R56E, S36, NW
1/4 section
T17N, R56E, S17
north h a lf
T17N, R56E, S5, SW
corner of NW 1/4 sec.
T17N, R56E, S2, NW
corner o f SW 1/4 sec.
T18N, R56E, S33, SW
1/4 section
T18N, R57E, S31, SW
corner o f NW 1/4 sec.
T18N, R57E, S20, SE
corner
T19N, R58E, S20, SW
corner o f SW 1/4 sec.
T20N, R58E, S34, NW
corner o f NW 1/4 sec.
T20N, R58E, S27, NW
1/4 section
T20N, R58E, S26, NW
1/4 section
T20N, R58E, S I* , NW
corner o f SE 1/4 sec.
T21N, R58E, S23
L
T21N, R58E, S13,
corner o f SW 1/4
T21N, R58E, S12,
corner of NW 1/4
M
T22N, R59E, S20, SW
corner
T22N, R59E, S10, NW
corner o f SE 1/4 sec.
17
18
Township, Range and
Section
Sidney
SW
sec.
NE
sec.
Cqmmunity
Types Examined
MC*,SB, GR*,
(GR*), W-S
SA,SE,TH*,YC,
MC1SH*,GR
GA
GA
GA
SA1SE1YC*,
SB*, GR
GA
SA*,SE*,TH*,
YC
SA*,SE*,TH,
YC1SH1GR
GA, W-S
MC*,SH*,GR*
SA*,SE*,TH*,
YC*,(MG*)
SA1SE1TH1YC
SA*,SE*,TH*,
YC*,MC*,SH*,
GR*,W-S.,PLW
GA
SA*,SE*,TH*,
YC*,MC*,SH*,
GR*
SA1SE1TH1SH,
GR
YC*
*Biomass was studied in community types marked with an a sterisk
Community types are sandbar (SA), seedling (SE), th ic k e t ( T H ) ,‘young
cottonwood (YC), mature cottonwood (MC), shrub (SB), grassland (GR),
willow-shrub (W-S), green ash (GA) and peach-leaved willow (PLW)
57
TABLE 6
Composite Vascular Plant Species1 L is t o f
the Lower Yellowstone River Floodplain
TREES
Acer negundo L.
Elaeagnus angustif o l ia L.
Fraxinus pennsylvanica
Marsh.
Jum'perus scopulorum Sarg.
Populus deltoides Marsh.
Salix amygdaloides
Anders.
Salix f l u v i a t i l i s N u tt.
Ulmus americana L .
p
O
(Frpe)d
boxelder maple
Russian-olive
green ash
(Pode)
(Saam)
Rocky Mountain ju nip er
eastern cottonwood
peach-leaved willow
(S a fi)
(Ulam)
sandbar willow
American elm
SHRUBS, CACTI AND VINES
Amelanchier a l n i f o l ia Nutt.
Artemisia cana Nutt.
(Area)
Artemisia dracunculus L.
Artemisia f r ig id a W illd .
Artemisia liidoviciana
(Arlu)
Nutt.
^
Cornus Stolonifera Michx.
(Cost)
Coryphantha v iv ip a ria (N u tt.)
B r i t t . & Brown
Cotoneaster sp.
Gutierrezia sarothrae
' (Pursh.) B.& R.
Jum'perus horizontal is
Moench.
Opuntia f r a g i l is (N u tt.)
Haw.
Parthenocissus quinquefolia (Paqu)
(L .) Planch
Prunus virginiana Marsh.
( P r v i)
Rhus t r ilo b a t e Nutt.
(Rhtr)
Ribes aureum Pursh,
(Riau)
Saskatoon serviceberry
s ilv e r sagebrush
tarragon sagebrush
fringed sagebrush
Louisiana sagewort
red-osier dogwood
cushion coryphantha
cotoneaster
broom snakeweed
creeping juniper
b r i t t l e pricklypear
V irg in ia creeper
common chokecherry
fragrant sumac
golden currant
I
S c ie n t ific names follow Booth (1972) and Booth and Wright (1966) or,
Pfo r species not lis t e d by Booth, VanBruggen (1976).
^Common names follow Scott and Wasser (1980).
dFour l e t t e r abbreviations of binomials presented in the te x t are also
li s t e d .
58
TABLE 6 - - C o n t i n u e d
Ribes setosum L in d l.
Rosa'woodsi i L in d l.
Shepherdia argentea Nutt.
Symphoricarpos occidental is
Hook;
Tamarjx chinensis Loureiro
Toxicondendron rydbergii
(Small) Greene
V it is r ip a r ia Michx.
(Rise)
( Rowo)
(Shar)
(Syoc)
redshoot gooseberry
woods rose
s ilv e r buffaloberry
western snowberry
(Tory)
tamarisk
common poison-ivy
(V iri)
riverbank grape
GRAMINOIDS
Agropyron cristatum ( I . )
crested wheatgrass
Agropyron repens (L .)
(Agre)
Beauv.
Agropyron sm ithii Rydb.
(Agsm)
Agropyron trachycaulum
(Link) Malte
Agrostis alba LAgrostis diegpensis Vasey
Andropdqon h a l l i i H Tack.
^eckmanhTjr syzigachne
(Steud.) Fernald
Bouteloua g r a c ilis (H .B .K .)
Lab.
Bromus inermis Leys.
(Brin)
Bromus .iaponicus Thunb,
Calamovil fa Io n q if o lia
(Calo)
(Hook.) S c ribn.
Carex brevior (Dew.) Macken.
Carex lanuginosa Michx.
D is tic h lis s t r ic t a (T o r r .)
Rydb.
Echinochloa c rusgalIi ( L . ) (Eccr)
Beauv.
E leocharis palustris ( L . )
R.& S.
E lvmus canadensis L.
(Elea)
Elvmus iunceus Fisch.
E lvmus viroinicus L.
Eragrostis hypnoides (Lam.)
B.S.P.
Eragrostis pectinacea
(Michx.) Nees.
Hordeum ,iubatum L.
quackgrass
western wheatgrass.
slender wheatgrass
redtpp beptgrass
sand bluestem
American sloughgrass
blue grama
smooth brome
Japanese brome
p r a ir ie sandreed
inland saltgrass
common barnyardgrass
common spikerush
Canada wildrye
f o x t a il barley
i
59
TABLE 6 — C o n t in u e d
Juncus balticus W ind.
Muhlenbprgia racemosa
(Mura)
(Michx.) B.S.P.
Oryzppsis hymenoides
(R.& S.) Ricker
Panicum c a p illa re L.
Phalaris arundinacea L.
Poa palus tris L.
Poa pratehsis L.
Poa reflexa Vasey & S c r ibn.
Polypogon monospeliensis
'
( L ) Desf.
Scirpus americanus Pers.
Scirpus v a lidus Vahl.
Setaria glauca ( L . ) Beauv.
Setaria v ir i d i s (L .) Beauv.
Spartina g r a c ilis T r i n.
Sporobolus cryptandrus (T o r r .)
Gray
Stipa comata T r i n . & Rupr.
Stipa v ir id u la T r in .
Vul pi a optoflora (W a lt.)
R yd b T "
B a ltic rush
Indian ricegrass
witchgrass panicum
reed canarygrass
Kentucky bluegrass
American bulrush
softstem bulrush
yellow bristlegrass
green bristlegrass
a lk a li cordgrass
sand dropseed
needle-and-thread
green needlegrass
sixweeks annual Fescue
FORBS
Abronia fraqrans Nutt.
Achillea mi11efo'li urn L.
Amaranthus powelli i Wats.
Ambrosia psilostachya DC.
Arabis hoib o e lii Horn.
Arctium lappa L .
Asclepias speciosa Torr.
Asclepias v e r t i c i l la ta L.
Asparagus o f f i c i n a l i s L.
Aster chilens is Nees.
Aster ericoides L.
Aster ob lonq ifolius Nutt.
Aster pansus (BTake) Cronq.
Bidens frondosa L.
Chenopodium aI bum L.
Chenopodium leptoohyllum Nutt.
Chrysopsis v illo s a (Pursh.)
N u tt.
Cirsium arvense ( L .) Scop.
Clematis I i q u s t i c i f o l ia Nutt.
common yarrow
western ragweed
rockcress
Asparagus
P a c ific aster
heath aster
devils beggarticks
lambsquarters goqsefoot
s lim le a f goosefoot
hairy goldenaster
Canada t h i s t l e
60
TABLE 6 _ _ c o n t i n u $ d
Cleome serrulata Pursh.
Convolvulus sepium- L.
Conyza canadensis ( L .) Cronq.
Cycloloma a t r i p iic ifo liu m
" (Sprang.) Coult .
Equisetum arvense L.
Equisetum laeyigatum A .Br.
Euphorbia esula L.~~
Euphorbia glyptosperma
Engelem.
Galium t r i f Iorum Michx.
Gaura coccinea PUrsh.
Gaiira p a rv iflo ra Dougl.
Glycyrrhiza lepidota
(GHe)
( N u t t . ) Pursh.
Gnaphalium palustre Nutt.
Grindelia~squarrosa (Pursh.)
D ural.
Helianthus p e t iplan's Nutt.
Iva a x i l l a r i s Pursh.
Iva xanthifoTia Nutt.
Kochia sgpparia ( L .) Schrad.
Lactuca~pulchella (Pursh.) DC.
Lactuca s e r r i o l a~L.
Lappula echinata G i lib .
Lappula redowskii (Hornem.)
Greene.
Lepidium densiflorum Schrad.
Linum perenne L.
Linum rigidum Pursh.
Lygodesmia juncea (Pursh.)
D. Don
Lysimachia c i l i a t a L.
Medieago lupulina L.
(Melu)
Medieago sativa L.
Meliio tu s aIba Desv.
Melilotus o f f i c i n a l i s ( L .)
(Meof)
Lam.
Mentha arvensis L.
Mentzelia nuda ( Pqrsh.) T.& G.
Nepeta c a ta ria L.
Oenothera a ! b icaul is Pursh.
Oenothera biennis L.
Physalis heterophylla Nees.
Plantago major L.
Plantago patagonica Jacq,.
Rocky Mountain bqeplant
bindweed
Canada horseweed
tunble ringwing
f i e l d horsetail
smooth horsetail
le a fy spurge
sweet-scented bedstraw
s c a rle t gaura
American lic o r ic e
cuflycup gumweed
p r a ir ie sunflower
poverty sumpweed
rag sumpweed
f i reweed summercypress
chicory lettuce
p ric k ly lettuce
European stickseed
western stickseed
p r a ir ie pepperweed
rush skeletonplant
black medic
a lfa lfa
white sweetclover
yellow sweetclover
f i e l d mint
common
clammy
common
woolly
evening primrose
groundcherry
plantain
plantain
61
TABLE
Polygonum aviculare L.
Polygonum coccirieum Muhl.
(Poco)
Polygonum IapathifOlium L.
P o te n tilla anserina L.
PotentiTla paradoxa Nutt.
Psoralea argophylla Pursh.
Ratibida columnifera (N u tt.)
. .
Woot. & Standi .
Rorippa sinuata (N u tt.)
Hitchc.
Rumex maritimus L.
