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Eawag_04793

J. Limnol., 65(2): 89-99, 2006

Lacustrine particle dynamics in high-altitude Estany Redó (Spain) - a high


resolution sediment trap study
Thomas KULBE*, Christian OHLENDORF1) and Michael STURM

Sedimentology Section, SURF-Eawag, CH-8600 Dübendorf, Switzerland


1)
GEOPOLAR, Institute of Geography, FB 8, University of Bremen, Celsiusstrasse, FVG-M, 28359 Bremen, Germany
*e-mail corresponding author: thomas.kulbe@eawag.ch

ABSTRACT
Particle fluxes were measured from 2000 to 2001 with 3 integrating open traps (O-traps) and a sequencing trap (S-trap) in the
73-m deep, oligotrophic, high-mountain Estany (Lake) Redó (2240 m a.s.l.) over a period of 558 days. O-traps were deployed at 26,
46, and 66 m water depth to measure overall sedimentation rates, while the S-trap was deployed at 66 m water depth to detect
dynamics of seasonal particle fluxes with a resolution of 4 days (during ice break-up, summer, ice formation) to 21 days (during ice
cover). Our results show a high degree of seasonal variability in particle dynamics. Total particle fluxes vary from almost zero to
more than 600 mg m-2 d-1. The highest fluxes occur during short time windows after ice-break-up (minerogenic particles), during
spring (planktonic biomass), and during fall overturn (chrysophycean cysts). Particle fluxes also differed markedly from year to year
in absolute values (2000: 644 mg m-2 d-1, 2001: 370 mg m-2 d-1) as well as in average values (2000: 76 mg m-2 d-1, 2001: 44 mg m-2
d-1). Annual and seasonal meteorological changes and events have a clear influence on the lake system and on the amount and
composition of particles. C/N ratios during April and May increased significantly from 2000 (6-14) to 2001 (>28), reflecting the
more intense soil erosion and transport of terrestrial plant remains into the lake caused by heavy precipitation in 2001. Air
temperature strongly influences the timing of the occurrence of the main bio-productivity peak. Strong wind events shorten the period
of ice cover. Our investigation shows that sediment trap studies lasting more than one limnological cycle are useful in studying the
effects of short-term meteorological changes and weather events on high mountain lakes. However, long-term particle flux
measurements would be necessary to determine amplitudes of natural seasonal cycles and for the interpretation of the decadal-scale
environmental changes occurring in such lakes.

Key words: sediment trap, high-resolution particle flux, high mountain lake, seasonality, bio-productivity

resolution sediment trap studies can reveal this other-


1. INTRODUCTION wise lacking information.
Sediment traps have been used widely in lakes since
Predicting the frequency and amplitude of future the 1950s (Bloesch & Burns 1980). A knowledge of the
climate changes, as well as supplying a precise answer quantity and quality of sedimenting particles and of the
to the question of how ecosystems might respond to timing of their formation is crucial a better understand-
these changes, are difficult problems to solve. The main ing of dynamic sedimentation processes and proxy cli-
reasons for this difficulty are, on the one hand, the mate data (Rathke et al. 1981; Sturm et al. 1982;
inherent complexity and variability of nature, and, on Ohlendorf & Sturm 2001; Wehenmeyer & Bloesch
the other hand, the influence of human activities, which 2001; Müller et al. 2005; Rose & Monteith 2005).
is often strong enough to mask natural variability. However, only automated, high-resolution sequencing
Remote mountain lakes are often influenced by direct traps can detect short-term flux events, caused by
atmospheric deposition and by input from a generally floodings, algal blooms, calcite precipitation etc.
small catchment area, and are not subject to significant (Bloesch & Sturm 1986; Ryves et al. 2003; Sturm et al.
influence from local human activities. In addition, 2003).
global environmental changes tend to leave strongly The geography, limnology and ecology of Estany
amplified signals in high-elevation regions (Beniston et Redó have been investigated since the 1980s. Snow and
al. 1997). Remote mountain lakes are thus thought to be ice development during wintertime, as well as other
excellent sensors of past environmental changes, the physical properties, have been studied to examine the
signatures of which are archived in the lake sediment coupling of physical and biological processes (Catalan
(Sturm et al. 2003). Moreover, many processes can 1988, 1989; Ventura et al. 2000). The lake came into
affect the transfer of climatic and environmental signals, the focus of biological studies because it is oligotrophic
from the atmosphere and the catchment area, through and remote, with little anthropogenic influence (Catalan
the water column and into the sediment. Even regularly & Camarero 1991; Catalan et al. 2002b). These authors
laminated sediments mask the heterogeneity of short- investigated biological processes to detect their potential
term particle fluxes that may occur within a year. High- as proxy data for weather and climate variability. In
90 T. Kulbe et al.

