Arctic, Antarctic, and Alpine Research, Vol. 40, No. 3, 2008, pp.

584–591

Evaluating the Response of Cladocera to Recent Environmental Changes in Lakes

from the Central Canadian Arctic Treeline Region

Jon N. Sweetman*{ Abstract

Elyse LaFace* Arctic and subarctic ecosystems have undergone considerable environmental
Kathleen M. Rühland* and changes in recent years as the result of climate warming. Fossil records of freshwater
diatoms in lakes throughout the circumpolar Arctic, including the central Canadian
John P. Smol* Arctic treeline region, have revealed marked directional shifts in diatom assemblages
*Paleoecological Environmental in recent lake sediments. These algal changes have been linked to longer growing
Assessment and Research Laboratory seasons, decreased duration of ice cover, and/or increased thermal stability. The
(PEARL), Department of Biology,
effects of these recent environmental changes on higher trophic levels, such as the
Queen’s University, Kingston, Ontario,
K7L 3N6, Canada Cladocera, are still unclear. Using cladoceran remains preserved in the sediments of
{Corresponding author: 50 lakes, which were previously examined for changes in diatoms, we show that
[email protected] significant changes in cladoceran species composition have occurred from pre-
industrial times to the present. However, these changes are considerably muted
compared to the more substantial changes observed in the diatom record. We found
no consistent patterns of change in planktonic cladocerans (i.e. Daphnia, Bosmina)
within our study lakes, and the response of the Cladocera to environmental changes
does not appear to be strongly coupled to recent changes in diatom communities,
thus further confirming that the previously observed diatom changes were related to
bottom-up limnological controls. These results highlight the complex response of
Arctic freshwater food webs to climate change, and the need for incorporating multi-
trophic studies into climate change investigations.

DOI: 10.1657/1523-0430(06-118)[SWEETMAN]2.0.CO;2

Introduction planktonic taxa (Sorvari and Korhola, 1998; Rautio et al., 2000;
Korhola et al., 2002; Sorvari et al., 2002.). Likewise, Rühland et
Freshwater lakes and ponds are one of the most ubiquitous al. (2003a) recorded similar shifts in a series of 50 lakes in the
features of Arctic and subarctic landscapes, and represent central Canadian Arctic treeline region, with a shift from benthic
environments that are highly susceptible to the effects of climate to planktonic taxa. These widespread shifts in Arctic diatom taxa
change. Global warming trends are expected to be amplified at have been attributed to a shorter duration of ice cover, an increase
higher latitudes (Serreze and Francis, 2006), and, because of the in the length of the growing season, and/or increased thermal
sensitivity of hydro-ecological processes (Rouse et al., 1997; stability resulting from climate warming (Smol et al., 2005).
Wrona et al., 2005), freshwater ecosystems in the north are The Cladocera (Crustacea: Branchiopoda) are an important
predicted to be particularly vulnerable to climate stressors. component of many high-latitude lakes and ponds, and are
Numerous studies have suggested that high-latitude freshwater considered to be key organisms in both pelagic environments (i.e.
ecosystems have undergone significant environmental changes the Daphniidae and Bosminidae families) as well as benthic
since pre-industrial times, including increasing lake temperatures, habitats (i.e. the Chydoridae family) (Frey, 1988). Because of their
increased fluxes of nutrients, reduced periods of ice cover, longer crucial role in aquatic food webs, and their abilities to respond
growing seasons, and substantial reorganization of aquatic food rapidly to changing environmental conditions, cladocerans have
webs (e.g. Serreze et al., 2000; Wrona et al., 2005; Hinzman et al., long been recognized as important ecological and paleolimnolo-
2005; Smol et al., 2005; Quinlan et al., 2005; Duguay et al., 2006; gical indicators (Korhola and Rautio, 2001; Jeppesen et al., 2001;
Schindler and Smol, 2006). Bennike et al., 2004). As the chitinous exoskeletons of these
Several striking examples of the impacts of climatic warming animals are well preserved in lake sediments, changes in the
on freshwater ecosystems in the Arctic have been provided by sediment record can be used to examine the past response of
paleolimnological studies of diatom communities. Douglas et al. cladoceran communities to changing environmental conditions
(1994), for example, recorded dramatic shifts in diatom assem- (Korhola and Rautio, 2001). The environmental conditions of
blages over the last ,200 years in a set of small High Arctic ponds northern lakes influence the distributions and abundances of
on Ellesmere Island, Canada, whereas relatively stable diatom cladocerans (e.g. Korhola 1999; Bos and Cumming, 2003;
assemblages were found to have persisted throughout the Jeppesen et al., 2003; Sweetman and Smol, 2006) and, as a result,
preceding millennia. Smol et al. (2005) documented a similar taxa in Arctic and subarctic lakes should also be expected to be
restructuring of diatom communities in post-industrial lake strongly affected by climate warming.
sediments throughout most regions of the circumpolar Arctic. In Compared to the well documented shifts in diatom communi-
subarctic Finnish Lapland, diatom assemblages shifted from being ties (e.g. Rühland et al., 2003a; Smol et al., 2005), however, the
primarily composed of benthic taxa to having high abundances of responses of higher trophic levels, including cladocerans, to recent