( Ruma)
S a g itta ria cuneata Sheld.
Salsola kali L.
Sisymbrium altissimum (L .)
B ritt.
Sisymbrium lo e s e lii L.
Smilacina s t e lla t a ( L . )
Desf.
Smilax herbacea L.
Solanum rostratum D ural.
■Solanum nigrum L.
Solidago qiqantea A it .
Solidago missouriensis' Nutt.
Solidago mollis B a r t l .
Solidago occidental is
(N u tt.) T.& G.
Sonchus asper ( L.) H i l l .
Tanacetum vulqare L .
Taraxacum Iaevigatum
( W illd .) DC.
Thaiictrum venulosum T r e l .
Thlaspi arvense L.
Tradescantia occidental is
( B r i t t . ) Smyth.
TragoPogon dubius Scop.
T r i f o l i urn fragiferum L.
Verbascum thapsus L.
Verbena bracteata L.& R.
Verbena hastata L.
Veronica americana Schwein.
Veronica catenate Penn.
Viola canadensis L.
Xanthium strumarium L.
G-TContinued
prostrate knotweed
marsh knotweed
curlytop ladysthumb
s i Iverweed cinquefoil
s i ly e r le a f scurfpea
common. Russianthistle
common tumblemustard
s ta rry solomonplume
carriopflower greenbrier
buffalobur nightshade
black nightshade
Missouri goldenrod
tansey
common dandelion
veiny meadowrue
f i e l d pennycress
p r a ir ie spiderwort
yellow s a ls ify
flannel mullein
bracted verbena
American speedwell
common cocklebur
62
TABLE 7
Ages of Representative PopuTus deltoides
and Salix f l u v i a t i l i s in Stands Studied^
^
Community types
Species2
Location number
Seedling
O
Pode
Safl
Thicket
Pode Safl
Young
Cottonwpod
Mature
Cottonwood
Pode
Pode
I
-
-
-
-
2
2
2
4
4
6
4
3
-
-
8
3
3
11
7
26
9
3
3
6
4
31
1,1
-
-
-
-
■
9&
12
3
3
7
5 '
-
85
13
4
2
6
6
-
14
3
3
9
9
23
95
16
4
2
7
7
43
751
17
3
2
7
4
T
18
-
-
-
■ -
-
108
44
-
39
1Ages were determined by sectioning or coring the trees and counting
gthe annual rings.
^Species are Populus deltoides (Pode) and Salix f l u v i a t i l i s ( S a f i) .
Locations are id e n t if ie d in 'T a b le 5.
r
63
TABLE 8
Elevation (m), Relative to the River
Surface, of Stands Studied^
Community
type ^
SA
SE
I
-
-
-
-
3.4
3.9
2
0.9
1.6
1.9
3.4
4.5
3.9
3.5
-
■
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
" TH
YC
MC
SH
GR
W-S
3 .5 /3 .3 ^
3.7
GA
Location
number3
3
! ■■
4
-
5
V
T
4 .0 /
4.0
T
5.0
-
-
5.0
6
1.0
1.4
-
2.7
-
3.7
3.6
-
-
7
-
-
-
-
-
-
-
-
3.7
8
1.3
1.4
2.4
2.4
-
-
-
-
_
9
0,3
1.1 ■
1.7
3.2
-
3.6
3 1
-
-
TT
4.0
4.4
10
-
-
-
-T
11
-
-
-
-
4.1
3.0
3.1
r
-
12
0.7
1.8
1.7
3.4
3.5
-
-
-
-
13
1.0
1.6
2.9
2.9
-
-
-
-
-
14
-
1.4
3.0
2.9
3.5
2 .8 /3 .3
3.3
-
-
15
0.8
-
-
-
-
-
-
-
4.3
16
0.8
0.9
3.2
3.3
3.4
3.5
2.9
17
0.4
0.7
3.0
-
-
3.1
3.6
-
-
18
-
-
3.4
-
-
-
-
-
-
- .
^Elevations (s o il height) were measured with a hand level and a rod.
^Community types are. sandbar (SA), seedling (SE), th ic k e t (TH), young
cottonwood (YC), mature cottonwood (MG), shrub (SH), grassland (GR),
-willow-shrub (W-S), and green, ash (GA).
^Locations are id e n tifie d in Table 5 .
Two transects were studied at the same l o c a l i t y . The elevations are
presented in the same order as in the 'Presence o f Species' Appendix
(Tables K k l 7 ).
64
TABLE 9
SE
Parameter^
ht
of the Upper Canopy Surface and Diameter (cm)
pf Populus deltoides in Stands Studied.
TH
ht
YC
dia
ht
MC
dia
ht
-
25.8
17.0
dia
0
SH
W-S
GA
ht
ht
ht
1.0
1.7
I
Community
typed
I
ro
45»
Height (m)
■
■
Location
number5______________________
I
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
-
-
-
0.3
-
2.0
-
3
-
0.4
-
-
-
-
-
-
-
-
-
-
-
2.5
2.0
0.3
r
0.8
0.3
i-
0.2
0.6
-
4
6
-
-
-
2.0
2.5
1.0
7
3
I
-
15.3
-
23
-
80
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
19.9
17.3
18
22
-
-
-
-
-
T
-
-
20.3
-
-
-
0.2
0.1
4.0
1.5
4
6
20.2
-
-
-
-
17.9
-
26
-
29
-
28,5
23.8
-
60
63
_
27.0
-
25.3
70
-
51
-
-
_
21
-
-
21.3/
13.9
10.7
11,3
-
,
-
_
12.0
-
_
-
-
4.0
0.9
—
r
_
■
«•
_
_
I .8/
1.7
■
-
0.9
-
-
8.7
ee
15.8
r
T
—
W
mm
^Height of the upper tree canopy surface was measured with an Abney
J e v e l.
^Diameter of representative Pppulus deltoides trees used fo r age were
measured a t t h e ir bases in the th ic k e t community type and a t breast
-height in the young cottonwood and mature cottonwood community types.
^Community types are seedling (SE), th ic k e t (TH), young cottonwood (YC),
mature cottonwood (MC), shrub (SH), grassland (GR), willow-shrub (W-S)
.and grepn ash (GA).
^Parameters are height (h t) and diameter ( d i a ) .
^Locations are id e n tifie d in Table 5.
6Two transects were studied at the same l o c a l i t y . The heights are pre­
sented in the same order as in the "Presence of Species" Appendix
(Tables 10 - 17).
65
TABLE 10
Presence
( %
C o v e ra g e )1 o f S p e cies
Location number'
Bare ground
L itte r
in
th e S e e d lin g
Community Typq
9
12
23
3
72
5
63
3
55
8
40
7
69
8
74
72
79
-
-
-
-
37
2
18
17
+
2
15
3
2
30
2
5
25
3
12
15
+
+
18
2
5
13
20
+
2
-
TREE SPP.
Popul us del toides
$aI i x amygdaloides
S alix f l u v i a t i l i s
+
SHRUB SPP.
Artemisia I udoviciana
Gutierrezia sarothrae
_
+
+
-
-
+
+
+
GRASS SPP.
Agropyron repens
Bouteloua g r a c ilis
P is t ic h lis s t r ic t a
EchinochToa crusqalI i
Eleocharis palustris
Elymus canadensis Eragrostis hypnoides
Eragrostis peetinacea
Hordeum jubatum
Muhlenbergia racemosa
Qryzopsis hymenoides
Panicum c a p ilIare
Poa pratensis
Polypogon monospeliensis
Setaria glauca
Setaria v ir id i s
Sporobol us cryptandrus
-
-
-
-
-
-
-
-
-
8
2
+
+
5
-
+
r
-
+
+
-
-
_
+
+
2
+
-
-
-
_
-
T
-
-
-
-
-
T
-
i
-
V
-
-
-
-
+
+
+
-
-
r
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
+
+
-
-
T
-
-
-
-
+
-
+
-
-
~
-
-
r
-
+
+
-
+
~
-
-
r*
-
-
-
-
-
_
+
•p
-
-
-
+
"
FORB SPP.
Aster chi lens is
Aster pansus
Bidens frondosa
Chenopodium album
Cleome serrulata 1
_
-
-
-
" '
2
+
-
-
-
-
-
+
-
2
-
-
+
-
-
+
-
-
1Grass and forb coverage were estimated by the step-point m ethod, shrub
cover by calculatin g canopy coverage ( p i - r 2) of each shrub present in
gdensity plots., and tree coverage was based upon ocular estimates
^Locations are id e n t if ie d in Table 5 ,
Species found in the same community type, but not sampled.
66
TABLE 1 0 —.C o n tin u e d
Location number
Convolvulus sepium
Conyza canadensis
Cycloloma a t r i p ! ic ifo liu m
Euphorbia glyptosperma
Glycyrrhiza lepidota
Gnaphalium palustre
Grindeiia squarrosa
Helianthus p e tio la ris
Iva a x i l l a r i s
Meli lotus alba
Melilotus o f f i c i n a l i s
Mentha arvensis
Oenothera biennis
Physalis heterophylla
Polygonum aviculare
Polygonum coccineum
Polygonum lapathifolium
P o te n tilla anserina
P o te n tilla paradoxa
Rorippa sinuata
Rumex maritimus
Salsola ka li
SdlIda^o gigantea
Sqlidaqo missouriensis
Solidago molI i s
Sonchus asper
Tanacetum vulqare
Vqrbena bracteata
Verbena hastata
Veronica catenata
Xanthium strumarium
I
9
12
+
+
-
-
16
13
8
2
+
-
-
-
2
-
-
-
_
+
-
-
17
2
6
-
T
14
Present
+
-
+
_
+
-
-
-
-
-
-
-
2
+
_
-
+
_
2
_
_
4
_
+
-
-
-
-
-
-
-
+
3
-
-
_
_
-
+
+
-
-
_
-
+
2
+
5
2
13
+
-
-
-
_
_
5
-
-
-
-
-
-
_
+
_
■
—
_
_
_
_
2
+
3
3
+
_
+
+
2
+
+
-
3
3
-
-
_
—
»
-
-
-
T
-
-
-
-
-
-
-
r
-
-
-
+
-
-r _
-
-
-
-
-
+
_
_
-
, _
T
-
-
-
-
-
_
7
+
-
-r
+
T
-
+
+
•
+
r
r
_
+
+
_
_
t
+
_
-
-
-
+
67
TABLE 11Presence (% Coverage) o f Species in the Thicket Community Type
Location number
16
8
17
2
14
13
12
Bare ground
L itte r
23
75
25
38
12
75
10
74
38
43
25
27
28
43
32
45
+
+
+
+
+
+.
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
H+
-
«■
-
3
-
+
-
T
-
+
-
H-
2
T
_
5
_
2
-T
+
2
5
-
+
_
H_
2
-
3
_
•
+
_
+
t
+
3
-
9" Present
_
-
TREE SPP.
Populus deltoides
S alix amyqdaloides
Salix f l u v i a t i l i s
r
SHRUB AND VINE SPP.
Artemisia f r iq id a
Artemisia ludoviciana
GRASS SPP.