Fig. 1. Estany Redó, showing location (inset), catchment area, and bathymetry. The location of the sediment trap mooring in the
centre of the lake is shown as a black dot. The dashed contour lines near the mooring indicate a small rise in the lake floor of up to 8 m.

addition, Estany Redó was used to study the effect of 35 cm thick (59%) (Camarero et al. 1999). The period
atmospheric pollution and acid rain on remote lake sites of ice cover can last up to 6 months starting in Novem-
(Camarero & Catalan 1993, 1998; Camarero et al. ber/December (Ventura et al. 2000; Catalan et al.
2004). 2002b). Estany Redó is fed mainly by precipitation-
Here we present high-resolution sediment trap data periods (snowmelt and rain), via non-permanent streams
collected from 15 April 2000 until 25 October 2001 in that dry up in summer. A permanent outflow stream
Estany Redó. The study aims to test if differences in the exists on the southern shore of the lake (Fig. 1, Catalan
seasonal weather pattern lead to high seasonal ecosys- 1988). Estany Redó is a dimictic lake and the water has
tem variability within the lake, and to investigate the a residence time of ~4 y. Electrical conductivity is very
influence of the catchment on the particles of the lake. low (~12 µS cm-1). The annual mean air temperature is
In comparison and combination with almost 20-year old 3.6 °C, and the annual mean precipitation is 1328 mm
"trap-like" data (Felip & Catalan 2000), we set out to y-1 (Appleby 2000; Ventura et al. 2000).
show the potential of perennial trap studies, that recover
inter-annual variability, to contribute to climate change 3. METHODS
research. There are the first published results from a 3.1. Mooring design and trap setting
study, which uses automated, high-resolution sediment
traps to determine particle dynamics in a remote high To assess the particle dynamics in Estany Redó, a
mountain lake. sediment trap mooring with three integrating Eawag-
130 open traps (henceforth O-traps: Ohlendorf & Sturm
2. THE SITE 2001) and one TECHNICAP® PPS5/2 sequencing trap
(henceforth S-trap) was deployed in April 2000 (Fig. 2).
Estany Redó is located at an elevation of 2240 m The S-trap, which had 24 sampling cups and an active
a.s.l. in the Spanish central Pyrenees (42°38'N, 0°46’E) area of 5000 cm2, was used to measure short-term, sea-
and was carved into granodiorite bedrock (Ventura et al. sonal fluxes. The rotation times of each of the sampling
2000; Catalan et al. 2002b). Estany Redó is a glacial cups were programmed individually by a microproces-
cirque lake (655 m × 565 m) with steep slopes and a sor, which allowed sampling to be conducted over any
maximum depth of 73 m. The lake covers 16% (0.24 given time interval. The sampling cups consist of PVC
km2) of the 1.5 km2 catchment area (Fig. 1). The rest of and each cup has a volume of 250 ml. The O-traps have
the catchment area consists of bare rock (25%) and an active area of twice 130 cm2 and an aspect ratio of
alpine meadows covering a poorly developed soil about 1:9. Total fluxes measured by O-traps were used to
Particle dynamics in Estany Redó 91

