Journal of Great Lakes Research 41 Supplement 2 (2015) 8190

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Journal of Great Lakes Research

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Decadal regulation of phytoplankton abundance and water clarity in a

large continental reservoir by climatic, hydrologic and trophic processes
R.J. Vogt a,, S. Sharma b, P.R. Leavitt a
a
Limnology Laboratory, Department of Biology, University of Regina, Regina, Saskatchewan, Canada
b
Department of Biology, York University, Toronto, Ontario, Canada

a r t i c l e i n f o a b s t r a c t

Article history: Large continental reservoirs are impacted by multiple stressors including climate change, watershed develop-
Received 13 July 2014 ment, food-web alteration, water extraction, hydroelectric-power generation, and industrial aquaculture. Such
Accepted 2 September 2014 complexity makes it difcult to identify the hierarchical relationships among regulatory processes, forecast
Available online 4 December 2014
effects of environmental change, or develop adaptive management strategies. Here we present a regression-
Communicated by Rebecca North
based analysis of 19 years of limnological change in a representative main-stem reservoir within central
North America to suggest that phytoplankton abundance and water clarity are regulated by hierarchical but in-
Index words: dependent mechanisms. The Qu'Appelle arm of Lake Diefenbaker, Canada, was monitored during summers
Reservoir (MayAugust) of 19952013 to evaluate the unique and interactive effects of continental climate systems,
Water quality regional meteorology, river hydrology and limnological characteristics on phytoplankton abundance and water
Transparency clarity. Regression models explained 4852% of historical variation despite the absence of pronounced temporal
Climate patterns in monthly or summer phytoplankton abundance and water clarity. Phytoplankton abundance (mainly
Multiple stressors diatoms, agellates) was correlated positively with soluble reactive phosphorus concentrations and inversely
Chl a
with the density of large herbivores, factors which were themselves correlated to variation in chemical conditions
(oxygen, dissolved inorganic carbon) and dissolved organic carbon content, respectively. In contrast, water clarity
varied directly as a function of climate systems (producing warm, dry winters) and inversely with river ow. We
conclude that anticipated climate change (warmer, less runoff) will improve water clarity in lacustrine regions of
large prairie reservoirs on decadal scales by reducing inorganic turbidity, while nutrient uxes associated with
economic development may independently regulate algal abundance.
2014 International Association for Great Lakes Research. Published by Elsevier B.V. All rights reserved.

Introduction Climate variability and directional warming are expected to have

As with natural lakes, water quality in large continental reservoirs is
impacted by climate change (Chen et al., 2007; Marce et al., 2010;
Sukenik et al., 2012), watershed development (Hall et al., 1999a;
Jackson et al., 2001; Leigh et al., 2010) and food-web alteration (Post
et al., 2002), all of which interact and vary in intensity through time. In
addition, reservoirs may be managed for water extraction (Baldwin
et al., 2008; Joeckel and Diffendal, 2004), hydroelectric power generation
(Soumis et al., 2004) and industrial aquaculture (Figueredo and Giani,
2005; Guo and Li, 2002), while ontogenetic changes in internal nutrient
supply can cause a decadal-scale upsurge in algal production (Hall et al.,
1999a; Thornton et al., 1990). Such complex interactions between multi-
ple stressors makes it difcult to identify the hierarchical relationships
among regulatory processes, forecast effects of environmental change,
or develop adaptive management strategies to balance economic and
environmental issues (Crossman et al., 2013; Marce et al., 2010).
Corresponding author at: Department of Biological Sciences, Trent University,
Peterborough, Ontario, Canada. Tel.: +1 705 748 1011 (6129)
E-mail address: RichVogt@gmail.com (R.J. Vogt).
pronounced effects on water resources in the northern Great Plains
(Barrow, 2009). Present-day climate variability arises from the interac-
tion of three major climate systems and three air masses (Arctic, Pacic,
Gulf of Mexico) that supply moisture into the continental interior
(Bonsal and Shabbar, 2008; Bryson and Hare, 1974). Variation in the
North Atlantic Oscillation during winter (NAOw) appears to regulate cy-
clonic activity over the Prairies early each year (Hurrell, 1995; Wang
et al., 2006) and is associated with changes in timing of ice melt, algal
production and development of the clear water phase during spring
(Drscher et al., 2009; Weyhenmeyer et al., 1999). Similarly, the Pacic
Decadal Oscillation (PDO) (Mantua et al., 1997) and El NioSouthern
Oscillation (ENSO) (Trenberth and Hurrell, 1994) regulate inux of Pa-
cic precipitation to the Prairies and runoff from western mountains
(Shabbar et al., 2011; St. Jacques et al., 2010) and can interact to produce
exceptionally warm, dry conditions in winter and spring (Mantua et al.,
1997; McCabe et al., 2004) and which, in turn, alter plankton phenology
(McGowan et al., 2005b; Winder and Schindler, 2004). Finally, general
circulation models predict that the mean annual temperature of the Ca-
nadian Prairies will increase ~4 C by 2050 (Barrow, 2009; Lapp et al.,
2012), leading to higher hydrological variability and intensication of

