© 2009 WILEY-VCH Verlag GmbH & Co.

KGaA, Weinheim 1434-2944/09/102-0103
Internat. Rev. Hydrobiol. 94 2009 1 103–125
DOI: 10.1002/iroh.200811071
Berlin University of Technology, Dept. of Water Quality Control, Straße des 17 Juni 135, Sekr. KF 4,
D-10623 Berlin, Germany; e-mail:guenter.gunkel@tu-berlin.de
Limnology of the Crater Lake Cuicocha, Ecuador,
a Cold Water Tropical Lake
key words: Andes, atelomixis, caldera lake, Ecuador, Lake Cuicocha, travertine
Cuicocha (3380 m a.s.l.) is a young, a few hundred years old volcanic lake in the western cordilleras
of the Ecuadorian Andes with some post-volcanic activities, such as emission of volcanic gases and
input of hydrothermal water. Water chemistry is influenced by the emission of CO
and weathering of
the young andesitic rocks in the water shed. A calcium cycle exists in the lake with intensive biological
Ca precipitation at the flanks and formation of travertine crusts, while in the hypolimnion dissolution
of Ca carbonate occurs. The crater lake is oligotrophic, biodiversity is low; the littoral flora and fauna
is more important than the pelagic species. In the littoral zone, a small Totora zone occurs, followed
by submerged macrophytes down to 35 m water depth. Phyto- and zooplankton occur down into the
hypolimnion. Phytoplankton is strongly influenced by down-welling of water (atelomixis) and by copre-
cipitation with detritial flocs.
1. Introduction
Volcanic lakes, built up in a caldera, are strongly influenced by volcanic activities such
as gas emissions and hydrothermal water springs. Thus, different lake types are formed,
and have been classified according to their physical water constraints by PASTERNACK and
VAREKAMP (1997). These authors distinguished volcanic lakes with different levels of activ-
ity, namely cool to hot acid-brine lakes, reduced to oxidized, acid-saline lakes, acid-sulphate
lakes and bursting to buoyant plume bicarbonate lakes; only neutral dilute volcanic lakes do
not show any activity. Besides this heterogeneity in water chemistry, a wide range of mor-
phometric characters occur as well as different lake genesis which influence the limnology
of the lakes (LARSON 1989; VAREKAMP et al., 2000; ARMIENTA et al., 2000; AGUILERA et al.,
2006). Some basic knowledge exists regarding the limnology of crater lakes (VÁZQUEZ et al.,
2004; SCHABETSBERGER et al., 2004) especially from the long research program on the Crater
Lake in Oregon, which was mainly concerned with water chemistry, stratification processes
and water currents (LARSON, 1996; LARSON et al., 1996; NELSON et al., 1996; CRAWFORD
and COLLIER, 1997).
Limnology of crater lakes is of high interest, because they are young lakes, situated in an
area with soils in a state of development. Many of the calderas are deep and have formed
lakes with a depth of a few hundred meters, mostly situated in high mountain regions. Some
* Corresponding author
© 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.revhydro.com
of these represent a scarce type of cold tropical lake (LÖFFLER, 1964; CASALLAS and GUNKEL,
2001; ROLDAN and RUIZ, 2001; SCHABETSBERGER et al., 2004), and a few investigations on
this type of lake have been carried out mainly concerning water mixing processes (TALLING,
1969; GUNKEL and CASALLAS, 2002, 2002a), primary production (KINZIE et al., 1998) and
occurrence of zooplankton (GREEN, 1995). The morphometry of caldera lakes is frequently
determined by the steep slopes of the flanks and non-eroded lake shores. Water chemistry is
strongly influenced by the source of water input (rainwater or hydrothermal water) and by
the weathering of volcanic deposits in the watershed. In some cases, the continuous input
of volcanic gases and/or hydrothermal water leads to ionic rich water (OHBA et al., 1994;
NELSON et al. 1996; RONDE et al., 2002), and a chemocline can build up (WOOD et al., 1984).
Inflow of hydrothermal water and the energy flux from the volcano strongly influences the
stratification processes (CAMERON and LARSON, 1993; CHRISTENSON, 1994; PASTERNACK and
VAREKAMP, 1997). This can lead to an irregular overturn due to inverse thermal stratifica-
tion, and convergent currents are observed (GOECKE, 1997; CRAWFORD and COLLIER, 1997).
Fauna and flora are determined by the young age of the lakes, the high mountain position,
and the mixing processes, and it must be assumed that the complexity of the biocoenosis
is reduced.
Crater lakes are poorly investigated worldwide. Since the CO
eruption of Lake Nyos,
Cameroon, followed by intense international research to analyse the phenomenon of limnic
eruptions (LE GUERN and SIGVALDASON, 1989, 1990; KUSAKABE, 1994; MARTINI et al., 1994),
there have been concentrated efforts to obtain detailed information on other volcanic lakes
with potentially dangerous CO
accumulation. However, there is still a deficit of knowledge
on volcanic lakes, especially in the Andes of South America, even though this is a region with
a great number of active volcanoes. In Ecuador, two lakes are known to be active volcanic
lakes, the Quilotoa (AGUILERA et al., 2000; GUNKEL et al., 2008) and the Cuicocha (von
HILLEBRANDT and HALL, 1988; GUNKEL et al., pers. com.), located in the high Andine region
> 3000 m above sea level (a.s.l.). Both the Quilotoa and Cuicocha volcanoes, have formed
large and deep caldera lakes and currently show post-volcanic activities in form of volcanic
gas emissions and input of geothermal water. The volcanic gases, mainly CO
, are accumu-
lated in the lake water but without the risk of a limnic eruption (GUNKEL et al., pers. com.).
Investigations on the Cotacachi-Cuicocha complex carried out by SAUER (1971), MOTHES
and HALL (1991) and GRUPE et al. (2008) provided a clear picture of the history of the
volcano and the petrogenesis of the erupted lavas, and VON HILLEBRANDT and HALL (1988)
developed a volcanic hazard map of the surrounding areas.
Lake Cuicocha was the focus of a detailed limnological study with the objective of a
comprehensive characterization of the volcanic lake and its watershed in respect to petrol-
ogy, morphometry, hydrophysics, water chemistry and limnology. Several scientific field
campaigns were carried out between 2003 and 2006; primary results have been published
regarding macrophytes (KIERSCH et al., 2004), Al polymerisation (GUNKEL et al., pers. com.)
and the risk assessment of a limnic eruption (GUNKEL et al., pers. com.).
2. Methods
2.1. Studied Site
Cuicocha is a parasitic volcano of the Cotacachi volcano, which was active in the Pleistocene period,
and is located in the western cordilleras of the Ecuadorian Andes, situated about 100 km north of Quito.
Lake Cuicocha was formed after the collapse of the Cuicocha domo and is a young caldera lake (Fig. 1).
Today, the caldera has a diameter of 3.2 km and a maximum depth of 450 m, filled by a lake to a water
depth of 148 m. Recent volcanic activities of the Cuicocha volcano include emission of volcanic gases
and some hydrothermal water inflow at the lake bottom as well as at the shore line.
Limnology of a Crater Lake 105
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The water input into the lake is of low quantity, due to the small catchment area, and to the moderate
annual precipitation of 1320 mm (2004–2006). The caldera has no continuous inflow and water inflow
occurs via a periodic waterfall, two cascades with low flow rates and the mentioned hydrothermal
entries; the lake has no superficial outflow. The soils consist of volcanic deposits, mainly andesite with
a high SiO
and Al
content (57–61 % SiO
, 17–18% Al
; GRUPE et al., 2008) and its weathering
products. The soils are in an early stage of development with a low clay content classified as andisols
covered by paramo vegetation (ZEHETNER, 2003).
2.2. Sampling
Lake Cuicocha was investigated using a regular monitoring program and sampling was carried out
twice a year for four years (2003–2006). The twice yearly investigation periods were February to April
(rainy period) and July to October (dry period), from 08/2003 to 04/2006. The data base for the water
chemistry included 6 vertical profiles (in 08/2003, 09/2003, 03/2003, 08/2004, 03/2005, 08/2005), and
lake profiler diagrams (T, pH, cond., E7) were based on 10 profiles (in 08/2003, 09/2003, 03/2004,
08/2004, 03/2005, 08/2005, 04/2006 with 2275 data points). Investigations on further parameters (water
physics and stratification stability at the crater rim and near the islands) were based on 29 lake profiles,
determined during the regular investigation periods.
For depth determination and detailed bathymetric mapping of the lake, a double frequency sonar
(50 and 200 kHz; Garmin Fishfinder 250 C) in combination with GPS (Garmin 60CS) was used; the
bathymetric map was developed by the sonar GPS data and digitalised topographic maps (1 : 15 000,
Souris, IRD). The use of sonar allowed detection of the lake floor as well as recognition of volcanic gas
emissions and resuspended sediments as a consequence of gas eruptions. Furthermore, the use of sonar
facilitated the positioning of the equipment in a specific water depth and thus allowed high precision
for data registration near the sediment.
Temperature, pH, conductivity, CO
and redox potential were determined by a lake profiler (Ocean
Seven 316, Idronaut, Italy) with extremely high accuracy of the probes (with values for temperature
+ 0.003 °C, for pH ± 0.01 pH units and for redox potential ± 1 mV). The CO
, probe was calibrated
immediately after utilization by chemical determinations of CO

