Hydrobiologia 501: 1–5, 2003.© 2003 Kluwer Academic Publishers. Printed in the Netherlands.

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Opinion

Re-assessment of Wundsch’s (1940) ‘H2S-Oscillatoria-Lake’ type using theeutrophic Lake Scharmützel (Brandenburg, NE Germany) as an example

Andreas KleebergBrandenburg Technical University of Cottbus, Chair of Water Conservation, Seestraße 45, D-15526 Bad Saarow,GermanyTel: 33631-8943. Fax: 33631-5200. E-mail: a.kleeberg@limno-tu-cottbus.de

Received 8 February 2001; in revised form 18 March 2003; accepted 16 June 2003

Key words: eutrophication history, sulfide, cyanobacteria, Oscillatoriales, Chironomidae

Abstract

Thienemann’s 1922 ‘biological lake’ classification indicates the increase in the trophic level of lakes. It is based onthe sucession of the disappearance of certain benthic indicator organisms in relation to the gradient of a decreasingO2concentration in the deep water, i.e. from O2-sensitive Chironomidae spp. (non-biting midges) to the lesssensitive Chaoboridae (phantom midges) larvae. As early as the mid 1930s, several lakes in Brandenburg, e.g. LakeScharmützel, belonged to the last category of this classification. They were O2-deficient in deep water during sum-mer and lacked the Chironomidae larvae. Simultaneously, filamentous cyanobacteria (Oscillatoriales) appeared.This lead to the replacement of Thienemann’s indicators, i.e. O2 by H2S and Chironomidae by Oscillatoriales, andto Wundsch’s 1940 new ‘H2S-Oscillatoria-Lake’ type. Since H2S and Oscillatoriales were not clearly identifiedas symptoms of eutrophication, it is not justified to use them to characterise a separate lake type. However, theseindicators are of ecological importance, since the regional and common creeping increase in SO4

2− concentrationfavoures the current high H2S formation. The successive deterioration in O2 conditions, the increase in the extentof H2S formation related to the decrease in species diversity clearly indicates that Lake Scharmützel reached aqualitatively new state of eutrophication within 60 years.

Introduction

While Thienemann (1922) was describing his systemof ‘biological lake types’, scientific fishery journalswere appealing for an increase in the fishery yields:‘Fishermen, fertilize your lakes!’ (e.g. Walter, 1925).A few years later, the first indications of increasedeutrophication became visible in various lakes in theMark Brandenburg, e.g. an ‘Oscillatoria disease’(Czensny, 1938) or a ‘Lake deterioration’ (Schäper-claus, 1940). Between 1934 and 1937, following ageneral decline in fishery yields as well as changesin fish communities (e.g. Schiemenz, 1934; Schäper-claus, 1940), Wundsch investigated at 41 lakes andreported his findings (Wundsch, 1940). In his German-language paper, he described ‘the development of aspecial type of lake (H2S-Oscillatoria-Lake) in the

river-lake region of River Spree and River Havel, andits importance for the fishery-biological conditions inthis region’. Wundsch concluded that the mass devel-opment of Oscillatoriales (filamentous cyanobacteria)and their rapid breakdown are, in part, responsiblefor the H2S formation, since the occurrence of sulfurdroplets in cyanobacterial cells (Hinze, 1903), theirsulfur content (Kolkwitz, 1914) and the sulphuretum’,i.e. a habitat containing H2S (Baas Becking, 1925),was already known. Subsequently, Hutchinson (1957)distinguished 11 main and 76 subtypes of lakes, butdid not include the ‘H2S-Oscillatoria-Lake’. Des-pite the considerable literature that has evolved fromthese early studies, the important question pertain-ing to the time of a distinct eutrophication remainsunanswered. Only a few mechanisms of water de-teriorations are documented (Müller, 1952; Scharf,

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1971), but clear evidence would be helpful in determ-ining the pristine stage of lakes in Brandenburg, i.e.a water state with no significant anthropogenic im-pact. Therefore, the aims of the present paper are,(1) to provide the historical background for the intro-duction of the ‘H2S-Oscillatoria-Lake‘ by Wundsch(1940), (2) to place Lake Scharmützel in Wundsch’sand Thienemann’s classification system, (3) to discussthe regional relevance of Wundsch’s lake type from acontemporary view, and (4) to compile evidence thatlakes in Brandenburg were not, as assumed, only at the1970s considerably suffering from eutrophication andits consequences, but already in the 1930s.

