RIVER RESEARCH AND APPLICATIONS

River. Res. Applic. 24: 698709 (2008)

Published online in Wiley InterScience
(www.interscience.wiley.com) DOI: 10.1002/rra.1136

TROPHIC GRADIENTS IN A LARGE-RIVER DELTA: ECOLOGICAL

STRUCTURE DETERMINED BY CONNECTIVITY GRADIENTS IN THE
DANUBE DELTA (ROMANIA)
HUGO COOPSa,b, LAURA L. BUIJSEe, ANTHONIE D. (TOM) BUIJSEa,b*, ADRIAN CONSTANTINESCUc,
SILVIU COVALIOVc, JENICA HANGANUc, BAS W. IBELINGSd,f, GEERT MENTINGb, ION NAVODARUc,
RO
Kc
WILLEM OOSTERBERGb, MIRCEA STARASc AND LILIANA TO
a
Deltares, PO Box 177, 2600 MH Delft, The Netherlands
RWS-Centre for Water Management, P.O.Box 17, 8200 AA Lelystad, The Netherlands
c
DDNI, 165 Babadag Str., 820112 Tulcea, Romania
d
EAWAG, Centre for Ecology, Evolution and Biogeochemistry, Seestrasse 79, 6047 Kastanienbaum, Switzerland
e
Alterra, Droevendaalsesteeg 3, 6708 PB Wageningen, The Netherlands
Netherlands Institute of Ecology, Centre for Limnology, Rijkssbraatweg 6, 3631 AC Nieuwersluis, The Netherlands
b

ABSTRACT
There are over 300 lakes interconnected by riverbranches and man-made canals in the Danube Delta (Romania). A multidisciplinary survey of these riverine lakes situated in large wetland complexes was made comprising hydrological modelling,
remote sensing and monitoring of water quality, plankton, aquatic vegetation and fish communities. Between-lake differences in
water quality and aquatic vegetation cover were inferred from satellite image. Based on channel and lake dimensions,
hydrological characteristics of the lakes in the delta were determined: hydrological distance from a riverbranch, residence time
and impact of reed water. Water-quality and biotic parameters (phyto- and zooplankton, submerged plants and fish) were
sampled in a comparative survey in June 19971999 in subsets of the lakes. In a follow-up study in 20012002, seasonality of
aquatic vegetation and fish were recorded.
A clear distinction was found in three lake types: (1) inflow lakes at short distance to the river, with a high flushing rate, high
load of suspended minerals but low chlorophyll concentrations, high cover of floating and submerged vegetation and dominance
of eurytopic fish; (2) large, relatively deep lakes with moderately long residence time, high Potamogeton-cover that collapses
during summer, and dominated by eurytopic fish; (3) smaller, isolated lakes at the longest distance from the river, with long
residence time, of which the water quality is strongly influenced by water flows through large floating reedbeds (blackwater),
with a high vegetation cover of Ceratophyllum demersum and Nitellopsis obtusa throughout the summer and a limnophilic fish
community tolerant for low oxygen conditions.
Aquatic vegetation showed a clear seasonality related to the interaction between plant development and light conditions.
Seasonal changes in fish distribution within the system merely followed the vegetation trends in different lakes and the
conditions in the river channels.
Past trends in the trophic gradient are discussed in the context of hydromorphological modifications in the Delta. There is a
high potential for restoring natural processes owing to the high connectivity and large scale of the system. Copyright # 2008
John Wiley & Sons, Ltd.
key words: connectivity; Danube Delta; ecohydrology; fish; hydrological model; macrophytes
Received 31 August 2006; Revised 30 May 2007; Accepted 25 September 2007