crosscheck sequencing fluxes of the S-trap. An "I-type" orological variables (e.g. air temperature, precipitation,
mooring string was used to deploy the traps in Estany wind speed) were attached to a mast ca 6-10 m above
Redó (Fig. 2, Sturm et al. 1982; Bloesch & Sturm the ground, which is 36-40 m above the lake level. For a
1986). The mooring consisted of a 50 kg anchor weight, more detailed description see Ventura et al. (2000).
a pre-stretched plastic rope 10 mm in diameter, and sev-
eral buoys at the upper end of the rope to provide the 3.3. Sample preparation and measurements
required buoyancy. The S-trap was deployed at 66 m After recovering the mooring, the supernatant water
water depth, 5 m above the sediment-water interface to from the S-trap samples was decanted for chemical
avoid the effects of resuspension. Three O-traps were analysis to check for dissolution within the sampling
attached to the mooring rope at water depths of 26 m cups. The samples were then transferred to the labora-
(below the epilimnion), 46 m and 66 m. In order to tory in cooling boxes and subsequently freeze dried.
avoid damage by ice and loss of buoyancy because of After determining the total dry-weight, the freeze-dried
large changes in lake level, the buoys were deployed trap material from both the O-traps and S-traps was
more than 2 m below the water surface. The mooring analyzed for inorganic carbon (Cinorg) using a coulom-
was deployed from the lake ice in early spring 2000 in eter (Coulometrix Inc. 5011 CO2-Coulometer). Total
order to obtain samples during ice-break up. The longest carbon (Ctot) and nitrogen (Ntot) was measured with a
exposure period included the winter period of Carlo Erba CHNS Elemental Analyzer (EA 1108).
2000/2001 and thus included a second stage of ice cov- Organic carbon (Corg) was calculated as Ctot – Cinorg.
erage. Detailed particle analyses of the trap samples were per-
formed with a Phillips XL30 scanning electron micro-
scope (SEM) with an attached energy dispersive spec-
troscopy system (EDS). 12 mg of a sample were placed
in a test-tube with 4 ml of ethanol. This suspension was
then sonicated for 2 minutes and sprayed onto the tar-
gets using a spray gun (Bollmann et al. 1999). In the
SEM, images were made at magnifications of 50×,
250×, and 1000×. Three analysis fields per image were
chosen to measure their concentrations of Na, Mg, Al,
Si, Cl, S, P, K, Ca, Fe, and Mn by SEM-EDS. To
account for sample inhomogenities and to assess ana-
lytical precision, the average values of these measure-
ments were used (Schloz 2000). Concentrations are
given in atom-% as determined by SEM-EDS with
respect to reference material.

4. RESULTS
4.1. Trap results and particle dynamics
4.1.1. O-traps
Particle fluxes were measured over a period of 558
days, from 15 April 2000 to 25 October 2001. This
period was split up into 4 intervals, i.e. 3 exchanges of
the traps:
• 1) 15.04.2000 – 13.08.2000: 120 days
• 2) 13.08.2000 – 19.11.2000: 98 days
• 3) 19.11.2000 – 15.06.2001: 208 days
• 4) 15.06.2001 – 25.10.2001: 132 days
Fig. 2. Detailed sketch map of the trap mooring in Estany Particle fluxes were determined separately for each
Redó in its winter setting showing the buoys 2 m under the
water surface. In summer the uppermost buoys are at the water of these periods and are shown in figure 3. Except for
surface. the lowest O-trap during period 4, total sediment fluxes
increased with water depth (Fig. 3). Total particle fluxes
were lowest during ice cover (58-82 mg m-2 d-1), and
3.2. Meteorological data
were 2 – 4 times higher than this under open water con-
Meteorological data were recovered from an auto- ditions. The highest total sediment fluxes - up to 278 mg
matic weather station, which was installed ca. 30 m next m-2 d-1 - were recorded in the lowermost O-traps during
to the southern shore (Fig. 1). The sensors for the mete- ice break up. The average particle flux within the expo-
92 T. Kulbe et al.

Fig. 3. Particle fluxes of O-traps in 26 m, 46 m, and 66 m


water depth during the four exposure periods.

Fig. 4. Seasonal particle fluxes in Estany Redó from 14 April 2000 to 25 October 2001 revealed by the S-trap in 66 m water depth.
X: mean sediment flux determined by O-trap at 66 m water depth during the four exposure periods.

sure period was half as high in the 26 m O-trap (89 mg 4). The fluxes varied from almost zero during the peri-
m-2 d-1) than in the lowest O-trap, at 71 m water depth ods of ice cover to more than 600 mg m-2 d-1 during the
(160 mg m-2 d-1). open water period. Five periods of high sediment accu-
mulation were observed (Fig. 4).
4.1.2. S-trap sediment fluxes and particle composition Ice breakup normally begins in May/June (Catalan
The sediment fluxes measured by the S-trap were 1988, 1992; Ventura et al. 2000) on the eastern shore,
highly variable during the period of investigation (Fig. where the main inflow enters the lake. In the very first
Particle dynamics in Estany Redó 93

Fig. 5. SEM images. (A) Initial phase of ice break-up, June 2000: detrital minerals and pollen. (B) Beginning of open-water period,
July 2000: large organic aggregates. (C) End of the high flux period, August 2000: more zooplankton remains less amounts of
organic aggregates. (D) October 2000: chrysophycean cysts, zooplankton remains, and organic aggregates. The scale bars are each
100 µm in length.