Description of site studied and routine measurements

Lake Scharmützel is a dimictic, eutrophic, sulfate-richlake (Table 1). Fortnightly vertical profiles of watertemperature (tw), pH, and dissolved oxygen (O2) weretaken using a multiprobe (H20�, Hydrolab) in 0.5 mincrements at the deepest location at Wendisch Rietz.At the same occasions, the concentration of sulf-ide (HS−) was determined photometrically (Rohde &Nehring, 1979). The concentration of non-dissociatedfree hydrogen sulfide (H2S), which is dependent on pHand tw, was calculated according to Kleeberg (1997).During the Wundsch campaign (1934–1937), O2 andH2S were determined according to Czensny (1926)and Stroede (1933), respectively. The lowest value ofH2S detected was 0.16 mg l−1 (Ohle, 1934).

Results and discussion

Historical background

Thienemann (1922) developed a lake classificationsystem of ‘biological lake types’ based on thebenthic summer profundic populations of variousO2-sensitive Chironomidae (non-biting midges) andChaoboridae (phantom midges) larvae and the cor-responding O2 conditions. He distinguished meso-trophic and oligotrophic lakes of the ‘Tanytarsus-lake type’, and eutrophic lakes of the ‘Chironomus-lake type’. The succession proposed by him, dir-ects from the ‘Tanytarsus-lake’, via the ‘Corethra-lacking Chironomus-lake’, to the ‘Corethra-occurringChironomus-lake’, and further via the ‘bathophilus-lake’ to the ‘plumosus-lake’, and finally to the‘Chironomus-lacking Corethra-lake’ and correspondsto the gradient of decreasing O2 concentration in thedeep water.

Figure 1. Seasonal course of dissolved oxygen (O2) and hydro-gen sulfide (H2S) concentration at the deepest location of LakeScharmützel 0.5 m above bottom in 1996, and O2 concentration atthat depth in 1934 and 1937 according to Wundsch (1940).

Already in the 1930s, other even shallower lakesin the region were also found to lack any macro-zoobenthos in the summer profundal (Czensny, 1938),as e.g. in Lake Riewend, with H2S concentrationsbetween 0.78 and 7.88 mg l−1 at 4 and 6 m depths,respectively. In addition, mass blooms of filament-ous cyanobacteria occurred (Czensny, 1938; Wund-sch, 1940). Consequently, Wundsch (1940) introduceda new classification, substituting the indicators O2and Chironomidae by H2S and the characteristic bluegreens (Oscillatoriales), resulting in his new ‘H2S-Oscillatoria-Lake’ type.

Position of Lake Scharmützel in Wundsch’s andThienemann’s classification system in the 1930s andin the 1990s

Distinct summer hypolimnetic O2 minima were foundbetween 1934 and 1937 (Fig. 1), but no H2S was de-tected by Wundsch (1940). The characteristic speciesof the profundal was Chironomus bathophilus Kieff.(Chironomidae, today: C. anthracinus Zett.). Hence,Wundsch classified the lake as a ‘bathophilus-lake’,i.e. as a ‘bottom-eutrophic’ lake type with relativelyclear water and low planktonic production comparedto the ‘eutrophic level’ of the ‘normal Baltic Lakes’.Moreover, the relatively O2-insensitive Corethra spp.(today: Chaoborus) were still found in 1934, andOscillatoriales, especially Oscillatoria redekei Meff.(today: Limnothrix redekei Meff.) were absent (Wund-sch, 1940). Around 1934, Lake Scharmützel wasqualified as a ‘Chironomus-free Corethra-lake’, andtherefore, according to Thienemann (1922) it be-longed to the last category of his range of ‘biologicallake types’.

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Table 1. Morphometrical and limnological characteristics (epilimnion, 1996) of Lake Scharmützel,NE Germany (52◦ 15′ N, 14◦ 03′ E)

Morphological parameter (unit) Limnological parameter (unit) Min. . . . max.