INTRODUCTION
Riverine wetlands and lakes in the lower reaches naturally have a high primary productivity supporting diverse and
abundant populations of, for example fish and birds. However, current nutrient loads that enter river deltas are far
beyond natural values, due to increased emissions and altered riverfloodplain connections. The increased nutrient
loading of delta wetlands may strongly have affected the trophic state of these ecosystems (Pringle, 2001; Dodds,
2006). The impact of increased nutrient loading may have increased further due to man-made hydromorphological
changes in riverine systems, such as channelization, dredging and loss of floodplain areas.
*Correspondence to: Tom Buijse, Deltares, PO Box 177, 2600 MH Delft, The Netherlands. E-mail: tom.buijse@deltares.nll

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ECOLOGICAL GRADIENTS IN THE DANUBE DELTA

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As a result, eutrophication effects may be more pronounced. Among the potential effects of eutrophication is the
loss of resilience and a switch from clearwater, macrophyte-dominated ecosystems to turbid ecosystems dominated
by algal blooms (Scheffer, 1998). Eutrophication effects may vary among and within riverine wetlands due to
differences in residence time, flood disturbance intensity, and processing of dissolved nutrients during water flow
through the system (Behrendt and Opitz, 1999). Generally, natural large riverfloodplain systems show a lateral
gradient of decreasing nutrient levels at longer distance from the inflow points (Gregory et al., 1991).
The present study was set up to explore the presence of trophic gradients in lakes situated in the complex delta of
the Danube River. The Danube Delta, at the outflow of the second largest European river into the Black Sea, has
undergone various man-induced changes but large-scale riverine processes remain relatively unaltered (Margesson,
1997; Buijse et al., 2002). The scale of the area is extensive (5640 km2), and the numerous lakes and vast reed
marshes still harbour a rich flora and fauna. Past efforts to enhance the economic exploitation of the area (fisheries,
reed harvesting, hunting, agriculture) and to improve accessibility (navigation channels) have resulted in an
increased water exchange between the main riverbranches and the alluvial lakewetland complexes (Romanescu,
1998).
In this study, variation in the trophic condition of delta lakes along the connectivity gradient was described and
analysed using remote-sensing information, hydrological modelling and surveys of water quality and ecosystem
components. In the first stage, a typology of lake types was derived that provided a framework for a more in-depth
study of the spatio-temporal dynamics of aquatic vegetation and fish.
Connectivity-related mechanisms in the aquatic system may be of prime importance for managing the delta to
maintain its production and biodiversity values. Modifying the degree of hydrological connectivity between the
river and lake wetlands in the floodplain may be a major tool for management.

STUDY AREA
The Danube Delta (Figure 1) is situated at the outflow of the Danube River, the second largest European river
(2860 km, catchment area 805 300 km2, average annual discharge at the base of the Delta 6750 m3 s
1), into the
Black Sea. The core of the Delta is roughly delimited by three major riverbranches, flowing through 5800 km2 of
wetlands, sand ridges and polders. The wetland complexes consist of vast reedbeds dotted with >300 lakes, situated
from close to river water inflow to strongly isolated. Water flow through the wetland complexes is greatly
accommodated by a network of natural and artificial channels.
The complexes represent different phases in the geological development of the delta, and are between c. 10 000
(fluvial part) and c. 5000 (marine part) years old (IUCN, 1992). The water level is basically determined by the levels

Figure 1. Map of the Danube Delta, showing the three main deltaic branches, the lakes and the network of natural and artificial channels. To the
south are several large lagoonal lakes
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H. COOPS ET AL.

Figure 2. Mean water levels (19942001) of the Danube River at Tulcea, at the head of the Delta (thick line); range over this period is shown, by
normal lines, water levels in 19961998 by hatched lines

of the Danube River; water levels at the upstream end of the delta at Tulcea are depicted in Figure 2. Lakes in the
marine part of the delta have an annual water level amplitude of max. 0.70 m, whereas the range in fluvial lakes is up
to 2.00 m. The lakes vary greatly in size (0.743 km2) and average depth (1.73.9 m), the fluvial lakes being on
average smaller and shallower than those situated more downstream.
The natural water flow through the delta has been altered by attempts to promote the production of reed and fish,
based on enhancing river water inflow and improving navigation conditions (Antipa, 1931) as well as a partially
conducted project to reclaim large part of the Delta.

