Abrupt downstream forest decline following river damming in southern Alberta

STEWART
B. ROODAND SIG HEINZE-MILNE
Department of Biological Sciences, University of Lethbridge, Lethbridge, Alta., Canada T l K 3M4
Received August 5, 1988

ROOD,S. B., and HEINZE-MILNE,

S. 1989. Abrupt downstream forest decline following river damming in southern Alberta.
Can. J. Bot. 67: 1744-1749.
The influence of river damming on the abundance of riparian poplar forests was investigated by comparing forest abun-
dances on the dammed St. Mary and Waterton rivers of southern Alberta with the neighbouring undammed Belly River.
Forest distributions were determined by estimating the linear river distance of forests from airphotos taken in 1961 and 1981,
both upstream and downstream from the dams. During the 20-year interval, a 48% reduction occurred downstream from
the St. Mary Dam, which was completed in 1951. Downstream from the Waterton Dam, completed in 1964, riparian forests
declined 23 %. Poplar abundance along the downstream region of the undammed Belly River was relatively unchanged over
the study period, showing less than a 1% reduction. The upstream (undammed) portions of all three rivers underwent a slight
Can. J. Bot. Downloaded from www.nrcresearchpress.com by 99.243.105.24 on 01/09/15

forest decline during the 20-year interval; declines of 4.7, 4.6, and 6.1 % occurred along the St. Mary, Belly, and Waterton
rivers, respectively. Thus, the damming of two rivers in southern Alberta has been followed by a rapid and dramatic down-
stream decline in riparian forests. Further, the close association between the location and extent of the forest decline supports
a causal relationship between the river damming and the forest decline. Possible causes of the decline are discussed, including
drought-induced mortality, particularly of seedlings.

ROOD,S. B., et HEINZE-MILNE,

S. 1989. Abrupt downstream forest decline following river damming in southern Alberta.
Can. J. Bot. 67 : 1744-1749.
L'influence de l'endiguement des rivibres sur l'abondance des peupleraies d'aval a kt6 CtudiCe en comparant les abondances
forestibres sur les rivibres endiguCes St. Mary et Waterton du sud de 1'Alberta avec la rivibre Belly, une rivibre voisine non
endiguke. Les distributions forestibres ont CtC dCterminCes par l'estimation de la distance fluviale IinCaire des forits ii partir
de photos atriennes prises en 1961 et 1981 en amont et en aval des digues. Au cours de I'intervalle de 20 ans, il y a eu une
rkduction de 48% en aval de la digue de St. Mary, qui a CtC achevCe en 1951. En aval de la digue de Waterton, achevte
en 1964, les forCts ripariennes ont accust un dCclin de 23%. L'abondance des peupliers le long de la rCgion d'aval de la
For personal use only.

rivikre Belly non endigute est demeurte relativement inchangee au cours de la pQiode d'ttude,montrant une rtduction de
moins de 1%. Les parties d'amont (non endigutes) des trois rivibres ont subi un dCclin au cours des deux dtcades; des dCclins
respectifs de 4,7, 4,6, et 6,l ont t t t notts le long des rivibres St. Mary, Belly et Waterton. Ainsi, l'endiguement de deux
rivibres dans le sud de 1'Alberta a caust un dtclin rapide et dramatique des forCts ripariennes. En outre, l'association Ctroite
entre la localisation et le degrt du dtclin forestier indique une relation causale entre l'endiguement et le dCclin forestier. Les
causes possibles du dtclin sont discuttes et comprennent peut-Ctre la mortalitC, en particulier celle des semis, causCe par la
sbcheresse.