http://dx.doi.org/10.1016/j.jglr.2014.11.007
0380-1330/ 2014 International Association for Great Lakes Research. Published by Elsevier B.V. All rights reserved.
82 R.J. Vogt et al. / Journal of Great Lakes Research 41 Supplement 2 (2015) 8190
both droughts (Lapp et al., 2013; van der Kamp et al., 2008) and pluvial
periods (Winter and Rosenberry, 1998). These events will further lead
to more variable water chemistry (Pham et al., 2009; Starks et al.,
2014), altered regional hydrology (Pomeroy et al., 2007; Schindler
and Donahue, 2006) and changes in heat budgets that regulate plank-
tonic production (Johnk et al., 2008; Paerl and Otten, 2013; Winder
and Sommer, 2012), community composition (Cantin et al., 2011;
Huisman et al., 2004; Magnuson et al., 1997), and outbreaks of
potentially-toxic cyanobacteria (Donald et al., 2011; Sukenik et al.,
2012).
Large reservoirs in continental interiors are often found in fertile
catchments subject to extensive anthropogenic inuence including agri-
culture (nutrients, erosion, pesticides, drainage), urbanization (microbes,
pharmaceuticals, metals, nutrients), and hydrologic modications
(Bolgrien et al., 2009; Fernandez et al., 2014; Hall et al., 1999a). For exam-
ple, ancestral European settlement of the Canadian Prairies since 1880
has transformed the grasslands due to mechanized cultivation (since
1920), fertilization (since the 1940s) and localized urban development
(Bunting et al., 2011; Hall et al., 1999b; Leavitt et al., 2006). Over 70% of
surface cover has been converted to grain and livestock farming while
permanent surface water cover has declined over 50% to 58% of land
area (Bunting et al., 2011; Hall et al., 1999b). Elevated uxes of nutrients
and suspended solids have increased algal productivity, reduced water
clarity, and intensied blooms of cyanobacteria in natural lakes (Leavitt
et al., 2006; Orihel et al., 2012) although less is known of the responses
of regional reservoirs (McGowan et al., 2005a; North et al., 2014).
Large-scale elimination of wetlands by farming has changed runoff of
snow and rain (Pomeroy et al., 2007; van der Kamp et al., 2008), an im-
portant control on the uxes of nutrients and other solutes to aquatic
ecosystems in the grasslands (Pham et al., 2008; Starks et al., 2014). Sim-
ilarly, reservoirs in continental interiors are often subject to regulated but
extensive (N 5 m) variation in lake water-level reecting the highly sea-
sonal nature of local snowmelt (MarchApril) (Akinremi et al., 1999;
Fang and Pomeroy, 2007), the importance of snowmelt from the Rocky
Mountains to the hydrological budgets of main-stem reservoirs (June)
(Saskatchewan Water Security Agency, SWSA, 2012), and the use of
large reservoirs for industrial, urban, and agricultural applications (Hall
et al., 1999a).
Complex, hierarchical, and temporally-variable interactions be-
tween multiple stressors complicate development of a mechanistic un-
derstanding of how humans and climate interact to regulate water
quality and ecosystem structure in reservoirs (Crossman et al., 2013;
Hart and Calhoun, 2010). To address this issue, we analyzed a 19-year
time series from a large lacustrine embayment (Qu'Appelle arm) of
the Lake Diefenbaker reservoir to quantify how environmental variabil-
ity associated with climatic regimes, regional meteorology, hydrologic
variability and intrinsic limnological properties related to human activ-
ity (nutrient content, food-web structure, etc.) inuences phytoplank-
ton abundance and water clarity in continental reservoirs. Such long-
term ecological research (LTER) is of particular use in developing robust
predictive models for changes in biological, chemical, and physical
properties of aquatic ecosystems, and allowing scientists and managers
to develop management strategies to adapt to future environmental
change (Adrian et al., 2009; Chen et al., 2007). The empirical models de-
veloped here serve as an important rst step to identifying targets for
potential future management action both in this region and in similar
sub-humid ecozones around the world (e.g., southern South America,
central Asia, central China, Australia).
Methods
Site description
Lake Diefenbaker is a 220-km long reservoir located ~554 m above
sea level (a.s.l.) on the South Saskatchewan River, Canada, and which
serves as headwater to Qu'Appelle River drainage basin (Hall et al.,
1999a; Hecker et al., 2012). The reservoir's catchment which lies mainly
within Alberta, Saskatchewan, and upstate Montana was originally
mixed-grass Prairie, but it is presently under predominantly (~65%) ag-
ricultural land use (cereal crops, pasture, livestock) (Hall et al., 1999a).
The reservoir lies within a broad glacial outwash valley formed by the
meltwaters of the Wisconsin ice sheet (Christiansen, 1960) and is
underlain by cretaceous marine sedimentary bedrock (Bearpaw forma-
tion) with a thick overburden of glacial till and predominantly dark-
brown chernozemic soils (Acton et al., 1998). There are no major
urban centers in the immediate vicinity of Lake Diefenbaker, although
Calgary (pop. 990,000), Red Deer (pop. 97,100) and Medicine Hat
(pop. 61,200) lie 200400 km upstream in the upper catchment within
Alberta while smaller Swift Current, Saskatchewan (pop. 15,500) is
located ~50 km south of the reservoir.
Regional climate is classied as sub-humid continental, exhibits a
moisture decit of 3060 cm/year, and is characterized by short warm
summers (mean 19 C in July), cold winters (mean 16 C in January),
and low mean annual temperatures (~1 C) with high seasonal variabil-
ity. Most regional runoff occurs in spring during a brief snowmelt period
in late March or early April (Pham et al., 2009; Pomeroy et al., 2007)
although monthly inows to Lake Diefenbaker are highest in June
and early July when melt-waters arrive from the Rocky Mountains
(Akinremi et al., 1999; Fang and Pomeroy, 2007; SWSA, 2012). Reservoir
hydrology varies widely among years with a four-fold difference be-
tween minimum and maximum annual inows since 1995 and a N 6-m
seasonal variation in lake level (Hall et al., 1999a; Quiones-Rivera
et al., 2015).
Lake Diefenbaker is a large (area = 394 km2), deep (Zmax ~ 62 m)
basin with a relatively short residence time (~1.3 years) characteristic
of large riverine reservoirs in central North America (Bolgrien et al.,
2009; Hall et al., 1999a). Surface waters in the Qu'Appelle arm (area
~ 50 km2; Zmax ~ 23 m) are cool and well oxygenated (Drscher et al.,
2009; Sadeghian et al., 2015) with low nutrient content and phyto-
plankton abundance relative to other regional lakes (Leavitt et al.,
2006; Patoine et al., 2006). In general, the embayment warms slowly
and is polymictic although in some years limited thermal stratication
is observed during late-July or August (Drscher et al., 2009; Hudson
and Vandergucht, 2015). Phytoplankton are characteristic of well-
mixed mesotrophic lakes with abundant diatoms in spring giving way
to later-summer assemblages of agellates and occasional
cyanobacteria (Hecker et al., 2012; McGowan et al., 2005b; Patoine
et al., 2006). The zooplankton community is composed mainly of
cyclopoid (Diacyclops thomasii) and calanoid (Leptodiaptomus siciloides)
copepods, but also includes large (Daphnia pulex, Daphnia galeata
mendotae, Diaphanosoma birgei, Holopedium gibberum) and small
(Bosmina longirostrus, Daphnia retrocurva) cladocerans at lower
densities (Patoine et al., 2006). Fish assemblages include 25 native
and stocked species, such as walleye (Sander vitreus), northern pike
(Esox lucius), rainbow trout (Oncorhynchus mykiss), lake char (Salvelinus
namaycush), yellow perch (Perca avescens), cisco (Coregonus artedi),
bigmouth buffalo (Ictiobus cyprinellus), and white sucker (Catostomus
commersonii) (for full list, see SWSA, 2012).
Paleolimnological analyses demonstrate that the downstream lacus-
trine portions of Lake Diefenbaker have experienced three intervals of
contrasting water quality since valley inundation in 1967 (Hall et al.,
1999a; Lucas et al., 2015; Tse et al., 2015). First, lake production in-
creased ca. 19681975 when diatoms were characteristic of eutrophic
conditions (Stephanodiscus niagarae, Stephanodiscus parvus) and colo-
nial and N2-xing cyanobacteria were present. Second, reservoir pro-
duction declined ca. 19751986, as mesotrophic diatoms (Tabellaria