584 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH E 2008 Regents of the University of Colorado
1523-0430/08 $7.00
 FIGURE 1. Locations of the 50
lakes across Arctic treeline. Study
sites are indicated by black cir-
cles, with the exception of Slipper
Lake, whose location is shown as
a white circle.

environmental changes are not as well understood. In their review of cladoceran assemblages from a series of 50 lakes, and to compare the
the effects of climate change on Arctic and subarctic freshwater responses of the cladoceran communities to previously observed
ecosystems, Rouse et al. (1997) predicted that, while increases in changes in diatom communities from the same lakes (Rühland et al.,
primary production will almost certainly be observed, the responses 2003a). Specifically, we were interested in knowing the following: (1)
of higher trophic levels are likely much more variable. As food Have any changes occurred to the community composition or
resources for planktonic grazers increase with higher levels of diversity of cladocerans between pre-industrial and present-day
primary productivity in these ecosystems (e.g. Michelutti et al., periods? (2) If so, was there a shift in abundances from benthic to
2005), the abundance of herbivorous cladocerans, such as Daphnia, planktonic cladoceran taxa similar to that observed in the diatom
might also be expected to increase. In some systems, this appears to taxa? (3) Are changes in cladoceran and diatom assemblages
be the case. For instance, in Lake Saanajärvi, located in contemporaneous? (4) Finally, do changes in grazing/predation
northwestern Finnish Lapland, Rautio et al. (2000) and Korhola regimes from cladocerans have a role in driving the response of
et al. (2002) found that increases in the planktonic cladoceran diatom communities to recent climate warming? If the latter is the
Daphnia were synchronous with the increases in planktonic diatom case, we would expect changes in cladoceran assemblages to closely
taxa described previously. O’Brien et al. (2005) also found that, correspond to the diatom changes across our 50 lake set.
following experimental fertilization of half of a partitioned lake in
northern Alaska, Daphnia longiremis abundance was significantly
higher in the treated half compared to the reference side. However,
Study Area
the response of food webs to climate warming is often difficult to Sediments were analyzed from 50 lakes located in the central
predict (Petchey et al., 1999; Callaghan et al., 2004; Baulch et al., Canadian Arctic treeline region, between Yellowknife, Northwest
2005). Climatic warming can potentially destabilize food webs, and Territories, and the northern boundary of the Thelon Game
responses of higher trophic levels may be much more complex and Sanctuary, Nunavut Territory (Fig. 1). The study lakes span a
unpredictable than in algal communities (e.g. Beisner et al., 1997; steep ecoclimatic gradient and are distributed across three major
Strecker et al., 2004). There is increasing evidence that in many ecozones, from the boreal forest zone in the south, through the
temperate lakes, a decoupling of the algae-zooplankton relationship forest-tundra transitional zone, and into the Arctic tundra zone in
can occur (Winder and Schindler, 2004; Adrian et al., 2006). In Lake the north (Rühland and Smol, 2002; Rühland et al., 2003a, 2003b).
Washington, for example, Winder and Schindler (2004) documented
a long-term decline in Daphnia, which they attributed to an
increasing temporal mismatch with food resources, due to an earlier Methods
spring diatom bloom as the result of climate warming. Relatively few Sediment cores were collected at approximately the deepest
studies, however, have examined the impacts of climate change on point of each of the lakes using a Glew gravity corer (Glew et al.,
Arctic freshwater food webs (e.g. Quinlan et al., 2005). 2001). The surface sediments (top 1.0 cm) and bottom sediments
The objectives of our study were to evaluate the response of (bottom 1.0 cm) of each core were extruded using a vertical
cladocerans to recent environmental changes in the central Canadian extruder (Glew, 1988). Additional details on water chemistry of
Arctic treeline region by comparing present-day and pre-industrial the lakes, the modern distribution of diatoms and Cladocera, as

