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LIMNOLOGY
in the Federal Republic of Germany

Edited by
Winfried Lampert und Karl-Otto Rothhaupt

for the 24th Congress of the International Association of

Theoretical and Applied Limnology, August 13-19, 1989, at Munich
LAKE CONSTANCE -
the largest German lake
W. Geller and H. Güde (eds.) *)

General limnological characterisation Morphometric data

Origin of the lake
Lake Constance is the second largest( to Lake Geneva)
European pre-alpine lake both by area and volume, and
the largest lake in Germany. lt is divided into two parts, the
deep "upper lake" and the shallow "lower lake" (Table 1).
The lake basin was formed by the erosive forces of the
Rhine glacier which excavated a 500 m deep valley during
its last extension, about 30,000 years b.p„ At the end of
this glaciation, 15,000 years ago, the lake area was more
than twice its present cover, including large parts of the
Rhine valley and both Lake Walen and Lake Zürich.
Fig. 1 Lake Constance and its catchment area (F. Kiefer).
The morphometric data of Lake Constance are presented
separately for the lower and the upper lake, in Table 1. The
upper lake is Lake Constance "sensu strictu". The catchment
area is 20 times the lake area, reaching to the Alpine ritt in
Switzerland, Austria and ltaly (Fig. 1). The annual water
budget is about 12 km3 , resulting in a water residence time
of 4 to 5 years. The main tributary is the "Alpenrhein"
contributing 75 % of the total inflow. Both water budget
and lake level reflect the seasonality of the Alps which
retain snow during the winter. The lake level is therefore
maximum in early summer and minimum during the winter.
Table 1 Morphometric Data of Lake Constance
( 47° 39'N 9°18'E)

Lower Lake Upper Lake

Altitude a.s.l.·) (m) 395.1 395.3

Area (km2 ) 63 476
Catchment area (km2 ) 10819
Shoreline (km) 90 165
Max. depth (m) 46 252
Mean depth (m) 13.2 100
Volume (km3 ) 0.83 47.7
Annual outflow (km3) (11.5) 11.5
Residence time (a) (0.072) 4.15
�;
(. *) German sea level referred to North Sea; + 32 cm =
Swiss s.l. referred to Mediterranean Sea (Marseille);+ 25
cm = Austrian s.l. (Adria).

km '--/
0 10 2.0 JO •O 50 60
Thermic behaviour

Lake Constance is the only warm-monomictic lake of con­

tinental Europe north of the alpine chain, with its holomixis
in late February or March, at a temperature of 4° C. During
the summer the maximum temperature may reach 22° to
23° C. The water temperatures follow air temperatures
with a delay of about one month, so that the seasonal
course of lake temperatures 'follows a "hysteresis curve"
(Fig. 2).

As in Lake Geneva or the lnsubric lakes of Northern ltaly,

mixing of the water column does not penetrate to maximum
depth in Lake Constance during winters with extraordinarily

·) Partitial contributions by statt members cf Limnological Institute, Con­

stance, and Institute for Lake Research and Fisheries, Langenargen

9
low wind speed. Usually, only the shallow lower lake has can easily be identified by its turbidity and low ionic
ice cover during winter. The upper lake, freezes only a few concentrations when it finds its path along the northern
times percentury, peculiar events which have been reported shore. lts branches form a main gyre in the lake centre and
for about eleven hundred years. The frequences of complete smaller gyres in the bight of Bregenz, east of the Rhine
ice cover reflect the climatic changes within the past inflow. A second gyre occurs in the "Überlinger Lake", the
millenium: fjordlike north-western bight of the upper lake.
875 895
1074 1076
1108 Seiches and water currents
1217 1227 1277
1323 1325 1378 1379 ·1383 Gyral surface currents, caused by wind forces and differ­
1409 1431 1435 1460 1465 1470 1479 ential air pressures, have been observed and are com­
1512 1553 1560 1564 1565 1571 1573 pensated by reverse flow of deeper water. With a north­
1684 1695 east wind the warm, epilimnetic water is directed towards
1788 the southern shore by gyral currents and cold water from
1830 1880 the hypolimnion upwells atthe northern shore. The resulting
1963 surface waves have been observed as seiches with
frequences of 55 and 39 minutes, 1st and 2nd order
oc Überlinger See 1977
oscillations of the lake's long axis. Oscillations in the short
20 Temperoture of Air A
axis of the 2 km wide Überlinger bight have durations of
ond Lake Surfoce Woter/0m only a few minutes. Interna! waves, between the epi- and
hypolimnic layers, attain amplitudes of more than 20 m.

15
Water chemistry

lonic composition basically corresponds to Rodhe's

"standard fresh water" with calcium and bicarbonate as
10 main components and a conductivity of 270 to 290 µS/cm
(Table 2). Compared with its natural, oligotrophic state,
retained until the beginning of this century, P and N levels,
and also chloride from road salt widely used in winter,
5 have increased anthropogenically.