stage of break-up in 2000, an initial input of allochtho- During summer, a second period of high particle
nous particles was observed, from 18.05. – 06.06. The accumulation occurred. In 2000 it lasted 36 days from
particles consisted mostly of detrital minerals, a few 14.10. – 18.11. and the particles were dominated by
pollen grains, and zooplankton remains (Fig. 5a). zooplankton remains. In contrast to 2001, there were
Detrital minerals and pollen grains were probably many chrysophycean cysts and few organic aggregates
released by melting lake ice and transported from the (Fig. 5d). In 2001, organic aggregates and zooplankton
catchment by snowmelt runoff. SEM images of particles remains dominated the particle composition during the
that entered the S-trap during the thawing of the ice high flux period from 28.07. – 10.09. (45 days). Minor
show that biological productivity was still limited in the components included chrysophycean cysts, as well as
lake at that time. This kind of particle input in the very very few pollen grains and diatoms. The final stage of
first stage of ice break-up did not appear in 2001. the open-water period (very low particle fluxes, no
The highest particle fluxes occurred when the lake peaks) was characterized by a few organic aggregates
became completely ice-free; commencement of the and zooplankton remains, and a few chrysophycean
open-water period was characterized by large autoch- cysts, pollen grains, and diatoms (Fig. 5d).
thonous organic aggregates (Fig. 5b). Minor compo- From May to October 2000, the overall sediment
nents included a few diatoms and pollen grains. This accumulation in the S-trap (7500 mg) was approxi-
high productivity peaks occured in 2000 from 09.07. – mately 2.5 times higher than the amount that accumu-
07.09. (61 days) and in 2001 from 03.04. – 29.05. (57 lated during the same time period in 2001 (3200 mg),
days). Towards the end of the high flux period, the large and the mean sediment flux (91 mg m-2 d-1) was more
organic aggregates became less abundant (Fig. 5c). than twice as high as in 2001 (44 mg m-2 d-1). The
94 T. Kulbe et al.

Tab. 1. Selected information on fluxes based on the S-trap material. Note that the sampling intervals in 2000
(15 April - 31 December) and 2001 (1 January - 25 October) differ. *The value in parentheses was calculated
after excluding the exceptionally high values measured from 2 April - 6 June 2001.
Total flux Corg flux N flux C/N ratio
[mg m-2 d-1] [mg m-2 d-1] [mg m-2 d-1]
2000 2001 2000 2001 2000 2001 2000 2001
Min. 0.1 0.2 0.0 0 0.1 0.0 6.1 5.5
Max. 644 370.4 79.4 51.4 7.4 2.3 13.7 60.8
Mean 75.6 43.7 13.5 7.6 1.9 0.4 8.7 27.3 (*12.4)

Tab. 2. Mean, minimum, and maximum values of chemical composition of the 96 S-trap
sediment samples calculated for the period from May 2000 to November 2001. Values are given
in atom % as determined by SEM-EDS.
Si Al S Na Fe Ca P K Mg Mn Cl
Minimum 14.8 3.4 2.4 0.0 2.4 0.7 1.0 0.8 0.7 0.0 0.0
Maximum 72.4 22.1 19.7 16.1 9.1 22.6 7.5 5.1 5.3 5.4 13.2
Mean 52.8 11.4 7.2 6.8 5.1 5.0 3.0 2.7 2.5 2.0 1.6