J. N. SWEETMAN ET AL. / 585

well as additional details about sediment core retrieval and Bonferroni-adjusted paired t-tests were used to compare
sampling of these lakes were previously presented in Rühland and differences in diversity between time intervals. All ordinations
Smol (2002) and Rühland et al. (2003a, 2003b). and Hill’s N2 calculations were carried out using CANOCO v.
In order to evaluate the response of cladocerans to recent 4.5 (ter Braak and Šmilauer, 2002); t-tests were calculated using
environmental change, we used a ‘top-bottom’ paleolimnological SYSTAT v. 9 (SPSS, 1998).
approach (Smol, 2008), identical to the approach used in the In order to assess how changes in cladoceran communities
Rühland et al. (2003a) diatom study. This ‘snap-shot’ approach matched previously observed changes in diatom communities
involves comparing subfossil assemblages from the surface lake within the same lakes (Rühland et al., 2003a), the degree of
sediments, which represent present-day conditions, to assemblages dissimilarity between top and bottom samples was calculated
deposited in downcore lake sediments representing pre-impact using a Bray-Curtis dissimilarity coefficient (Clarke et al., 2006).
conditions (ca. 1850). The goal of the comparisons made between Cladoceran Bray-Curtis dissimilarity coefficients were compared
these two discrete points in time (i.e. top vs. bottom) is to assess to the matching Bray-Curtis values for the diatom assemblages. In
whether environmental conditions during the pre-industrial era are addition, we compared the relative abundance of the dominant
substantially different than present. These data can then be used to planktonic diatom taxa (Cyclotella) to changes in pelagic
evaluate the magnitude, direction, and nature of cladoceran Cladocera species, using simple Pearson correlations with Bon-
assemblage changes over the last ca. 200 years on a regional scale. ferroni-adjusted p-values for multiple comparisons. Correlations
In addition to the 50-lake regional top-bottom analyses, we were calculated using SYSTAT v. 9 (SPSS, 1998).
examined the detailed changes in cladoceran communities from a For the continuous sediment core data from Slipper Lake, a
sediment core from Slipper Lake (64u359650N, 110u509070W), a detrended canonical correspondence analysis (DCCA) was used
site also examined for changes in diatom communities by Rühland to calculate a quantitative estimate of compositional turnover (b-
et al. (2003a; 2005). As there was insufficient sediment from the diversity) following the procedures used by Birks (2007). An
original Slipper Lake core used by Rühland et al. (2003a), we estimate of the total amount of compositional change for the last
examined cladoceran remains from a second core that was used by ,150 years was obtained by constraining the DCCA by using the
Rühland and Smol (2005) to verify the trends in the primary core. 210
Pb-based sample ages as the sole constraint. Significance was
Both sediment cores were collected at the deepest point in the lake measured using Monte Carlo permutations (999 permutations).
in March 1997 and sectioned into 0.5 cm intervals. For this analysis, species data were square-root transformed,
Analysis of cladoceran remains from the sediments of each there was no down-weighting of rare taxa in the ordination,
lake followed standard procedures, outlined in Korhola and detrending was by segments, and scaling was nonlinear. DCCA
Rautio (2001). Approximately 2–4 g wet weight of sediment from was performed using CANOCO v. 4.5 (ter Braak and Šmilauer,
each sample was deflocculated in 150 mL of a 10% KOH solution 2002).
at 80uC for 30 minutes, after which they were rinsed through a
37 mm sieve. For Slipper Lake, the 0.5 cm intervals were combined
into 1.0 cm intervals because of the low abundance of cladoceran Results
remains. Slipper Lake samples were separated into .90 mm and
37–90 mm subsamples to facilitate identification of remains. Most cladoceran taxa in our study lakes did not demon-
Material retained on the sieves was stored in distilled water and strate a uniform pattern of change from the pre-industrial to the
ethanol. Safranin-glycerine solution was added to stain the present-day assemblages (Fig. 2). This is in contrast to the
cladoceran remains. Slides were prepared by pipetting 50 mL diatom communities in the same lakes that showed a large and
aliquots onto slides, and mounting in glycerin jelly. Cladocera consistent increase in the planktonic species Cyclotella stelligera,
were identified at 100–4003 magnification using a Leica DMRB and a corresponding decline in benthic Fragilaria taxa (Fig. 3;
compound microscope. For each taxon, the most abundant Rühland et al., 2003a). The cladoceran assemblages show no
remains (i.e. head shield, carapace, postabdomen) were used to corresponding increase in planktonic species, not withstanding a
calculate the number of individuals within a sample. A minimum few lakes that showed changes in the dominant planktonic taxa
of 50 individuals were counted per sample. Three lakes (TK-13, (i.e. Bosminidae, Daphniidae). Despite no consistent increases or
TK-21, and TK-40) contained insufficient numbers of Cladocera, decreases in cladocerans across the study area, relatively subtle
and were excluded from subsequent analyses. shifts were apparent. For example, the benthic cladoceran
Changes in the cladoceran communities between top and Pleuroxus trigonellus, which was only found in lake sediments
bottom samples were quantified using detrended correspondence from boreal forest lakes, decreased in abundance in modern
analysis (DCA). DCA is an unconstrained ordination technique, sediments compared to pre-industrial periods (negative values in
and has the advantage that the sample scores are scaled in Fig. 2), whereas a second benthic chydorid taxon (Camptocercus
standard deviation (SD), or units of species turnover (b- spp.) increased in relative abundance in several lakes across the
diversity) (Smol et al., 2005; Birks, 2007). Prior to analysis, all study region (positive values in Fig. 2). However, most taxa in
species relative abundance data were square-root transformed to our lakes did not show a synchronous shift in their relative
stabilize variances. Bonferroni-adjusted paired t-tests were used abundance from pre-industrial assemblages to modern sedi-
to compare both axis 1 and axis 2 sample scores between the two ments.
time periods. Changes in the sample scores between top and The detailed paleolimnological record from the Slipper Lake
bottom sediment samples were calculated following Quinlan et sediment core is consistent with the regional top-bottom survey.
al. (2003). Briefly, the axis 1 sample score for the pre-industrial Throughout the sediment record, the cladoceran assemblage
assemblage from each lake was subtracted from the axis 1 sample remains relatively stable, and the pelagic cladocerans (Daphnia,
score of the present-day assemblage to calculate a net change in Bosmina) present in the sediment record show no apparent change
DCA sample scores along that axis. This calculation was also in their abundance (Fig. 3). This is in contrast with the large
performed for axis 2 ordination scores. Differences in a-diversity directional changes occurring in the diatom record (Fig. 3).
between present-day and pre-industrial cladoceran communities Beginning at about 5 cm (A.D. ,1800), there is a shift in the
were also compared, using Hill’s N2 as a measure of a-diversity. diatom taxa, with pelagic Cyclotella stelligera becoming more