[Traduit par la revue]

understanding of the ecological consequences of river dam-

Introduction
Although it is unquestionable that human influence has con-
siderable impact on terrestrial ecosystems, the complex web of
biological and physical interactions complicates analyses of
the ecological consequences of specific human activities. With
the increasing demand for water for agricultural, industrial,
and domestic uses, rivers throughout the world are being
dammed, diverted, and altered. Whereas it is certain that alter-
ation of riverflows influences riparian ecosystems, the specific
influences are generally poorly understood since riparian eco-
systems may be complex and respond dynamically to artificial
as well as natural events, including catastrophic floods (Kon-
dolf et al. 1987). Few damming and diversion projects have
been preceded by riparian ecosystem assessments that would
allow for subsequent analysis of the ecological impacts of the
river damming or water diversion. Further, the damming
projects are typically coincidental with intensification of
agricultural, urban, and recreational developments, also con-
founding the issue.
With a warm summer, fertile soil, and limited rainfall,
southern Alberta has become the focus for agricultural irriga-
tion in Canada. Damming and diversion projects are planned
or have been implemented on all of the rivers in southern
Alberta; on rivers where projects have been implemented,
flow regimes have been altered to varying degrees. Thus, an
Printed in Canada 1 Imprime au Canada
ming is particularly relevant to this region.
The St. Mary, Belly, and Waterton rivers flow from the
Rocky Mountain foothills in northern Montana and south-
western Alberta along parallel northeasterly paths character-
ized by generally similar topographical features. The St. Mary
Dam was completed in 1951, replacing the smaller Kimball
Dam. The Waterton River was dammed in 1964. The Belly
River is situated between the St. Mary and Waterton rivers and
only has diversion weirs, which have less impact on down-
stream hydrology than the large dams.
The presence of these three rivers provides an ideal situation
to investigate the consequences of river damming on the distri-
bution of riparian poplars. The three were reasonably similar
in the past with respect to riparian vegetation (Dawson 1885).
Two of the three rivers have been dammed, whereas the Belly
River, located in the middle of the three, is less altered. As
well as permitting comparisons across the three rivers, ade-
quate river segments exist upstream and downstream from the
dams and allow for comparisons within each river. All three
rivers originate in the Waterton - Glacier International Peace
Park (Waterton Lakes National Park, Alberta, and Glacier
National Park, Montana) and two flow along Indian reserves;
thus, the three river valleys and their watersheds are relatively
pristine, minimizing other artificial alterations that might con-
found the study of the impact of damming on riparian forests.
 ROOD AND HEINZE-MILNE
Can. J. Bot. Downloaded from www.nrcresearchpress.com by 99.243.105.24 on 01/09/15
WATERTON RESERVOIR
For personal use only.
FIG. 1. Location of the dams and rivers in southwestern Alberta from which poplar abundances were determined from air photos. Gauging
stations are represented by (*).
Methods
The study area (Fig. 1) consists of three river valleys in the south-
west corner of Alberta. The Belly, St. Mary, and Waterton rivers
originate in alpine tundra (elevation 2000-3000 m) in the Lewis
Range of the Rocky Mountains and flow northeasterly through mon-
tane forest and the Rocky Mountain foothills (aspen parkland) before
cascading into the treeless stretches of fescue prairie (elevation 900-
1000 m). All three rivers originate from mountain snowmelt and rain-
fall runoff, are relatively steep in gradient, and contain generally
clear and cold water. The river discharges are quite erratic, respond-
ing quickly to changes in precipitation and temperature.
Hydrological data were collected by the Water Survey of Canada
(Environment Canada 1985, 1986). Figures 3 -5 represent typical
data for the study area and originate from hydrometric gauging sta-
tions 05AD003 at the Waterton River near Waterton Park (upstream
data), 05AD008 at the Waterton River near Stand Off (downstream,
1950- 1966), and 05AD028 at the Waterton River near Glenwood
(downstream, 1967- 1985). Figure 6 represents data from stations
05AD005 at the Belly River near Mountain View (upstream) and
05AD002 at the Belly River near Stand Off (downstream).
The riparian ecosystem along southern Alberta prairie rivers is
floristically relatively simple with the dominant, and generally exclu-
sive, large woody plants present being poplars (Shaw 1976). Populus
angustifolia, P. balsamifera, and P. deltoides comprise a unique,
natural, disectional, trispecific hybrid swarm (Brayshaw 1965; Rood
et al. 1986).
