Structure of invertebrate communities in oak ...

Ekológia (Bratislava)

Vol. 24, Supplement 2/2005

Structure of invertebrate communities in oak-hornbeam forest ecosystem

Published with the financial support of VEGA (Scientific Grant Agency of the Ministry of Education and the Slovak Academy of Sciences), grant number 1/0119/03 Contributors: Comenius University in Bratislava, Faculty of Natural Sciences, Department of Zoology, Department of Pedology, Department of Ecosozology Slovak Academy of Sciences in Bratislava, Institute of Zoology Technical University in Zvolen, Faculty of Ecology and Environmental Sciences, Department of Biology and General Ecology Palacký University in Olomouc, Faculty of Science, Department of Ecology and Environmental Science State Phytosanitary Administration, Jičín Masaryk University in Brno, Faculty of Medicine and Faculty of Natural Sciences, Centre for Biostatistics & Analyses Special editor: Milada Holecová

ECOLOGY

(Bratislava)

International Journal for Ecological Problems of the Biosphere

CONTENTS Zlinská J., Šomšák L., Holecová M.: Ecological characteristics of studied forest communities of an oak-hornbeam tier in SW Slovakia . . . . . . . 3 Tirjaková E., Vďačný P.: Ciliate communities (Protozoa, Ciliophora) in treeholes and influence of selected environmental factors on their structure . 20 Bartošová P., Tirjaková E.: Selected ecological characteristics of ciliate communities (Protozoa, Ciliophora) in decaying wood mass in the Malé Karpaty Mountains . . . . . . . . . . . 37 Mrva M.: Diversity of active gymnamoebae (Rhizopoda, Gymnamoebia) in mosses of the Malé Karpaty Mts (Slovakia) . . . . . . . . . . . . . . . . . 51 Degma P., Šimurka M., Gulánová S.: Community structure and ecological macrodistribution of moss-dwelling water bears (Tardigrada) in Central European oak-hornbeam forests (SW Slovakia) . . . . . . . . . . . . . . . . . . 59 Christophoryová J., Krumpál M.: Communities of pseudoscorpions (Pseudoscorpiones, Arachnida) in epigeon of oak-hornbeam forests in the Malé Karpaty Mts and Trnavská pahorkatina hills (SW Slovakia) . . . . . . . . 76 Krumpálová Z.: Epigeic spiders (Araneae) in ecosystems of oak-hornbeam forests in the Malé Karpaty Mts (Slovakia) and their ecological characterisation . . . . . . . . . . . . . 87 Fenďa P., Ciceková J.: Soil mites (Acari, Mesostigmata) of oak forests in the Malé Karpaty Mts (W Slovakia) . . . 102 Tuf I.H., Tufová J.: Communities of terrestrial isopods (Crustacea: Isopoda:

EKOLÓGIA

(Bratislava)

Medzinárodný časopis pre ekologické problémy biosféry

OBSAH Zlinská, J., Šomšák L., Holecová, M.: Ekologická charakteristika študovaných lesných spoločenstiev dubovo-hrabového vegetačného stupňa v.oblasti JZ Slovenska . . . . . . . . . 3 Tirjaková E., Vďačný P.: Spoločenstvá nálevníkov (Protozoa, Ciliophora) v.dendrotelmách a vplyv vybraných environmentálnych faktorov na ich štruktúru . . . . . . . . . . . . . . . . . . 20 Bartošová P., Tirjaková E.: Vybrané ekologické charakteristiky spoločenstiev nálevníkov (Protozoa, Ciliophora) v.odumretej drevnej hmote na území Malých Karpát . . . . . . . . . . 37 Mrva M.: Diverzita nahých meňaviek (Rhizopoda, Gymnamoebia) v.machoch Malých Karpát (Slovensko) . 51 Degma P., Šimurka M., Gulánová S.: Štruktúra spoločenstva a ekologická makrodistribúcia pomaliek (Tardigrada) žijúcich v.machoch stredoeurópskych dubovo-hrabových lesov (JZ Slovensko) . . . . . . . . . . . . . . 59 Christophoryová, J., Krumpál, M.: Spoločenstvá šťúrikov dubovo-hrabových lesov Malých Karpát.a Trnavskej pahorkatiny . . . . . . . . . . . . . . . . 76 Krumpálová Z.: Epigeické pavúky (Araneae) v.ekosystémoch dubovo-hrabových lesov Malých Karpát (Slovensko) a ich ekologická kategorizácia . . . . . . . . . . . . . . . . . . 87 Fenďa P., Ciceková J.: Pôdne roztoče (Acari, Mesostigmata) dubových lesov Malých Karpát (Západné Slovensko) 102 Tuf I.H., Tufová J.: Společenstva stejnonožců (Crustacea: Isopoda) v.epigeonu dubo-habrových lesů JZ Slovenska . . . . . . . . . . . . . . . . . 113

Országh I., Országhová Z.: Structure of centipede communities (Antennata: Chilopoda) in oak-hornbeam forests of the Malé Karpaty Mts and Trnavská pahorkatina hills (SW Slovakia) . . . Stašiov S.: Millipede communities (Diplopoda) of oak-hornbeam ecosystems (the Malé Karpaty Mts, Trnavská pahorkatina hills, SW Slovakia) . . . Országh I.: Phenology of Chelidurella acanthopygia (Dermaptera: Forficulidae) nymphal instars in oak-hornbeam forests of the Malé Karpaty Mts and Trnavská pahorkatina hills (SW Slovakia) . . . . . . . . . . . . . . . . . . Hradil K.: Bug assemblages in epigeon of.oak-hornbeam forests in SW Slovakia . . . . . . . . . . . . . . . . . . Holecová M., Némethová D., Kúdela M.: Structure and function of weevil assemblages (Coleoptera, Curculionoidea) in epigeon of oak-hornbeam forests in SW Slovakia . . . . . . . . . Holecová M., Krumpál M., Országh I., Krumpálová Z., Stašiov S., Fedor, P.: Biodiversity of selected invetrebrate groups in oak-hornbeam forest ecosystem in SW Slovakia ......

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Országh I., Országhová Z.: Štruktúra taxocenóz stonožiek (Myriapoda: Chilopoda) dubovo-hrabových lesov Malých Karpát.a.Trnavskej pahorkatiny (JZ Slovensko) . . . . . . . . . . . Stašiov S.: Mnohonôžky (Diplopoda) v.dubovo-hrabových ekosystémoch (Malé Karpaty, Trnavská pahorkatina, JZ Slovensko) . . . . . . . . . . . Országh I.: Fenológia nymfálnych instarov ucholaka Chelidurella acanthopygia (Dermaptera: Forficulidae) v.dubovo-hrabových lesoch Malých Karpát a Trnavskej pahorkatiny (JZ Slovensko) . . . . . . . . . . . . . . . . Hradil K.: Společenstva ploštic v.epigeonu dubo-habrových lesů v.oblasti JZ Slovenska . . . . . . . . . . . . . . . Holecová M., Némethová D., Kúdela M.: Štruktúra a funkcia spoločenstiev nosáčikov (Coleoptera, Curculionoidea) v.epigeóne dubovo–hrabových lesov JZ Slovenska . . . . . . Holecová M., Krumpál M., Országh I., Krumpálová Z., Stašiov S., Fedor, P.: Biodiverzita vybraných skupín bezstavovcov v.ekosystéme dubovo-hrabového lesa v.oblasti JZ Slovenska . .

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Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 3–19

ECOLOGICAL CHARACTERISTICS OF THE STUDIED FOREST COMMUNITIES OF AN OAK-HORNBEAM TIER IN SW SLOVAKIA JANKA ZLINSKÁ1, LADISLAV ŠOMŠÁK2, MILADA HOLECOVÁ3 Martinčekova 20, 820 04 Bratislava, The Slovak Republic Department of Soil Science, Faculty of Natural Sciences, Comenius University, Mlynská dolina B-2, 842 15 Bratislava, The Slovak Republic 3 Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská dolina B-1, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected] 1 2

Abstract Zlinská.J., Šomšák.L., Holecová.M.: Ecological characteristics of the studied forest communities of an oak-hornbeam tier in SW Slovakia. Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 3–19. The paper deals with characterization of the study area with its sampled plots more in details, including geology, climatic conditions, pedological and phytocoenological aspects of the 10 oak-hornbeam forest stands. For 4 seasons we were focused on selected invertebrate groups inhabiting epigeon, mosses and dendrotelmae from the coenological point of view. The study forest stands with 40–100 years of age are situated in 2 orographic units: the Malé Karpaty Mts and Trnavská pahorkatina hills. Corresponding with the 3 recorded forest types (Carpinion betuli, Quercion confertae-cerris, Q. pubescentis-petraeae) we have analysed 3 soil types with different genesis and features: Rendzic Leptosols, Cambisols and Planosols. The studied stands differ in their age, anthropogenous impact and fragmentation (they represent fragmented forests in cultural environment as well as more continuous forest complexes). Key words: forest ecosystem, oak-hornbeam vegetation tier, climate, pedology, phytocoenology, Malé Karpaty Mts, Trnavská pahorkatina hills, SW Slovakia

Introduction In 1999–2002 we realised complex coenological research of the selected invertebrate groups (ciliates, naked amoebae, water bears, soil mites, pseudoscorpions, spiders, terrestrial isopods, millipedes, centipedes, ear-wigs, bugs, weevils) in an oak-hornbeam forest ecosystem in SW Slovakia. The presented supplement includes analyses from soil as well as some other microhabitats such as mosses, decaying wood and dendrotelmae, which have not been intensively studied yet. 3

In the introduction we provide detailed characteristics of the study area with a special emphasis on geology, climate, soils and phytocoenology.

Material and methods The names of soil types and subtypes are presented according to the Morphogenetic soil classification system of Slovakia (Šály et al., 2000). The other soil analyses refer to the methods by Hraško et al. (1962). The soil samples were obtained in the same depth like the zoological material from fallen leaves and upper humus horizon (epigeon). The nomenclature of vascular plants is presented according to Ehrendorfer (1973). The phytocoenoses were analysed according to the Central-European Zürich-Montpellier school (Braun-Blanquet, 1964).

Study area Invertebrate communities of the mentioned forest stands (Carpinion betuli, Quercion confertae-cerris, Q. pubescentis-petraeae) were analysed in southern, central and northern parts of the Malé Karpaty Mts and nearby NW points of Podunajská nížina lowland (Trnavská pahorkatina hills) from Pezinok to Naháč (Fig. 1). We selected 10 following study plots, which were analysed for 4 vegetation periods (1999–2002) from the zoological point of view. 1. Cajla (CA), 48°20' N, 17°16' E, GRN (Grid Reference Number of the Databank of the Fauna of Slovakia) 7669c; 2. Vinosady (VI), 48°19' N, 17°17' E, GRN 7669d; 3. Fúgelka (FU), 48°22' N, 17°19' E, GRN 7669b; 4. Lindava (LI), Nature Reserve, 48°22' N, 17°22' E, GRN 7670a; 5. Horný háj grove (HH), 48°29' N, 17°27' E, GRN 7570b; 6. Lošonec–lom quarry (LL), 48°29' N, 17°23' E, GRN 7570b; 7. Lošonský háj grove (LH), Nature Reserve, 48°28' N, 17°24' E, GRN 7570b; 8. Naháč–Kukovačník (NA), 48°32' N, 17°31' E, GRN 7471c; 9. Naháč–Katarínka 1 (NK1), Nature Reserve, 48°33' N, 17°33' E, GRN 7471a; 10. Naháč–Katarínka 2 (NK2), Nature Reserve, 48°33' N, 17°32' E, GRN 7471a. The study area of the Malé Karpaty Mts geomorphologically belongs to the subunit of Pezinské Karpaty Mts, being predominantly built by crystalinicum. Its main part is formed by 2 massives: Bratislava massive, built by two mica garnet rocks and Modra massive formed by biotite granodiorite. Their contact zone has been built by crystalic shales, known as Pezinok-Pernek crystalinicum. Except for shales the bedrock consists of biotites phyllites and biotite-garnet mica schist gneisses. This actually occurs at the study sites of Cajla, Vinosady, Fúgelka, Lošonský háj grove, Naháč–Kukovačník, Naháč–Katarínka 1. On north and west the crystalinicum is bordered by a narrow strip of cover Mesozoic unite with the main rocks of Lower Triasic quartzites, siliceous and calcareous sandstones, variegated shales, limestones and dolomites. Carbonates appear on surface at the study sites of Lošonec–lom quarry and Naháč–Katarínka 2. Of 4

Neogene sediments deposed in Trnavská pahorkatina hills in depression under the slopes of the Malé Karpaty Mts. Pliocene clay and claystone bads, sands and sporadically gravel are the most frequent. Clays occur under the site Lindava, sandstones and gravels are at Horný háj grove (Buday et al., 1962; Maheľ, Cambel, 1972). From the altitudinal point of view the study sites are situated in the hilly zone from 240 to 350 m a.s.l.

Fig. 1. Study area and position of the ten study plots. Abbreviations of study sites see chapter Study area.

Climate According to the climatic classification the area of the Malé Karpaty Mts up to the altitude of 400 m a.s.l. belongs to the moderately warm climatic region. The southern 5

part is classified under the moderately warm and moderately moist climatic district A5 with mild winter, the northern part under moderately warm and moderately moist hilly district B3 (Petrovič, 1972). The main climatic character of the area refers to perpendicular position towards dominating NW falling winds on both sides of the mountains. Average annual temperature varies between 7–9 °C, in January 2–3 °C and in July 18–20 °C (Table 1). Average temperatures in vegetation season (April–September) reach 14–15 °C. Average number of summer days (more than 25 °C) is 40–60 annually while there are 30–40 winter days (less than -0,1 °C) a year (Petrovič, 1968; Tarábek, 1980). T a b l e 1. Average monthly and annual air temperature (°C) in 1931–1960. Meteorological station Pezinok–Myslenice Jaslovské Bohunice

1 1.2 2.1

2 1.3 0.3

3 5.2 4.4

4 5 6 7 8 9 10 10.4 15.4 18.5 20.3 19.5 15.7 10.1 9.7 14.5 17.6 19.2 18.7 14.9 9.6

11 4.5 4.1

12 0.6 0.2

year 10.2 9.2

Long-term average monthly and annual precipitations (1931–1960) from the studied region are given in Table 2. Precipitations of 320 mm refer to the vegetation period (April–September). Snow cover takes 90–100 days annually (Petrovič, 1968). T a b l e 2. Average monthly and annual precipitations (mm) in 1931–1960. Meteorological station Pezinok–Myslenice Jaslovské Bohunice

1 44 33

2 44 33

3 36 29

4 38 38

5 57 57

6 68 67

7 57 59

8 60 62

9 37 40

10 37 36

11 58 52

12 54 43

year 590 549

Precipitations are influenced by numerous factors, e.g. altitude. These two variables positively correlate (Modra 172 m a.s.l. – 687 mm, Limbach 181 m a.s.l. – 881 mm) (Petrovič, 1968). Monitoring of long-term averages of temperature and precipitations appears as important for forest vegetation. However for research on invertebrates and their dynamics actual average annual temperatures and precipitations during sampling have to be taken into account. (Tables 3, 4). T a b l e 3. Average monthly and annual air temperature (°C) in individual years of the research. Meteorological station Pezinok–Myslenice 1999 Pezinok–Myslenice 2000 Pezinok–Myslenice 2001 Pezinok–Myslenice 2002 Jaslovské Bohunice 1999 Jaslovské Bohunice 2000 Jaslovské Bohunice 2001 Jaslovské Bohunice 2002

6

1 2 -0.2 1 -1.3 3.4 0.4 1.8 0.2 4.4 -0.6 -0.1 -2.7 2.6 0.4 1.8 -0.9 4

3 7.2 6 5.8 6.9 6.8 5 5.8 6.2

4 11.8 14 9.9 10.5 11.5 13.6 8.9 9.9

5 15.6 17.4 17.2 18.1 15.2 16.6 16 17.4

6 18.2 20.4 17.5 20.5 17.6 19.5 16.5 19.4

7 21.2 18.9 20.9 22 21 18.3 20.5 21.9

8 18.9 21.7 21.8 20.8 18.8 21.3 21.3 20.6

9 18.1 15.2 13.9 15 18.1 14.7 13.4 14.3

10 10.4 12.5 13.1 9.3 10.1 12.8 12.6 8.6

11 3.8 7.9 3.7 7.6 3.3 7.7 2.9 7.3

12 0.5 2.3 -3.7 -0.4 -0.5 1.6 -4.6 -1.2

year 10.5 11.5 10.2 11.2 10.1 10.9 9.6 10.6

T a b l e 4. Average monthly and annual precipitations (mm) in individual years of the research. Meteorological station 1 2 3 4 5 6 7 8 9 10 11 12 Pezinok–Myslenice 1999 16.7 36.6 24.2 28.7 45.8 74.9 106.5 94.4 67.7 89.2 53.9 42.4 Pezinok–Myslenice 2000 64.5 60.6 109.3 14.0 27.4 12.1 65.0 47.5 66.7 46.6 56.6 58.0 Pezinok–Myslenice 2001 14.3 20.9 62.9 29.5 14.6 27.9 82.8 33.5 85.9 14.4 45.0 35.9 Pezinok–Myslenice 2002 19.1 40.2 31.3 22.0 29.5 56.4 53.8 161.7 40.9 87.8 70.3 74.6 Jaslovské Bohunice 1999 13.9 40.8 15.9 91.4 39.2 103.2 78.6 49.2 13.0 21.2 45.3 50.5 Jaslovské Bohunice 2000 38.5 29.4 87.4 11.2 32.7 8.1 66.0 21.4 53.9 32.6 78.9 46.7 Jaslovské Bohunice 2001 14.3 20.9 62.9 28.1 54.8 34.8 107.8 16.9 109.0 15.3 39.2 35.4 Jaslovské Bohunice 2002 16.7 36.6 24.2 28.7 45.8 74.9 106.5 94.4 67.7 89.2 53.9 42.4

year 745.5 410.9 467.6 687.6 532.2 506.8 539.5 681.0

According to both the meteorological station data the average annual temperature in 2000 and 2002 (Tables 1, 3) were significantly higher comparing with the long-term average. Annual precipitations appeared significantly higher in the southern part of the study area than the average in 1999 and 2002, in the northern part in 2002 (Tables 2, 4). Therefore the years 1999 and 2002 were warmer and in summertime more humid than averagely. The years 2000 and 2001 were warmer and drier.

Soils In accordance with the recorded vegetation of Carpinion betuli, Quercion confertae-cerris and Q. pubescentis-petraeae we analysed 3 soil types with different genesis and features. As for their genetic features the development has been influenced by warm climate and carbonate bedrock (Rendzic Leptosols), warm climate and crystalic bedrock, being spatially and successionally impacted by forest stands and increased water overflow in valleys and slopes or surface humidity in plain areas with clays (Cambisols, Planosols). Physical features of soils refer to higher aeration due to numerous stones, gravel and sand and with lack of higher amount of dust as well as clay in soil mater (Rendzic Leptosols, Cambisols). This fact reflects in very high or middle resistance to compaction. Clays situated at the site of Lindava are markedly affected by compaction and are weakly aerated. On the contrary shallow carbonatic soils with a high portion of skelete avoiding compaction are very resistant to compaction (Rendzic Leptosols). This resistance is even enhanced by frequent occurrence of compact rocks on the surface and is supported by presence of carbonates, which provide coagulation of soil mater. Cambisols are partially resistant soils with higher portion of dust and clay components. Planosols have the lowest resistance due to high portion of clay components. Moreover its resistance against compaction is weakened by absence of carbonates. However they have higher content of humus therefore their resistance to trampling is sufficient. The soils usually occur on gentle slopes or in lowlands and are not potentially threatened by water erosion. Only Rendzic Leptosols at Naháč–Katarínka 2 cover steeper SW slope. This fact synerged by occasional rainfalls and remarkable spring streams of rain water on the surface (app. 650 mm of atmospheric rainfalls annually) represents significant and potential threat for soils due to water erosion. It is 7

T a b l e 5. The survey of pedological variables at the study sites. Study site/horizont

Soil type by WRB98*, sign.

pH/ H2O

pH/KCl

%C

%N

C:N

% Cox % humus

SOB

EA

CEC

% BS

(mval/100 g) 1. Cajla O-horizon +2–0 cm Ao-horizon 0–5 cm 2. Vinosady O-horizon +2–0 cm Ao-horizon 0–5 cm 3. Fúgelka O-horizon +2–0 cm Ao-horizon 0–5 cm 4. Lindava O-horizon +2–0 cm A/Bg-horizon 0–5 cm 5. Horný háj grove O-horizon +2–0 cm Ao-horizon 0–5 cm 6. Lošonec–lom quarry O-horizon +2–0 cm Amc-horizon 0–5 cm 7. Lošonecký háj grove O-horizon +2–0 cm Ao-horizon 0–5 cm 8. Naháč–Kukovačník O-horizon +2–0 cm Ao-horizon 0–5 cm 9. Naháč–Katarínka 1 O-horizon +2–0 cm Ao-horizon 0–5 cm 10. Naháč–Katarínka 2 O-horizon +2–0 cm Amc-horizon 0–5 cm

Dystric Cambisols KMma 4.83 4.29 Dystric Cambisols KMma 4.48 4.18 Dystric Cambisols KMma 3.90 3.76 Dystric Planosols PGm 4.68 3.97 Dystric Cambisols KMma 5.00 4.17 Rendzic Leptosols RAk 6.74 6.55 Dystric Cambisols KMma 4.29 4.37 Dystric Cambisols KMma 4.18 4.08 Dystric Cambisols KMma 4.24 3.94 Rendzic Leptosols RAm 6.45 6.37

4.21 3.43

6.50 2.80

0.48 0.20

13.54 14.00

6.50 4.83

18.40 4.20

9.10 8.50

27.50 12.70

67.00 33.00

3.82 3.31

8.00 2.55

0.67 0.22

11.94 11.59

7.99 4.40

16.00 4.80

11.50 9.40

27.50 14.50

58.00 34.00

3.37 3.02

14.40 3.50

1.01 0.29

14.26 12.07

14.40 6.03

12.60 2.60

13.10 10.30

25.70 12.90

49.00 20.00

3.98 3.20

8.40 1.80

0.63 0.18

13.33 10.00

8.39 3.10

22.60 2.20

12.50 11.10

35.10 13.30

64.00 17.00

4.54 3.48

9.60 2.30

0.73 0.19

13.15 12.11

9.59 3.97

31.80 3.00

9.50 8.30

41.30 11.30

77.00 27.00

6.14 5.75

3.80 2.70

0.35 0.29

10.86 9.31

3.79 4.65

22.60 17.40

1.80 2.50

24.40 19.90

93.00 87.00

3.54 3.48

7.40 3.75

0.60 0.31

12.33 12.10

7.40 6.47

15.40 7.40

13.70 10.90

29.10 18.30

53.00 40.00

3.60 3.39

7.20 1.80

0.53 0.20

13.58 9.00

7.19 3.10

15.40 4.40

12.80 9.40

28.20 13.80

55.00 32.00

3.68 3.35

7.40 1.70

0.87 0.17

8.51 10.00

7.40 2.93

19.60 3.00

14.90 9.80

34.50 12.80

57.00 23.00

5.90 5.55

5.40 3.20

0.52 0.35

10.38 9.14

5.40 5.52

49.40 36.20

3.00 3.70

52.40 39.90

94.00 91.00

Explications: * according to comparison in Šály et al. (2000), SOB – sum of exchangeable bases, EA – exchangeable acidity, CEC – cation exchangeable capacity, BS – base saturation

followed by local soil denudation. Soils with accumulated layers of sediments occur down the hill and in deeper valleys. While in upper parts of slopes we often record shallow to middle deep soils with rocky bedrock in depth of 0.3 to 0.6 m, in lower parts the compact bedrock lies in depth of 2–3 m. Real water erosion effects are slowed down by vegetation cover. Forest with its undergrowth has weaker resistance to water erosion. Soils particularly developed from crystalic rocks (sites Vinosady, Cajla, Fúgelka, Horný háj grove, Lošonský háj grove, Naháč–Kukovačník, Naháč–Katarínka 1), in 2 cases on carbonates (sites Lošonec–lom quarry, Naháč–Katarínka 2) and once on tertiary clays (site Lindava). The studied natural forest soils contain sufficient amount of humus in a relatively shallow humus (Ao, Amc, Aoq) horizon (2.93–6.47%). However in comparison with farmland this concentration appears low (Table 5). Concentration of Cox is relatively balanced in fallen leaf horizon, the lowest amount of organic matter refers to fallen leaf horizon in Rendzic Leptosols (3.79–5.40%), where leaves are markedly mineralised due to carbonates. In the other soils (Cambisols, Planosols) the carbon concentration in a fallen leaf horizon varies between 6.50–14.40%, what hints at various degree of organic mater decay. All the studied soils may be considered as significantly anthropized. Many forests covered former vineyards in coline zone, where former vegetation had been destroyed and soils rigoled. At present forests are usually economically exploited what leads to partial damage on soil profile. The Rendzic soil at Naháč–Katarínka 2 was affected by the monastery and a part of the locality has been covered by ruins. At most of the study sites (7) in communities of Carpinion betuli (Galio sylvaticae-Carpinetum a Querco petraeae-Carpinetum) we recorded Cambisols, which are developed, brown soils, weakly to partially eroded. These soils develop in hillier or even mountainous areas from sloap compact bedrock and have suitable biologically active form of humus in saturated to acid mul. A-horizon reaches thickness of several centimeters. This is an alternation soil with dominant Cambic Bv-horizon under surface, formed in brunification process. Moreover in the study area we recorded District Cambisols, with no more diagnostic horizons under surface. C-horizon represents eroded crystalic rocks or even compact rock. Development of Cambisols is accompanied by acidification leaching, what actually slows down biological circulation of nutrients. Acid and unsaturated District Cambisols occur at Vinosady, Cajla, Fúgelka, Horný háj, Lošonecký háj, Naháč–Kukovačník, Naháč–Katarínka. Two of the soils at Lošonec–lom quarry and Naháč–Katarínka 2 in communities of Corno-Quercetum pubescentis and Querco petraeae-Carpinetum are Rendzic Leptosols, being developed on limestone. These soils belong to the category of Rendzic soils with sod pedogenetical process to processes of accumulation and stabilization of humus, hence the soils with Mollic Am-horizon, sometimes even Ochric horizon (Ao) with no more diagnostic horizons (Šály et al., 2000). According to Němeček et al. (1990) they belong to weakly developed postlitogene soils, which were formed on eroded carbonates with higher content of skeleton (more than 30%). They are indicated by a weak degree of soil profile development and low production potential, which is enhanced by extreme climatic conditions and location on slopes. Content of humus and acidification of soil profile depend on hydrothermic regime. From the physical point of view the soil is well aerated due to frequent 9

occurrence of stones, gravel and sand. It is very resistant to compaction. In the stand of Corno-Quercetum pubescentis (site Naháč–Katarínka 2) we recorded the subtype Rendzic Leptosols (RAm) with no more diagnostic horizons or their indication, with remarkable transformation into C-horizon. In the forest stand of Querco petraeae-Carpinetum (site Lošonec–lom quarry) we observed Rendzic Leptosols (RAk) with indication of Cambic Bv-horizon and presence of carbonates. In Rendzic Leptosols the exchangeable pH is weakly acid and they are very saturated. The study site Lošonec–lom quarry (LL) is situated just over the quarry and during working days it is intensively impacted by carbonatic dust, what actually reflects in the highest value of active as well as exchangeable pH in decaying horizon and the highest degree of organic matter decay. At Lindava we detected Dystric Planosols (PGm) in the community of Quercetum petraeae-cerris in deluvial plain. In the depression under the slopes of the Malé Karpaty Mts these soils occur on tertiary clays. A shallow (3 cm) Ochric silicate Aoq-horizon is followed by deeper Bgv-horizon, which is formed under remarkable periodical saturation of soil mater by surface water in horizons with decreased resp. low hydraulic conductibility. Alternation of stagnation and water circulation, reduction and oxidation processes leads to forming colourful horizon with thickness of more than 15 cm. This is a mosaic coloration, alternation of rusty, pale brown and grey colours in the matrix. At the study site we recorded the subtype Dystric Planosols (PGm) with no more diagnostic horizons under surface. The soil is strongly acid, saturated in Aoq/Bg (Table 5).