Table 2 Chemical analysis of L. Constance water

D
0 J Conductivity 270-290 µS/cm
7 8 9 70 11 12 13 14 15 16 17 Dissolved solids 190 mg/L
Doy length [h] pH 7.8

Fig. 2 Monthly means of air and lake temperature versus Carbonate alkalinity 2.45 meq/L
daylength, as measured at the Überlinger See 1977 (W. Sulfate 0.71 meq/L
Geiler). Nitrate 0.056 meq/L
Chloride 0.13 meq/L
Flow of the Rhine through the lake Anions 3.35 meq/L

lnflow of the Alpenrhein into the lake was "corrected" to its Calcium 2.50 meq/L
present course by an artificial canal in 1900. In August the Magnesium 0.70 meq/L
Rhine water may be 10° C cooler than the lake surface Sodium 0.17 meq/L
waters resulting in downward flow of river water within the Potassium 0.03 meq/L
lake until it finds a layer of equal density in the thermocline Cations 3.40 meq/L
(Fig. 3). The course of the Rhine water through the lake

Fig. 3 Seasonal development of the thermocline in Lake Long-term development of the trophic state
Constance and of the depth layer of through-flowing
Alpenrhein water. Ranges and means of depth of 1st and In its natural state, L. Constance was a typical oligo­
2nd order thermocline 1963-1976 (H. Lehn). trophic pre-alpine lake with low concentrations of nu­
trients (total P 10 mg/m3, total N 1000 mg/m3), low
0 r-.---..;;i;::::==�;;::=:::r:=�=:-,--r---r---.----,-7
1 s1 rh.
·-·•:-•-•,-::•:❖:❖:-:-.--. densities of phytoplankton, high water transparency and
m ,-,:::::::•:-:,,,...... high hypolimnic oxygen concentrations. With an increase
I
I
in human population and fundamental changes in the eco­
20 I nomy and human social behaviour, the trophic state of the
1
'I lake was changed over the past four decades. At present,
the 1.5 million people living in the catchment area, local
'
1

L.O
1
industry and agriculture together discharge sewage water
'' equivalent to 3.2 million inhabitants. Until the early 1970s,
'1 the major part of this sewage entered the lake without any

2ndTh '
treatment, resulting in a strong increase of the P-load,

''
1
60 additionally enhanced by increasing inputs from agricultural
sources (fertilization) and from precipitation.
'
''
1
The total nutrient input from the tributaries reflects the
80
runoff conditions of particular years rather than the long­
s
1

J F M A M J J A 0 N D term charges in sewage loading (Table 3). lt is mainly due

10
to the nutrients bound to suspended solids of rivers, which to the response time required for the lake to reach a new
increase exponentially with run-off and are almost all equilibrium alter changes in P-loading. Thus, initially, the
deposited in the lake sediments. Apart from the bound process of eutrophication proceeded further to a maximum
fraction, however, the nutrient input correlates weil with in the late 1970s. However, since the beginning of the
the development of nutrient concentrations in the lake 1980s, the effectiveness of the measures against nutrient
(Fig. 4, Table 3). This suggests that the long-term trophic loading has become apparent:
trend results mainly from loading with ecologically effective
dissolved nutrients. This reached a maximum in the early - A strong decrease in the phosphorus load in sewage
1970s in the case of phosphorus, but continues to increase treatment plant inflows and outflows.
for dissolved nitrogen. - A significant decrease in the phosphorus (without
suspended solids) load in the inflow streams (Table 3).
- A reduction in the concentration of PO4 -P during overturn,
Table 3 Comparison of loading of Lake Constance by its from a maximum of 87 mg/m3 in 1979 to 47 mg/m3 in
inflow for years (1965/66 extremely high, 1971/72 extremely 1987 (Fig. 4).
low run-off) - Hypolimnic oxygen concentrations always >5 mg/L.
65/66 67/68 71/72 78/79 85/86 At present, 75% of the sewage phosphorus is chemically
precipitated in treatment plants. Further reduction of the
Run-oft (km3 ) 14.7 12.5 7.4 11.0 11.5 P-load is expected after completion ofthe sewage purification
Susp. solids programme including the use of combined flocculation
in rivers (Mio t) 4.7 1.1 2.6 2.6 and filtration in large treatment plants. Depending on the
Posphorus (103 t) remaining P-load, new lower equilibrium concentrations
total >3.1 3.9 2.1 3.0 2.3 will be reached in the lake. Thus, this provides an example
without susp.sol. ca 1.1 1.3 1 .1 0.6 of a successful restoration programme beyond national
Nitrogen (103 t) borders.
total 16.6 12.0 19.2 21.0
without susp.soL 13 10.4 14.5 18.0

