Biological Recovery of the Bohemian Forest Lakes from Acidification
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Biologia, Bratislava, 61/Suppl. 20: S453—S465, 2006 Section Zoology DOI: 10.2478/s11756-007-0071-y Biological recovery of the Bohemian Forest lakes from acidification Linda Nedbalová1,2, Jaroslav Vrba3,4,JanFott1, Leoš Kohout1,Jiří Kopáček4,3, Miroslav Macek5,4 & Tomáš Soldán6,3 1Charles University in Prague, Faculty of Science, Department of Ecology, Viničná 7,CZ-12844 Prague 2, Czech Republic; e-mail: [email protected] 2Institute of Botany AS CR, Dukelská 135,CZ-37982 Třeboň, Czech Republic 3University of South Bohemia, Faculty of Biological Sciences, Department of Ecology and Hydrobiology, Branišovská 31, CZ-37005 České Budějovice, Czech Republic 4Biology Centre AS CR, Institute of Hydrobiology, Na Sádkách 7,CZ-37005 České Budějovice, Czech Republic 5Universidad Nacional Autónoma de México, campus Iztacala, Av. de los Barrios 1, Los Reyes Iztacala, Tlalnepantla, 54090 Edo. México, México 6Biology Centre AS CR, Institute of Entomology, Branišovská 31,CZ-37005 České Budějovice, Czech Republic Abstract: A limnological survey of eight small, atmospherically acidified, forested glacial lakes in the Bohemian Forest (Šumava, B¨ohmerwald) was performed in September 2003. Water chemistry of the tributaries and surface layer of each lake was determined, as well as species composition and biomass of the plankton along the water column, and littoral macrozoobenthos to assess the present status of the lakes. The progress in chemical reversal and biological recovery from acid stress was evaluated by comparing the current status of the lakes with results of a survey four years ago (1999) and former acidification data since the early 1990s. Both the current chemical lake status and the pelagic food web structure reflected the acidity of the tributaries and their aluminium (Al) and phosphorus (P) concentrations. One mesotrophic (Plešné jezero) and three oligotrophic lakes (Černé jezero, Čertovo jezero, and Rachelsee) are still chronically acidified, while four other oligotrophic lakes (Kleiner Arbersee, Prášilské jezero, Grosser Arbersee, and Laka) have recovered their carbonate buffering system. Total plankton biomass was very low and largely dominated by filamentous bacteria in the acidified oligotrophic lakes, while the mesotrophic lake had a higher biomass and was dominated by phytoplankton, which apparently profited from the higher P input. In contrast, both phytoplankton and crustacean zooplankton accounted for the majority of plankton biomass in the recovering lakes. This study has shown further progress in the reversal of lake water chemistry as well as further evidence of biological recovery compared to the 1999 survey. While no changes occurred in species composition of phytoplankton, a new ciliate species was found in one lake. In several lakes, this survey documented a return of zooplankton (e.g., Cladocera: Ceriodaphnia quadrangula and Rotifera: three Keratella species) and macrozoobenthos species (e.g., Ephemeroptera and Plecoptera). The beginning of biological recovery has been delayed for ∼20 years after chemical reversal of the lakes. Key words: Acidification, lake recovery, phytoplankton, zooplankton, bacteria, ciliates, macrozoobenthos. Introduction sponding biological recovery of these areas has, how- ever, been significantly delayed or uncertain (Jeffries Small glacial mountain lakes with water of low ionic et al., 2003; Skjelkv˚ale et al., 2003; Wright et al., strength water usually host less complex pelagic com- 2005). munities compared to larger water bodies. Such head- Like the whole region of Central Europe, the Bo- water lake ecosystems are greatly vulnerable to at- hemian Forest (Šumava, B¨ohmerwald) was exposed to mospheric acidification (e.g., Kalff, 2002), which has heavy atmospheric pollution during the last century, caused a dramatic reduction of biodiversity followed by followed by a significant drop in both S and nitrogen disruption of food web structure in many lake districts (N) depositions during the last two decades (Kopáček since the middle of the 20th century (e.g., Schindler, et al., 2001, 2002). Due to severe acidification, the Bo- 1988). Deposition of acidifying pollutants has signifi- hemian Forest lakes became unique ecosystems, with cantly declined in Europe and North America, partic- bacterioplankton and/or phytoplankton largely domi- ularly due to reduced sulphur (S) emissions since the nating their pelagic biomass, the complete absence of 1980s, leading to a partial reversal of surface waters fish, and significantly reduced or even extinct crus- from acidification (Stoddard et al., 1999). A corre- tacean zooplankton (Vrba et al., 2003a, b). Accord- c 2006 Institute of Zoology, Slovak Academy of Sciences S454 L. Nedbalová et al. Table 1. Location and main characteristics of the Bohemian Forest lakes and their catchments. Lake CN CT RA PL KA PR GA LA Latitude N 49◦11 49◦10 48◦58 48◦47 49◦08 49◦05 49◦06 49◦07 Longitude E 13◦11 13◦12 13◦24 13◦52 13◦09 13◦24 13◦07 13◦20 Elevation m a.s.l. 1008 1027 1071 1087 918 1079 935 1085 Lake area ha 18.8 10.7 5.7 7.6 9.4 4.2 7.7 2.6 Catchment area km2 1.24 0.89 0.58 0.67 2.79 0.65 2.58 1.02 Max. depth m 40 35 13 19 9 17 16 3 Lake volume 106 m3 2.92 1.86 0.18 0.61 0.25 0.35 0.45 0.05 Explanations: Codes and names of the lakes: CN – Černé jezero; CT – Čertovo jezero; PL – Plešné jezero; PR – Prášilské jezero; RA – Rachelsee; GA – Grosser Arbersee; KA – Kleiner Arbersee; LA – Laka. For locations see Fig. 1; for more detailed information on lakes and their catchments see VRBA et al. (2000) and JANSKÝ et al. (2005). 