occulosa str. IIIp, Fragilaria crotonensis, Aulacoseira ambigua) replaced
eutrophic species (Hall et al., 1999a; Lucas et al., 2015). Finally, the tro-
phic status of Lake Diefenbaker increased again after ca. 1986, with ele-
vated abundance of productive diatoms (A. ambigua), chlorophytes and
colonial cyanobacteria (Hall et al., 1999a; Tse et al., 2015). During this lat-
ter transition, water-column chlorophyll (Chl a) increased from ~1 g/L
 R.J. Vogt et al. / Journal of Great Lakes Research 41 Supplement 2 (2015) 8190
in 1984 to ~5 g/L in 1994 (Environment Canada, 1988). Recent publica-
tions suggest that modern phytoplankton abundance has remained ele-
vated (McGowan et al., 2005b; Patoine et al., 2006) and that colonial
cyanobacteria may be problematic (Hecker et al., 2012), although there
has been no comprehensive evaluation of water quality change since
the mid-1990s. Whole-basin carbon budgets demonstrate that Lake Die-
fenbaker has been net heterotrophic (respiration N primary production)
during the past 20 years (but see Quiones-Rivera et al., 2015), although
CO2 sequestration has increased since ca. 2003 in this and other basins
(Finlay et al., 2009, 2010).
Climate data
Time series of climate variables (19952010) were assembled from
public archives, including the US National Oceanic and Atmospheric Ad-
ministration (NAO), the Australian National Climate Centre (Southern
Oscillation Index, SOI), and the University of Washington Joint Institute
for the Study of the Atmosphere and Ocean (PDO). Additionally, we es-
timated the known synergistic interaction between El Nio phases of
ENSO and positive phases of the PDO as the product of their respective
indices (Bonsal and Shabbar, 2008; Shabbar and Yu, 2012). The calendar
day of year (DOY) of spring ice melt from Lake Diefenbaker was
obtained from Vogt et al. (2011).
Meteorology
Meteorological measurements (19952013) within the catchment
were obtained from the Environment Canada National Climate Archives
(http://www.climate.weatherofce.gc.ca/ climateData; April 2014) for
the Outlook, Saskatchewan, weather station situated ~ 28 km north of
Lake Diefenbaker (5129N, 10703W). Mean monthly air temperature
(C), precipitation (mm) and daily wind speed (km/h) were computed
for an ice-free (summer) period that overlapped with limnological
sampling (01 May31 August). Snow accumulation (mm) was estimat-
ed as the sum of monthly precipitation from JanuaryMarch in each
calendar year.
Hydrology
Hydrologic data (19952010) were collected by the Saskatchewan
Water Security Agency (SWSA) and used to assess the monthly or
total summer inow to Lake Diefenbaker (m3/d). Inow was estimated
using standard governmental procedures, based on data from discharge
gauges on streams, measurements of lake levels, and estimates of net
evaporation (Saskatchewan Water Security Agency, SWSA, 2012 and
unpublished).
Limnological characteristics
The Qu'Appelle arm of Lake Diefenbaker is one of two large lacustrine
embayments in the reservoir and was sampled bi-weekly during sum-
mer 19952013 using standard protocols as part of a regional long-
term ecological monitoring program (QU-LTER) (Finlay et al., 2009;
McGowan et al., 2005b; Patoine et al., 2006). Briey, depth-integrated
samples were collected at midday at a standard geo-positioned station
(5101.260N, 10629.840W) by pooling casts of a 2-L Van Dorn water
bottle deployed at 1-m intervals to 15 m depth. Integrated samples
were ltered through a 0.45-m pore membrane lter and analyzed at
the University of Alberta Water Chemistry Laboratory for concentrations
of soluble reactive phosphorus (SRP, g P/L) and total dissolved phos-
phorus (TDP, g P/L), as well as total dissolved nitrogen (TDN), ammoni-
um (NH+ 4 ), and the sum of NO2
2
and NO2 3 (all g N/L) (Patoine et al.,
2006). Total inorganic (TIC) and dissolved organic carbon (DOC) concen-
trations (mg C/L) in ltered samples were quantied using a Shimadzu
model 5000A (Shimadzu, Kyoto, Japan) total carbon analyzer following
Finlay et al. (2009). Water temperature (Twater; C) and oxygen content
83
(mg O2/L) were computed as average values of depth proles collected
every 1 m using a YSI model 85 meter (YSI, Yellow Springs, Ohio,
USA). Surface water pH was measured at 0.5-m depth using a calibrated
handheld probe, while water clarity was measured using a 20-cm diam-
eter Secchi disk. Zooplankton densities were estimated bi-weekly, May
August for 19952012, from vertical tows of a 20-cm diameter Wiscon-
sin net (243-m mesh) at the standard sampling station. Invertebrate
samples were preserved and enumerated according to Patoine et al.
(2006). Individual species densities (ind./L) were summed as total zoo-
plankton, copepods, large cladocerans and small cladocerans (see
above for species). Rotifers were not enumerated.
Phytoplankton abundance and community composition in the Qu
Appelle arm was estimated 19952012 using standard trichromatic
and high performance liquid chromatographic (HPLC) analysis of carot-
enoid and chlorophyll pigments (Leavitt and Hodgson, 2001; Vogt et al.,
2011). Briey, particulate organic matter (POM) from depth-integrated
water samples (see above) was collected on GF/C glass ber lters
(nominal pore 1.2-m) and extracted using either pure acetone
(trichromatic) or a mixture of acetone:methanol:water (80:15:5, by
volume) (HPLC). Trichromatic estimates of chlorophyll a (Chl a) con-
centrations were used as an index of total phytoplankton abundance
(g Chl/L). In contrast, carotenoids, chlorophylls and their derivatives
were isolated and quantied (nmol pigment/L) using an Agilent
model 1100 HPLC system (Agilent Technologies Inc., Mississauga,
Canada) tted with photodiode-array and uorescence detectors and
calibrated using authentic standards (Leavitt and Hodgson, 2001).
Historical changes in phytoplankton community composition were
estimated using pigment biomarkers from siliceous algae (mainly
diatoms and chrysophytes; fucoxanthin), cryptophytes (alloxanthin),
dinoagellates (peridinin), chlorophytes (Chl b), colonial cyanobacteria
(myxoxanthophyll), and N2-xing cyanobacteria (aphanizophyll). As
shown by Donald et al. (2013), trichromatic- and HPLC-based analyses
provide equivalent estimates of gross phytoplankton abundance and
composition as do enumerations of cellular biovolume using light
microscopy.
Data analysis
For modeling purposes, historical changes in water quality were
assessed using estimates of total phytoplankton abundance (Chl a)
and water clarity (Secchi depth) during 19952010, the interval in
which complete predictor data were available. First, linear regression
models were developed for both explanatory and response variables
to assess the mode of variation of each time series. Time series were
detrended using rst-difference procedures if temporal autocorrelation
was detected. Second, multiple regression models were developed for
both water quality indices using forward selection of predictors with
two stopping criteria ( and R2adj; 9999 permutations) from the full
suite of climatic, meteorological, hydrological and limnological variables
(Blanchet et al., 2008). As our goal was to identify potential regulatory
mechanisms rather than develop the most parsimonious model, nal
model composition was not based solely on Akaike's Information Crite-
rion (AICc), although typically the nal model also exhibited the lowest
AICc (analysis not shown). The relative contribution of each predictor
was quantied using a type III sum of squares analysis of variance
(ANOVA) in which variation explained by each predictor was summed
into classes representing climate, meteorology, hydrology and internal
lake characteristics. Subsequently, regression models were developed
for the statistically signicant predictors of water quality and clarity,
thereby allowing us to evaluate the hierarchical relationships among
potential regulatory variables. This approach avoids over-tting with
complex models of poor predictive capabilities (Sharma et al., 2013). Fi-
nally, regression analyses were repeated using time series with monthly
resolution to assess seasonal variation in potential regulatory processes.
Variables were transformed (log10) as necessary to produce normal dis-
tributions and co-linear parameters were excluded from nal models.
84 R.J. Vogt et al. / Journal of Great Lakes Research 41 Supplement 2 (2015) 8190