Structure and functioning of the pelagic ecosystem

3
mgP/m Phytoplankton and primary productivity
100

Long-term development of phytoplankton community struc­

80
ture: Since the earliest records in 1896 until the 1930s
phytoplankton development in Lake Constance indicated
an oligotrophic state, characterized by low algal densities
60
with one or two annual maxima predominantly of centric
and pennate diatoms (Table 4). The first indications of
eutrophication appeared in the 1950s: Algal densities
40
increased and showed two regular maxima per year.
Simultaneously, the community structure of the
phytoplankton changed significantly: (1) The previously
20
important group of Cyclotella spp. disappeared and (2)
other algal groups such as cryptophyceans, or the diatom
Tabellaria fenestratabecame predominant. With accelerating
0 eutrophication in the 1960s phytoplankton density further
1950 1960 1970 1980 1990
increased, exhibiting three to four maxima per year. The
strongest increase was observed for the spring bloom
_
Fig. 4 Long-term development of the PO4 -P concentration
at holomixis in Lake Constance. Table 4 List of dominant species observed in Lake
Constance at different trophic states. (v), (e), (a) = vernal,
The response of the lake to the increased P-loading was estival, and autumnal species, respectively.
already apparent in the 1950s from increased plankton
densities, composition changes and enhanced oxygen prior 1950- 1965- 1987/
deficiencies in the hypolimnic waters. Faced with these 1950 65 80 88
and other threats to the ecological balance of the lake,the
International Commission for the Protection of Lake Cyclotella spp. e
Constance (IGKB) was founded in 1959 by the adjacent Fragilaria crotonensis v v,e e e
states. After intensive discussions on the causes of, and Stephanodiscus
possible measures against, eutrophication, the commission hantzschii v V V
decided to reduce P-loading of the lake by intensive Stephanodiscus
sewage purification measures including chemical P­ astraea V V
precipitation over the entire catchment area. Over the Stephanodiscus
past years, partial or complete substitution of phosphorus binderanus e
in detergents and more economic use of washing pow­ Asterionella formosa v v e V
ders due to increasing environmental consciousness in Synedra acus var.ang. v
the public, have achieved additional reductions in P­ Tabellaria fenestrata e e
loading. Rhodomonas minuta v V V
Cryptomonas ovata e,a V
These decisions were not followed by immediate effects, Mougeotia thylespora e,a e
because the realization of this programme needed con­ Ceratium hirundinella e
siderable time and money (4 billions of Swiss Francs, Anabaena planctonica e,a
about 3 billions of US $ by 1985). Further delays are due

11
which was dominated by Stephanodiscus hantzschii and
Rhodomonas minuta. From 1963 on the occurrence of a
clear-water phase, caused by intensive zooplankton grazing,
was also observed. During summer and autumn,
phytoplankton showed an increased portion of chloro­
phyceans and blue-green algae. Recent evidence suggests
that autotrophic picoplankton (<2 µm) are an additional
important component of the phytoplankton community.
However, there is no corresponding information on the
long-term development of this group in the lake.
Up to now, the remarkable decline of P-concentration
alter 1981 (Fig. 4) has caused no significant decrease of
phytoplankton biomass. However, the return to a more
oligotrophic state is indicated by some conspicuous changes
in plankton community composition: (1) the reappearance
of Cyclotella species, (2) the shift in the occurence of
Asterionella formosa toward the spring bloom, (3) the
return of Tabellaria fenestrata in summer, and (4) the
decline of filamentous diatoms and blue-green algae.
Light and nutrients
During the time between 1979 and 1984, Secchi disc
transparency varied between 1 m and 14 m, and euphotic
depth (1%-surface light level) between 5 and 22 m (Fig. 6).
The ratio of Secchi depth to euphotic depth was ca. 1.5
under clear-water conditions but increased to >5 with
turbidity due to high phytoplankton abundance. About
80% of the seasonal variation of the vertical light attenua­
tion coefficient can be explained by changes in euphotic
phytoplankton abundance.
Phosphorus is the single most important nutrient for
plankton growth in L. Constance, but only a part of the total
P is incorporated into phytoplankton biomass, the rest can
Fig. 5 Seasonal succession of phytoplankton community composition in Lake Constance according to the relative
strength of varying environmental factors: light, lake stratification, nutrients and zooplankton grazing (U. Sommer).
winter spring clear
b loom wa ter h.
surface light
day length &
____________________, -----
stratification NO �--------------------,_
M I X I N G DE PT H O - 3 m
..____
100
SRP
1000
SR Si
grazing
dominant
a lgae
be attributed to detritus, zooplankton and bacteria. During
short periods in July and August, C:P ratios of algae
indicate P-limitation. Significant portions (ca. 18 % of
PPR) are lost by sedimentation from the euphotic zone
both by incorporated P and particle-adsorbed P. For short
periods when diatoms dominate the phytoplankton, silicon
can be growth limiting.
Seasonal change of the phytoplankton community composi­
tion: From 1979 to 1983 the pattern of phytoplankton
species composition was explained by both environmental
factors and intracommunity interactions (Fig. 5). Seven
successional stages can be distinguished. The onset of
the spring bloom is triggered by establishment of thermal
stratification. During the spring bloom small cryptophyceans
and small centric diatoms (Table 4) dominate, and nutrient
concentrations, especially SRP, decrease rapidly.
Overgrazing by rapidly reproducing Daphnia in early June
decimate the phytoplankton within a few days, thus causing
a clearwater phase. With the resulting nutrient regeneration,
a summer assemblage of slower growing algae builds up
which is less susceptible to grazing. As thermal stability
increases and phosphorus is declining, large diatoms
(Asterionella, Fragilaria, Stephanodiscus binderanus)
outcompete other algal groups as long as silicon is present.
After decrease in silicon large dinoflagellates (Ceratium)
dominate, followed by filamentous cyanobacteria and
diatoms (Anabaena, Aphanizomenon, Diatoma), and then
filamentous green algae (Mougeotia, Ulothrix). In late
autumn, the deeper mixing leads to the recurrence of
large diatoms.
The seasonal cycle of primary productivity
The light requirements of the phytoplankton community
change seasonally. During winter and early spring, phyto-
su mmer september autumn
bloom m1n1mum
ZMIX INCREASING
1 1
 120
CILIATES
,"O 40
,_
90
N
'E 30 a 60
z
E 20 30