586 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH

FIGURE 2. The percentage of change of the main cladoceran species from pre-industrial to modern lake sediments in the 47 central
Canadian Arctic treeline lakes that contained sufficient numbers of fossil Cladocera for analysis. Lakes are arranged in order of the distance of
each lake from current treeline. Distance to treeline (km) follows the delineation by Rühland et al. (2003a) and is based on percentage tree
cover, derived from field observations, topographical maps, aerial photographs, and maps derived from satellite imagery. Positive distances
along the x-axis indicate lakes north of treeline, and negative distances indicate lakes south of treeline. Positive values on the y-axis indicate an
increase in the relative abundance (%) from pre-industrial to present-day lake sediments, while negative values on the y-axis indicates a
decrease in the relative abundance (%) from pre-industrial to present-day lake sediments.

abundant and Aulacoseira and Fragilaria spp. decreasing in change in b-diversity (species turnover) (Fig. 4). Both DCA axis 1
abundance towards present day. and 2 sample scores were significantly different between present-day
Although there is no clear directional change in individual and pre-industrial times (DCA axis 1, paired t-test, p , 0.0001, t 5
cladoceran taxa across our regional lake set, there is a significant 4.470; DCA axis 2, paired t-test, p , 0.0001, t 5 5.519), indicating a

FIGURE 3. Relative abundances of littoral cladocerans, pelagic cladocerans, and dominant diatom taxa in Slipper Lake, Northwest
Territories. The two Chydorus species, C. biovatus and C. brevilabris, were grouped together, as the remains of these taxa were difficult to
distinguish from each other, as were the Alona species, A. guttata and A. barbulata. Diatom data are summarized from Rühland et al. (2003a)
and Rühland and Smol (2005).