Surface area (km2) 12.09 total phosphorus (µg l−1) 36–102

Volume (106 m3) 108.23 total nitrogen (µg l−1) 44–912

Mean depth (m) 8.8 sulfate (mg l−1) 54.7–58.3

Maximum depth (m) 29.5 secchi depth (m) 0.8–3.6

Mean residence time (a) 16 seston (mg d.w. l−1) 1.8–8.8

Drainage area (km2) 112 chlorophyll a (µg l−1) 11–53

Later on, the O2 concentration was zero below15 m water depth only in September 1949, and H2Swas detected for the first time by Müller (1952).At present, the annual H2S formation usually startsin June and lasts until October/November (Fig. 1).H2S moves up from lower strata to depths between8 and 9 m, leading to HS− dominated hypolimneticchemistry (Kleeberg, 1997). Consequently, the currentO2 and H2S conditions during the summer months,compared to those of 1934 and 1937 (Fig. 1), nolonger allow inhabitation of the profundal by Chiro-nomidae. Even the more resistent predatory Corethraspp. (today: Chaoborus) are rarely found in thesummer profundal. Moreover, 76–83% of the sum-mer phytoplankton is dominated by cyanobacteria,mainly Oscillatoriales (Zippel, 1996). Therefore, LakeScharmützel would at present theoretically be classi-fied as a ‘H2S-Oscillatoria-Lake’.

H2S is a reduced, very reactive compound. Itssolubility in water is more than twice as high thanthat of CO2. H2S is toxic to fish and macroinver-tebrates, especially at pH values <7 if more than50% HS− occurs in undissociated form, i.e. as freeH2S. Even concentrations of 0.015 mg H2S l−1 aredemonstrably chronically toxic to aquatic organisms(Oseid & Smith, 1975). Furthermore, the distributionof benthic invertebrates is controlled by HS− (Iwasakiet al., 1986). For Lake Scharmützel, all of this is reflec-ted by a 80% decrease in macrozoobenthos abundanceand diversity also decreased from 1937 to the present(Ordóñez, 2001). The disappearance of fish speciesas Coregonus albula L. (Vendace) and Leucaspiusdelineatus (Heck.) (Sunbleak) is also significant.

‘H2S-Oscillatoria-Lake’ type as one potential finalstate of progressive eutrophication

Wundsch (1940) stated: ‘The H2S-Oscillatoria-Laketype is not to be taken as the natural final state of aprogressive eutrophication of certain lake individuals,but as a special formation due to a particular coin-cidences of factors, which have a regional characterand which are not attained everywhere’. The first partof this assertion is not unrestrictedly valid. Czensny(1938) could show that lakes of Brandenburg ‘laidup with’ Oscillatoria (including H2S) are enriched bynutrients in comparison to ‘normal eutrophic lakes’(Table 2). Moreover, vertical profiles of an increasingphosphate concentration with depth (Schweng, 1937;Wundsch, 1940) clearly indicate an internal phos-phorus loading. Hence, Schäperclaus (1940) statedthat ’solely the richness of plant nutrients, light andheat are responsible for the development of the phe-nomena of an Oscillatoria-dominated lake’. The factthat 25 lakes (i.e. 69.4% of the 36 lakes studied) showan Oscillatoria dominance (Wundsch, 1940) also sup-ports the hypothesis that numerous lakes in the regionwere already eutrophied in the 1930s. At present, 965lakes of >0.05 km2 are registered for Brandenburg.Of the 784 lakes which have already been studied 728(92.8%) are eu- to polytrophic (Mietz et al., 1996).Cyanobacterial blooms are common place, i.e. also the11 Oscillatoria-free lakes out of the 36 lakes studiedby Wundsch (1940) are today in part sulphureta anddominated by Oscillatoriales (Zippel, 1996). Successof Oscillatoriales is due to their ability to live on aminimal income of light energy in near-continuouslymixed, turbid systems of exposed, shallow and oftenenriched lakes as well as in cold, deep (and, thus, well-mixed) lakes (Reynolds, 1997). Also for the deep LakeScharmützel the appearance of the not N2-fixing cy-anobacteria Limnothrix redekei Meff. and Planktothrix

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Table 2. Span of chemical parameters of lakes of Brandenburg‘laid up with’ Oscillatoria and H2S (n = 2) in comparison to‘normal eutrophic lakes’ (n = 4) for April, July and Novemberbetween 1934 and 1937. Data were drawn from Czensny (1938)

Parameter (Unit) ‘Normal eutrophic ‘Sick lakes’

lake’

pH 7.1–8.9 7–8.5

CaO (mg l−1) 63–103 82–131

KMNO4 32–57 53–110

consumption

(mg l−1)

NH4+ (mg l−1) 0.05–0.065 . . . 5.5

NO3− (mg l−1) 8–9 . . . 1.2

P2O5 (mg l−1) 0.012–0.460 0.001–0.190

Cl− (mg l−1) 36–57 46–60

SO42− (mg l−1) 70–80 45–140

agardhii Anagn. & Kom. (former: Oscillatoria redekeiMeff. and O. agardhii Gomont) (Zippel, 1996) is at-tributed to the increase in nutrients clearly indicatingits present state of eutrophication.