f 10 0
H N F
0 LLL.L.L..L.L.L.L..L.LL...L..L..L..l....L..1....L.L......_.....L..1...L..1...L..1....1....1L....L..J'--'-L...L......,
\i
N
45

:: 30
M
Z 15

BACTERIA
+
,_
"8
0/
.c 10
u
� :::
10 O> M
5
E z
30

0
15
F M A M A 0 N D
20

20
Fig. 7 Seasonal course of microbial abundance: bacteria,
heterotrophic nannoflagellates (HNF), and ciliates.

25 Much information exists on the bakterioplankton although

its specific composition is still very poorly known. However,
for trophic interactions, heterotrophic bacteria may be
regarded as a uniform assemblage consisting of more
Fig. 6 Seasonal course of primary production, Secchi than 90% of freely suspended single cells. Heterotrophie
depth, euphotic depth, and chlorophyll content in Lake bacterial biomass and production change throughout the
Constance. growing season, closely following primary production and
phytoplankton biomass which provide the major substrate
sources. Fluctuations in bacterial abundance and activity
plankton are able to efficiently utilize low irradiances, and are restricted to the water layers near the euphotic zone.
during the summer months light-saturated photosynthetic In the hypolimnion, numbers and production remain constant
rates are high. Therefore, seasonal variations in light and low (106 cells/ml, 1-3 µg C L 1 d 1). Highest bacterial
intensities can be compensated for by changing light numbers and production occur during the phytoplankton
requirements, and apparent variations in production rates spring bloom, reaching 107 /ml (Fig. 7) and 20-50 µgC
are mainly due to changing proportions of underwater light L-1 d-1, with population doubling times of 10-30 h. Thereafter,
harvesting by the photosynthetic pigments and variations bacterial abundance and production decline strongly (clear­
in day-length. Lake Constance phytoplankton are water phase). Bacterial numbers and production usually
characterized by unusually wide seasonal variations in rise again in July and fluctuate at higher levels until the
both biomass and daily production rates (Fig. 6). They can breakdown of the thermal stratification in the autumn.
be explained as follows: (1) The daily radiant energy input
varies widely because of the combined effects of fluctuations Bacteria comprise a major portion of the total biomass in
in daily surface irradiance and mixing depth. During deep L. Constance, equalling phytoplankton biomass on an
winter mixing, the production process is limited by energy annual mean (Fig. 8). Even when phytoplankton biomass
supply leading to extremely low biomass and productivity. is high, bacterial biomass accounts for 25-50% of total
(2) During summer, biomass accumulation is high, even
though the carrying capacity predicted from winter nutrient Fig. 8 Seasonal course of epilimnetic particulate organic
levels is never reached. The measured production rates carbon, algal and bacterial biomass, measured as cell
are indicators of phytoplankton growth potential rather carbon.
than of actual biomass growth. This is because apparent
changes in biomass are due to shifts in the balance of
growth and losses, and only in species that are subject to 16 ·· - ---POC
little loss can changes in population densities be directly
related to in situ growth rates. The measured annual --algal biomass
cumulative primary productivity varied between 270 to ----bact. biomass
330 g C m·2 a-1 during 1980 and 1983. 12
N
'E
u
Heterotrophie microbial community
cn 8
Over recent years a new concept of the aquatic food web
has evolved, according to which heterotrophic microor­
ganisms contrib�te to a large extent in the flow of matter 4
through the planktonic community ("microbial loop"). As
this was mainly a result of the development of new
methods (especially the use of epifluorescence micro­
scopy), we only started to learn about the significance of
the heterotrophic microbial community in L. Constance M A M J J A S O N D J F M A M J
since the early 1980s. 1982 1983