maximum particle fluxes of the S-traps were approxi- (2000) and 117 mg m-2 d-1 (2001). These fluxes are
mately twice as high (644 mg m-2 d-1) in 2000 (meas- within the range found in other small high mountain
ured from 8. – 12. July) as in 2001, with only 370 mg m- lakes. Lower mean annual particle fluxes of 48 mg m-2
2 -1
d in 2001 (measured from 29 May to 2 June; Fig. 4, d-1 have been measured in Gosseköllesee/Austria
Tab. 1). (EMERGE final report). Gosseköllesee is small (150 ×
As summarized in table 1 and figure 4, the highest 80 m), shallow (10 m), has a small catchment area (0.59
fluxes of Corg (79 mg m-2 d-1) and Ntot (7.4 mg m-2 d-1) km2), and is on a higher elevation (2417 m a.s.l.).
occurred from 8 – 20 July in 2000, coinciding with the Higher particle fluxes 703 mg m-2 d-1 (average over 2.5
period of maximum total particle flux. In 2001, the y) have been observed in Hagelseewli/Switzerland (cor-
maximum flux (51 mg m-2 d-1) occured simultaneously rected after Ohlendorf & Sturm 2001). This lake is 250
with the maximum flux of 29 May – 2 June 2001, × 150 m in size, 19 m deep, has a very small catchment
within the first productivity period. However, the high- of 0.3 km2 and is located at an elevation of 2339 m a.s.l.
est input of nitrogen occurred within the second period Larger high mountain lakes like Sil-
of high productivity in August (2.3 mg m-2 d-1). Mini- vaplanasee/Switzerland located at 1791 m a.s.l. show
mum values of the C/N ratio of the trap material were much higher mean particle fluxes of 3900 mg m-2 d-1
~6 in both 2000 and 2001. For the period April to May (Troxler 2005; Bluszcz et al., submitted). Silvaplanasee
2000 the C/N ratios found (6 -14) were substantially is 77 m deep, but in contrast to Estany Redó,
lower than those found in 2001 (32 - 61). Excluding this Gosseköllesee, and Hagelseewli it differs very much in
time period from the statistics, the mean C/N ratio was 9 size (3100 m × 1400 m) and catchment area (129 km2)
in 2000 and 12 in 2001, indicating autochthonous and represents a proglacial lake.
source material (Wetzel 1983; Leenheer 1988). Wehenmeyer & Bloesch (2001) investigated 11
Swedish and 9 Swiss lakes of different sizes, water
4.1.3. Chemical composition of particles depth and topographical settings, but all on a much
The semi-quantitative chemical composition meas- lower elevation. They measured particle fluxes of 400 –
ured by SEM-EDS of the trap particles is shown in table 7900 mg m-2 d-1 in generally shallow Swedish lakes
2. The mean values are typical for bedrock dominated (<21 m water depth), some of whichare influenced by
by crystalline rocks in the catchment of the lake. Si and re-suspension, sediment sliding and eutrophication. In
Al dominated the element composition. The samples Swiss lakes they reported particle fluxes of 1500 – 4800
also contained some S, Na, Fe, and Ca whereas P, K, mg m-2 d-1; however, these lakes are much larger, have
Mg, Mn, and Cl showed the lowest values. prominent tributaries and are often eutrophied.
The comparatively low particle fluxes in Estany
5. DISCUSSION Redó as well as in other high mountain lakes are caused
by their high elevation, relatively small catchment areas
5.1. Comparison of the 2000/2001 sediment trap results together with a lack of permanent inflows and low
with other lakes nutrient content. Additionally, the long duration of ice
The mean particle flux measured in the lower O-trap cover also causes a reduction of particle fluxes.
from 2000 to 2001 was 160 mg m-2 d-1. Mean particle In Estany Redó, Ventura et al. (2000) describe
fluxes from May to September in the lowest sediment maxima of Daphnia pulicaria during the period of ice
O-trap in Estany Redó differed between 251 mg m-2 d-1 cover, and maxima of Daphnia cyaneus during the
Particle dynamics in Estany Redó 95

Fig. 6. Variations in total sediment flux and air temperature (7-days running mean) in 2000 and 2001. The shaded bars display
periods with high autochthonous particle fluxes. The peak of June 2000 displays allochthonous material.