All data manipulation and statistical analyses were performed in the the greatest water clarity occurred during August and the least during
R-language environment using the lm (linear regression), acf (auto- June in most years (Fig. 1c).
correlation) and forward.sel (packfor) (forward selection) functions
(R Development Core Team, 2014). Phytoplankton community composition

HPLC analysis of phytoplankton community composition revealed

Results that siliceous algae (diatoms and chrysophytes) composed 5080%
total algal biomass in the Qu'Appelle arm of Lake Diefenbaker in each
Water quality time-series year (Fig. 2). In general, relative abundance of these taxa peaked during
May, before being partly replaced by cryptophytes (most years), dino-
Linear regression analysis revealed no signicant linear trends in agellates (9 years) and colonial cyanobacteria (14 years). Overall,
total phytoplankton abundance or water clarity (Fig. 1a, Table 2). diazotrophic cyanobacteria were abundant early (19961999) and
When highly productive 1995 was not considered, a small but signi- late (20072012) in the time series, whereas non-N2-xing colonial
cant increase in total phytoplankton abundance was recorded for the in- cyanobacteria were more common in other years (Fig. 2). Cyanobacteria
terval 19962012 (R2adj = 0.17, p = 0.05) (Fig. 1a). Regression analysis were uncommon in 2001, 2004 and 2006, whereas this group exhibited
of time series with monthly resolution showed the increase in phyto- its maximum abundance within the QU-LTER during 2012.
plankton abundance was restricted to July (R2adj = 0.26, p = 0.019)
only if 1995 was eliminated, and was not observed in any analyses Environmental time series
of data derived solely from May, June or August (all R2 adj = 0.00,
p N 0.34). In contrast, there were no signicant linear trends in water Regression-based analysis of time series of physico-chemical, cli-
clarity recorded in any month (R2adj = 0.000.07, p N 0.15), though matic, meteorological and hydrological parameters (Table 1) revealed

20.0
Total Phytoplankton (g Chl L-1)

A)
15.0
Water Clarity (m)

10.0

5.0

0.0
1995 1997 1999 2001 2003 2005 2007 2009 2011 2013

Total Phytoplankton Water Clarity

Total Phytoplankton (g Chl L-1)

30.0
B)
25.0

20.0

15.0

10.0

5.0

0.0
1995 1997 1999 2001 2003 2005 2007 2009 2011 2013

8.0
C)
7.0
6.0
Water Clarity (m)

5.0
4.0
3.0
2.0
1.0
0.0
1995 1997 1999 2001 2003 2005 2007 2009 2011 2013

May June July August

Fig. 1. Historical changes in mean (standard error) total phytoplankton abundance (as g Chl a/L) and water clarity (as Secchi depth, m) in the Qu'Appelle arm of Lake Diefenbaker,
Canada, based on annual (MayAugust) (A) and monthly monitoring (B, C) during 19952013. There were no statistically signicant trends in any time series, though there was a modest
increase in total algae for the interval 19962013.
 R.J. Vogt et al. / Journal of Great Lakes Research 41 Supplement 2 (2015) 8190 85

100%

90%

Relative abundance (% pigment sum)

80%

70%

60%

50%

40%

30%

20%

10%

0%
MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA MJ JA
1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012
N2-fixing cyanob. colonial cyanob. chlorophytes siliceous algae cryptophytes dinoflagellates

Fig. 2. Mean monthly (MayAugust) relative abundance of the major phytoplankton groups in the Qu'Appelle arm of Lake Diefenbaker, Canada (19952012). Absolute and relative abun-
dance of phytoplankton groups was determined by high performance liquid chromatography of their diagnostic pigments. Phytoplankton groups (and pigments) include N2-xing
cyanobacteria (aphanizophyll), colonial cyanobacteria (myxoxanthophyll), chlorophytes (chlorophyll b), siliceous algae (diatoms and chrysophytes; fucoxanthin), cryptophytes
(alloxanthin), and dinoagellates (peridinin).