13
biomass. Due to this high biomass and to low C:P ratios, Table 5 Species of the crustacean plankton community
bacteria also bind high amounts of phosphorus often ex­ before and alter eutrophication of Lake Constance.
ceeding 50% of the total phosphorus in the euphotic zone
in summer. Bacterial production is estimated as c. 60% of pre 1940s 1950s-1980s
primary production on an annual mean.
Diaphanosoma brachyurum
The taxonomically heterogeneous group of heterotrophic (-1957)
nannoflagellates (HNF) is an important consumer of bac­ Daphnia hyalina Daphnia hyalina
teria and of autotrophic picoplankton (<2µm). lts abun­ Daphnia galeata
dance varies between 102/ml and 104 /ml. Highest numbers Bosmina longispina Bosmina longispina
occur above the thermocline in spring and summer. In the Bosmina mixta kessleri
hypolimnion, abundances remain at the low winter levels, Bosmina longirostris
also true for ciliates. Whereas HNF abundance is domina­ Bythotrephes longimanus Bythotrephes /ongimanus
ted by cells <2µm, organisms 2-10 µm in size contribute Leptodora kindtii Leptodora kindtii
the major part of the HNF biomass. On an annual mean, Heterocope borealis (-1963)
50% bacterial production is estimated to be removed by Eudiaptomus gracilis Eudiaptomus gracilis
HNF. The grazing impact of HNF on bacteria is highest in Acanthodiaptomus
late spring when bacterial production is almost com­ denticornis
pletely consumed. During the subsequent clear-water Cyclops abyssorum Cyclops abyssorum
phase, the bacterial and HNF abundances are mainly con­ Cyclops vicinus
trolled by grazing daphnids. Thereafter, HNF numbers
rise again to higher, but rather fluctuating levels until the
winter decline. During the cold months, the copepods dominate by numbers
with Cyclops vicinus being the most important single
Of the heterotrophic microorganisms, the ciliates are the species, but at onset of the growing season in May,
least studied; regular monitoring of their abundance, dominance shifts to the cladocerans (Fig. 9). During the
biovolume and species composition has only been carried growing season, the two Daphnia species are the most
out since 1986. Their abundance varies from 1 to 100 important consumers of the phytoplankton. Crustacean
cells/ml with two distinct maxima in spring and in summer. biomass maximum is reached in June, with ca. 15 g DW/
Estimates of biovolumes range between 10 and 1000 m2 , thereby surpassing the phytoplankton in early summer.
mm3/m3 . The ciliate community is numerically dominated The mean standing stock throughout the growing season
by the orders Protostomatida and Oligotrichida, species is about 5 g DW/m2 .
known to ingest food of the size of small algae. To date
little is known on the grazing impact of ciliates on bacteria. The migratory behaviour of the different crustaceans is
shown in Fig. 10, including a representative scenario for
At present, the quality and quantity of the trophic relationships the time from early summer to autumn: Daphnia galeata
within the heterotrophic microbes, and to other components lives in the warm epilimnion, the bosminids live in the
of the planktonic community of L. Constance can only be deeper, cold layers of the hypolimnion, while D. hyalina
roughly evaluated. However existing evidence is sufficient migrates between the epilimnion (night) and deep water
to conclude that they play a major role in the planktonic (day). lts specific daytime position is defined by a narrow
theatre of this lake, mainly in the regeneration of nutrients band around the 5.7 ° C isotherm. The copepods show a
required for primary production and in energy transfer to similar migratory behaviour, but with differing migration
higher trophic levels which would otherwise be lost. amplitudes.

The zooplankton community and its trophic role
Crustaceans
The crustacean community of Lake Constance consists of
copepods and cladocerans. Species composition changed
from the oligotrophic lake until the 1940s to the present,
eutrophic state (Table 5).
Fig. 9 Seasonal course of community composition of
planktonic crustaceans (% of total numbers of indivi­
duals): cladocerans (Cl), Cyclops vicinus (Cv) and other
copepods (Cp), excluding naupliar stages.
100
¾

Species composition and population dynamics of the

planktonic crustaceans have been monitored since 1962.
This included taxonomic distinction of the copepodite Cp
stages to the species level, regular population census
over the year, and observation of generation cycles,
diapause and reproductive stages. By annual averages,
the total number of crustaceans did not change very much
over the lang term, but irregular variability between years 50
was observed with minima of about 0.6·106 , and maxima
of 1.5·106 crustaceans per m2 . The species showe.d different
long-term development. The abundances of the cladocerans
and C. vicinus followed the biomass trend of the
phytoplankton, whereas C. abyssorum declined
continuously, and Mesocyclops showed a long-term trend
to a minimum in 1979, against phosphorus and phytoplankton
Cv
trend. The recently immigrated Daphnia ''galeata"showed
great morphological and genetic variability. The highest
population densities of planktonic crustaceans were found Cl
Oo/0
xn
in the Bregenz bay at the eastern end of the lake, especially
of those species which appeared alter 1950. VI IX
14
 20 July 1982 ;; 8
1 14