+
-
8
2
_
+
+
+
3
+
-
2
-
w
_
_
-T
2
8
r
_
_
+
+
■
+
_
+
—
+
_
10
5
-
2
+
.
T
+
W
.
_
3
_
•
._
-
-
4r
+
2
FORB SPP,
Ambrosia osi lostachva
Ascleoias v e r t i c i l lata
Aster oblongifolius
Aster oansus
T
8
2
+ +
Aqroovron repens
Aqroovron sm ithii
Aqropyron trachycaulum
Aqrostis dieqoensis
Andropoqon h a l l i i
Bromus inermis
Calamovilfa lo n q ifo lia
Carex brevior
Carex lanuginosa
D is tic h lis s t r ic t a
EchinochToa crus qa lli
Elvmus canadensis
Hordeum iubatum
Juncus balticus
Muhlenberqia racemosa
Phalaris arundinacea
Poa spp.
Poa p alu stris
Poa pratensis
Poa r e f l exa
Polvpogon monosoeliensis
Scirous americanus
Scirous v a lidus
Setaria v ir i d i s .
Sborobol us cryptandrus
r
2
+
2
2
-
-
-
15
68
TABLE ! ! - - Continued
Location number
Chenopodium aI bum
Chenopodium le p to p h y llurn
Cirsium arvense '
Conyza canadensis
Cycloloma a t r i p li c if o li u m
Equisetum arvense
Equisetum laevigatum
Euphorbia glyptosperma
Galium triflo ru m
Glycyrrhiza lepidota
Gnaphalium palustre
Hel ianthus peti'olaris
MeliIo tu s alba
MeliIo tu s o f f ic i n a l is
Oenothera a l bicaul is
Plantago major
Plantago patagonica
Polygonum lapathifolium
P o te n tilla anserina
P o te n tilla paradoxa
Rorippa sin'uata
Piumex m'aritimus
Salsola~kali
Sisymbrium lo e s e lii
Solidago gigantea
Solidago occidental is
Sonchus asper
Tanacetum v u lgare
Taraxacum laevigatum
Tradescantia occidental is
Verbena bracteata
Veronica americana
16
8
17
2
14
13
12
9
-
-
+
-
-
-
-
-
-
-
-
-
+
-
-
-
+
-
-
-
-
-
-
-
-
3.
3
Present
-
+
-
T
-
-
-
-
+
-
-
-
-
+
+
+
-
+
-
-
_
'
_-
-
-
-
-
-
-
+
+
3
-
-
-
-
3
-
-
-
-
-
-
-
-
-
-
+
-
-
_
-
-
+
3
_
10
-
-
-
-
_
-
-
2
+
-
-
.
_
3
2
-
-
-
-
+
-
2
13
3
-
3
-
-
-
-
-
+
-
-
-
-
-
-
-
-
-
-
-
+
-
+
-
-
2
T
-
-
-
-
-
-
-
-
+
-
-
_
-
-
+
+
-
2
-
-
-
-
+
-
-
-
-
-
-
-
-
T
-
-
+
+
-
-
I
-
-
+
-
+
-
-
+
-
-
-
-
-
-
-
-
-
t
-
'
,
.
7
-
-
-
-
-
+
3
7
-
-
-
-
-
+
_
+
-
+ .
-
-
-
-
-
-
-
-
-
-
-
-
-
+
-
-
+
-
-
-
-
-
-
-
-
i
-
-
-
T
-
+
-
-
-
-
-
2
'
-
7
-
-
-
-T
-
-
-
_
69
TABLE 12
Presence
{%
C overage)
Location number____________
Bare ground
L itte r
o f S p e c i e s i n t h e Young C o tto n w o o d
Community Type
9
14
8
18
2
12
13
6
Present
2
62
16
57
. 2
82
74
13
67
2
77
3
69
20
57
3
58
-
_
65
+
+
60
-
+
t
75
+
+
75
-
+
45
+
+
75
+-
-
-
'60
-
+
60
+
75
+
r
+
+
+
+
-
+
+
+
+
+
+
+
+
+
-
-
■
r>
+
T
+
+
8
8
+
-
TREE SRP.
ETLaeagnus an g u stifo lia
Fraxinus pennsyIvanica
Juniperus scopulorum
Populus del toides
S alix amygdaloides
I
SHRUB SPP.
Amelanchier aI n i f o l i a
Artemisia I udoviciana
Cornus s tolo nifera
Parthenocissus quinquefolia
Rhus t r ilo b a t e
Ribes aureum
Ribes setosum
Rosa woodsi i
Shepherdia argentea
Symphoricarpos occidental is
Toxicodendron rydbergii
V it is rip a ria
+
+
2
+
+•
+'
+
I
+
T
+
+
-
+
~
+
+
22
-
8
3
-
5
-
-
7
I
5
-
8
-
2 •+
3
5
2
+
13
2
-
r
2
7
13
T
-
2
-
T
+
T
+
-
_
r
IT
-
+
•
n
+
+
t
-
-
-
-
-
_
r
+
+
-
+
-
_
+
-
2
—
_
GRASS SPP.
Aqropyron repens
Agropyron sm ithii
Agrostis aTba
Bromus inermis
Calamoyilfa lo n g ifo lia
Carex brevior
Carex lanuginosa
Elymus canadensis
El vmus .iunceus
Elymus virginicus
Muhlenbergia racemosa
Poa spp.
Poa palustris
Poa pratensis
Polypogon monosoeliensis
Setaria v ir id i s
-
5
15
-
-
-
7
2
T-
-
+
-
-
10
5
+
-
3
3
- ‘
_
2
-
-
_
■
12
_
8
-
12
_
13
2
-
+
■
+
+
+
4r
70
TABLE 1 2 — C o n t in u e d
L o c a tio n
number________________
9
14
IB
8
18
2
12
13
6
Present
FORB $PP.
Ambrosia pgilostachya
Asclepias speci osa
' Asclepias v e r t i c i l la ta
Aster pansus
Chrysopsis v i Ilosa
Cirsium arvense
Clematis l i g u s t i c i f o l i a
Conyza canadensis
Equisetum arvense
Equisetum laevigatum
Euphorbia esula
Glycyrrhiza lepidota
Grindelia squarrosa
Helianthus p e tio la ris
Lactuca pulchella
Medicago lupulina
MeliIo tu s aIba
MeliIo tu s o f f i c i n a l is
Plantago major
Sdl idago' gigahtea
' Thaiictrum venulosum
Verbascum thapsus
Viola canadensis
2
+
-
+
-
+
T
-
T
+
+
+
—
—
—
—
—
+
+
2
2
+
+
+
7
-
+
-
-
—
—
—
—
3
+
.+
+
71
TABLE 13
Presence
( %
C overage) o f S pe cies in
Community Type
Location number____________
Bare grpund
L itte r
t h e M a t u r e C o tto n w o o d
I
2
-
«■
47
14
12
16
11
Present
60
63
3
85
67
2
45
-
+
+
50
+
+
+
25
+
+
50
+
— 4+
+
+
40 30
TREE SPP.
Acer negundo
El agagnus T h g u s ti fol i a
Fraxinus pennsyIvanica
Jumperus spopplorum
Populus deltoides
S alix amygdaIoides
+
+
+
45
_
-
SHRUB AND VINE SPP.
Artemisia Iudoviciana
Cornus stolo nifera
Parthjenocissus quinquefol ia
Rhus t r llo b a t a
Ribes~aureum
Ribes setosum
Rosa woodsi i
ShepherdTa 'argentea
Symphoricaroos occidental is
Toxicodendron rydbergii
V it is r ip a r ia
+
2
19.
+
+
5
I
T
5
—
7
20
13
+
+
+
+
—
—
2 .2
I
+
T
—
—
—
+
-
+
14
—
7
2
3
-
4
+
- 4
10
8
20
+
16
25
+
-
3
-
—
+
2
3
2
-
T
3
5
+
3
15
+
5
7
-
3
3
_
8
5
2
2
7
3
10
13
13
-
—
—
-
■
+
+
-
-
+
-
-
—
—
-
T
+
-*
T
f
—
_
GRASS SPP.
Aqropyron repens
Agropyron sm ithii
Aqropvron trachvcaulurn
Bromus inermis
E lvmus canadensis
Muhlenberoia racemosa
Poa spp.
Poa palustris
Poa oratensis
Setaria v i r id is
Sti pa comata
-
+
-
-
10
2
2
11
-
+
+
FORB SPP.
Abronia fra grans
Asclepias v e r t i c i l la ta
Asparagus o f f i c i n a l i s
+
*
72
TABLE 13 --Continued
Location ngmber______ ;_______________
Aster dblongifolius
Cirgium arvense
Euphorbia esula
Euphorbia glyptosperma
Glycyrrhiza lepidota
Lygodesniia juncea
Lysimachia ciliatum
Melilotus o f f ic in a l is
Ratibida columm'fera
Smilax herbacea
Splahvm nigrum
Sol idago gig'antea
Thalictrum venulosum
Tragopogoh dubius
1 2
14
12
16
11
Present
-
+
-
-
-
-
-
+
_
-
-
-
-
-
-
+
+
-
-
-
-
-
-
2
+
+
-
I
-
-
-
-
-
-
-
+
-
-
-
-
-
-
+
-
-
-
-
-
3
-
-
-
-
-
-
+
-
-
-
-
-
-
+
-
-
-
-
-
+
-
+
-
+
-
-
-
-
-
+
+
-
-
-
-
,
-
-
-
-
-
-
*
•
73
TABLE 14
Presence (% Coverage) o f Species in the Shrub Community Type
Location number
14
6
2
I
17
Bars grpupd
L ifte r .
69
82
T
84
64
+
+
16
14
11
9
Pres
B4
89
95
82
80
I
+
-
+
-
-
_
TREE SPP,
Fraxinus pennsylvanica
+
SHRUB AND VINE SPP.
Artemisia f r iq id a
Artemisia Tudoviciana
+
— — Coryphantha v iv ip a ria
—
—
Opuritia' f r a q i l i s
^
Parthenocissus quinquefolia +
I
—
—
_
Rhus t r ilo b a t e
+
Ribes aureum
+
+
— +
Ribes setosum
+
Rosa woodsi i ’
12 11 20 20
Symphoricarpos occidental is 10
9 13
6
Toxicodendron, rydberqii '
10
2
8
V i t f s 'r i p a r i a
(Dead shrubs)'
- 3 - 40 20
_
+
r
n
I
+
+
_
+
T
+
+
17
6
I
4
+
+
3
I
_
14
3
+
14
8
+
10
14
4
7
5
_
30
-
r
25
30
+
2
5
3
+
T
GRASS SPP. •
Aqropyron repens
Aqropyron smithi i
Brbmus inermis
Oalambvilfa Ib n q ifo lia
Elymus canadensis
MuhTehberqia racemosa
Poa spp. " " 1
Pba palustris
Poa riratensis
Setaria v ir id i s
2
12
+
+
—
5
+
_
+
5
+
+
2
8
3
3
+
+
17
3
2
5
3
_
3
2 ' 2
2
5
+
3
7
+
+
-
-
-
-
-
+
T
+
2
-
FORB SPP.