14º E Poland Germany 50º N Czech Republic GERMANY Černé jezero (CN) Čertovo jezero (CT) Austria Slovakia Kleiner Arbersee (KA) Laka (LA) Grosser Arbersee (GA) Prášilské jezero (PR) Rachelsee (RA) CZECH REPUBLIC Plešné jezero (PL) AUSTRIA 06020 40 km Fig. 1. The map of location of the Bohemian Forest lakes (their codes used throughout the text are in parentheses). ing to their acidification status in 1999, eight Bo- steep, forested (Norway spruce) catchments with acid sensi- hemian Forest lakes were divided into three categories: tive bedrock (mica-schist or granite); for detailed morpho- (i) strongly acidified lakes (Černé jezero, Čertovo jezero, logical data and land use history, see VESELÝ (1994), WEIL- Rachelsee, Plešné jezero), (ii) moderately acidified lakes NER (1997), VRBA et al. (2000) and JANSKÝ et al. (2005). (Prášilské jezero, Kleiner Arbersee), and (iii) slightly Samples of lake water were taken at the deepest points acidified lakes (Grosser Arbersee, Laka) (Vrba et al., of the lakes during 10 days in early September 2003. The 2000, 2003a). samples for chemical analyses were taken from the epil- imnion (0.5 m depth) and were immediately filtered through This study presents both current species composi- a 200 µm polyamide sieve. The samples for microbial analy- tion and carbon biomass of the plankton, together with ses were sampled with a van Dorn sampler (1 m length, 6.4 L relevant chemistry of the Bohemian Forest lakes. In ad- volume) from specific depths (5–6 samples along the vertical dition, we present the first comparison of littoral macro- profile in the deep lakes, except for the shallow KA and LA zoobenthos assemblages in all the lakes. Our main ob- with 3 and 2 samples, respectively). The samples were fixed jective is to compare the new (2003) data with a former with either formaldehyde (for bacteria) or acid Lugol’s so- survey from 1999 and older acidification data since the lution (for phytoplankton and ciliates). Large zooplankton early 1990s, in order to evaluate biological recovery of were sampled by several vertical hauls with a quantitative µ the lakes from acid stress. net (200 m mesh size) of the Apstein type, while small zooplankton were sampled with the van Dorn sampler from the same depths as microorganisms, and concentrated by µ Material and methods a plankton net (40 m mesh size); both samples were pre- served by formaldehyde. Temperature and dissolved oxygen Eight small, glacial lakes (see Tab. 1 for lake names and were measured with a DataSonde 4 (Hydrolab, USA) at 0.5- their codes used further in the text) in the Bohemian Forest mintervals. (C Europe) were studied. The lakes are situated along the Main tributaries were sampled near their inlets to the border between the Czech Republic and Germany (Fig. 1) in lakes and the water discharges were estimated by means of Biological recovery of Bohemian Forest lakes S455 a bucket and stopwatch. Samples were filtered immediately Samples for ciliate analysis were postfixed with Bouin’s through a 40 µm polyamide sieve to remove coarse particles fluid. The quantitative protargol staining was applied re-suspended from the streambed during sampling. Each in- (MONTAGNES &LYNN, 1993); samples were dehydrated in let sample was analysed separately. The chemistry and dis- ethanol, phenol-xylol and xylol series, and neutral Canada charge of the inlets were used to calculate the discharge- balsam-mounted. The whole filter area was inspected at weighted mean chemical composition of total water input 500× to 1250× on microscopes equipped with Nomarski to each lake. (DIC) contrast (Olympus, Nikon, or Leica). FOISSNER In the laboratory, water samples were filtered with ei- (1994), FOISSNER et al. (1999) and literature cited therein ther membrane filters (pore size of 0.45 µm) for the de- were used for identification. For all observed specimens, indi- termination of ions (ion chromatography) and silica (Si) or vidual cell volume was calculated (using simple shape mod- with glass-fibre filters (pore size of 0.4 µm) for other anal- els) from dimensions measured in the protargol stained sam- yses, except for samples for pH, acid neutralizing capacity ples. (ANC, determined by Gran titration), and total concentra- The zooplankton samples were fixed by formaldehyde tions of aluminium (Al), phosphorus (P), carbon (C), and N, (4% final concentration), and counted on an inverted mi- which were not filtered beyond the field pre-filter.