"/1/lf'/
0 2 4 6 8 10 12 14 16 18 20 22 24 h
0 E
X 7
i::. 35 26
6
,._
Cf)

Q)
.D
5
E
::::J
_ 20
E Bosmina coregoni
C 4
CJ)
..2
3
Bosmina longirostris
J F M A M J J A s 0 N D

40
Fig. 11 Seasonal succession of rotifers in Lake Con­
stance, selectively shown by their specific phases of
maximum growth of population. Initial and final density of
50'----'--------_L_-'---------'--L----'----------'------------'-----'---------'--L----'--------------' the exponentially growing populations are connected by
straight lines.
Fig. 1 O Diel migratory cycle of the crustacean plankton 1: Ascomorpha ecaudis 2: Ascomorpha ovalis
with their vertical migration, shown by the populations' 4: Asplancha priodonta 14: Conochilus unicornis
centres of density in the water column. 17: Gastropus stylifer 18: Kellicottia longispina
19: Keratella cochlearis 23: Lepadel/a patella
The different migratory behaviour leads to turnover times 26. Notholca caudata 35: Pompholyx sulcata
(B/P) which are 2.5 times langer in 0. hyalina than in D. 37: Synchaeta pectinata 43: Trichocerca capucina
galeata, averaged over a growing season of 200 days.
The seasonal mean standing stock of 0. hyalina was 1.6 rotifers (1984) was about 300:18:9:6 g C m· 2 a· 1. The
g DW/m2 , versus 0.9 g DW/m2 in 0. galeata. These annual secondary production of crustaceans and rotifers
standing crops have turnover rates of 11x and 27x per is up to 11% of the annual primary production. From this
growing season, respectively. In terms of maintenance estimate of the available energy flow through the eco­
costs per day and per unit of standing stock biomass, this system, an annual grazing lass of about 40% of the pri­
means that daily rates of energy throughput and food mary production to the metazooplankton seems probable.
demands are 2.5 x higher in 0. galeata than in D. hyalina
to maintain a given population density during the summer. Table 6 Annual averages of biomass and annual cumulative
The visible result is that D. hyalina dominates in seasons productivity of phytoplankton and of the dominant
with low primary productivity, but 0. galeata predominates heterotrophic plankton groups in L. Constance; values
in years of higher food supply. from different years through 1980s.
The annual copepod productivity is small compared with Biomass Productivity
Daphnia, mainly because of their langer development. (g C/m2) (g Cm2 a· 1)
Their seasonal occurrence appears to vary inversely with Phytoplankton 1.1 300
the cladocerans: when population densities of the cope­
Bacteria 1.0 200
pods are low, the cladocerans are numerous, and vice
Protoza 0.23 20
versa. The most important copepod species, C. vicinus,
Rotifera 0.15 7.5
ceases its development between June to September,
Crustacea 1.2 26
living in diapause on the lake bottom as a dormant C-IV
Fish ? 5
stage.
Sum of heterotrophs >2.6 260
Rotifers
During the course of the year, about 50 species (from 20 Fish community
genera) of rotifers are found in Lake Constance. The The fish fauna of Lake Constance comprises about 30
phases of maximum population growth are shown for 12 species (Table 7) including three "forms" of coregonids
species in Figure 11: several "vernal" species grow du ring (Coregonus lavaretus): "Sandfelchen" are confined to the
the spring, simultaneously with the annual maximum of shallower, eutrophic L. Constance-Untersee, while an
phytoplankton, whereas the "aestival" and "autumnal" inshore-spawning ("Gangfisch") and a pelagic-spawning
species or groups are clearly separated in time. Within a form ("Blaufelchen") live in the mesotrophic main basin
season, the rotifers may reach or even surpass the (Obersee). Non-native species are the pikeperch
phytoplankton as biomass. The same is seen for the (Stizostedion lucioperca) and the rainbow traut (Salmo
daphnids and the copepods, in late spring. This transi­ gairdneri).
tory, high density of herbivorous zooplankton leads to
"overgrazing" of the phytoplankton, and, consequently, to Whitefish (C. lavaretus) and perch (Perca fluviatilis) are
very low algal densities in June, called the "clear-water the economically most importantspecies, while eel (Anguilla
phase". anguilla), pikeperch (S. lucioperca) and northern pike
(Esox lucius) and cyprinids are of less importance.
Biomass and productivity relations among the plankton
The average biomass and the annual production of the Fisheries regulations in the lake are identical for all lake­
different plankton components are compared in Table 6. bordering countries. They are decided by the IBK (Inter­
In both 1984 and 1985, the average biomass of the phyto­ nationale Bevollmächtigtenkonferenz), which includes
plankton and the total consumer community were in the delegates from all countries. Fisheries in littoral waters
same range: there was 25% to 30% more phytoplankton down to a depth of 25 m is under national control, whereas
than zooplankton. Rotifers and ciliates were present at the deeper regions of the lake are open to fishermen of all
about 10% of the crustacean biomass, while the herbivorous bordering countries. Fish are caught with gill nets (operated
cladocerans had about half the copepod biomass. In as bottom or dritt nets), trap nets, and fyke nets by about
terms of annual productivity, the relationship primary 170 professional fishermen. In addition more than 10,000
productivity: daphnids (both 1980-82): copepods (1988): anglers spend time on the lake.