Achillea m illefolium
Ambrosia psilostachva
Ascleoias soecibsa
Asclepias' v e r t i c i l la ta
Asparaqiis o f f i 'c i nalis
Aster ericbides
'
Aster pahsus
+
_
+
3
7
+
+
_
__
_
_
+
+
_
_
Tf
-
-
,
3
No data fo r dead shrubs fo r the transects with dashes.
-
+
-
74
TABLE 14— Continued
Location number
Cirsium arvense
Conyza canadensis
Euphorbia esula
Glycyrrhiza Iepidota
Helianthus p e tio la ris
Meli lotus o f f i c i n a l i s
Physalis heterophylla
Polygonum avicuTare
Ratibida c o lumnifera
Sisymbrium lo e s e lii
Smilacjna s t e lla t a
Solidago gigantea
Viola canadensis
14
2
6
2
I
17
16
+
+
2
-
14
11
9
- • -
T
Present
+
f
75
TABLE. 15
■
Presence (%'Coverage) of Species in the Grassland Community Type
Location number
Bare groupd
L itte r
6
17
7
67
2
I
I
9
14
11
16
45
82
45
38
5
40
53
45
12
38
4
+
-
-
_
I . 2
2
+
I
10
+
+
-
_
_
+
I
T
+
+
+
-
-
-
Present
SHRUB AND VINE SPP.
Artemisia cana
Artemisia dracunculus
Artemisia friq id a
Artemisia Iudoviciana
Coryphantha Viv ipa ria
Opuntia f r a q i i i s
Parthenocissus quinquefolia
Rosa woodsi i
Symphbricarpos occidental is
Toxicodendron rydberqii
2
-
2
_
■
3
+
+
+
+
3
-
-
-
_
-
-
+
+
+
+
-
-
+'
I
+
-
2
+
-
-
•T
+
+
_
GRASS SPP.
10
3
3
n
5
15
_
+
15
10
-
+
+
2
+
+
- ■ T
+
+
_
5
+
_
10
_
_
+
20
_
+
2
20
7
8
+ 28 10
_
— _
_
+
+
5
+
+
+
T
+
— _
_
_
_
_
■
+
_
_
7
2
+
45
2
12
2
13
8
8
35
rr
+
+
+
30
+
+
-
+
+ I
Aqropyron cristatum
Agropyron repens
Aqropyron smithi i
Bouteloua q r a c ilis
Bromus Inermist'
Bromus .iaponicus
Calamovilfa lo n q ifo lia
Carex brevior
Echinochloa crusqalI i
Elymus canadensis
Muhlenberqia racemosa
Panicum c a p illa re
Poa spp.
Poa palustris
Poa pratensis
Setaria v ir id i s
Spartina q ra c ilis
Sporobblus cryptandrus
Stipa comata
Stipa v ir id u la
Vulpia octoflo ra
+
2
5
18
+
+
+
2
+
_
_
+
FORB SPP.
Achillea m illefolium
Amarantbus powelli i
Ambrosia psiTostachva
Arabis ho lboelii
AscTepias speciosa
+
-
-
-
+
-
+
-
-
+
T
_
+
+
r-
+
-
-
+
76
TABLE 15—Continued
Location number
Aster oblongifolius
Aster pansus
Chenooodium a l bum
Chenooodium leptoohyllum
Conyza canadensis
Euphorbia esula
Glycyrrhiza lepidota
Grindelia squarrosa
He!ianthus p e tio la ris
Heterotheca v lllo s a
Iva a x i l l a r i s
Kochia scoparia
Lactuca pulchella
Laooula echinata
Laooula redowskii
Leoidium densiflorum
Linum oerenne
Medicaoo sativa
MeliIo tu s aIba
MeliIo tu s o f f ic i n a l is
Mentzelia nuda
Nepeta cataria
Oenothera biennis
Physalis heteroohvlla
Polygonum aviculare
PsOralea argoohvlla
Ratibida columnifera
Rumex maritimus
Salsola kali
Sisymbrium altissimum
Sisymbrium lo e s e lii
Smilacina s t e lla t a
Solanum rostratum
Solidaoo gigantea
Solidago mol I is
Thlasoi arvense
Tradescantia occidental is
Traqoooqon dubius
Veronica americana
6
17
+
2
+
-
2
I
I
9
14
11
16
Present
+
—
+
—
2
-
7
+
—
-
+ .
r-
5
+
+
12
-
+
+
+
2
+
+
+
+
+
+
+
7
-
-
+
+
77
TABLE 16
Presence {% Coverage) q f Species in the Green Ash Community Type
Location number___________
4
5
15
3
Bare ground
L itte r
-
-
53
58
10
55
—
20
+
+
20
+
+
+
70
+
+
-
-
-
+
+
+
84
3
10
7
2
28
27
47
17
27
65
+
75
+
T
-
-
75
+
+
-
-
-
+
+
+
+
+
■
-
Present
-
TREE SPP.
Elaeagnus angustif o l ia
Fraxinus pennsylvanica
Juniperus scopulorum
PopuTus~deltoides
Sa lix amygdaloides
Salix f l u v i a t i l i s
Ulrnus americana
80
-
r
-
SHRUB AND VINE SPP.
Acer negundo
Amelanchier aI n i f o lia
Artemisia ludoviciana
Cornus stolo nife ra
Cotoneaster sp.
Parthenocissus quinquefolia
Rrunus virginiana
Rhus tr ilo b a te
Ribes aufeum
Ribes setosum
Rosa woodsi i
Shepherdia argentea
Symphoricarpos occidental is
Toxicodendron rydbergii
V it is r ip a r ia
T
-
+
-
+
+
-
-
-
-
2
I
+
I
2
5
7
I
I
-
28
3
2
+
+
+
-
-
_
-
_
-
-
+
+
3
-
-
-
+
-
‘ -
-
_
-
_
2
+
+
-
-
—
-
-
I
-
-
-
T
-
-
I
11
+
16
2
+
+
+
+
+
-
-
7
-
10
+
I
-
3
2
17
2
8
5
+
-
+
-
-
,
-
+
I
6
-
-
-
-
_
-
I
-
6
+
-
-
-
-
-
-
-
_
_
GRASS SPP.
Agropyron repens
Agropyron sm ithii
Agrostis^aTba
Bromus inermis
Bromus japonicus
Carex brevior
Carex lanuginosa
Elymus~canadensis
MuhTenbergia racemosa
Poa spp.
Poa palustris
Stipa v irid u la
3
5
5
T
3
7
+
+
-
-T 3
2
T+
12 69
- ■ -
+
-
+
2
_
5
5
-
10
-
- ,
15
2
18
-
-
+
-T
78
TABLE 16— Continued
Location number
4
5
15
3
3
10
7
+
+
3
_
+
+
Present
FORB'SPP.
'
_
_
-
_
_
_
-
-
_
_
+
+
"
-
-
T
_
-
-
+
5
_
3
_
+
+
-
+
+
_
_
_
_
-
-
_
_
_
3
-
-
-
_
■
■
2
+
■
-
-
-
+
_
-
-
-
-
-
-
-
_
-
-
-
-
+
_
3
3
-
3
-
5
_
+
-
+
_
_
+
-
-
-
F-
CM
+
+
,
+
2
+
+
+
+
•7
.
I
Achillea m illefolium
Asclepias v e r t i c i l la ta
Asparaqus o f f i c i n a l i s
Aster pansus
Chenopodium album
Cirsium arvense
Convolvulus sepium
Euphorbia esula
Galium triflo ru m
Glycyrrhiza lepidota
Helianthus p e tio la ris
Lactuca s e r r i o l a
Lappula redowskii
Medicaqo lupulina
Medicaqo sativa
Melilotus alba
MeliIo tu s o f f i c i n a l is
Mentha' arvensis
"""
Oenothera 'al bicaul is
Plantago major
Rorippa sinuata
Smilacina s t e lla t a
Solidaqo qiqantea
Taraxacum Ia e v i qatum
Thalictrum venulosum
Tragopoqon diibius
Verbascum thapsus
Viola canadensis
10
+
+
+
+
+
2
-
79
TABLE 17
Presence (% Coverage) of Species in the Willow-Shrub,
Peach-Leaved Willow and Marsh Community Types
Community type
Willow-!Shrub
Location number
I
_
38
33
50
45
43
_
-
20
_
15
+
+
25
+
60
_
-
28
9
I
+
+
_
_
_
_
-
23
23
-
Bare gound
L itte r
10
14
Peach-leaved willow
Marsh
14
.
TREE SPP.
Elaeagnus a ngu stifolia
Fraxinus pennsylvaniea
Salix amygdaloides
SHRUB AND VINE SPP.
Artemisia I udoviciana
Cornus stolo nife ra
Parthenocissus quinquefolia
Prunus virginiana
Ribes setosum
Rosa woodsi i
Sal ix "fT'uviatil is
Symphoficarpos occidental is
Toxicodendron rydbergii
V itis r i pari a
2
2
+
+
6
9
3
+
21
2
2
2
■ I
+
7
+
8
+
+
8
5
5
-
2
-
28
-
-
5
5
2
-
-
-
I
-
GRASS SPP.
Agropyron repens
Aqrostis alba
Beckmannia syzigachne
Bromus ine'rmis
Carex lanuginosa
Echinochloa crusgalIi
Eleocharis palustris
Elymus canadensis
Muhlenberqia racemose
Panicum c a p illa re
Phalaris arundinacea
Poa palustris
Scirpus americanus
Scirpus validus
-
2
5
-
-
-
-
-
-
3
-
3
-
+
+
-
2
3
-
-
-
-
-
-
-
-
FORB SPP.
Arctium lappa
Asclepias speciosa
Asclepias v e r t i c i l la ta
Asparagus o f f i c i n a l i s
_
+
+
~
—
+ - ’
+
_
+
+
+.
