Biologia, Bratislava, 61/Suppl. 18: S181—S184, 2006 Section Zoology DOI: 10.2478/s11756-006-0129-2

Macrozoobenthos of two different catchment areas of the Tatra Mountain lakes with a special reference ontheeffectsofacidification

Iľja Krno

Department of Ecology, Faculty of Sciences, Comenius University, Mlynská doolina B-2,SK-84215 Bratislava, ; e-mail: [email protected]

Abstract: The structure of macrozoobenthos of two different high-mountain Tatra lakes, namely Starolesnianske pleso (strongly acidified) and Nižné Terianske pleso (reference site) is described. The latter is characterised by a large catchment and total area, and is relatively deep with strong inlet and outlet. It is not affected by acidification. Univoltine species of mayflies, stoneflies and caddisflies dominate in the littoral, mainly scrapers. Predators are represented by stoneflies. Shredders dominate in the lake outlet. From the zoogeographic point of view, Carpathian endemites and Central European species prevail. Forty percent of species are sensitive to acidification. The former has a smaller catchment and total area, and is relatively shallow, without inlet. It is strongly affected by acidification. Semivoltine species, mainly predators () and shredders prevail. From the zoogeographic point of view Palaearctic species dominate. Species sensitive to acidification are missing. Emergence of water has two peaks during spring and fall circulation. Key words: Mayflies, stoneflies, caddisflies, beetles, zoogeography, trophic guilds, emergence, phenology, mountain lakes, acidification, High Tatra Mts, Slovakia.

Introduction els for acidity, pollutant flux and climate variability. The aim of this contribution is a comparative The macrozoobenthos of the Tatra Mountain lakes, as study of mayflies, stoneflies, caddisflies and aquatic bee- well as of their inlets and outlets, has been the sub- tles inhabiting alpine aquatic ecosystems in two sig- ject of interest of many entomologists and hydrobiol- nificantly different and naturally oligotrophic moun- ogists. Surveys on these works are published in the tain lakes – Starolesnianske pleso (strongly acidified) publications of Krno (1988a, 1991a, b). The Tatra and Nižné Terianske pleso (reference site). Attention Mountain lakes were negatively affected by acidifica- was focused on the emergence of imagoes from the lit- tion at the end of the last century (Stuchlík et al., toral and the adjacent lake outlet, and the influence of 1985). Acidification significantly affects the structure temperature regimen. New knowledge on macrozooben- of macrozoobenthos in the lake littoral (Krno, 1988b, thos distribution in the littoral of these two lakes was 1991a). A decrease in pH below 5.5 results in the re- introduced by the studies of Kasprzak & Šporka lease of phosphorus from sediments (Stuchlík et al., (1991) (shellfish), Kodada (1990), Chvojka (1992) 1993; Vyhnálek et al., 1994), with a subsequent in- (caddisflies), Zaťovičová (2002) (macrozoobenthos), crease in lake trophics and the growth of macrozooben- Hamerlík (2004) and Bitušík (2004) (chironomids). thos biomass (Krno, 1991a) in strongly acidified lakes Bitušík (1994) and Bitušík & Kubovčík (1999) such as Starolesnianske pleso, Sesterské pleso, and oth- analysed the occurrence of subfossil chironomids in sed- ers. In non-acidified alpine lakes there are acidosensi- iments of Starolesnianske pleso and Nižné Terianske tive species, like Crenobia alpina (Dana, 1766), Amele- pleso, respectively. tus inopinatus, Electrogena lateralis (Curtis 1834), Ar- cynopteryx compactata, Diura bicaudata, Capnia vidua, Characteristics of monitored sites Apatania fimbriata (Pictet, 1834), and Drusus trifidus. Only two species, namely Crenobia alpina and Capnia Starolesnianske pleso is situated at an altitude of 2,000 vidua, survive in acid-sensitive lakes. These species do m a.s.l. and has an area of 0.71 ha, a maximum depth not penetrate to acidified mountain lakes. of 4.2 m and average annual pH is 4.9. It lies on gra- The project MOLAR (Štefková & Šporka, nodiorites and has phosphorus (P) concentrations > 7 2001) allowed detailed limnological studies on remote µgL−1.There is an increased amount of toxic metals mountain lakes with an aim to develop predictive mod- (Al, Cd, Pb, Hg); maximum water temperature in the

c 2006 Institute of Zoology, Slovak Academy of Sciences S182 I. Krno

Table 1. Littoral macrozoobenthos of Starolesnianske pleso in 1993–1997.