15
Table 7 List of fish species occuring in L. Constance Artificial prapagation of fish in L. Constance started about
100 years ago. Eight hatchery plants operate around the
Brown traut Salmo trutta
lake, incubating up to 400 million whitefish eggs per year,
Lake traut Salmo trutta
Rainbow traut Salmo gairdneri as weil as smaller amounts of lake traut and northern pike.
Grayling Thymal!us thymallus Some of the larvae are pre-fed on zooplankton (whitefish,
Arctic char Salvelinus alpinus northern pike), others on artificial diets (lake traut, whitefish)
Whitefish Coregonus lavaretus prior to stocking the lake. In the case of whitefish, a
Common carp Cyprinus carpio correlation can be demonstrated between number of pre­
fed larvae stocked and yields two years later in L. Constance­
Barbel Barbus barbus
Untersee, while in L. Constance-Obersee effects of stocking
Chondrostoma nasus
Minnow are less evident.
Phoxinus phoxinus
Chub Leuciscus cephalus
Dace Leuciscus leuciscus The littoral and profundal zones
Bream Abramis brama
Silver bream 8/icca bjoerkna Due to the large depth of L. Constance, the greatest part
Crucian carp Carassius carassius of its benthic habitat lies in the profundal zones whereas
Bleak Alburnus alburnus the littoral area comprises only about 10% of the lake
Roach Rutilus ruti!us surface area. Until now, most studies on benthic community
Rud Scardinius erythrophta!mus structure focussed on the macroflora and macrozoobenthos,
Rhodeus sericeus while little is known on the meso- and microfauna and the
Tench Tinca tinca microflora. The available data are also too scarce for a
Perch Perca f!uviatilis realistic estimate of the intensity of praduction and
Pike perch Stizostedion lucioperca mineralization processes in the respective zones and their
Burbot Lota Iota partition within the lake.
Gudgeon Gobio gobio
Bullhead Cottus gobio Historical aspects
Threespine stickleback Gasterosteus aculeatus
Eel Anguilla anguilla The first notes on benthic organisms inhabiting L. Con­
Northern pike Esox lucius stance date from 1886. Until 1942 most benthic graups
Spined loach Cobitis taenia had been studied in some detail, providing valuable
Catfish Silurus glanis information on community structure in the oligotrophic
lake. During the eutrophication up to the end of the 1970s,
many taxonomic groups had been treated in monographic
Average annual yield (official statistics, excluding angler's studies, and structural changes during this period are weil
catches for L. Constance-Obersee increased from 320 documented. Parallel to eutraphication processes, the
metric tons (1914-1920) to 1,128 tons (1981-1987) (Fig. recent spread of some species caused drastic changes in
12), which corresponds to about a 4-fold increase. During the benthic food web. Most important was the arrival of the
the period of eutrophication, the fish community changed mussel Dreissena polymorpha at about 1966.
fram a formerly "coregonid lake" to a "coregonid-perch"
lake. The present strang fluctuations in annual yield directly Macrophytes and eutrophication
reflect the varying year-class strengths of coregonids and
perch; generally only one year class is exploited at a given Reed and reed decay
time. Recent declines in total phosphorus loading have not
yet been reflected in fish yields. Eutraphication provoked an increase in Phragmitesdensity,
but a concomitant weakening of stem stability. These
In contrast to smaller lakes, zooplankton are the main food symptoms led to reduced water exchange between lake
for most fish in L. Constance during the grawing season. and reed belt. At the same time, great masses of filamentous
Even many species which normally consume other food, algae grew in the littoral zone but died in autumn. As is
become planktivoraus during the summer, e.g. trout, adult indicated from biotests, the decay products of these algae
perch, eel and raach. The influence of fish predation may have contributed to the decline of the reeds by
pressure on the zooplankton community is being studied. preventing shoot development and inhibiting water uptake
by roots. In other parts of the lake reed stands were
reduced by increased erasion. This was caused by changes
Fig. 12 10-year averages of annual yield of professional in current patterns due to man-made buildings (walls,
fishery at L. Constance, and proportion of different fish harbours, etc.) in the littoral region. Recent activities have
species. attempted to stabilize the reed belts and to recover lost
areas by apprapriate management.
T0!',6
1500 Submersed vegetation
0 OTHERS The eulittoral zone in the oligotrophic phase was char­
[J PERCH acterized by an association of submersion-tolerant plants
1000 II COREGONIDS (e.g. Litorella uniflora) which became extinct with
eutraphication due to the excessive grawth of filamentous
algae and accumulation of plant debris at the shore-line in
autumn. The sublittoral vegetation shifted from Chara
spp. and braad leafed Potamogeton spp. to a grass like
500 association of Zannichel/ia and Potamogeton pectinatus.
In contrast to the former, the latter began to decay as early
as September. Afterwards the sublittoral was free of
vegetation and exposed to wave action during autumn
and winter which contributed to enhanced erasion. Recent
1914-20 21-XI 31-40 51 - fil 61- 70 71-00 81-87 evidence indicates that the macraphyte community