The second assertion of Wundsch (1940), that‘the H2S-Oscillatoria-Lake is a regional special caseof particular geological conditions’ is significant. Invarious lakes, the concentration of sulfate (SO4

2−)increased by a factor between 1.6 and 5.7 within thelast decades (Table 3). This is significantly more thanthe global increase for freshwaters, where human im-pact has caused a rise from a mean, pre-industrialvalue of 6.72 mg l−1 to a present mean value of 11.52SO4

2− mg l−1, i.e. by a factor of 1.7 within roughly100 years (Caraco et al., 1993). The SO4

2− in the lakes(Table 3) has been originated from natural (e.g., LakeScharmützel: geological background), and anthropo-genic sources (e.g., Lake Scharmützel: fertilizer, wastewater; Lake Flaken: lime stone open pit mining, andLake Dämeritz: lignite mining).

The increase in SO42− concentration can lead to

an increase in SO42− reduction (Kelly et al., 1982),

which, in turn, results in enhanced formation of HS−.In addition, phytoplanktic organic matter is proneto rapid mineralization by microorganisms and mayundergo anaerobic decomposition and/or putrefaction(desulfuration) to form also HS− (Dunette, 1989).However, in Lake Scharmützel only 7.9–10.2% of theformed HS− originated from desulfuration (Kleeberg,1997, 2000). Hence, the HS− originates mainly fromSO4

2− reduction, and due to the significant increasein SO4

2− concentration, at a certain level of pro-

Table 3. Examples for the increase in sulfate concentration in Lakesof Brandenburg. Data were drawn from Schweng (1937), Wundsch(1940), Müller (1952), Bursche (1955), Scharf (1971), Behrendt &Böhme (1994), and Kleeberg (unpubl.)

Lake Sulfate concentration in . . . Years Factor

Year (mg l−1) Year (mg l−1)

Werl 1951 39.4 1993 61.1 42 1.6

Flaken 1933 47.5 1993 202.4 60 4.3

Dämeritz 1933 31.2 1993 136.8 60 4.4

Scharmützel 1934 10.0 1996 56.6 62 5.7

ductivity, the extent of HS− formation might be morepronounced than in other, SO4

2−-poorer regions.

Conclusions

As shown for Lake Scharmützel as early as the early1930s, increasing eutrophication placed numerouslakes in Brandenburg theoretically at the last categoryof Thienemann’s (1922) scale, i.e. without any O2 orthe corresponding benthic indicator organisms in deepwater during the summer.

In the present view, the clear symptoms of eu-trophication (HS−, Oscillatoriales) – from Wundsch(1940) – logically lead to the introduction of the new‘H2S-Oscillatoria-Lake’ type. However, the latter isnot justified as a separate lake type in general, since theincreasingly Oscillatoriales occurrence due to nutrientrichness is not necessarily related to a HS− forma-tion. But this lake type is of a regional and ecologicalimportance due to the creeping increase in SO4

2− con-centration which favoures the current extent of theHS− formation. The successive deterioration in O2conditions, the increase in the extent of HS− forma-tion related to the decrease in species diversity clearlyindicates that Lake Scharmützel reached a qualitat-ively new state of eutrophication within 60 yearsonly which, due to its expansion, was delayed incomparison to other smaller lakes in the region.

Acknowledgements

I sincerely thank B. Nixdorf (BTU Cottbus) and T.Rohrlack (Humboldt Univ. at Berlin) for making someolder literature sources available. The linguistic im-provements by U. Riebow (BTU Cottbus) are much

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appreciated. Comments by J. Padisák (Univ. Vesz-prém, Hungary), I. Chorus (UBA Berlin), and bya second anonymous reviewer greatly improved themanuscript.

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