Differences in benthic macroinvertebrate structure of headwater streams with extreme hydrochemistry

Biologia 68/2: 303—313, 2013Section ZoologyDOI: 10.2478/s11756-013-0156-8

Differences in benthic macroinvertebrate structureof headwater streams with extreme hydrochemistry

Jakub Horecký1, Jan Rucki2, Pavel Krám3, Josef Křeček4, Peter Bitušík5, Jan Špaček6

& Evžen Stuchlík1

1Institute for Enviromental Studies, Faculty of Science, Charles University in Prague, Benátská 2, CZ-12844 Prague, CzechRepublic; e-mail: [email protected] River Authority, State Public Enterprise, Holečkova 8, CZ-15024 Prague, Czech Republic3Department of Geochemistry, Czech Geological Survey, Klárov 3, CZ-11821 Prague, Czech Republic4Department of Hydrology, Czech Technical University in Prague, Thákurova 7, CZ-16629 Prague, Czech Republic5Research Institute and Faculty of Science, Matthias Belius University, SK-97401 Banská Bystrica, Slovakia6Labe River Authority, State Public Enterprise, Víta Nejedlého 951, CZ-50003 Hradec Králové, Czech Republic

Abstract: A synoptic study of acidified mountain streams covering six Czech sites was performed. The aim was to providebiological data from small mountain streams in catchments with historically high acid atmospheric deposition, which have sofar been subject of intensive long-term monitoring of hydrology and hydrochemistry only, in order to follow the developmentof the structure of benthic macroinvertebrates in the course of recovery from acidification. We focused on small headwaterstreams with minimum human influence in the catchment and relatively low concentrations of dissolved organic matter. Thesites were classified according to their water pH status: three of them were strongly acidified (pH 4.07–4.57, concentrationof reactive aluminium R-Al 448–1913 µg L−1), two were moderately acidified (pH 5.08–6.38, R-Al 52–261 µg L−1) and onenon-acidified (pH 7.63–7.89, R-Al 12–59 µg L−1). The largest biotic difference detected by PCA was in the presence of thecaddisflies Drusus, Rhyacophila, and Chaetopteryx villosa and stoneflies Leuctra pseudocingulata and Diura bicaudata. Theresults indicate that at the most acidified sites (the Lysina and the right branch of the Litavka), the process of biologicalrecovery has not started yet due to an insufficient increase in stream pH. Indeed, MAGIC modelling published earliershows that significant increases cannot be expected in the following decades. An average pH of at least 4.5 is needed forreturn of less acid-tolerant taxa such as Diura bicaudata, Leuctra major, L. pseudocingulata, L. pseudosignifera, Drusus orRhyacophila. However, at the Sklářský potok – Jizerka site, both the clear-cut of mature spruce plantations in 1984–1990and the regional drop in SO2 emissions in the 1990s resulted in a decline of acid deposition and rising streamwater pH.Mean annual pH at the Sklářský potok – Jizerka outlet increased from 4.0 (1982–1985) to 5.3 (1990–1994), but episodicacidification has still resulted in a delay in recovery of the biota, particularly acid-sensitive species, which may be expectedwithin a couple decades.

Key words: mountain catchments; acid atmospheric deposition; acidification; stream waters; low pH; benthic macroinver-tebrates; biological recovery; aluminium

Abbreviations: R-Al reactive aluminium; TOC total organic carbon; DOC dissolved organic carbon; PCA PrincipalComponent Analysis, MAGIC Model of Acidification of Groundwater in Catchments

Introduction

The most significant cause of anthropogenic acidifica-tion of surface waters in Central Europe is acid atmo-spheric deposition of sulphur and nitrogen oxides andammonia (Kopáček et al. 2001). The strong effects ofair-borne acidification in the Czech Republic was ob-served in hundreds of acid soft water streams with lowmineralization and hence low acid neutralizing capacity(Veselý & Majer 1998). The natural acidity of these wa-ters based on humic substances was often overwhelmedby anthropogenic acidity (Stuchlík et al. 1997), whichresulted in decreases of pH in water and significantchanges in water chemistry. These changes could lead to

aquatic ecosystem collapse (Psenner & Catalan 1994).The mobility of many metallic elements increases withdecreasing pH. Their toxic effects depend on the formof chemical compounds and can be reduced by higherconcentrations of humic acids (Kullberg 1992; Stuch-lík 2003) or higher concentrations of ions in the wa-ter (Winterbourn & McDiffet 1996). The most toxicelement in anthropogenically-acidified waters is alu-minium, which is highly toxic to aquatic organisms,primarily in its ionic form Al3+ (Hermann 1987).In the second half of the 1980s, a significant de-

crease in the emissions of sulphur oxides and certaindecrease in the emissions of nitrogen oxides in CentralEurope (Kopáček et al. 2002) resulted in a decrease of

c©2013 Institute of Zoology, Slovak Academy of Sciences

304 J. Horecký et al.

Fig. 1. Location of the study sites in the Czech Republic, Brdy Mts (LIR = Litavka – right branch, LIL = Litavka – left branch),Slavkov Forest (LYS = Lysina, PLB = Pluhův Bor) , and Jizera Mts (JER = Jeřice, JIZ = Sklářský potok – Jizerka).