and CO
according to the
German Standard Methods for pK
and pK
analysis (DEV 2005). The density of the water was cal-
culated under consideration of salinity, local pressure and temperature using the formula of CHEN and
MILLERO (1986) for natural waters and is expressed as ρ
(S, T, Psurface)
(= density 1000 kg/m
); salinity was
calculated by the total ionic content. Oxygen concentration within the carbonate precipitates was deter-
mined using optical oxygen sensor of ∅ 2 mm with a Fibox 3 oxygen meter (PreSens, Germany). Water
samples were taken using a Ruttner water sampler (Hydrobios, Germany) at 78 m (9 samples in 1–78 m;
N 0°18´11.4˝ W 78°22´10.9˝) and 148 m depth (13 sample in 1–148 m; N 0°17´42.3˝ W 78°21˝26.7˝;
Fig. 2), portions of the water samples were filtered immediately, using 0.45 μm cellulose-acetate filters
for cation and anion analyses. Water samples for chemical analyses were preserved by HNO
as well
as HCl (both at pH ~ 1), using HDPE bottles.
Gas sampling was done directly with GC vials at the lake shore and on Island Yerovi.
Samples of suspended material for Scanning Electron Microscopy was obtained by filtration of
250 mL lake water from the water sampler, immediately after sampling in the boat, using 0.4 μm poly-
carbonate filters (Nuclepore). The wet polycarbonate filters were stored in plastic bags with taps soaked
with formaldehyde (37%) for preservation.
Sediment samples were collected by a sediment gravity corer, 5 cm ∅ with a sediment capturer. Sedi-
ment sampling was recognized to be very difficult due to only a thin sediment layer on the stony floor
and due to sediment oversaturation by gases, which led to the loss of sediments while degassing during
lifting of the equipment. Sediment samples were prepared using an HCl/HNO
acid digestion method
(VDLUFA 1991) and chemically analysed using the analytical methods mentioned below. Sedimentation
behaviour of seston was studied using 8 sediment traps, 8 cm ∅, 50 cm length, which were exposed
in the western bay of the lake for two periods, each one for 2 weeks at depths of 15, 30, 50 and 70 m
(N 0°18΄15.3˝ W 78°22´14.9˝). Samples were preserved with 4% glutaraldehyde solution for Scanning
Electron Microscopy with Energy Dispersive Spectroscopy (SEM-EDS) analyses.
A SONY HCR-HC16E digital video camera was modified as an underwater camera and was pro-
tected by a purpose-built aluminium housing and equipped with 4 underwater lamps (NEMO 8C Xenon,
14 watt). With this camera, sediment type and gas emissions with sediment resuspension were regis-
© 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.revhydro.com
tered as well as the penetration of the sediment core sampler into the sediment. During two campaigns,
divers were used for sampling and underwater filming, and the experiences of the divers were helpful
for further investigations.
Phytoplankton samples were collected from the water sampler (11 depths in 0–145 m), preserved with
Lugol’s solution. Zooplankton was collected using a plankton net (55 μm) with trapping mechanism (a
profile was made by 8 sections from 0–145 m), and preserved with HANEY and HALL solution. Plants
were collected using a macrophyte anchor (KIERSCH et al., 2004) in 1998 within 6 profiles from the lake
surface down to 40 m. Sampling of the invertebrates was done by a fine sieve and a plankton net at
6 different positions in the littoral zone, surficial and down to a few meter water depth, preserved with
4% formaldehyde. Fish control was done by using a sonar (Garmin Fishfinder 250 C).
2.3. Chemical Analyses
Nitrite was determined using the Merck test (Aquaquant, 0.005 – 0.1 mg L