METHODS
Remote sensing
A supervised classification of all lakes in the Danube Delta was made based on an atmospherically corrected
Landsat-TM image of July 1996. Features of reflectance in spectral bands 14 were combined in feature spaces
using the image analysis programme ERDAS-Imagine. Characterization of categories was made using available
field knowledge of aquatic vegetation and transparency sampled during field surveys in June 1996 and 1997. The
classified image was accurate in distinguishing between floating aquatic vegetation and submerged vegetation, and
between clear water and turbid water with suspended sediments or dense phytoplankton biomass, respectively.
Hydrological modelling
Hydrological characteristics of lakes in the Danube Delta were modelled using the one-dimensional model
SOBEK (WL/Delft Hydraulics and Rijkswaterstaat, 1995), which is based on a nodal frame representing lakes and
interconnecting channels (including throughflow marshes), using their dimensions (size, depth) for calculating the
distribution of water through the system. The model calculates water levels and flow velocities in any of the
modelled lakes (n 79) based on the upstream (Tulcea, depending on river discharge) and downstream (Black Sea,
fixed) levels at any given time.
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ECOLOGICAL GRADIENTS IN THE DANUBE DELTA

Based on the data the following hydrological variables were applied for each lake:
-

Travel time: average time (days) for water from one of the riverbranches to reach a lake
Residence time: volume of a lake divided by the flow in or out of the lake
Cumulative residence time: sum of travel time and residence time
Reed water: relative proportion of water flowing into a lake through reed marsh (as opposite to channel flow).
Sediment type of each lake was obtained from soil maps of the Danube Delta.

Water chemistry, macrophytes and fish

Water quality during 19961998 was sampled in a monitoring programme covering 23 lakes in the delta (David
and Hulea, 2001). Monthly water samples were taken in the centre of each lake and analysed according to ISO
standards. Hydrochemical parameters (inorganic suspended solids, P-total, P-ortho (flow analysis, ISO 15681),
NOx (flow analysis and spectrometric detection, ISO 13395), NH4 (flow analysis and spectrometric detection, ISO
11732), chlorophyll a (Romanian standard using UV detection), water temperature and Secchi-disk transparency
were determined.
Biotic parameters of lakes were obtained by sampling of phytoplankton, zooplankton, aquatic vegetation and
fish.
Chlorophyll a concentrations were measured in triplicate after filtration over Whatman GF/F filters (Maidstone,
UK); ethanol was used as extraction fluid and acid was used to correct for phaeopigments (Moed and Hallegraef,
1978). Additionally, phytoplankton composition was determined in a sub-set of 13 lakes in the monitoring
programme. On every occasion, a 1 L sample was taken from the surface layer and fixed in lugol. From the
composition of 1 ml concentrated sub-samples, the distribution of major phytoplankton groups (Chlorophyta,
Bacillariophyta, Cyanophyta) was determined.
For zooplankton, 10 L samples were filtered over a 55 mm mesh plankton net. Samples were fixed in
formaldehyde. Microscopic counts were made of sub-samples.
Aquatic macrophyte vegetation was determined in 21 lakes in June of different years, In each lake, vegetation
cover and species abundances were estimated at c. 10 sites of c. 100 m2 representative for the lake. The macrophyte
vegetation was classified using the typology presented in Coops et al. (1999). From the data of the individual lakes,
the proportion of the lake covered by vegetation was calculated.
Fish sampling was done in 11 lakes in June 1996, 1997 and 1998. Multi-mesh gillnets were used. Additionally,
fishes were sampled using electrofishing. Species, length and weight of all individual fish were determined.
Estimates were made of total fish biomass and species composition (percentage of total biomass). See Navodaru
et al. (2002) for further details.
Lake typology and seasonality
Ordination of lakes based on hydrological, hydrochemical and biotic variables was performed based on Principal
Components Analysis, using CANOCO (Ter Braak and Silauer, 1998). The (indirect) ordination was based on
biotic components and then correlated with the abiotic factors. Missing values in the dataset were replaced by
averages. For each biotic component parameters describing abundance and composition were selected (Table I).
Seasonal changes of biotic condition of lakes were studied by additional surveys of aquatic vegetation and
sampling of fish in June and September 20012002, focussing on the central lake complex (IsacUzlina) of the
Danube Delta.