J. N. SWEETMAN ET AL. / 587

 FIGURE 4. Changes between
(top) DCA axis 1 sample scores;
(center) DCA axis 2 sample
scores, a measure of b-diversity;
and (bottom) cladoceran species
diversity (Hill’s N2), a measure of
a-diversity, between pre-industrial
lake sediments and modern lake
sediments in the 47 central Cana-
dian Arctic treeline lakes. Lakes
are arranged in order of the
distance of each lake from current
treeline.

directional shift in the overall composition of species assemblages. record. The cladoceran fauna is comprised primarily of benthic
There was no significant change in a-diversity (Hill’s N2) between chydorids and is relatively stable throughout the core (Fig. 3).
pre-industrial and present-day cladoceran assemblages (Fig. 4;
paired t-test, p , 0.636, t 5 0.477). a-diversity refers to the diversity
(i.e. species richness) within a particular area and is usually expressed Discussion
as a measure of the number of species. Alternatively, ß-diversity is a The lowermost cladoceran assemblages from all of our study
measure of the change in species diversity between areas, allowing a sites represent environmental conditions pre-dating ca. 1850 (pre-
comparison between ecosystems or sites. industrial times). Several of our study lakes have undergone
A comparison of the Bray-Curtis dissimilarity coefficients detailed paleolimnological analyses that involved establishing a
(DC) for diatom taxa (mean DC 5 0.46) and cladocerans (mean chronology based on 210Pb dating techniques (including Slipper
DC 5 0.33) suggests that the species composition of diatom Lake). These sedimentary cores have consistently shown that
assemblages are more dissimilar between pre-industrial and background conditions (A.D. ,1850) occurred within the top 6 cm
present-day periods than the cladoceran fauna (Fig. 5). This is of the core, which is consistent with the typically low rates of
not surprising, as the diatom assemblages show striking changes in sedimentation in Arctic lakes (Rühland et al., 2003a, 2003b). The
their relative abundances (Rühland et al., 2003a). mean core depth for all of the bottom sedimentary intervals from
A comparison of the modern distributions of Cyclotella and our lake set was 19.4 m (Table 1), suggesting that most, if not all,
planktonic cladocerans reveals that the relative abundance of of our bottom intervals pre-date the onset of the industrial era.
Cyclotella spp. in our 47-lake set is uncorrelated with Daphnia Rühland et al. (2003a) found that increases in planktonic
relative abundance (r 5 0.19, p 5 0.188), Bosminidae relative Cyclotella abundance were most pronounced in the deeper lakes
abundance (r 5 0.13, p 5 0.377), or total pelagic Cladocera (r 5 (.6 m) within our study area. They suggested that, among other
0.21, p 5 0.159). Both Daphnia and Bosmina remains occur at limnological changes, deeper lakes likely experienced increased
relatively low abundances throughout the Slipper Lake core, and water column stability as the warmer temperatures and longer
neither pelagic taxa appear as dominant in the paleolimnological growing season likely allowed longer and stronger periods of
thermal stratification. One possible explanation why these deeper
lakes did not show comparable increases in planktonic Cladocera is
because of strong top-down predation pressure from fish or
invertebrate predators. While data on the occurrence of potential
predators in our study lakes were not available, it is likely that these
deeper lakes sustained fish populations. In Arctic tundra lakes in
Alaska, O’Brien et al. (2004) and Yurista and O’Brien (2001) found
that deeper lakes tend to contain planktivorous fish populations,
and in lakes with fish, large planktonic zooplankton species such as
Daphnia were generally limited in their abundance. Lauridsen et al.
(2001) also reported that in a survey of 56 lakes in Arctic Greenland,
Daphnia pulex only occurred in lakes without fish.
Invertebrate predators may also have limited the response of
Daphnia. O’Brien (2001) found that when the predatory copepod
Heterocope septentrionalis was introduced into a pond that lacked
FIGURE 5. Scatter plot comparing Bray-Curtis dissimilarity both fish and invertebrate predators, D. pulex was eliminated
coefficient values for diatoms and cladocerans for the 47 study from the pond within a year. Members of the Daphnia pulex
lakes. The majority of lakes lie above the 1:1 line, indicating that complex (i.e. D. middendorffiana and D. pulex) may not be able
diatom assemblages are more dissimilar than cladoceran assemblag- to increase in abundance in the deeper lakes, despite increased
es between pre-industrial and modern lake sediments. algal food resources, due to suppression by predators.