No attempt was made to investigate possible preferential survival
of the poplar species or of certain hybrids. Rather, in the present
study only overall forest abundances were determined. Since the
three species freely interbreed forming a continuous hybrid swarm
1745
R Y RESERVOIR
(Rood et al. 1986), there was less rationale for investigating specific
differences in the decline of the different poplar genotypes.
Two sets of air photos were used to identify riparian poplar forests:
1961 (scale = 1 : 3 1 680) and 1981 (scale = 1 : 60 000). Transparen-
cies were placed on one of a stereoscopic pair of air photos, and
rivers and riparan poplar forests were traced on the transparency. The
abundance of poplar forests was measured from the transparencies by
a lineal ticking method measured to the nearest rnillimetre. In this
method, the tracing of the rivers was broken into 1-mm segments and
the number of segments associated with forest was counted. This
procedure did not consider the two-dimensional area of the forests,
but estimated the lineal distance of forest abundance, a one-dimen-
sional representation.
Ten 4-km river reaches were evaluated upstream and downstream
from the St. Mary and Waterton dams and a corresponding 80-km
stretch of the Belly River was also studied (twenty 4-km segments for
each river). For each 4-km study segment, the proportion of river
with a poplar forest associated with at least one river bank was deter-
mined, and this value is referred to as the "forest abundance." Mean
(* SE) forest abundances were calculated from the 10 segments of
each of the six river sections (three rivers, upstream and down-
stream). The 80-km sections upstream and downstream from the
dams comprised almost the entire rivers; hence, this sampling evalu-
ated most of the river valley forests rather than providing a sample
for extrapolation to a larger population.
Results and discussion
Along undarnrned segments of rivers in southern Alberta
(i.e., the Belly River and upstream portions of the St. Mary
and watertonrivers), forests generally occurred along
 1746 CAN. J . BOT. VOL. 67, 1989
UPSTREAM DOWNSTREAM
Can. J. Bot. Downloaded from www.nrcresearchpress.com by 99.243.105.24 on 01/09/15
BELLY
For personal use only.
RIVER DISTANCE (km)
FIG. 2. River valley forest abundance along ten 4-km river seg-
ments above and below the St. Mary and Waterton dams and along
a corresponding portion of the neighbouring Belly River. Abun-
dances were determined from air photos taken in 1961 (a) and 1981
(El), and the offset between the two lines represents the change over
the 20-year interval.
70-80% of the river length (Fig. 2). Some river segments,
such as the upper Waterton River near km 16, were less
favorable for the poplar groves and this probably reflected
river valley morphology. For example, steep-walled canyons
exist along certain river segments and are undesirable for the
riparian poplar forests.
Downstream from the St. Mary Dam (Fig. 2, upper right
graph) there was a considerable and consistent decline in
forest abundance over the 20-year interval from 1961 to 1981.
In all of the nine river segments studied (excluding km 36,
which was void of trees in 1961), forest decline was observed.
Represented as a fraction of 1961 abundance, this decline
averaged almost one-half (-47.8%, Table 1). There was a
trend towards lower forest abundances along the furthest
downstream portions of the St. Mary River both in 1961 and
in 1981, a pattern that had already been reported in 1885
(Dawson 1885).
Substantial forest decline was also observed downstream
from the Waterton Dam (Fig. 2, lower right graph). The
decline was observed in all of the 10 river segments and was
very consistent; parallel but offset variation was observed in
TABLE1. Percent change in forest abundance between 1961
and 1981 along rivers of southern Alberta upstream and down-
stream from dams
Downstream
River Upstream Downstream -upstream
St. Mary -4.7k 1.8 -47.8,4.0 -43.1
Belly (undammed) -4.6+1.5 -0.1 k 2 . 6 4.5
Waterton -6.1k1.9 -22.9k1.7 -16.8
NOTE: Values are means t SE of 10 values, except downstream St. Mary for
which nine values were included.
the forest abundance plots for 1961 and 1981 (Fig. 2). The
average decline during the 20-year interval was almost one-
quarter (-22.9%) and was consistently lower than the decline
observed downstream from the St. Mary Dam (Table 1). A
greater decline downstream from the St. Mary Dam might be
expected because this dam was completed in 1951, whereas
the Waterton Dam was completed in 1964.