Vegetation Mixed forest stands at the 10 study sites in SW the Malé Karpaty Mts and N promontory of the Podunajská nížina lowland – Trnavská pahorkatina hills, occurring in a coline zone at the altitude of 250–350 m a.s.l., may be classified into mesophilous oak-hornbeam forests of Carpinion betuli I s s l e r 1931 em. M a y e r 1937 (sites of Vinosady, Cajla, Fúgelka, Horný háj grove, Lošonský háj grove, Lošonec–lom quarry, Naháč–Kukovačník, Naháč–Katarínka 1), subxerothermophilous Balkan mixed forests of Quercion confertae-cerris H o r v a t 1954 (Lindava) and oak xerothermophilous submediterranean forests of Quercion pubescentis-petraeae B r. - B l. 1931 (Naháč–Katarínka 2). In the area of the Malé Karpaty Mts and nearby Podunajská nížina lowland the oak-hornbeam forests have been floristically and pedologically differentiated. For all the cases included in the research the oak-hornbeam forests are situated on sandy-loam skeletal mesotrophic to oligotrophic Dystric Cambisols on poor crystalic bedrock (granodiorite). Floristically they are indicated by Carex pilosa, Galium sylvaticum, G. schultesii, Stellaria holostea, Carpinus betulus, Quercus dalechampii, Pulmonaria officinalis, Symphytum tuberosum of Carpathian oak-hornbeam forest suballiance Carici pilosae-Carpinenion betuli J. et M. M i c h a l k o 1986 (Michalko et al., 1986). Moreover the species spectrum includes many representatives of Fagetalia P a w ł. 1928, such as Galium odoratum, Arum alpinum ssp. besserianum, Melica nutans, M. 10

uniflora, Asarum europaeum, Bromus benekenii, Dentaria bulbifera, Festuca heterophylla, Cornus mas, Euphorbia polychroma, Tanacetum corymbosum, Pulmonaria obscura, Campanula trachelium, etc. In warmer and drier habitats the communities of Carpinion betuli (Naháč–Kukovačník and Naháč–Katarínka 1) are sporadically enriched in some species of subxerothermophilous oak forests, such as Quercus cerris, Melittis melissophyllum, Cornus mas, Vicia cassubica, Pulmonaria mollis, Clinopodium vulgare, Carex montana and Vincetoxicum hirundinaria. Domination of Quercus cerris in stands of Carpinion betuli as well as Quercion confertae-cerris is supposed to correspond with human impact in a synergy with good sprouting capacity. In several stands on oligotrophic cambisols acidophilous species, e.g. Luzula luzuloides, Avenella flexuosa and Melampyrum pratense occur occasionally. Oak-hornbeam forests in SE part of the Malé Karpaty Mts belong to the communities of Galio sylvaticae-Carpinetum O b e r d. 1957 and Querco petraeae-Carpinetum S o ó et P ó c s 1957. The associations are not indeed significantly differentiated and are floristically changed as the whole SE part of the Malé Karpaty Mts has been considerably managed (vineyards, economically managed forests). High anthropization has been proved by a big proportion of nitratophilous species (Alliaria petiolata, Galium aparine, Geranium robertianum, Chaerophyllum temulum, Geum urbanum, Glechoma hirsuta, Lamium maculatum, Urtica dioica, Sambucus nigra, Viola odorata, Mycelis muralis, Fallopia convolvulus, Lapsana communis, etc.) in all the study sites. Subxerothermophilous (xerothermophilous) forests with a more significant portion of Quercus cerris indicate climatically warm locations and are interacted with ilimerized Luvisols on loess cover (in the study area Dystric Planosols on tertiary clays). The soils are dried up in summer or during intensive droughts. In spring under heavy rains they are humid, loamy and hence heavy, gently acid to acid. Such the conditions have been observed at the site Lindava being situated in Podunajská nížina lowland (geomorphological subunit of Trnavská pahorkatina hills). According to floristic structure and soil type the stand may be classified as Balkan mixed forests of Quercion confertae-cerris, which are climatically zone in the continental part of Balkan Peninsula with effect towards Pannonian region and even slopes of W Carpathians. Quercus cerris is accompanied by Q. dalechampii and sporadically Acer campestre. The shrub tier, relatively rich, however poor at damaged phytocoenoses, includes Ligustrum vulgare, Cornus mas, C. sanguinea, Prunus spinosa, Rosa gallica, Crataegus laevigata, Rhamnus catharticus. In our case it has been eliminated due to human impact. The undergrowth is formed by Carex montana, Potentilla alba, Poa pratensis var. angustifolia, Pulmonaria mollis, Lathyrus niger, Primula veris, Melittis melissophyllum, occasionally followed by some species of Quercion pubescentis-petraeae B r. - B l. 1931, such as Festuca rupicola, Hypericum perforatum, etc. or acidophilous species, e.g. Veronica officinalis, Luzula luzuloides, L. pilosa, Avenella flexuosa, Genista tinctoria, Silene nutans. The less depredated phytocoenoses are enriched in some species of mesophilous deciduous forests – Poa nemoralis, Dactylis polygama, Carpinus betulus, which are in contact with the Malé Karpaty Mts. All the xerothermophilous forests have been classified into more concipated association of Quercetum petraeae-cerris S o ó 1957. 11

The forest stand at Naháč–Katarínka 2 on the soil subtype of Rendzic Leptosols belong to the group of xerothermophilous submediterranean oak forests (Quercion pubescentis-petraeae) which are interacted with warm southern, south-western and south-eastern slopes in hills, on carbonates, dolomites, calcareous conglomerates, flysches and basaltic eruptive rocks. They inhabit extreme forms of surface, such as ridges of mountains and steep slopes. On carbonates in the Malé Karpaty Mts the communities contact nearby oak-hornbeam forests in fragments up to the altitude of 400 m a.s.l. The stands are dominated by (Quercus pubescens), which reaches the height of 5 m as well as Q. cerris (8 m maximally). Limetree (Tilia cordata) has been planted in Nature Reserve Katarínka right under the monastery of St. Catherine. The trees are also represented by Acer campestre, shrubs by Cornus mas, Crataegus monogyna, Fraxinus excelsior, Viburnum lantana, Euonymus verrucosa, E. europaea and Sambucus nigra. The undergrowth is built by Buglossoides purpurocaerulea, Pulmonaria mollis and numerous species of Festuco-Brometea B r. -B l. et R. T x. 1943. For the long time the study forest has been anthropically impacted by the nearby monastery and at present by tourism particularly. This obviously correlates with high dominance of nitratophilous species, locally significantly dominating. The xerothermous forest belongs to the association of Corno-Quercetum pubescentis J a k u c s et Z ó l y o m i 1958. The community forms a relatively narrow wreath in SE part of the peak. Down the hill on deeper soil and less steep slope it transforms into Quercetum petraeae-cerris S o ó 1957.

The study sites with their phytocoenological records 1. The Malé Karpaty Mts, Cajla (CA), SE slope of Malá cajlanská homola hill, 280 m a.s.l. Oak-hornbeam forest Galio sylvaticae-Carpinetum, economically managed forest neighbouring with meadows and vineyards on S and E, from N and W closed forest complexes. Analysed area in m2: 400, exp.: S, slope 7°, date: June 6, 2005. Age of a stand: 80–100 years, average height: 27–30 m, average thickness at chest height: 40 cm, coverness: E3: 75%, E2: 2%, E1: 90%, E0: 5%. E3: Quercus dalechampii 5, Carpinus betulus +, Sorbus torminalis (8 m) subdominant +. E2: eliminated artificially, Acer campestre +, Quercus dalechampii r. E1: Poa nemoralis 3, Quercus dalechampii 3, Galium odoratum 2, Brachypodium sylvaticum 1, Melica uniflora 3, Acer campestre +, Pulmonaria officinalis +, Fragaria moschata +, Carex pilosa +, Galium sylvaticum +, Campanula persicifolia +, Rhamnus catharticus +, Scrophularia nodosa r, Melampyrum pratense +, Hieracium sabaudum +, Tilia platyphyllos.+, Euonymus verrucosa.+, Hieracium sylvaticum 1, Viola reichenbachiana +, Prunus avium +, Carpinus betulus +, Fraxinus excelsior +, Dactylis polygama +, Rosa canina agg. +, Impatiens parviflora +, Rubus fruticosus agg.+, Lysimachia nummularia +, Viola mirabilis r, Geranium robertianum +, Galium aparine +, Alliaria petiolata 1, Impatiens noli-tangere +, Mycelis muralis +, Galeopsis 12

pubescens r, Juglans regia r, Lathyrus vernus +, Luzula luzuloides +, Veronica officinalis +, Ranunculus ficaria 1, Euphorbia amygdaloides +, Geum urbanum +, Rubus caesius 1, Corydalis solida +, Stellaria nemorum +, Scilla bifolia +, Prunella vulgaris +, Hedera helix +. 2. The Malé Karpaty Mts, Vinosady (VI), NW slope of Kamenica hill, 280 m a.s.l. Oak-hornbeam forest Querco petraeae-Carpinetum variant with Poa nemoralis, economically managed forest, former vineyard, neighbouring from W with drier subxerophilous meadows and shrub complexes. Analysed area in m2: 400, exp.: NW, slope: 5–7°, date: June 23, 2005. Age of a stand: 60–80 years, average height: 25–27 m, average thickness at chest height: 35–40 cm, coverness: E3: 70%, E2: 30%, E1: 80%. E3: Quercus dalechampii 4, Quercus cerris +, Prunus avium +, Acer campestre r. E2: Acer campestre 2, Tilia cordata +, Ligustrum vulgare +, Fraxinus excelsior +, Cornus sanguinea +. E1: Poa nemoralis 3, Melica uniflora 3, Acer campestre 2, Cephalanthera alba 1, Quercus dalechampii 1, Tilia cordata +, Prunus avium +, Ligustrum vulgare +, Sorbus torminalis +, Quercus cerris +, Pyrus pyraster r, Arum alpinum ssp. besserianum +, Rhamnus catharticus +, Rosa canina agg. +, Hieracium sylvaticum +, Sedum maximum +, Euonymus europaea +, Fraxinus excelsior +, Hedera helix +, Cornus sanguinea +, Hieracium sabaudum +, Brachypodium sylvaticum +, Dactylis polygama +, Solidago virgaurea r, Lilium martagon +, Clinopodium vulgare +, Erigeron annuus subsp. septentrionalis +, Sambucus nigra +, Lamium maculatum 2, Geranium robertianum 2, Alliaria petiolata 2, Galium aparine 1, Mycelis muralis +, Glechoma hirsuta 1, Geum urbanum 1, Viola odorata 1, Fallopia convolvulus 1, Anthriscus cerefolium ssp. trichosperma 1, Chaerophyllum temulum 1, Stellaria nemorum +, Gagea pratensis +, Lamium purpureum +, Rubus caesius +, Pulmonaria officinalis +, Lathyrus vernus +, Arabis turrita +, Hordelymus europaeus +, Corydalis cava 3, C. solida 1, Scilla bifolia 1, Ranunculus ficaria 1, Geum urbanum +, Allium oleraceum 1, Veronica hederifolia 1, Pulmonaria mollis r. 3. The Malé Karpaty Mts, Fúgelka (FU), appr. 3 km NW from the village of Dubová, 350 m a.s.l. Oak-hornbeam forest Galio sylvaticae-Carpinetum facies with Rubus fruticosus and R..hirtus (oceanic species). Economically exploited forest with appr. 20 year old underplanting of Acer pseudoplatanus in lines due to transformation from a low forest to high-trunked forest. There are several older game-keeping trails in the forest. On S in contact with vineyards, on E with road, on N and W with more continuous forest complexes. Analysed area in m2: 400, exp.: S, slope: 5°, date: June 23, 2005. Age of a stand: 80–100 years, average height: 30–35 m, average thickness at chest height: 40 cm, coverness: E3 80%, E2: 50%, E1: 40%, E0: 3%.

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E3: Quercus dalechampii 4, Prunus avium +, Fraxinus excelsior +, Acer pseudoplatanus (8–10 m subdominant) r, Larix decidua r. E2: Corylus avellana +, Acer pseudoplatanus 3, Crataegus monogyna +, C. laevigata r, Ulmus minor +, Rosa canina agg. +. E1: Ribes uva-crispa (wild) +, Melica uniflora 3, Poa nemoralis +, Cephalanthera alba +, Carex pilosa 1, Symphytum tuberosum +, Viola reichenbachiana +, Galium sylvaticum +, G. odoratum 1, Prunus avium 1, Acer pseudoplatanus 2, Stachys sylvatica +, Scrophularia nodosa +, Melampyrum pratense +, Avenella flexuosa +, Luzula luzuloides +, Polygonatum odoratum +, Pulmonaria officinalis +, Rubus fruticosus agg. 2, R. hirtus 2, Hieracium sabaudum +, Lysimachia punctata +, Quercus dalechampii 2, Ulmus campestre 1, Pyrus communis r, Acer platanoides +, Fraxinus excelsior +, Circaea lutetiana +, Dryopteris filix-mas +, Athyrium filix-femina +, Geranium robertianum 1, Urtica dioica 1, Galium aparine 1, Impatiens noli-tangere 1, I. parviflora 1, Sambucus nigra +, Alliaria petiolata r, Ranunculus ficaria 3, Euphorbia amygdaloides +, Arum alpinum ssp. besserianum +, Polygonatum odoratum +, Glechoma hirsuta +, Carex digitata +, Lathyrus vernus +. 4. The Trnavská pahorkatina hills, Lindava (LI), appr. 1 km on E from the village of Píla, nature reserve, 240 m a.s.l. Oak-hornbeam forest Quercetum petraeae-cerris on Dystric Planosols, former economically exploited forest. Large complex of island forest surrounded by fields and road. Analysed area in m2: 400, date: June 23, 2005. Age of a stand: 80–100 years (or even 120 and more), average height: 27–30 m, average thickness at chest height: 44 cm, coverness: E3: 65%, E2: 1%, E1: 80%, E0: 5%. E3: Q. cerris 4, Quercus dalechampii +. E2: eliminated by human impact, Tilia x vulgaris +, Ribes uva-crispa (wild) +,. E1: Melica uniflora 4, Poa nemoralis 1, Quercus dalechampii 2, Q. cerris 1, Sorbus aria +, Carex montana 1, Galium odoratum 1, Campanula rapunculus +, Festuca rupicola 1, Hypericum perforatum +, Clinopodium vulgare +, Veronica chamaedrys +, Rubus fruticosus agg. 1, Carpinus betulus +, Symphytum tuberosum +, Prunus spinosa r, Rosa canina agg. +, Rhamnus catharticus +, Acer campestre +, Solidago virgaurea +, Cephalanthera alba +, Viola reichenbachiana +, V. mirabilis +, Athyrium filix-femina r, Dryopteris filix-mas +, Hieracium sylvaticum +, Genista tinctoria 1, Veronica officinalis +, Avenella flexuosa 1, Luzula luzuloides +, L. pilosa +, Silene nutans 1, Digitalis grandiflora +, Scrophularia nodosa 1, Carex divulsa +, Geranium robertianum +, Galium aparine +, Vicia sepium +, Fallopia convolvulus +, Alliaria petiolata +, Impatiens parviflora 1, Lapsana communis 1, Chaerophyllum temulum +, Vincetoxicum hirundinaria r, Ajuga reptans +, Mycelis muralis +, Ranunculus ficaria 2, Ajuga reptans +, Geum urbanum +, Carex pilosa +, Cardamine impatiens +, Viola alba +, Fragaria moschata +, Lathyrus vernus +.

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5. The Trnavská pahorkatina hills, Horný háj grove (HH), in the cadastral area of Horné Orešany, 240 m a.s.l. Oak-hornbeam forest Querco petraeae-Carpinetum variant with Melica uniflora, economically exploited forest, former vineyard. It forms a larger complex of isolated forest surrounded by vineyards and farmland. Analysed area in m2: 400, exp.: W – SW, slope: 5°, date: June 23, 2005. Age of a stand: 60–80 years, average height: 27–30 m, average thickness at chest height: 40 cm, coverness: E3: 80%, E2: 10%, E1: 75%, E0: 2%. E3: Quercus cerris 3, Carpinus betulus 2, Quercus dalechampii 2, Fraxinus excelsior 1, Tilia cordata +, Sorbus torminalis r, Prunus avium r. E2: Tilia cordata 1, Sambucus nigra +, Sorbus torminalis +, Acer campestre +, Fraxinus excelsior +. E1: Melica uniflora 3, Lamiastrum galeobdolon 1, Fraxinus excelsior 1, Quercus cerris 1, Acer campestre +, Allium ursinum 3, Polygonatum multiflorum +, Arum alpinum ssp. besserianum+, Symphytum tuberosum +, Tilia cordata +, Ligustrum vulgare +, Scrophularia nodosa +, Viola odorata 2, Geum urbanum +, Chaerophyllum temulum 2, Alliaria petiolata 3, Galium aparine 2, Impatiens parviflora 1, Geranium robertianum 1, Mycelis muralis +, Fallopia convolvulus 1, Sambucus nigra +, Mercurialis paxii +, Anemone ranunculoides +, Gagea pratensis +, Euphorbia amygdaloides +, Lamium purpureum 1, Allium oleraceum +, Pulmonaria officinalis +, Ajuga reptans +, Corydalis cava 2, C. solida 1, Veronica hederifolia 3, Lathyrus vernus +, Mercurialis paxii 1, Ranunculus ficaria +, Isopyrum thalictroides 1 . 6. The Malé Karpaty Mts, Lošonec–lom quarry (LL), 340 m a.s.l. Oak-hornbeam forest Querco petraeae-Carpinetum caricetosum pilosae, economically exploited forest, intensively and systematically impacted by dust from the quarry, from E neighbouring with mesophilous meadows and pastures. Analysed area in m2 400, exp.: NE, slope: 8–10°, date: June 23, 2005. Age of a stand: 80–100 years, average height: 25 m, average thickness at chest height: 44.cm, coverness: E3: 65%, E2: 25%, E1: 100%, E0: 5%. E3: Quercus dalechampii 3, Quercus cerris 2. E2: Carpinus betulus 2, Crataegus monogyna 1, Acer campestre 1, Prunus spinosa +. E1: Melica uniflora 3, Galium odoratum 2, Carpinus betulus 1, Pulmonaria officinalis 1, Carex pilosa 1, Poa nemoralis 2, Dactylis polygama 1, Fragaria moschata +, Veronica chamaedrys +, Hedera helix +, Fraxinus excelsior +, Quercus cerris 2, Lysimachia nummularia 1, Quercus dalechampii 1, Brachypodium sylvaticum +, Hieracium sylvaticum +, H. sabaudum +, Luzula luzuloides +, Melampyrum pratense +, Malus sylvatica +, Rubus fruticosus agg. +, Acer platanoides +, Crataegus monogyna +, Ajuga reptans +, Geum urbanum 1, Rosa canina agg. +, Sorbus torminalis +, Viola reichenbachiana +, Lathyrus vernus +, Hypericum perforatum +, Stachys sylvatica +, Polygonatum multiflorum +, Scrophularia nodosa +, Dentaria bulbifera +, Galium aparine 1, Alliaria petiolata 1, Geranium robertianum +, Viola odorata +, Impatiens noli-tangere +, I. parviflora 1, Mercurialis paxii 1, Urtica dioica +, Mycelis muralis +, 15

Anemone ranunculoides +, Arum alpinum ssp. besserianum +, Ranunculus ficaria 2, Corydalis solida +, Rubus caesius +, Glechoma hirsuta +, Euphorbia amygdaloides +. 7. The Malé Karpaty Mts, Lošonský háj grove (LH) being situated near the village of Lošonec close to Smolenice in deluvium under the slope, 260 m a.s.l., from 1984 nature reserve Oak-hornbeam forest Querco petraeae-Carpinetum caricetosum pilosae, economically exploited forest, surrounded by closed forest complexes Analysed area in m2: 400, exp.: NE, slope 1°, date: June 23, 2005. Age of a stand: 80–100 years, average height: oaks 25 m, hornbeams 20 m, average thickness at chest height: oaks 44 cm, hornbeams 32 cm, coverness: E3: 80 %, E2: 1 %, E1: 75 %, E0: 5%. E3: Carpinus betulus 3, Quercus dalechampii 2, Q. cerris +. E2: eliminated by human impact, Carpinus betulus +. E1: Carex pilosa 3, Melica uniflora 2, Galium odoratum 3, Allium ursinum +, Quercus dalechampii 1, Quercus cerris 1, Hedera helix 2, Viola reichenbachiana 2, Pulmonaria officinalis 1, Dentaria bulbifera 2, Dactylis polygama +, Fraxinus excelsior +, Carpinus betulus +, Acer platanoides r, Oxalis acetosella +, Fragaria vesca +, Hieracium sylvaticum +, Sanicula europaea +, Lathyrus vernus 1, Acer campestre +, Polygonatum multiflorum +, Hieracium sabaudum +, Rosa canina agg. r, Lilium martagon +, Stachys sylvatica +, Rubus fruticosus agg. 1, Campanula persicifolia +, Alliaria petiolata 1, Geranium robertianum 1, Impatiens noli-tangere 1-2, Mycelis muralis +, Galium aparine 2, Impatiens parviflora 3, Fallopia convolvulus +, Vinca minor +, Corydalis solida +, Viola alba +, Neotia nidus-avis r, Crataegus monogyna 1, Rubus caesius +, Silene nemoralis +, Euphorbia amygdaloides +, Ranunculus ficaria 1, Anemone ranunculoides +, Isopyrum thalictroides 1, Carex sylvatica +, Prunus avium +. 8. The Malé Karpaty Mts, Naháč–Kukovačník (NA), 300 m a.s.l. Oak-hornbeam forest Querco petraeae-Carpinetum variant with Melica uniflora, young isolated stand amongst farmland areas, economically exploited forest. Analysed area in m2: 400, exp.: SE–E, slope: 2–3°, date: June 23, 2005. Age of a stand: 40–60 years, average height: 23 m, average thickness at chest height: 25.cm, coverness: E3: 75%, E2: 20%, E1: 100%, E0: 1%. E3: Quercus dalechampii 2, Q. cerris 2, Carpinus betulus 1, Prunus avium r, Robinia psedacacia r. E2: Carpinus betulus +, Sorbus torminalis +, Cornus mas +, Cornus saguinea +, Corylus avellana r, Malus sylvestris r, Prunus avium +, Sambucus nigra 1, Fraxinus excelsior (underplanted) 1, Ligustrum vulgare +, Acer campestre +, Crataegus laevigata r, Robinia psedacacia +. E1: Melica uniflora 5, Poa nemoralis +, Quercus cerris 1, Q. dalechampii 1, Prunus avium +, Crataegus monogyna +, Acer campestre +, Galium odoratum 2, Polygonatum multiflorum 1, Melittis melissophyllum 1, Vicia cassubica +, Vincetoxicum hirundinaria +, Pulmonaria mollis +, Galium schultesii +, Fragaria moschata +, Cornus sanguinea 16

+, Malus sylvestris +, Sorbus torminalis +, Campanula trachelium +, Carex montana +, C. muricata +, Rubus fruticosus agg. 3, Viola reichenbachiana +, Rosa canina agg. +, Lilium martagon +, Fraxinus excelsior +, Geranium robertianum 1, Urtica dioica +, Sambucus nigra 2, Alliaria petiolata +, Arum alpinum ssp. besserianum 1, Geum urbanum +, Viola odorata +, Chelidonium majus +, Robinia psedacacia +, Allium oleraceum +, A. ursinum 1, Symphytum tuberosum +, Corydalis solida +, Ranunculus ficaria +, Veronica hederifolia 1. 9. The Malé Karpaty Mts, Naháč–Katarínka 1 (NK1), 340 m a.s.l. Oak-hornbeam forest Querco petraeae-Carpinetum, economically exploited forest, nature reserve, surrounded by closed forest complexes. Analysed area in m2: 400, exp.: NW, slope 3–5°, date: June 23, 2005. Age of a stand: 40–60 rokov, average height: 17 m, average thickness at chest height: 22.cm, coverness: E3: 80%, E2: 1%, E1: 80%, E0: 3%. E3: Quercus dalechampii 4, Carpinus betulus 2, Quercus cerris +, Prunus avium +, Fagus sylvatica r. Tree tier consists of two parts: the main stratum with Quercus dalechampii and Carpinus betulus under it (coverness of 40 %). E2: Carpinus betulus 1, Sorbus torminalis +, Cornus mas +, Crataegus monogyna +. E1: Poa nemoralis 1, Galium odoratum 2, Quercus dalechampii 2, Q. cerris 1, Carex pilosa +, Pulmonaria officinalis +, Lathyrus vernus +, Crataegus monogyna +, Prunus avium +, Malus sylvestris +, Melittis melissophyllum +, Polygonatum multiflorum +, Clinopodium vulgare +, Fragaria moschata +, Campanula trachelium +, Hieracium sabaudum +, Vincetoxicum hirundinaria +, Euphorbia amygdaloides +, Ligustrum vulgare +, Euonymus europaea +, Acer campestre +, Dryopteris filix-mas +, Sorbus domestica r, Rosa canina agg. +, Viola reichenbachiana +, V. riviniana r, Rubus fruticosus agg. +, Vicia cassubica +, Lilium martagon +, Festuca gigantea +, Scrophularia nodosa +, Ajuga reptans +, Carex muricata +, Circaea lutetiana +, Acer platanoides juv. +, Urtica dioica +, Impatiens parviflora +, Alliaria petiolata +, Fallopia convolvulus +, Geum urbanum +, Mycelis muralis +, Geranium robertianum +, Sambucus ebulus +, Allium ursinum 1, Neotia nidus-avis r, Arum alpinum ssp. besserianum +, Ranunculus ficaria 1, Corydalis solida 1, Gagea lutea +. 10. The Malé Karpaty Mts, Naháč–Katarínka 2 (NK2), old forest stand under the monastery ruins, nature reserve, 320 m a.s.l. Xerothermophilous oak forest Corno-Quercetum pubescentis on the top of the hill, forming a wreath. Analysed area in m2: 400, exp. SE, slope: 45°, date: July 18, 2005. Age of a stand: 80–100 years, average height: 5–8 m, average thickness at chest height: 20 cm, coverness: E3: 45%, E2: 35%, E1: 90%, E0: 5%. E3: Quercus pubescens 3, Q. cerris 1, Tilia cordata (culture) +, Acer campestre +. E2: Crataegus monogyna 2, Cornus mas +, Tilia cordata +, Fraxinus excelsior +, Viburnum lantana +, Euonymus verrucosa 1, E. europaea +, Acer campestre +, Sambucus nigra +. 17

E1: Buglossoides purpurocaerulea 2, Melica uniflora 3, Cornus mas +, Ligustrum vulgare 2, Brachypodium pinnatum 1, Euonymus verrucosa 1, Viola hirta +, V. mirabilis 1, V. odorata +, Fragaria viridis +, Festuca valesiaca 2, Anthericum ramosum 1, Coronilla varia 1, Poa pratensis var. angustifolia +, Verbascum austriacum +, Euphorbia cyparissias +, Stachys recta +, Teucrium chamaedrys 1, Sedum maximum +, Achillea collina 1, Hypericum perforatum +, Clinopodium vulgare 1, Vincetoxicum hirundinaria +, Inula conyza +, Pulmonaria mollis +, P. officinalis +, Alliaria petiolata 3, Urtica dioica +, Galium odoratum 1, Euonymus europaea +, Geranium robertianum 1, Geum urbanum 1, Corydalis solida 2, Arum maculatum ssp. besserianum 1, Ranunculus ficaria +, Allium flavum 1, Gagea pratensis +, Campanula trachelium +, Cardaminopsis arenosa r, Veronica hederifolia 1, Centaurea stoebe +, Seseli osseum +, Erophila verna +, Sanguisorba minor +, Mercurialis paxii 1, Anemone ranunculoides +, Muscari comosum +, Potentilla arenaria +, Lathyrus vernus +. Translated by P. J. Fedor Acknowledgement Our thanks belong to E. Bednárová for the chemical analysis of the soil samples obtained from the study sites and to M. Šimurka for elaboration of the map. This research was supported by VEGA (Scientific Grant Agency of the Ministry of Education and the Slovak Academy of Sciences), grant number 1/0119/03 and 1/2344/05.

References Braun-Blanquet, J., 1964: Pflanzensoziologie. Grundzüge der Vegetationskunde. 3 Aufl. Springer Verlag, Wien–New York, 866 pp. Buday, T., Cambel, B., Maheľ, M. (eds), 1962: Explications to the detailed geological map of C.S.S.R. 1:200.000 (in Slovak). Wien – Bratislava. Geofond, Bratislava, 248 pp. Ehrendorfer, E., 1973: Liste der Gefässpflanzen Mitteleuropas. G. Fischer Verlag, Stuttgart, 320 pp. Hraško, J., Červenka, L., Facek, Z., Komár, J., Pospíšil, F., Sirový, V., 1962: Soil analyses (in Slovak). SVPL, Bratislava, 342 pp. Maheľ, M., Cambel, B., 1972: Geological map of Malé Karpaty Mts. 1:50 000 (in Slovak). Geologický ústav Dionýza Štúra, Bratislava. Michalko, J., Berta, J., Magic, D., 1986: Geobotanical map of Czechoslovak Socialist Republic. Slovak Socialistic Republic. Text part (in Slovak). VEDA, Bratislava, 168 pp. Němeček, J., Smolíková, L., Kutílek, M., 1990: Pedology and palaeopedology (in Czech). Academia, Praha, 552 pp. Petrovič, Š. (ed.), 1968: Climatic and phenological characteristics of Western Slovakia (in Slovak). Hydrometeorologický ústav, Praha, 343 pp. Petrovič, Š., 1972: Weather and climate. In Lukniš, M. (ed.): Slovakia 2. Nature (in Slovak). Obzor, Bratislava, p. 203–282. Šály, R., Bedrna, Z., Bublinec, E., Čurlík, J., Fulajtár, E., Gregor, J., Hanes, J., Juráni, B., Kukla, J., Račko, J., Sobocká, J., Šurina, B., 2000: Morphogenetic soil classification system of Slovakia (in Slovak). VÚPOP, Bratislava, Societas pedologica slovaca, 76 pp. Tarábek, K., 1980: Map 43. Climatic-geographical types. In Mazúr, E. (ed.): Atlas of Slovak Socialistic Republic (in Slovak). Geografický ústav SAV, Bratislava, p. 64. Received 30. 7. 2005

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Zlinská J., Šomšák L., Holecová M.: Ekologická charakteristika študovaných lesných spoločenstiev dubovo-hrabového vegetačného stupňa v oblasti JZ Slovenska. V práci podávame podrobnejšiu ekologickú charakteristiku skúmaného územia (vrátane geológie, klimatických pomerov, pedológie a fytocenológie). Na 10 študijných plochách sme počas 4 vegetačných sezón (1999–2002) realizovali komplexný synekologický výskum vybraných skupín bezstavovcov v epigeóne, v.odumretom dreve v rôznom štádiu rozkladu, v machoch, v dendrotelmách. Študované lesné porasty veku 40–100 rokov sa nachádzajú v orografických celkoch Malé Karpaty a Trnavská pahorkatina. Zhodne s tromi zistenými typmi dubovo-hrabových (Carpinion betuli), dubovo-cerových (Quercion confertae-cerris) a.xerotermofilných dubových lesov (Q. pubescentis-petraeae) sme analyzovali tri rôzne pôdne typy vyznačujúce sa odlišnou genézou a vlastnosťami: redziny, kambizeme a pseudogleje. Študované porasty sa líšia vekom, mierou antropického vplyvu a mierou fragmentácie. Reprezentujú jednak lesné fragmenty v kultúrnej krajine, ako aj súvislejšie lesné komplexy.

19

Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 20–36

CILIATE COMMUNITIES (Protozoa, Ciliophora) IN TREE-HOLES AND INFLUENCE OF SELECTED ENVIRONMENTAL FACTORS ON THEIR STRUCTURE EVA TIRJAKOVÁ, PETER VĎAČNÝ Department of Zoology, Comenius University, Mlynská dolina B-1, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected], [email protected] Abstract Tirjaková E., Vďačný P.: Ciliate communities (Protozoa, Ciliophora) in tree-holes and influence of selected environmental factors on their structure. Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 20–36. The structure and influence of selected ecological factors on ciliate communities in tree-holes of 3 tree species (Acer campestre, Carpinus betulus and Quercus dalechampii) have been analysed. In 136 samples 94 taxa of ciliates were found. Ciliates were not detected in 2 tree-holes only. An average of 3–6 species were presented in a single tree-hole, but the diversity was very irregular (0–15 species). Similarly, abundance was very irregular too and rapidly oscillated between 0–200,000 ex/ml. The frequency of individual species in the samples did not overlap the value of 30%. The highest frequency was recorded at the species Leptopharynx costatus (28.68%) and Sathrophilus mobilis (27.21%). Specifically, subclass Peritrichia was over-represented (notably species of the genera Propyxidium and Scyphidia). On the basis of Canonical Correspondence Analysis (CCA) (15 nominal and 2 gradient variables were analysed, 9 of them were statistically significant) 4 principal communities of ciliates have been distinguished. A tree species, volume and age of a tree-hole, absence of rotifers and other metazoans were revealed as critical factors with an influence on composition of the ciliate taxocoenoses. The results were evaluated by hierarchical classification (complete linkage) as well. Tree-holes of Quercus dalechampii have reached the highest species richness and were completely different from the others (Acer campestre and Carpinus betulus). Tree-holes with a volume under 1 ml and those over 500 ml had the lowest species diversity and a special position among the others. According to the age of tree-holes, the youngest and the oldest ones were linked in one cluster by Wishart’s index, meanwhile the special position in the freshest tree-holes only was validated by Sørensen’s index. Thus the sense of time factor on a species structure of ciliate communities in tree-holes was revealed. Key words: ciliates, communities, environmental factors, Slovakia, tree-hole (dendrotelma)

20

Introduction Ciliates are a protistan group, well adapted on surviving in a variety of freshwater habitats or the others being characterised by extreme fluctuations of moisture. Particular species are selectively tolerant to different factors like temperature, specific water chemistry, periods of drying etc. Tree-holes (dendrotelmae), cavities in tree trunks filled and supplied with rain water and dead organic matter, belong to the habitats characterised by extreme conditions, particularly as for their water chemistry (e.g. high concentration of dissolved organic compounds and low content of oxygen dissolved in water), deficiency of nutrition sources, fluctuation of temperature (overheating) and change of different moisture periods with possibility to dry up. Tree-holes are periodically or permanently filled up by rainwater. Apart from protozoans, rotifers and nematodes, variety of insect larvae, notably dipterans (Culicidae, Chironomidae and Ceratopogonidae), often occur in tree-holes (Záruba, 2004). Paradise, Dunson (1997a, b) studied relationships between insects and Protozoa in tree-holes. Considerable attention was devoted to investigation of relation of parasite and host between larvae of mosquitoes and ciliates Lambornella clarki in dendrotelmae (Wasburn et al., 1991). In spite of a relatively rich presence of Protozoa, tree-holes have not been comprehensively studied from the faunistic and ecological point of view. Data that assume the occupation of ciliate species in tree-holes are available in several papers only, e.g. Kahl (1935) noticed Colpoda cavicola, the same species was recorded by Novotny et al. (1977) as C. spiralis. Foissner (1993) mentioned other species that occurred in tree-holes. The only faunistic data on tree-holes from the territory of Slovakia with regard to Protozoa were published just by Mrva (2003) dealing with naked amoebae (Gymnamoebia). The data based on other types of telmae, with similar factors that influence inhabitation of dendrotelmae, are very insufficient. For instance phytotelmae were examined by Addicott (1974), cavernicolous Protozoa were investigated by Gittleson, Hoover (1969) and sporadic data on Diplites telmatobius in lithotelmae come from Foissner (1998). Various kinds of rainwater pools were investigated, e.g. in Austria (territory of Grossglocknergebiet) (Dingfelder, 1962; Foissner, 1980; Foissner et al., 1982), Hungary (Gelei, 1954) and in Slovakia (Matis, 1966, 1975). The investigation was focused mainly on a species spectrum of ciliates in dendrotelmae and an influence of chosen abiotic and biotic factors on a structure and dynamics of ciliated protozoa communities (e.g. species of tree, pH, volume and age of a tree-hole, inhabitation of tree-holes by other Protozoa and microscopic Metazoa). This paper is an introduction to analysis on tree-hole problems. Our research originated in a grant being concentrated on animal communities in oak-hornbeam forests of Malé Karpaty Mts (Slovakia). Several studies of various animal groups in this territory have been already published (Bulánková, Holecová, 1998; Tirjaková, 2000; Holecová, Sukupová, 2002).