588 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH

 TABLE 1
Geographical and limnological data on the 50 study lakes.

Core Length
Lake Lat (N) Long (W) Area (ha.) Depth (m) Cond. (mS cm21) DOC (mg L21) TP (mg L21) Zone (cm)

SL 64u37.009 110u51.009 19.0 14.0 33.8 5.0 9.3 AT 18.5

TK-6 66u41.319 104u55.869 5.3 4.5 17.0 1.7 4.2 AT 15.0
TK-7 66u41.239 104u55.899 1.8 9.5 12.3 1.3 8.4 AT 14.0
TK-9 64u53.369 102u45.079 35.7 5.5 36.3 2.7 13.4 AT 10.0
TK-10 65u29.409 103u22.019 36.6 3.0 14.8 3.0 5.3 AT 12.0
TK-11 65u33.629 103u23.269 6.3 3.5 9.7 2.4 5.6 AT 13.0
TK-13 64u36.159 107u26.239 2.6 3.1 16.1 2.3 5.9 AT 13.0
TK-14 64u35.909 107u05.939 2.8 4.9 9.2 1.7 4.3 AT 15.0
TK-15 64u35.749 107u05.499 0.1 3.0 9.4 1.3 4.6 AT 9.0
TK-16 64u35.609 107u05.309 4.2 6.0 7.3 1.3 4.2 AT 25.0
TK-17 64u35.699 107u05.019 10.3 5.9 9.3 1.7 2.7 AT 23.0
TK-18 64u09.399 107u49.489 10.6 7.3 8.7 3.3 31.3 AT 14.5
TK-19 64u09.399 107u49.489 8.4 0.8 9.7 5.3 28.1 AT 8.5
TK-20 64u09.009 107u49.009 9.7 8.8 8.7 2.3 6.2 AT 15.0
TK-21 64u09.009 107u49.009 9.7 1.0 17.1 10.3 28.3 AT 10.5
TK-29 63u25.259 112u39.819 18.0 10.1 14.6 138.0 4.2 FT 21.0
TK-30 63u33.709 112u21.219 10.7 1.5 15.0 30.4 28.5 FT 12.0
TK-31 63u37.039 112u17.779 3.5 5.0 13.4 27.3 9.2 FT 15.0
TK-32 63u29.059 112u12.639 5.9 1.5 13.7 17.5 24.5 FT 16.5
TK-50 63u18.489 111u58.279 4.3 1.8 26.2 13.4 4.5 FT 21.5
TK-51 63u32.919 111u42.349 8.8 1.8 18.4 11.0 9.4 FT 8.0
TK-52 63u35.619 112u25.129 9.0 1.9 18.1 14.0 6.0 FT 10.5
TK-53 63u42.549 112u34.409 4.6 19.0 19.8 13.4 4.2 FT 17.0
TK-54 64u30.689 112u41.479 9.9 3.4 12.3 5.8 3.1 FT 34.0
TK-55 64u06.019 113u01.549 9.8 2.6 20.6 10.4 5.3 FT 17.0
TK-56 63u41.429 113u26.689 13.2 6.0 25.2 11.4 7.8 FT 22.5
TK-22 62u42.039 113u59.199 8.8 2.8 62.0 12.8 16.2 BF 24.0
TK-23 62u43.739 113u49.539 6.9 2.0 24.2 15.2 7.9 BF 26.0
TK-24 62u54.209 113u13.159 16.8 8.0 102.2 24.3 11.0 BF 22.0
TK-26 62u54.149 113u23.229 20.9 16.0 56.9 6.2 6.9 BF 22.0
TK-27 62u49.239 113u33.759 10.1 3.0 64.6 11.0 23.0 BF 25.0
TK-28 63u15.319 113u00.229 8.8 6.1 74.8 165.0 10.8 BF 17.0
TK-33 63u12.979 112u36.059 6.1 10.0 2.5 116.0 6.1 BF 24.0
TK-34 63u07.649 112u47.679 3.6 5.0 25.9 15.0 12.1 BF 28.0
TK-35 63u02.729 113u06.469 4.3 7.0 44.1 332.0 15.6 BF 22.0
TK-36 62u45.759 113u28.639 4.2 1.5 39.1 80.1 23.3 BF 22.0
TK-38 62u33.599 114u01.349 17.4 2.1 64.4 26.5 8.7 BF 24.0
TK-39 62u32.899 113u57.139 14.4 1.0 271.1 69.9 48.6 BF 28.5
TK-40 62u32.879 113u56.099 7.9 2.2 238.9 30.2 11.2 BF 21.5
TK-41 62u34.099 114u01.749 4.3 0.5 149.5 51.6 13.0 BF 16.5
TK-42 62u32.759 113u53.409 8.4 0.9 138.5 44.8 34.8 BF 23.0
TK-43 62u31.499 113u50.209 7.3 7.3 42.8 13.4 3.7 BF 23.5
TK-44 62u31.499 113u50.209 2.9 1.4 134.0 26.2 5.7 BF 21.5
TK-45 62u31.019 113u46.769 8.0 6.5 228.9 24.8 8.7 BF 22.7
TK-46 62u30.449 113u39.919 2.6 0.7 405.1 61.1 10.9 BF 10.5
TK-47 62u40.699 113u51.039 4.6 2.5 32.8 17.4 8.5 BF 28.0
TK-48 62u48.279 113u26.129 18.5 1.5 100.2 24.6 22.1 BF 31.5
TK-49 63u10.999 112u22.839 12.2 3.7 97.4 9.6 5.5 BF 28.5
TK-57 63u20.219 113u42.309 3.0 4.6 27.6 16.4 6.0 BF 14.0
TK-58 62u53.379 114u04.249 2.6 3.2 76.9 26.2 10.8 BF 32.5
Mean 9.3 4.7 58.4 31.2 12.0 19.4