RESULTS
Remote sensing
Supervised classification of the Landsat-TM image produced spectral reflectance patterns that could be
interpreted as different types of surface water. It was possible to discern at pixel level clear water without or with
Copyright # 2008 John Wiley & Sons, Ltd.

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Copyright # 2008 John Wiley & Sons, Ltd.

Bream Abramis brama,

Gibel carp Carassius gibelio,
Roach Rutilus rutilus

>8 km2
>2 m
1080 days
Sand/silt
7.8  5.5; 35.2  22.8
0.89  0.70; 0.64  0.59
0.09  0.04; 0.13  0.07
15.3  12.1; 62.7  50.0
Potamogeton spp.,
filamentous algae

<10 km2
12 m
<40 days
Clay
11.5  8.8; 16.7  14.7
0.68  0.22; 0.32  0.28
0.11  0.06; 0.15  0.06
11.8  9.8; 19.7  19.0
Trapa natans,
Potamogeton spp.,
Elodea nuttallii
Bleak Alburnus alburnus,
White bream Blicca bjoerkna

Tench Tinca tinca,

Rudd Scardinius
erythrophthalmus, Pike
Esox lucius

<4 km2
1.53 m
>50 days
Organic
9.4  10.1; 21.3  23.5
0.77  0.64; 0.33  0.22
0.09  0.05; 0.13  0.05
10.10  6.7; 32.0  21.7
Nitellopsis obtusa,
Ceratophyllum demersum

Remote lakes

Navodaru et al., 2002

Coops et al., 1999;

Covaliov et al., 2003

Further details

Data from lake monitoring in 19961998, encompassing various numbers of lakes. Average  SD of suspended solids, total nitrogen, total phosphporus and chlorophyll a in spring (AprilJune)
and summer (JulySeptember), respectively, are shown. For aquatic vegetation and fish only a number of characteristic species are listed.

Fish

Size
Depth
Flushing rate
Soil type
Suspended solids mg/L
Total nitrogen mg/L
Total phosphorus mg/L
Chlorophyll a mg/L
Aquatic vegetation

Large lakes

Inflow lakes

Table I. Characteristics of lakes with differing connectivity (Inflow lakes, Large lakes and Remote lakes) in the Danube Delta

702
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submerged macrophytes, floating-leaved macrophytes, and turbid, vegetation-poor water. The larger lakes in the
central and marine part of the Delta were predominantly turbid, while the many small lakes and lakes in the fluvial,
upstream part of the delta were clear and macrophyte dominated.

Hydrological modelling
Water levels of the Danube vary from year to year, but there is a strong tendency of high water levels in MayJune
followed by steadily decreasing levels during summer and autumn (Figure 2). In the survey period, this pattern was
observed also although in 1997 an unusually high flood in August occurred. Average water-level fluctuations were
relatively high in lakes near the head of the delta (up to 1.68 m) and low in lakes near the mouth (0.69 m).
Although the shortage of validation data may have caused spurious results, indicative values were derived for
individual lakes of the travel time, hydraulic residence time and the cumulative residence time. Hydrological
characteristics of lakes varied extensively: average residence time between March and October could be as low as
5 days and as high as 300 days. Likewise, travel time ranged between 0.2 days for a lake immediately adjacent to the
river and 93 days for a highly isolated lake. Cumulative residence time ranged between 9 and 322 days.
Special attention was paid to the flow-path of water between the main riverbranches and lakes. A significant part
of the water flow into isolated lakes may be through reedbeds. Floating reedbeds are present over large areas of the
lake complexes, resulting in hydraulic connectivity of lakes even in periods without channel connections. In the
schematization frame for the delta, estimates were made of the contribution of the water flow under the floating
reedbeds to the water budget in the lakes.
Lake water levels depend on water discharge of the main river. Levels at the apex of the delta are shown in
Figure 2; generally there is a high water level in spring (May), followed by a strong decline during the summer
months. Lake water levels follow this pattern with a delay of maximally a few days, but fluctuations diminish
towards the Black Sea. In 1996, the river water level was above average in May and October and below in
JulyAugust, in 1997 an unusually high flood occurred in August, and in 1998 started dry in MayJune and had a
high water peak in October.
Ordination of lakes based on hydrological and soil characteristics revealed a major influence of substrate type
(first axis of PCA). There was a clear separation between the lakes classified as turbid and clear by remote
sensing (Figure 3).