acid atmospheric deposition, and subsequently startedthe process of recovery from anthropogenic acidifica-tion (Raddum & Fjellheim 2003; Stuchlík et al. 2005).However, large amounts of sulphur accumulated in soilhorizons in the period of increased acid depositions.Nowadays, this reserve is gradually leaching (Alewellet al. 2000), resulting in only slight increases of pH anddecreases of ionic concentrations forms of aluminium(so called hysteresis; Kopáček et al. 2002). Results ofdynamic hydrochemical modelling of small catchments(Hruška et al. 2002; Hardekopf et al. 2008) show thatacidification, despite decreasing trends in atmosphericdeposition, will still affect freshwater ecosystems for asignificant period of time.Few studies on the influence of acid atmospheric

deposition on the structure of macroinvertebrates inrunning waters have been published from the CzechRepublic (Růžičková 1998; Scheibová & Helešic 1999;Horecký et al. 2002, 2006; Fricová et al. 2007). Unfortu-nately, biological data from most long-term monitoredsites do not exist. The main task of this study was toelucidate the main differences in benthic macroinverte-brate structure of streams affected to different degreesby acid atmospheric deposition, and based on estab-lished and published differences, indicate the possibledirection of expected biological recovery. A secondaryobjective was to obtain complete biological data fromheadwater mountain streams in catchments with histor-ically high acid atmospheric deposition that have so farbeen the subject of intensive long-term monitoring ofabiotic parameters only, in order to compare and tracethe development of the structure of benthic macroin-vertebrates in the course of recovery from acidification.

Material and methods

Site characteristicsThis study focused on six headwater catchments situatedin three mountain regions of the Czech Republic with min-

imum human influence in the catchment except for formerhigh acid atmospheric depositions: the Brdy Mts (Litavka –right branch = LIR and Litavka – left branch = LIL),Slavkov Forest (Lysina = LYS and Pluhův Bor = PLB)and the Jizera Mts (Sklářský potok – Jizerka = JIZ andJeřice = JER; see Fig. 1). Sklářský potok – Jizerka is inten-sively monitored within the Mountain Waters project of theEarthwatch Institute. Lysina and Pluhův Bor (Krám et al.1997, 2009; Hruška & Krám 2003; Shanley et al. 2004) are inthe Czech GEOMON network of small forested catchmentsand in the International Long-Term Ecological Researchnetwork. Moreover, Lysina is in two international networksof the International Cooperative Programme – ICP Inte-grated Monitoring and ICP Waters, both organized underthe Economic Commission for Europe of the United Na-tions. Litavka – right branch (referred to as the Litavkarain-fed branch by Hardekopf et al. 2008) belongs to theCzech GEOMON network of small forested catchments.

The basic catchment characteristics are shown in theTable 1, for further details see Hardekopf et al. (2008), Krámet al. (1997), and Křeček et al. (2006).

All studied catchments are part of managed forests,particularly plantations of Norway spruce (Picea abies). Inthe JIZ catchment, after a clear-cut of spruce stands (1984–1990), herbaceous vegetation still dominates.

Sampling and field measurementsSamples of benthic macroinvertebrates and water samplesfor hydrochemical analyses were taken in April, July, andOctober 2005 from the sites LIL, LIR, LYS, and PLB; thesite JIZ in April and July 2005, and October 2004; the siteJER was sampled in July and October 1999 and in May2000. Supplementary samples of chironomid pupal exuviaewere taken at the sites LYS and PLB in July 2006 andOctober 2007.

In the field, basic environmental characteristics of the100 m long sampling stretch (mean slope, stream widthand depth, turbulence and substrate) were identified. Ateach sampling stretch, the composition of substrate (per-centage of mud, sand, gravel, stones, or rocks) and thestream turbulence composition (percentage of still, lotic,or riffled segments) were visually estimated and then thepercentage categories were converted to single index values

Benthic macroinvertebrate structure of headwater streams 305

Table 1. Basic environmental characteristics of the studied streams and catchments.

Site characteristics Catchment characteristics

Altitute Latitude Distance from source Area Altitude SlopeRegion Orientation Main recipient

◦ (N) ◦ (E) km km2 m a.s.l. %

LIR Brdy Mts 49.39 13.52 1.00 1.85 695–843 8.6 NE BerounkaLYS Slavkov Forest 50.02 12.40 0.90 0.27 829–949 11.5 NE TepláJER Jizera Mts 50.51 15.09 0.35 0.10 800–886 9.0 NW Lužická NisaLIL Brdy Mts 49.40 13.52 0.65 0.90 700–840 8.0 NE BerounkaJIZ Jizera Mts 50.49 15.21 0.57 1.00 860–993 7.5 NE JizeraPLB Slavkov Forest 50.04 12.48 0.70 0.22 690–804 13.0 SE Teplá