open-water period. This is consistent with the fact that remobilization of phosphorus. According to Catalan &
analyses of the trapped particles in 2000 and 2001 Camarero (1991) phosphorus can be remobilized when
revealed zooplankton remains and fecal pellets (Fig. the lower boundary of the mixed epilimnion water layer
5b). Overall, particles of autochthonous organic origin reaches phosphorus-rich fine-grained sediment layers at
dominate the particle sedimentation during that time approximately 25 m water depth. However, the differ-
period as well as during the end of the high flux period ence in the amount of biomass produced in 2000 as
(Figs 5b, c). Total sediment fluxes and the flux of compared to 2001 did not necessarily reflect a differ-
organic carbon are very well correlated (r2 94.2%, p ence in the phosphorus pool or in other nutrients, but
<0.001). However, the preservation of this kind of parti- was more likely the result of natural variability. Sea-
cle is limited because they decompose very easily and sonal particle flux variations within a single year in
thus will not be preserved in the sedimentary archive. Estany Redó exceeded those between years, a finding
The trapped material during fall overturn agrees with that corresponds with the work from Wehenmeyer &
the results of Felip & Catalan (2000), who reported Bloesch (2001).
blooms of flagellate chrysophytes in the hypolimnion.
After fall overturn fecal pellets, chrysophycean cysts, 5.2. 2000/2001 sediment trap results and concurrent
zooplankton, and diatoms characterized the sedimenta- meteorological data
tion during the final stage of the open-water period (Fig. In ice-covered lakes, the main relevant factor is the
5d). timing of ice break-up, which, is both closely linked to
The biological cycle in Estany Redó is controlled by ambient air temperature (e.g., Livingstone 1997) and a
the availability of phosphorus (Catalan & Camarero major determinant of the timing of the spring phyto-
1991; Ventura et al. 2000; Catalan et al. 2002b), i.e. plankton bloom (e.g., Wehenmeyer et al. 1999). In
once the phosphorus pool is consumed, the food chain Estany Redó the ice cover started to break up in late
collapses. The sediment trap data implies that this hap- May in 2000, and early May in 2001. Taken into
pens after approximately 2 months of high biological account the uncertainties of an exact dating of the break
productivity. As shown in figure 6, the duration of the up due to the remoteness of the lake and its harsh
period of high sediment fluxes during the open-water weather conditions in winter, the data agree with the
period was ca 2 months and was essentially the same in meteorological information. A comparison of the mete-
2000 and 2001. orological data from 2000 and 2001 shows that average
The occurrence of a second bloom in fall (Fig. 6), meteorological conditions in 2000 and 2001 were com-
shorter in duration (36 and 45 days in 2000 and 2001, parable (Tab. 3), although differences did occur in the
respectively) and with lower productivity than the first timing, frequency, and amplitude of weather events
bloom in both years, is presumably the result of the (Figs 6 and 7a-c).
96 T. Kulbe et al.

Tab. 3. Comparison of meteorological data measured at Estany Redó from 2 April


to 1 November in 2000 and 2001. The data were measured at an automatic
weather station located 30 m from the south shore of the lake. The time period
chosen represents the time during which data is available for both years.
2000 2001
Temperature [°C]
Minimum -6.5 -9.3
Maximum 17.0 19.1
Mean 6.6 7.0
Incident solar radiation [W m-2]
Minimum 16.5 6.3
Maximum 390.3 390.8
Mean 219.4 231.8
Wind speed [m s-1]
Minimum 1.0 0.5
Maximum 17.0 17.6
Mean 5.7 5.4
Precipitation [mm d-1]
Minimum 0.0 0.0
Maximum 76.8 56.8
Mean 4.4 4.3
Sum 930.7 911.4

Fig. 7. Meteorological parameters (air temperature, wind speed, precipitation) and C/N-ratio (atomic) in 2000 and 2001. The
meteorological data and C/N ratios are averaged over the trap exposure intervals. Total sediment flux rates (TF) are indicated in gray
for comparison purposes.
Particle dynamics in Estany Redó 97