There was very little variation in forest abundance along the
corresponding (control) downstream portion of the Belly River
over the study interval (Fig. 2; Table 1). This argues against
the downstream decline being due to confounding factors
rather than the damming, since the Belly River is located
between the St. Mary and Waterton rivers (Fig. 1).
Upstream portions of all three rivers showed a slight reduc-
tion in forest abundance over the 20-year interval (Fig. 2).
Quantitatively, the upstream forest decline was similar along
all three rivers and indicated that a minor forest decline
occurred, even without damming, over the interval from 1961
and 1981 (Table 1).
The patterns reported above were very consistent; hence,
the effects of both upstream versus downstream and specific
river were statistically highly significant (two-way ANOVA:
upstream versus downstream, F = 68.0, df = 1, P < 0.001;
rivers, F = 40.6, df = 2, P < 0.001). A highly significant
interaction of upstream -downstream versus river ( F = 35.8,
df = 2, P < 0.001) confirms the differential forest decline on
the dammed versus undammed rivers (Table 1). Downstream
portions of the dammed St. Mary and Waterton rivers under-
went greater forest declines than the upstream reaches of those
same rivers. In contrast, along the undammed Belly River the
downstream portion displayed less forest decline than the
upstream reach between 1961 and 1981.
Thus, the data indicate three patterns with respect to river
damming and forest abundance: (i) forest decline occurred
downstream from two dams but not along a corresponding
downstream portion of an undammed river; (ii) forest decline
was greater downstream than upstream from dams on two
rivers; and (iii) the duration of damming may be related to the
degree of forest decline. Collectively, these data demonstrate
a close relationship between river damming and poplar forest
decline in southern Alberta. Further, the spatial and quantita-
tive associations involving three types of comparisons support
a causal relationship between the river damming and forest
decline.
Although these data clearly demonstrate that an abrupt
decline in poplar forest abundance is associated with river
damming in southern Alberta (Fig. 2; Table l), the ecophysio-
logical basis for the decline is less clear. Declines in poplar
forest abundance have recently been reported downstream
from dams from two other rivers in the western North Ameri-
 ROOD AND HEINZE-MILNE
T UPSTREAM
DOWNSTREAM
I v,
Can. J. Bot. Downloaded from www.nrcresearchpress.com by 99.243.105.24 on 01/09/15
1950 19 6 0 1970 19 8 0
YEAR
For personal use only.
FIG.3. Maximum daily discharge for the Waterton River upstream
and downstream from the Waterton Dam for the period 1941-1981
(data from Environment Canada 1985).
can prairies (Bradley and Smith 1985; Johnson et al. 1976).
The proposed causes for the poplar decline in the previous
studies can be partially tested with the replicated, controlled
experimental design of the present study. Such an analysis
must occur within the context of the ecophysiology of southern
Alberta poplar forests.
The poplars indigenous to southern Alberta are typical of
riparian poplars in their regeneration requirements and life
cycle. Although there is some clonal propagation through
suckering (new shoot production from existing roots), they
probably regenerate principally through seedling establish-
ment. The poplars are relatively fast growing and quite short-
lived. Thus, for the riparian poplar groves to be replenished,
replacement seedling establishment must occur on an ongoing
cycle of about 50 years (Shaw 1976).