Geology Dominant Main source ofvegetation streamwater

LIR sandstone, quartz conglomerates, deluvial sediments spruce shallowLYS leucogranite, deluvial sediments spruce shallowJER granite, deluvial sediments spruce shallowLIL sandstone, quartz conglomerates, deluvial sediments spruce deepJIZ granite, deluvial sediments grass shallow

Calamagrostis sp.PLB serpentinite, deluvial sediments spruce shallow

Fig. 2. Comparison of pH, specific conductivity (SC25), sulphate SO2−4 , and reactive aluminium R-Al of the strongly acidified (JER,

LIR, LYS), moderately acidified (JIZ, LIL) and non-acidified (PLB) catchments. Mean and limit values recorded during the dates ofmacroinvertebrate sampling are given for the each site.

as follows: Substrate Index = 1 (% mud) + 2 (% sand)+ 3 (% gravel) + 4 (% stones) + 5 (% rocks); TurbulenceIndex = 1 (% still) + 2 (% lotic) + 3 (% riffled). The al-

titude range, area and bedrock of the catchment, and thedistance of the site from source were identified from 1:50 000maps. Water samples for chemistry were taken prior to bio-


Table2.Selectedparametersofwaterchemistryatthestudiedsites:mean(min–max)forthehydrologicalyear2005.*DetaileddataarenotavailableforJER,thereforevaluesofthe

watersamplestakeninyear1999/2000areshowninstead.

LIR

LYS

JER*

LIL

JIZ

PLB

pH

4.15(4.07–4.24)

4.25(4.11–4.53)

4.55(4.51–4.57)

5.34(5.08–5.98)

5.46(5.18–6.38)

7.71(7.63–7.89)

SC25

µScm

−1

77.4(74.7–82.5)

54.5(53.4–55.8)

67.8(60.1–75.5)

56.3(50.8–59.8)

21.5(17.3–25.9)

161.6(119.4–203.7)

ANC

µeqL−1

−57(−57.3–−56.7)

−50.2(−60.8–−31.6)

−32.8(−41.5–−27)

6.6(0–17.5)

39.7(6–76.4)

1214.9(883.9–1756.4)

H+

µeqL−1

70(57.8–84.5)

56.4(29.2–78.4)

28.3(26.9–31)

4.6(1–8.4)

3.5(0.4–6.6)

0(0–0)

Ca2+

µeqL−1

47.8(38.7–54.5)

68.7(47.3–86.7)

208.7(189.6–219.1)

129.2(120–137.4)

60.9(34.2–76.9)

154.8(131.2–177.7)

Mg2+

µeqL−1

71.1(56–97.5)

35.3(20.7–61.9)

96.5(84.7–106.9)

174.3(125.1–263.4)

31.2(28.2–34.5)

1963.5(952.1–3309.4)

Na+

µeqL−1

42.5(39.5–47.6)

91.2(66.9–120.3)

108(95.7–116.1)

65.7(60.4–69.5)

61(38.4–82.5)

53.3(39.5–63.1)

K+

µeqL−1

18.5(17–21.1)

9.6(6.1–15.9)

12.2(10–14.3)

23.7(21.5–25.1)

5.7(4.1–7.9)

6.9(3.5–10.1)

SO2−4

µeqL−1

393.9(363.6–413.6)

241.3(163.3–296.9)

386.7(321.9–438.3)

304.3(287.8–324.7)

43(41–45.2)

298(222.3–350.1)

NO

− 3µeqL−1

7(4.8–10.4)

3.4(1.9–5.4)

20.7(17.9–23.7)

47.8(32.9–57.2)

19.5(9.4–32.4)

63.3(14.2–109.9)

Cl−

µeqL−1

36.7(33.9–39.8)

29.9(25.7–35.9)

30.5(26.2–37)

36.7(35.2–38.4)

22.5(17.5–27.1)

38.7(32.1–44.4)

F−

µeqL−1

7.6(6.6–8.2)

9.8(7.4–11.9)

8.9(7.4–10)

2.2(1.4–3.4)

1.9(1.7–2)

4.7(3.2–5.9)

TOC

mgCL−1

3.9(2.5–4.8)

12.2(7.2–15.2)

5.1(3.4–7)

3.2(1.5–5.7)

7.5(6.8–8.2)

11.1(5.6–15.7)

R-Al

µgL−1

1717(1530–1913)

513(448–550)

625(616–638)

112(52–221)

250(239–261)

36(12–59)

Mn

µgL−1

334.6(306.8–360.2)

98.9(54.9–146.4)

44.9

15(3.4–31.7)

10.3(9.5–11)

3.7(0.8–6.6)

Co

µgL−1

17.9(15.61–20.98)

1.02(0.6–1.42)

0.39

0.37(0.07–0.88)

0.12(0.11–0.13)

0.32(0.14–0.5)

Ni

µgL−1

6.28(5.34–7.02)

1.38(1.01–1.66)

0.79

3.11(2.59–4.14)

0.3(0.25–0.34)

83.47(52–114.4)

Cu

µgL−1

1.36(0.89–2.22)