The ice-cover period was much shorter in 2001 than Catalan et al. (2002a) reported a warming trend in
in 2000, probably because of at least two periods of the central Pyrenees from the early 1970s onwards,
high wind, which accelerated the initial ice break-up especially in summer and autumn (June to November).
(Fig. 7b). The open water period began ca 50 days ear- Felip & Catalan (2000) investigated the phytoplankton
lier in 2001 (end of May) in comparison to 2000 (begin- community from May 1984 to August 1985, taking
ning of July). The onset of the first and second phase of samples at intervals of 6-30 days from 9 different water
bio-productivity also shifted towards an earlier date in depths. Their 20-yr-old data has been compared with
2001 compared to 2000, by 98 and 59 days, respec- our results from 2000 and 2001 to test whether the pos-
tively. This suggests that the onset of the main period of tulated warming trend has influenced the biological
biological productivity may be linked to internal lake productivity of Estany Redó, even if the interpretation
factors that are governed primarily by climate. of both investigation periods is hampered by inadequate
A comparison of the organic material with the pre- long-term data series. During both time periods, the first
cipitation data reveals a different picture (Fig. 7d). C/N productivity peak occurred under the ice in spring (Fig.
ratios from April to May differ substantially between 4). This phenomenon has recently been observed also in
2000 (6-14) and 2001 (>28). The high values in the lat- other ice-covered lakes in the Alps, Scandinavia and
ter case reflect the enhanced transport of particles from Siberia (Marchetto et al. 2004; Sturm et al. 2005). In
soil erosion and enhanced transport of the remains of 1984, small first productivity peaks occurred during the
higher plants to the lake (Wetzel 1983). Camarero et al. open-water phase in July/August, followed by maxi-
(1999) also suggest an external source of particulate mum productivity peaks in November/December. This
organic matter for Estany Redó, because mobilization of succession within the year 1984 is also apparent in 2000
allochthonous organic material is possible during ice and 2001. However, at present, the first phase of bio-
break-up, as it is incorporated into soil material from the productivity is greater than the second (Fig. 4). We
shore or deposited on the snow and then transported into attribute these differences to natural variation within the
the lake by streams of meltwater. Another reasonable lake ecosystem. There is no evidence to support the
explanation for higher C/N values is heavy rainfall and hypothesis that the productivity of Estany Redó has
initial meltwater in early spring. In April 2001 (Fig. 7c, reacted to climate warming in the central Pyrenees.
d), C/N ratios of trapped material increased parallel with
higher precipitation, implying that surface runoff and 6. CONCLUSIONS
erosion within the catchment area was transporting Sedimentation in Estany Redó is dominated by
organic material from soils into the lake. autochthonous particles based on the strong correlation
of total sediment fluxes and the flux of organic carbon
5.3. Comparison of the years 2000-2001 with the years of 94.2 (r2, in percent, p <0.001). The elemental
1984-85 composition of the trapped minerogenic particle reflects
According to Felip & Catalan (2000), dinoflagellates the crystalline bedrock in the catchment.
are the main phytoplankton species in Estany Redó. In The small catchment area of the lake and its lack of
1984 and 1985, they analyzed water samples from dif- permanent inflows as well as a very low nutrient supply
ferent water depths in the lake and observed the first result in particle fluxes that are similar to other small
algal bloom to occur in July in both years, although dif- high mountain lakes. From 2000-2001 the mean particle
ferences in the amount of productivity from one year to flux of high alpine Estany Redó was 160 mg m-2 d-1.
another do occur. Felip et al. (1999) report highest pri- Mean fluxes from May to September differ between 251
mary-productivity during the same time in 1984 and mg m-2 d-1 (2000) and 117 mg m-2 d-1 (2001). This is an
1985 but this peak was dominated by chrysophycean order of magnitude smaller than in lakes which are on a
cysts and dinoflagellates. lower altitude, bigger in size, with a bigger catchment,
By contrast, SEM images of sediment trap samples and have permanent inflows.
during this time interval (the beginning of the open The particle fluxes of Estany Redó show a seasonal
water period) in 2001 do not clearly reflect this bloom. pattern consisting of a long period with low sedimenta-
Chrysophycean cysts are very rare and the trapped tion, when the lake is ice-covered, and short periods
material consists mainly of organic aggregates about with high flux rates during ice break-up and during the
100 µm in size, which have a similar structure to fecal open-water period. In 2000 a distinct particle flux of
pellets, although they differ in shape (Fig. 5b). detrital minerals occurred directly after the initial ice-
As seen in figure 4, the open-water productivity break-up. The highest particle fluxes consist of autoch-
peaks differed in terms of their shape and height at least thonous material and, although the inter-annual pattern
as much from 2000 to 2001 as they had done from 1984 is highly comparable, the seasonal variability is high. In
to 1985. For this reason, excluding the timing of the 2000 and 2001 two phases of enhanced bio-productivity
peaks, both studies document the high inter-annual envi- were recorded. The first peak in both years marked the
ronmental variability of the lake, but do not provide a highest sedimentation period and lasted ca 2 months.
clear conclusion on the issue of climatic forcing. The trapped material consisted mostly of zooplankton
98 T. Kulbe et al.