* AT 5 Arctic tundra; FT 5 forest-tundra; BF 5 boreal forest

Cladocerans make up only a component of the zooplankton flora within the phytoplankton may have increased significantly,
of lakes. Other zooplankton taxa, such as copepods or rotifers other algal groups may not have responded to recent environ-
populations, may have increased their abundance and prevented a mental changes to the same degree (Prowse et al., 2006). In a
response in the planktonic Cladocera because of competitive study of food sources and feeding rates in shallow high-latitude
exclusion (MacIsaac and Gilbert, 1989; Dzialowski and O’Brien, lakes and ponds, Rautio and Vincent (2006) found that
2004). zooplankton and phytoplankton communities were not strongly
The paleolimnological record indicates that diatoms have coupled, and that 89–98% of the zooplankton resources
shown a large shift in favor of planktonic taxa. While the diatom consisted of detritus. Despite the apparent changes in planktonic

J. N. SWEETMAN ET AL. / 589

algae, cladocerans may not have shown a similar response Bennike, O., Brodersen, K. P., Jeppesen, E., and Walker, I. R.,
because of a decoupling of the food web (i.e. Brönmark and 2004: Aquatic invertebrates and high latitude palaeolimnology.
Weisner, 1996). In Pienitz, R., Douglas, M. S. V., and Smol, J. P. (eds.), Long-
term environmental change in Arctic and Antarctic lakes.
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Birks, H. J. B., 2007: Estimating the amount of compositional
Conclusions
change in late-Quaternary pollen—Stratigraphical data. Vege-
Despite evidence for marked ecological shifts in the diatom tation History and Archaeobotany, 16: 197–202.
communities within the past ,150 years, the cladoceran assem- Bos, D. G., and Cumming, B. F., 2003: Sedimentary cladoceran
blages within our treeline lakes do not appear to have experienced remains and their relationship to nutrients and other limnolog-
comparable ecological changes. Rühland et al. (2003a) document- ical variables in 53 lakes from British Columbia, Canada.
Canadian Journal of Fisheries and Aquatic Sciences, 60:
ed striking shifts in the fossil diatom records within these same
1177–1189.
lakes, with the relative abundances of planktonic diatom taxa Brönmark, C., and Weisner, S. E. B., 1996: Decoupling of
increasing across our study region. Although we found that there cascading trophic interactions in a freshwater, benthic food
was a significant unidirectional shift in the composition of chain. Oecologia, 108: 534–541.
cladoceran species between pre-industrial and modern periods Callaghan, T. V., Bjorn, L. O., Chernov, Y., Chapin, T.,
within the same study lakes, we found no consistent pattern of Christensen, T. R., Huntley, B., Ims, R. A., Johansson, M.,
change in the planktonic cladocerans (i.e. Daphnia, Bosmina). The Jolly, D., Jonasson, S., Matveyeva, N., Panikov, N., Oechel, W.,
lack of a strong coupling between these two trophic levels suggests Shaver, G., Schaphoff, S., and Sitch, S., 2004: Effects of changes
in climate on landscape and regional processes, and feedbacks to
that cladoceran and diatom communities may be responding
the climate system. Ambio, 33: 459–468.
independently to climate change signals within our lakes, and that
Clarke, K. R., Somerfield, P. J., and Chapman, M. G., 2006: On
top-down controls (i.e. grazing) do not appear to have a strong resemblance measures for ecological studies, including taxo-