0.82(0.4–1.05)

0.28

1.13(0.52–2.23)

0.73(0.35–1.12)

2.38(1.14–3.62)

Zn

µgL−1

45(43.1–47.2)

11.2(9.2–12.8)

49.5

13.6(11.1–18.1)

5.6(4.4–6.9)

1.4(0–4)

As

µgL−1

0.51(0.31–0.64)

2.21(0.93–3.15)

0.55

0.38(0.22–0.62)

0.96(0.81–1.12)

1.97(1.85–2.09)

Cd

µgL−1

0.95(0.87–0.99)

0.19(0.16–0.22)

0.3

0.2(0.18–0.22)

0.1(0.09–0.11)

0.02(0.01–0.04)

Pb

µgL−1

2.85(1.51–4.5)

1.64(0.72–2.2)

1.54

0.63(0.21–1.27)

1.88(1.59–2.17)

1.36(0.53–2.2)

Fe

µgL−1

176(95–303)

782(634–1073)

172

117(49–239)

340(269–411)

515(159–820)


logical samples and filtered through 40 µm inert mesh fab-ric.

The samples of benthic macroinvertebrates were takenby kicking technique (Frost et al. 1971) with a hand netof mesh-size 500 µm from 6 habitats 30 s giving adequateattention to all microhabitats. Then, the material was care-fully sieved through a 300 µm net, and preserved with an80% ethanol. This was supplemented by a 20–minute in-dividual picking off the submerged stones and woods. Sam-pling of pupal exuviae was performed by skimming the watersurface with a hand net (frame diameter 25 cm, mesh size250 µm), especially in accumulation areas and behind ob-stacles, along a distance of ca. 200–300 m (chironomid pupalexuviae technique – CPET, Wilson & Ruse 2005). Detaileddescriptions of chemical analytical methods and statisticalmethods are described in Horecký et al (2006).

Results

The basic results of hydrochemical analyses are shownin Table 2, and selected parameters are presented inFig. 2. According to the acidity status, the streams weredivided into 3 groups: strongly acidified (LIR, LYS,and JER with pH 4.07–4.57, reactive aluminium ormonomeric aluminium R-Al 448–1913 µg L−1); mod-erately acidified (LIL and JIZ with pH 5.08–6.38,R-Al 52–261 µg L−1); and non-acidified (PLB withpH 7.63–7.89, R-Al 12–59 µg L−1). The measured val-ues from the actual dates of benthic macroinvertebratesampling are consistent with data from a given hydro-logic year, if available (the intensively monitored sitesLIR, LYS, JIZ, and PLB). In our case, TOC (total or-ganic carbon) was a good estimation of DOC (dissolvedorganic carbon) because of the low content of particu-late organic matter in the stream water.Chironomid larvae dominated in strongly acidi-

fied LIR and LYS and acidification unaffected PLB,while at other sites the stonefly (Plecoptera) larvae pre-vailed. The dominance of chironomids and stoneflieswas more pronounced at the strongly acidified sites,where they represented 76 % of all organisms; in con-trast, at moderately acidified sites it was approx. 10 %less. Other groups except Oligochaeta and fly larvae(Diptera) were less represented or completely absent atthe strongly acidified sites. The strongly acidified sitesshowed smaller taxonomic richness of caddisflies (Tri-choptera) and an absence of mayflies (Ephemeroptera),crustaceans and molluscs (see Table 3).Differences between the groups of sites are also vis-

ible when evaluating the sites using ecological indices.The Shannon-Wiener diversity index incorporates bothspecies richness and equitability. The highest averagevalue was calculated for the site JIZ (3.09) and low-est for LYS (1.63), while the minimum actual valuewas calculated for the autumn collection of LIR (1.40)and maximum for summer collection of JIZ (3.28).The Simpson dominance index showed highest averagevalues for strongly acidified sites (0.17 to 0.22), low-est values for moderately acidified sites (0.06 to 0.14),and intermediate value for non-acidified PLB (0.15; seeFig. 3).

Table 3. The number of identified taxa (taxonomic richness) foreach group of benthic organisms at the observed sites.

Taxa/Site LIR LYS JER LIL JIZ PLB

Turbellaria 1 1 1 1Nematoda 1Oligochaeta 2 2 3 5 4 1Mollusca 1Hydracarina 1Crustacea 1Ephemeroptera 2 3 4Odonata 1Plecoptera 7 7 13 9 20 7Megaloptera 1 1Heteroptera 2 2 1 1Trichoptera 5 5 4 10 17 13Diptera excl. Chironomidae 10 8 12 8 10 12Chironomidae 4 4 4 5 5 5Coleoptera 8 3 1 4 6 11