Figure 3. PCA diagram of lakes in the Danube Delta. Lakes are ordinated according to hydromorphological variables. Curestm, Cumulative
residence time; Reedw, Reed water; Amplit, Yearly water-level amplitude
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Water chemistry, macrophytes and fish

The concentrations of phosphorus and nitrogen found in the lakes were generally similar, although nitrogen
tended to be higher in large lakes in summer. No clear patterns of hydrochemical data could be discerned in the July
19961998 data. However, there were clear patterns of ecosystem characteristics of lakes.
Vegetation occurrence was related to the gradient of sediment deposition. Silt deposition zones in inflow lakes
were indicated by vegetation of Trapa natans. Deep lagoonal lakes were characterized by narrow-leaved
Potamogeton species. Lakes on organic (peaty) sediments, enclosed by vast reedbeds showed dense vegetations
with Ceratophyllum demersum and Nitellopsis obtusa.
In vegetation-dominated, large lakes with mineral sediments, a mixture of limnophilic (Rudd Scardinius
erythrophthalmus, Pike Esox lucius) and eurytopic fish species (Perch Perca fluviatilis, Gibel carp Carassius
gibelio, Roach Rutilus rutilus) were found. Blackfish species (Weatherfish Misgurnus fossilis, Mudminnow
Umbra krameri, Tench Tinca tinca, Crucian carp Carassius carassius) were predominantly found in the lakes
enclosed by reedbeds.
Lake typology
Principal Components Analysis revealed a major gradient determined by lake size and depth, and turbidity.
Smaller, shallower lakes tended to have clear water and were characterized by low concentrations of suspended
matter, high submerged vegetation cover and filamentous algae, whereas larger, deeper lakes tended to be turbid,
characterized by high densities of phyto- and zooplankton and dominance of cyanobacteria. The second axis of the
PCA was related to hydraulic characteristics and the type of vegetation and fish community: lakes with a short
residence time that had a Potamogeton-dominated vegetation with a high proportion of eurytopic fish versus
isolated lakes with a Nitellopsis-vegetation and a Blackfish-type community. Based on the multivariate analysis,
three lake types may be discerned (Figure 4): (1) Large lakes, (2) Inflow lakes and (3) Remote lakes.
Environmental characteristics of the three lake types are listed in Table I.
Seasonality
Compared to values measured in the river channel itself, the seasonal variation of phosphorus was more
pronounced in lakes: in spring, total P concentrations were 0.070.11 mg L
1 (dissolved P: 0.020.04 mg L
1),

Figure 4. PCA ordination of surveyed lakes in the Danube Delta based on biotic variables: EUF, percentage eurytopic fish; BLF, percentage
Blackfish; NIT, percentage cover Nitellopsis; AQV, dominance of aquatic vegetation; POT, percentage cover Potamogeton; CLS, presence of
clear water/low suspended solids; FIL, percentage cover filamentous algae; CLA, cladocerans; ZPL, zooplankton; CYA, cyanobacteria; PPL,
phytoplankton. Clusters: 1, Large lakes; 2, Inflow lakes; 3, Remote lakes
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while during the summer months, total and dissolved P concentrations increased in all lakes to values of
0.130.20 mg L
1. In several lakes total nitrogen concentrations showed a considerable increase from spring to
summer; NOx concentrations in spring were 0.070.25 mg L
1, and in summer 0.241.54 mg L
1, and were almost