16

Fig. 13 Effects of riverine loadings upon zoobenthos and tubificid density
of total organic carbon by rivers is given by arrows of proportional w1dth. Elevated dens1t1es of tub1f1c1ds are shown by
small and by large dots (104 and 105 tubificids/m2 , respectively), based on a gnd of 1 x 1 km.
composition is changing back to the former state. Within
a sublittoral, hard-bottom association, dominated by
Rhodophyta, one of the few freshwater Phaeophyta species
was described, Bodane/la lauterborni(Zimmermann 1928).
This alga seems to be reduced by the space competitor
Dreissena.
The zoobenthos community
Littoral
In the past, riverine inflows into the eastern part of L.
Constance brought high organic loadings into the lake and
to its sediments. These resulted in high densities of
tubificids up to >3·106 /m2 (Fig. 13). In areas dominated by
primary production and not by organic loadings, many
structural changes are weil documented: While the
Arthropoda dominated the littoral macrofauna in the 1920s,
at present the Tubificidae are the most important group.
The sessile filter-feeding Spongillidae and Bryozoa
disappeared and were functionally replaced by Dreissena
polymorpha. Many !arge taxa with an annual or biennial
life cycle disappeared too, indicating a shift from K- to r­
strategists. Numbers of species and diversity in littoral
Cladocera declined, but for littoral Tubificidae the opposite
was observed.
Profundal
Even in areas not.influenced by riverine inputs, Tubificidae
dominate the macrofauna, followed by chironomid larvae
(mainly Micropsectra) and by the turbellarian Dendrocoelum.
Mean annual biomass of the macrofauna is ca. 4 g fresh
weighVm2 . In terms of energy flow through the profundal
communities, and with respect to microbial and microfaunal
activity, the macrofaunal contribution is one to a few
percent of total benthic catabolism. The tubificids seem to
be more important with respect to bioturbation, transporting
deposited organic matter from the anaerobic sediment
layers. In this way, sediment equivalent to 10-20% annually
sedimented matter can be re-exposed to the microbial
and chemical processes of early diagenesis.
Lake Constance as a source of drinking water and
recreation
In addition to fisheries, the lake has become important for
other economic reasons. Situated in a densely populated
region, large amounts of drinking water are required and
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of Lake Con�tance during the 1970s. Input
_ _ _
intense demands for recreational activities have to be
satisfied.
In spite of the eutrophication problems described above,
the water quality of L. Constance has always been
significantly better than the minimum standards required
for drinking water. Drinking water is pumped from the lake
at 25 sites (always from deep water below 30 m) equivalent
to 170·106 m3 of water per year, which is less than 2% of
the annual outflow. Supplying more than 4 million people
with drinking water, L. Constance is the most important
reservoir in Central Europe. The largest drinking water
plant is situated at Sipplingen, where the water is pumped
from a depth of more than 60 m and then distributed via a
network of pipelines into 500 cities and villages of South­
West Germany.
Surrounded by alpine mountains in the southeast and by
gentle hills with vineyards, hops and apple-trees in the
north, the lake has become a major tourist area, with more
than 2 mio visitors registered per year. This creates an
additional burden for the lake ecosystem: not-withstanding
the increased sewage problems (which now are partly
solved by an efficient sewage 1=1urification), the main
problems result from structural changes arising from
tourism (land use for parking, harbours, camping,
appartments etc.). More than 50,000 boats registered at
the lake require large areas of the littoral for anchorage.
Moreover, they cause additional water pollution, threatening
the drinking water quality with the emission of toxic
substances from motors (e.g. aromatic and aliphatic
hydrocarbons). Therefore, an administrative order te regulate
the emission levels of motors is in preparation.
Concerned with preserving the L. Constance ecosystem
and its surrounding region, the International Commission
for the Protection of L. Constance (IGKB) reacted to this
development by publishing memoranda on the current
state of the ecosystem and guidelines for the maintenance
of the purity of L. Constance water, e.g.,
- to treat the lake and its catchment area as an ecological
entity,
- to avoid further population growth in the lake region,
to prevent any further increase in sailing activities,
- to stop structural changes for touristic purposes
around the lakeshore,
- to take care that the existing laws and rules for the
protection of nature and landscape are followed and
promoted.
17