21

Material and methods The samples of dendrotelmae were obtained in 2000–2002 from 10 localities of the Malé Karpaty Mts (Slovakia). Totally 136 samples of tree-holes in 3 tree species: Acer campestre (15 samples), Carpinus betulus (42 samples) and Quercus dalechampii (79 samples) were investigated. The majority of the tree-hole samples were analysed repeatedly in month intervals. Apart from a tree species, a pH value was detected with litmus paper (LACHEMA) (Table 1). Dendrotelmae have been divided into 5 size categories (as was reflected in volume of a tree-hole) as followed: (1) minimum content of rain-water (1 ml) or only moist sediment; (2) from 2 to 10 ml; (3) from 10 to 200 ml; (4) from 200 to 500 ml and (5) over 500 ml. Furthermore tree-hole water was divided into 5 colour categories, from light-coloured to dark-coloured that reflected the age of dendrotelmae (a light-coloured tree-hole water occurs in fresh dendrotelmae and dark-coloured water in long-term filled tree-holes). Samples with sediment were obtained by polyvinyl pipette and were processed in vivo immediately after the arrival to the laboratory up to 24 h. after collection. The samples were quantitatively analysed by a direct counting method using micropipette (vol. 50, 20 and 10 μl) and the data were recounted to 1 ml of sediment. The samples were not homogenised and were analysed by micropipette from 20 spot sites of a surface layer of sediment. The species of ciliates were investigated, using the live observation and protargol impregnation for permanent preparation (Foissner, 1991). The ciliates were identified using the following standard identification keys and publications: Foissner (1993), Foissner et al. (1991, 1992, 1994, 1995, 2002) and other ones cited in the references. Cluster Analysis (complete linkage) was used to evaluate a similarity of ciliate communities, according to Sørensen’s and Wishart’s indices of dissimilarity in tree-holes situated at various tree species of variable volume and age. The cluster analysis of communities was realised using the computer program NCLAS (Podani, 1993). Relationship between ciliate communities in tree-holes and chosen environmental variables was revealed in accordance with Canonical Correspondence Analysis (CCA) in the program of Canoco (Ter Braak, Šmilauer, 1998). Only 13 nominal variables: tree species – Acer campestre, Carpinus betulus and Quercus dalechampii; coloration of tree-hole water – 5 categories, from light-coloured to dark-coloured; presence or absence of flagellated protozoans, rotifers and other metazoans and 2 gradient variables (pH and volume of tree-hole) were investigated.

Results Species diversity and abundance of ciliates Totally 94 taxa of ciliates have been determined in the examined dendrotelmae: 82 species and 12 taxa identified only to a genus level (Table 2). Inhabitation of tree-holes was very irregular, often with a poor amount of species and diverse values of abundance. The frequency of ciliate species in the samples in comparison with other freshwater and terrestrial habitats reached low values only and no species overlapped the value of 30%. The highest frequency was recorded at the species Leptopharynx costatus (28.68%) and Sathrophilus mobilis (27.21%). 41 out of the 94 (43%) identified ciliate species occurred in one sample only. Ciliates were not detected (but metazoans occurred) only in 2 from a total of 136 tree-holes, in 11 samples only 1 ciliate species was recorded and in 22 samples only 2 ciliate species occurred. The highest amount (15) of ciliate species was detected in one dendrotelma only. Abundance of ciliates in the samples was very diverse as well and rapidly oscillated from 0 to 200 000 ex/ml. 22

T a b l e 1. Distribution of pH values in the investigated tree-holes. Year/Month Locality Naháč–Kukovačník Naháč–Kukovačník Lindava Horný háj grove Horný háj grove Vinosady Cajla Naháč–Katarínka 1 Naháč–Katarínka 2 Lošonský háj grove Lošonec–lom guarry Fúgelka

Tree C Q Q Q C A C Q Q C Q Q

2000 6 + +

8

5

6

7

2001 8

+++

5.5

5.5 5.6

5.8

5.1

6.5 5.9 7.4 5.4 5.1

7.2 5.8 6.1 + 4.9

+ 5.6 7.2 ++ 5.5

9 5.1

6.8 6.8 5.4 7.9 6.9;7.1 5.8 5.4 7.8 6.9 6.2 5.1 4.8 4.8 5.6 5.4

10 5.1 5.4 6.9 6.2 6.8 6.6 6.2

11 7.1 5.9 7.2 5.1 7.2 5.9 6.1 5.8

3 5.2

4 5.1

5 5.1

6 5.2

5.4 7.2

6.1

5.8

5.1 6.9

8.2 5.6 7.1 8.1 5.7 5.9 5.4

8.9 4.9 5.9 5.1 5.9

5.2 7.1 5.8 8.4 6.1 6.4 6.0

Notes: A – Acer campestre; C – Carpinus betulus, Q – Quercus dalechampii; + = samples without detected pH value

7.2 5.1 7.6 7.2 5.0 6.2

2002 7

8

10

7.1 5.4 7.6

6.7 5.3 7.1

6.6 5.2 6.9

6.0 6.9

6.6 5.1 6.6

8.2 5.2 6.7 5.3 6.2 6.1

6.8 5.3 7.3 6.1 5.8 6.1

6.6 5.5 7.1 6.2 6.2 6.1

5.6 6.2

11 5.3

12 5.9 6.2

5.9 6.9 6.9 6.3 6.4 5.4

Dominance of several species in some tree-holes was significant: Colpoda steinii, Colpidium colpoda, Dexiostoma campylum, Paramecium caudatum, Tetrahymena pyriformis, T. rostrata, Sathrophilus mobilis and Drepanomonas obtusa. Paramecium caudatum surprisingly occurred only in communities with more ciliate species and then it reached the dominance of over 70%. The most frequent ciliates in the tree-holes include (listed in the decreasing presence): Leptopharynx costatus (28.68%), Sathrophilus mobilis (27.21%), Drepanomonas obtusa (22.79%), Tetrahymena rostrata (22.06%) and Halteria grandinella (21.32%). From the qualitative point of view mainly the subclass of Peritrichia was over-represented; notably genera Propyxidium and Scyphidia achieved the highest frequency. In total 8 colpodid species, characteristic in soils and semiterrestrial environment, were recorded. Only 2 of them occurred in a frequency over 10%: Colpoda steinii (14.71%) and C. cavicola being found almost exclusively in tree-holes. The later species occurred in 11% of the investigated dendrotelmae. The number of recorded ciliate species in the tree-holes differed significantly; however in fact there was dependence between number of species and number of taken samples (Table 2). On average 3–6 species were presented in each tree-hole, but only 1 species was recorded in 8% of the investigated samples and only 2 species were recorded in 16% of the samples. Higher number of ciliate species in tree-holes was very rare with maximum of 15 species in a tree-hole.

Structure of ciliate communities On the basis of Canonical Correspondence Analysis (CCA), 4 principal communities of ciliates have been distinguished. A tree species, volume and age of a tree-hole, absence of rotifers and other metazoans were revealed as critical factors with influence on composition of ciliate taxocoenoses (Fig. 1). The community A includes the species with strongest affinity to tree-holes being taken from Quercus dalechampii, to light-coloured tree-hole water (age 1–3) and big dendrotelmae. This community may be characterised by higher species richness, composed predominantly of freshwater species (e.g. Acineria uncinata, Colpidium colpoda, Dexiostoma campylum, Frontonia leucas, Paramecium caudatum, Philasterides armatus, Steinia platystoma, Cyclidium glaucoma and Dexiotrichides centralis) and terrestrial species (e.g. Colpoda inflata, C. maupasi and Tetrahymena rostrata). Epistylis entzii and Vorticella aquadulcis complex, common in limnetic environments, have been classified as characteristic peritrichs of the community. The community B was formed in tree-holes of Carpinus betulus, in dark-coloured tree-hole water (age 5) and small dendrotelmae. This community was poor in ciliate species (16) and was characterised by predominance of euryoecious and terrestrial species (e.g. Cinetochilum margaritaceum, Drepanomonas revoluta, D. obtusa, D. exigua, Chilodonella uncinata and Colpoda henneguyi) and minimal presence of limnetic species (e.g. Tetrahymena pyriformis complex). Vorticella infusionum, common in limnetic environments and ordinary in soils, has been considered as a characteristic peritrich here. 24

T a b l e 2. Distribution and frequency (%) of the 95 indentified ciliate taxa being found in 136 samples from tree-holes at 10 sites. Species Acineria uncinata T u c o l e s c o, 1962 Acropisthium mutabile P e r t y, 1852 Blepharisma sp. Cinetochilum margaritaceum (E h r e n b e r g, 1831) P e r t y, 1852 Coleps hirtus (O.F. M u e l l e r, 1786) N i t z s c h, 1827 Colpidium colpoda (L o s a n a, 1829) Colpoda cavicola K a h l, 1935 Colpoda cucullus (O.F.M u e l l e r, 1773) G m e l i n, 1790 Colpoda henneguyi F a b r e-D o m e r g u e, 1889 Colpoda inflata (S t o k e s, 1884) K a h l, 1931 Colpoda maupasi E n r i q u e z, 1908 Colpoda reniformis K a h l, 1931 Colpoda steinii M a u p a s, 1883 Cyclidium elongatum (S c h e w i a k o f f, 1889) S c h e w i a k o f f, 1896 Cyclidium glaucoma O.F.M u e l l e r, 1773 Cyclidium muscicola K a h l, 1931 Cyrtohymena candens K a h l, 1932 Cyrtohymena quadrinucleata (D r a g e s c o et N j i n é, 1971) F o i s s n e r, 1989 Cyrtolophosis muscicola S t o k e s, 1885 Dexiostoma campylum (S t o k e s, 1886) J a n k o w s k i, 1967 Dexiotrichides centralis (S t o k e s, 1885) K a h l, 1931 Drepanomonas dentata F r e s e n i u s, 1858 Drepanomonas exigua P e n a r d, 1922 Drepanomonas obtusa P e n a r d, 1922 Drepanomonas revoluta P e n a r d, 1922 Drepanomonas sphagni K a h l, 1931 Enchelys gasterosteus K a h l 1926 Epispathidium sp. Epistylis entzii S t i l l e r, 1935 Epistylis sp. Euplotes affinis (D u j a r d i n, 1841) K a h l, 1932 Euplotes muscicola K a h l, 1932 Frontonia angusta K a h l, 1931

1 Q + + + + + + -

2 C + + + + + + -

3 Q + + + + + + + + -

C + + + + -

Q + + + + + + -

4 A + + + + + + + + +

5 C + + + -

6 Q + + + + + + + + + -

7 Q + + + + + + + + + + -

8 C + + + + + + + + + -

9 10 % Q Q + - 0.73 - 0.73 - 0.73 + - 4.41 + - 0.73 + - 5.88 - + 11.03 + - 2.21 - 0.73 - 1.47 + - 2.21 - 0.73 + - 14.71 + - 2.21 + - 11.76 - 0.73 - 0.73 - 0.73 - 2.94 + - 2.21 - 0.73 - 0.73 - 0.73 + - 22.79 - 5.88 - 1.47 - 0.73 - 0.73 - 0.73 - 0.73 + - 1.47 - 1.47 - 1.47

T a b l e 2. (Continued) Species Frontonia depressa (S t o k e s, 1886) K a h l, 1931 Frontonia leucas (E h r e n b e r g, 1833) E h r e n b e r g, 1838 Gastrostyla steini E n g e l m a n n, 1862 Glaucoma scintillans E h r e n b e r g, 1830 Gonostomum affine (S t e i n, 1859) S t e r k i, 1878 Gonostomum kuehnelti F o i s s n e r, 1987 Halteria grandinella (O.F.M u e l l e r, 1773) D u j a r d i n, 1841 Hemicycliostyla sphagni S t o k e s, 1886 Hemisincirra gellerti (F o i s s n e r, 1982) F o i s s n e r, 1984 Hexotricha caudata L a c k e y, 1925 Histriculus vorax (S t o k e s, 1891) Holosticha muscorum (K a h l, 1932) F o i s s n e r , 1982 Holosticha sp. Homalogastra setosa K a h l, 1926 Chilodonella uncinata (E h r e n b e r g, 1838) S t r a n d, 1928 Leptopharynx costatus M e r m o d, 1914 Leptopharynx eurystomus (K a h l 1931) F o i s s n e r, 1988 Metopus hasei S o n d h e i m, 1929 Metopus minor K a h l, 1927 Odontochlamys gouraudi C e r t e s, 1891 Ophryoglena flava E h r e n b e r g, 1833 Ophryoglena oblonga G a j e v s k a j a, 1927 Ophryoglena sp. Opisthonecta henneguyi F a u r e-F r e m i e t, 1906 Oxytricha setigera S t o k e s, 1891 Oxytricha similis E n g e l m a n n, 1862 Oxytricha sp. Paramecium caudatum E h r e n b e r g, 1833 Paraurostyla macrostoma F o i s s n e r, 1982 Peritrichia gen. sp. Philasterides armatus (K a h l, 1926) K a h l, 1931 Pithothorax processus K a h l, 1926 Plagiocampa metabolica (K a h l, 1926) K a h l, 1930

1 Q + + + + + -

2 C + + + + -

3 Q + + + + + + + + + + + + + + -

C + + -

Q + + + + + + + + + -

4 A + + + + + + + + + + + + + -

5 C + + + + + -

6 Q + + + + + + + + + + + + + +

7 Q + + + + + + + + + + + + + -

8 C + + + + + + + + + +

9 10 % Q Q - 0.73 + - 0.73 + - 0.73 - 8.82 + - 5.14 - 2.21 - 21.32 - 0.73 - + 0.73 - 0.73 + - 0.73 - 1.47 - 0.73 - 2.94 + - 17.65 + - 28.68 - 3.68 - 2.21 - 13.97 - 2.21 - 0.73 - 0.73 - 0.73 + - 8.09 - 2.21 - 6.62 - 0.73 + - 11.03 - 1.47 - 3.68 - 1.47 - 0.73 - 1.47

T a b l e 2. (Continued) 1 2 3 4 5 6 7 8 9 10 % Q C Q C Q A C Q Q C Q Q Platyophrya spumacola K a h l, 1927 - + - 0.73 Platyophrya vorax K a h l, 1926 - + - + + - + - 2.94 Plesiocaryon elongatum (S c h e w i a k o f f, 1892) F o i s s n e r, A g a t h a et B.e.r.g.e.r, 2002 + + + - + + + + 9.56 Propyxidium spp. - + + + + + + + - 11.03 Protospathidium vermiforme F o i s s n e r, A g a t h a & B e r g e r, 2002 - + - 0.73 Pseudocohnilembus pusillus (Q u e n n e r s t e d t, 1869) F o i s s n e r et W.i.l.b.e.r.t, 1981 - + - 0.73 - + - 0.73 Pseudoglaucoma muscorum K a h l, 1931 Pseudoholophrya terricola B e r g e r, F o i s s n e r et A d a m, 1984 + - 0.73 Pseudochilodonopsis mutabilis F o i s s n e r, 1981 + - 1.47 Pseudochilodonopsis sp. - + + - 0.73 Pseudomicrothorax agilis M e r m o d, 1914 - + - 0.73 Rhabdostyla pyriformis P e r t y, 1852 - + - 0.73 Sathrophilus mobilis (K a h l, 1926) C o r l i s s, 1960 + - + + + + + + + + + - 27.21 Scyphidia spp. + + + - + + + + + + + - 18.38 Steinia platystoma (E h r e n b e r g, 1831) D i e s i n g, 1866 - + - 0.73 Sterkiella histriomuscorum (F o i s s n e r, B l a t t e r e r, B e r g e r et K.o.h.m.a.n.n, 1991) F.o.i - + - + + - 2.94 s s n e r, B l a t t e r e r, B e r g e r et K o h m a n n, 1991 Stylonychia pustulata (O.F.M u e l l e r, 1786) E h r e n b e r g, 1835 + - + + - + - + - + + - 13.23 Tachysoma pellionellum (O.F. M u e l l e r, 1773) B o r r o r, 1972 - + - + - 1.47 Telotrochidium sp. - + - 1.47 Tetrahymena edaphoni F o i s s n e r, 1986 - + - 0.73 Tetrahymena pyriformis complex - + + + + - + - 5.88 Tetrahymena rostrata (K a h l, 1926) C o r l i s s, 1952 - + + + + - + + + + - 22.06 Urotricha globosa S c h e w i a k o f f, 1892 - + - 0.73 Vorticella aquadulcis complex - + - 0.73 Vorticella astyliformis F o i s s n e r, 1981 - + - + - + - 0.73 Vorticella infusionum D u j a r d i n, 1841 - + - + - 2.94 Vorticella microstoma E h r e n b e r g, 1830 + - + - + - 3.67 Vorticella spp. - + + + - 4.41 Number of taxa 16 12 32 11 22 32 12 33 40 30 32 3 Number of samples 7 8 19 4 7 15 15 16 17 16 11 2 Notes: 1 – Naháč–Kukovačník; 2 – Lindava; 3 – Horný háj grove; 4 – Vinosady; 5 – Cajla; 6 – Naháč–Katarínka 1; 7 – Naháč–Katarínka 2; 8 – Lošonský háj grove; 9 – Lošonec–lom quarry; 10 – Fúgelka; A – Acer campestre; C – Carpinus betulus, Q – Quercus dalechampii; + = present, - = absent Species

Fig. 1. Ordination diagram based on Canonical Correspondence Analysis. Eigenvalues of two first axes of CCA are λ1 = 0.340 and λ2 = 0.298. The first axes account for 42.2% of the total variance of the species–environment relation. Explanations: nominal environmental variables are represented by squares and gradient environmental variable – volume of a tree-hole by arrow. A, B, C, D – community; Ac – Acer campestre; Cb – Carpinus betulus; Qr – Quercus dalechampii; Rotifera A – absence of rotifers; M – absence of metazoans; Age 1, 3 and 5 – coloration of a tree-hole (light-coloured, moderately-coloured and dark-coloured).

The community C refers to tree-holes in Acer campestre, and volume and coloration of tree-hole water may be classified as crucial environmental variables for its formation. A species spectrum was rather low (12 species) with equipollent number of freshwater (e.g. Coleps hirtus, Euplotes affinis, Stylonychia pustulata and Urotricha globosa) and terrestrial (e.g. Colpoda cucullus, Gonostomum affine, Leptopharynx eurystomus, Oxytricha similis and O. setigera) species. Vorticella microstoma represented the characteristic peritrich of this community. The fourth community D included a poor spectre of species (13 species), where the key role in its formation refers to environmental variables such as absence of rotifers and other metazoans; tree species, volume and coloration of tree-hole water appeared irrelevant. The community was formed by bacteriovorous limnetic and terrestrial species (e.g. Glaucoma scintillans, Ophryoglena spp., Scyphidia sp., Opisthonecta hennegyui and Cyclidium elongatum, Colpoda steinii, Leptopharynx costatus, Halteria grandinella, Platyophrya vorax, P. spumacola, Cyclidium muscicola, Gonostomum kuehnelti, Tetrahymena edaphoni and Euplotes muscicola). Vorticella astyliformis, the most frequent peritrich in soil world-wide, was the species characteristic for the community. 28

Influence of chosen environmental factors on a structure of ciliate communities On the basis of CCA ordination, we assume that the most important environmental variables were represented by a tree species and volume of a tree-hole. The communities lined along the first axis in the ordination diagram according to tree species. This axis could be characterised as an axis of specific substances leached out of wood. The second axis represents a change in volume of a tree-hole and position along this axis indicates an increased or decreased ratio of freshwater and terrestrial species in the communities. This is a reason of a position of the community A (predominance of limnetic species) on the middle bottom of the ordination diagram and of the community B (predominance of terrestrial species) in the middle above (Fig. 1).

Fig. 2. Cluster analysis, Sørensen’s (A) and Wishart’s (B) indices of dissimilarity, Complete linkage of the ciliate communities in tree-holes from various tree species. Ac – Acer campestre; Cb – Carpinus betulus; Qd.– Quercus dalechampii.

A tree species has been considered as one of the crucial factor in formation of ciliate communities in dendrotelmae (Fig. 2). The community in telmae of Quercus dalechampii was detached on a high level of faunistic dissimilarity (55% by Sørensen’s index and 75% by Wishart’s index) in accordance Hierarchical Cluster Analysis (complete linkage). Totally, 75 ciliate species were recorded in tree-holes of this tree, with predominance of limnetic (e.g. Cyclidium glaucoma, Glaucoma scintillans, Opisthonecta henneguyi, Paramecium caudatum and Vorticella microstoma) and ubiquitous species (e.g. Colpoda steinii, Halteria grandinella, Chilodonella uncinata, Leptopharynx costatus, Metopus minor, Sathrophilus mobilis and Vorticella infusionum). The following enumerated species were recorded in oak tree-holes only: Paramecium caudatum, Opisthonecta henneguyi, Acineria uncinata, Acropisthium mutabile, Enchelys gasterosteus, Epistylis entzii and Frontonia leucas. The dendrotelmae of trees Carpinus betulus and Acer campestre represented a separate cluster on approximately 45% of qualitative and quantitative level of faunistic dissimilarity. Totally 40 and 36 species were recorded in both the hornbeam and maple tree-holes, some of them are common in soils (e.g. Cinetochilum margaritaceum, Colpoda inflata, C. henneguyi, Drepanomonas revoluta, D. sphagni, D. dentata, D. obtusa, Gonostomum kuehnelti and Metopus hasei) some autochthonous in soil (e.g. Protospathidium vermiforme and Epispathidium sp.). 29

Fig. 3. Cluster analysis, Sørensen’s (A) and Wishart’s (B) indices of dissimilarity, Complete linkage of the ciliate communities in various volume of tree-holes. (1) minimum content of rain-water (1 ml) or only moist sediment; (2) vol. from 2 to 10 ml; (3) vol. from 10 to 200 ml; (4) vol. from 200 to 500 ml and (5) vol. over 500 ml.

Volume of a tree-hole was shown as an important factor with an influence on structure of ciliate communities, especially in extreme cases: dendrotelmae with a minimum content of rain-water (1 ml) or with only moist sediments and dendrotelmae with volume over 500 ml. They were evaluated by Cluster Analysis as well (Fig. 3): small tree-holes (vol. 1 ml) were detached on a high level of faunistic dissimilarity (72% by Sørensen’s index and 88% by Wishart’s index) because of a low species richness (14) and presence of very rare ciliates in dendrotelmae (e.g. Protospathidium vermiforme and Hemisincirra gellerti). Occurrence of Colpoda species (e.g. C. cavicola, C. henneguyi and C. steinii) in lower abundance in small tree-hole was characteristic. The big tree-holes may be considered as a special category (vol. over 500 ml) because of a lower number of recorded species (27), than in dendrotelmae with volume from 10 to 500 ml and because of occurrence of a Vorticella aquadulcis complex, being recorded in big tree-holes only. A separate cluster refers to the tree-holes with volume less than 10 ml, 10–200 ml and 200–500 ml. The category mentioned above includes the typical variety of peritrichs and hymenostomes and decreasing number of colpodid species in dendrotelmae with volume 200–500 ml. In the 10 ml tree-holes the highest abundance of Vorticella astyliformis was achieved. The maximum number of peritrich species (e.g. Epistylis entzii, Opisthonecta henneguyi, Propyxidium sp. and Vorticella infusionum) was recorded in the tree-holes with their volume from 10 to 200 ml. Vorticella microstoma may be classified as the dominant peritrich in the tree-holes with a volume from 200–500 ml. Maximum number of colpodids (e.g. Colpoda steinii, C. maupasi, C. reniformis and C. cavicola) occurred in the dendrotelmae with their volume of 10–200 ml. The 200–500 ml tree-holes were detached on 46% level of faunistic dissimilarity by Sørensen’s index from the tree-holes with their volume of 10 ml and 10–200 ml, because of presence of rare species, such as Gastrostyla steinii, Epispathidium sp., 30

Ophryoglena spp. and Drepanomonas dentata. Further the tree-holes with a volume of 10–200 ml were detached on 50% level of faunistic dissimilarity by Wishart index from the tree-holes with their volume of 10 ml and 200–500 ml. Peritrich species Epistylis entzii and several other species (e.g. Metopus hasei, Colpoda reniformis, Hexotricha caudata, Holosticha spp. and Plagiocampa metabolica) occurred in the 10–200 ml dendrotelmae only, whereas they were absent in the above mentioned volumes of tree-holes.

Fig. 4. Cluster analysis, Sørensen’s (A) and Wishart’s (B) indices of dissimilarity, Complete linkage of the ciliate communities in various coloured tree-holes (reflected in age of tree-holes). (1) light-coloured water appears in fresh dendrotelmae and (5) dark-coloured in long-term filled tree-holes.

Hierarchical Cluster Analysis (complete linkage, Sørensen’s and Wishart’s indices) according to a species spectrum was used to evaluate dissimilarity in age of dendrotelmae (as reflected in coloration of tree-hole water) (Fig. 4). The predominance of peritrichs (e.g. genus Scyphidia achieved the highest abundance in the dendrotelmae of age 2, genus Vorticella in the tree-holes of ages 2–3 and genus Epistylis in the holes of age 3) was characteristic for the light-coloured (ages 1–3) tree-hole water. The amount of colpodid species (e. g. Colpoda cavicola, C. maupasi and C. steinii) increased in older dendrotelmae (age 4) and presence of colpodids and hypotrichs (e.g. C. inflata, C. reniformis, C. steinii, Gonostomum affine, Histruculus vorax, Hemisincirra gellerti, Cyrtohymena quadrinucleata and Paraurostyla macrostoma) was characteristic to dark-coloured (age 5) tree-hole water. The ciliate communities, being formed in tree-holes of ages 1 and 3, were detached on the 57% level of faunistic similarity by Sørensen’s index using Cluster Analysis. Predominance of freshwater species (e.g. Dexiostoma campylum and Enchelys gasterosteus) and peritrich species (e.g. Epistylis entzii, Opisthinecta henneguyi, Propyxidium sp. and Vorticella infusionum) is typical for the dendrotelmae being aged above. The communities in tree-holes of ages 2 and 4 were detached on a higher level of faunistic similarity (64%) and the community formed in tree-holes of age 5 was attached on 62% level of similarity. The similarity of ages 2 and 4 was caused by presence of euryoecious species of the genera Colpoda and 31

Drepanomonas (e.g. Colpoda steinii, Drepanomonas revoluta and D. obtusa) and freshwater species (e.g. Colpidium colpoda, Glaucoma scintillans and Paramecium caudatum). The tree-holes of ages 1 and 5 were detached according to a qualitative and quantitative structure of ciliate communities (Wishart’s index); characterised by presence of several common r-selective species (e.g. Leptopharynx costatus and Drepanomonas obtusa), but under their different abundance (58% dissimilarity). The communities being formed in the dendrotelmae of ages 2 and 4 were attached on high level of faunistic similarity (approximately 63%) by Sørensen’s and Wishart’s indices, as it seems from the Fig. 4. The community in the tree-holes of age 3 was attached to the tree-holes of ages 2 and 4 on 52% level of dissimilarity (Wishart’s index), because of occurrence of following peritrichs: Opisthonecta henneguyi, Vorticella infusionum and V. astyliformis and absence of V. microstoma, being common in the dendrotelmae of ages 2 and 4.