1
always higher than NH
in spring to 0.080.54 mg L
1 in summer). Chlorophyll
4 concentrations (0.020.09 mg L
concentrations increased in summer strongly in the large lakes, whereas increases were highly variable in remote
lakes and lowest in inflow lakes. Water transparency showed a distinctly seasonal pattern. In spring, the Secchi
depth was generally >1 m in all sampled lakes. The Secchi depth remained high during the summer in some lakes
(inflow lakes and most remote lakes), but decreased to 0.5 m or less in others (particularly large lakes; Figure 5).
Suspended solids and phytoplankton contributed in approximately equal amount to the turbidity of the sampled
lakes.
Phytoplankton composition was in general dominated by diatoms (especially in spring, generally the larger
taxa). Cyanobacteria increased after June (in summer mostly Planktothrix). Blooms of cyanobacteria and diatoms
are not mutually exclusive. No distinct spring bloom of zooplankton was observed.
Comparison between early and late summer vegetation showed a distinct seasonality of the vegetation in the
large lakes: these lakes were almost entirely covered by Potamogeton spp. vegetation in June, and devoid of
macrophytes in late summer. Concomitantly a strong decrease in transparency had occurred with the development
of algal blooms. In the other two lake types (inflow, remote) the submerged vegetation was much more stable
throughout spring and summer.
Limnophilic and eurytopic fish species were present over the entire gradient, but abundance and dominance
varied. Distribution patterns in June and September suggested opposite patterns of seasonal migration, depending
on preferred habitat for spawning (June) and feeding (September). For instance, in June Bleak Alburnus alburnus
occurred predominantly at close distance from the river, but in September it had moved into the large lakes which
were then devoid of vegetation. In contrast Rudd S. erythrophthalmus was present in remote lakes in June, whereas
its occurrence shifted towards inflow lakes in September.

DISCUSSION
The Danube Delta forms a large and complex hydrological system through which water flows in three
riverbranches, hundreds of lakes, connecting channels, under floating reedbeds and, during high river stages, over
levees and reedbed surfaces. We consider it inappropriate to examine the hydrology of individual lakes within the
larger complexes because of the temporally and spatially variable interconnections within the system. We used a
rather crude schematization of the dimensions of the channellake network in the delta to assess water-level
fluctuation, retention time and flow velocities of water. Even with a limited accuracy of the model results, the
model of the delta allowed us to position lakes in a gradient of hydrological connectivity, expressed by water age
(travel time, residence time) and influence of reedbeds.
Several components of hydrological connectivity can be distinguished in riverine systems (Tockner et al., 1999;
Amoros and Bornette, 2002). Within the Danube Delta, the river shows a lateral connectivity gradient of decreasing
exchange between lake water and river water. Such gradients of chain-set lakes exhibit differences in the average
age of water, or residence time, that may determine state transitions (Lesack et al., 1998; Aspetsberger et al., 2002;
Squires and Lesack, 2003). On the other hand, there is an aspect of temporal connectivity promoting the
seasonal changes in lake complexes, steered by the water discharge pattern of the Danube River, though
hydrological connections will be maintained throughout the year since even at low river discharge flow into the
wetland complexes occurs; however, the flow of water through the system would be much higher during high than
during low river stages.
The impact of hydrological connectivity may vary between different complexes of the delta. The most upstream
lakes in the fluvial part of the delta exhibit a greater water-level fluctuation than the lake complex close to the Black
Sea (the so-called marine part). Apart from these hydrodynamic gradients, morphological characteristics vary
within and between complexes, due to diagenetic and sedimentary differences between lakes. The typology
presented here reflects these differences: highly connected inflow lakes show high concentrations of mineral
suspended matter, a high sedimentation rate and a rather shallow, mineral bottom substrate. The large lakes with an
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Figure 5. Seasonal course of transparency (Secchi Disk measurements) of three lakes in the central Danube Delta representing inflow (lake
Uzlina), large (lake Isac) and remote (lake Cuibul cu Lebede) during 19971998