Species Taxonomic Habitat Faunistical Life cycle Food guild Acidification Dominance group element number (%)

Nemurella pictetii Klapálek,1900 Plecoptera littoral Pa Sv shredder 0 28 Limnephilus coenosus Curtis, 1834 Trichoptera littoral Pa Sv shredder 0 40 Agabus bipustulatus (L., 1761) Coleoptera littoral Pa Sv predator 0 22 palustris (L., 1761) Coleoptera littoral Pa Sv predator 0 5 Hydroporus incognitus Sharp, 1869 Coleoptera littoral Pa Sv predator 0 2 Hydroporus ferrugineus Stephens, 1827 Coleoptera littoral Pa Sv predator 0 3

Key: Pa – Palaearctic; Sv – semivoltine.

12 sect imagoes were caught by means of sweeping from vegeta- Air tion, and gathered from the surface and from beneath rocks 10 Water surface on the lake shore. Additionally, in 1997 they were regularly caught in six formaldehyde traps (yellow Moericke dishes, 10 8 × 10 cm) that were exposed in the field for 3–4 weeks. The

C relative abundance of adults caught was calculated for three- o 6 weeks interval of three dishes. Water and air temperature 4 (Fig. 1) at Nižné Terianske pleso were regularly recorded in half-hour intervals by an automatic weather station (AWS) 2 (ŠPORKA et al., 2002). Temperature

0 Results and discussion -2 V VI VII VIII IX X XI -4 In Starolesnianske pleso, increased sedimentation con- nected with soil runoff into this shallow lake in the Fig. 1. Seasonal variation in mean weekly air and water temper- past changed its initial oligotrophic status before 1930 ature at Nižné Terianske pleso in 1997. (recession of species of the genera Corynoneura, Eu- kiefferiella, Drusus). Simultaneously, acidification took place, resulting in a significant change in the zoo- littoral was 14 ◦C during the study period. The amount plankton structure (Stuchlík et al., 1985) and pro- of calcium is (Ca) < 1mgL−1, total retention time fundal chironomid assemblages (Bitušík, 1994). The is only 75 days. The proportion of alpine meadows to littoral was characterized by a high biomass of ben- rocky moraines is 3 : 1 within the total catchment area. thic organic material and macrozoobenthos. The moun- The lake has no inlet and outlet sometimes dries during tain lake is characterized by widely spread Palaearctic summer. There is relatively rapid sedimentation rate, species showing more than one year larval development more than 1 mm within two years. The lake is strongly (during the whole study, these species were represented acidified (Stuchlík et al., 1993). by different cohorts of larvae whose imagoes emerged in Nižné Terianske pleso is situated at an altitude of different years). Detritivores and predators dominated 1,941 m a.s.l. and has an area of 4.83 ha, a maximum (Tab. 1). Species which had an acidification number depth of 44.4 m and pH 6.9. It also lies on granodiorites higher than zero did not occur there (Raddum et al., and has a maximum water temperature in the littoral 1988; Krno, 1991b). of 10 ◦C. The amount of Ca is > 3mgL−1, total P 2 µg On the contrary, rare Carpathian endemic species L−1. Retention time in the lake is more than a half-year. and Central European mountain species with one-year The proportion of alpine meadows to rocky moraines is development were typical of the catchment of the non- 1 : 1. The lake has both an inlet and an outlet. It is acidified Nižné Terianske pleso. The semivoltine Diura a classical alpine non-acidified oligotrophic lake. There bicaudata species has one-year diapause of eggs and sub- is a characteristically low biomass of benthic organic sequent one-year larval development. Shredders domi- material in the littoral. Sedimentation is very slow in nated in the littoral while detritivores dominated in the the lake, 1 mm per two years (Šporka et al., 2002). inlet and outlet. In contrast to Starolesnianske pleso, species with acidification number higher than zero, oc- curred there (Raddum et al., 1988; Krno, 1991b) Material and methods (Tab. 2). After thawing of the lake edges in June, Macrozoobenthos was collected from the littoral in July or spring species such as Capnia vidua and Acrophylax August 1993–1996 by means of the kicking method (FROST, vernalis (its pupae actively swim to holes in the ice) 1971). In 1997, samples were collected at regular monthly emerged out of the water. Females of A. vernalis lay intervals in the period from May to November. Aquatic in- their eggs in dry moss on the lake shore. Emergence Zoogeography and phenology of the Tatra lakes macrozoobenthos S183

Table 2. Macrozoobenthos of the littoral, inlet, and outlet of Nižné Terianske pleso in 1993–1997.