Discussion Tree-holes represent an extraordinary interesting habitat, because of broad qualitative and quantitative differences in inhabitation compared with other limnetic habitats, but amongst tree-holes as well. Tree-holes can be characterised as habitat, where freshwater and terrestrial communities of ciliates meet together and present specific habitat with very low frequency of presence of individual species. No species were recorded in more than 30% of investigated dendrotelmae, so it seems difficult to establish a specific community of ciliates in tree-holes generally. An interesting fact hints at the absence of euryoecious genus Aspidisca, with a high frequency in limnetic habitats. This genus was in tree-holes substituted by genera Drepanomonas and Leptopharynx, preferring terrestrial habitats. On the contrary, typical freshwater species such as Paramecium caudatum, Dexiostoma campylum and Colpidium colpoda with their frequency at the values of 2.21–11% were recorded, however they did not occur in soils (Foissner, 1981, 1987; Foissner et al., 2002 etc.). The species of genera Trochilia and Chlamydonella, common in limnetic habitats, were absent as well. In aspect of several similar abiotic factors with influence on periodical rain-pools and dendrotelmae, significant similarity of these two habitats could be expected. But our results indicate relevant differences of ciliate communities in these two specific habitats. Periodical rain-pools possess a more diverse species spectrum, e.g. Foissner et al. (1982) recorded 143 ciliate species in 37 rain-pools in Alps (Austria), of which only 17 occurred in the investigated dendrotelmae. Furthermore, in 77 samples from small, predominantly periodical waters of various types (little standing water – Kleingewässer) in the territory of Hohe Tauern (Austria) Foissner (1980) recorded 194 species of ciliates, of which only 24 occurred in our investigated samples. He also stated a relatively low presence of ciliate species in the samples (only 8 species with the presence over 40%, but Oxytricha falax with even 69%). According to Gelei (1954) and Dingfelder (1962) the typical genera of periodical pools include the following: Phascolodon, Paruroleptus and Astylozoon. None of them occurred in our samples. The 32

characteristic community in dendrotelmae may contain Colpoda spp., Leptopharynx costatus, Satrophilus mobilis and various peritrichs. In terms of the recorded number and abundance of ciliate taxa, the inhabitation of tree-holes was very different. In spite of this, ciliates represent rather a stable component of a protozoan community in dendrotelmae. Only in 2 of all the investigated samples (i.e. 1.47%) no ciliates were recorded. Compared with occurrence of active gymnamoebae in dendrotelmae, Mrva (2003) stated that almost 40% of the samples had contained no amoebae. Parallelly, he referred to a low diversity of naked amoebae compared with other water habitats (maximum 9 taxa per individual dendrotelma). Furthermore, Mrva (2004) stated that particularly ubiquitous species of amoebae were characteristic in tree-holes. He supposed that in aspect of unsuitable conditions (overheating, possibility to dry up, high concentration of dissolved organic compound etc.) euryoecious species from other groups of Protozoa would be predominant, what has been actually proved by our results. Most of the species with high frequency of occurrence belong to euryoecious ciliates (e.g. Leptopharynx costatus, Sathrophilus mobilis, Drepanomonas obtusa, Colpoda steinii and Tetrahymena rostrata), being well adapted to extreme conditions of environment. Rather a low diversity of ciliates in tree-hole was recorded in a comparison with other limnetic habitats (0–15 species). Only 3–6 ciliate species occurred in most of the dendrotelmae, what are relatively low values compared with other habitats. In terms of abundance, significant differences in dendrotelmae are present; Mrva (2004) stated 0–1,850 ex/ml at naked amoebae, more extreme values were achieved by ciliates (0–200,000 ex/ml). Abundance of ciliates was very diverse, there was a frequent ciliate mass outbreak, however a statistically significant factor with an influence on this outbreak was not recovered. Apart from ciliates, the other groups of Protozoa or rotifers at the same time in the same tree-hole were in outbreak. The other statistical significant dependence between mass outbreak of ciliates and other protozoan groups or microscopic metazoans was not proved too, in spite of a nutrition competition. This effect was observed in simulated tree-holes, big ciliates and flagellates were selectively eliminated by water beetles and dipteran larvae (Paradise, Dunson, 1997b). Záruba (2004) stated that pH values and chemistry in a tree-hole were rather stable. In spite of this, a wide rang of pH values (4.8–8.9) and significant oscillation during a year were recorded in dendrotelmae. Acid reaction (pH 4.8–6.6, mostly about pH 5) was characteristic in hornbeam tree-holes; in oak tree-holes it achieved the highest deviations (pH 4.8–8.4), but slightly acid or neutral reaction. Maple tree-holes provide neutral or alkaline reaction (pH 6.5–9.9). It is very interesting, that in aspect of significant oscillation of pH values, this factor was not statistically significant and does not participate on a structure of ciliate communities in tree-holes, which is very important. An indirect influence of pH can be expected through relation between ciliate communities and trees, being characterised by predominance of particular pH value. Paradise, Dunson (1997a) found out a significant influence of pH value in simulated tree-holes on formation of protozoan communities with interaction of insects. Ciliates achieved higher values of density at high pH values and flagellates at low pH values. In some cases, mass outbreak of several ciliate species at a high pH values was recorded, but we can not generalise this phenomenon. 33

Tree species turned to be statistically significant, with important influence on formation of communities. Tree-holes formed in Quercus dalechampii were distinguished by the richest species spectrum, highest quotient of freshwater species and occurrence of species being absent in other kinds of tree-holes. This can be connected with quality of wood; oak wood may be characterised as hard and low intensity of leaching the resin and other substances can be expected. Nevertheless, dendrotelmae in 3.tree species only were investigated. This is supposed to be an introduction to the next analyses. The age of a tree-hole, derived from coloration, turned to be a statistically important variable, with an influence on formation of ciliate communities. Freshwater species were predominant in freshly filled (light-coloured) tree-holes. Later with further leaching wood and with growing concentration of organic matter, these species were driven out by better adapted small terrestrial species of the genera Colpoda, Drepanomonas and Leptopharynx. After the filling, r-selective species of the genus Colpoda appeared in a short period, and were soon driven out by K-strategists of the genera Glaucoma, Paramecium, Stylonychia and Tetrahymena. In a period of decreasing volume and ageing of a tree-hole, number of r-strategists rose. These results well correspond with succession in terrestrial and semiterrestrial habitats (Foissner, 1987; Tirjaková, 1997; Bartošová, Tirjaková, 2005 etc.). Small-sized species (e.g. Leptopharynx costatus, Colpoda steinii, Drepanomonas obtusa etc.) were over-represented in dark-coloured dendrotelmae water and big species (e.g. Paramecium caudatum, Stylonychia pustulata and other hypotrichs) were predominant in light-coloured tree-hole water and big tree-holes. Paradise, Duton (1997b) noticed, that this phenomenon referred to predation or trophic competition. Totally, 11 taxa (11.6%) were identified to the genus level only. So we suppose that several ciliate species from tree-holes are still undescribed. Because of a very rare occurrence, low abundance of these species and low number of impregnated specimens, we were unable to describe them. In spite of that, more environmental factors influence the formation of ciliate communities in tree-holes; each dendrotelma is remarkable and unique by inhabitation and environmental parameters. In conclusion, we state that our results represent the introduction to this problem and tree-holes are perspective habitats for further investigations in field of ecology and taxonomy of Protozoa. Translated by authors Acknowledgement The study was realised with the support of grant No. 1/0119/03 provided by the Slovak Scientific Grant Agency VEGA. We would like to thank M. Holecová, Z. Krumpálová and I. Országh for collecting the material. Further we are grateful to D. Matis and M. Mrva for valuable comments on the text.

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Kahl, A., 1935: Urtiere oder Protozoa I: Wimpertiere oder Ciliata (Infusoria) 4. Peritricha und Chonotricha. Tierwelt Dtl., 30, p. 651–886. Matis, D., 1966: Bemerkung über die Infusorienfauna der Regentümpel vom St. Georg–Moorwald. Acta Fac. Rerum. Nat. Univ. Comenianae, Zool., 12, p. 235–239. Matis, D., 1975: Infusoria of the order Hypotrichida in some localities in the vicinity of Bratislava. Biológia (Bratislava), 30, p. 583–588. Mrva, M., 2003: Diversity of active gymnamoebae (Rhizopoda, Gymnamoebia) in dendrotelmae of oak-hornbeam forests in Malé Karpaty Mts (Western Slovakia). Protistology, 3, p. 121–125. Mrva, M., 2004: Naked amoebae (Rhizopoda: Lobosea, Filosea) of chosen forest habitats in Malé Karpaty Mts (in Slovak). Ph.D. Thesis, Comenius University, Bratislava, 101 pp. Novotny, R.T., Lynn, D.H., Evans, F.R., 1977: Colpoda spiralis sp. n., a colpodid ciliate found inhabiting treeholes (Colpodida, Ciliophora). J. Protozool., 24, p. 364–369. Paradise, C.J., Dunson, W.A., 1997a: Effects of pH and sulphfate on insects and protozoans inhabiting treeholes. Arch. Environ. Contamin. Toxicol., 33, p. 182–187. Paradise, C.J., Dunson, W.A., 1997b: Insects species interactions and resource effects in treeholes: are helodid beetles bottom-up facilitators of midge populations? Oecologia, 109, p. 303–312. Podani, J., 1993: Syn-tax. Version 5.0. Computer programs for Multivariate Data Analysis in Ecology and Systematics. User’s guide. Scientia Publishing, Budapest, 104 pp. Ter Braak, C.J.F., Šmilauer, P., 1998: CANOCO References Manual and User’s Guide to Canoco for Windows: Software for Canonical Community Ordination (version 4). Microcomputer Power. New York, 352 pp. Tirjaková, E., 1997: Structure and dynamics of communities of ciliated protozoa (Ciliophora) in field communities. Succesion, genus–species relationships, nutrition. Ekológia (Bratislava), 16, p. 243–252. Tirjaková, E., 2000: Occurrence of Bunonema richtersi Jägerskiöld, 1905 (Nematoda, Bunonematidae) on some localities of Little Carpathians (Slovakia) (in Slovak). Folia Faunistica Slovaca, 5, p. 55–59. Washburn, J.O., Anderson, J.R., Mercer, D.R. 1991: Parasitism of newly-hatched Aedes sierrensis (Diptera: Culicidae) larvae by Lambornella clarki (Ciliophora: Tetrahymenidae) following habitat flooding. J. Invertebr. Pathol., 58, p. 67–74. Záruba, P., 2004: Tree-holes – special freshwater insect’s habitat (in Czech). Živa, 5, p. 221–222. Received 30. 7. 2005

Tirjaková E., Vďačný P.: Spoločenstvá nálevníkov (Protozoa, Ciliophora) v dendrotelmách a vplyv vybraných environmentálnych faktorov na ich štruktúru. Sledovali sme spoločenstvá nálevníkov v dendrotelmách 3 druhoch drevín (Acer campestre, Carpinus betulus, Quercus dalechampii). V 136 vzorkách sme určili 94 taxónov nálevníkov. Iba v 2 dendrotelmách sa nálevníky nevyskytli. Diverzita bola veľmi nepravidelná od 0–15 druhov v.jednej dendrotelme, väčšinou sa vyskytovalo 3–6 druhov. Podobne i početnosť bola veľmi nepravidelná a prudko kolísala od 0–200 000 ex/ml. Frekvencia výskytu vo vzorkách ani u jedného druhu neprekročila hodnotu 30%. Najvyššiu frekvenciu sme zaznamenali u.druhov Leptopharynx costatus (28,68%) a Sathrophilus mobilis (27,21%). Bohato zastúpená bola podtrieda Peritrichia (najmä zástupcovia rodov Propyxidium a.Scyphidia. Na základe CCA anylýzy (testovaných 15 nominálnych a 2 gradientové premenné, z.ktorých 9.bolo štatisticky významných) boli rozlíšené štyri základné spoločenstvá nálevníkov. Ako rozhodujúce faktory ovplyvňujúce zloženie taxocenóz pôsobili druh dreviny, veľkosť (objem) a vek telmy, prítomnosť Rotifera a iných Metazoa. Tieto výsledky boli potvrdené aj hierarchickou klasifikáciou. Druhovo najbohatšie a od ostatných najodlišnejšie boli telmy odobraté z Quercus dalechampii, druhovo najchudobnejšie a osobitné postavenie mali telmy s objemom pod 1 ml a telmy s objemom nad 500 ml. Wishartov index na základe veku telmy zlúčil do jedného zhluku najmladšie a najstaršie telmy. Sørensenov index tiež potvrdil osobitné postavenie najmladších teliem a tým aj význam časového faktora na formovanie spoločenstiev nálevníkov v dendrotelmách.

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Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 37–50

SELECTED ECOLOGICAL CHARACTERISTICS OF CILIATE COMMUNITIES (Protozoa, Ciliophora) IN DECAYING WOOD MASS IN THE MALÉ KARPATY MOUNTAINS PAVLA BARTOŠOVÁ, EVA TIRJAKOVÁ Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská dolina B-1, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected], [email protected] Abstract Bartošová P., Tirjaková E.: Selected ecological characteristics of ciliate communities (Protozoa, Ciliophora) in decaying wood mass in the Malé Karpaty Mountains. Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 37–50. In total 28 samples of 14 tree species were collected from 13 localities of Slovakia during the research of ciliate communities in decaying wood mass in 2001–2004. Generally 58 ciliate species (Protozoa: Ciliophora) were determined in our samples and 9 species were recorded in Slovakia for the first time. Apart from the species diversity the research has included analyses on structure of systematic and feeding groups, genus–species relationships and forming the communities in dependence on tree species. Colpodids predominated in systematic categories and bacteriovores were dominant in feeding groups. The maximum of species (8) was noted in the genus Colpoda. Hierarchical classification based on identity of ciliate species on trees has differentiated two large groups of tree communities. However the tree species probably does not seem to play a very important role in forming ciliate communities. Even no special relation (or differences of species spectrum) has been proved between deciduous and coniferous trees. Key words: ciliates, communities, wood mass, decay, Malé Karpaty Mountains

Introduction Rocks, mushrooms, mosses and lichens growing at trees and decaying wood mass have their own characteristic fauna. The common feature of the mentioned microhabitat fauna has been derived from general soil fauna (Wallwork, 1976). A lot of studies have been published on decaying processes in wood. They are mainly concentrated on the role of various arthropod groups in decaying wood (e. g. Wallwork, 1976; Speight, 1989; Dajoz, 2000). The importance of Protozoa (ciliates too) in decaying processes has been studied sporadically. As the majority of ciliates living in terrestrial habitats are 37

bacteriovores (Brunberg Nielsen, 1968; Laminger, 1980; Nikolyuk, 1965, 1969 etc.), their role in the wood decay is mainly releasing the nutriments after the consumption by bacteria. Several authors have hinted at decaying processes in soil (Bamforth, 1973; Stout, 1973). An increased supply of nutriments positively influences the abundance of microfauna, which is trophicaly associated with successive wood decay caused by macrofauna. Despite the rapid progress of terrestrial and semiterrestrial ciliate research in the recent years, there have not been published any entire data on ciliates living in decaying wood mass. Just several studies present some data on occurrence of ciliates in wood, however mainly with no details on studied tree species (Wenzel, 1953; Blatterer, Foissner, 1988; Foissner, 1993a). In fact there are only several authors determining relevant tree species (Miteva, 1992; Foissner, 1993b; Foissner, 1994; Foissner et al., 2002). Some authors paid attention to the occurrence of ciliates living in the terrestrial habitats of the Malé Karpaty Mts, for instance Mrva, Matis (2000) and Mrva (2003) studied gymnamoebae (Protozoa: Amoebozoa) in the leaf-litter and dendrotelmae of oak-hornbeam forests. The large research on litter, soil and moss has contributed to the knowledge on species diversity of ciliates, amoebas (Tirjaková et al., 2002) and arthropods (Holecová et al., 2002, 2005) in oak-hornbeam forests of the Malé Karpaty Mts.

Material and methods In total 28 samples of bark and wood mass in various stages of decay were collected from 13 localities of Slovakia during the years 2001–2004 (Table 1). The stage of wood decay (SD) was considered according to the following scale: 1 – bark of living steady tree, 2 – decaying wood mass without contact with soil, 3 – decaying wood mass in contact with soil and with well-preserved wood structure, 4 – decaying wood mass in contact with soil and in far-gone stage of decay, 5 – decaying wood mass in contact with soil without preserved wood structure. The material was collected from 14 tree species, as followed: Acer campestre (Ac), A. pseudoplatanus (Aps), Alnus glutinosa (Ag), Betula pendula (Bp), Carpinus betulus (Cb), Cerasus avium (Ca), Fagus sylvatica (Fs), Fraxinus excelsior (Fe), Picea excelsa (Pe), Pinus sylvestris (Ps), Quercus dalechampii (Qd), Q. petraea (Qp), Q. pubescens (Qpu) and Q. robur (Qr). Resting cysts of ciliates from air-dried samples were reactivated using the „non-flooded Petri dish method” (Foissner, 1987c). Ciliate species were determined by combining „in vivo“ observations and protargol impregnation (Foissner, 1991). Mainly the publications of Foissner (1981b, 1982, 1984, 1987a, b, 1993a, b, 1999) and Foissner et al. (2002) were used to determine the ciliate species. The frequency of ciliate taxa was categorised according to Schwerdtfeger (1975): species of the 1st class (0–10%), 2nd class (11–25%), 3rd class (26–45%), 4th class (46–70%) and 5th class (71–100%). Communities of ciliates were compared to each other using hierarchical classification, based on identity of species representation (Sørensen’s index of dissimilarity, complete linkage method). The cluster analysis was made with the help of the program Syntax (Podani, 1993).

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T a b l e 1. The list of studied localities. No. Locality 1 Borinka–under Pajštún reservation 2 Bratislava–Bystrické forest park 3 Bratislava–Karlova Ves under Sitina 4 Bratislava–Koliba 5 Bratislava–Železná studienka 6 Devínska Kobyla above telecast 7 Fúgelka 8 Lozorno–Rusniaky 9 Naháč–Katarínka 10 Pohanská NPR 11 Stupava–Lintavy 12 Vinosady–environment 13 Zbojníčka

DFSa 7768 7768 7868 7868 7868 7867 7669 7668 7471 7569 7768 7669 7768

Tree species b Cb,Qr Fs Ca Ca,Cb,Fe,Pe, Ps,Qp Ps Qpu Aps,Qd Ag,Cb Ca Ac,Fe Bp,Cb,Fs,Qd Ac Cb

No. of samples 2 1 1 7 1 2 2 3 1 2 4 1 1

Notes: a - square of the Databank of Slovak fauna; b - abbreviations of tree species see chapter Material and methods

Results and discussion Generally 58 ciliate species (Protozoa: Ciliophora) being classified into 7 classes, 9.subclasses, 16 orders and 27 families were identified in 28 samples of decaying wood mass of 14 tree species. The highest species richness (33) (in sensu Spellerberg, Fedor, 2003) was recorded in the samples of Carpinus betulus (Table 2). All the ciliate species have been observed worldwide from various terrestrial habitats for instance in soil, litter, dry mosses and sand (e. g. Foissner, 1987a, b; Foissner et al., 2002; Tirjaková, Matis, 1987 etc.). Nine species of the total number of ciliates were recorded in Slovakia for the first time: Anictostoma terricola, Arcuospathidium japonicum, Bakuella pampinaria, Enchelys polynucleata, Ilsiella elegans, Litonotus muscorum, Oxytricha granulifera quadricirrata, Platyophrya macrostoma and Spathidium turgitorum. The first records of ciliates refer to wood mass of 7 tree species in 5 localities. All the first records were confirmed in 1–2 localities and in the samples of 1–3 tree species. Maximum of the records corresponds with the locality Stupava–Lintavy (Tirjaková, Bartošová, 2004). 18 from a total of 58 ciliate species identified occurred in one sample and 7 of them were recorded in two samples. This fact reflects a high proportion of rare species (Fig..1), being also proved by Fig. 2 with 44% of species with frequency ≤ 10%. None of the species occurred in all the samples. Leptopharynx costatus – the most frequent euryoecious species occurred in 82.1% of the samples (Fig. 1, Table 2). It has been classified in the 5th frequency category together with Colpoda steinii (Fig. 2). The 4th and 5th frequency categories have included just 5 species (Colpoda cucullus, C. inflata, C. steinii, Gonostomum affine and Leptopharynx costatus) with their frequency > 45% (Fig..2, Table 2). All these species are common in terrestrial habitats. Colpodids belong to r-strategists and fast-growing species well adapted to utilise very short and wet 39

T a b l e 2. List of recorded ciliate species (Protozoa, Ciliophora) on the studied localities and in trees in years 2001–2004. Ciliate species Locality numbera Tree species b Anictostoma terricola F o i s s n e r, 1993 8, 11 Ag, Qd Apospathidium atypicum (B u i t k a m p et W i l b e r t, 10 Ac, Fe 1974) Arcuospathidium australe F o i s s n e r, 1988 11 Cb Arcuospathidium japonicum F o i s s n e r, 1988 8 Cb Bakuella pampinaria E i g n e r et F o i s s n e r, 1992 11 Fs Blepharisma hyalinum P e r t y, 1849 4, 7, 8, 11, 12 Ac, Ag, Cb, Fe, Fs, Pe, Qd, Qp, Bresslaua vorax K a h l, 1931 3 Ca Colpoda aspera K a h l, 1926 4 Fe Colpoda cucullus (M ü l l e r, 1773) 1, 2, 3, 4, 5, 8, 9, 10, 11 Ac, Ag, Bp, Ca, Cb, Fe, Fs, Pe, Ps, Qd, Qp, Qr Colpoda edaphoni F o i s s n e r, 1980 4, 6, 8, 11 Ag, Cb, Fs, Ps, Qpu Colpoda henneguyi F a b r e-D o m e r g u e, 1889 2, 11 Bp, Fs Colpoda inflata (S t o k e s, 1884) 1, 2, 3, 4, 5, 6, 7, 8, 10, 11, 13 Ac, Ag, Bp, Ca, Cb, Fe, Fs, Ps, Qd, Qpu, Qr Colpoda lucida G r e e f f, 1888 8, 11, 13 Ag, Cb, Fs, Qd Colpoda maupasi E n r i q u e z, 1908 6, 8, 9, 10, 11, 13 Ag, Ca, Cb, Fe, Fs, Qd, Qpu Colpoda steinii M a u p a s, 1883 1, 2, 4, 6, 7, 8, 9, 10, 11, 12, 13 Ac, Ag, Aps, Bp, Ca, Cb, Fe, Fs, Ps, Qd, Qp, Qpu, Qr Cyclidium muscicola K a h l, 1931 2, 4, 5, 7, 8, 10, 11, 12 Ac, Ag, Bp, Ca, Cb, Fs, Pe, Ps, Qd Cyrtohymena muscorum (K a h l, 1932) 11 Fs Cyrtolophosis acuta K a h l, 1926 4, 6, 8, 10, 11 Ac, Ag, Cb, Fe, Qp, Qpu Cyrtolophosis elongata (S c h e w i a k o f f, 1892) 2, 4, 6, 8, 9, 10, 11 Ac, Bp, Ca, Cb, Fe, Fs, Qpu Cyrtolophosis mucicola S t o k e s, 1885 1, 4, 8 Cb, Qp, Qr Dileptus breviproboscis F o i s s n e r, 1981 8 Cb Drepanomonas pauciciliata F o i s s n e r, 1987 4, 11 Bp, Cb, Qp Drepanomonas revoluta P e n a r d, 1922 1, 4, 8,11 Ag, Cb, Fe, Fs, Ps Enchelys polynucleata F o i s s n e r, 1984 10 Ac, Fe Epispathidium amphoriforme (G r e e f f, 1888) 1, 8 Cb, Qr Euplotes muscicola K a h l, 1932 8, 10, 11, 12 Ac, Bp, Cb, Fe, Fs Frontonia angusta K a h l, 1931 11 Bp Frontonia depressa (S t o k e s, 1886) 4, 5, 8, 11, 13 Bp, Cb, Ps, Qp Gonostomum affine (S t e i n, 1859) 1, 2, 4, 5, 8, 10, 11, 13 Ac, Ag, Bp, Cb, Fe, Fs, Pe, Ps, Qd, Qr, Qp Hausmanniella patella (K a h l, 1931) 2 Fs Hemisincirra gellerti (F o i s s n e r, 1982) 4, 11 Cb, Fe, Qd Hemisincirra interrupta (F o i s s n e r, 1982) 4, 13 Cb, Fe

SD c 3.5 3.4

Fd 10.71 7.14

5 4 2,3,4,5 5 4 2,3,4,5 3,4,5 3 1,2,3,4,5 2,3,4 1,2,3,4 1,2,3,4,5

3.57 3.57 3.57 32.14 3.57 3.57 67.86 17.86 7.14 67.86 14.29 32.14 75.00

2,3,4,5 4 1,3,4,5 1,3,4,5 3.4 2,3,5 3,4,5 3.4 4 3.4 3 2,3,5 2,3,4,5 3 3,4,5 2.4

42.86 3.57 25.00 32.14 10.71 3.57 14.29 21.43 7.14 10.71 17.86 3.57 21.43 67.86 3.57 10.71 7.14

T a b l e 2. (Continued) Ciliate species Locality numbera Holosticha tetracirrata B u i t k a m p et W i l b e r t, 1974 10 Homalogastra setosa K a h l, 1926 10 Chilodonella uncinata (E h r e n b e r g, 1838) 11 Chilodontopsis muscorum K a h l, 1931 8, 11 Ilsiella elegans F o i s s n e r, A g a t h a et B e r g e r, 2002 4, 10 Kahlilembus attenuatus (S m i t h, 1897) 8 Leptopharynx costatus M e r m o d, 1914 1, 2, 3, 4, 5, 7, 8, 10, 11, 12, 13 Litonotus muscorum (K a h l, 1931) 7 Odontochlamys gouraudi C e r t e s, 1891 8 Opercularia arboricolum (B i e g e l, 1954) 4, 8, 11 Oxytricha granulifera quadricirrata B l a t t e r e r et 11 F.o.i.s.s.n.e.r, 1988 Oxytricha setigera S t o k e s, 1891 4, 8, 11 Platyophrya macrostoma F o i s s n e r, 1980 8 Platyophrya spumacola K a h l, 1927 2, 4, 7, 8, 11 Platyophrya vorax K a h l, 1926 2, 8, 11 Pseudocohnilembus pusillus (Q u e n n e r s t e d t, 1896) 5, 7 Sathrophilus muscorum (K a h l, 1931) 1, 4, 8, 10, 11, 13 Spathidium bavariense K a h l, 1930 4 Spathidium muscicola K a h l, 1930 2, 4, 8, 11, 13 Spathidium spathula (M ü l l e r, 1773) 4, 8, 11 Spathidium turgitorum F o i s s n e r, A g a t h a et 11 B.e.r.g.e.r, 2002 Sphaerophrya terricola F o i s s n e r, 1986 11 Sterkiella histriomuscorum (F o i s s n e r, B l a t t e r e r, A 4, 8, 11 u g u s t i n et K o h m a n n, 1991) Telotrochidium cylindricum F o i s s n e r, 1978 4, 8, 11 Urosomoida agiliformis F o i s s n e r, 1982 2, 11 Vorticella astyliformis F o i s s n e r, 1981 4, 7, 8, 10, 11

Fd

Tree species b Ac, Fe Ac Fs Cb, Fs Ac, Fe, Qp Cb Ac, Ag, Aps, Bp, Ca, Cb, Fe, Fs, Pe, Ps, Qd, Qp, Qr Qd Cb Ag, Pe, Qd Cb

SD c 3.4 3 4 4 3.4 2,3,4,5 2,3,5 5

3.57 3.57 14.29 3.57

Ag, Fe, Fs, Qd Ag Ag, Aps, Bp, Cb, Fs, Ps, Qd Ag, Fs, Qd Aps, Ps, Qd, Ac, Ag, Ca, Cb, Fe, Fs, Ps, Qd, Qp Fe, Qp Ag, Cb, Fs, Qd, Qp Cb, Fs, Ps Fs

3.4 3 3,4,5 3.5 2,3,4,5 2.4 2,3,4 4.5 4

14.29 3.57 32.14 10.71 10.71 46.43 7.14 25.00 14.29 3.57

Qd Ag, Bp, Fs, Qd, Qp

3 3.4

3.57 17.86

Ag, Cb, Fs, Pe Fs, Qd Cb, Fe, Fs, Pe, Qd, Qp

2,3,4,5 3.4 2.4

14.29 10.71 21.43

7.14 3.57 3.57 7.14 10.71 3.57 82.14

Notes: a – see Table 1.; b, c – abbreviations of tree species and consideration of stage of wood decay see chapter Material and methods; d – frequention of ciliate occurrence in studied samples (%)

periods. The cell division in cysts is considered as an r-selective life strategy and serves for the fast multiplying in a short time, what enables ciliates their wide distribution and to colonise various terrestrial habitats (Lüftenegger et al., 1985; Foissner et al., 2002).

20 18 16

species number

14 12 10 8 6 4 2 82.1

75

67.9

46.4

42.9

32.1

25

21.4

17.9

14.3

10.7

7.14

3.57

0

frequention (%)

Fig. 1. Frequentions of 58 ciliate species recorded in 28 samples of decaying wood mass.

5%

3%

10%

I. (0–10%) 44%

II. (11–25%) III. (26–45%) IV. (46–70%) V. (71–100%)

38%

Fig. 2. Percentual representation of ciliate species in frequention groups (categorization in groups according to Schwerdtfeger, 1975).

42

Blatterer, Foissner (1988) show 23 species with frequency ≥ 50% in soil samples from Australia. The total of 20 species occurred with the frequency ≥ 50% in Germany (Foissner, 2000). 17 and 13 species were recorded in Austria (Tullnerfeld or alpine habitats) (Foissner et al., 1985; Foissner, 1981a) and 12 species with the same frequency occurred in South America (Foissner, 1997). Totally 11 species with frequency ≥ 48% were found in samples from Namibia (Foissner et al., 2002) and 19 species with their frequency > 55% were noted in Kenya (Foissner, 1999). 15 species belonged to the most frequent in Antarctica (Foissner, 1996). Many samples collected in Antarctica did not include any species, what might refer to very low frequency values. Therefore species with their frequency ≥ 5.1% have been classified into the category of the most frequent species.

Structure of systematic classes The percentual structure of ciliates recorded in the samples from decaying wood is demonstrated on the Fig. 3 and 4 according to their systematic classes. From this point of view we can submit following ascendant rank: Heterotrichea (1.7%), Phylopharyngea (5.2%), Nassophorea (6.9%), Oligohymenophorea (17.2%), Spirotrichea and Litostomatea (19.0%) and Colpodea (31.0%) (the system accords to Lynn, Corliss, 1991, Fig. 3). Being based on the older system by Corliss (Corliss, 1979) (Fig. 4) the structure appears as followed: Polyhymenophora (20.7%), Oligohymenophora (24.1%) and Kinetofragminophora (55.2%). 19.0%

Colpodea 31.0%

Heterotrichea Litostomatea

5.2%

Nassophorea Oligohymenophorea 1.7%

17.2%

Phylopharyngea 6.9%

19.0%

Spirotrichea

Fig. 3. Percentual representation of ciliate species in systematic groups (the system according to Lynn, Corliss, 1991).

Tirjaková (1988), who has been studying soil ciliates in field communities, found out, that Kinetofragminophora and Polyhymenophora (43%) had been dominantly 43

represented. Soil habitats are mainly characteristic by distinct prevalence of Kinetofragminophora, actually proved by Foissner (1981a), Buitkamp (1977), etc. According to Foissner (1985), the high proportion of colpodids is one of the most important characteristics of soil ciliate community in contrast to limnic habitats. This has been proved by many other studies, for instance by Foissner et al. (2002). Oligohymenophora and Polyhymenophora (mainly Spirotrichea) keep the dominant position in limnic habitats e.g. Baláži, Matis (2002) considered Spirotrichea as a dominant class with prevalence of hypotrichs (49%). Oligohymenophora with 57% of peritrichs were the second dominant class, what actually refers to the Foissner’s arguments (1981a, 1987c; Foissner et al., 2002), that peritrichs and sessile forms are predominant in limnic habitats and in soils they are substituted by Kinetofragminophora and Polyhymenophora. 20.7%

Kinetofragminophora

Oligohymenophora 55.2%

Polyhymenophora 24.1%

Fig. 4. Percentual representation of ciliate species in systematic groups (the system according to Corliss, 1979).

Trophic groups The ingestion of 10 various feeding groups were studied to set the food preference of ciliates: bacteria, ciliates, diatoms, zooflagellates, phytoflagellates, hyphae and fungal spores, gymnamoebae, thecamoebae, rotifers and detritus. Ciliates split into 3 trophic categories according to the above-mentioned food sources: bacteriovores, predators and omnivores (Fig. 5). Bacteriovores include species feeding on bacteria. However, if a species feeds on minimally 2 various food sources apart from bacteria, it is classified as an omnivore. Generally, omnivores included ciliate species feeding minimally on 3 various food sources. Predators were defined as the species feeding on ciliates, flagellates, amoebas, rotifers and nematodes. During the evaluation of ciliate food preference we established, that the highest percentage of the recorded ciliate species fed on bacteria (51.7%). The prevalence of 44

bacteriovores in the samples may be explained by sufficient food supply as bacteria considerably participate on wood mass decay. In accordance with couple of authors as well as our results most of ciliate species living in soil are bacteriovores (e. g. Brunberg Nielsen, 1968; Laminger, 1980; Nikolyuk, 1965, 1969). Foissner (1981a) shows as many as 86 resp. 94% bacteriovores in alpine soils. Tirjaková (1997) generally recorded 66% ciliate species feeding on bacteria in agrocoenoses. However, some of the recorded omnivores were classified into bacteriovores as well. Gellért (1956) noted only 27% portion of exclusive bacteriovores (without omnivores) in the humus horizon under mosses. 20.7% 27.6%

Omnivores Bakteriovores Predators

51.7%

Fig. 5. Percentual representation of ciliate species in feeding groups.