Total 40 33 38 46 68 55

The core of the macroinvertebrate community inthe studied streams is quite similar. Widely representedmacroinvertebrates included the chironomid subfami-lies Chironominae, Orthocladiinae, and Tanypodinae,the stoneflies Leuctra nigra (Olivier, 1811), Nemurellapictetii Klapalek, 1900, and Protonemura sp., the cad-disflies Plectrocnemia conspersa (Curtis, 1834) andLimnephilidae juv., the flies Dicranota sp., and lar-val beetles of the genus Agabus. Exclusively in thestrongly acidified streams, there occurred a group ofnon-pigmented planarians (Phagocata sp. and Dendro-coelum sp.), the alder fly Sialis fuliginosa Pictet, 1836and the caddisflyMicropterna nycterobia Mac Lachlan,1875. Mayflies of the families Baetidae [Baetis rhodani(Pictet, 1843–1845), Baetis vernus Curtis, 1834, andCentroptilum luteolum (Muller, 1776)], Leptophlebi-idae [Habrophlebia lauta Eaton, 1884 and Leptophlebiamarginata (L., 1767)], Siphlonuridae [Siphlonurus la-custris (Eaton, 1870)], and Ameletidae (Ameletus in-opinatus Eaton, 1887), the crustacean Niphargus sp.,and the bivalve mollusc Pisidium casertanum (Poli,1791) were present exclusively at sites with pH above5. The oligochaete species Stylodrilus heringianus Cla-parede, 1862, the caddisflies Apatania fimbriata (Pictet,1834), Sericostoma personatum (Spence, 1823), gen-era Rhyacophila, Drusus, and Potamophylax and thechironomid subfamilies Diamesinae and Prodiamesinaewere missing in the strongly acidified streams. Taxathat preferred moderately acidified and non-acidifiedsites, although in low abundance in samples fromstrongly acidified streams, were for example Diura bi-caudata (L., 1758), Chelifera sp. or Eloeophila sp.In the Principal Component Analysis (PCA), the

first and second gradient (axes) explained 40.9% and24.2% of the biological variability of the studied sites,respectively. The benthic macroinvertebrate structureof the strongly acidified sites is similar (Fig. 4). The firstgradient, which shows the biggest difference in biologi-cal variability of the samples, is formed mainly by the


Fig. 3. Shannon-Wiener diversity index (H′) and Simpson dominance index (Ds) of the strongly acidified (JER, LIR, LYS), moderatelyacidified (JIZ, LIL) and non-acidified (PLB) sites.

Fig. 4. Results of the Principal Component Analysis (PCA). The squares represent the strongly acidified sites (JER, LIR, LYS),triangles represent the moderately-acidified (reference) sites (JIZ, LIL) and the circle represents the non-acidified site (PLB). Underthe designation JIZ-SPEC and PLB-SPEC are taxa found at those sites only:JIZ-SPEC: Nematoda g.sp, Tubificidae g.sp., Hydracarina g.sp., Niphargus sp., Ameletus inopinatus, Siphlonurus lacustris, Perlodidaeg.sp., Siphonoperla torrentium, Siphonoperla sp. juv., Brachyptera seticornis, Nemoura cinerea, Protonemura intricata, Leuctra aurita,Leuctra digitata, Leuctra hippopus, Leuctra inermis, Leuctra teriolensis, Rhyacophila sk. dorsalis, Apatania fimbriata, Drusus sp. juv.,Drusus discolor, Allogamus auricolis, Chaetopterygopsis maclachlani, Prosimulium tomosvaryi, Tabanidae g.sp, Prodiamesa olivacea,Limnius sp.PLB-SPEC: Baetis rhodani, Centroptilum luteolum, Leptophlebiidae g.sp., Nemoura cambrica, Rhyacophila praemorsa, Plectrocnemiacf. geniculata, Tinodes rostocki, Crunoecia irrorata, Sericostoma personatum, Dixa sp., Thaumalea sp., Dolichopus sp., Hydraenabritteni, Hydraena sp., Limnebius sp., Limnius perrisi

taxa present at moderately acidified sites JIZ and LIL[e.g., Drusus annulatus (Stephens, 1837), Rhyacophilapolonica Mac Lachlan, 1879, Stylodrilus heringianus,Chaetopteryx villosa (F., 1798), Leuctra pseudocingu-lata Mendl, 1898 and Diura bicaudata] or at JIZ only(see legend of Fig. 4). The second gradient is com-posed mainly by the species occurring only at PLBand by differences in the abundance of some beetlegenera [Elodes sp., Limnius sp., Deronectes platyno-tus (Germar, 1834), Agabus sp. juv.], the chironomid

subfamily Diamesinae, Ceratopogoninae, the stoneflyNemurella pictetii, the caddisfly Plectrocnemia con-spersa, and the dagger fly Wiedemannia sp. (Empidi-dae).The environmental variables can be projected on

to the results of the PCA based on macroinvertebratestructure (indirect analysis). The strongest correlationwith the first gradient was shown by the sulphate con-centration, turbulence, and forest cover at the sites. Thesecond gradient was mostly correlated with pH and de-


pendent concentrations of R-Al and some metals (Zn,Cd, Mn, Mg, Cu).