intermediate connectivity usually are the larger, relatively deep lagoonal lakes. The remote lakes, enclosed by
reedbeds, are mostly smaller sized, isolated, shallow lakes with highly organic sediments.
In several studies, nutrient levels have been shown to decrease towards the isolated end of the connectivity
gradient (Van den Brink et al., 1993; Heiler et al., 1995), which is correlated with an increase in transparency of the
water (Squires and Lesack, 2003). Phosphorus concentrations in the lakes were generally lower than in the river. On
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the one hand, nutrients imported with suspended particles may sediment directly after entering a lake complex, on
the other hand, large quantities may be immobilized in algal and plant biomass. The high nutrient loading of the
lakes and rising temperatures in springtime promote massive macrophyte growth, especially in the large, highly to
intermediately connected lakes. Because of the coincident high river stage, water retention time in these lakes may
be too short to support strong phytoplankton development; due to the severity of winters the spring inoculum may
be small as well. Also, the dense macrophyte beds may prevent algal growth. Although a clear spring peak of
zooplankton grazers was not observed, facilitation of grazers within the vegetation may be an important mechanism
by which phytoplankton development is suppressed (Muylaert et al., 2006). The absence of a distinct zooplankton
development may be due to short residence times in spring resulting in washing out of the large cladoceran
population (Baranyi et al., 2002), or a high predation by planktivorous fish (Jeppesen et al., 2000).
Additionally, resuspension of sedimented algae may not occur within densely vegetated lakes, resulting in a
severe loss factor for algae. In some cases, phosphorus limitation may have occurred. Flushing of (some of the)
lakes during the flood pulse in spring may have contributed to low phytoplankton biomass in spring. Other
mechanisms, that is the exhaustion of nutrients by the growing macrophytes, would be less likely as we found no
potentially growth-limiting reductions in lake phosphorus and nitrogen concentrations.
The distinct increase in dissolved phosphorus observed in summer may result from mobilization due to decay of
macrophytes, and the release from reedbeds surrounding the lakes and lake-bottom sediments. In the large lakes,
filamentous algae started to overgrow the fully developed submersed vegetation in early summer. The release of
nutrients from macrophytes has been shown to be primarily dependent on tissue decay, rather than excretory
processes (Barko and Smart, 1980). The concurrently developing phytoplankton bloom may be stimulated by
nutrient mobilization. In the larger and deeper lakes of the Danube Delta, phytoplankton generally forms the
dominant contributor to underwater light extinction (Cristofor et al., 1994).
Reedbeds surrounding the remote lakes may be an important source of dissolved nutrients in summer, in
particular in those cases where the surface becomes exposed (Baldwin and Mitchell, 2000; Biddanda and Cotner,
2002), for example when floating reedbeds around the more shallow lakes settle on the mineral bottom. On the other
hand, the beds of aquatic vegetation and reed may act as extended sites for denitrification, especially in summer
when the temperatures are high. In this way nitrogen may be lost from the system in summer (Hilbricht-Ilkowska,
1999). However, in some lakes total nitrogen concentrations showed a considerable increase in summer relative to
spring; since NOx and NH
4 did not show a coincident increase, it is concluded that the particulate N must come
from an internal source of N, for example N-fixing cyanobacteria or release from decomposing aquatic vegetation,
followed by rapid incorporation in the biomass of the growing phytoplankton population.
Contrary to the inflow and large lakes (high intermediate connectivity), phytoplankton biomass remained low
throughout the summer in the remote lakes, while a high aquatic vegetation cover (characterized by extensive
charophyte beds) persisted (Covaliov et al., 2003). In the latter group of lakes, a long residence time in combination
with the water exchange between reedbed and lake may have produced high concentrations of dissolved organic
carbon. High amounts of humic substances may potentially inhibit phytoplankton growth due to either a direct toxic
effect or by the strong alteration of light attenuation; in contrast, the availability of nutrients may be enhanced,
resulting in an opposite effect on phytoplankton growth (Jones, 1992). The scarcity of algal blooms in remote lakes,
and the similarity of phytoplankton composition among the lake types allows no clear conclusion regarding the
impact of humic substances in the system.
The water in the remote lakes had a brown colour (also referred to as blackwater) and often showed signs of
sulphide production, indicating anoxic conditions in the sediment. The decomposition of organic material will be
incomplete, resulting in oxygen depletion. Daily oxygen fluctuations in plant beds may be large as well, resulting in
an adverse environment for fish, that may rise to the surface and either remain there or use it as a corridor to travel to
oxygenated pockets (Miranda et al., 2000). In the Danube Delta, all lakes are interconnected and fish may migrate
within the system to sites where conditions are most suitable. Especially in late summer, the fish present in reed
lakes consisted for a large part of Blackfish species that tolerate low oxygen conditions.
The lake complexes form an open system in which fish are distributed according to their habitat preferences and
competitive interactions. Fishes may migrate freely both from and to the river, and to and from the remote
blackwater lakes. The generally declining water levels in early summer (June) may be simultaneous with the
spawning period of most fish species. The presence of dense vegetation in the same period would be advantageous
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708