'
7
-
+
-
-
-
-
-
-
-
2
-
-
-
+
-
r,
80
TABLE 1 7 - - C o n t i n u e d
Community type
Willow-Shrub
Location number
■I
Aster pansus
Cirsium arvense
Convolvulus sepium
Conyza canadensis
Equisetum laeviqatum
Galium triflo ru m
Glycyrrhiza lepidota
Helianthus p e tio la ris
Iva a x i l l a r i s
Medicaqo sativa
Melilotus o f f i c i n a l i s
Mentha arvensis
Plantaqo major
Polyqonum coccineum
P o te n tilla paradoxa
S a q itta ria cuneata
Sm ilacina. s t e lla t a
Solidaqo mollis
Solidaqo occidental is
Sonchus asper
Trifo liu m fraqiferum
Viola canadensis
Peach-leaved Willow
10
14
14
2
+
12
2
+
2
+
+
+
+
+
-
-
-
+
-
-
-
-
_
-
_
-
+
_
+
-
2
-
-
-
-
-
-
-
12
5
2
+
+
_
-
-
-
-
-
_
-
-
_
-
-
_
-
-
-
-
-
-
+
-
-
-
_
-
— ,
8
-
Marsh
-
-
-
-
-
-
-
2
_
+
+
+
+
+
+
+
+
TABLE 18
Density (number/10 m ) of Trees, by Diameter Class , in the Seedling and Thicket Community Types
Community type
Location number^
Original plot -size
Species
Pode^
Saam
Seedling
8
17
1.5 1.5
16
1.5
9
1.5
12
1.5
13
1.5
573
-
520
533
807
207
-
-
-
-
-
-
-
-
-
-
-
-
-
-
2
„1.5
6
1.5
14
1.5
153
673
207
660
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
16
12
8
18
17
18
Thicket
2 14
18 18
13
18
12
18
9
18
18
6
2
I
3
I
77
13
5
3
D ia.Cl ass(cm)
0 -0.5
0.5-1
UZ
2-3
3-4
4-5
5-6
0-0.5
0.5-1
1-2
2-3
D
-
-
_
58
15
6
I
I
14
3
3
I
. mm
89
2
-
33
2
I
-
38
17
-
_
-
I
I
•
20
-10
40
60
_
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
30
-
90
-
10
-
10
-
20
84
2
I
-
I
I
-
-
I
-
6
-
-
-
-
I
/I
-
2
I
-
4-5
Safl
0-0.5
0.5-1
1-2
2-3
3-4
190
'
70
'
210
-
10
-
170
—■
7
I
-
90
12
2
-
15 156
50 64
26
3
2
-
93
12
_
-
86 92
24 . 9
2
-
I
I
I
I
46
2
2
-
Diameter classes from 0 cm to 6 cm are basal diameters and diameters 6+ are diameters a t breast
^height.
^Locations are id e n tifie d in Table 5.
Species are Acer negundo (Acne), Elaeagnus angustifol ia (E la n ), Fraxinus pennsyl vanica -(Frpe).
Juniperus scopulorum (Jusc), Populus deltoides (-Rode), Salix amygdaloides (Saairi), Salix f l u v i a t i l i s
( S a f i ) and Ulmus amerlcana ( Ulam).
~
'
TABLE 19
Density (number/1000 m2 ) o f Trees, by Diameter Class, in the Young
Cottonwood and Mature Cottonwood Community Types
Community type
Location number
Original plot size
Species
Acne
Elan
Frpe
Jusc
Pode
Saam
9
108
14
75
16
75
-
_
_
_
_
Young cottonwood
8 18
2 12
120
75
92
75
13
84
6
75
2
1200
Mature cottonwood
2
14
12
16
360 1200 1200 1200
11
1200
D ia .Class(cm)
4-5
6-10
0.5-1
1-2
2-3
0-0.5
1-2
2-3
6-10
20-30
1-2
6-10
10-20
20-30
6-10
10-20
20-30
30-40
40-50
50-60
60-70
0.5-1
1-2
2-3
4-5
5-6
6-10
_
_
_
27
13
_
-
-
-
13
-
-
_
_
-
-
_
_
_
_
-
-
_
-
40
-
_
-
_
—
49
100
19
-
■
_
_
_
-
-
-
I
I
-
-
5
-
-
-
_
-
-
-
-
-
-
2
-
-
-
-
-
-
-
-
-
-
4
-
-
-
-
-
-
-
-
108
87
21
-
_
_
-
-
45-9
-
_
-
-
-
17
2
2
-
-
13
106
27
_
-
4
-
-
_
_
-
-
50
7
I
_
I
OO
rv>
3
_
_
-
-
_
_
_
53
120
-
-
_
_
_
-
-
-
3
-
175
—
-
_
I
5
-
12
319
160
-
-
120
101
20
80
53
106
-
-
-
-
-
-
-
-
-
-
-
-
-
-
_
I
I
8
8
3
12
3
—
I
_
I
2
7
8
2
9
7
3
-
_
_
-
-
-
-
-
-
_
-
15
5
-
-
-
-
-
-
-
-
-
-
-
-
-
—
-
-
-
-
-
-
-
-
-
-
+
I
I
I
2
-
-
1
3
6
11
3
2
2
2
I
2
2
2
I
TABLE 20
p
Density (number/10 m ) o f Trees, by Diameter Class,
i n the Shrub Community Type
Community Type
Location number
Original plot size
Species
Frpe
14'
15
6
15
2
15
I
14
Shrub
17
16
14
14
14
30
11
30
Dia.Class(cmj
0-0.5
0.5-1
5
-
I
-
1-2
-
-
-
2-3
4-5
6-10
I
2
I
-
-
-
-
I
-
I
—
I
I.
-
28
8
-
-
9
30
TABLE 21
9
Density (number/1000 m“ ) of Trees, by Diameter Class, in the Green Ash,
Willow-Shrub and Peach-Leaved Willow Community Types
Community type
Location number
Original plot size
Species
4
360
Green ash
3
3
240 240
10
360
7
284
-
_
-
-
-
-
-
-
—
-
206 1289
38
8
76 105
34
42
21
-
120
3
20
25
14
11
17
3
-
5
160
15
360
_
-
-
3
11
Elan
0 -0.5
0.5-1
3-4
-
-
3
-
Frpe
0 -0.5
0.5-1
1-2
. 2-3
3-4
5-6
6-10
10-20
20-30
30-40
218 3093
_
25
_
_
6
12
32
14
95
14
6
3
-
118
I
-
-
-
-
3
0.4
0.4
+
- -
-
-
+
•:
Jusc
Ppde
Peach-leaved willow
14
120
Dia.Class(cm)
0-0.5
6-10
Acne
Willow-shrub
I
10
14
HO
60 220
0 -0.5
6-10
10-20
4
-
- I
-
6-10
+
+
-
-
8
101
3
-
-
11
-
7
18
32
25
91
53
-
-
-
-
-
. -
-
9
5
-
3590
18
27
-
170
-
104
36
18 '
-
75
—
-
-
-
-
-
-
-
-
TABLE 21 - - C o n t i n u e d
Community type
Location number
Original plot size
Species
Saam
Safl
Olam
4
360
5
160
15
360
Green ash
3
3
240 240
10
380
7
284
.
Willow-shrub
I
10 14
HO 60 220
Peach-leaved willow
14
120
D ia .Class(cm)
0.5-1
1-2
2-3
3-4
4-5
5-6
6-10
10-20
20-30
30-40
0.5-1
1-2
■0-0.520-30.
_
_
_
-
—
-
-
_
-
-
-
_
_
_
_
-
-
_
-
_
-
-
-
-
-
_
_
_
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
_
-
-
-
-
-
-
-
-
-
-
■-
109
-
6
11
22
-
-
1285
4
—
.
+
-
18
45
27
36
189
117
136
■425
323
221
34
17
51
102
36
126
90
63
18
18
9
-
-
-
-
-
-
-
-
-
-
-
-
-
4
—
-
36
32
-
—
-
8
-
42
16
-
-
TABLE 22
O
I
,Density (number/100-m ) o f Shrubs, by Canopy Diameter Class , in the Young Cottonwood
and Mature Cottonwood -Community Types
Tlommunity type Locati on number
O rig inal p lo t size
Species
9
120
14
90
16
120
-Young cottonwood
12
2
8
18
37 L60
60.
28
13
46
6
120
I
15
Mature cottonwood
2
14
12
16
15
15
30
15
11
20
Size Class(dm)
Amal ^
0-1
-
-
-
-
7
-
-
-
-
-
T-
-
-
-
-
Cost
0-1
1-2
O
L-OO
3-4
5-6
-
37
2
-
14
- '
~
-
-
-
-
-
3
-
-
+
-
“
-
-
-
-
-
-
-
39
8
4
2
4
-
-
-
-
-
-
-
_
-
_
-
_
-
Rhtr
Riau
0-1
1-2
2-3
345-6
0-1
1-2
2-3
3-4
4
5
-
-
-
I
-
4
-
-
-
-
-
-
-
10
-
-
-
-
97
-
-
-
-
-
-
-
-
-
-
60
20
13
13
7
.7
207
20
7
-
6
-
-
-
7
-
-
10
10
-
-
-
-12
9
-
■*"Canopy diameter is the average o f the widest crown measure and the crown measure perpendicular to
p it a t it s m idpoint.
-locations are id e n tifie d in Table 5.
dSpecies a re Amelanchier a ln if o lia {Am al), Artemisia iCana (A re a ), Artemi si a dracunculus (A rd r),
Artemisia ludovicfana (Arl-u), Cornus s to lo n ife ra .{-Cost), Cotoneaster sp. (Cosp)-, Pranus
virg in ian a (P r-vi), Rhus tr ilo b a te (R h tr ), Ribes atireum (R ia u ), Ribes setosum (Rise) , Rosa
woodsi i (Rowo) , Shepherdia argentea (Shar) and Symphoricarpos occidental is (Syoc).
TABLE 22--Continued
-Gommunity type
Location -number
O riginal ^ lo t s iz e
Species '
Rowo
9
120
14
90
Y-oong cottonwood
8
18
2 J12
60.
28
37 IGO
16
120
13
46
6
120
I
15
• Tlature cottnnwnnrl
2
14
12
16
15
30 45
15
11
20
Size Class(dm)
.0-1
1-2
2-3
3-4
4-5
5-6
+
—
-
' +
r-
4
4
“
-
— ■ -
7-8
9-10
-
10
8
—
-
Shar
0-1
-
I
-
.
-
-
Syoc
0-1
1—2
2-3
3-4
3
I
-
4
I
-
-
217
27
-
218
28
D-/
5
-
-
39
2
-
_
2
-
113
- 227
- 180
40
13
—
-
-
' -
-
_
41
. —'
-
102
41
4
_
-
400
234
20
7
200
80
47
13
-
30
40
47
43
13
3
7
-
874 286
47
67
- ■ 47
10
127
87
40
27
7
-
20
60
20
20
_
-
145
100
35
40
15
10
5
5
5
634
153
7
-
980
465
80
15
_
374
60
_
-
TABLE 23
Density Xnumber/100 m )■ of "Shrubs, by "Cajiapy Diameter -Class, in the "Shrub
and Grass!andXommunity Type's.
Community type
lo c a tio n number
O rig inal -p lat size
Species
'
Rhtr
Ri.au
Rise
Shrut
17
16
15
15
14
30
11
30
9
30
-
-
_
-
_
-
_
-
6
6
6
-
-
7
-
*-
-
-
-
-
3
-
-
-
-
-
-
-
-
-
6
15
2
15
I
■ 15
-
-
-
-
.-
-
-
-
-
-
Grassland
2
6 17
I
I
9
120 12-0 120 120 120 120
14 11
50 120
16
35
14
2
9
4
6
I
I
83
"6
Size Class(dm)
Area
Ardr
14
15
D -I
1-2
2-3
3-4
4-5
5-6
6-7
7-8
0-1
1-2
234-5
+
-
-
67
47
39
8
3
2
2
“
-
_
-
-
—
—
-
3
4
0-1
1-2
2 -3
3-4
-
-
-
-
-
-
0-1
1-2
2-3
27
.13
-
47
20
-
-
7
13
7
-
13
-
0-1
-
20
-
-
-
7
_
—
6
9
7
-
9
20
9
5
_
2
2
2
2
-
20
20
12
6
-
15
15
3
-
3
-
_
_
—
_
_
-
‘ 7
_ -
-
-
-
“
—
-
T-
-
-
TABLE 2 3 - - C o n t i n u e d
Community -type
Location number
OrTganaJ p lo t szaze
Species
Rowo
14
6
15 ■ 15
2.