) immediately after
the sample was collected. Chemical analyses for CO

and CO
were undertaken on the sam-
pling day applying the German Standard Methods for pK
and pK
determination (DEV 2005). Oxygen
concentration was analysed using the WINKLER method, soluble reactive phosphorus (SRP), ammonia
and nitrate were analysed according to US APHA Methods (1998) in the Laboratory of Chemistry,
University Central, Quito, Ecuador. Water chemistry of the non-reactive cations and anions were carried
out in the laboratory of the Berlin University of Technology, Dept. of Water Quality Control, Germany,
using the acid-preserved water samples. Water samples for the determination of total amounts of ions
were digested in an autoclave under acidic conditions using K
(121 °C, 1.3 bars for 2 h). Total
phosphorus and total nitrogen were analysed photometrically by flow injection analysis in accordance
with EN ISO 15681-1 (2004) and 11905-1 (1998), respectively (Foss Tecator FIAstar 5000) with a
detection limit of 0.005 μg L
The cations Ca
, Na
, Mg
, K
, As
and Fe
were analysed by flame AAS (GBC Scientific Equip-
ment, Pty. Ltd. Victoria, Australia), lower concentrations of Li
, Fe
and Mn
(< 0.1 mg L
) were ana-
lysed by graphite furnace ASS (Varian Spectra A-400). The anions Cl

, SO
and NO

were analysed
using an ionic chromatograph (AS 50 Dionex) with CD 20 detector, GD 50 gradient pump and an AS
11 column for separation. Boron was determined photometrically in accordance with DIN 38405-D17
(DEV 1981; Dr. LANGE LCK 307). Analyses of the gases CO
, CO, O
, N
, N
O, and CH
were carried
out using a GC with FID and TCD detectors at the Leibniz-Institute of Freshwater Ecology and Inland
Fisheries, Berlin, Germany.
C-analyses of soils were carried out by the Leibnitz Institute for Applied
Geosciences, Hannover, Germany.
2.4. Biological and Microscopic Analyses
The determination of seston (bacteria, precipitations, detrital flocs) was done by Scanning Electron
Microscopy as with Energy Dispersive Spectroscopy. For SEM-EDS analyses the wet polycarbonate
filters were fixed on a bracket, air dried and then sputtered with gold or carbon. A SEM-EDS Hitachi
S 2700 electron microscope was used with an acceleration voltage of 20 kV and an IDFix hardware
and software from SAMx for analysis.
Phytoplankton was determined qualitatively using a Zeiss Laboval 4 and quantitatively according
to the Utermöhl method (DEV 2007) using a Olympus microscope CK30, about up to 5 transects of
the Utermöhl chamber (2 to 50 ml) were counted for quantitative determination, for the calculation of
the biovolume 20-100 cells were measured in the Zeiss using an ocular with a reticle. The biovolume
was calculated using related geometrical bodies (PADISÁK and ADRIAN, 1999). Diversity, abundance and
biovolume were determined for each of the 11 sampling depths.
Determination of macroinvertebrates was carried out by Dr. CORREOSO, Catholic University of
Limnology of a Crater Lake 107
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3. Results
3.1. Lake Formation and Morphometry
The strato volcano Cotacachi was active during the middle Pleistocene period up to
630 000 years ago (INECEL 1983), which documents the end of volcanic activities. Cui-
cocha, a parasitic volcano of the Cotacachi, began its activity with a series of eruptions
4500–3000 years BP followed by the collapse of the dome and a caldera was formed.
Final eruptions took place more or less simultaneously about 1450–1330 BP and led to
the formation of four domes within the caldera, nowadays the islands of Yerovi and Wolf
(GRUPE et al., 2008; Fig. 1), and the differentiation of the caldera into two main basins of
different depth. Following the extinction of the Cuicocha volcano, the caldera was partly
filled with sediment from the catchment area; fine grained and clay material sealed existing
fissures and fractures within the caldera and after the end of the colmation process, lake
water accumulation began, probably a few hundred years ago.
Nowadays Lake Cuicocha is situated about 3072 m a.s.l. and has a maximum water depth
of 148 m, a mean depth of 72 m; the volume amounts to 0.28 km
and the lake surface
3.78 km
(Table 1). The lake morphometry is determined by the caldera flanks with steep
inclinations, and in some parts of the crater rim, a water depth of 100 m is reached only 20 m
from the shore line. The bathymetric map (Fig. 2) was developed on the basis of 1250 data
points and shows two lake basins, one with a maximum depth of 148 m extending east of
the islands, and the other with a depth of 78 m, situated in the western area of the lake.
From the catchment area, water inflow into the caldera occurred until recently, however,
the watershed is small (18.2 km
) with a surrounding factor of only 5.9. Additional hydro-
thermal water has been entering into the caldera and mainly influences the water chemistry.
The lake’s water conductivity clearly points to a significantly increased ionic content com-
pared with the inflow water of the catchment, and a rough estimate shows in total about
30% hydrothermal water, based on rain water inflow and recent hydrothermal water quality
(see below).
3.2. Lake Stratification and Mixing Processes
Lake Cuicocha is a weakly thermal stratified lake with a monomictic cycle, and overturn
occurs during the windy dry period from July to October (Fig. 3). The epilimnion stretches
down to 40 m. Temperature differences between surface waters (approx. 16–18 °C) and the
deep water body were small and amounted to about 2 °C during stratified periods. The stabil-
ity of the stratification in Lake Cuicocha was further reduced due to nocturnal cooling and
deep lake mixing, the so called atelomixis, which was analysed in detail in the nearby Lake
San Pablo (GUNKEL and CASALLAS, 2002, 2002a). During a 24 hour period in March during
stratification period (Fig. 4), heating occurred in the upper water body of 50 m, mainly as
a consequence of daily radiation input; and in the deeper water layer of 70 to 100 m depth
a decrease in temperature was registered due to convection currents which occurred during
the night after cooling of the surface layer.
An input of hydrothermal water was detected in the 78 m lake basin, and near the lake
bottom both temperature and conductivity increased significantly (ΔT = about 0.04 °C;
ΔCond = about 25 μS cm
; Fig. 5), resulting in a decreasing density of Δρ
= 0.00915 kg m