Poplar seeds are extremely small and consequently lose
viability quickly (Fenner et al. 1984; McComb and Lovestead
1954); Alberta poplar seeds die within 2-4 weeks (Moss
1938). They require moist soils that lack competing vegetation
and sites well exposed to sunlight for successful germination
and initial seedling establishment (Read 1958). These condi-
tions exist on an irregular basis following spring flooding
(Everitt 1968). Since dams often alter spring flooding down-
stream, previous authors have proposed this as a basis for the
negative impact of damming on riparian forests (Bradley and
Smith 1985; Fenner et al. 1985). However, the irrigation
reservoirs of the present study are not managed in a manner
that effectively alters spring flooding. In most years, flooding
persists downstream from the dams and peak discharges are
not substantially reduced (Fig. 3). Major flood events, such as
those of 1964 and 1975 (Fig. 3), occur at about 20-year inter-
1747
1001 UPSTREAM
I
0
DOWNSTREAM
+F A
MAY JUNE JULY AUG. SEPT.
FIG. 4. Mean daily discharge for the Waterton River upstream and
downstream from the Waterton Dam for the period May 1 to Septem-
ber 30, 1985 (data from Environment Canada 1986).
vals (Phinney 1982) and are relatively unaltered by the present
dams in southern Alberta. Intermediate-intensity flood peaks
are slightly moderated by the dams, and during dry years
minor flood peaks are substantially reduced (Fig. 3). The
timing of peak discharge is generally unaltered or delayed
slightly. Because the dams in southern Alberta have very little
impact on hydrological aspects of spring flooding, it is
unlikely that attenuation of flooding is the cause of the
observed downstream forest decline.
Whereas flooding is essential for seed germination and
establishment, subsequent seedling survival is another limita-
tion to poplar replenishment (McBride and Strahan 1984). Due
to small seed size, and, consequently small seedling root sys-
tems, poplar seedlings are particularly prone to drought-
induced mortality (Read 1958). Thus, conditions of adequate
soil moisture must be maintained throughout the summer if
poplar seedlings are to survive.
This provides a second aspect of poplar replenishment that
could be disturbed by the dams. Summer flows downstream
from southern Alberta dams are dramatically reduced as water
is diverted through canals for irrigation (Fig. 4). The dams are
presently managed to maintain a minimum downstream flow
of 1 m3 . s-', a fraction of the natural flow (Fig. 4). Particu-
larly in July and August, downstream discharge is minimal in
most years (Fig. 5).
The reduced downstream discharge throughout the growing
season is expected to lead to generally reduced plant vigor
(Reily and Johnson 1982) and probably to increased mortality.
Following the prolonged drought of the 1930s and correspond-
ingly reduced stream flows, extensive riparian poplar mortal-
 1748 CAN. J. BOT. VOL. 67, 1989
30-
UPSTREAM
-- 2 0 -
V)
ro
-E
-l
10
cn
3
C)
3 0
a C)
Can. J. Bot. Downloaded from www.nrcresearchpress.com by 99.243.105.24 on 01/09/15
DOWNSTREAM
For personal use only.
YEAR
FIG.5. Mean discharge in August for the Waterton River upstream
and downstream from the Waterton Dam for the period 1950- 1984
(data from Environment Canada 1985).
ity was observed in the central and northern American prairies
(Albertson and Weaver 1945). Similarly, it is likely that the
extensive diverison of water away from the rivers results in an
artificial drought condition in the river valleys.
Whereas this second hydrological alteration, i.e., the reduc-
tion in downstream discharge during the summer, is possibly
the cause of the poplar forest decline, an analysis of flow
characteristics of the Belly River complicates this argument.
Although the Belly is undammed, a diversion weir was con-
structed in 1935 at about the same elevation as the Waterton
and St. Mary dams. The weir has almost no impact on spring
flooding but does considerably reduce the late season flow as
water is diverted for irrigation (Fig. 6). Poplar forests down-
stream from the Belly River weir have not experienced the
dramatic declines observed downstream from the St. Mary and
Waterton dams (Fig. 2; Table 1). Hence, it is not solely the
reduction in August flow that has caused the forest decline
downstream from these two dams.
One aspect of the downstream hydrology that generally
differs between the lower Belly and St. Mary or Waterton
rivers is the abrupt reduction in downstream discharge after
the spring peak (Fig. 4). Whereas operation of the dam creates
a sharp reduction in flow as the sluice gates are closed, the
flow reduction downstream from weirs is usually a more grad-
ual tapering.