The rest portion was formed by omnivores (27.6%) and predators (20.7%) in our samples. Tirjaková (1997) and Gellért (1956) published higher values in representation of predators (33.5% and 37.8%) in field communities or humus. Foissner (1981a) recorded only 7 resp. 2.8% of predators in alpine soils. Predators were represented by a relatively high percentage (about 21%) in decaying wood mass.

Genus–species relationships The 58 identified ciliate species have been classified into 39 genera. Most of them were represented by one species (76.9%, Fig. 6) what may hint at similar food requirements of the same genus species, which are in mutual competition. The results from various authors have been considerably different. Foissner (1981a) recorded 67% genera with one species only in the ciliate community research of alpine soils. The maximum values did not exceed 80% in agrocoenoses studied by Tirjaková (1997). Foissner et al. (2002) noted just 20.2% genera with one species in their samples from Namibian terrestrial habitats. 45

80

76.92

70

% of genera

60 50 40 30 20

12.82 5.13

10

2.56

2.56

4

8

0 1

2

3 species number

Fig. 6. Genus–species relationships of ciliates.

In the samples five genera were represented by two species (12.8%), two genera by three (5.1%), one genus by four (2.6%) and one (Colpoda) by eight species (2.6%). The prevalence of colpodids in the samples corresponds with the general statement, that the genus Colpoda is commonly represented in soils world-wide, for instance Foissner (1981a) recorded six colpodid species in alpine soils, Tirjaková (1988) recorded 8.species in field communities and Foissner et al. (2002) 17 colpodids from the genus Colpoda.

The similarity of ciliate communities Communities of ciliates were mutually compared using hierarchical classification, based on identity of species representation (Sørensen’s index of dissimilarity, complete linkage method). Totally 58 ciliate species recorded in the samples from 14 tree species were included in the analysis. Hierarchical classification (Fig..7) has differentiated two large groups of tree communities, which split up into several smaller groups. The first group of the communities has divided into two smaller clusters. The first most diversified cluster consists of the following tree species: Acer campestre, Fraxinus excelsior, Quercus petraea, Alnus glutinosa, Quercus dalechampii, Fagus sylvatica and Carpinus betulus. It includes 56 ciliate species, actually 96.6% of the total number of species recorded in the decaying wood mass samples. The second cluster is formed by four tree species (Betula pendula, Pinus sylvestris, Quercus robur and Picea excelsa) and is characterised by presence of 25 ciliate species. Two of them appear with a low frequency (Colpoda henneguyi and Frontonia angusta).

46

Fig. 7. Hierarchical classification of ciliate communities at tree species (Sörensen’s index, complete linkage) (vertical axis – dissimilarity, horizontal axis – tree species: 1– Acer campestre, 2 – Fraxinus excelsior, 3 – Quercus petraea, 4 – Alnus glutinosa, 5 – Quercus dalechampii, 6 – Fagus sylvatica, 7 – Carpinus betulus, 8 – Betula pendula, 9 – Pinus sylvestris, 10 – Quercus robur, 11 – Picea excelsa, 12 – Acer pseudoplatanus, 13 – Cerasus avium, 14 – Quercus pubescens).

The second group of the communities has split up into the other two clusters. The first one determined by Acer pseudoplatanus has been established at a relatively high value of dissimilarity. Only 4 ciliate species were recorded in the relevant samples. They may be considered as common terrestrial ciliates (Colpoda steinii, Leptopharynx costatus, Platyophrya spumacola and Pseudocohnilembus pusillus). The second cluster joins to the first one at the lower value of dissimilarity. It includes 11 ciliate species being recorded in the samples of Cerasus avium and Quercus pubescens. The rare species Bresslaua vorax was confirmed in the sample of Cerasus avium. Although splitting the communities on the basis of a tree species structure is relatively distinct, we do not have to overrate its effect. Confrontation with other papers (Bartošová et al., 2005) hints at the contradiction. From this point of view a tree species structure is not supposed to play a significant role in forming ciliate communities. Any

47

special relation or differences in species diversity have not been proved even between deciduous and coniferous trees. Translated by authors Acknowledgement The research was supported by the grant No. 1/0119/03.

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Foissner, W., 1993a: Colpodea (Ciliophora). Protozoenfauna Vol. 4/1. Gustav Fischer Verlag, Stuttgart–Jena–New York, 798 pp. Foissner, W., 1993b: Corticocolpoda kaneshiroae n. g., n. sp., a new colpodid ciliate (Protozoa, Ciliophora) from the bark of Ohia trees in Hawaii. J. Euk. Microbiol., 40, p. 764–775. Foissner, W., 1994: Pentahymena corticicola nov. gen., nov. spec., a new colpodid ciliate (Protozoa, Ciliophora) from bark of Acacia trees in Costa Rica. Arch. Protistenk., 144, p. 289–295. Foissner, W., 1996: Faunistics, taxonomy and ecology of moss and soil ciliates (Protozoa, Ciliophora) from Antarctica, with description of new species, including Pleuroplitoides smithi gen. n., sp. n.. Acta Protozool., 35, p. 95–123. Foissner, W., 1997: Soil ciliates (Protozoa: Ciliophora) from evergreen rain forests of Australia, South America and Costa Rica: diversity and description of new species. Biol. Fertil. Soils, 25, p. 317–339. Foissner. W., 1999: Notes on the soil ciliate biota (Protozoa, Ciliophora) from the Shimba Hills in Kenya (Africa): diversity and description of three new genera and ten new species. Biodiv. Conserv., 8, p..319–389. Foissner, W., 2000: A compilation of soil and moss ciliates (Protozoa, Ciliophora) from Germany, with new records and descriptions of new and insufficiently known species. Europ. J. Protistol., 36, p. 253–283. Foissner, W., Agatha, S., Berger, H., 2002: Soil ciliates (Protozoa, Ciliophora) from Namibia (Southwest Africa), with emphasis on two contrasting environments, the Etosha Region and the Namib Desert. Part I: Text and line drawings. Denisia, 5, p. 1–1063. Gellért, J., 1956: Ciliaten des sich unter dem Moosrasen auf Felsen gebildeten Humus. Acta. Biol. Acad. Sci. Hung., 6, p. 337–359. Holecová, M., Zach, P., Kardošová, J., 2002: Epigaeic weevils (Coleoptera, Curculionidea) of oak-hornbeam forest in a vicinity of Bratislava (SW Slovakia) (in Slovak). Folia Faunistica Slovaca, 7, p. 39–48. Holecová, M., Lukáš, J., Haviar, M., Harakaľová, E., 2005: Ants (Hymenoptera, Formicidae) as an important part of the epigeic fauna in Carpathian oak-hornbeam forests. In Tajovský, K., Schlaghamerský, J., Pižl, V. (eds): Contributions to Soil Zoology in Central Europe I. ISB AS CR, České Budějovice, p. 31–35. Laminger, H., 1980: Bodenprotozoologie. Microbios, 1, p. 1–142. Lüftenegger, G., Foissner, W., Adam, H., 1985: R- and K-selection in soil ciliates: a field and experimental approach. Oecologia, 66, p. 574–579. Lynn, D. H., Corliss, J.O., 1991: Phyllum Ciliophora. In Harrison, F.W., Corliss, J.O. (eds): Microscopic Anatomy of Invertebrates, New York, p. 333–467. Miteva, D., 1992: A study of Protozoa in communities of Quercus frainetto Ten. and Q. cerris L. in the northern part of the Strandja Mountain. Godišnik na Sofijiskija Universita „Kliment-Ochridski“ Biologičeski Fakulti, 81, p. 127–132. Mrva, M., Matis, D., 2000: Rhizopoda in Leaf-litter of Some Localities of Oak-hornbeam Forest in Malé Karpaty Mts (Western Slovakia) (in Slovak). Folia Faunistica Slovaca, 5, p. 1–9. Mrva, M., 2003: Diversity of active gymnamoebae (Rhizopoda, Gymnamoebia) in dendrotelmae of oak-hornbeam forests in Malé Karpaty Mts (Western Slovakia). Protistology, 3, p. 121–125. Nikolyuk, V.F., 1965: Antagonistic interrelationship between some soil protista and certain phyto-pathogenous fungi affecting cotton plants. Progress in Protozool. (Proc. 2nd Int. Congr. Protozool.), 2, p. 118–119. Nikolyuk, V.F., 1969: Some aspects of the study of soil protozoa. Acta Protozool., 7, p. 99–109. Podani, J., 1993: Syn–tax. Version 5.0. Computer programs for Multivariate Data Analysis in Ecology and Systematics. User’s guide. Scientia Publishing. Budapest, 104 pp. Schwerdtfeger, F., 1975: Ökologie der Tiere. Band III - Synökologie. Verlag Paul Parey, Hamburg–Berlin, 451.pp. Speight, M.C.D., 1989: Saproxylic invertebrates and their conservation. Council of Europe: Publications and Documents Division, Strasbourg, 81 pp. Spellerberg I.F., Fedor, P.J., 2003: A tribute to Claude Shannon (1916–2001) and a plea for more rigorous use of species richness, species diversity and the Shannon-Wiener’s index. Global Ecology and Biogeography, 12, p. 177–179. Stout, J.D. 1973: The relationship between protozoan populations and biological activity in soils. Am. Zool., 13, p. 193–201. Tirjaková, E., Matis, D., 1987: Ciliated protozoa (Ciliophora) from submerged, wet, moist and dry mosses in selected localities of Slovenský raj. Acta Fac. Rerum Nat. Univ. Comenianae Zool., 29, p. 1–16.

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Tirjaková, E., 1988: Structures and dynamics of communities of ciliated protozoa (Ciliophora) in field communities. 1. Species composition, group dominance, communities. Biológia, Bratislava, 43, p..479–503. Tirjaková, E., 1997: Structure and dynamics of communities of ciliated Protozoa (Ciliophora) in field communities. Succession, genus-species relationships, nutrition. Ekológia (Bratislava), 16, p. 243–252. Tirjaková, E., Mrva, M., Hlúbiková, D., 2002: Ciliophora and Rhizopoda in soil, leaf-litter and mosses of oak-hornbeam forests in the Malé Karpaty Mts (Western Slovakia). In Tajovský, K., Balík, J, Pižl, V. (eds): Studies on Soil Fauna in Central Europe, ISB AS CR, České Budějovice, p. 233–240. Tirjaková, E., Bartošová, P., 2004: The first record of ciliated protozoa (Protozoa, Ciliophora) from Slovakia in the decaying wood mass (in Slovak). Folia Faunistica Slovaca, 9, p. 11–20. Wallwork, J.A., 1976: The distribution and diversity of soil fauna. Academic Press, London, 355 pp. Wenzel, F., 1953: Die Ciliaten der Moosrasen trockner Standorte. Arch. Protistenk., 99, p. 70–141. Received 30. 7. 2005

Bartošová P., Tirjaková E.: Vybrané ekologické charakteristiky spoločenstiev nálevníkov (Protozoa, Ciliophora) v odumretej drevnej hmote na území Malých Karpát. V rokoch 2001–2004 sme v rámci výskumu spoločenstiev nálevníkov v odumretej drevnej hmote odobrali 28 vzoriek drevnej hmoty 14 druhov drevín z územia Malých Karpát. Vo vzorkách sme zaznamenali 58 druhov nálevníkov, z ktorých 9 sme zistili na území Slovenska prvýkrát. Okrem druhového spektra sme študovali zastúpenie systematických tried, potravné, rodovo–druhové vzťahy a formovanie spoločenstiev v závislosti od druhu dreviny. Zo systematických tried prevažovali zástupcovia triedy Colpodea a z potravných skupín bakteriovory. Maximálny počet druhov (8) sme zaznamenali v.rode Colpoda. Hierarchická klasifikácia podľa identity druhového zastúpenia nálevníkov na jednotlivých druhoch drevín vyčlenila dve výrazné spoločenstvá. Napriek tomu druh dreviny pravdepodobne nezohráva významnú úlohu pri formovaní spoločenstiev nálevníkov. Nezistili sme dokonca ani osobitnú väzbu resp. odlišnosť druhového spektra medzi listnatými a ihličnatými drevinami.

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Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 51–58

DIVERSITY OF ACTIVE GYMNAMOEBAE (Rhizopoda, Gymnamoebia) IN MOSSES OF THE MALÉ KARPATY MTS (SLOVAKIA) MARTIN MRVA Department of Zoology, Comenius University, Mlynská dolina B-1, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected] Abstract Mrva M.: Diversity of active gymnamoebae (Rhizopoda, Gymnamoebia) in mosses of the Malé Karpaty Mts (Slovakia). Ekológia (Bratislava), Vol. 24, Supplement 2, p. 51–58. In the period of 2000–2002 the fauna of active naked amoebae (Rhizopoda, Gymnamoebia) was studied in mosses at five sites in oak-hornbeam forests of the Malé Karpaty Mts (Slovakia). The dry sample material was moistened by distilled water and after 5 days of incubation the amoebae were identified by direct examination. Identification of amoebae was performed on the base of morphological characters of the active stages. Relatively high diversity of 32 taxa of naked amoebae was recorded. The diversity at the sites varied from 17 to 23 taxa. The highest richness appeared in the family Thecamoebaidae (9 species), however some other families – Hartmannellidae, Vannellidae and Paramoebidae were significantly represented as well. The observed community of species of naked amoebae in mosses indicate considerable similarity to freshwater communities. Key words: Gymnamoebia, moss, diversity, Slovakia, oak-hornbeam forests

Introduction Terrestrial habitats are recognised as specific freshwater ecosystems because active stages of protists always depend on presence of water (e.g. Bamforth, 1980; Finlay et al., 2000). Generally, higher moisture enhances the species diversity (Bamforth, 1973). Modern studies of diversity of naked amoebae were focused mainly on water habitats. Some of them refer to freshwater (Smirnov, Goodkov, 1996) or sea (Butler, Rogerson, 2000). In terrestrial habitats quantity of amoebae has been analysed (Singh, 1946; Bischoff, Anderson, 1998; Anderson, 2000), however their systematic diversity in these habitats remains practically unknown. Recently only Brown, Smirnov (2004) have brought several results from a study on diversity of Gymnamoebia in soil. The differences between the fauna in freshwater and in terrestrial habitats are well known for ciliates, which have formed specific morphological and physiological 51

adaptations (Fenchel, 1987; Foissner, 1987; Cowling, 1994). Similarly, the fauna of testate amoebae is different in freshwater and soil habitats (Foissner, 1987). However, such information is unavailable for naked amoebae though some of the species were found both in freshwater and soil (Page, 1991). Until now, mosses as habitats of the naked amoebae were studied scarcely or marginally. Some information can be found in the works of Bartoš (1940, 1947, 1949, 1963), Ertl (1955), Matis et al. (1997), Matis, Mrva (1998) and Page (1991). Practically there are no modern works directed to naked amoebae from these habitats and the diversity of naked amoebae in mosses is almost unknown. For several years the Malé Karpaty Mts have been an object of research on both micro- and macrofauna of various habitats, dealing with Protozoa (Tirjaková et al., 2002; Mrva, 2003a), Tardigrada (Degma et al., 2005) and Arthropoda (Holecová, Sukupová, 2000; Majzlan et al., 2000; Štepanovičová, Országh, 2002; Krumpálová, Szabová, 2003; Holecová et al., 2005). This article brings new information on diversity of the naked amoebae in the habitat of mosses from the Malé Karpaty Mts in Western Slovakia.

Material and methods Samples of mosses growing on soil were collected monthly in the period of 2000–2002 at five sites in oak-hornbeam forests of the Malé Karpaty Mts (Western Slovakia): Fúgelka, Naháč–Katarínka 1, Naháč–Katarínka 2, Lindava, Lošonec-lom quarry. For detailed ecological characteristics and situation of sites see Zlinská et al. (2005). The sampled material was analysed according to modification of method used for ciliates (e.g. Foissner, 1987; Matis, Tirjaková, 1994; Aescht, Foissner, 1995): dry sampled material of moss was flooded with distilled water and incubated 5 days on undirected light and laboratory temperature. The amoebae were directly examined in the suspension pipetted from the flooded sample. Observations were made using the Nikon Labophot microscope with phase contrast equipment. Identification of amoebae was performed on the base of morphological criteria according to Page (1988, 1991) with the help of other works, which included detailed descriptions of the species (Page, 1969a, b, 1977, 1983; Sawyer, 1975; Smirnov, Goodkov, 1994; Smirnov, 1995, 1999; Michel, Smirnov, 1999).

Results Naked amoebae occurred in all the 78 samples examined. Totally 32 taxa of 3 orders, 8 families and 16 genera of naked amoebae were recorded: 23 species, 8 taxa being identified into the genus level only and 1 unidentified leptomyxid amoeba (Table 1). Unidentified heterolobosean amoebae were found at all the sites. The diversity at the study sites varied from 17 to 23 taxa. The highest diversity appeared in the family Thecamoebaidae (9 species). Of the species observed, 7 were recorded at all the sites: Dermamoeba minor, Thecamoeba quadrilineata, Platyamoeba stenopodia, Korotnevella stella, Flamella sp. 1, Acanthamoeba sp. 1, Acanthamoeba sp..2. 52

T a b l e 1. Naked amoebae recorded from localities. Taxon

F

Locality K1 K2 Lin LQ

EUAMOEBIDA Amoebidae Deuteramoeba algonquinensis (B a l d o c k, R o g e r s o n & B e r g e r, 1983) P.a.g.e, 1987 Hartmannellidae Hartmannella cantabrigiensis P a g e, 1974 Hartmannella vermiformis P a g e, 1967 Saccamoeba limax (D u j a r d i n, 1841) P a g e, 1974 Saccamoeba stagnicola P a g e, 1974 Thecamoebidae Dermamoeba granifera (G r e e f f, 1866) P a g e & B l a k e y, 1979 Dermamoeba minor (P u s s a r d, A l a b o u v e t t e & P o n s, 1979) P a g e, 1988 Paradermamoeba levis S m i r n o v & G o o d k o v, 1994 Paradermamoeba valamo S m i r n o v & G o o d k o v, 1993 Sappinia diploidea (H a r t m a n n & N ä g l e r, 1908) A l e x e i e f f, 1912 Thecamoeba quadrilineata (C a r t e r, 1856) L e p ş i, 1960 Thecamoeba sphaeronucleolus (G r e e f f, 1891) S c h a e f f e r, 1926 Thecamoeba striata (P e n a r d, 1890) S c h a e f f e r, 1926 Thecamoeba terricola (G r e e f f, 1866) L e p ş i, 1960 Vannellidae Platyamoeba stenopodia P a g e, 1969 Vannella sp. Vannella lata P a g e, 1988 Vannella platypodia (G l ä s e r, 1912) P a g e, 1976 Paramoebidae Korotnevella bulla (S c h a e f f e r, 1926) G o o d k o v, 1988 Korotnevella diskophora S m i r n o v, 1999 Korotnevella stella (S c h a e f f e r, 1926) G o o d k o v, 1988 Mayorella penardi P a g e, 1972 Mayorella vespertilioides P a g e, 1983 LEPTOMYXIDA Flabellulidae Flamella sp. 1 Flamella sp. 2 Leptomyxidae Leptomyxa reticulata G o o d e y, 1914 Rhizamoeba sp. Unidentified leptomyxid amoeba ACANTHOPODIDA Acanthamoebidae Acanthamoeba sp.1 Acanthamoeba sp.2 Acanthamoeba sp.3 INCERTAE SEDIS Stygamoeba sp. Total

+

+

+

+

+ + +

+ + + + +

+

+ +

+

+

+ + +

+ + +

+ +

+

+ +

+ +

+ +

+

+

+

+

+ +

+

+ +

+ + +

+

+

+

+ + +

+ + +

+

+

+

+

+

+ +

+

+ +

+ + +

+

+ + + + + +

+ + + +

+

+ +

+ + +

+ 17

+ 18

+ +

+ +

+ +

17

+ 17

23

Notes: F – Fúgelka, K1 – Naháč–Katarínka 1, K2 – Naháč–Katarínka 2, Lin – Lindava, LQ – Lošonec–lom quarry

53

Discussion Since amoebae were examined only by light microscopy, the determination depended largely on the morphological criteria according to Page (1988, 1991). The results were consulted with other works containing descriptions and illustrations of trofozoits of naked amoebae published before but also after Page’s monographs (Page, 1969a, b, 1977, 1983; Sawyer, 1975; Smirnov, Goodkov, 1994; Smirnov, 1995, 1999; Michel, Smirnov, 1999). The emphasis was given to detailed observations of locomotive and floating forms. The shape of the body and its dimensions, the type, number and dimensions of nuclei and the presence of crystals were key characters for identification. Some of the recorded species are difficult to be identified. Saccamoeba limax was differentiated from S. stagnicola by absence of crystalic inclusions (Page, 1991). Dermamoeba granifera possess a zone of fine granules in anterior hyaloplasma (Page, 1977). This zone lacks in D. minor, which is a smaller species with more rounded anterior end than the previous one. Paradermamoeba valamo differs from P. levis by shape, larger size and floating form. Amoebae of the genus Korotnevella were typical with finger-like dactylopodia and by lack of the double crystalic inclusions that were characteristic for amoebae of the genus Mayorella. Of the genus Korotnevella, K. bulla was the largest species with floating form often with several very long (more than 4 times of diameter of central mass) thin subpseudopodia. K. stella formed typical floating form with about 6–8 bent pseudopodia and K. diskophora, the smallest species, formed floating form similar to Vannella amoebae with up to 4 straight thin pseudopodia. Mayorella penardi was typical with frequent absence of any subpseudopodia in its locomotive form. The similar species, M. vespertilioides, lacked all the subpseudopodia only rarely (Page, 1991). If we compare the obtained results with literary data, the observed diversity of 32 taxa is relatively high. From the older literature, Bartoš (1940) detected 6 species of naked amoebae in mosses from Karpaty Mts (Slovakia), in later works he reported 2 species from mosses near Prague (Czech Republic) (Bartoš, 1947), 2 species in Šumava mosses (Czech Rrepublic) (Bartoš, 1949) and 2 species of gymnamoebae in moss from China (Bartoš, 1963). Fantham, Porter (1945) found 12 species of naked amoebae in mosses in Canada. Unfortunately many of their species can not be identified at present and some of them have invalid names. Ertl (1955) noted 2 species in moss of the peat -bog Bór (Slovakia). From mosses of the Slovenský raj Mts (Slovakia) 4 species of the family Thecamoebidae have been known (Matis et al., 1997). Matis, Mrva (1998) found 6 species of ameboid protists in mosses in Bratislava (Slovakia). Finally, we should note that the low number of species in the cited works is a result of focusing mainly on testate amoebae or of few samples investigated. Closer number of species was reported by Brown, Smirnov (2004) who revealed 48 species in grassland soil. Interestingly, in the present study the richness of 32 species is distinctly higher or similar to the data on diversity of naked amoebae in freshwater habitats. In lakes, 15 species were found by O’Dell (1979), 29 by Smirnov, Goodkov (1996), 14 by Matis et al. (1997), 23 by Butler (1999a), 16 by Butler (1999b), 13 by Butler et al. (2000), 20 and 54

30 by Smirnov (2003). In rivers, 17 species were identified by Ertl (1984) and 14 by Mrva (2003b). Please note that cited works are established on enrichment cultivation with exception of Mrva (2003b) who used the direct examination of the samples. The opinion that the enrichment cultivation is the only method for investigating the species diversity, is widespread among researchers working with amoebae (e.g. Butler, Rogerson, 2000; Smirnov, 2003; Smirnov, Brown, 2004). Despite of this, the study revealed relatively high diversity of active gymnamoebae by long-term direct and detailed examination of the samples. Recently, treeholes were examined by this method with 19 species of Gymnamoebia recovered (Mrva, 2003a) and similarly a river with 14 recorded species (Mrva, 2003b). However, the present study probably did not exhaust the whole community of species of gymnamoebae. The distribution of gymnamoebae in habitats is heterogeneous (Smirnov, 2003) and many amoebae are adhered to substrate particles, so some of them could be omitted. Until now we do not know how the fauna of naked amoebae differs in freshwater and terrestrial habitats though there are indications that the fauna is similar as many freshwater species were isolated from soil, moss or leaf litter (Page, 1991; Brown, Smirnov, 2004; Smirnov, Brown, 2004). The exception includes probably only large species of the family Amoebidae that were not reported from terrestrial samples (Smirnov, Brown, 2004). From this family only Deuteramoeba algonquinensis, known as „typical soil“ species, was recorded in the paper. Members of the family Thecamoebidae seem to be the dominant group of Gymnamoebia in mosses, in this study 9 species of this family were revealed. Though it is known that amoebae of the genus Thecamoeba are frequent in soil and mosses (Page, 1977), surprisingly Brown, Smirnov (2004) in their study of grassland soil did not find any members of this genus. They suppose mainly the influence of enrichment methods. Their next assumption, that the amoebae were destroyed by drying the soil samples, is less probable. In the present study the moss samples were air-dried and all the recorded species had surely formed cysts or pseudocysts. Further, species of the families Hartmannellidae, Vannellidae and Paramoebidae that are well known from freshwater (Ertl, 1984; Smirnov, Goodkov, 1996; Matis et al., 1997; Butler, 1999a, b; Butler et al., 2000; Smirnov, 2003), were numerously represented (Table 1) but at present it is impossible to consider gymnamoebae as typical for habitat of mosses because the knowledge on the occurrence of gymnamoebae in various habitats is not sufficient. Translated by the author Acknowledgement The research was supported by the grant No. 1/0119/03.

55

References Aescht, E., Foissner, W., 1995: Microfauna. In Schinner, F., Öhlinger, R., Kandeler, E., Margesin, R. (eds): Methods in soil biology. Springer, Berlin-Heidelberg-New York, p. 316–337. Anderson, O.R., 2000: Abundance of terrestrial gymnamoebae at a northeastern U. S. site: A four year study, including the El Nino winter. J. Eukaryot. Microbiol., 47, p. 148–155. Bamforth, S.S., 1973: Population dynamics of soil and vegetation Protozoa. Amer. Zool., 13, p. 171–176. Bamforth, S.S., 1980: Terrestrial Protozoa. J. Protozool., 27, p. 33–36. Bartoš, E., 1940: Studien über die moosbewohnenden Rhizopoden der Karpaten. Arch. Protistenkd., 94, p..93–160. Bartoš, E., 1947: The moss-dwelling Rhizopoda of the neighbourhood of Prague (in Czech). Sb. Kl. Přír. v Brně, 28, p. 1–10. Bartoš, E., 1949: Microscopical fauna of the Šumava-mosses. – I. Mosses from neighbourhood of Lac-Plešné (in Czech). Acta Soc. Zool. Bohemoslov., 13, p. 10–29. Bartoš, E., 1963: Die Rhizopoden einiger Moosproben aus China. Acta Soc. Zool. Bohemoslov., 27, p. 85–96. Bischoff, P J., Anderson, O.R., 1998: Abundance and diversity of gymnamoebae at varying soil sites in northeastern U.S.A. Acta Protozool., 37, p. 17–21. Brown, S., Smirnov, A.V., 2004: Diversity of gymnamoebae in grassland soil in southern Scotland. Protistology, 3, p. 191–195. Butler, H.G., 1999a: Seasonal dynamics of the planktonic microbial community in a maritime Antarctic lake undergoing eutrophication. J. Plankton Res., 21, p. 2393–2419. Butler, H.G., 1999b: Temporal plankton dynamics in a maritime Antarctic lake. Arch. Hydrobiol., 146, p..311–339. Butler, H.G., Edworthy, M.G., Ellis-Evans, J.C., 2000: Temporal plankton dynamics in an oligotrophic maritime Antarctic lake. Freshwater. Biol., 43, p. 215–230. Butler, H.G., Rogerson, A., 2000: Naked amoebae from benthic sediments in the Clyde Sea Area, Scotland. Ophelia, 53, p. 37–54. Cowling, A.J., 1994: Protozoan distribution and adaptation. In Darbyshire, J. F. (ed.): Soil Protozoa. CAB International, Wallingford. p. 5–42. Degma, P., Šimurka, M., Gulánová, S., 2005: Community structure and ecological macrodistribution of moss-dwelling water bears (Tardigrada) in Central European oak-hornbeam forests (SW Slovakia). Ekológia (Bratislava), 24, Suppl. 2, 59–75. Ertl, M., 1955: Rhizopoda of the peat-bog Bór (Orava) (in Slovak). Práce II. sekcie SAV, 1, 12, p. 1–39. Ertl, M., 1984: Die Besiedlung verschiedener Typen des Grundes und des hyporheischen Interstitials des Flusses Belá durch Sarcodina. Práce Lab. Rybár. Hydrobiol., 4, p. 63–98. Fantham, H.B., Porter, A., 1945: The microfauna, especially the Protozoa, found in some Canadian Mosses. Proc. Zool. Soc., 115, p. 97–174. Fenchel, T., 1987: Ecology of Protozoa. Science Tech Publishers, Madison. 197 pp. Finlay, B.J., Black, H.I.J., Brown, S., Clarke, K.J., Esteban, G.F., Hindle, R.M., Olmo, J.L., Rollett, A., Vickerman, K., 2000: Estimating the growth potential of the soil protozoan community. Protist, 151, p..69–80. Foissner, W., 1987: Soil Protozoa: fundamental problems, ecological significance, adaptations in ciliates and testaceans, bioindicators and guide to the literature. Prog. Protistol., 2, p. 69–212. Holecová, M., Sukupová, J., 2000: Weevils (Coleoptera, Curculionoidea: Attelabidae, Apionidae, Curculionidae) of the Nature reserve Lošonecký háj (SW Slovakia) (in Slovak). Folia faunistica Slovaca, 5, p. 123–134. Holecová, M., Lukáš, J., Haviar, M., Harakaľová, E., 2005: Ants (Hymenoptera, Formicidae) as an important part of the epigeic fauna in Carpathian oak-hornbeam forests. In Tajovský, K., Schlaghamerský, J., Pižl, V. (eds): Contribution to Soil Zoology in Central Europe I. ISB AS CR, České Budějovice. p. 31–35. Krumpálová, Z., Szabová, S., 2003: Epigeic araneocoenoses of oak-hornbeam forest in the Nature Reserve Katarínka – Malé Karpaty Mts (in Slovak). Entomofauna Carpathica, 15, p. 49–55. Majzlan, O., Gajdoš, P., Fedor, P., 2000: The chosen arthropod groups (Arthropoda) in the suggested Vrchná Hora Reserve near town of Stupava (Protected landscape area CHKO Malé Karpaty) (in Slovak). Acta Fac. Paed. Univ. Tyrnaviensis, 4, p. 3–14.