(P < 0.0001) in the 70 m water depth compared to the 55 m depth. In the whole lake density
of the water increased from the surface (ρ = 999.0191) to the ground (ρ = 999.2731 in 78 m
depth, ρ = 999.2986 in 148 m depth), but no chemocline was build up and the hydrothermal
water was quickly mixed with the lake water. The inflow of hydrothermal water at the lake
floor could not be quantified. Well known is the hydrothermal water inflow at the shore line
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of Island Yerovi, which was used for water chemical analyses, in 2005 an additional inflow
of hydrothermal water was observed near the Island Yerovi in 5–10 m water depth.
The conductivity profile in the 148 m lake basin showed no increase, and an inflow of
hydrothermal water must be excluded.
In the western basin of Lake Cuicocha, at 78 m depth, a permanent emission of CO
place detected by sonar and few gas bubbles reached the lake surface. The composition of
the gas (see below) clearly pointed to the volcanic origin. The emission of the gas led to a
resuspension of sediments proved by sonar and underwater filming and to a partial mixing
of the water body by billows during upraising of gas bubbles.
Figure 1. Lake Cuicocha with domos Yerovi (left) and Wolf (right).
Table 1. Morphometric data of Lake Cuicocha.
Parameter Lake Cuicocha
Lake coordinates
North–South N0 18 49.9 W78 21 40.1 – N0 17 32.6 W78 21 28.8
West–East N0 18 22.1 W78 22 33.1 – N0 18 03.9 W78 20 52.8
Watershed (km
) 18.2
Surrounding factor 5.9
Lake water level (m a.s.l.) 3,072
Length (m) 3,238
Width (m) 2,232
Shore line without islands (km) 10.14
Shore line with islands (km) 14.43
Shore line development without islands 2.9
Shore line development with islands 4.2
Surficial area (m
) 3,781,012
Maximum depth (m) 148
Mean depth (m) 72
Relative depth (%) 3.3
Volume (m
) 282,053,575
Volume of the islands underwater (m
) 103,648,907
Volume of Jerovi above water line (m
) 9,011,699
Volume of Wolf (m
) above water line (m
) 29,381,720
Limnology of a Crater Lake 109
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3.3. Water Chemistry and Post-Volcanic Activities
Gas analyses showed that the main composition was CO
(51%) and N
(23%) with
smaller amounts of O
(3.0%) CH
(1.7%) and CO (0.3%). Gaseous emissions of boron
compounds led to increased BO
concentrations in the lake water, a typical component of
volcanic gases.
The pH of Lake Cuicocha in the epilimnic water (0–40 m) was weakly alkaline, the pH
was raised to 8.3 as a consequence of loss of CO
from the oversaturated water to the atmos-
phere and primary production. The hypolimnion (> 40 m) exhibited neutral conditions with
a pH down to 7.0 (Fig. 6), caused by CO
input from volcanic gases from the lake floor in
the western basin (GUNKEL et al., pers. com.).
The carbon dioxide concentrations were elevated in the whole water column with the
highest concentrations in the deep water layers > 60 m, a consequence of the hypolimnic
gas input in the shallow basin, combined with high wind-induced horizontal water cycling
between the different parts of the lake. In the deep water, an oversaturation of CO
of up
to 78 times, compared with the atmosphere and the local pressure was observed, this cor-
responded to a CO
concentration of 40 mg L
or 31 mL CO
per litre water. Nevertheless,
the in situ CO
saturation of the hypolimnic water was low, due to the high pressure of
the water column in the deep water, where great quantities of CO
are soluble. Taking into
consideration the partial pressure of the gas, leaving the volcano (xi = 0.39%), the in situ
saturation of CO
in the bottom-near water layers amounted to only 0.08%.
Figure 2. Bathymetric map of Lake Cuicocha, ♦ = sampling points, ∇1 = cascades, ∇2 waterfall.
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Figure 3. Temperature isopleths of Lake Cuicocha, based on 10 profiles.
Figure 4. Temperature profiles and temperature differences over a 1 day period during lake stratifica-
tion period, Lake Cuicocha, 148 m depth position, 30/31.3.2004.
Limnology of a Crater Lake 111
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The dissolved ionic concentration of Lake Cuicocha resulted in a conductivity of about
810 μS cm
; this was elevated compared with the water inflow of the cascades (∼ 260 μS
) and a nearby rain water-filled caldera lake, Lake Mojanda (mean 43 μS cm
). Thus the
Cuicocha lake water must be influenced significantly by the input of hydrothermal water.
Lake Cuicocha is a sodium hydrogen carbonate water (60.6 mg L
, 189 mg L


) with significant amounts of magnesium and calcium as cations (29.8 mg L
46.8 mg L
) and chloride as anion (70.5 mg L

, Table 2). Ions of minor concen-
trations were the cations K
, Li
, Fe
, Mn
, Al
and the anions SO
, SiO
, B(OH)

In Lake Cuicocha the trophic level is very low, and a median phosphorus concentration of
12 μg L
was observed. The nitrogen concentrations were also shown to be low (mean
96 μg L
). The N/P ratio (median = 10.0, mass basis) for most data showed P limita-
tion. Due to the volcanic origin of the lake, the siliceous concentration (median = 21 mg L

) was very high and exceeded the normal range in natural lakes.
Pelagic calcium concentration varied between 39.8–56.1 mg L
(5- and 95-percentile),
and the epilimnic Ca saturation index of + 0.75 (0–30 m) showed a slight oversaturation.
Nevertheless, precipitation of calcium carbonate was registered at the shore line of Lake
Cuicocha. These precipitations were continuously built up by epilithic algae and cyanobac-
teria (see below), leading to CaCO
crusts of about 1 cm in thickness, which reached down
deep into the epilimnion. Wave action partly destroyed these crusts, forming fine carbonate
debris, which sank to the lake bottom, where dissolution of the CaCO
precipitates occurred
under conditions of negative Ca saturation index (S = –20.2 at 45–135 m, see Fig. 8). An
up-welling of hypolimnic water by the lake floor heating (see 3.2.) led to an internal calcium
Figure 5. Increases in temperature and conductivity with decrease of the density near the lake bot-
tom in the 78 m lake basin; (n = 107; variances R
for temperature = 0.8298, conductivity = 0.8913,
density = 0.8297).
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The inflow of hydrothermal water was analysed at the shore of Island Yerovi. The water
was of relatively low temperature (23–26 °C), rich in sodium (247 mg L
), calcium
(120 mg L
) and iron (7.4 mg L
) as cations and sulphate (193 mg L
chloride (171 mg L