Discussion

The macroinvertebrate sampling at the outlet of thebasins occurred during three different stages of runoff:snow-melt (April), autumnal base-flow (October) andsummer saturation with moderate direct-flow (July).The observed hydrochemistry at JIZ, LYS, LIR, andPLB corresponds to the runoff regime, with a markeddecrease of pH (and its consequences) with increasingrunoff volume; for LIL and JER we expect a similartrend.The water chemistry of strongly acidified streams

LIR, JER, and LYS reflects the leaching of sulphatethat accumulated in soils during former high depo-sitions (Kopáček et al. 2002), acid-sensitive bedrock,and the acidification effects of spruce monocultures. Atthese sites, the water pH was between 4.07–4.57 andalkalinity was negative. This shows an exhausted acidneutralizing capacity and such streams are consideredpermanently strongly acidified (Braukmann 2001). Thesites LIL and JIZ are characterized by large fluctua-tions in pH, and after the Braukmann classification fallinto the group of periodically acidified streams withacid episodes during snowmelts or long-term intensiverains. Occurring on comparable acid-sensitive bedrockwith slow weathering rates, the actual pH of these twostreams may be similar to their natural status. There-fore these sites could reflect a reference status for thestrongly acidified streams, and suggest the maximumrecovery potential. The site PLB has different bedrock(non-sensitive to acidification), and so is a non-acidstream with pH averages between 7 and 8 only sporad-ically falling to values around 6.8 (Krám et al. 2012),though having high concentrations of Mg, Cu and Ni.Elevated concentrations of R-Al were recorded in

all three strongly acidified streams. An important fac-tor affecting the toxicity of elevated concentrations ofaluminium to aquatic organisms is the form, in whichit is present in stream water. If there is enough DOCin water, toxic aluminium ions can be bound by the or-ganic complexes (Havas & Rosseland 1995) leading to asubstantial reduction of toxicity (Fott et al. 1994). How-ever, at the streams investigated here this process waslikely not significant, since even the highest recordedconcentrations of organic carbon were lower than thelevel claimed for such biotic responses (Kullberg 1992).At the strongly acidified streams, elevated concentra-tions of other toxic metals (Mn, Co, Zn, Cd), especiallyat LIR, were found, in accordance with the literature(e.g., Hermann et al. 1993; Lampert & Sommer 1997).Very high concentrations of copper and nickel at PLBare caused by specific geological conditions of the site– the bedrock is composed of serpentinite, which is anultramafic metamorphic rock composed mainly of mag-nesium silicate with elevated rates of chemical weath-ering, and this leads to a faster release of base cationscompared to less weatherable granite (Hruška & Krám

Fig. 5. Streamwater chemistry and forest change at the Sklářskýpotok – Jizerka catchment. The catchment clear-cut (in %) isrepresented by dashed line and the streamwater pH and sulphates(in mg L−1) by black line and grey line, respectively.

2003). This also explains the extremely high concentra-tion of magnesium, high alkalinity and pH above 7 thatwere recorded at this site.Very low concentrations of sulphate at JIZ com-

pared to other sites can be explained as a consequenceof the low current proportion of forest in this catch-ment. Křeček et al. (2006) reported a recovery in waterquality for this stream, including an increase of meanannual pH values from 4 to the range 5–6 and a drop inaluminium concentrations to the range 0.2–0.5 mg L−1.However, this improvement can be explained not onlyby the reported decreased air pollution since 1990s, butalso by the stabilization of forested wetlands by grasscover after clear-cutting of spruce stands between 1984and 1990 (Křeček et al. 2006). It is well known thatnot only acid atmospheric deposition but also intensiveforestry contributes to the anthropogenic acidificationof ground and surface waters (Weatherley et al. 1989).Forests, especially coniferous, greatly increase deposi-tion (both horizontal wet and dry) and thus the sup-ply of sulphate ions into the catchment (Wilpert et al.1996), and can significantly slow down or prevent theprocess of recovery from acidification (Hruška & Krám2003; Kopáček et al. 2002). In addition, the uptake ofbase cations by trees from the soil plays a very impor-tant role, which can influence the acidity of freshwaters,especially in the case of intensive forestry (Bredemeier1988; Binkley & Giardina 1998). The rapid water qual-ity recovery at the JIZ catchment (see Fig. 5) probablyreflects the synergetic effect of all the above-mentionedfactors together with the fact that deforestation startedalready one decade before air pollution started to de-cline. The catchments of the remaining streams aremore markedly forested and thus the streamwater con-centration of sulphate at these sites was much higher.Decreased species diversity is a well-documented

effect of acid atmospheric deposition to aquatic biota.It is caused by the fact that organisms are negativelyinfluenced by the toxicity of H+ ions or ionic forms oftoxic metals (especially Al), problems with ion regula-tion, corrosion of calcium from shells, changes in foodavailability, etc. (e.g., Herrmann et al., 1993). The mostaffected taxonomic groups are mayflies, crustaceans and