few signicant trends during the past 20 years. Total zooplankton den-
sities declined signicantly and substantially since the mid-1990s
(R2adj = 0.38 p = 0.004), with a particularly steady rate of decrease
since 2000 (R2adj = 0.78 p b 0.001) (Fig. 3a). This trend was driven by
concomitant reductions in mean summer densities of large-bodied cla-
docerans (R2adj = 0.69, p b 0.001) and copepods (R2adj = 0.47, p =
0.002), but not small cladocerans (R2adj = 0.00, p = 0.46) (Fig. 3b). Sim-
ilarly, the pH of Lake Diefenbaker surface waters increased nearly
1.0 unit since 1995 (R2adj = 0.37, p = 0.008), similar to patterns record-
ed in other prairie lakes (Finlay et al., 2009; Vogt et al., 2011). Of the 20
remaining variables, only concentrations of oxygen (increase; R2adj =
0.31, p = 0.015) and NH+ 2
4 (decline; R adj = 0.37, p = 0.007) exhibited
signicant directional changes during the monitoring period.
Models of summer water quality
Intrinsic limnological characteristics were the strongest predictors of
total phytoplankton abundance during 19952010, accounting for 48%
of historical variation in concentrations of Chl a (Fig. 4a). Specically,
mean phytoplankton abundance during summer was correlated posi-
tively to SRP concentration (29% variation explained), and negatively
to total zooplankton abundance (19% variation) (Fig. 4b, Table 2). Anal-
ysis of models based on monthly data (not shown) suggested that SRP
was a signicant predictor (p b 0.05) of phytoplankton abundance late
in summer (JulyAugust), while effects of zooplankton were restricted
to July (analysis not shown). In contrast, changes in mean summer
water clarity (52% variation explained) were predicted by hydrologic
inow (37% variation) and the prevalent climate system (ENSO-PDO;
15% variation), but not by limnological or meteorological parameters
(Fig. 4b, Table 2).
Additional regression models suggested that the controls of water
quality (SRP, zooplankton) were themselves under independent regula-
tion. For example, water-column concentrations of SRP were correlated
negatively to oxygen content and positively to concentrations of inor-
ganic C (R2adj = 0.48 p = 0.006), whereas total zooplankton abundance
was correlated negatively to concentrations of DOC (R2adj = 0.21 p b
0.05) (Table 2). Regression models were not developed for the principal
predictors of water clarity (hydrologic inow, ENSO-PDO), because
those factors are regulated by processes outside the spatial domain of
our data collection (snowmelt in Rocky Mountains and Pacic sea-
surface temperature, respectively) (Lapp et al., 2013; St. Jacques et al.,
2010).
Discussion
Analysis of a comprehensive suite of limnological and environmen-
tal variables from the lacustrine zone of a major continental reservoir
(Table 1) demonstrated that phytoplankton abundance, community

Table 1
Physico-chemical, morphometric, climatic, hydrological and meteorological variables for the Qu'Appelle arm of Lake Diefenbaker. Mean physico-chemical and meteorological values are
calculated for May-August during 19952012. Ranges are standard deviations of the mean. Variables include soluble reactive phosphorus (SRP), total dissolved P (TDP), dissolved organic
carbon (DOC), total inorganic carbon (TIC), depth-integrated water temperature (Twater), oxygen concentration (O2), and indices of the Pacic Decadal Oscillation (PDO), El NioSouthern
Oscillation (ENSO), and North Atlantic Oscillation (NAO). DOY = calendar day of year.