) and silicate (110 mg L
) as anions.
3.4. Littoral Zone and Travertine Formation
At the littoral zone of Lake Cuicocha the reed plant Totora (Schoenoplectus totora) built
up a dense but narrow vegetation zone, followed by a few submerged plants including
Myriophyllum quitense (from 0–2.3 m depth), Potamogeton illinoensis (2–8 m), P. pectina-
tus (2–14 m), Chara rusbyabana (27–29 m), Ch. globularis (8–30 m) and Nitella acuminata
(20–35 m). As a new proof Drepanocladus capillifolius, a Bryophyta, Fam. Amblystegiace-
ae, was found at 20–28 m depth (KIERSCH et al., 2004). The biomass of the submerged
macrophytes was very high and they built up dense stands with very long plants with several
meters lengths on the high flanks.
The invertebrates of the littoral zone were dominated by Hyalella cf. dentata, Fam.
Malacostraca; GONZALEZ and WATLING (2002). Other species with regular mass develop-
ment were Dugesia sp. and some Gastropoda species, Biomphalaria sp. and Potamopyrgus
antipurvarum (Table 3). The biodiversity was very low and the few species occurred in high
Only one fish species was observed and filmed by divers, the Andean catfish Astroblepus
ubidiai (Siluriform), living in the upper part of the littoral zone was a new proof for this
lake. This fish is a resident species of the high Andes, with a few remnant populations in
this area (VÉLEZ-ESPINO and FOX, 2005).
The colonization of littoral interfaces by benthic algae and bacteria was of high ecological
importance. A dense biofilm with intensive photosynthetic activity induced the precipita-
tion of travertine on the caldera flanks and consisted mainly of motile pennate diatoms
(Epithemia argus, Mastogloia smithii, Cymbella pusilla and Rhoicosphenia abbreviata as
dominant species) and one frequent cyanobacteria species, Calothrix parietina, which lived
Figure 6. pH isopleths of Lake Cuicocha, based on 10 profiles.
Limnology of a Crater Lake 113
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Table 2. Water chemistry of Lake Cuicocha, median and percentiles of 2003–2005;
n = 84.
Parameter 5 % Median 95 %
7.65 8.14 8.58
7.18 7.37 7.86
Conductivity (μS cm
) 797.2 812.3 818.2
(mg L
) 55.9 60.6 69.0
(mg L
) 39.8 46.8 56.1
(mg L
) 4.7 5.5 6.4
(mg L
) 0.100 0.110 0.135
(mg L
) 25.3 29.8 41.2
(mg L
) 0.000 0.006 0.022
(mg L
) 0.004 0.0120 0.073
(mg L
) 0.001 0.005 0.083
(mg L
) 0.003 0.009 0.139
(mg L
) 2.4 3.9 5.8

(mg L
) 138 189 351

(mg L
) 64.0 70.5 74.4
(mg L
) 9.2 13.4 18.3
(mg L
) 17.1 20.0 23.3
(mg L
) 17.0 21.0 23.4
(mg L
) 0.029 0.096 0.296
(mg L
) 0.004 0.011 0.027
SRP (mg L
-P) 0.002 0.005 0.013
N/P (weight based) 2.5 10.0 28.9
(mg L
) 5.9 6.7 7.0
(mg L
) 0.8 2.0 3.2
Table 3. Fauna in Lake Cuicocha (rare to mass describes the relative abundance in a seven
grade scale).
Class/Subclass/Family Taxon Abundance
Littoral fauna:
Turbellaria Dugesia sp. frequent
Gastropoda, Planorbidae Biomphalaria sp. frequent
Gyraulus sp. less frequent
Physidae Physa sp. less frequent
Hydrobidae Potamopyrgus antipurvarum mass
Bivalvia, Spaeridae Sphaerium forbesi less frequent
Crustacea, Ostracoda Cypris sp. frequent
Malacostraca Hyalella cf. dentata mass
Insecta, Zygoptera Oxyallagma dissidens less frequent
Osteichthyes, Siluridae Astroblepus ubidiai rare
Crustacea, Phyllopoda Daphnia pulex pulex 0.2–3 n L
Crustacea, Copepoda Metacyclops mendocinus WIERZEJSKI 0.11–7 n L
Malacostraca Hyalella cf. dentata 0.001–0.06 n L
© 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.revhydro.com
at the surface and inside the pore system of the CaCO
crusts (Fig. 7). The intensive primary
production in this biofilm was proved using oxygen measurements, which showed a mean O

oversaturation of 199% (min. = 152%, max. = 216%, n = 13) 5 mm inside the CaCO
resulting in a microenvironment with increased pH and favoured carbonate precipitation; the
crusts were composed of pure calcium and contained only traces of magnesium.
3.5. Plankton Community
The phytoplankton community was characterized by Diatomeae (Aulacoseira granulata,
Fragilaria ulva var. acus, Fragilaria sp.), Crytophyceae (Rhodomonas and Cryptomonas),
Euglenophyceae (Trachelomonas volvocina) and some Chlorococcales species (Oocystis
lacustris, O. naegli, O. parva and Scenedesmus linearis, S. quadricauda, S. acuminatus,
Figure 7. Calcium carbonate precipitations within the biofilm at the lake shore, REM picture of the
calcium crusts with some diatoms embedded in CaCO
Limnology of a Crater Lake 115
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Table 4. Occurrence of phytoplankton in Lake Cuicocha, abundance of algae (n mL
), n.d. = not
determined. The abundance is given as the maximum cell number in one of the 11 sampling depths
(sampling depths: 0, 5, 15, 30, 45, 60, 75, 90, 105, 120, 135, 145m), *) biomass without Mougeotia sp.
and Spirogyra sp.
03/2004 08/2004 03/2005 08/2005
Cyanobacteria n.d. 3.306
Asterionella formosa < 4
Fragilaria sp. < 4 72
F. ulva var. acus 18 22
Navicula sp < 4 < 4
Nitzschia sigmoides 4
Pennales n.d. < 4 < 4 91 4
Melosira sp. < 4
Aulacoseira granulata 117 49 108
Aulacoseira sp. 108
Peridinium sp. < 4 4
Gymnodinium sp. 9 < 4
Cryptophyceae 4
Rhodomonas minuta 9
Rhodomonas sp. 90
Cryptomonas sp. 9 4 13 31
Trachelomonas volvocina 1.054 22 40 9
Trachelomonas verrucosa 31 13 4
Chlamydomonas sp. 81 40 439
Monoraphidium komarkovae 251 58
Oocystis lacustris 157 175 27
Oocystis naegeli 9 4 22 9
Oocystis parva 117
Chlorella sp. < 4
Lagerheimia wratislaviensis 4
Planktosphaeria gelatinosa < 4
Pediastrum boryanum 144
Scenedesmus linearis 36
Scenedesmus quadricauda × 332
Scenedesmus acuminatus 72
Tetraedron minimum 4
Botryococcus braunii < 4
Dictyosphaerium ehrenbergianum < 4
Lagerheimia longiseta 18
Neglectella sp. 13
Schroederia setigera < 4
Cladophora sp. 583
Mougeotia sp. 193
Spirogyra sp. 108
Cosmarium sp. < 4 4
Abundance (n mL
) 1094 193 3485 363
Biomass (μm
) 784,000 218,000 3,808,000 118,000
Depth of abundance maxima 0–75 m 0–75 m 130–140 m 45–75 m
© 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.revhydro.com
Table 4). In general, the abundance (< 1–100 cells mL
) and biomass (< 400 000 μm
were small, and corresponded with the oligotrophic character and the Secchi depth of
13.0– 14.9 m. On one occasion the occurrence of Mougeotia and Spirogyra at 140 m depth
led to a maximum biomass of 3 800 000 μm
The vertical algae distribution showed continuously occurrence of algae in the whole water
column down to 140 m, and cell number as well as biovolume did not had any significant
maximum in the epilimnic zone, while other parameters such as E
, conductivity, pH, CO