molluscs, which are usually missing in acidified waters;nevertheless other families such as stoneflies and cad-disflies are also severely reduced in the number of sur-viving species or genera. Streams which are affected byacidification tend to have higher index of dominancecompared to non-acidified streams, while the Shannon-Wiener diversity index is higher at non-acidified sites(Scheibová & Helešic 1999). These effects on aquaticbiota are in concordance with the results presented here(see Fig. 3). The same relationship between low waterpH and Shannon-Wiener diversity index was also no-ticed by Niyogi et al. (2002) who studied influence ofacid drainage waters on primary producers.Mayfly nymphs were only found at the moderately

and non-acidified sites, which is not surprising giventheir proven sensitivity to low pH (Raddum et al. 1988;Hermann et al. 1993). The mayfly Ameletus inopinatusprobably does not occur at LIL (and in the Brdy Mts atall), because it inhabits streams above 750 m a.s.l. (Sol-dán et al.1998). The absence of the other two mayfliesat this site could be due to low alkalinity compared toJIZ as it is considered a similarly important parameteras pH (Braukmann 2001).Molluscs were represented by a single species, Pi-

sidium casertanum, and only at the sites LIL and PLB.This species is considered to be the most acid-tolerantmollusc and occurs in waters with low calcium contentdown to 2.7 mg L−1, respectively 135 µeq L−1 (Horsák& Hájek 2003). The concentration of calcium at the siteJIZ, however, is so low that it does not even meet thesecriteria.The dominant component of benthic macroinverte-

brates in the forest streams with average pH above 7 isoften formed by the crustacean Gammarus (e.g., Dan-gles & Guérold 2000). At PLB, the absence of thesecrustaceans, together with the limited occurrence ofmolluscs, is apparently caused by extreme concentra-tions of magnesium, which can be toxic to inverte-brate organisms, especially at low calcium concentra-tions (Camilleri et al. 2003).In mountain streams, and especially acidified ones,

the stoneflies (Plecoptera) usually form the dominantgroup of benthic macroinvertebrates (Guérold et al.1995; Szczesny 1998). Besides the abundant repre-sentatives of the families Nemouridae and Leuctridae(mainly Leuctra nigra) the predatory stonefly Diura bi-caudata is also highly resistant to acidification (Brauk-mann 2001). Consistent with previous results (Horeckýet al. 2006), this species was recorded only at siteswith pH higher than 4.5. The very low stonefly diver-sity at PLB, even in comparison with strongly acidifiedstreams, can also be explained by the extreme concen-trations of magnesium.In addition to the acid-tolerant species Plectrocne-

mia conspersa (Scheibová & Helešic 1999), caddisflies(Trichoptera) are also often represented by the fam-ily Limnephilidae in acidified streams (Guérold et al.1995; Braukmann 2001). Especially some species of thetribes Chaetopterygini and Stenophylacini [e.g., Pseu-dopsilopteryx zimmeri (Mac Lachlan, 1876) and Alloga-

mus uncatus (Brauer, 1857)] are very resistant to lowpH. On the other hand, the sensitive representatives ofthe family Limnephilidae (Drusus annulatus and Ap-atania fimbriata) and Rhyacophilidae were present onlyin the reference (moderately acidified) streams. Appar-ently it is the combination of low pH, elevated con-centrations of R-Al and heavy metals and low concen-trations of dissolved organic and inorganic matter thathad adverse effect on these caddisflies. The occurrenceof Micropterna nycterobia at LIR could be evidence ofepisodic occasional drying of the stream, because thisspecies is often found in intermittent waters.Chironomids are a major component of the macro-

invertebrate fauna in all freshwater ecosystems. Theyare considered to be good indicators of water ecosystemquality as they respond to different environmental andanthropogenic disturbances. Chironomid fauna in acid-ified streams has been much less studied than in lakes,but it is known that chironomid assemblages becomevery simplified, with only resistant species able to copewith the acid conditions (Lindegaard 1995). Changesin chironomid assemblages reflect rather the changes,for example, in food resources (e.g., Allard & Moreau1987; Rasmussen & Lindegaard 1988) and in intraspe-cific relationships (e.g., de Bisthoven et al. 2005) thanthe direct physiological effects of low pH.In this study, significant changes in chironomid

fauna could be expected due to the strong pH gradientamong investigated streams. However, since the larvaewere identified to the family level only, crucial infor-mation on the effects of acidification on chironomid as-semblages are lacking, as different species and/or gen-era can react to acidification stress differently. Furtherresearch is needed to examine the potential of Chirono-midae as bioindicators of different acid conditions inthe investigated streams and the recovery process fromacidification.Nine chironomid species/taxa were identified from

the pupal exuviae material taken from LYS and PLB:Zavrelimyia signatipennis (Kieffer, 1924), Corynoneuracf. coronata Edwards, 1924,Corynoneura Pe2a Langton1991, Eukiefferiella brevicalcar (Kieffer, 1911), Krenos-mittia boreoalpina (Goetghebuer, 1944), Parametri-ocnemus boreoalpinus Gowin et Thienemann, 1942,Parametriocnemus Pe1 Langton 1991, Micropsectraaristata Pinder, 1976, Tanytarsus heusdensis Goetghe-buer, 1923. Two species, Zavrelimyia signatipennis andMicropsectra aristata, both collected from PLB, wererecorded firstly in the Czech Republic. Most recordedspecies are known to be cold-stenothermic, preferringspring brooks and montane streams. The pupal exuviaematerial was rather poor; but nevertheless it resulted inmore detailed and valuable information on chironomidsliving in the extreme conditions studied.The rich fauna of adult beetles at strongly acidified