Physico-chemistry Morphology Climate, hydrology, meteorology

SRP (g/L) 10.5 12.3 Lat (N)long (W) 51.12106.63 Ice out date (DOY) 120.7 8.8
TDP (g/L) 18.9 18.1 Elevation (m) 552 PDO 0.03 0.74
NO3 (g/L) 167.2 93.7 Lake area (km2) 394 ENSO 0.54 6.24
NH4 (g/L) 18.6 20.9 Mean depth (m) 33 NAO 4.82 12.79
DOC (mg/L) 6.8 3.2 Max depth (m) 62 Annual inow (m3) 7.0 2.6 109
TIC (mg/L) 33.6 4.9 Volume (m3) 9.40 109 Summer temperature (C) 15.9 1.0
pH 8.7 0.5 Water residence time (year) 1.3 Mean monthly precipitation (MayAug) (mm) 50.5 17
Twater (C) 13.2 1.0 Gross drainage area (km2) 1.36 105 Mean monthly precipitation (JanDec) (mm) 29.5 8.6
O2 (mg/L) 9.0 1.6 Effective drainage area (km2) 8.69 104 Winter snowfall (mm) 133 55
Total zoopl. (ind/L) 23.8 14.5 Wind speed (km/h) 11.9 3.7
Chlorophyll a (g/L) 5.4 2.5
Secchi depth (m) 3.4 0.4
86 R.J. Vogt et al. / Journal of Great Lakes Research 41 Supplement 2 (2015) 8190
A)
70.00
60.00
Total Zooplankton (ind L-1)
50.00
40.00
30.00
20.00
10.00
0.00
1995 1998 2001
Fig. 3. Mean summer density of (A) total zooplankton density (ind./L) and (B) copepods (Leptodiaptomus siciloides, Diacyclops thomasii), large cladocerans (Daphnia spp.) and small
cladocerans collected from the Qu'Appelle arm of Lake Diefenbaker, Canada, during MayAugust 19952012.
composition and water clarity have been resilient to climatic and an-
thropogenic stressors during the past 20 years (Figs. 1, 2). All three pa-
rameters show little evidence of directional change during 19952013,
consistent with recent paleolimnological quantication of phytoplank-
ton abundance in lacustrine regions since 1967 (Tse et al., 2015) and
30 years of Landsat imagery (Yip et al., 2015), but in contrast to contin-
ued increases in regional population, agricultural production and cli-
mate warming, and to declining river discharge (Bunting et al., 2011;
Leavitt et al., 2006; St. Jacques et al., 2010). Instead, half of observed
inter-annual variation in mean-summer phytoplankton abundance
(mainly diatoms, agellates) was explained by trophic processes (nutri-
ent availability, herbivory) (Table 2, Fig. 3), whereas a similar propor-
tion of changes in water clarity were explained mainly by physical
controls (ENSO-PDO, river inow) (Table 2). Given that the food-web
structure and hydrological cycle of Lake Diefenbaker are similar to
those observed for many of the largest reservoirs in the United States
(Bolgrien et al., 2009; Bowersox et al., 2014; Lynott et al., 1995), we sug-
gest that our ndings provide a useful model for lacustrine regulatory
processes in reservoirs in much of central and western North America.
Specically, we forecast that future declines in river inow from moun-
tain snowelds (Lapp et al., 2013; St. Jacques et al., 2010) should reduce
the inorganic turbidity, whereas continued economic development that
elevates nutrient supply will likely favor increased phytoplankton
abundance and possibly cyanobacterial outbreaks in lacustrine embay-
ments (Donald et al., 2011; Leigh et al., 2010). Consequently, selection
of the appropriate management strategy will depend on whether such
reservoirs are exploited principally for industrial, recreational or
domestic uses.
Phytoplankton abundance
Moderate phytoplankton abundance (Fig. 1) and diatom-rich com-
munity composition (Fig. 2) are consistent with observations in other
large temperate reservoirs in the Great Plains of North America
(Bolgrien et al., 2009; Joeckel and Diffendal, 2004). In these large
lakes, hydrologic and nutrient inux (Finlay et al., 2009; Patoine et al.,
2006) is controlled mainly by snowmelt in westerly mountain ranges
during early summer (Lapp et al., 2013; St. Jacques et al., 2010), rather
than local snowmelt during March and April (Fang and Pomeroy,
2007; Pham et al., 2009; Quiones-Rivera et al., 2015). Typically,
water from mountain ranges is cold, has lower nutrient content than
seen in natural prairie lakes (Pham et al., 2008, 2009), and exhibits
high turbidity associated with suspended inorganic particulates, leading
B)
50.00
45.00
Copepods
40.00
Large Cladocerans
Zooplankton (indL-1)
35.00 Small Cladocerans
30.00
25.00
20.00
15.00
10.00
5.00
0.00
2004 2007 2010 2013 1995 1998 2001 2004 2007 2010 2013
Year
to low algal production (Dubourg et al., 2015). Diatoms predominate
when turbulence related to inow is greatest (spring, early summer)
(Hall et al., 1999a), while agellates (cryptophytes, chlorophytes, dino-
agellates) are increasingly common as inow slows, turbulence de-
clines, and thermal stratication may occur (Abirhire et al., 2015;
Drscher et al., 2009; McGowan et al., 2005b). These patterns are in
marked contrast to many natural lakes in the prairie ecozone (Donald
et al., 2013; Leavitt et al., 2006; Pham et al., 2008), as well as some
large reservoirs from continental interiors of China (Lv et al., 2014),
Australia (Bowling et al., 2013), and South America (Fernandez et al.,
2014), where high turbidity reects dense blooms of cyanobacteria aris-
ing from excessive supply of P and N (Leigh et al., 2010; Sukenik et al.,
2012). Differences in production between the large reservoirs of western
North America and natural prairie lakes may reect differences in the
predominant source of nutrients to the water-column of articial (al-
lochthonous, riverine) and natural aquatic ecosystems (autochthonous,
sedimentary) (e.g., Leavitt et al., 2006; Abirhire et al., 2015; Patoine
et al., 2006).
Absence of decadal-scale trends in water quality is consistent with
observations from other large prairie lakes (Bunting et al., 2011;
Patoine et al., 2006), Landsat analysis of this reservoir (Yip et al.,
2015), and recent paleolimnological analysis of phytoplankton fossils
in Lake Diefenbaker sediments (Yip et al., 2015). Although phytoplank-
ton abundance has increased N300% above baseline in many large prai-
rie lakes, most cultural eutrophication occurred by 1990 mainly due to
intensication of grain and livestock production during the 20th centu-
ry (Bunting et al., 2011; Hall et al., 1999a). Similarly, although the down-
stream reaches of Lake Diefenbaker experienced a small trophic surge
for several years following inundation of the valley in 1967 (Hall et al.,
1999a; Lucas et al., 2015), phytoplankton abundance declined before
expanding again to a stable plateau in the late 1980s and early 1990s
(Tse et al., 2015). Although speculative, recent research suggests that
modern water quality conditions may be the result of an interaction be-
tween long-term land-use patterns and a sudden persistent change in
climatic forcing within the northern Great Plains (McCullough et al.,
2012).
Multiple regression models suggest that nutrient supply from
reservoir sediments during summer may be an important control of
decadal variability in total phytoplankton abundance. Specically,
depth-integrated estimates of phytoplankton abundance were correlat-
ed positively with that of SRP, whereas the latter was correlated
negatively with water-column oxygen level and positively with DIC
content (Table 2). Bottle bioassays conducted biweekly during summers
 R.J. Vogt et al. / Journal of Great Lakes Research 41 Supplement 2 (2015) 8190 87

A) Water Quality Models

100

90

80

70
% Variation Explained
Internal
60
Hydrology
50
Climate
40
R2adj = 0.48

R2adj = 0.52
30

20

10

0
Total Phytoplankton Water Clarity

B) Specific Predictor Variables

Total
Water Clarity
Phytoplankton

+ - + -
60% 40% 30% 70%

SRP Zoop. PDO-ENSO Inflow

Fig. 4. (A) Proportion of total explained variation (open bars) and sources of explained variation (lled bars) in total phytoplankton abundance (as g Chl a/L) and water clarity (as Secchi
depth, m) as identied in multiple regression analysis of mean summer (May-August) conditions in the Qu'Appelle arm of Lake Diefenbaker, Canada, 19952010. Model performance was
evaluated by adjusted coefcient of determination (R2adj). Signicant (p b 0.05) explanatory variables were selected by forward selection multiple regression based on 9999 permutations
and were classied into categories associated with variation in climate systems (black), river hydrology (dotted), and internal lake characteristics (waves). (B) Potentially-causal relation-
ships among water quality variables and independent predictors were identied from multiple regression models (Table 2). Predictors include soluble reactive phosphorus (SRP), mean
total zooplankton density (Zoop.), indices of interactions between the Pacic Decadal Oscillation (PDO) and El Nio-Southern Oscillation (ENSO), and riverine inow to Lake Diefenbaker
(Inow). Numbers and signs reect the proportion of explained variation in water quality parameters (ovals) attributable to variation in predictor variables (squares) due to inhibitory (-)
or stimulatory (+) interactions.