Species Taxonomic Habitat Faunistical Life Food guild Acidification Dominance group element cycle number (%)

Protonemura brevistyla (Ris, 1902) Plecoptera inlet CE Uvs shredder 0 3 Leuctra rosinae Kempny, 1900 Plecoptera inlet CE Uvs gather 0 53 Capnia vidua Klapálek, 1904 Plecoptera inlet Aa Uvsp shredder 0.25 6 Drusus monticola McLachlan, 1876 Trichoptera inlet CE Uvs scraper 0.5 28 Allogamus starmachi Szcz˛esny, 1967 Trichoptera inlet C Uva scraper 0 10 Ameletus inopinatus Eaton, 1887 Ephemeroptera outlet Aa Uvs scraper 0.5 6 Protonemura brevistyla (Ris, 1902) Plecoptera outlet CE Uvs shredder 0 5 Leuctra pusilla Krno,1985 Plecoptera outlet C Uvs gather 0 3 Capnia vidua Klapálek, 1904 Plecoptera outlet Aa Uvsp shredder 0.25 10 Diura bicaudata (L., 1758) Plecoptera outlet Saa Uvs predator 0.5 22 Drusus discolor (Rambur, 1842) Trichoptera outlet CE predator 0.5 1 Allogamus starmachi Szcz˛esny, 1967 Trichoptera outlet C Uva scraper 0 10 Chaetopteryx polonica Dzi˛edzielewicz, 1889 Trichoptera outlet C Uva shredder – 4 Prosimulium latimucro (Enderlein, 1925) Diptera outlet CE Uvs filterer 0 37 Orimargula alpigena Mik, 1883 Diptera outlet – predator – 1 Ameletus inopinatus Eaton, 1887 Ephemeroptera littoral Aa Uvs scraper 0.5 12 Leuctra rosinae Kempny, 1900 Plecoptera littoral CE Uvs gather 0 1 Capnia vidua Klapálek, 1904 Plecoptera littoral Aa Uvsp shredder 0.25 22 Arcynopteryx compacta (McLachlan, 1872) Plecoptera littoral Aa Uvs predator 0.5 1 Diura bicaudata (L., 1758) Plecoptera littoral Saa Uvs predator 0.5 9 Drusus trifidus McLachlan, 1868 Trichoptera littoral CE UVS scraper 0.5 6 Allogamus starmachi Szcz˛esny, 1967 Trichoptera littoral C Uva scraper 0 41 Acrophylax vernalis Dzi˛edzielewicz, 1912 Trichoptera littoral C Uvsp scraper – 8

Key: Aa – Arcto-Alpine, Saa – Subarctic-alpine, C – Carpathian, CE – Central European; life cycle: Sv – semivoltine, Uvsp – univoltine (spring emergence), Uvs – univoltine (summer emergence), Uva – univoltine (autumnal emergence).

Fig. 2. Seasonal variation in length of adults (mm) of selected species in Nižné Terianske pleso in 1997. of both species suddenly ended in the second half of the second half of August – the period of the highest wa- July. The size of the last emerging imagoes (Fig. 2) was ter and air temperatures (Fig. 3). The autumn species similar to those which emerged in May, and significantly Chaetopteryx polonica and especially Alogammus star- smaller than imagoes emerging in mid-June. Vannote, machi emerged until the lake was frozen (November). & Sweeney (1980) indicated that aquatic insect ima- The number of adults emerging in the warm period goes reach their maximum size under optimum tem- of summer stagnation was lower than during the sig- perature conditions during their larval development. nificant autumn drop of water and air temperatures Other species were summer ones, with the exception of and subsequent autumn circulation, and also in partic- Chaetopteryx polonica and Alogammus starmachi,and ular during the period of melting and spring circula- the maximum emergence of their imagoes occurred in tion (Figs 2, 3). Zaťovičová (2002) recorded a simi- S184 I. Krno

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Received June 2, 2005 Accepted May 9, 2006