The importance of the gradual reduction of downstream dis-
charge may relate to seedling mortality. Seedling roots must
be able to grow downwards and maintain contact with the
water table. Further, the seedlings probably become hardened
-
UPSTREAM
T
DOWNSTREAM
1950 1960 1970 1980
YEAR
FIG.6. Mean discharge in August for the Belly River upstream and
downstream from the main weir for the period 1950-1984 (data
from Environment Canada 1985).
to water stress as the water table recedes and the evapotranspi-
rational demand increases with increasing midsummer temper-
atures. It is possible that a gradual reduction of river flow
could enable hardening of the seedlings and seasonal harden-
ing of established trees, whereas the abrupt restriction of water
downstream from dams would be more-lethal.
While the abrupt restriction of downstream discharge is a
logical partial cause of the poplar decline, it is probably not
the only factor underlying the failure of poplar replenishment.
The large St. Mary and Waterton reservoirs slow the water
flow almost to a halt, resulting in deposition of all sediment
but the fine clay. The downstream discharge is relatively clear
and impoverished of bed load and suspended material; conse-
quently, downstream sedimentation is considerably reduced
(Bradley and Smith 1984; Williams and Wolman 1984). Given
the importance of the dynamic meandering of prairie rivers
and the need for new point bars for poplar replenishment, the
consequence of this reduced sedimentation would be a less
favorable habitat for poplar recruitment (Bradley and Smith
1985; Everitt 1968; Johnson et al. 1976; Noble 1979). The
capture of sediment in reservoirs and consequent reduction in
downstream sedimentation might contribute to the observed
poplar forest decline.
Previous studies have indicated that recovery of suspended-
sediment loads may require up to hundreds of kilometres
downstream from major dams (Williams and Wolman 1984).
In the present study, there was no evidence of recovery of the
f o r e ~ t ~ w i t hthe
i n 40-km segments downstream from dams; no
convergence of the 1961 and 1981 forest-abundance plots
occurred downstream (Fig. 2). Both the abrupt flow restriction
in midsummer and the reduced sedimentation would be
expected to have influence for at least 40 km downstream. The
 ROOD AND HEINZE-MILNE
flow alteration would only be moderated when another tribu-
tary or other drainage compensates for the flow restriction. In
the case of the St. Mary and Waterton rivers, there are no
major tributaries within 40 krn downstream of the dams.
The poplars of southern Alberta comprise the principal, and
generally only, large woody plants in the riparian forests.
Consequently, the fate of the entire forest is ultimately tied to
the fate of the poplars. The poplar groves provide a forest
canopy allowing for the development of an understory of
shrubs and herbs. Arboreal bird nesting is common and the
forests support a rich terrestrial fauna. The central role of
poplars in the riparian ecosystem is undeniable, and as the
poplars die, so dies the riparian forest ecosystem.
In summary, a close relationship was established between
river damming and riparian forest decline in southern Alberta.
Can. J. Bot. Downloaded from www.nrcresearchpress.com by 99.243.105.24 on 01/09/15
Some possible causes of the decline based on altered hydrolog-
ical patterns are discussed and these might contribute to a
failure of poplar replenishment due to drought-induced mortal-
ity, particularly of seedlings.
Acknowledgements
Preliminary data was gathered with the enthusiastic assis-
tance of Dennis Wollersheim, Marie Wilfong, MaryLee
Naydiuk, and Nic Bate. Useful discussions about sampling
methodology were provided by Dr. John S. Campbell and
Mr. John Mahoney. Dr. Alex Johnston and Mr. Ron Middle-
ton provided useful comments on historical aspects of flooding
For personal use only.
and damming in southern Alberta. This study was supported
by Alberta Environmental Research Trust grant TO1016 and a
University of Lethbridge research grant to S.B.R.