since 1994 as part of the QU-LTER program reveal that while most down-
stream Qu'Appelle lakes exhibit pronounced algal limitation by supply of
N (Donald et al., 2011; Leavitt et al., 2006), instantaneous algal growth in
the lacustrine portion of Lake Diefenbaker is regulated mainly by P sup-
ply (Dubourg et al., 2015; Hecker et al., 2012; Quiones-Rivera et al.,
2015). Further, diazotrophic cyanobacteria are rare (Fig. 2) (Abirhire
et al., 2015; McGowan et al., 2005b) and there is no net xation of atmo-
spheric N2 in most years (Patoine et al., 2006). Instead, Lake Diefenbaker
is a substantial net sink for P ux through the South Saskatchewan River
drainage basin (North et al., 2015). As shown by Nrnberg (1998),
water-column P concentrations are thought to be inuenced by the de-
velopment of deepwater hypoxia (but see Hupfer and Lewandowski,
2008), mainly due to declines in the effectiveness of the oxidized
microzone as a P trap with loss of O2 (reviewed in Wetzel, 2001).
Although not directly measured in this study, the positive correlation
between SRP and DIC concentrations is also consistent with the effects
of organic matter decomposition in sediments both reducing oxygen
tensions and increasing water-column DIC levels, as respiratory CO2
reacts with insoluble sedimentary carbonate to produce two dissolved
bicarbonate molecules (Wetzel, 2001). Consistent with this hypothesis,
sediments of the Qu'Appelle arm of Lake Diefenbaker are over 50%
inorganic carbonate by mass (Finlay et al., 2010) and appear to be a
signicant source of P to the water column (North et al., 2015).
Internal nutrient loading is also hypothesized to be responsible for
decade-long surges in planktonic production in reservoirs. In this case,
ooding of dry river valleys leaches soluble nutrients from formerly
terrestrial soils, erodes newly formed shorelines, and decomposes
inundated vegetation (reviewed in Thornton et al., 1990). Oxygen con-
sumption during vegetation decomposition favors deepwater anoxia,
further increasing nutrient ux from sediments (see above). However,
continued erosion of shorelines also buries submerged organic sub-
strates, reduces oxygen demand and nutrient release, and reduces pro-
duction by benthic communities in favor of planktonic assemblages,
leading to a decline in lake production (Hall et al., 1999a; Hewlett
et al., 2015; Thornton et al., 1990). Although this ontogenetic pattern
can be modied by differences in the original habitat type (terrestrial,
88 R.J. Vogt et al. / Journal of Great Lakes Research 41 Supplement 2 (2015) 8190

Table 2
Summary of regression models predicting total phytoplankton abundance (as g Chl/L), water clarity (Secchi depth, m), soluble reactive phosphorus concentration (g P/L) and total
zooplankton density (individuals/L). Models are based on 16 years of monitoring in the Qu'Appelle arm of Lake Diefenbaker, Canada (MayAugust, 19952010). Model performance is
summarized using an adjusted coefcient of determination (R2adj), and models were signicant with a probability level of p. Multiple regression models include predictor variables for
each water quality index selected using a Monte Carlo forward selection. Predictors selected include river inow (m3/month), indices of Pacic Decadal Oscillation (PDO) and El Nio
Southern Oscillation (ENSO), oxygen concentration (mg O2/L), inorganic carbon concentration (TIC, mg C/L), and dissolved organic C content (mg C/L).

Water quality variable Linear regression Multiple regression

R2adj p Slope R2adj p Predictor Coefcient % total variation % explained

explained variation

Total phytoplankton 0.02 0.09 0.60 SRP total zooplankton 0.48 0.005 SRP 1.67 29.0 60.4
(g Chl/L) Total zoop 2.29 19.0 39.6
Residual 52.0
Intercept 8.97
Water clarity 0.00 0.57 0.00 Inow + PDOENSO 0.52 0.003 Inow 2.74 108 36.6 70.5
(m) PDOENSO 7.5 103 15.3 29.5
Residual 48.1
Intercept 1.68
Soluble reactive phosphorus 0.001 .33 0.05 O2 + TIC 0.48 0.006 O2 0.42 34.6 72.6
(g P/L) TIC 0.09 13.0 27.4
Residual 52.4
Intercept 2.95
Total zooplankton 0.38 0.004 0.07 DOC 0.21 0.04 DOC 0.09 21.0 100.0
(ind./L) Residual 79.0
Intercept 3.66