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Matis, D., Tirjaková, E., 1994: Contribution to the knowledge of Ciliophora of the river Morava. Ekológia (Bratislava), Suppl. 1, p. 55–61. Matis, D., Tirjak, J., Tirjaková, E., 1997: Synopsis of amoebae of Slovakia (in Slovak). Folia Faunistica Slovaca, 2, p. 1–6. Matis, D., Mrva, M., 1998: New amoebae for fauna of Slovakia (in Slovak). Folia Faunistica. Slovaca, 3, p..1–8. Michel, R., Smirnov, A.V., 1999: The genus Flamella Schaeffer, 1926 (Lobosea, Gymnamoebia), with description of two new species. Europ. J. Protistol., 35, p. 403–410. Mrva, M., 2003a: Diversity of active gymnamoebae (Rhizopoda, Gymnamoebia) in dendrotelmae of oak-hornbeam forests in Malé Karpaty Mts. (Western Slovakia). Protistology, 3, p. 121–125. Mrva, M., 2003b: Notes on active gymnamoebae (Rhizopoda, Gymnamoebia) in Turiec river. Folia Faunistica Slovaca, 8, p. 23–26. O’Dell, W D., 1979: Isolation, enumeration and identification of amoebae from a Nebraska Lake. J. Protozool., 26, p. 265–269. Page, F C., 1969a: Hartmannella limax: the original limax amoeba? Trans. Am. Microsc. Soc., 88, p..199–204. Page, F.C., 1969b: Platyamoeba stenopodia n. g., n. sp., a freshwater amoeba. J. Protozool., 16, p. 437–441. Page, F C., 1977: The genus Thecamoeba (Protozoa, Gymnamoebia). Species distinctions, locomotive morphology, and protozoan prey. J. Nnat. Hist., 11, p. 25–63. Page, F.C., 1983: Three freshwater species of Mayorella (Amoebida) with a cuticle. Arch. Protistenkd., 126, p. 201–221. Page, F C., 1988: A new key to freshwater and soil Gymnamoebae. Freshwater Biological Association, Ambleside. 122 pp. Page, F.C., 1991: Nackte Rhizopoda. In Page, F. C., Siemensma, F. J.: Nackte Rhizopoda und Heliozoea. G. Fischer Verlag, Stuttgart-New York. p. 1–170. Sawyer, T.K., 1975: Marine amoebae from surface waters of Chincoteague Bay, Virginia: two new genera and nine new species within the families Mayorellidae, Flabellulidae and Stereomyxidae. Trans. Am. Microsc. Soc., 94, p. 71–92. Singh, B.N., 1946: A method of estimating the numbers of soil Protozoa, especially amoebae, based on their differential feeding on bacteria. Ann. Aappl. Biol., 33, p. 112–119. Smirnov, A.V., Goodkov, A.V., 1994: Freshwater gymnamoebae with a new type of surface structure Paradermamoeba valamo and Paradermamoeba levis sp. n. (Thecamoebidae), and notes on the diagnosis of the family. Acta Protozool., 33, p. 109–115. Smirnov, A V., 1995: Stygamoeba regulata n. sp. (Rhizopoda) – A marine amoeba with an unusual combination of light-microscopical and ultrastructural features. Arch. Protistenkd., 146, p. 299–307. Smirnov, A.V., Goodkov, A.V., 1996: Systematic diversity of gymnamoebae in the bottom sediments of a freshwater lake in Karelia (Lobosea, Gymnamoebia). Zoosyst. Rossica, 4, p. 201–203. Smirnov, A. V., 1999: Korotnevella diskophora n. sp. (Gymnamoebia, Paramoebidae) – small freshwater amoeba with peculiar scales. Protistology, 1, p. 30–33. Smirnov, A.V., 2003: Optimizing methods of the recovery of gymnamoebae from environmental samples: a test of ten popular enrichment media, with some observations on the development of cultures. Protistology, 3, p. 47–57. Smirnov, A.V., Brown, S., 2004: Guide to the methods of study and identification of soil gymnamoebae. Protistology, 3, p. 148–190. Štepanovičová, O., Országh, I., 2002: To the occurence of bug Myrmedobia exilis (Heteroptera, Microphysidae) in Slovakia (in Slovak). Folia Faunistica Slovaca, 7, p. 35–38. Tirjaková, E., Mrva, M., Hlúbiková, D., 2002: Ciliophora and Rhizopoda in soil, leaf-litter and mosses of oak-hornbeam forests in the Malé Karpaty Mts (Western Slovakia). In Tajovský, K., Balík, V., Pižl, V. (eds): Studies on soil fauna in Central Europe. ISB AS CR, České Budějovice. p. 233–240. Zlinská, J., Šomšák, L., Holecová, M., 2005: Ecological characteristics of studied forest communities of an oak-hornbeam tier in SW Slovakia. Ekológia (Bratislava), 24, Suppl. 2, p. 3–19. Received 30. 7. 2005

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Mrva M.: Diverzita nahých meňaviek (Rhizopoda, Gymnamoebia) v machoch Malých Karpát (Slovensko). Počas rokov 2000–2002 som študoval faunu nahých meňaviek (Rhizopoda, Gymnamoebia) machov na piatich lokalitách dubovo-hrabových lesov Malých Karpát. Vysušený materiál som navlhčil destilovanou vodou a po piatich dňoch kultivácie som sledoval zastúpenie meňaviek. Determinácia prebehla na základe morfologických kritérií aktívnych štádií. Zaznamenal som pomerne vysokú celkovú diverzitu 32 taxónov nahých meňaviek, ktorá sa pohybovala na jednotlivých lokalitách od 17 po 23 taxónov. Najviac bola druhovo zastúpená čeľaď Thecamoebidae (9 druhov), pomerne vysoké počty druhov boli zistené aj u čeľadí Hartmannellidae, Vannellidae a Paramoebidae. Zistené druhové zastúpenie nasvedčuje tomu, že spoločenstvo nahých meňaviek v skúmaných machoch je podobné spoločenstvám v sladkovodných biotopoch.

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Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 59–75

COMMUNITY STRUCTURE AND ECOLOGICAL MACRODISTRIBUTION OF MOSS-DWELLING WATER BEARS (Tardigrada) IN CENTRAL EUROPEAN OAK-HORNBEAM FORESTS (SW SLOVAKIA) PETER DEGMA, MAREK ŠIMURKA, SLÁVKA GULÁNOVÁ Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská dolina B-1, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected] Abstract Degma P., Šimurka M., Gulánová S.: Community structure and ecological macrodistribution of moss-dwelling water bears (Tardigrada) in Central European oak-hornbeam forests (SW Slovakia). Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 59–75. The structure of tardigrade communities in mosses of Central European oak-hornbeam forests was studied at 10 sites located in the Malé Karpaty Mts and Trnavská pahorkatina hills (SW Slovakia). A total of 3, 050 tardigrade specimens of 21 species and 2 families were gathered from 79 quantitative samples taken from 2000 and 2002. Kruskal-Wallis’ tests and regression analyses showed no statistically significant influence between the 12 studied environmental variables and the number of Tardigrada specimens or number of species in the samples. A chi-square goodness of fit test suggested that the number of Tardigrada species in samples of moss Hypnum cupressiforme was random within the investigated area. A t-test of tardigrade species diversity resulted in significant differences between study sites. The group of communities with the lowest diversity does not differ mutually. Majority of differences in species diversity were caused by randomly found species. Results of cluster analysis as well as CCA point out that distribution of tardigrades and their colonisation of particular substrata is a random process. Although the results are affected by restricted number of samples, we believe that Tardigrada as passively dispersed organisms are without significant relationship to ecological variables related to their distribution amongst substrata. Key.words:.Tardigrada, community structure, oak-hornbeam forests, Central Europe

ecological

macrodistribution,

mosses,

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Introduction Tardigrades living in mosses are organisms passively dispersed predominantly by wind but also by rain, floodwaters and melting snow. Occasionally animals visiting mosses can transport their specimens or eggs. Density of populations in colonized substrata can depend on several abiotic and biotic factors (Ramazzotti, Maucci, 1983). Community structure or ecological macrodistribution (distribution between substrata) of moss-dwelling tardigrades have been already discussed in more papers (Dastych, 1988; Ito, 1999; Peters, Dumjahn, 1999 etc.). Although these studies were based on quantitative data only few of them have used statistical methods so far (Nelson, 1975; Kathman, Cross, 1991; Wright, 1991; Nelson, Adkins, 2001; Nichols et al., 2001; Romano et al., 2001; Jönsson, 2003). The present study is a part of the project focused on microfauna and arthropod communities in oak-hornbeam forests (predominantly in soil) in SW Slovakia. From moss fauna, only tardigrades and active gymnamoebae (Mrva, 2005) were studied within this project. Only one study (Guoth, 1986) and several published records including the description of Echiniscus pajstunensis Bartoš, 1941 have referred to Tardigrada of this region (Bartoš, 1937a,b, 1941; Degma, 2003; Degma et al., 2004, 2005). The goal of this study is to characterize the structure of the tardigrade community in mosses of Central European oak-hornbeam forests using the data from SW Slovakia as well as to identify significant environmental variables which affect the structure.

Material and methods All the studied taxonomic groups including Tardigrada were sampled at the following study plots located in the Malé Karpaty Mts and Trnavská pahorkatina hills (SW Slovakia): Cajla, Vinosady, Fúgelka, Lindava Nature Reserve, Horný háj grove, Lošonec–lom quarry, Lošonský háj grove Nature Reserve, Naháč–Kukovačník, Naháč–Katarínka 1 Nature Reserve, Naháč–Katarínka 2 Nature Reserve (to locate study plots and their characterization see Zlinská et al., 2005). Quantitative samples of mosses (79 samples) were taken from each study plot in two periods: in 2000 (May, 31 or August, 15) and in 2002 (November, 7 or November, 8). Each quantitative sample was cut from substrate using metallic cylinder with surface of 10 cm2 with minimal addition of soil. Eight samples were taken from each the study plot with one exception (for number of samples in each study plot see Table 3). Immediately after cutting, each sample was put into a labelled paper bag for gradual drying. Laboratory processing of the samples also followed the procedure according to Dastych (1980) including submersion of a sample in a tap water for a period of 20–24 hours, detaching material from a substrate by agitation, squeezing and sedimentation. Extraction of tardigrades, their eggs and exuvia from sediment in Petri dish was done under the stereo-microscope (magnification 40x) using a micropipette. Specimens were mounted in Hoyer’s medium and the presence or an absence of eyes was noted before they dissolved in medium. Cover slips were sealed with asphalt varnish after medium desiccation. Specimens were identified under light microscope with phase contrast and oil immersion using species descriptions of Ramazzotti, Maucci (1983) as translated by Beasley (1995), original species descriptions and using the keys by Biserov (1990), Bertolani, Rebecchi (1993), Pilato, Binda (1999) and Pilato et al. (2000). Specimens of the genus Macrobiotus remained unidentified when their eggs were absent or only simplex specimens were found in a sample. Some specimens of the genera Hypsibius and Diphascon also remained unidentified as only simplex specimens were found.

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Environmental variables were measured or noted at each study plot (Zlinská et al. 2005) and the values 9.of them were used for comparisons (Table 1). Three additional variables were noted for each sample (Table.2).

pH of litter in H 2O

Forest fragmentation [%]

2 30 50 1 10 25 1 20 1 35

75 70 80 65 80 65 80 75 80 75

4.83 4.48 3.90 4.68 5.00 6.74 4.29 4.18 4.24 6.45

20 30 10 0 15 5 15 50 0 20

Forest stand age [years]

7 80–100 90 6 60–80 80 5 80–100 40 0 80–100 80 5 60–80 75 9 80–100 100 1 80–100 75 3 40–60 100 4 40–60 80 45 100–120 90

Slope [°]

E3 cover [%]

270 S 280 NW 350 S 240 none 240 W-SW 340 SW 260 NE 300 NE 340 NW 320 SE

E 2 cover [%]

Cajla Vinosady Fúgelka Lindava Nature Reserve Horný háj grove Lošonec–lom quarry Lošonský háj grove Nature Reserve Naháč–Kukovačník Naháč–Katarínka 1 Nature Reserve Naháč–Katarínka 2 Nature Reserve

E 1 cover [%]

CA VI FU LI HH LL LH NA NK1 NK2

Exposition

Study plot

Altitude [m a.s.l]

T a b l e 1. Values of chosen environmental variables at different study plots.

Constancy is defined as the number of study plots where the tardigrade species was found divided by total number of study plots and expressed as a percentage. Frequency is defined as the number of samples in which the tardigrade species occurred in the entire studied area divided by the total number of samples and also expressed as a percentage (Nelson, 1975). The calculation of Tardigrada species dominancy in study plots as well as in the whole area was based on all specimens found including the unidentified ones. Species diversity was measured using Shannon-Wiener’s index (natural logarithms were used) and the diversity of each community at the study sites was compared with a t-test (Poole, 1974; Spellerberg, Fedor, 2003). Kruskal-Wallis’ tests were done to evaluate the impact of nominal variables on the total number of Tardigrada specimens as well as on the number of species at the sites, underbeds, study plot exposure, age of a forest stand and month of sampling using all 79 samples. Species of mosses did not enter into any Kruskal-Wallis’ test as their distribution between the samples was very unequal, most samples –54 of them– were taken from the dominant species Hypnum cupressiforme and number of samples taken from rest of moss species varied from 1 to 5. Chi-square Goodness of fit test with Poisson distribution was performed to test whether Tardigrada species in the samples of moss Hypnum cupressiforme were distributed randomly in the studied area. Simple linear regression analyses were done to test relationships between dependent number of tardigrade specimens or number of their species in all 79 samples and each of 7 gradient variables. Stepwise regression analysis with forward selection of gradient variables was performed to identify variables with significant influence on the number of specimens or the number of species in the samples. Kruskal-Wallis’s tests as well as regression analyses and Chi-square Goodness of fit test were performed using computer programme Statgraphics Plus V.7.0 (Manugistics, 1993). The cluster analysis of the communities at all the study sites were done using the computer program Nclas from the package Syn-Tax (Podani, 1993). The complete linkage clustering method in combination with Wishart’s similarity ratio index for clustering according to abundance of species transformed with log-transformation was used. Canonical Correspondence Analysis (CCA) with forward selection of environmental variables using Monte Carlo permutation tests for significance of each variable as well as for significance of the first canonical axis was performed using the program Canoco (ter Braak, Šmilauer, 1998). The goal was to find significant

61

T a b l e 4. Results of t-tests of Shannon-Wiener’s species diversity H’ for the Tardigrada communities in couples of study plots. Study plot Study plot/H' CA VI FU LI HH LL LH NA NK1 NK2

CA 0.529 1.600ns 2.017* 0.377ns 13.514*** 1.699ns 12.882*** 15.478*** 6.043*** 8.125***

VI 0.677 169 0.011ns 1.401ns 9.504*** 1.225ns 5.908*** 7.901*** 2.524* 5.979***

FU 0.676 489 271 1.763ns 10.622*** 1.246ns 7.337*** 10.546*** 3.031** 6.457***

LI 0.548 1663 169 488 13.324*** 1.644ns 12.527*** 15.926*** 5.765*** 7.981***

HH 1.891 75 148 120 75 2.287ns 6.184*** 19.949*** 8.549*** 1.954ns

LL 1.099 3 3 3 3 4 0.437ns 3.296* 0.472ns 1.364ns

LH 1.245 675 195 533 663 86 3

NA 0.000 931 126 304 753 59 3 291

28.418*** 4.194*** 16.045*** 2.554* 12.648***

NK1 0.939 385 241 521 385 108 3 433 220

NK2 1.584 44 77 60 44 78 4 48 38 56

4.669***

Notes: for abbreviations of study plots see Table 1; t-values are under the diagonal and corresponding degrees of freedom are above it; ns non significant – P > 0.05, * 0.01 < P < 0.05, ** 0.001 < P < 0.01, *** P < 0.001; P is probability for t-value

environmental variables influencing the structure of the Tardigrada community. Pooled abundance of each species at particular study plots transformed with log-transformation Y = log (Y + 1) entered this analysis together with the values of 9 environmental variables from the 13 mentioned above. Variables ‘study plot’ (defined analysed samples) as well as underbed, substrate species and month of sampling were omitted from the analysis (the later three ones had various values in particular samples). CCA was also performed for examination of the relationship between Tardigrada species and all 13 environmental variables to identify significant ones influencing species assemblages in the 79 samples expressed by transformed abundance of each species with log-transformation. Downweighting of rare species was used to suppress the influence of randomly found species that were collected in low numbers on the overall result of this analysis. Only the identified specimens were taken into all the statistical analyses with exception of Kruskal-Wallis’ tests and regression analyses when total numbers of all Tardigrada specimens in the samples were involved. Nomenclature of tardigrades is according to Guidetti, Bertolani (2005) and that of mosses according to Kubínska, Mišíková (1998).

Results and discussion Structure of Tardigrada communities in oak-hornbeam forests Abundancy and number of Tardigrada species A total of 3, 050 specimens of water bears were found and identified except for 164 (5.4%) which lacked eggs or were only found in simplex stage. One class, 2 families and 21 species were represented in the 79 quantitave samples. Tardigrades were absent in 24.(30.4%) of the 79 analysed samples. The number of samples without tardigrades varied between 0 (CA) and 6 (LL, 75% of samples) at the study sites. The lowest number of specimens was recorded in LL (21 specimens in two positive samples) and the highest in CA (953 specimens in 8 positive samples). Minimal numbers of species were detected at study site NA (1 species) and LL (3 species) while maximum species richness was in HH (9 species) (Table 3). The study plot NA is the most isolated (a small forest surrounded by fields) and could be influenced by dust rising during agricultural work. Moreover the site LL was covered with calcareous dust from a near by limestone quarry. Out of 21 identified species 12 (Hypsibius convergens, H. dujardini, H. pallidus, Isohypsibius lunulatus, I. prosostomus, Diphascon pingue, D. belgicae, D. prorsirostre, D. scoticum, Astatumen trinacriae, Macrobiotus hufelandi and Minibiotus intermedius) were found at the same altitudes in adjacent Poland. However the Poland species were found in a significantly broader range of altitudes. On the other hand, Astatumen bartosi and Macrobiotus pallarii were recorded in different altitudinal zones in Poland (Dastych, 1988). We have found only Isohypsibius lunulatus from all the upland species which have the center of their distribution in the zone of 201–500 m a.s.l. according to his classification of tardigrades. In the same altitudinal zone in Newfoundland, only seven species were found (Collins, Bateman, 2001). Out of those, four species (Macrobiotus cf. harmsworthi, M. hufelandi, Minibiotus intermedius and Diphascon scoticum) were also recorded at our study sites. Out of the 8 species which prefer (although some of them only slightly) altitudinal zone of 0–500 m a.s.l. based on their 63

T a b l e 3. Dominancy [%] of species at the study sites, their total dominancy (D), constancy (C) and frequency (F). Taxon

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

Class: EUTARDIGRADA Richters, 1926 Family: Hypsibiidae P i l a t o, 1969 Hypsibius convergens (U r b a n o w i c z, 1925) Hypsibius dujardini (D o y e r e, 1840) Hypsibius cf. morikawai I t o, 1995 Hypsibius pallidus T h u l i n, 1911 Isohypsibius lunulatus (I h a r o s, 1966) Isohypsibius prosostomus T h u l i n, 1928 Diphascon (D.) brevipes (M a r c u s, 1936) Diphascon (D.) pingue (M a r c u s, 1936) Diphascon (A.) belgicae R i c h t e r s, 1911 Diphascon (A.) prorsirostre T h u l i n, 1928 Diphascon (A.) scoticum M u r r a y, 1905 Astatumen bartosi (W e g l a r s k a, 1959) Astatumen trinacriae (A r c i d i a c o n o, 1962) Family: Macrobiotidae T h u l i n, 1928 Macrobiotus cf. harmsworthi M u r r a y, 1907 Macrobiotus hufelandi C.A.S. S c h u l t z e, 1834 Macrobiotus pallarii M a u c c i, 1954 Macrobiotus cf. seychellensis B i s e r o v, 1994 Macrobiotus cf. vanescens P i l a t o et al., 1991 Macrobiotus sp.1 Minibiotus intermedius (P l a t e, 1889) Minibiotus sp. 1 Total Number of specimens Total Number of samples (Nr. of negative ones) Number of moss species

Note: for abbreviations of study plots see Table 1

CA

VI

FU

12.51 3.13

LI

Study site HH LL

D [%]

C [%]

F [%]

0.39

4.94

0.8 0.1 0.1 3.3 0.4 0.2 0.5 7.9 0.0 1.3 0.1 0.0 0.5

40.0 10.0 10.0 20.0 20.0 30.0 20.0 80.0 10.0 30.0 10.0 10.0 70.0

7.6 1.3 1.3 2.5 5.1 3.8 3.8 26.6 1.3 6.3 1.3 1.3 10.1

2.63 11.0 59.1 1.2 28.95 0.4 4.76 0.6 0.2 39.38 0.45 6.9 0.0 21 353 167 224 38 8 (6) 8 (1) 8 (2) 8 (1) 7 (2) 3 2 2 4 7

40.0 50.0 20.0 10.0 20.0 10.0 30.0 10.0

15.2 11.4 2.5 1.3 2.5 1.3 7.6 1.3

LH

NA

NK1 NK2 0.45 5.26

78.91

0.28 11.11

0.52 5.14

7.89 4.76

3.91

5.79

1.05

7.40 0.64

0.10

1.29

5.30

1.23 4.94

0.13

3.70

0.45 7.93

34.21 40.18 18.42 0.45 2.68 2.63

5.88 84.99

4.76

1.13

0.89 96.41 52.68

81.03 82.56 12.34 11.11 22.22 1.61 8.91

1.23 953 128 311 774 81 8 (0) 8 (3) 8 (3) 8 (3) 8 (3) 2 2 3 3 6

25.78 7.93

frequency in the samples according to Maucci (1981), only Macrobiotus hufelandi, Minibiotus intermedius, Hypsibius pallidus and Diphascon scoticum were present in our material. Finally, Guoth (1986) identified four species (Macrobiotus hufelandi, Hypsibius dujardini, H. microps and Diphascon scoticum) in oak-hornbeam forest in southern parts of the Malé Karpaty Mts (study plot Borinka). Most of Gouth’s (1986) species were found in our material as well. It is necessary to say that species like Hypsibius dujardini were identified only in a single sample and the limited number of recorded species may be caused by limited samples. No single tested nominal variable was found significant (P > 0.05 for all values of Kruskal-Wallis’ statistics) neither for Tardigrada specimens number nor for number of their species when each variable was tested separately by Kruskal-Wallis’ test. This result partly reflects relatively small number of values of specimens or species numbers in each category (7–8 values in categories of a study plot, 13–23 values in each underbed category, 7–16 values in study plot exposure categories, 7–40 values in each age of a forest stand and 19–38 values in each month of sampling) as a small number of values makes confidence intervals for median (as well as for mean) broader and these intervals overlap in spite of appropriate mean values are very different from each other. For example average number of specimens in one sample in LL is less than one what is in contradiction with mean in CA when it is more than 116. Steiner (1994) noticed the highest abundance of Tardigrada in May and a population decline in November and March. The average of Tardigrada quantity in our samples was 52.9 in May, 22.3 in August and 33.3 in November, so as it might seem our research partially confirmed Steiner’s findings. It is necessary to mention that the same comment about the differences among study plots can be applied in this case. Due to broad confidence intervals for these medians there is no statistical difference between those seasonal ones in our numbers of specimens per sample. In contradiction to us, Nichols et al. (2001) ascertained higher number of specimens as well as species in spring than in summer and fall of the same year using statistical methods in Alabama. However similarly they did not find a significant difference in these parameters among the sampling stations. The statistical results of Romano et al. (2001) indicated fully inverted distribution as they discovered no significant difference in numbers of Tardigrada specimens during season but these numbers were significantly different among the sampling sites in the other locality in Alabama. These two antagonistic results illustrate how some most different samples – the only sample from spring 1977 in Nichols et al. (2001) and two samples from two sites in Romano et al. (2001) can dramatically influence overall result of statistical analysis. That is why the authors of both the studies emphasized a need of more replicate samples to reduce the significant variability due to random patchiness in tardigrade populations. From these reasons, we do not believe the results of both authors reflect real habitat or seasonal differences. From all seven gradient variables, only cover of E2 is significant but only for number of Tardigrada species and not for number of their specimens in moss samples (F = 4.336, P = 0.041, R2 = 5.3%) when each variable impact was separately tested by simple linear regression analysis. Coefficient of determination R2 is too small hence we do not consider even this variable to be significant. We did not obtain sufficiently good general linear model explaining tardigrade specimen number in the samples when stepwise regression analysis was performed 65

because the coefficient of determination was too small. For number of species (NS), the best obtained model is NS = 0.0192*E3 – 0.0175*E2 (t = 2.812, P = 0.032 for coefficient of E2; t = 7.585, P < 0.001 for coefficient of E3; F = 40.110, P < 0.001, R2 = 50.7% for model). As we can see, this model explains only 51% of species number variability in moss samples and so major part of this variability remains unexplained. We think that Tardigrada species number in mosses is either largely random or it is influenced by variables not included into our study. The fact, that altitude does not appear to be important variable for specimen or species number per sample, is not surprising with respect to too small interval of altitudes of our study plots (240–350 m a.s.l.). But interesting is, that the results of two multivariate statistical methods (principal components analysis and cluster analysis) also indicated that the abundance of moss-dwelling tardigrades on Vancouver Island was not dependent upon the altitude (Kathman, Cross, 1991) when they were sampled even in wider range of altitudes (150–1525 m a.s.l.). In our opinion, the mentioned results indicate that moss-dwelling tardigrades either do not significantly prefer any of the tested nominal variable category or interval of gradient variable values or preference of some categories by any species is statistically balanced by preference of other categories by other species in a community. We could consider these results to support the hypothesis that distribution of Tardigrada on mosses (as well as on other substrata) is a random process and due to their broad ecological valence they survive in colonized substrata whereas their population density and microdistribution can be simply various in time or under influence of some ecological variables. For example, according to Wright (1991) three species – Macrobiotus hufelandi, M. richtersi and Isohypsibius prosostomus are in negative associations. Our samples support this statement as two of three mentioned species (without Macrobiotus richtersi) were present in them and they never occurred together. As it concerns tardigrades migration in moss cushion (thus their microdistribution), Wright (1991) discovered vertical migration during dehydration of cushion in M.richtersi while according to Nelson, Adkins (2001) migration within the moss cushion was not detected as a result of changes in moisture conditions. Finally, population density may be the duration which elapsed since concrete substratum was colonized by tardigrades. Number of tardigrade species in samples of moss Hypnum cupressiforme from all the study plots varied between 0 and 4. The result of Chi-square Goodness of fit test with Poisson distribution for number of tardigrade species in these samples (χ2 = 1.198, P = 0.549) allowed us to not reject the null hypothesis that the number of species in moss sample is random and with good agreement with Poisson distribution. So Tardigrada species colonize this moss species cushions in oak-hornbeam forests in sampled area randomly. It is very probable that the same is valid for other substrata with regard to mode of tardigrade distribution. From this reason it is not surprising that the greatest number of tardigrades species was found in the moss species which most samples were taken from. From others moss species, it is notable that even four species of Tardigrada were found in a single sample of Dicranum scoparium as well as even seven of them in only two samples of Anomodon viticulosus. But this is not in contradiction with our presumption that number of Tardigrada species is random also in these moss species.

66

This hypothesis can be confirmed or refused only by testing of sufficiently large number of samples. Dominancy, constancy, frequency and species diversity Nearly 70% of samples were positive in Tardigrada in the study area while in Poland there were only 29% positive samples in these types of forests (Dastych, 1988). The most dominant species in the study area were Macrobiotus hufelandi, M. cf. harmsworthi, Diphascon pingue, Minibiotus intermedius and Hypsibius pallidus. But the dominancy of these species varied at different study sites considerably (Table 3). All these species were relatively frequently present in the samples but only four species of all occurred in more than 10% of the samples: Macrobiotus cf. harmsworthi, M. hufelandi, Diphascon pingue and Astatumen trinacriae. Of them, Diphascon pingue, Astatumen trinacriae and Macrobiotus hufelandi have even the highest constancy values (Table 3). Species diversity was lowest (beyond that on the study plot NA, when it was equal to zero with respect to only one species found) at the study plot CA and the highest on HH. T-tests of species diversity (Table 4) allowed us to differentiate a group of communities with their diversity as not significantly different. Communities with the lowest species diversity (CA, LI, FU and VI) with 4–6 Tardigrada species present and simultaneously one of them having very high dominancy (around 80%) belonged into this group. M. hufelandi was the dominating species at all the mentioned study plots with exception for the study plot VI where it was „substituted“ by Hypsibius pallidus. The mentioned study plots do not differ from the others in values of the studied environmental variables hence it is not possible to explain their difference in species diversity on this basis. Species diversity at the other sites mostly differ from those mentioned above as well as among each other. These differences are mostly caused by species which were present in only one or two samples within series from each study plot. Only four species (Diphascon pingue, Macrobiotus cf. harmsworthi, M. hufelandi and Minibiotus intermedius) occurred in more than two samples in some of the study plots. So due to restricted number of samples at each study site, the occurrence of most of species seem to be random. Tardigrada communities at the study sites The results of Tardigrada community clustering according to pooled log-transformed abundance of species at the study sites is presented in Fig. 1. Five clusters can be recognized on high dissimilarity level (from left to right in Fig. 1): 1 – communities on study plots CA, FU, LI, LH and HH, 2 – community on NK2, 3 – community on VI, 4 – community on LL and 5 – communities on NA and NK1. The considerable component of the communities in the first cluster was Macrobiotus hufelandi (not found in the other communities). Also the species Astatumen trinacriae (dominancy up to 4%) and Diphascon pingue (dominancy app. 5–8 %) were found in all these communities. We can recognize 3 subgroups within this cluster: a – communities in CA and FU (relatively significant representation of Diphascon prorsirostre), b.–.communities in LI and LH (nearly all the found specimens of Minibiotus 67

T a b l e 4. Results of t-tests of Shannon-Wiener’s species diversity H’ for the Tardigrada communities in couples of study plots. Study plot Study plot/H' CA VI FU LI HH LL LH NA NK1 NK2

CA 0.529 1.600ns 2.017* 0.377ns 13.514*** 1.699ns 12.882*** 15.478*** 6.043*** 8.125***

VI 0.677 169 0.011ns 1.401ns 9.504*** 1.225ns 5.908*** 7.901*** 2.524* 5.979***

FU 0.676 489 271 1.763ns 10.622*** 1.246ns 7.337*** 10.546*** 3.031** 6.457***

LI 0.548 1663 169 488 13.324*** 1.644ns 12.527*** 15.926*** 5.765*** 7.981***

HH 1.891 75 148 120 75 2.287ns 6.184*** 19.949*** 8.549*** 1.954ns

LL 1.099 3 3 3 3 4 0.437ns 3.296* 0.472ns 1.364ns

LH 1.245 675 195 533 663 86 3

NA 0.000 931 126 304 753 59 3 291

28.418*** 4.194*** 16.045*** 2.554* 12.648***

NK1 0.939 385 241 521 385 108 3 433 220

NK2 1.584 44 77 60 44 78 4 48 38 56

4.669***

Notes: for abbreviations of study plots see Table 1; t-values are under the diagonal and corresponding degrees of freedom are above it; ns non significant – P > 0.05, * 0.01 < P < 0.05, ** 0.001 < P < 0.01, *** P < 0.001; P is probability for t-value

intermedius), c – community in HH (nearly all found specimens of Macrobiotus cf. vanescens – the richest community with 9 species). This first cluster is nearly identical with a group of study plots with statistically same species diversity (CA, VI, FU and LI). Community in VI does not belong to this cluster due to the absence of M. hufelandi. The differences between results of species diversities t-tests and cluster analysis are caused by the fact that species composition plays a role in cluster analysis while species may be mutually substituted in calculation and testing of species diversities.

Fig. 1. Dendrogram of Tardigrada communities at the study sites (for abbreviations of study plots see Table 1; scale of dissimilarity in vertical axis).