15
I
15
Shrub
17
16
15
15
14
30
Grassl an cl
6 1 - 7 2 1 1 9
120 1-20 120 120 120 120
11
30
9
30
293 160 123
-93 213 . 103
153
67
63
13
50
20
7
5
3
7
3
3
3
-
210
77
27
3
3
28
15
-
534
53
13
586
220
33
+
-
14 11
.50 120
16
35
Saxe Glass (dm)
0-1
1-2
2-3
3-4
4 -5
5-6
6-7
400
354
73
13
-
-
-
-
-
-
-
-
-
Shar
0-1
—
I
Syoc
0-1
1 -2
2-3
"3-4
4 -5
5-6
—
454
293
87
7
-
280
200
107
73
20
2828 2428 1067
227 260 460
20
—
—
-
-
—
—
-
-
133
-
67
153
187
40
27
340
347
153
20
7
880 1834
133 153
46
-
-
-
—
-
-
-
-
-
-
793 1272
190 240
37
43
10
20
13
3
-
-
+
- ' -
7
-
-
f
—
-
53
29
-
-
30
23
19
"5
61
17
+
2
76
26
2
7
4
-
69
9
-CO
LO
TABLE
-2 4
p
Density .(number/100 m ) o f .Shrubs, by Canopy Diameter Class, in the Green Asli
Uj I Iow-Slirub and -Peach-Leaded4}i11 ow Community Types
Community type
LocatTon number
O riginal p lo t size
"Species
-Araal
Gost
Co sp.
Prvi
Rhtr
-Riau
Green ash
3
3
120
70
4
15
5
15
15
39
-
7
8
-
- .
—
—
—
r
-
20
-
r
—
-
-
15
2
-
7
7
7
7 . .13
-
-
C -I
1-2
2-3
+
—
-
7
7
-
_
-
C -I
1-2
4-5
-
7
7
7
_
-
TC
15
7
17
-
_
-TIil I ow-shrub
I
10
14
15. 15
15
"Peach-Iaaved wi 11 ow
14
15
Size Class(dm)
0-1
1-2
0-1
1-2
2-3
3-4
6-7
7-B
C -I
I -"2
0-1
1-2
2-3
-
.
.
,
-
-
-
7
14
I
40
7
14
-
-
-
-
54
-40
33
-20
27
-
-
-
-
_
—
-
-
-
■ _
-
'
-
-
-
—
-
—
—
.
_
_
-
-
-
_
■~ . . _
_
--
-_
-
—
4_
—
-
—
_
-
_
TABLE 24 — C o n t i n u e d
-Community type
Location .number
O riginal p lo t s3ze
Species
Rise
-0-1
23-
Syoc
5
15
-20
67
13
34
34
13
i5reen a-sb
3 3
TG
12-0
70
15
15
39
7
17
Hi 11 ow-sTi rub
I 10
14
15 15
15
Peach-leaved -willow
14
15
Size Class(dm-)
1-2
-Rowo
4
15
3
4
-0-1
1-2
2345-6
6-7
7-8
0-1
1-2
23-
3
4
5
-
3
4
-T-
7
7
3
27
40 1487
20
67
20
40
7
7
194
844
214
20
067
523
94
.7
3
20
18
'
34
30
6
-
80
114
73
34
77
142
34
94 370
25 536
-3
87
13
133
165
59
13 101
80. 174
60 223
13
60
13
27
13
1005
1-61
30
7
101
54
27
27
54
47
20
7
47
114
80
20
7
40
47
27
34
7
7
7
7
20
13
to
9%
T W E 25
Constants
and E s t i m a t e s o f R e l a t i v e E r r o r ? f o r R e g r e s s i o n s
R e l a t i n g L o g a r i t h m Pl a n t , W e i g h t s t;o T h g i r L o g a r i t h m P i a m g t e r s
Qiamgter
basis
Crown qapopy
Crown canopy
Crqwn ganppy
Trunk base
Syoc
Area
Safl
Leaf
b
m
r
E
1.46
0.23
0.93
X.73
1.44 1.59
0.30 ■ 0.47
0.92 0.86
1,73 9,07
b
m
r
E
1.48
0.18
0.94
1.37
1.59 1.93
0.3-1 -0 .0 9
0.98 0.95
1.20 1,30
b
m
r
E
E
2.31 3.21
0.43 -0.02
0.95 0.97
1:63 1.99
2.136 ’
0,9$%
T
eT
Q,?9
1,39
b
m
r
E
2.09
0.93
0.98
2.46
b
1P
r
E
2.53
q,7§
0.99
I . $7
b
m
r
•E
%.53$
-1.29
0.97
1.27
b
m
r
Trunk base
Tmnk be so
DBH
Saam
Pode
Rode
Woqd
$
cm
Wood
O r, 5
cm
Cqn?tant$
:Species .+ e rro r
W
0.46
0,99
1.33
'
_
-
=■7
-
-
T
r
-
-
7
I
FT
7
-
-
r
r
-
-
-
•
-
r
-
-
-
-
-
r
-
Wood
o -l
cm
Wood
InlO
cm
Wood
10+
cm
$pqts
Icm+
•7
T
T
T
--
-
-
-
_
-
T
r
7
TT
■
rr
T
-
r
_
H
r
-
-
I
JT
'T
2.43
0.89
0.97
2.69
3.35
1,96
0,92
1.56
2.22
1.0$
0.97
2.99
2,9$
0.91
0,92
1.75
2.38
0,88
0.98
2.33
2.85
0.87
0.99
1.09
1,93
-1 .7 8 '
0.99
1.24
_
-
T-
-
_
TT
_
-
-T
r
-
-
r
'T *
TT
-
p
-
-
7
ir
-r
Tf
r
-
_
T
r
-
1,87 2.83
-0 .8 5 -1.67
0.99 0.99
1.18 1.18
2.33
0.62
0.99
1.11
2,45
1.14
0.99
1,41
i
2m slopg of thg regression lin g , p = y in tercep t p f rggressipn l]n e .
Tr Tf c o rre la tio n c o e ffic ie n t. E s estimate o f r e la tiv e e rro r fo r a
logarithm ic regres$iQn. Ap p of 2 indicates th at the 'predicted value
31 i'es between one h a lf and twice the true value.
Plant species are Rosa woodsi i (Rowo), Symphoricargos occidental is
(Syqc), Artemisia cana TArqiJT S a lix f l u v i a t j l i s ( S a f i) , S alix
amygdalqides (Saami2
) "and Pppulus' del toides (Podg),
' ^
93
TABLE 26
2
Bulk Density (g/m ) of Soils from Three Horizons
in Seven Community Types1
SoilI
horizon
Community
type^.
K
L
0-10 cm
SA
SE
TH
YC
MC
SH
GR
0.80
0.90
1.20
1.08
0.97
0.95
0.90
1.05
1.Q5
1,89
1:31
1,31
1.55
1.10
1.25
1.25
1.24
0.99
1,21
1,29
1.19
1.25
1.25
1,23
149
l.io
0.97
1.15
1,23
1.23
1.07
1.26
1,12
1.02
1.05
10-30 cm
SA
SE ■
TH
YC
MG
SH
QR
1.42
1.42
0,79
1.09
1.28
1,31
1.00
0.95
0.95
1.37
1.12
0.99
0.99
1,29
0.98
0.98
1 .04
1.02
Q.9?
1.34
. 1,29
1.04
1.04
1 . 19
1.19
1.04
1,19
1.45
1.15
1.15
149
1.15
1.051.07
I rI l
?0-150 qm
SA
SE
TH
YC
MC
SH
GR
0.87
0.87
0.71
1.06
.1.09
1,10
0.84
0,7%
0.78
1.11
0.68
0.99
1,14
1,20
1,23
1.23
1.08
1.16
1.06
0.9?
1.15
141
141
1.04
1.12
1.06
143
149
1,29
1:20
0.92
0,90
0.98
1.16
1.23
•
Location'3
N
M
Il / I
1•' J
0
^Bulk densities were estimated hy weighing dried samples from fiv e
pooled cores and dividing by the volume of the coring tube segment.
Weights o f s o ils (gm/nr) in each horizon are calculated by m ultiplying
by the volume o f the horizon fo r 0-10 cm =? 100,000; fo r 20-3Q cm s
200,000; fo r 30-15 cm = I , 2 0 0,Q00'.
^Community types.are sandbar (SA), seedling ( 5 | ) , th ic k e t (TH ), youncj
.,cpttontyood (YC), mature cottonwood (MQ), shrub1(SH) and grassland (GR).
^Locations are id e n tifie d in Table 5:
94
TAPLE 27
Elemental Consents (%) o f Riparian Plants o f
the Lower Yqllowgtone River1
P
Species^
Part^ Number 4
Pod^
If
■ 0-1
1-10
10+
Safl
Rqwq
$yoc
Area
If
XiSD
Ash
XiSD
0,20+0,04
1.32+9.03
9,P3±P.P1
9±1
0.0810.00
0.4610.08
0,03+0.01
5 il
PrOOiO1Pt
O.Q4+0.02
4+2
0.0610.05
3 il
XlSD
5
1 . 64+p49
5
0 .42±0.11
.5
Na
XlSD
X1
IbSDp
4 ,
4
E lements
_ K
0.07±0.02
0.01+0.01
0 ,3 0 *0 4 3
0.1810.04
2 46±P.39
0.28±prp,3
1,2410.28
0.0510,03
Oil
0.65+0.29
0.0810.03
0.58+0.31
0.0310.01
3 il
0-1
5 .
HO
if
3
. 0,34+0.17
0 .04±01:00
0,20+0,00
9 .0 2 + 0 .QO
2i l
P
$34040
0.2510.06
p.35+0.06
0.02+0.00
6+1
0 -0 .5
5
0.55+0.11
0.07+0.03
0.1410.04
0.0210.00
3i0
0.5+
5
0.32+0.07
q.Q4+Q.Q0
0,0 7 + 0 .Ojl' 9r0jl±O.OQ
2*1
If
5
1.65±0.18
0.23+0.08
1.7010.20
0.0210.00
7+1
OrrO. 5
0,48+0.06
0,05+0.01
9.99+0.11
0 ^ 2 *9 ,9 0
4+0
0.5+
§
4
■0.3510.08
0 0410.01
:
0.7010.11
O.OliO.OO
3l0
If
5 ’
1.8410.38
0 25 0 . 0 4
, +
4.64+0 4 3
9 .0 2 + 0 .QO
7*1
0 -0 .5
5
0.84+0.96
Q.0910.02
1.05+0,41
0.03+0.01
5 il
0.5+
4
0.53+0.08
0.05+0.01
0.47+0.18
0.02+0.00
311
Methods used fo r determining elemental content? ere described in
pBremner (.1965,' N ), Olsen and Dean (1965, P) and P ra tt (1965, K and N a),
species studied were Populus deltoide? ( Pode) , S alix f l u v i a t i l j s
( S a f i) , Rosa woodsi i (Rowo), Symphoricarpps occidental i s ' (Syoc).