H. COOPS ET AL.

for certain fish species that are adapted to low oxygen conditions. Furthermore, intense daily migrations of fish may
take place within the system (Navodaru et al., 2002).
The intense interactions between river, lake and reedbeds in the Danube Delta appear to play a key steering role
for the water quality and the food web of the lakes. Lakewater flowing through or below the reedbeds becomes less
nutrient rich owing to nutrient extraction, and is loaded with humic substances before it is released again into the
lakes and channels. Hence the annual flood cycle may be the primary driving force of the ecosystem of the delta and
the changes in water quality along the gradient of lake types.
Over the past 50 years, inputs of river water into the lakereedbed complexes have been greatly altered by
digging channels to improve the accessibility and the fish production in the area. The impacts of this opening-up of
the delta were aggravated by the severe eutrophication and pollution of the Danube River since the 1970s. The
estimated water inflow into floodplain enclosed by the major riverbranches increased from 160 m3 s
1 in 1920 to
620 m3 s
1 currently. In the same period the annual loading with sediments increased from 1.3  106 to 2.5  106
tonnes, the annual total-P loading from 0.2  103 to 3.2  103 tonnes and the annual total-N loading from 4  103 to
59  103 tonnes (Oosterberg et al., 2000). Furthermore, significant parts of the floodplain (312 km2, c. 10% of the
delta) have been embanked in the past 50 years. Both the historical trend to open up the delta complexes by cutting
channels and the polder construction reducing the floodplain area, as the restoration measures taken so far can to
some extent be regarded as large-scale hydrological experiments as they show the effect of this hydrological
interference. It is now recognized that the pathway of rehabilitation should first and foremost include hydrological
measures that restore the natural flow of water through the system (Gore and Shields, 1995; Schmidt, 2001; Buijse
et al., 2002).
The findings could be of use in other lowland river systems throughout Europe, since current management
according to the EU Water Framework Directive (WFD)is targeted at restoring natural conditions, as far as
possible in the context of man-made changes. To some degree the Danube Delta acts as a natural reference site,
highlighting the importance of connectivity at a landscape scale. Although our studies started conducted prior to the
WFD coming into force (EU, 2000), its multidisciplinary approach based on hydrological modelling, satellite
images and monitoring data of water quality, plankton, aquatic vegetation and fish communities turned out to be an
extremely useful tool to develop a typology for the hundreds of lakes in the delta and to classify their present
ecological status. Another main achievement of our study is the integrated use of various sources of spatial and
temporal data in a complex riverine lake system. The results facilitate the design of a monitoring programme, and
may also serve to identify the need and the potential for restoration.

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DOI: 10.1002/rra