Community in NK2 was distinguished from the others by presence of some species which were found only here (Hypsibius cf. morikawai, Astatumen bartosi, Macrobiotus cf. seychellensis) as well as by concentration of nearly all the specimens of Diphascon brevipes. Community in VI is different from the others in presence of all the specimens of Hypsibius dujardini and nearly all the specimens of H. pallidus as well as with absence of Macrobiotus cf. harmsworthi and M. hufelandi. Community in LL is the second poorest one with only three identified species (all present in some other communities) and also with absence of M. cf. harmsworthi as well as M. hufelandi. Finally, the communities in NA and NK1 are joined by meaningful proportion of M. cf. harmsworthi specimens however the community in NA contained only this species (the purest one) on the contrary to community on NK1 with 8 species (second richest one). As we can see the studied communities embody relatively large structural differences. However, the presence of concrete species in the only sample was the cause of their mutual differentiation in some cases: Diphascon prorsirostre on CA and FU, 69

Macrobiotus cf. vanescens on HH, Hypsibius cf. morikawai, Astatumen bartosi and Macrobiotus cf. seychellensis on NK2 and Hypsibius dujardini and H. pallidus on VI. Therefore we believe that the result of cluster analysis is largely caused by random distribution as well as by apparently small number of samples at different study plots. Environmental variables and community structure Only two gradient variables – slope of study plot and pH of litter in H2O – from amongst 7 gradient and 2 categorial variables were significant as explanatory (their P < 0.05 when tested by Monte Carlo permutation test) in the CCA when the pooled log-transformed abundances of particular species at the study plots were used as species data (Fig. 2). Eigenvalues of the two first canonical axes are λ1 = 0.614 and λ2 = 0.340. The first two canonical axes account for 35% of the total variance of the species data and 100% of the species-environment relation. First canonical axis is significant (F = 2.030, P = 0.015) when tested by Monte Carlo permutation test.

Fig. 2. Biplot of CCA of the communities at the study sites according to pooled abundance of species (for abbreviations of study plots see Table 1; A species 3, 12 and 17, B species 11 and 19; for species codes see Table 3; species found only in one or two communities are in ellipse together with appropriate community).

70

Variable slope was significant as it had expressively extreme value of 45° at the study plot NK2. Similarly, pH of litter in H2O had the second maximum value 6.45 at the same study plot while the maximum value of 6.74 was recorded at the study plot LL due to calcareous dust which is produced by the nearby stone quarry. That is the reason why communities on NK2 and LL have extreme position on biplot. Among them, the community on HH with the third maximum pH value is situated. The next two communities (LL and HH) have no significant relation to slope regarding its low values being compared with that on NK2. The rest of communities are located at the other end of its scale and their location on the biplot is due to species representation and relation of their species to two gradient variables. Similarly, the species position on biplot is not only due to their relation to two significant variables but also to their communities. For example, three species Hypsibius cf. morikawai, Astatumen bartosi and Macrobiotus cf. seychellensis (position A on biplot) were found only in NK2. Two other species were detected as mutual in communities in NK2 and HH: Isohypsibius lunulatus and Diphascon brevipes (see Table 3 and biplot on Fig. 2). Though it seems that three species found only in NK2 have a real relationship to localities with steep slopes it does not have to be, as each of these species was found only in one sample so it concerned accidental findings. The relation of Isohypsibius lunulatus and Diphascon brevipes to higher values of litter pH is rather different as the second species mentioned was found in three while the first one was found in four samples (on each occasion in two samples) from these two study plots. However, we are not fully convinced that dependence of these species on less acid substratum is hereby demonstrated as they lacked on study plot LL. The result of CCA is in good agreement with the result of cluster analysis as groups of communities we can recognize in both the graphs are practically the same (Figs 1 and 2). We believe that the result of CCA was affected not only by extreme values of two significant gradient variables but also by abundance of the same species which determined the result of cluster analysis.

Ecological macrodistribution of Tardigrada species in oak-hornbeam forests Only three categories of two nominal variables – study plot VI, two age intervals of forest stand 40–60 and 80–100 years – from amongst 13 variables were significant as explanatory (their P < 0.05 when tested by Monte Carlo permutation test) in the CCA when the log-transformed abundances of particular species in the samples were used as species data (Fig. 3). Eigenvalues of the two first canonical axes are λ1 = 0.524 and λ2 = 0.444. The first two canonical axes account only for 9% of the total variance of the species data and 73% of the species-environment relation. First canonical axis is significant (F = 2.278, P = 0.042 when tested by Monte Carlo permutation test). 40–60 year old forests were represented only at study sites NA and NK1 while the study plot NK2 was covered by an 80–100 year old forest. The study plot VI, together with HH, are the only plots in which the forest was 60–80 years old. So it seems that age 71

of a forest stand is the only one, of all the studied variables, affecting quantitative structure of tardigrade species in different moss samples.

Fig. 3. Biplot of CCA of the communities at the study sites according to abundance of species in samples (VI study site Vinosady, 40–60 and 80–100 = age intervals of forest stand; A species 3, 5, 7, 12, 17 and 21; for species codes see Table 3; species found only in one or two communities are in ellipse together with appropriate community).

There were no remarkable relationships between species distribution and forest stands of a certain age interval except for a few exceptions which were seen in several species found in one random sample (Diphacon belgicae in 40–60 year old forest stand, 72

Diphascon scoticum and Macrobiotus sp. 1 in 80–100 year old forest stand, Hypsibius dujardini and Minibiotus sp. 1 in 60–80 year old stand and Hypsibius cf. morikawai, Astatumen bartosi and Macrobiotus cf. seychellensis in 100–120 year old stand). Thus for example Macrobiotus cf. harmsworthi, Diphascon prorsirostre, Isohypsibius prosostomus and Minibiotus intermedius were found in 40–60 year old stands as well as in those of 80–100 years of age. Similarly, Macrobiotus hufelandi, M. pallarii and M. cf. vanescens were discovered in 60–80 years old forests as well as in those of 80–100 years of age. The other species occurred in a broader age spectrum of forest stands. From those we can mention the most constant species such as Astatumen trinacriae (age interval 40–100 years) and Diphascon pingue (all the age categories studied). It is necessary to emphasize that most of the mentioned species were found in only one sample at the sites (for example Macrobiotus cf. harmsworthi on NK2, Diphascon prorsirostre on FU, Isohypsibius prosostomus on CA, LL and NK1, Macrobiotus pallarii on HH and LH, M. cf. vanescens on HH and LL). On distant position A (Fig. 3), we find those species having no affinity to any from mentioned three significant categories. In addition also every from these species was found in each occasion only in one sample. Those are: Hypsibius cf. morikawai, Isohypsibius lunulatus, Diphascon brevipes, Astatumen bartosi, Macrobiotus cf. seychellensis and Minibiotus sp. 1. We believe that two significant categories of forest stand age substitute study plots covered by those forests. Thus category 40–60 aged forest substitutes study plots NA and NK1 as well as category 80–100 aged stand substitutes study plots CA, FU, LI, LL and LH. From this visual angle, the result of this CCA is in quite agreement with the result of cluster analysis as well as with that obtained by previous CCA of tardigrades communities in study plots (Figs 1, 2). We do not consider any of the variables to be really significant for uniqueness of findings of many tardigrade species responsible for the result of CCA. Thus we believe that water bears as passively dispersed organisms have no significant relationship to a majority (if any) of ecological variables since they dispose of sufficiently broad ecological valence to maintain in randomly colonized substrate in larger or negligible density. Variables potentially having significant influence to population density changes (species associations, humidity) were partly discussed in this study. Translated by P. Degma Acknowledgement We would like to thank to B. Nichols (U.S.A.) for his valuable comments and the correction of the English version of the manuscript. We also give thanks to E. Miadoková and E. Gálová (Slovakia) who were so kind to provide the microscope with phase contrast available to us, to K. Mišíková (Slovakia) for identification of mosses and to Daphne (Institute of Applied Ecology, Slovakia) for providing the programme Canoco. This study was partly supported by the Slovak Scientific Grant Agency VEGA as a Project No. 1/0119/03.

73

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Pilato, G., Binda, M.G., Napolitano, A., Moncada, E., 2000: The specific value of Macrobiotus coronatus De Barros 1942, and description of two new species of the harmsworthi group (Eutardigrada). Boll. Acc. Gioenia Sci. Nat., 33, 358, p. 103–120. Podani, J., 1993: Syn-Tax. Version 5.0. Computer Programs for Multivariate Data Analysis in Ecology and Systematics. User’s guide. Scientia Publishing, Budapest, 104 pp. Poole, R.W., 1974: An Introduction to Quantitative Ecology. McGraw-Hill, 532 pp. Ramazzotti, G., Maucci, W., 1983: Il Philum Tardigrada. (III. edizione riveduta e aggiornata). Memoire dell’ Istituto Italiano di Idrobiologia, 41, 1016 pp. Romano, III, F.A., Barreras-Borrero, B., Nelson, D.R., 2001: Ecological distribution and community analysis of Tardigrada from Choccolocco Creek, Alabama. Zool. Anz., 240, p. 535–541. Spellerberg, I.F., Fedor, P.J., 2003: A tribute to Claude Shannon (1916-2001) and a plea for more rigorous use of species richness, species diversity and the "Shannon-Wiener" Index. Global Ecology and Biogeography, 12, 3, p. 177–179. Steiner, W. A., 1994: The influence of air pollution on moss-dwelling animals: 4. Seasonal and long-term fluctuations of rotifer, nematode and tardigrade populations. Rev. Suisse Zool., 101, p. 1017–1031. Ter Braak, C.J.F., Šmilauer, P., 1998: CANOCO References Manual and User’s Guide to Canoco for Windows: Software for Canonical Community Ordination (version 4). Microcomputer Power (Ithaca, NY, USA), 352 pp. Wright, J. C., 1991: The significance of four xeric parameters in the ecology of terrestrial Tardigrada. J. Zool., 224, p. 59–77. Zlinská, J., Šomšák, L., Holecová, M., 2005: Ecological characteristics of studied forest communities of an oak-hornbeam tier in SW Slovakia. Ekológia (Bratislava), 24, Suppl. 2, p. 3–19. Received 30. 7. 2005

Degma P., Šimurka M., Gulánová S.: Štruktúra spoločenstva a ekologická makrodistribúcia pomaliek (Tardigrada) žijúcich v machoch stredoeurópskych dubovo-hrabových lesov (JZ Slovensko). Študovali sme štruktúru spoločenstiev pomaliek (Tardigrada) v machoch stredoeurópskych dubových lesov na 10 študijných plochách situovaných v Malých Karpatoch a Trnavskej pahorkatine (JZ Slovensko). Zo 79 kvantitatívnych vzoriek odobratých v rokoch 2000 a 2002 sme získali 3 050 jedincov pomaliek 21 druhov a dvoch čeľadí. Kruskal-Wallisove testy a regresné analýzy ukázali, že žiadna z 12 študovaných environmentálnych premenných nemá štatisticky významný vplyv na počet jedincov alebo druhov Tardigrada vo vzorkách. Chí-kvadrátový test dobrej zhody naznačil, že počet druhov Tardigrada vo vzorkách machu Hypnum cupressiforme bol v.rámci skúmanej oblasti náhodný. Výsledkom t-testu druhovej diverzity boli významné rozdiely medzi študijnými plochami. Skupina spoločenstiev s najnižšou diverzitou sa vzájomne nelíši. Väčšinu odlišností v druhovej diverzite spôsobili náhodne nájdené druhy. Výsledky zhlukovej analýzy ako aj CCA poukazujú na to, že šírenie pomaliek a ich osídľovanie jednotlivých substrátov je náhodný proces. Hoci sú výsledky ovplyvnené obmedzeným počtom vzoriek, domnievame sa, že Tardigrada ako pasívne sa šíriace organizmy majú nevýznamnú väzbu na ekologické premenné súvisiace s ich distribúciou medzi substrátmi.

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Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 76–86

COMMUNITIES OF PSEUDOSCORPIONS (Pseudoscorpiones, Arachnida) IN EPIGEON OF OAK-HORNBEAM FORESTS IN THE MALÉ KARPATY MTS AND TRNAVSKÁ PAHORKATINA HILLS (SW SLOVAKIA) JANA CHRISTOPHORYOVÁ, MIROSLAV KRUMPÁL Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská dolina, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected], [email protected] Abstract Christophoryová J., Krumpál M.: Communities of pseudoscorpions (Pseudoscorpiones, Aarachnida) in epigeon of oak-hornbeam forests in the Malé Karpaty Mts and Trnavská pahorkatina hills (SW Slovakia). Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 76–86. Research on pseudoscorpion communities was performed at four study plots of the Malé Karpaty Mts – Lošonský háj grove, Lošonec–lom quarry, Cajla, Vinosady and at two study plots of the Trnavská pahorkatina hills – Horný háj grove and Lindava during the years 2000–2002. The soil macrofauna was collected by the square method combined with sifting. A.total of 949 specimens belonging to 10 taxa were examined, seven of them were identified on species level. The species Neobisium muscorum (L e a c h, 1817) was the only species common for all the study plots. The Sørensen’s and Wishart’s indices of similarity and PCA have confirmed the presence of four community types, which differed in species composition, dominance, equitability, diversity and mean abundance of pseudoscorpions. We suggest that the community type of the 80–100 year old forests – Lošonský háj grove and Lošonec–lom quarry was the most stable, because of the highest diversity and equitability recorded there. The most unstable community type was represented by the community of the younger (60–80 year old) oak-hornbeam forest Vinosady, which was the most arid and it had a very poor leaf litter layer. The lowest species richness, the lowest mean abundance and diversity were recorded there. In all the study plots, 15 selected environmental variables were measured with aim to find out environmental variables influencing the community. Unfortunately, none of environmental variable had the statistically significant influence on the occurrence of pseudoscorpions. Key words: Pseudoscorpiones, false-scorpions, Slovakia, communities, oak-hornbeam forests

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Introduction Like most other arachnids, pseudoscorpions are predators. They may be found in many habitats and microhabitats, such as woodland, fields, grassland, soil, leaf litter, underneath stones or in nests of birds. Some species are synanthropic. Their small size, life stages and the fact that pseudoscorpions have to be mounted on permanent slides for their identification have contributed to insufficient knowledge on their taxonomy, species richness and ecology. In the Slovak and Czech Republic, the research of pseudoscorpions is still at beginning. Verner (1959) published a study on arthropods of oak-hornbeam forest of the Karlštejnsko (Czech Republic), recording four species of pseudoscorpions. Ducháč (1988) investigated the occurrence of pseudoscorpions in oak forests on two localities of České středohoří Mts (Czech Republic). He found 15 specimens only, which belonged to 3 species. Ducháč (1993) recorded 4 species of false-scorpions from oak hollows in Třeboň in the Czech Republic. Šťáhlavský (2001) published the occurrence of false-scorpions in many microhabitats in the Praque region (Czech Republic). In his work, a special attention was given onto tree hollows (including Carpinus betulus, Quercus pubescens and Q. robur). Systematic faunistic and ecological investigation on pseudoscorpions of the Poloniny National Park (Slovak Republic) was carried out by Krumpál, Krumpálová (2003). With regard to deciduous forests, the occurrence of pseudoscorpions was recorded mainly in beech and oak forests. In other countries, the main subject of the research on pseudoscorpions from deciduous forests was their life cycle. Life cycles of some British pseudoscorpions on beech, oak and sycamore-ash forests were studied by Gabbutt, Vachon (1965), Gabbutt (1969), Goddard (1976). Wood, Gabbutt (1978) evaluated the phenology and seasonal vertical distribution of Neobisium muscorum (L e a c h, 1817) (i n s e n s u Gabbutt, Vachon, 1965), Chthonius ischnocheles (H e r m a n n, 1804) and Roncus lubricus L. K o c h, 1873 from beech litter in Oxon, England. Meyer et al. (1985) dealt with the population structure and dynamics of Neobisium carcinoides (H e r m a n n, 1804) in oak forests in North Tirol (Austria). The other predators were studied in oak-hornbeam forests in Slovakia as well. Országh, Országhová (2005) investigated the structure of centipede communities in oak-hornbeam forests of the Malé Karpaty Mts and Trnavská pahorkatina hills. Krumpálová, Bartoš (2002) analysed the spider communities, especially lycosids, in four types of oak-hornbeam forests near the city Modra. Krumpálová, Szabová (2003); Krumpálová, Szabová (2005) compared the epigeic spider communities in natural and human impacted oak-hornbeam forests. The aims of the present study were (i) to describe the quantity and quality of the pseudoscorpion communities, (ii) to study the pseudoscorpion community structure – dominance, diversity and equitability, (iii) to compare the communities of six study plots, (iv) to find out environmental variables influencing the communities.

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Material and methods Study plots: The investigation of the pseudoscorpion communities was performed at four study plots of the Malé Karpaty Mts – Lošonský háj grove (LH), Lošonec–lom quarry (LL), Cajla (CA), Vinosady (VI) and at two study plots of the Trnavská pahorkatina hills – Horný háj grove (HH) and Lindava (LI) during the years 2000–2002. The study plot Lošonský háj grove (48°28' N, 17°24' E, Grid Reference Number of the Databank of the Fauna of Slovakia – GRN 7570b, 260 m a.s.l.) and Lošonec–lom quarry (48°29' N, 17°23' E, GRN 7570a, 340 m a.s.l.) represent 80–100 year old forests. The vegetation of these forests belongs to the subassociation Querco-Carpinetum caricetosum pilosae. The study plot Lošonec–lom quarry is covered by the calcareous dust from the nearby quarry. The vegetation of the old forest (80–100 year old) of study plot Cajla (48°20' N, 17°16' E, GRN 7669c, 260-280 m a.s.l.) belongs to the subassociation Querco-Carpinetum poetosum nemoralis. The study plot Lindava (48°22' N, 17°22' E, GRN 7670a, 240 m a.s.l.) is also situated in the old forest (80–100 year old) of the subassociation Quercetum petraeae - cerris. The study plot Vinosady (48°19' N, 17°17' E, GRN 7669d, 280 m a.s.l.) represents younger forest (60–80 year old), its vegetation belongs to the subassociation Querco-Carpinetum poetosum nemoralis. The 60–80 year old forest represents the study plot Horný háj grove (48°29' N, 17°27' E, GRN 7570b, 240 m a.s.l.) with the vegetation of the subassociation Querco-Carpinetum melicetosum uniflorae. More detailed characteristics of these study plots have been described by Zlinská et al. (2005). The soil macrofauna was collected by the square method combined with sifting. At app. 1-month intervals from March to November, the material was collected from the leaf litter and the upper part of soil from 16 squares at each study plot. Each square has included 25x25 cm of the area, i.e. altogether an area of 1 m2 was sifted, representing one sample. The samples were extracted using xereclectors of the Moczarski’s type. The specimens were preserved in 96% ethyl alcohol and then mounted on permanent slides to be identified. The palpal chela, leg I and leg IV of all the specimens of all stages were removed. The material is deposited at the Department of Zoology, Comenius University in Bratislava. The mean abundance is calculated as a fraction of total number of specimens obtained at the study plot and number of samples realised at the study plot. The species dominance is characterised by the scale proposed by Tischler (1949) and completed by Heydemann (1955): ED = eudominant, D = dominant, SD = subdominant, R = recedent, SR = subrecedent. The cluster analysis of the pseudoscorpion communities was realized using the computer program NCLAS (Podani, 1993). The clustering method complete linkage in combination with Sørensen’s index and Wishart’s similarity ratio was used. Species diversity was evaluated according to Odum (1977). All the couples of diversity values were compared with a t-test, to find out their significant difference (Poole, 1974). Principal Component Analysis (PCA) was performed to assess the relationship among the communities of pseudoscorpions at 6 study plots using the program Canoco (Ter Braak, Šmilauer, 1998). For each PCA a canonical ordination (RDA) with forward selection of environmental variables was used to find out environmental variables influencing the community. At all the study plots following environmental variables were measured (Range of values in gradient variables or categories of categorical ones are in brackets): pedological and chemical characteristics of leaf litter such as total organic carbon (3.80–14.40%); total nitrogen content (0.35–1.1%); content of exchange bases (Ca2+, Mg2+, Na+, K+) (12.60–49.40 mval/100g); total acidity (1.8–14.9 mval/100g); pH in H2O (3.90–6.74); humus content (6.55–24.83%); age of a forest stand (40–100 years); cover of E3: (65–80%); cover of E2 (1–50%); cover of E1 (40–100%); lying and standing dead wood (2–20%); forest fragmentation (0–50%), exposition of study plots (E, W, S, NE, NW, SW, SE), slope (0–45°); pollution by calcareous dust (present, absent).

Results and discussion A total of 949 specimens belonging to 10 taxa were examined at four study plots of the Malé Karpaty Mts and at two study plots of the Trnavská pahorkatina hills, during three years. Seven of them were identified on the species level. The maximum number of 78

species belonged to the family of Neobisiidae Neobisium erythrodactylum, N. muscorum and Roncus lubricus. Moreover we have recorded 2 species of Chthoniidae – Chthonius boldorii and Ch. fuscimanus. Pselaphochernes scorpioides and Chernes similis represented the family of Chernetidae (Table 1). We have recorded one tritonymph of Chernetidae at the study plot Lošonský háj grove, different from the tritonymphs of Chernes similis and Pselaphochernes scorpioides. We have found several different specimens of subgenus Chthonius (Ephippiochthonius) at five study plots (Table 1). According to morphological differences we suppose that these specimens belonged to two new species from subgenus Chthonius (E.), which have not been recorded in Slovak republic yet. It is necessary to elaborate a detailed description of these specimens. Neobisium muscorum was the only species common for all the study plots. This species belonged to the most widespread and common pseudoscorpions (Beier, 1963; Ducháč, 1997; Krumpál, Krumpálová, 2003). T a b l e 1. General survey of pseudoscorpion species found in the epigeon of oak-hornbeam forests in SW Slovakia. Study plots Taxon Chthoniidae Chthonius (Ephippiochthonius) boldorii B e i e r, 1934 Chthonius (Ephippiochthonius) fuscimanus S i m o n, 1900 Chthonius (Ephippiochthonius) sp. 1 Chthonius (Ephippiochthonius) sp. 2 Neobisiidae Neobisium (Neobisium) erythrodactylum (C. L. K o c h, 1873) Neobisium (Neobisium) muscorum (L e a c h, 1817) Roncus lubricus C. L. K o c h, 1873 Chernetidae Chernes similis C. L. K o c h, 1873 Pselaphochernes scorpioides (H e r m a n n, 1804) Chernetidae gen. sp.

Total number of specimens Mean abundance (ex.m-2)

LH

LL

HH

CA A D [%]

VI

LI

27 20.77 24 18.46 5 3.85 -

18 18.00 10 10.00 -

11 6.96 5 3.16 -

30 12.20 5 2.03

4 10.53

131 47.29 25 9.03

21 16.15 43 33.08 -

29 29.00 18 18.00 -

1 0.63 26 16.46 3 1.9

20 8.13 190 77.24 -

34 89.47 -

1 0.36 116 41.88 -

9 6.92 1 0.77

25 25.00 -

110 69.62 2 1.27 -

1 0.41 -

-

4 1.44 -

130 5.42

100 4.17

158 10.53

246 9.84

38 1.58

277 11.08

Note: LH – Lošonský háj grove, LL – Lošonec–lom quarry, HH – Horný háj grove, CA – Cajla, VI – Vinosady, LI – Lindava. D – dominance, A – abundance (number of specimens)

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Four species were eudominant at the study plot Lošonský háj grove – Neobisium erythrodactylum, N. muscorum, Chthonius boldorii and Ch. fuscimanus. We have also recorded a.dominant Pselaphochernes. scorpioides and subdominant Chthonius sp. (Table 1). The.material from Lošonec–lom quarry has included five eudominant species – Neobisium. erythrodactylum, N. muscorum, Chthonius boldorii, Ch. fuscimanus and Pselaphochernes scorpioides. Neobisium muscorum and Chernes similis were eudominant pseudoscorpions in Horný háj grove, with dominant Chthonius boldorii and subdominant Chthonius sp. 1. Neobisium muscorum and Ch. boldorii belonged to the eudominant pseudoscorpions in Cajla. Neobisium erythrodactylum may be classified as dominant and Chthonius sp. 2 as subdominant. Only two species were found at the study plot Vinosady, both of them as eudominant. Neobisium muscorum and Chthonius boldorii were eudominant and Chthonius sp. 2 was dominant in Lindava.

Fig. 1. Cluster analysis based on Sørensn’s index (Hierarchical classification, Complete linkage). For codes of study plots see Table 1.

The hierarchical cluster analysis based on a species identity (Sørensen’s index) has determined four diverse clusters in the communities (Fig. 1). Amongst the compared communities of pseudoscorpions a high similarity was recorded in Lošonský háj grove and Lošonec–lom quarry, however without statistically significant differences. The community in Horný háj grove was represented by the second cluster because the species Chernes similis and Roncus lubricus were found only there. The most similar 80

communities of false-scorpions were recorded in Cajla and Lindava and they were defined by the third cluster. Only two species were found at the study plot Vinosady and this fact has differentiated the community from the others. The hierarchical cluster analysis (Fig. 2) has separated four diverse clusters of the pseudoscorpion communities being based on the qualitative-quantitative similarity (Wishart’s index). The community in Vinosady appeared to be the most different from all others because of the lowest number of specimens and only two species present there (Table 1).

Fig. 2. Cluster analysis based on Wishart’s index (Hierarchical classification, Complete linkage). For codes of study plots see Table 1.

The PCA ordination diagram of the communities at 6 studied plots is shown on Fig..3. The species data were logarithmically transformed because of significant differences in species abundances at the study plots. The first two ordination axes account for 72.6% of the variance of the species data. Eigenvalue of the axis I is λ1 = 0,400, of the axis II λ2.=.0,326. The ordination diagram has confirmed the presence of four community types. The first community type refers to the 80–100 year old forests – Lošonský háj grove and Lošonec–lom quarry. They show low mean abundance, but the highest diversity and equitability (Table 2). The most eudominant species were found at these study plots. As all the recorded species in Lošonec–lom quarry were eudominant, we suggest that the calcareous dust has no significant influence on the occurrence of pseudoscorpions. The species Chthonius fuscimanus were found only at these two study plots. This species prefers leaf litter in moister biotopes (Beier, 1963). 81

Fig. 3. Ordination diagram based on Principal Component Analysis (PCA). First two axes of PCA as a biplot of taxa and study plots. For codes of study plots see Table 1. Codes of taxa: 1.– Chernes similis, 2 – Pselaphochernes scorpioides, 3 – Chthonius fuscimanus, 4 – Chthonius boldorii, 5 – Neobisium muscorum, 6 – Neobisium erythrodactylum, 7 – Roncus lubricus, 8.–.Chernetidae gen. sp., 9 – Chthonius sp. 1, 10 – Chthonius sp. 2. T a b l e 2. The comparison of the couples of diversity values of study plots by a t-test.

LH LL HH CA VI LI

e c H' 1.65 1.55 1.01 0.76 0.34 1.02

LH 0.846 0.219 1.646 0.000 1.452ns 6.230*** 10.708*** 10.838*** 9.340***

LL 0.965 0.221 1.553 212.891 0.000 5.810*** 11.051*** 10.707*** 10.056***

HH 0.517 0.518 1.006 256.140 204.048 0.000 2.277* 4.818*** 0.112ns

CA 0.473 0.618 0.762 365.483 341.438 309.269 0.000 3.403*** 3.432***

VI 0.485 0.812 0.336 58.211 46.105 94.184 66.543 0.000 5.897***

LI 0.632 0.407 1.017 275.510 332.237 226.582 427.354 49.172 0.000

Note: e - Pielou’s index of equitability, c – Simpson’s index of dominance, H’ – Shannon’s index of species diversity. T-test values are under the diagonal and degrees of freedom are above it. Significance levels: *** P < 0.001; ** 0.001 < P < 0.01; * 0.01 < P < 0.05; ns 0.05 < P (non-significant). For codes of study plots see Table 1.

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The second community type also referred to old oak-hornbeam forests (80–100 year old) at the study plots Cajla and Lindava. Higher mean abundance, lower diversity and smaller equitability were recorded there (Table 2). Neobisium muscorum and Chthonius boldorii were eudominant at both the plots. In these oak-hornbeam forests an identical species composition was recorded, but a higher number of specimens was found in Lindava than in Cajla. Horný háj grove is a younger oak-hornbeam forest (60–80 year old) with the highest humus content (16.55%). The pseudoscorpion community of this study plot had very similar mean abundance, diversity and equitability to that in Lindava. The comparison of the couples of diversity values of these two study plots by a t-test showed no statistically significant differences (Table 2). However, they differed in a species composition – Chernes similis and Roncus lubricus were found only in Horný háj grove. The fourth community type was the community of the younger (60–80 year old) oak-hornbeam forest Vinosady. This study plot was inhabited by the poorest and unstable community. The lowest species richness (in sensu Spellerberg, Fedor, 2003), the lowest mean abundance and diversity were as recorded there. The oak-hornbeam forest of this study plot was the most arid and had a very poor layer of the leaf litter. We may suggest that this was the reason for the occurrence of the lowest number of specimens belonging to only two taxa – Neobisium. muscorum and Chthonius. sp. 2. T a b l e 3. The measured environmental variables at all the study plots. Environmental variables / study plot Slope (in °) Age of a forest (years) NE exposition NW exposition S exposition W exposition SW exposition SE exposition E exposition Cover of E3 (%) Cover of E2 (%) Cover of E1 (%) Dead wood (%) Fragmentation (%) Calcareous dust (present, absent) pH in H2O Humus content (%) Total organic carbon (%) Total nitrogen content (%) Content of exchange bases - Ca, Mg, Na, K (mval/100g) Total acidity (mval/100g)

LH LL CA VI LI HH 0 5 1 10 7 7 80–100 80–100 80–100 60–80 80–100 60–80 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 80 65 75 70 65 80 1 25 2 30 1 10 75 100 90 80 80 75 10 3 5 12 3 5 15 5 20 30 0 15 0 1 0 0 0 0 4.29 6.74 4.83 4.48 4.68 5.00 12.76 6.55 11.21 13.79 14.48 16.55 7.40 3.80 6.50 8.00 8.40 9.60 0.60 0.35 0.48 0.67 0.63 0.73 15.40 22.60 18.40 16.00 22.60 31.80 13.70

1.80

9.10

11.50

12.50

9.50

For codes of study plots see Table 1.

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At all the study plots, 15 selected environmental variables were measured (Table 3). Unfortunately, none of the measured environmental variables showed a significant relationship with the organisms. This result supports our suggestion, that the calcareous dust in Lošonec–lom quarry had no influence on the occurrence of the pseudoscorpions.

Conclusion During the 3 year long research of the pseudoscorpion communities of the oak-hornbeam forests in SW Slovakia we recorded 949 pseudoscorpion specimens of 10 taxa. The species Neobisium muscorum occurred at each the study plot. According to the Sørensen’s and Wishart’s indices of similarity and PCA four community types of false-scorpions were formed. We suggest that the first community type (LL, LH) is the most stable, because of the highest diversity and equitability recorded there. The most unstable community represents the fourth community type (VI) with the lowest species richness and the lowest mean abundance, diversity and equitability. The results of canonical ordination showed that none of the 15 measured environmental variable did not havde the statistically significant influence on the occurrence of pseudoscorpions. Translated by J. Christophoryová Acknowledgement We would like to thank M. Holecová and P. Vďačný for assistance in the statistical evaluation of results. We are very grateful to P. Fedor for the revision of the English text and to M. Behuliak for review of the style of manuscript. The research was supported by the Slovak Grant Agency VEGA – No. 1/0119/03.