and the Ca saturation index showed an analogous stratification to temperature (Fig. 8).
Algal species were regularly found in deep water and maximum abundance occurred
down to 75 m, and an extreme maximum abundance of Mougeotia and Spirogyra occurred
once at 130–140 m (Fig. 9). The depth distribution during stratification periods point out
clearly that most species occurred in the whole water column (Chlamydononas, Scenedesmus
linearis, Oocystis lacustris, Trachelonomas volvocina, T. verrucosa, Aulacoseira sp.), but a
few species were found only in the upper water body down to 80 m such as Scenedesmus
acuminatus, Lagerheimia longiseta and Monoraphidium komarkovae. Other species, mostly
filamentosus forms occurred only in depth >> 80 m such as Aulacoseira granulata, Pedias-
trum boryanum, Mougeotia sp. and Spirogyra sp. (Fig. 9).
In the plankton some iron-oxidising bacteria such as Siderocapsa occurred frequently, and
Metallogenium-like structures were also found.
The zooplankton biocoenosis of Lake Cuicocha was formed by two species, the filter
feeder Daphnia pulex pulex and the carnivorous Metacyclops mendocinus (Table 3). Zoo-
Figure 8. Vertical stratification of physical-chemical water parameters (redox potential, conductivity,
temperature, pH, CO
, calcite saturation index) and depth distribution of plankton (phytoplankton cell
number and biomass, abundance of Daphnia pulex, Metacyclops mendocinus and Hyalella cf. dentata)
in the 148 m depth position of Lake Cuicocha, 18.3.2004.
Limnology of a Crater Lake 117
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plankton abundance was low and the maximum number of animals amounted to 7 n L
the mean abundance was 1.8 n L
(Metacyclops mendocinus) and 0.7 n L
(Daphnia pulex
pulex), respectively. The vertical distribution of the zooplankton stretched down to a depth
of 140 m (Fig. 8).
In the pelagic water body, the invertebrate found in the littoral zone, Hyalella cf. dentata
was found regularly. Hyalella occurred in the whole water column with a few animals per
litre (Fig. 8), but was frequent at the lake bottom at 78 m depth, proved by under water
filming; Hyalella was also found at 148 m depth in the plankton net samples, however, at
this depth no intensive filming was performed.
In the pelagial of Lake Cuicocha no fish were found by use of sonar, Astroblepus ubidiai
occurred only in the littoral zone. A few decades before, the trout (Oncorhynchus mykiss)
was introduced into Lake Cuicocha as well as into the nearby Lake San Pablo and Lake
Mojanda, however, in Lake Cuicocha the population has broken down, while in the other
lakes, the fish populations have increased.
3.6. Sediments and Sediment Resuspension
The sediments in Lake Cuicocha formed a very thin layer of up to 10–15 cm on volcanic
rocks a few decimetres in diameter, proved by underwater photography (see GUNKEL et al.,
2008). The sediments consisted mainly of mineral compounds and sedimented diatom frus-
tules. These sediments were poor in Ca (10–20 g kg
ds, = dry substance) but rich in P
(24–46 g kg
ds). The sediments were anaerobic with sulphide formation in the form of
Figure 9. Vertical distribution of some dominate algae species in the 148 m depth position of Lake
Cuicocha (17.03.2005).
© 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.revhydro.com
pyrites, only the surface layer was oxidized down to a few millimetres. In particular, the sur-
face layer in the 78 m deep area consisted of brown flocs a few millimetres in diameter.
The sediment traps showed extraordinary high sedimentation rates with the higher rates
in deeper exposed traps (1–3 mm fresh material within 2 weeks), in contrast to the low
abundance of plankton in the pelagial. The trapped material consisted of the same type of
flocs as were found at the lake bottom, recognizable by a similar chemical composition. The
settled material of both origins was poor in Ca (12 g kg
ds) but rich in Fe (74 g kg
ds) and
consisted of algae, detritus and masses of iron oxidising bacteria (Fig. 10). Observations with
Figure 10. Sediment flocs in Lake Cuicocha, detritus, diatom cell is embedded in flocs with iron-
oxidising bacteria, SiO
precipitations and filamentosus fungi structures, free water, 40 m depth,
18.08.2005, in situ filtration.
Figure 11. Formation of amorphous SiO
in Lake Cuicocha sediment on Nuclepore filters,
148 m depth, 31.08.2004; EDS analyses verified the Si composition with traces of Al and Fe.
Limnology of a Crater Lake 119
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sonar and the underwater camera registered resuspension of sediment and sediment flocs by
gas bubbles rising up to 20 m.
Sediment forming processes were supported by two linked water chemistry processes, the
precipitation of polymeric Al(OH
and the formation of amorphous SiO
; due to the
atomic characteristics mixed precipitations are common. In the water as well as in the sedi-
ment, Si precipitation occurred and formed amorphous structures (Fig. 11). Al precipitation
occurred in the water column due to a decrease in pH to about 7, which meant that down-
welling of epilimnic water leading to oversaturation of the Al polymer, and gelatinous Al
polymers as well as Al microcrystals like gibbsite were observed (GUNKEL et al., 2008b).
Both processes were frequent and promoted the coagulation of flocs in the water, formed by
detritus, bacteria and living algal cells.
4. Discussion
4.1. Characteristics of Caldera Lakes
Crater lakes are a special type of lake, which are classified by the lake formation pro-
cess (caldera as an intrusion of the magma chamber of a volcanic dome, maar as explosive
craters), without consideration of any geological or climatic settings. Thus caldera lakes pos-
sess a wide scattering of ecological characteristics. The physical and chemical parameters of
caldera lakes were summarized by PASTERNAK and VAREKAMP (1997) and VAREKAMP et al.
(2000). Nevertheless caldera lakes are characterized by properties of ecological interest:
caldera lakes are young lakes in a state of development, mostly located at higher altitudes;
only maar lakes and some marine atoll lakes are found in the plain. The watershed is nor-
mally covered by young volcanic deposits, with weathering processes and the beginning
of soil formation. The watershed is small, the lakes do not have a regular outflow; caldera
lakes have no connection with the ground water table. Especially in young caldera lakes the
accumulated sediment layer is very small.
Lake Cuicocha is characterized by the young age of about 500 years with weathering of
the rocks in the water shed and the lake succession processes such as rock falls and land
slides from the crater rim, deposition of sediments, colmation and nutrient accumulation,