Artemisia' cgna (A rea), Toxicodendron rydbergii (TO ry), vine and 1A r t e - •
,,mi?ia ludoviciana (ArluTT
dPlant parts analyzed were leaves ( I f ) and ty/igp of the dimensions
,s p e c ifie d : 0 -0 .5 cm, 0.5+ cm, 6-1 cm, 1-10 cm, and over 10 cm.
gThe number o f transect? examined.
dMpan 4 standard d e v ia tio n .
95
TABLE 2 7 - C p n t i n u e d
Species
Part
Number
• N
X"±SD
P
I+SD
E lements
K
I+SD
■
i+so
Ash
^+SD
1341
Tory
4.
Vine
2
0 . 60±0.14
Arlu
,5
1,42+0,50
SE6
3
0.99±0.35 ■0.25±0.02
1.91+1.10
0.0840.02
TH
5
0.98+0.05
0.17+0.03
1.09+0.55
0,01+0
7+?
YC
5
1.00±0.03
0.17+0.03
0.91+0.41
o.oi±o
8+1
MC
3
1,00+0.09
Q,14i0.O5
0.61+0.06
0,0140
74%
SH
4
■1.00±0.01
0.12+0.04
0.62+0.30
O.pl+o
741
GR
5
0.98+0.06
0.10+0.05
0.49+0.25
0.0140
74%
SE
?7
1.08+0.25
0,3040,06
Q,pB*Q.Q6
0.1340.07
114%
TH
3
2.01+0.65
0.23+0.00
0.81+0.03
0,0640.04
15+3
VQ
4
0 . 19±Q.05
9,45+p.08
0 .2 5 :0 . !I!
m
0,64 • '
0.05
0.50
0.04
5
0.6840,41
0.04+0,0%
Crass
forb
MQ
I1I t t q r
SH
4
1 ,9 2 + 0 .^
0.31+0,08
GR
3
2.01+1.33
0.13+0.13 ' 1.09+1.49
•0.024o!oi 11+11
SA
2
0.46+0
0.0440.01
Pj J H O .04
0 .0 9*0.06 15+10
SE
5
0.91+0.41
0.10±0.04
0.34+0.18
0.1040.12 19412
TH .
5
P1^BlO.I?
0. Q8+Ot 02
0,23+0.05' jp.02+9
14+%
YC
5
0.63±0.17
0.0740.0%
0.2040.02
0.02+0
1645
MC
4
0 .6 7 + 0 ,to
p .07 + 0 .Ql
0,25+0,03
9.02+0
2048
SH
5
0.55+0.11
0.0540.01
p.2740.09
0.0240
1244
GR
5
0.94+0.15
0.07+0.Q1
0.20+0.04
0.91+0
ig+2
1143
Community types are sandbar (SA), seedling (S E ), th ic k e t (TH), young
yCottonwpod (YC), mature cottonwood (MC), shrub (SH) and grassland ((?R),
Fqr nitrogen, tfie number q f Ramplqs tqstqd is two in th% needling and'
grassland community types.
96
TABLE 38
Orggnip M atter Qohtpnts (%) o f Sqilp from Thrpe,Hon'zpop
in Sqven Community Types!
Soil
horizon
Community
typp^ ____________ •
I
prlO cm
'U
‘ I JM' if
K
I 'I I I ' I'., 'JJj Hlj'' jl JJJ 'I JI1' I . «
SA
SE
TH
YC
<0.1
0.3
1.0
I,?
6 ,0 '
MQ
SH
GR
lOnSO pm
SA
SE
TH
YC
MC
SH
GR
30^50 ^
SE.
TH
YC
MC
SH
GR
4:6
■
.
3.1
<0.1
<0.1
I.?
1.9
2.0
2,2
Z-?
10,1
<04
0.6
0.9
r
1.5
LQ
L
<0.1
Q.4
<0.1
0,7
3 .Si
1.0
1.5
Location
M
3
0.2
L2
L6
2.3
1.8
1.2
2.0
<0.1
40,1
l.l
1.2
Q,;
L*
1.6
H
L 7 . 2.1 ■ ■
. L4
L
S
L4
L7
Q.6
IQ
,I .
<0 . 1
.Li
0.3
L
?
1.5
1.2
Q,5
0.8
1.1
1,5
1.5
Q,3
N
0
<0 , 1
L r?
1.2
0,8
2,2
2,2
3.3
40.1
1.4
L5
O r?
4-3
3:8
4.4
1.0
Q.5
1.3 . 1.6
1,5
1.6
0,6
LO
1.0
2,0
2i?
?.2
1.8
2.8
0,1
0.-2
0.2
L
Q
IP,I
0.2
0.6
0.9
L
0.8
1.2
0,9
LP
1Organic master contents were c p lp rim e trip a lly determined a f t e r dichror,
2mate oxidation (Sims an^ Maby J97QI).
Community types are sancjbar (QA), 's e a lin g
, th ig k e t (ilifi), young
-cottpnv/opd (YC), mature ppttqnwood (MC), shrub (SM) and grassland (GR).
■Locations are id e n t if ie d 'in Table 5.'
97
TABLE 29
Kjeldahl Nitrogen Contents (%) o f Soils from Three Horizons
in Seven Community Types-*Soil
horizon
•3
Community
type '
K
L
Location
M
■N
0
0-10 cm
SA
SE
TH
YC
MC
SH
GR
.002
.019
.050
.051
.253
.149
.138
.002
.023
.014
.034
.114
.035
.062
.030
.012
.063
.089
.062
.069
.074
.010
.028
.022
.029
.073
.101
.107
.012
.039
.064
.023
.169
.167
.202
10-30 cm
SA
SE
TH
YC
MC
SH
GR
.007
.006
.045
.068
.084
.103
.104
.012
.034
.009
.036
.062
.072
.055
.005
.036
.049
.07%
.083
.057
.075
.043
.040
.048
.037
.052
.097
.080
.028
.047
.062
.020
.069
.126
.163
.009
.002
.028
.040
.020
.036
.014
.055
.053
.043
.018
.012
.026
.028 '
.039
.053
.061
.023
.017
.013
.014
.053
.044
.019
.032
.012
.008
.025
.046
.033
.043
.039
30-150 cm
SA
SE
TH
YC
MC
SH
GR
■
-
.056
.034
I
pThe method i s d e s c r i b e d i n Br emne r ( 1 9 6 5 ) .
Community t y p e s a r e s a n d b a r ( S A ) , s e e d l i n g ( S E ) , t h i c k e t ( T H ) , young
, c o t t o n w o o d ( Y C ) , m a t u r e c o t t o n w o o d ( M C ) , s h r u b ( SH) and g r a s s l a n d (GR)..
^L o c atio n s a r e i d e n t i f i e d in T a b le 5.
98
TABLE 30 .
Total Phosphorus Content {%) of. Soils in Three Horizons
in Seven Community Types1
Soil
horizon
0-10 cm
Community
type
O
Location
K_______ L_______ M______ N
SA
SE
TH
YC
MC
SH
GR
.057
.05.2
,065
.071
.090
.080
.083
10-30 cm
SA
SE
TH
YC
MC
SH
GR
30-150 cm
SA
SE
TH
YC
MC
SH
GR
.046
.065
.045
0
.085
.055
.065
.059
.032
.073
.075
.073
.073
.073
.056
.064
.048
.059
.064
.074
.072
.052
.062
.068
.058
.074
.083
.088
.037
.053
.071
.076
.076
.071
.080
.045
.071
.052
.055
.076
.065
.065
.035
.071
.067
.063
.063
.063
.065
.072
.070
.066
.061
.060
.073
.071
.060
.068
.072
.052
.064
.074
.072
.050
.046
.061
.071
.065
.069
.063
.046
.069
.055
.073
.071
.073
.059
.041
.033
.062
.066
.068
.072
.060
.048
.054
.048
.068
.064
.048
.064
.046
.048
.062
.058
.064
.062
.064
gThe method is described in Olsen and Dean (1965).
Community types are sandbar (SA), seedling (SE), th ic k e t (TH), young
-cottonwood (YC), mature cottonwood (MC), shrub (SH) and grassland (GR).
Locations are id e n tifie d in Table 5.
i
99
TABLE 31
B i c a r b o n a t e E x t r a c t a b l e Pho s ph o r u s C o n t e n t s { % ) O f 1S o i l s
i n T h r e e H o r i z o n s i n Seven Community T y p e s 1
Soil
hdnzoh
Community
type^
K
L
Location
M
N
0 -1 0 .cm
SA
SE
TH
YC
MC
SH
GR
.013
.009
.015
.015
.038
.031
.029
.004
.006
.005
.012
.0004
.006
.011
.0004
.020
.020
.018
.016
.013
.003
.001 .
.003
.001
.007
.010
.011
10-30 cm
SA
SE
TH
YC
MC
SH
GR
.005
.017
.017
.020
.017
,020
.025
.005
.016
.006
.008
.023
.001
.008
.006
.009
.008
.010
.006
.006
.008
.001
.001
.005
.003
.001
.015
.011
SA
SE
TH
YC
MC
SH
GR
.010
.005
.008
.015
.012
.014
.015
.004
.023
.004
.020
.007
.009
.010
.008
.004
.006
.012
.001
.015
.010
.00003
.001
.004
.007
.007
.001 '
.007
30-150 cm
0
.008
.007
.011
.008
.021
.019
.025
.006
.007
.009
.002
.012
.013
. .015
.002
.002
.003
.001
.003
.001
.003
i
pThe met hod i s d e s c r i b e d i n P r a t t ( 1 9 6 5 ) .
Community t y p e s a r e s a n d b a r ( S A ) , s e e d l i n g ( S E ) , t h i c k e t ( T H ) , young
, c o t t o n w o o d ( Y C ) , m a t u r e c o t t o n w o o d ( M C ) , s h r u b ( SH) and g r a s s l a n d ( GR) . .
L o c a tio n s a r e i d e n t i f i e d in T a b le 5.
100
TABLE 32
Ammonium A c e t a t e E x t r a c t a b l e P o t a s s i u m C o n t e n t ( % ) o f S o i l s
f r o m T h r e e H o r i z o n s i n Seven Community T y p e s 1
Soil
horizon
Community
type^_____________________Kj______ L
Location3
M
N
O
0-10 cm
SA
SE
TH
YC
MC
SH
GR
.001
.005
.015
.014
.054
.032
.036
.002
.008
.004
.031
.011
.016
.008
.002
.021
.022
.021
.014
.029
.004
.009
.010
.012
.024
.022
.029
.003
.006
.028
.012
.044
.032
.051
10-30 cm
SA
SE
TH
YC
MC
SH
GR
.002
.003
.010
.018
.026
.020
.028
.001
.010
. .005
.010
.022
.015
.018
.001
.009
.012
.024
.018
.015
.016
.008
.010
.013
.012
.019
.027
.024
.006
.009
.016
.009
.026
.031
.039
30-150 cm
SA
SE ■
TH
YC
MC
SH
GR
.002
.002
.007
.011
.010
.016
.014
.004
.004
.005
.016
.021
.012
.007
.002
.001
.007
.010
.021
.019
.009
.003
.004
.004
.021
.014
.006
.010
.004
.004
.006
,011
.014
.012
.013
- T h e me t h o d i s d e s c r i b e d i n P r a t t ( 1 9 6 5 ) .