ALBERTSON, F. W., and WEAVER, J. E. 1945. Injury and death or
recovery of trees in prairie climate. Ecol. Monogr. 15: 395-433.
BRADLEY, C. E., and SMITH,D. G. 1984. Meandering channel
response to altered flow regime: Milk River, Alberta and Montana.
Water Resour. Res. 20: 1913- 1920.
1985. Plains cottonwood recruitment and survival on a
prairie meandering river floodplain, Milk River, southern Alberta
and northern Montana. Can. J. Bot. 64: 1433- 1442.
BRAYSHAW, T. C. 1965. Native poplars of southern Alberta and their
hybrids. Can. Dep. For. Publ. No. 1109.
DAWSON, G. M. 1885. Map showing wooded and prairie districts,
etc., in the region of the vicinity of the Bow and Belly rivers. In
Maps, etc., to accompany rePo& of progress for 1882- 83 - 84.
1749
Geological and Natural History Survey and Museum of Canada.
Dawson Bros., Montreal, Que.
ENVIRONMENT CANADA. 1985. Historical streamflow summary:
Alberta, to 1984. Environment Canada, Ottawa.
1986. Surface water data: Alberta, 1985. Environment
Canada, Ottawa.
EVERITT, B. 1968. Use of cottonwood in an investigation of the recent
history of a floodplain. Am. J. Sci. 266: 417-439.
FENNER, P., BRADY, W. W., and PATTON, D. R. 1984. Observations
of seeds and seedlings of Fremont cottonwood. Desert Plants, 6:
55-58.
1985. Effects of regulated water flows on regeneration of
Fremont cottonwood. J. Range Manage. 38: 135- 138.
JOHNSON, W. C., BURGESS, R. L., and KEAMMERER, W. R. 1976.
Forest overstory vegetation and environment on the Missouri River
floodplain in North Dakota. Ecol. Monogr. 46: 59-84.
KONDOLF, G. M., WEBB,J. W., SALE,M. J., and FELANDO, T. 1987.
Basic hydrologic studies for assessing impacts of flow diversions on
riparian vegetation: examples from streams of the eastern Sierra
Nevada, California, USA. Environ. Manage. 11: 757 -769.
MCBRIDE, J. R., and STRAHAN, J. 1984. Establishment and survival
of woody riparian species on gravel bars of intermittent streams.
Am. Midl. Nat. 112: 235-245.
MCCOMB, A. L., and LOVESTEAD, H. S. 1954. Viability of cotton-
wood seeds in relation to storage temperatures and humidities.
USDA For. Serv. Tree Planters Notes, 17: 9- 11.
Moss, E. H. 1938. Longevity of seed and establishment of seedlings
in species of populus. Bot. Gaz. 99: 529-542.
NOBLE,M. G. 1979. The origin of Populus deltoides and Salk
interior zones on point bars along the Minnesota River. Am. Midl.
Nat. 102: 59-67.
PHINNEY, R. B. 1982. Flood of June 1975 in the Oldman River Basin,
Alberta. Environment Canada, Water Resources Branch Report.
READ,M. A. 1958. Silvicultural characteristics of Plain's Cotton-
wood. Rocky Mt. For. Range Exp. Stn. Pap. No. 33.
REILY,P. W., and JOHNSON, W. C. 1982. The effects of altered hydro-
logic regime on tree growth along the Missouri River in North
Dakota. Can. J. Bot. 60: 2410-2423.
ROOD,S. B., CAMPBELL, J. S., and DESPINS, T. 1986. Natural poplar
hybrids from southern Alberta. I. Continuous variation for foliar
characteristics. Can. J. Bot. 64: 1382- 1388.
SHAW,R. K. 1976. A taxonomic and ecologic study of the riverbot-
tom forest on St. Mary River, Lee Creek and Belly River in South-
western Alberta, Canada. Great Basin Nat. 36: 243-271.
WILLIAMS, G. P., and WOLMAN, M. G. 1984. Downstream effects of
dams on alluvial rivers. Geol. Surv. Prof. Pap. (U.S.), No. 1286.