and water chemistry is strongly influenced by these processes as well as by post volcanic
activities such as gas emission and hydrothermal water inflow. One effect is the resuspension
of sediments is a process which determines the water chemistry as well as the development
of the phytoplankton (WEYHENMEYER, 1998).
The colmation of the lake basin is unstable or insufficient due to the small sediment layer
and the post-volcanic activities. Triggered by an earthquake in 1987, the lake colmation layer
was destroyed and from this time a decrease in the water level of about 30 cm per year
occurred. Water percolation leads to a higher risk of phreatic-magmatic eruption, meaning
an explosive eruption, when the infiltrating water contacts the magma of hot rocks (MASTIN
and WITTNER, 2000; GUNKEL et al., 2008).
Post-volcanic emissions including gases or hydrothermal springs are frequent in volcanic
lakes and affect lake stratification and water chemistry. In Lake Cuicocha the ionic content
of the water was increased, compared with a nearby inactive caldera lake, Lake Mojanda,
which clearly points to the high significance of the hydrothermal inflow.
4.2. Physical Characteristics of Lake Cuicocha: Mixing Processes
Besides wind-induced water cycling, lake mixing processes of Lake Cuicocha are deter-
mined by post-volcanic activities like the emission of volcanic gases, hydrothermal underwa-
© 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.revhydro.com
ter springs and a heat flux via the lake floor. Gas emissions and hydrothermal water springs
at the lake floor lead to a mixing of the lake due to billows with upraising gas bubbles or
divergent currents of heated water. These processes can be detected only by the application
of special monitoring (use of divers, sonar, underwater camera and a lake profiler).
Thus the stability of the thermal stratification of Lake Cuicocha is diminished, and atelo-
mixis occurs, that means daily or frequent periodic mixing down into the hypolimnion e.g.
by nocturnal cooling in periods of low density gradients (TALLING, 1969). Increased losses of
phytoplankton by sedimentation take place, and the atelomixis leads to a more oligotrophic
status of the lake (TAVERA and MARTÍNEZ-ALMEIDA, 2005), who postulated that atelomixis
was the driving force in determining phytoplankton composition. In the nearby Lake San
Pablo, atelomixis was proved and led to hypolimnic phytoplankton maxima (GUNKEL and
CASALLAS, 2002, 2002a).
4.3. Water Chemical Characteristics of Lake Cuicocha
Water chemistry of the caldera lake is determined by rain water interacting with vol-
canic deposits in the catchment area and represents a new (young) terrestrial-aquatic linked
ecosystem. The export of nutrients and other ions from the watershed, being composed of
volcanic deposits, lava, volcanic rocks or ashes, which undergo an intensive mineralization
and soil forming process determine the ionic composition. Typical for the Andean zone is
the high export of Al and Si as characteristic compounds of the andesites, while the content
of Ca must be regarded as low. Lake Cuicocha, only a few hundred years old, reflects these
conditions: Al and Si are increased, Al polymerisation and microcrystal formation (GUN-
KEL et al., pers. com.), amorphous SiO
precipitation as a process of sediment build-
ing. Al–Si polymerisation processes promote the formation of flocs and it must be assumed
that the stability of flocs is also increased.
Lake Cuicocha is an oligotrophic lake, due to low P and N input from the watershed with
young soils, the andisols (ZEHETNER et al., 2003) and due to no accumulation of nutrients
in the young lake. These nutrients are further reduced by intensive precipitation with iron
oxides being a consequence of hydrothermal water inflow.
In Lake Cuicocha CO
accumulation with pH change and changes in the lime carbonic
acid equilibrium occur.
The water of Lake Cuicocha has a low Ca and Mg concentration, however, in the epilim-
nion, the calcite saturation index regularly shows a positive value, meaning oversaturation,
whereas the hypolimnion values decrease to levels < 0. Travertine formation occurs as a
biological CaCO
precipitation in an extended biofilm of a few millimetres thickness with
high abundance of diatoms and median abundance of cyanobacteria. It is of interest that
only traces of magnesium occurs in the precipitates. The productivity of the biofilm, which
stretched down to more than 15 m is very high (sampling at deeper areas of the crater flanks
was not possible due to diving limitations). Part of the travertine crusts are eroded by wave
action and form a fine carbonate sand in the littoral zone, which sink to the lake floor,
where it is dissolved. This Ca cycle – Ca precipitation in the epilimnic zone, sedimentation
of Ca sands, re-dissolution in the hypolimnic zone and introduction of dissolved Ca in the
epilimnion by regular partial lake mixing is the basis of the intensive development of the
biofilm with travertine formation.
Similar travertine formation is a frequent process at the crop-out of hydrothermal springs,
where the water flow is a source of Ca, Mg and corresponding anions. In lakes it has also
been observed at a small rate, induced by photosynthetic pH changes (GOLUBIĆ, 1969). In
crater lakes intensive travertine formation is known e.g. from Lake Quilotoa, Ecuador (up
to 50 cm thickness, own observations) and the Ries crater, Germany (RIDING, 1979), but in
Lake Quilotoa an abiotic formation process with extreme Ca over-saturation and aragonite
Limnology of a Crater Lake 121
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precipitates can also become of high significance, and under these conditions Ca–Mg-car-
bonate precipitates occur.
In the hypolimnion the Ca concentration is in the range of undersaturation, due to decreas-
ing pH caused by increased CO
concentrations. This Ca undersaturation in the hypolimnion
leads to re-dissolving of biogenic precipitated Ca, resulting in a very small amount of Ca
in the lake sediments of < 1%. Consequently the sediment building rate is reduced and the
lake colmation process is hindered and slowed, and the stability of the colmating layer must
be regarded as low.
4.4. Biocoenosis of Lake Cuicocha
The phytoplankton community is characterized by a low diversity, which corresponds to
the nearby glacial Lake San Pablo (Lake San Pablo: 31 species, Cuicocha: 37 species), but
only 8 species occur in both lakes (GUNKEL and CASALLAS, 2002). Most of the species in
Lake Cuicocha are wide spread, endemic species of Latin America were not found, espe-
cially of the group of diatoms (LANGE-BERTALOT, pers. com. after sample revision).
The dominant species in 2004/2005 are Aulacoseira granulata, Fragilaria ulva var. acus,