impact in controlling diatom communities within these lakes. If nomic dissimilarities and a zero-adjusted Bray-Curtis coefficient
this is true, and given the relatively muted shift in cladoceran for denuded assemblages. Journal of Experimental Marine
assemblage composition, it is probable that the cladoceran fauna Biology and Ecology, 330: 55–80.
in subarctic Canadian lakes may not have reached an equivalent de Senerpont, L. N., Mooij, W. M., Hülsmann, S., van Nes, E. H.,
environmental threshold which the diatom communities have and Scheffer, M., 2006: Can overwintering versus diapausing
apparently surpassed. Cladoceran communities may be relatively strategy in Daphnia determine match-mismatch events in
zooplankton-algae interactions? Oecologia. DOI 10.1007/
more resilient to the impacts of recent climate change. These
s00443-006-0549-2.
results highlight the complex response of Arctic freshwater food
Douglas, M. S. V., Smol, J. P., and Blake, W. Jr., 1994: Marked
webs to climate change. Understanding the responses of organisms post-18th century environmental change in high arctic ecosys-
at multiple trophic levels will be essential for a holistic tems. Science, 266: 416–419.
understanding of the impacts of climate warming on freshwater Duguay, C. R., Prowse, T. D., Bonsal, B. R., Brown, R. D.,
ecosystems. Given that Arctic ecosystems are predicted to Lacroix, M. P., and Menard, P., 2006: Recent trends in
experience an additional 4–6uC warming over the next century, Canadian lake ice cover. Hydrological Processes, 20: 781–801.
substantial reorganizations of freshwater food webs will undoubt- Dzialowski, A. R., and O’Brien, W. J., 2004: Is competition
edly continue to occur in the future, particularly if higher trophic important to arctic zooplankton community structure? Fresh-
levels surpass ecological thresholds. water Biology, 49: 1103–1111.
Frey, D. G., 1988: Littoral and offshore communities of diatoms,
cladocerans and dipterous larvae, and their interpretation in
paleolimnology. Journal of Paleolimnology, 1: 179–191.
Acknowledgments Glew, J. R., 1988: A portable extruding device for close interval
This research was supported by a Natural Sciences and sectioning of unconsolidated core samples. Journal of Paleo-
Engineering Research Council of Canada (NSERC) grant to J. P. limnology, 1: 235–239.
Smol, an NSERC special collaborative grant to the Paleoecolog- Glew, J. R., Smol, J. P., and Last, W. M., 2001: Sediment core
ical Analysis of Circumpolar Treeline (PACT), and the Northern collection and extrusion. In Last, W. M., and Smol, J. P. (eds.),
Studies Training Program (NSTP). Logistical support during Tracking Environmental Change Using Lake Sediments. Vol. 1:
sample collection was given by the Polar Continental Shelf Basin Analysis, Coring, and Chronological Techniques. Dor-
Project. Tammy Karst, Tamsin Laing, Brent Wolfe, and Dave drecht: Kluwer Academic Publishers, 73–105.
Porinchu assisted in field work. Assistance in the field was also Hinzman, L. D., Bettez, N. D., Bolton, W. R., Chapin, F. S.,
provided by the Department of Indian Affairs and Northern Dyurgerov, M. B., Fastie, C. L., Griffith, B., Hollister, R. D.,
Development (DIAND). Hope, A., Huntington, H. P., Jensen, A. M., Jia, G. J.,
Jorgenson, T., Kane, D. L., Klein, D. R., Kofinas, G., Lynch, A.
H., Lloyd, A. H., McGuire, A. D., Nelson, F. E., Oechel, W. C.,
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