aquatic), area of ooded land, degree of shoreline development, valley
geometry, water turbidity and magnitude of water level uctuations
(Hecky and Guildford, 1984; Thornton et al., 1990), studies generally
agree that autothchonous nutrient sources can be important controls
of decadal-scale development of reservoir productivity independent of
allochthonous sources (Thornton et al., 1990). Better estimation of the
importance of internal nutrient sources is essential for Lake Diefenbaker
and other large reservoirs in central North America (North et al., 2015),
as these water bodies supply much of the potable water to urban cen-
tres in the continental interior (e.g., Saskatchewan Water Security
Agency, SWSA, 2012).
Recent changes in the grazing regime in Lake Diefenbaker may have
also had an important effect on phytoplankton abundance during the
past 20 years (Table 2, Fig. 3). While quantitative estimates of sh abun-
dances are not available, the marked and persistent decline in density of
large zooplankton (Fig. 3) coincides with a documented escape of
~ 360,000 rainbow trout from a local sh farm in the year 2000
(Saskatchewan Water Security Agency, SWSA, 2012; Saskatchewan
Environment, unpublished data). This event was one of a series of
unintended releases during the past 15 years. Rainbow trout have a
varied diet in large reservoirs of central North America, but prefer
large-bodied zooplankton and other invertebrates when sh are up to
46 cm in length (Lynott et al., 1995). Consistent with the effects of
size-selective predation by visually-orienting sh, densities of large-
bodied Daphnia and copepods declined nearly 75% between 2000 and
2012 (Fig. 3a), whereas the abundance of small cladocerans was un-
changed (Fig. 3b). Similarly, research on other large western lakes
(e.g., L. Tahoe) reveals that stocked rainbow trout prefer D. thomasii
(Elser et al., 1995), one of the two predominant copepods in eastern
Lake Diefenbaker. As noted by Vogt et al. (2011), D. thomasii and
the equally large-bodied Leptodiaptomus siciloides are the biomass-
dominant taxa in the Qu'Appelle arm and other regional lakes, and
both taxa are strongly omnivorous (Patoine et al., 2006). Finally, analy-
sis of time series with monthly resolution demonstrates that phyto-
plankton abundance increased only during July, a month in which
edible agellates are most common (Fig. 2), densities of large herbi-
vores are normally greatest (Drscher et al., 2008, 2009), and relaxation
of predation by sh would be expected to have large effects on phyto-
plankton abundance. Thus while variation in invertebrate grazing ap-
peared to explain less variation (19%) in phytoplankton abundance
than does inter-annual variation in SRP supply (29%), our regression
analysis and supporting evidence suggest that decadal-scale variation
in phytoplankton abundance is under mainly trophic control.
Water clarity
Multiple regression analysis shows that inter-annual variation in
water clarity is regulated mainly by changes in climatic conditions
that regulate river discharge and inorganic turbidity (Table 2). Speci-
cally, the Qu'Appelle arm of Lake Diefenbaker was more transparent in
years with dry winters (ENSO-PDO) and low discharge of the South
Saskatchewan River (inow). Runoff to reservoirs in the Northern
Great Plains is derived from two main water sources; local snowmelt
during MarchApril (Fang and Pomeroy, 2007; Pham et al., 2009), and
inux of alpine meltwaters via east-owing rivers in June (e.g., South
Saskatchewan, Missouri) (Quiones-Rivera et al., 2015; St. Jacques
et al., 2010). While the former combines with regional groundwater
sources to sustain natural lakes in the Canadian Prairies (Pham et al.,
2009; van der Kamp et al., 2008), discharge of large prairie rivers that
regulate reservoir hydrology is controlled mainly by release in alpine
snow packs and upstream hydrologic management (Lapp et al., 2013).
Timing and magnitude of release of water from mountain sources
appears to be controlled in part by variation in the PDO (Lapp et al.,
2012, 2013; Schindler and Donahue, 2006), with anomalously low dis-
charge occurring when ENSO and PDO systems reinforce each other to
split the winter jet stream and induce warm dry winters in western
Canada and northern USA (Mantua et al., 1997; McCabe et al., 2004;
Shabbar et al., 2011). In general, years with low river discharge result
in lower turbidity associated with suspended solids (Thornton et al.,
1990), because of both lower inux and reduced erosion of reservoir
shorelines (Coakley and Hamblin, 1969; Hewlett et al., 2015).
Interestingly, although both ENSO and PDO systems are known to regu-
late spring climate, trophic interactions, phytoplankton composition
and primary production (Drscher et al., 2009; Weyhenmeyer et al.,
1999), our regression analysis revealed no evidence of direct effects of
climate, hydrology or regional meteorological condition on mean sum-
mer phytoplankton abundance or composition within the Qu'Appelle
arm (Table 2). Instead, we conclude that water clarity and phyto-
plankton abundance in large lacustrine reaches are under parallel
but largely independent regulatory mechanisms (physical and trophic,
respectively).
 R.J. Vogt et al. / Journal of Great Lakes Research 41 Supplement 2 (2015) 8190
Forecasts of reservoir response to climate and anthropogenic stressors
General circulation models and downscaled regional climate models
predict that the North American grasslands will warm ~4 C within 40
years (Barrow, 2009). Such effects may be particularly noticeable in cen-
tral Canada, where minor changes in mean annual temperatures (~1 C)
change the state of water (solid, liquid). Although precipitation on the
northern Great Plains is expected to increase and become more variable
in coming decades, elevated evaporation associated with atmospheric
warming is expected to offset increased receipt of snow and rain, pro-
ducing a more arid climate (Lapp et al., 2012). We infer that river inow
to large prairie reservoirs (Diefenbaker, Oahe, Sakakawea, Fort Peck)
will remain a key regulatory mechanism in the future, given that base
ow in prairie rivers has already declined nearly 50% since 1900 due
to climate change and industrial use (St. Jacques et al., 2010), future
PDO systems may be weaker than at present (Lapp et al., 2012), and
the snowmelt in the Rockies is expected to occur earlier and decline in
magnitude (St. Jacques et al., 2010). Based on our analyses, we antici-
pate that these declines will favor less turbid water suitable for domestic
and agricultural consumption, particularly in downstream lacustrine
regions.
Continued strong urban and industrial growth in much of the Cana-
dian Prairies, combined with expected economic recovery in the central
United States, may increase nutrient inux to regional reservoirs. For
example, application of agricultural fertilizers has risen steadily since
1960 and is expected to nearly double again by 2050 (Bogard et al.,
2012; Donald et al., 2011). Similarly, stronger commodity prices
(grain, oil, potash), combined with plentiful regional reserves, suggest
that anthropogenic nutrient inux to large reservoirs will continue to
increase due to future economic development. While these nutrient
sources do not appear to have degraded water quality in eastern Lake
Diefenbaker during the past 20 years (Fig. 1, Table 2) (Tse et al., 2015;
Yip et al., 2015), modern and paleolimnological analysis of large region-
al water bodies suggest that anthropogenic effects may develop slowly,
before reaching a tipping point beyond which water quality changes
suddenly (Bunting et al., 2011). Further research is needed to determine
whether increased phytoplankton production after 1990 reects such a
threshold effect (Bunting et al., 2011), the result of a trophic upsurge
during reservoir ontogeny (Hall et al., 1999a; Thornton et al., 1990), or
a sudden change in regional climatic forcing mechanisms
(McCullough et al., 2012).
Regression analysis of nearly two decades of limnological monitor-
ing suggests that optimal management strategy for lacustrine reaches
of large reservoirs within central North America may depend on the rel-
ative importance of industrial, recreational and domestic uses of the
ecosystem, as well as whether the reservoir is managed to optimize
phytoplankton abundance or water clarity. Longer reservoir residence
times will favor less turbid water suitable for consumption by livestock
and humans, although this strategy will likely reduce generation of hy-
droelectric power (Saskatchewan Water Security Agency, SWSA, 2012)
and may favor potentially-toxic blooms of cyanobacteria (Fig. 2) be-
cause of reduced dilution of internal nutrient sources and increased
water clarity (Dubourg et al., 2015). Similarly, management for industri-
al aquaculture will likely increase nutrient supply (Figueredo and Giani,
2005; Guo and Li, 2002), while reducing herbivore control of phyto-
plankton (Fig. 3, Table 2). Finally, managers must recognize that specic
strategies are likely to interact with continued economic growth and cli-
matic variability to produce complex, hierarchical pathways of ecosys-
tem regulation (Leavitt et al., 2009).
Acknowledgments
We thank the members of the Limnology Laboratory for assistance
with data collection since 1994, and Curtis Halborg and Bart Oegema
at the Saskatchewan Water Security Agency for estimates of river
discharge. This work was supported by the Natural Sciences and
89
Engineering Research Council of Canada Discovery Grants (NSERC),
the Canada Research Chair Program, Canada Foundation for Innovation,
the Province of Saskatchewan, and the University of Regina, York
University and Trent University. We thank two reviewers and editor
Rebecca North for valuable comments that improved this paper.
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