Trachelomonas volvocina, Oocystis lacustris, O. parva and Clamydomonas sp., which have
to be classified as meso- to eutraphent species (trophic value 3–3.5) with low to median
saprobe character (saprobic value 2), these species are cosmopolite with a wide ecological
tolerance (LUB 2005). Only Monoraphidium komarkovae is a more oligotraphent species
(trophic value 1.5). The more eutrophic species in Lake Cuicocha do not correspond to the
high water transparency and the low phosphorous concentrations, but it must be considered,
that the algae biomass is low and the continuity is less.
Of high interest is the vertical distribution of the algae with a significant distribution
below the epilimnic zone, the calculated euphotic depth is 18–20 m (TILZER, 1988), but
most of the algae we found in depth > 20 m; this is a phenomenon already investigated in
the nearby Lake San Pablo, and atelomixis as deep nocturnal mixing caused the temporary
residence in the aphotic zone (GUNKEL and CASALLAS, 2002, 2002a). A significant part of
the population is even in more than 80 m depth, some species such as Aulacoseira granu-
lata, Pediastrum boryanum, Mougeotia sp. and Spirogyra sp. had a maximum distribution in
130–148 m. This paradox seems to be a consequence of the intensive flocculation processes
due to Al and Si precipitation, the flocs trap algae cells and promote sedimentation processes
and lead to an accumulation of algae near the sediment.
The biocoenosis of the zooplankton is characterized by some ubiquists which are highly
abundant, however, the biodiversity is strictly reduced. Only one filter-feeding Phyllopoda,
one carnivorous Copepoda and one Malacostraca form a pelagic food web with low phyto-
plankton and bacterioplankton at the primary trophic level. Top predation by fish is missing,
a catfish being the only inhabitant of the littoral zone. A very restricted food web was also
observed in the nearby Lake San Pablo, a glacial lake of 38 m depth (CASALLAS and GUNKEL,
2001). Without doubt high mountain lakes in this tropical area show infrequent colonization
by aquatic organisms, and the fauna and flora are characterized by introduced species (direct
introduction of fish, and by ballast water from water planes). Tropical Middle America is a
barrier for migration to cold water species from North to South America, and inversion of
species occurred only from the south of South America via the Andes mountain range.
Little is known about the occurrence of Hyalella cf. dentata, a species of the new world,
and even the description of some Hyalella species must be revised (GONZALEZ and WATLING,
2002). Hyalella cf. dentata found in many South American lakes, even in salt lakes, and in
some North American lakes (GONZALEZ and WATLING, 2002). However, information on the
habitat and ecology of this organism is insufficient. Although Hyalella cf. dentata is a her-
bivorous littoral species, its occurrence in the pelagial and at the lake bottom was regularly
© 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.revhydro.com
registered at 148 m, and at the 78 m depth a high abundance was observed. However, due
to the extension of the lake, no leaves or similar particulate organic material can accumulate
in the pelagial and on the lake floor. It can be assumed, that filamentous algae such as the
desmids Mougeotia and Spirogyra, observed in deep water in high biomass (Table 4), play
a role in the feeding behaviour of these organisms.
This investigation did not include any microbiological studies, however, some interesting
effects were observed. Iron-oxidizing bacteria were frequent, mostly of the genus Side-
rocapsa as well as Metallogenium-like structures (KLAVENESS, 1977), which were classified
by EMERSON et al. (1989) as manganese-oxidizing fungi. These bacteria, mainly the filamen-
tous forms of Metallogenium-like structures promote the build-up of flocs and increase their
stability. Flocs are cycled in the water body by sediment resuspension and in turn strongly
influence water quality. The role of flocs in chemical and biological processes in lakes must
be regarded as very important, however, available knowledge is still insufficient (WEYHEN-
MEYER, 1998). In Lake Cuicocha flocs played a significant role in the occurrence of algae,
serving as a substrate for adhesion and as a nutrient source in the oligotrophic lake.
The lack of fish within the pelagial may have been caused by the high CO
however, it should be noted that in the nearby high Andean lakes only a few species such
as a hybrid (Carassius carassius, C. auratus) called carp, trout (Oncorhynchuss mykiss) and
pike (Micropterus salmoides) occur. Fish biodiversity is extremely low in these lakes and
little is known about the native biocoenosis (e.g., Atroblepus ubidiai), because trout, carp
and pike were introduced into all these lakes.
The littoral macrophytes had a high biomass, and invertebrates in the littoral zone were
frequent, whereas the planktic community was of low abundance and biomass. This increased
importance of the littoral biocoenosis has also been observed in other tropical high moun-
tain lakes, and one reason for this seems to be the increased losses of pelagic species due
to deep diurnal mixing processes (GUNKEL and CASALLAS, 2002, 2002a). In Lake Cuicocha
this process is forced by the formation of flocs and the coagulating effect of polymeric Al
precipitation, by the formation of amorphous SiO
and filamentous Metallogenium
like structures, and by the stabilization of flocs by the abundant occurrence of Siderocapsa
5. Acknowledgements
This study is part of a cooperation project of the Berlin University of Technology, Germany, Dept.
Water Quality Control with support from the Dept. of Environment and Marine Science and Technology
(VWS), Dipl. Geol. B. GRUPE, and the Central University of Quito, Facultad de Ingeniería en Geología,
Minas, Petróleos y Ambiental (FIGEMPA), Ecuador, Ing. F. VITERI. It was financed by the Deutsche
Forschungsgemeinschaft, Germany (DFG, German Scientific Society) and the Ministry for Economic
Development and Cooperation, Germany (BMZ). Logistic support was provided by the National Park of
Cotacachi, Ibarra, Ecuador. Dr. P. CASPER, Leibniz-Institute of Freshwater Ecology and Inland Fisheries,
Germany, carried out gas analyses. Dr. GABRIELE GUNKEL, Berlin, Germany, carried out the phytoplank-
ton analyses. Dr. CORREOSO, Catholic University of Quito carried out the systematic identification of the
C-analyses of soils were carried out by the Leibnitz Institute for Applied Geosciences,
Hannover, Germany.
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Manuscript received March 6th, 2008; revised July 11th, 2008; accepted August 19th, 2008