^Community t y p e s a r e s a n d b a r ( S A ) , s e e d l i n g ( S E ) , t h i c k e t ( T H ) , young
- c o t t o n w o o d ( Y C ) , m a t u r e c o t t o n w o o d ( M C ) , s h r u b ( SH) and g r a s s l a n d ( G R ) .
Locations are i d e n t i f i e d in Table 5.
101
.TABLE 33
Ammonium A c e t a t e E x t r a c t a b l e
from Three Horizons
Soil
horizon
Community
ty p e * '
0-10 cm
SA
SE
TH
YC
MC
SH
GR
•
Sodium C o n t e n t ( m e q / l O O g l o f S o i l s
i n Seven Community T y p e s 1
K
.1
.3
.1
.2
.3
.1
.2
10-30 cm
SA
SE
TH
YC
MC
SH
GR
.1
.1
.3
.4
.3
.2
.3
30-150 cm
SA
SE
TH
YC
MC
SH
GR
.1
.01
.6
1.1
.4
1.4
.4
.
3
L
Location
M
.01
.3
.01
.4
.2
.1
.3
.2
.9
.3
.5
.1
.3
.01
.2
• Q1
.1
.3
• 01
.1
.1
.5
1.2
.5
.3
.2
.3
.2
.3
.7
.3
.2
.2
.3
.3
.4
.9
.2
.5
2.0
1.4
.1
.4
.03
.6
2.2
.7
.1
.2
.1
1.0
1.7
2.2
.7
.3
.3
.3
.4
1.5
.4
.3
1.2
.2
.3
.8'
.4
2.2
4.1
2.2
N
0
.4
.3
.1
.01
.01
.1
.1
.2
.5
.3
.1
.3
.3
.3
pThe method i s d e s c r i b e d i n P r a t t ( 1 9 6 5 ) .
^Community t y p e s a r e s a n d b a r ( S A ) , s e e d l i n g ( S E ) , t h i c k e t ( T H ) , young
^ c o t t o n w o o d ( Y C ) , m a t u r e c o t t o n w o o d ( M C ) , s h r u b ( SH) and g r a s s l a n d ( G R ) .
L o c a t i o n s a r e i d e n t i f i e d i n T a b l e 5.
TABLE 34
2
Biomass ( g/m ) Present in ttie Sandbar and Seedling Community Types
Community type
Locati on
Component
K
L
Total Soil DM2
0—
10 cm ■
10-30 em
30-150 cm
14-12
80
283
1049
5932
105
191
5635
Total Roots
0-1 mnr
I -10 rnim
-
-
-
-
-
-
Total L it t e r
Total Herbs
Grass
Forb
Total S a fl5 " 6
Leaves
0-1 cm
Total Pode
Leaves
0-1 cm
Sandbar
M
8
-
1917
250
196
1471
-N
0
K
L
3537
125
2082
1330
2685
123
1150
1442
1572
240
283
1049
12853
422
2098
10333
--
218
90
128
2
-
-
-
-
44
-
-
-
-
-
-
—
»
-
.
-
-
_
_
_
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
■-
Seed! incj
M
N
0
5323
1500
2352
1471
6870
1502
2707
2661
8286
1-721
3681
2884
406
2%8
128
223
95
128
514
386
128
358
230
128
102
9
5
114
+
+
-
23
13
10
3
+
3
20
4
16
19
8
11
18
9
9
58
29
29
2
I
I
17
8
9
96
48
48
11
5
6
168
84
8.4
34
17
17
28
14
14
92
46
46
^Locations are id e n tifie d in Table 5.
Organic m atter {DM) c o lo rim e tric a lly determined a f t e r d ichromate oxidation (Dims and
^1970) included roots less than one centim eter in diameter.
^Soil horizons sampled included the depths 0-10 cm, 10-30 cm and 30-150 cm.
gRoots were separated into the diameter classes 0-1 mm„ 1 -1 0 -mm and 10 mm+.
Shrub wood was separated into the diameter classes 0 -0 .5 cm and 0.5 cm+; for tree or
gwood 0-1 em, 1-10 cm and 10 cm+.
V la n t species are Dymphoricarpos occidental is (Syoc), Rosa woodsi i -(Rowo), Artemisia
{A re a ), Toxicodendron rydbergii (T o ry ), Artemisia Iudoviciana (A r lu ) , S alix f l u v ia til
S a lix amygdaloides (Saam) and Popolus deltoides (Pode).
Haby
w illow
cana
is F T S a fl) .
TABLE
35
2
Biomass Xg/m ) Pres-ent. in tLie TLiickst and ifoung T)o-ttonwooti "Community Types
K
L
M
Total Soil -0M
O-^lO cm
10-30 cm
30-150 .cm
8246
1197
1908
5141
4748
188
547
4013
15226
1976
2923
10327
Total Roots
O -I mm
1-10 mm
10 mm+
1085
786
296
3
566
249
296
21
Aboveground Dead
-499
L it te r
499
Dead trees ■
-
T tic k e t
N
Young -cottonwood
M
N
0
K
L
7524
1476
3541
2507
12088
1-60-7
3821
6660
16962
1298
4174
11490
15436
918
2246
12272
20864
2288
3253
15323
24759
953
2383
21423
12580
377
1377
1-0826
617
254
296
67
708
392
296
20
1380
107-9
.296
■5
6298
540
558
5200
9531
1003
558
7970
13215
977
558
11680
5673
1335
558
.3780
6046
4960
"532
532
-
144
144
-
445
445
- -
315
"315
-
1559
612
947 .
-1779
514
1265
2060
1913
147
2287
301
1986
388
251
4
4
+
55
33
22
57
17
40
. 21
14
I
1.6
4
12
8
8
d8
42
6
45
38
7
_
_
_
-
_
3
+
+
+
I
•
I
I
11
4
5
2
+
+
12
4
5
2
.3+
2
Total Herbs
Grass
Fo r t .
34
34
-
17
15
2
Total Struts
Syoc leaves
0 - 0 .5cm
0.5 cm+
Rowo leaves
O-O-5cm
0.5 cm+
Arl u
13
-
-
-
-
_
-
-
_
-
-
_
-
-
-
-
+
-
-
+
-
"0
528
558
639
I
I
' 103
-.Community type
Location
Component
OBLE
-Gommtinj -ty type
.locatjon
Component
To tal Wjj low
5 aam loaves
D -I cm
I cm*
Safl lea-v.es
0-1 -cm
I -cm*
Jodaj Pode
I -eaves
0-1 cm
1-10 -cm
10 cm+
K
112
+
+
138
+
*
-
-
40
32
40
-18
5
6
7
-
Jtijcket
JJ
T
4r
-38
5.3
.
108
30
3839
-
35--Continued
Jj
0
125
I
2
' 4
■29
26
-63
206
*
+
725
-*
--*$■ ■
-559
141
159
.25:9
1-03
34
39
30
-
-
O
_
-
_
-
-59
50
97
"Young cottonwood
%
M
N
K
107 ;
1:74
35.5
39
10
13
11
-
__
-
. 12421
579
501
3420
7821
" 15230
43.7
• 486
3477
10830
26965
■ 891
991
6620
18463
-
8843
394
378
2675
5337
-
10122
628
533
3621
5340
TABLE 36
Biomass '{■g/m2 ) -Present in the Mature -Cottonwood and Shrub Community Types
Community type
Location
5Gompoflent
Mature cottonwood
M '
N
I
T<
43
K
26749
3883
6228
16638
"1609
854
755
-
1436
681
755
-
1549
794
755
-
2023
1268
755
407
407
-
867
867
-
380
380
882
882
684
684
39
29
10
8
4
4
22
16
6
14
14
5
I
4
5804
6148
15890
2417
2077
41396.
23203
4817
4240
14-146
"30020
4374
5759
19887
6967
641
946
4380
6758
542
"946
527-0
12052
' 1036
946
10070
8757
1411
946
6400
8876
IOlS
946
6920
1320
565
755'
-
Aboveground -Dead - 987
987
Li t i e r
Dead trees
2328
1074
1254
.
1232
795
437
1553
1237
316
632
609
23
I
I
-
8
8
+
-19
7
12
TotaT Tierbs
rfirass
Jorb
8
8 '
-
30
30
+
0
18237
. 2128
5218
10891
25487
2-184
4154
19149
Total Roots
0 -1 mm
1-10 mm
TO mm-+
Tl .
24493
20569
1541
1547 ■
5078
2594
17874
16428
25804
4574
3354
17879
"Total Soil OM
-Q-TD cm
10-30 cm
30-150 cm
Shrub
-"M
I
TABlTE 36 — Continued
Community type
lo ca tio n
Component
-Mature cottonwood
I
M
N
K
156
Total Shrubs
.24
Syoc --leaves
0 -0 .5 e-m 34
Q. 5 cm+
16
^Rowo leaves
20
0 -0 .5 cm 23
0.5 cm+
39
Tory
-Ar! u
-
83
7
10
5
15
17
30
24
+
44
8
11
4
5
6
10
18
-
Vines
29
-
-
Total Pode
I eaves
0-1 cm
1-10 cm
10 cm+
15.270
T64
27-6
1790
13040
15792
244
354
2337
12857
27003
381
571
3754
22297
"0
135
11
15
7
25
30
48
-
2
17270
226
■350
2287
14407
K
50
-4
5
2
TO
11
18
41
4
20755
303
450
2962
17040 -
113
21
29
14
12
14
23
;S:tirub
M
-I
-
87
■5
7
3
18
21
32
I
-
-
I
-
0
' 126
21
27
10
18
21
29
113
14
' 18
7
19
23
32
168
22
* 30
14
25
30
47
-
-
-
_
— .
_
--
_
_
-
_
-
_
-
N
-I
-
TABLE 37
■Biomass "t^/m ) Present i'n "the 5Gnassland Bommumty Type
Community type
Location
Component
K
L
Grassland
M
N
0
Total Soil BM
B-IO cm
TB-30 jcm
'3 0 -1SB -cm
17251
TT778
M l2
IBO 61
13034
1648
4181
7205
170-65
2372
3611
11082
-21902
3787
5214
12901
2558-1
4641
6228
14712
€B58
1.369
4B9
1252
763
489
1529
1840
489
1951
1462
489
1939
1450
489
Total L it t e r
185
455
152
506
385
Total Herbs
Brass
Ferb
114
108
6
154
132
2-2
144
124
-20
-114
114
+
147
147
31
+
+
+
7
13
11
23
I
2
I
2
3
3
11
4
I
I
64
I
2
I
5
12
13
30
47
Total Roots
B -I mm
.1-10 mm
Total Shrubs
Syoc
leaves
B-O .'5 cm
B . 5 -cm+
Area
leaves
B-O.5 cm
B .5 -cm+
Arl u
-
>
I
■ I
+
-
-
—
-
13
1-9
11
. 4
MONTANA STATE UNIVERSITY LIBRARIES
Condition Nntec;