LIR compared to other sites can be explained by thefact that they breathe atmospheric oxygen and are pro-tected by a solid wing case (Havas & Rosseland 1995).On the other hand, Elodes (Scirtidae) and Limnius(Elmidae), which were recorded only at the moderately


acidified JIZ and non-acidified PLB, are considered lessacid-resistant genera (Braukmann 2001).The reduction of acid atmospheric deposition has

allowed chemical and biological recovery processes ofwaters from anthropogenic acidification to begin. Agradual return of acid-sensitive species is expected withchanges in water chemistry, particularly pH increasesand R-Al concentration decreases, as reported, e.g., byTipping et al. (2002) and Raddum & Fjellheim (2003).Marked decreases of sulphate and calcium concentra-tions have been observed at the strongly acidified sites,though the values of pH and aluminium concentrationhave not yet changed significantly. This suggests thatthe process of biological recovery has not yet started,which is supported by the results of benthic macroin-vertebrate analyses given here. No species consideredby Braukmann (2001) as acid-sensitive and only fewspecies considered partially acid-tolerant (third classacid) were found in the strongly acidified sites dur-ing the studied period. A comparison of the benthiccommunity structure found at LIR with the results ofprevious studies (Horecký et al. 2006; Hardekopf et al.2008) also does not show any significant difference. Thenewly found taxa at the site LIR [e.g., Notonecta glaucaL., 1758, Hybomitra sp. Anacaena lutescens (Stephens,1829)] can hardly be considered as evidence of a com-munity recovery from acidification.In contrast, there are reasonable grounds to believe

that recovery of the benthic macroinvertebrate com-munity has already started at JIZ. Both the clear-cutof mature spruce plantations (1984–1990) and the re-gional drop in SO2 emissions (particularly in the 1990s),resulted in the decline of acid deposition and risingstreamwater pH at this site. Mean annual pH at JIZincreased from 4.0 in years 1982–1985 to 5.3 in years1990–1994 (Křeček & Hořická, 2001). Species such asthe mayflies Baetis vernus and Ameletus inopinatusand stoneflies Siphonoperla torrentium (Pictet, 1841)and Brachyptera seticornis (Klapálek, 1902), along withsome other there found taxa, do not survive in waterswith long-term pH values below 5 (Braukmann 2001;Raddum et al. 1988; Scheibová & Helešic 1999; Krno etal. 2006); therefore, the JIZ site had to be recolonizedby acid-sensitive species already in the 1990s. This alsocorresponds with the results of a faunistic study fromthe years 1997–2000 (Preisler & Špaček 2001), whichalready found imagos of the above listed stoneflies inthis stream.Based on these and previously published results

(Horecký et al. 2002, 2006; Hardekopf et al. 2008), areturn of some species that are less tolerant to streamacidity to the strongly acidified sites may be expected,if pH increases at least over 4.5. In poorly mineral-ized mountain streams these could be e.g. the stone-flies Diura bicaudata, Leuctra major, L. pseudocingu-lata or L. pseudosignifera Aubert, 1954 and caddisfliesof the genera Drusus or Rhyacophila. Published resultsof dynamic modelling of the processes in strongly acidi-fied catchments (Hruška & Krám 2003; Hardekopf et al.2008) do not predict streamwater pH values which are

necessary for the return of acid sensitive taxa such asthe mayflies Baetidae and Siphlonuridae, or the bivalvemollusc Pisidium.These results indicate that at the strongly acidified

sites LIR and LYS, the process of biological recovery hasnot yet begun due to insufficient increases in streamwa-ter pH. In contrast, reduced load of acidity initiated byextended clear-cuts of spruce plantations at JIZ in the1980s and further enhanced by the large reduction ofemissions in the 1990s resulted in a drop of acid deposi-tion and a significant increase in mean streamwater pH(approximately one unit, from 4 to 5, over five years).These factors support the opportunity for the recov-ery of acid-sensitive species even in environment withacid-vulnerable bedrock within a couple decades.

Acknowledgements

This research was supported by the Czech Science Foun-dation (GACR grant No. 526-09-0567), the EnvironmentProject of the European Commission, EURO-LIMPACS(GOCE-CT-2003-505540), and the Ministry of the Environ-ment of the Czech Republic in the framewok of the Conven-tion on Long-Range Transboundary Air Pollution (LRTAP).In addition, we would like to thank Dr. David Hardekopf forrevision of the English text and colleagues Dr. Pavel Chvo-jka, Edita Šípková, MSc. Martin Fikáček, and Dr. Petr Pařilfor their contribution to this research.

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Received May 2, 2012Accepted November 10, 2012

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Differences in benthic macroinvertebrate structure of headwater streams with extreme hydrochemistry