remains and organic aggregates (fecal pellets). The sec- Bluszcz, P., C. Ohlendorf, B. Zolitschka & M. Sturm. (2006).
ond period was comprised of the same particles but was High-resolution sediment trap studies in Silvaplanasee,
Switzerland. Aquatic Science: (submitted).
shorter in duration. One difference between 2000 and Bollmann, J., B. Brabec, M.Y. Cortés & M. Geisen. 1999.
2001 was that much higher amounts of chrysophycean Determination of absolute coccolith abundances in deep-
cysts were deposited in 2000. Both periods of high bio- sea sediments by spiking with microbeads and spraying
productivity, within the year, together covered ca 100 (SMS-method). Mar. Micropaleontol., 38(1): 29-38.
Camarero, L. & J. Catalan. 1993. Chemistry of bulk precipita-
days. tion in the central and eastern Pyrenees, northeast Spain.
Estany Redó is one of the very few lakes in which Atmos. Environ., 27A: 83-94.
detailed particle flux data from recent years can be Camarero, L. & J. Catalan. 1998. A simple model of regional
compared with older water sample data that was col- acidification for high mountain lakes: application to the
Pyrenean lakes (North-East Spain). Water Res., 32: 1126.
lected almost 20 years ago. According to this compari- Camarero, L., M. Felip, M. Ventura, F. Barthumeus & J.
son, there is no clear sign of a warming trend in the Catalan. 1999. The relative importance of the planktonic
Pyrenees affecting Estany Redó over the past two dec- food web in the carbon cycle of an oligotrophic mountain
ades. Our data do, however, show a link to meteorologi- lake in a poorly vegetated catchment (Redó, Pyrenees). J.
cal data: the onset of the main period of biological pro- Limnol. 58: 203-212.
Camarero, L., R.F. Wright, J. Catalan & M. Ventura. 2004.
ductivity in the lake appears to be associated with air Application of MAGIC to Lake Redó (Central Pyrenees):
temperature, and the duration of ice cover and partial ice an assessment of the effects of possible climate driven
cover is affected by air temperature and perhaps high changes in atmospheric precipitation, base cation deposi-
wind speeds. Soil erosion during periods of high pre- tion, and weathering rates on lake water chemistry. J.
Limnol., 63: 123-132.
cipitation is documented by changing C/N ratios in Catalan, J. 1988. Physical properties of the environment rele-
2001. Although sediment trap data can thus be used for vant to the pelagic ecosystem of a deep high-mountain
studies of environmental change and weather events in lake (Estany Redó, Central Pyrenees). Oecologia
high mountain lakes on a very short time scale, our aquatica, 9: 89-123.
Catalan, J. 1989. The Winter Cover of a High-Mountain
results strongly suggest that long-term investigations of Mediterranean lake (Estany Redó, Pyrenees). Can. J.
such lakes are necessary to clarify the existence, rate, Earth Sci., 25: 519-527.
duration, and amplitude of environmental changes. Catalan, J. 1992. Evolution of Dissolved and Particulate Mat-
ter during the Ice-Covered period in a Deep, High-Moun-
ACKNOWLEDGEMENTS tain Lake. Can. J. Fish. Aquat. Sci., 49: 945-955.
Catalan, J. & L. Camarero. 1991. Ergoclines and biological
We would like to express our thanks to Ingrid processes in high mountain lakes: Similarities between
Holderegger, Yvonne Lehnhard, Brian Sinnet and Alois summer stratification and the ice-forming periods in Lake
Zwyssig (all at EAWAG), and to Lluis Camarero, Jordi Redó (Pyrenees). Verh. int. Ver. Limnol., 24: 1011-1015.
Catalan, J., S. Pla, M. Rieradevall, M. Felip, T. Buchaca, L.
Catalan and Marc Ventura (all at Dept. of Ecology, Camarero, S. Brancelj, P.G. Appleby, A. Lami, J.A. Gryt-
Univ. of Barcelona, Spain) for their help during the field nes, A. Agustí-Panareda & R. Thompson. 2002a. Lake
work, laboratory work, sediment sampling and helpful Redó ecosystem response to an increasing warming in the
discussions. The manuscript benefited substantially Pyrenees during the twentieth century. J. Paleolimnol., 58:
from comments from David Livingstone, who also 129-145.
Catalan, J., M. Ventura, A. Brancelj, I. Granados, H. Thies, U.
improved the English of the manuscript. We further Nickus, K. A., A.F. Lotter, A. Barbieri, E. Stuchlík, L.
thank two referees for their valuable comments. This Lien, P. Bitusik, T. Buchaca, L. Camarero, G.H.
study was funded by the Swiss Federal Office of Edu- Goudsmit, J. Kopácek, G. Lemcke, D.M. Livingstone, B.
cation and Science within the framework of the Euro- Müller, M. Rautio, M. Sisko, S. Sorvari, F. Sporka, O.
Strunecky & M. Toro. 2002b. Seasonal ecosystem vari-
pean Union Environment and Climate project ability in remote mountain lakes: implications for detect-
"EMERGE" (European Mountain lake Ecosystems: ing climatic signals in sediment records. J. Paleolimnol.,
Regionalisation, diaGnostics and socio-economic 58: 25-46.
Evaluation), grant BBW no.99.0394. Felip, M., F. Barthumeus, S. Halac & J. Catalan. 1999. Micro-
bial plankton assemblages, composition and biomass,
during two ice-free periods in a deep high mountain lake
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Received: May 2006


Accepted: September 2006

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