References Beier, M., 1963: Ordnung Pseudoscorpionidea (Afterskorpione). Bestimmungsbücher zur Bodenfauna Europas 1. Akademie-Verlag. Berlin, 313 pp. Ducháč, V., 1988: A contribution to the knowledge on the pseudoscorpions in České středohoří Mts (in Czech). Fauna Bohemiae Septentrionalis, 13, p. 103–108. Ducháč, V., 1993: Pseudoscorpions (Pseudoscorpionidea) in tree hollows in Třeboňsko (in Czech). Sbor. Jihočes. Muz. v Čes. Budějovicích, Prír. Vědy, 33, p. 65–69. Ducháč, V., 1997: Used occurrences of pseudoscorpions (Pseudoscorpiones) in Adršpašsko-teplické skály (in Czech). Vč. Sb. Přír.-Práce a Studie, 5, p.117–119. Gabbutt, P.D., Vachon, M., 1965: The external morphology and life history of the pseudoscorpion Neobisium muscorum. Proc. Zool. Soc. Lond., 145, p. 335–358. Gabbutt, P.D., 1969: Life-histories of some British pseudoscorpions inhabiting leaf litter. The Soil Ecosystem, Systematics Association Publication, 8, p. 229–235. Goddard, S.J., 1976: Population dynamics, distribution patterns and life cycles of Neobisium muscorum and Chthonius orthodactylus (Pseudoscorpiones: Arachnida). J. Zool., Lond., 178, p. 295–304. Heydemann, B., 1955: Die Frage der topographischen Überlinstimmung des Lebensraumes von Pflanzen- und Tiergesellschaften. Verh. Dtsch. Zool. Ges., Erlangen, p. 444–452.

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Krumpál, M., Krumpálová, Z., 2003: Pseudoscorpions - Pseudoscorpiones (in Slovak). In Mašán, P., Svatoň, J. (eds): Arachnids of the Poloniny NP (Arachnida: Araneae, Pseudoscorpiones, Opiliones, Acari-Parasitiformes). Štátna ochrana prírody SR Banská Bystrica a Správa Národného parku Poloniny Snina., Balada press Humenné, p. 115–126. Krumpálová, Z., Bartoš, D., 2002: Lycosid spiders (Araneae) of the oak forests of the Malé Karpaty Mts. near Modra. In Tajovský, K., Balík, V., Pižl, V. (eds): Studies on Soil Fauna in Central Europe, ISB AS CR, České Budějovice, p. 105–111. Krumpálová, Z., Szabová, S., 2003: Epigeic araneocoenoses of oak-hornbeam forest in the Nature Reserve Katarínka – Malé Karpaty Mts (in Slovak). Entomofauna Carpathica, 15, p. 49–55. Krumpálová, Z., Szabová, S., 2005: Spiders (Araneae) of oak-hornbeam forest – influenced by the human activities in Malé Karpaty Mts (in Slovak). Entomofauna Carpathica, 17, p. 55–60. Meyer, E., Wäger, H., Thaler, K., 1985: Struktur und jahreszeitliche Dynamik von Neobisium–Populationen in zwei Höhenstufen in Nordtirol (Österreich) (Arachnida: Pseudoscorpiones). Rev. Écol. Biol. Sol, 22, 2, p..221–232. Odum, E.P., 1977: The basis of ecology (in Czech). Academia. Praha, 987 pp. Országh, I., Országhová, Z., 2005: Structure of centipede communities (Antennata: Chilopoda) in oak-hornbeam forests of the Malé Karpaty Mts. and Trnavská pahorkatina hills (SW Slovakia). Ekológia, 24, Suppl. 2, p. 124–142. Podani, J., 1993: Syn-Tax. Version 5.0. Computer programs for Multivariate Data Analysis in Ecology and Systematics. User’s guide. Scientia Publishing, Budapest, 104 pp. Poole, R.W., 1974: An introduction to quantitative ecology. McGraw-Hill. New York, 532 pp. Spellerberg, I.F., Fedor, P.J., 2003: A tribute to Claude Shannon (1916-2001) and a plea for more rigorous use of species richness, species diversity and the "Shannon-Wiener" Index. Global Ecology and Biogeography, 12, 3, p. 177–179. Šťáhlavský, F., 2001: Pseudoscorpions (Arachnida: Pseudoscorpiones) of Prague (in Czech). Klapalekiana, 37, p. 73–121. Ter Braak, C.J.F., Šmilauer, P., 1998: CANOCO References Manual and User´s: Guide to Canoco for Windows: Software for Canonical Community Ordination (version 4). Microcomputer Power (Ithaca, NY, USA), 352 pp. Tischler, W., 1949: Grundzüge der terrestrischen Tierökologie. Friedrich Vieweg, Braunschweig, 219 pp. Verner, P.H., 1959: Study on arthropods living in soil of oak-hornbeam stands in Karlštejnsko (in Czech). Bohemia Ccentralis, A 1, 7, p. 345–408. Wood, P.A., Gabbutt, P.D., 1978: Seasonal vertical distribution of pseudoscorpions in beech litter. Bull. Br. Aarachnol. Soc., 4, p. 176–183. Zlinská, J., Šomšák, L., Holecová, M., 2005: Ecological characteristics of studied forest communities of an oak-hornbeam tier in SW Slovakia. Ekológia (Bratislava), 24, Suppl. 2, p. 3–19 Received 30. 7. 2005

Christophoryová J., Krumpál M.: Spoločenstvá šťúrikov dubovo-hrabových lesov Malých Karpát a.Trnavskej pahorkatiny. Počas rokov 2000–2002 sme uskutočnili výskum spoločenstiev šťúrikov na štyroch študijných plochách Malých Karpát – Lošonský háj, Lošonec–lom, Cajla, Vinosady – a na dvoch študijných plochách Trnavskej pahorkatiny – Horný háj a Lindava. Na zber pôdnej makrofauny sme použili kvadrátovú metódu kombinovanú s metódou presevu pôdnej hrabanky. Celkovo sme našli 949 šťúrikov patriacich do 10 taxónov, 7 z nich bolo určených na úroveň druhu. Druh Neobisium muscorum (L e a c h, 1817) bol jediným spoločným pre všetky študijné plochy. Klastrová metóda complete linkage v kombinácii so Sørensenovým indexom a Wishartovým indexom podobnosti a PCA potvrdili prítomnosť štyroch typov spoločenstiev, ktoré sa líšili druhovým zložením, dominanciou, ekvitabilitou, diverzitou a priemernou abundanciou šťúrikov. Za najviac stabilné považujeme spoločenstvá šťúrikov 80–100 ročných dubovo-hrabových lesov Lošonský háj a Lošonec–lom,

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pretože tu bola zaznamenaná najväčšia diverzita a ekvitabilita šťúrikov. Najmenej stabilné spoločenstvo predstavuje spoločenstvo šťúrikov mladšieho (60–80 rokov) dubovo-hrabového lesa Vinosady, ktorý bol najviac arídny a mal veľmi redukovanú vrstvu hrabanky. Zaznamenali sme tu najmenšie druhové bohatstvo, priemernú abundanciu a diverzitu šťúrikov. Na všetkých študijných plochách sme zmerali 15 environmentálnych premenných s cieľom nájsť tie premenné, ktoré ovplyvňujú spoločenstvá šťúrikov. Ani jedna z environmentálnych premenných nemala štatisticky významný vplyv na výskyt šťúrikov.

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Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 87–101

EPIGEIC SPIDERS (Araneae) IN ECOSYSTEMS OF OAK-HORNBEAM FORESTS IN THE MALÉ KARTATY MTS (SLOVAKIA) AND THEIR ECOLOGICAL CATEGORISATION ZUZANA KRUMPÁLOVÁ Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, 845 06 Bratislava, The Slovak Republic, e-mail: [email protected] Abstract Krumpálová Z.: Epigeic spiders (Araneae) in ecosystems of oak-hornbeam forests in the Malé Karpaty Mts (Slovakia) and their ecological categorisation. Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 87–101. 6,233 spider species were obtained in the oak-hornbeam forests at seven study sites in the Malé Karpaty Mts. The spider communities of the oak-hornbeam forests were rich in species diversity. Trochosa terricola, Pardosa lugubris and Tenuiphantes mengei may be considered as the eudominant species. CCA analysis has proved the influence of undergrowth on the spider communities. The nominal variable – phragmentation did not influence the assemblages. We did not notice any negative influence of the lime dust on the spiders, except for the impact on species diversity. Migration of the spiders from the surrounding was noticed. The migration effect of the non-specific species was noticed in the spider community in the young oak-hornbeam forest. Being based on the analysis of the spiders we have specified spiders into four categories - leaders, typical spiders, specific spiders and tourist species. Key words: epigeic spiders, communities, oak-hornbeam forests, Araneae, Malé Karpaty Mts, Slovakia

Introduction The Protected Landscape Area Malé Karpaty Mts belongs to the most deeply analysed areas of Slovakia from botanical as well as zoological point of view. Specialists paid attention to the investigation of spiders in the Malé Karpaty Mts, mainly in 80’s. The spiders of the oak forests in southern part of the Malé Karpaty Mts were studied by Žitňanská (1979), Gajdoš (1981) and Gajdoš, Krumpál (1986, 1988), actually, the research was carried out in the National Nature Reserve Devínska Kobyla near Bratislava. Gajdoš (1989) analysed the spiders from the oak forests near the village 87

Cerová-Lieskové. Gajdoš (1992) investigated spiders from the oak forest influenced by emissions from the nearby limestone-quarry near the village Rohožník. Krumpálová, Bartoš (2002) analysed the spider communities, especially lycosids, in four types of oak-hornbeam forests near the town Modra. Krumpálová, Szabová (2003) compared two spider communities of oak-hornbeam forests from the point of view of geographical exposition of the slopes and the age of forest stands at two study sides in the Malé Karpaty Mts, but these two parameters did not influence the communities. Krumpálová, Szabová (2005) analysed the spider communities of the oak-hornbeam forest near the village Lošonec, which was influenced by emissions from the nearby limestone quarry. Krumpálová (2005) compared the epigeic spider communities in natural and human impacted ecosystems in the Malé Karpaty Mts. Žitňanská (1973) analysed the araneofauna of the oak forest in the village Báb near Nitra. In the central part of the Malé Karpaty Mts, we carried out the research at four study sites – Cajla (CA) and Fúgelka (FU) the oak-hornbeam forests, Lošonec–lom quarry (LL) – the forest near a lime-stone quarry and Lošonský háj grove (LH). The spider communities of two study sites – Katarínka I. – the young forest (KI.) and Katarínka II – the old oak-hornbeam forest (KII.) represented the northern part of the Malé Karpaty Mts. The study site Lindava (LI) belongs to the unit of the Trnavská pahorkatina hills, however, this site adjoining the Malé Karpaty Mts. The more detailed data in the sites are included in the paper by Zlinská et al. (2005). This paper deals with the comparison, analysis and categorisation of the spider communities at seven study sites in oak-hornbeam forests in the Malé Karpaty Mts.

Methods The research was carried out at seven study sites in the period of 2000–2002. Spiders were collected by six pitfall formaldehyde traps set in a line. The traps were exposed from March to November and emptied monthly. The collection of material was interrupted for several times at the Lošonský háj grove due to human impact. We have used five categories of dominance according to the Tischler’s (1949) and Heydemann’s (1955) classifications. The complete linkage cluster analyses using PC-ORD (Mc Cune, Mefford, 1999) – were used to evaluate the spider communities. The assemblages were characterised by the Pielou’s index of equitability, Margaleff’s index of species richness, and Shannon’s index of diversity (Ludwig, Reynolds, 1988). Nomenclature of the spiders follows Platnick (2005). The spider communities were analysed according to the indirect gradient analysis (PCA) (Ter Braak, 2003). The first axis explains 59.3% of variable data, the second axis 26.2%. Cumulative variance of species data reached 96.4%. The spider species and their interactions to the environment was analysed in accordance with direct gradient analysis CCA. The gradient variables were important – the herbal and tree stratum (P-value of the Monte Carlo test was 0.0001) and the variable of the shrub stratum had lower P-value. These selected variables explained together 71.5% ((0.479/0.670)*100) of total variance of species data. The cumulative percentage of the species variance was 71.4%. The nominal variable represents phragmentation of the study sites.

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Results Epigeic spiders in the Malé Karpaty Mts During the three years of investigation (2000–2002) 6,233 spider individuals, belonging to 158 species and 24 families were obtained in the oak-hornbeam forests in the Malé Karpaty Mts (Table 1). T a b l e 1. Dominance of spiders on the study sites in the Malé Karpaty Mts. Study sites Taxon Atypidae Atypus piceus (S c h u l z e r, 1776) Segestriidae Segestria senoculata (L i n n a e u s, 1758) Dysderidae Dysdera erythrina (W a l c k e n a e r, 1802) Harpactea rubicunda (C. L. K o c h, 1838) Mimetidae Ero furcata (V i l l e r s, 1789) Theridiidae Enoplognatha oelandica (T h o r e l l, 1875) Enoplognatha ovata (C l e r c k, 1757) Enoplognatha thoracica (H a h n, 1833) Enoplognatha sp. Episinus angulatus (B l a c k w a l l, 1836) Euryopis flavomaculata (C.L. K o c h, 1836) Robertus lividus (B l a c k w a l l, 1836) Robertus sp. Steatoda albomaculata (D e G e e r, 1778) Steatoda bipunctata (L i n n a e u s, 1758) Steatoda phalerata (P a n z e r, 1801) Steatoda sp. Theridion tinctum (W a l c k e n a e r, 1802) Linyphiidae Abacoproeces saltuum (L. K o c h, 1872) Anguliphantes angulipalpis (W e s t r i n g, 1851) Asthenargus paganus (S i m o n, 1884) Bathyphantes nigrinus (W e s t r i n g, 1851) Centromerus arcanus (O. P. C a m b r i d g e, 1873) Centromerus brevivulvatus D a h l, 1912 Centromerus incilium (L. K o c h, 1881) Centromerus sellarius (S i m o n, 1884) Centromerus sylvaticus (B l a c k w a l l, 1841) Centromerus sp. Ceratinella brevis (W i d e r, 1834) Ceratinella major K u l c z y ń s k i, 1894 Ceratinela scabrosa (O. P. C a m b r i d g e, 1871) Diplocephalus latifrons (O. P. C a m b r i d g e, 1863)

CA FU LI LL LH KI. KII. D% D% D% D% D% D% D% 0.84 0.38 0.13 0.17

0.47 0.27 0.14

0.13 1.57 3.12 1.66 0.70 0.98 0.68 0.70 1.17 1.52 0.27 0.27

0.23 0.08

0.09 0.09 0.18

0.17 0.30 0.13 0.35 0.08 0.23

0.18 0.17 0.08 0.23 0.67 0.17 0.08 1.79 0.55 1.20 1.05 0.30 0.83 0.47 0.71 0.41 0.08 0.13 0.15 0.23 0.38 0.13 0.17 0.08 0.30 0.08 0.13 0.23 0.23 0.09 0.14 0.53 5.07 0.08 0.53 3.04 4.11 1.92 0.08 0.13 0.08 0.08 0.23 0.14 0.08 0.52 0.91 0.08 0.27 0.40 0.17 0.23 0.70 0.27 0.14 0.23 0.23 0.27 5.90 0.23 0.09 0.27 0.23 1.60 0.17 0.47 0.63 0.68 0.40 0.94 1.97 0.14

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T a b l e 1. (Continued) Study sites Taxon Diplocephalus picinus (B l a c k w a l l, 1841) Diplostyla concolor (W i d e r, 1834) Drapetisca socialis (S u n d e v a l l, 1833) Moebelia penicillata (W e s t r i n g, 1851) Lasiargus hirsutus (M e n g e, 1869) Lepthyphantes minutus (B l a c k w a l l, 1833) Linyphia hortensis S u n d e v a l l, 1830 Linyphia triangularis (C l e r c k, 1757) Linyphia sp. Macrargus rufus (W i d e r, 1834) Megalepthyphantes collinus (L. K o c h, 1872) Meioneta affinis (K u l c z y n s k i, 1898) Micrargus herbigradus (B l a c k w a l l, 1854) Micrargus sp. Microlinyphia pussila (S u n d e v a l l, 1830) Microneta varia (B l a c k w a l l, 1841) Minicia marginella (W i d e r, 1834) Neriene clatrata (S u n d e v a l l, 1830) Oedothorax retusus (W e s t r i n g, 1851) Palliduphantes alutacius (S i m o n, 1884) Palliduphantes pallidus (O. P. C a m b r i d g e, 1871) Panamomops fagei M i l l e r et K r a t o c h v í l, 1939 Pocadicnemis juncea L o c k e t et M i l l i d g e, 1953 Pocadicnemis pumila (B l a c k w a l l, 1841) Sintula corniger (B l a c k w a l l, 1856) Tapinocyba biscissa (O. P. C a m b r i d g e, 1872) Tapinocyba insecta (L. K o c h, 1869) Tenuiphantes mengei (K u l c z y ń s k i, 1887) Tenuiphantes tenbricola (W i d e r, 1834) Tenuiphantes flavipes (B l a c k w a l l, 1854) Tenuiphantes tenuis (B l a c k w a l l, 1852) Tenuiphantes zimmermani (B e r t k a u, 1890) Tenuiphantes sp. Tapinopa longidens (W i d e r, 1834) Thyreostenius biovatus (O. P. C a m b r i d g e, 1875) Trichoncus affinis K u l c z y n s k i, 1894 Walckenaeria antica (W i d e r, 1834) Walckenaeria atrotibialis (O. P. C a m b r i d g e, 1878) Walckenaeria corniculans (O. P. C a m b r i d g e, 1875) Walckenaeria cucullata (C. L. K o c h, 1836) Walckenaeria cuspidata B l a c k w a l l, 1833 Walckenaeria dysderoides (W i d e r, 1834) Walckenaeria furcillata (M e n g e, 1869) Walckenaeria incisa (O. P. C a m b r i d g e, 1871) Walckenaeria mitrata (M e n g e, 1868) Walckenaeria monoceros (W i d e r, 1834) Walckenaeria nudipalpis (W e s t r i n g, 1851) Walckenaeria obtusa B l a c k w a l l, 1836 Walckenaeria vigilax (B l a c k w a l l, 1853) Walckenaeria sp.

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CA FU LI LL LH D% D% D% D% D% 2.27 2.97 0.45 4.68 0.27 5.42 0.30 7.73 0.17 0.08

KI. KII. D% D% 0.63 3.29 1.61 6.03 0.09

0.18 0.08 0.23 0.38 0.98 0.41 0.53 0.52 0.15 0.94 0.71 0.14 0.40 0.38 0.08 0.08 0.14 0.09 0.93 1.05 0.46 0.45 1.41 0.71 1.23 0.08 0.09 0.54 0.09 0.52 0.15 0.15 1.17 0.18 0.23 0.40 0.52 0.08 0.30 0.00 0.09 0.27 6.54 3.32 3.20 2.80 3.75 4.83 6.99 0.14 0.30 0.17 0.47 0.18 0.27 0.13 0.87 0.08 0.91 0.70 0.27 0.14 0.67 1.22 0.68 0.15 0.94 0.63 0.55 16.96 7.69 1.29 7.87 17.56 13.76 18.36 0.13 3.85 0.38 0.61 1.17 0.18 0.82 0.53 0.52 0.68 0.68 1.22 0.68 0.91 0.23 0.63 0.55 0.13 0.45 0.27 0.17 1.14 1.06 0.23 0.27 1.23 0.23 0.09 0.23 0.09 0.14 0.53 1.57 0.15 0.83 0.63 0.14 0.13 0.38 0.70 0.63 0.41 0.08 0.09 0.13 1.92 0.23 1.06 0.47 0.09 0.27 0.17 0.08 0.08 0.76 0.36 0.27 0.27 0.70 0.30 0.30 0.70 0.89 0.14 0.08 0.38 0.70 0.30 0.45 0.36 0.08 0.40 0.35 0.38 2.86 0.27 0.15 0.23 0.94 0.27 0.27 0.09 0.46 0.61 0.27

T a b l e 1. (Continued) Study sites Taxon Tetragnathidae Pachygnatha degeeri S u n d e v a l l, 1830 Pachygnatha listeri S u n d e v a l l, 1830 Araneidae Araneus diadematus C l e r c k, 1757 Araneus quadratus C l e r c k, 1757 Araneus sp. Cercidia prominens (W e s t r i n g, 1851) Lycosidae Alopecosa aculeata (C l e r c k, 1757) Alopecosa pulverulenta (C l e r c k, 1757) Arctosa lutetiana (S i m o n, 1876) Pardosa agrestis (W e s t r i n g, 1861) Pardosa amentata (C l e r c k, 1757) Pardosa hortensis (T h o r e l l, 1872) Pardosa lugubris (W a l c k e n a e r, 1802) Pardosa monticola (C l e r c k, 1757) Pardosa sp. Trochosa ruricola (D e G e e r, 1778) Trochosa terricola T h o r e l l, 1856 Trochosa sp. Pisauridae Pisaura mirabilis (C l e r c k, 1757) Zoridae Zora nemoralis (B l a c k w a l l, 1861) Zora silvestris K u l c z y ń s k i, 1897 Zora spinimana (S u n d e v a l l, 1833) Agelenidae Agelena gracilens C. L. K o c h, 1841 Agelena sp. Histopona torpida (C. L. K o c h, 1837) Tegenaria campestris C. L. K o c h, 1834 Tegenaria ferruginea (P a n z e r, 1804) Tegenaria silvestris L. K o c h, 1872 Tegenaria sp. Textrix denticulata (O l i v i e r, 1789) Cybaeidae Cybaeus angustiarum L. K o c h, 1868 Hahnidae Hahnia helveola S i m o n, 1875 Hahnia ononidum S i m o n, 1875 Dictynidae Cicurina cicur (F a b r i c i u s, 1793) Dictyna uncinata T h o r e l l, 1856 Amaurobiidae Coelotes inermis (L. K o c h, 1855) Coelotes terrestris (W i d e r, 1834) Coelotes sp. Liocranidae Agroeca brunnea (B l a c k w a l l, 1833) Apostenus fuscus W e s t r i n g, 1851

CA FU LI LL LH KI. KII. D% D% D% D% D% D% D% 0.17 0.52 0.40 0.13

0.15 0.15

0.09

0.00 0.70 0.09 0.09 0.35 0.23 0.09 0.52 0.84 0.68 0.36 0.08 0.08

0.27

0.27 0.09 15.89 2.27 1.87 14.42

0.08 0.14 3.15 3.33 9.44 21.77 4.01 14.29 13.49 20.00 0.00 0.00 0.17 0.53 0.17 0.15 12.41 31.35 14.67 6.79 11.44 4.52 4.84

0.13

0.23 0.23

0.09

0.46 0.00 0.27 1.20 0.52 0.15 0.15 0.47 0.89 0.82 0.53 0.17 0.76 0.08 0.18 0.27 0.23 0.36 0.09 3.74 3.32 0.08 5.82 3.75 3.22 5.62 0.13 1.40 1.37 0.91 0.47 0.36 0.55 0.08 0.23 0.14 0.30 0.08 0.30 0.14 0.17

0.40

0.15

0.18

0.08 0.08

0.18

0.15 0.17 0.08 0.08

0.45 0.55

7.88 4.72 11.57 5.60 4.45 5.54 5.62 1.87 2.45 4.77 1.64 3.93 1.51 0.40 0.70 0.46 3.48 1.20 2.97 0.91 1.97 1.87 1.16 1.23 0.40 0.23 1.43 0.14

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T a b l e 1. (Continued) Study sites Taxon Scotina celans (B l a c k w a l l, 1841) Corinnidae Cetona laticeps (C a n e s t r i n i, 1868) Phrurolithus festivus (C. L. K o c h, 1835) Zodariidae Zodarion germanicum (C. L. K o c h, 1837) Clubionidae Clubiona caerulescens L. K o c h, 1867 Clubiona corticalis (W a l c k e n a e r, 1802) Clubiona comta C. L. K o c h, 1839 Clubiona terrestris W e s t r i n g, 1851 Clubiona sp. Gnaphosidae Drassyllus praeficus (L. K o c h, 1866) Drassyllus villicus (T h o r e l l, 1875) Haplodrassus silvestris (B l a c k w a l l, 1833) Micaria dives (L u c a s, 1846) Trachyzelotes pedestris (C. L. K o c h, 1837) Zelotes apricorum (L. K o c h, 1876) Zelotes aurantiacus M i l l e r, 1967 Zelotes erebeus (T h o r e l l, 1871) Zelotes subterraneus (C. L. K o c h, 1833) Zelotes sp. Philodromidae Philodromus dispar W a l c k e n a e r, 1826 Philodromus sp. Thanatus formicinus (C l e r c k, 1757) Thomisidae Diaea dorsata (F a b r i c i u s, 1777) Ozyptila praticola (C. L. K o c h, 1837) Ozyptila simplex (O. P. C a m b r i d g e, 1862) Xysticus cristatus (C l e r c k, 1757) Xysticus lanio C. L. K o c h, 1835 Xysticus luctator L. K o c h. 1870 Xysticus sp. Salticidae Ballus chalybeius (W a l c k e n a e r, 1802) Carrhotus xanthogramma (L a t r e i l l e , 1819) Euophrys frontalis (W a l c k e n a e r , 1802) Euophrys sp. Evarcha falcata (C l e r c k , 1757) Marpisa sp. Neon reticulatus (B l a c k w a l l , 1853) Number of individuals R1 H´ E

CA FU LI LL LH KI. KII. D% D% D% D% D% D% D% 1.47 1.45 1.17 0.89 0.82 0.14 0.17

0.15

1.74 1.57 2.59 1.36 3.28 2.06 1.23 0.23 0.09 0.14 0.08 0.09 0.13 0.15 0.08 0.09 0.55 0.13 0.35 0.08 0.47 1.52 1.64 0.08 0.40 0.52 0.53 0.61 0.47 0.27 0.82 0.93 2.80 1.67 3.25 0.47 0.98 0.82 0.08 1.05 0.08 0.15 0.18 0.08 0.08 0.00 0.52 0.08 0.08 0.47 0.27 0.08 0.08 0.23 0.08 0.13

0.08

0.09 0.00

0.15 0.08 0.23 0.40

0.54 0.41 0.17

0.30 0.15

0.47 0.27 0.14 2.80 3.50 4.03 3.93 2.58 1.79 3.15 0.13 0.53 0.76 0.47 0.36 0.14 0.13

0.08 0.08 0.30

0.09 0.09 0.09

0.13 0.35 0.08 0.13 749 572 1,314 1,322 9.7 9.8 9.2 14.1 3.0 3.4 2.5 3.5 0.7 0.8 0.6 0.8

0.09 0.27 427 1,119 730 9.4 12.7 9.7 3.2 3.4 3.0 0.8 0.7 0.7

Notes: CA – Cajla. FU – Fúgelka, LI – Lindava, LH – Lošonský háj grove, LL – Lošonec–lom quarry, KI. – Katarínka I., KII. – Katarínka II., R1 – richness index, E – evenness index, H´ – Shannon’s index of species diversity

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Cajla – 65 spider species were identified. Eudominant spiders include Tenuiphantes mengei, Pardosa lugubris and Trochosa terricola; Panamomops fagei and Coelotes inermis were dominant species. The spider community was stabilised and balanced. The epigeic araneocoenose responds with the stabile oak-hornbeam forests and habitat conditions as well. Fúgelka – 63 species were obtained at this study site. Trochosa terricola was eudominant; Anguliphantes angulipalpis, Diplostyla concolor, Tenuiphantes mengei and Pardosa lugubris may be classified as dominant species. The spider community was balanced. The epigeic spider community corresponds to the stabile oak-hornbeam forests and habitat conditions as well. Lošonec–lom quarry – 102 spider species were recorded at this study site. Eudominant species include Trochosa terricola; dominant spiders were represented by Tenuiphantes mengei, Histopona torpida and Coelotes inermis. The spider community was relatively stabilised, but the species spectrum was non-balanced and higher than at the other study sites. This rich epigeic araneocoenose corresponds to the habitat conditions with influence of lime dust (adjacent lime-stone pit). The dust affects mainly undergrowth, which may provide wider food offer at this study site. Two more facts influenced the spider community – presence of the brooklet and the surrounding corn field as well. Lošonský háj grove – 58 species were found there. Eudominant spiders were represented by Tenuiphantes mengei and Pardosa lugubris; dominant species included Diplostyla concolor and Trochosa terricola. The spider community was stabilised and balanced. The epigeic araneocoenose responds with the stabile habitat conditions in this oak-hornbeam forest as well. Lindava – 67 spider species were obtained at this study site, including eudominant Pardosa lugubris, Trochosa terricola and Coelotes inermis. The spider community was balanced and stabilised. The epigeic spider community corresponds to the stabile habitat conditions in this oak-hornbeam forest as well. Katarínka I. – 90 spider species were recorded there. Eudominant species were represented by Tenuiphantes mengei, Pardosa lugubris, Trochosa terricola; Coelotes inermis was dominant. The epigeic spider community was rich in species diversity that corresponds with diverse habitat conditions in this young forest. Katarínka II. – 65 species were obtained at this study site including eudominant Tenuiphantes mengei and Pardosa lugubris and dominant Diplostyla concolor, Panamomops fagei, Histopona torpida as well as Coelotes inermis. The spider community was balanced and stabilised. The epigeic spider community corresponds to the stabile habitat conditions in this oak-hornbeam forest as well. Considering to the eudominant and dominant species, Pardosa lugubris was the most dominant spider during the study period (14% on average); this spider achieved the highest dominance (22%) at the study site Lindava. Trochosa terricola was the second spider with high dominance in the communities (almost 14% on average), the highest value refers to Lindava (31%). The third dominant spider – Tenuiphantes mengei; (12% on average) achieved the highest dominance at the study site Katarínka II. (18%). The species representation in the spider community was different at the study sites. In general, the highest species richness was noticed in the forest near the limestone quarry – Lošonec–lom quarry (102 species) and at the young oak-hornbeam forest 93

Katarínka I.; relatively balanced and lower number of species was noticed at the other sites. The value of species diversity was high at all the study sites (Table. 1). The highest value of species diversity refers to the site of Lošonec–lom quarry (H´ = 3.5198). The lowest value corresponds with Lindava. The highest value of richness index was noticed in Lošonec–lom quarry (E = 14.053), as well as the above mentioned species diversity at this study site. The high values of evenness index were observed at the study sites – Lošonec–lom quarry, Lošonský háj grove and Fúgelka. Generally, the spider communities reached high values of species diversity and species richness (Table 1).

Fig. 1. Hierarchical cluster analysis (complete linkage) of the studied spider communities in the Malé Karpaty Mts according to Sørensen's index of similarity. Notes: CA – Cajla, FU – Fúgelka, LI – Lindava, LH – Lošonský háj, LL – Lošonec quarry, K1 – Katarínka I., K2 – Katarínka II.

The species structure was evaluated according to the Sørensen’s index (Fig. 1). High species similarity was noticed amongst the spider communities throughout the years. Hierarchical classification divided the spider communities into two groups and one separate line. The separate line represented the epigeic spider community at Lošonec– lom quarry, in which the species spectrum was the richest of all the study sites. The spider communities of Katarínka I., Lindava and Cajla forest represented the first group. The similarity between the study sites Lindava and Cajla was high in the compared communities. The non-balanced spider communities characterize this first group. The second group of the hierarchical classification represented the spider communities of the study sites Katarínka II., Fúgelka and Lošonský háj grove. The highest similarity refers to the study sites Katarínka II. and Lošonský háj grove. The second group characterized the structure of the stable spider community typical for the balanced oak-hornbeam forest. Differences in the spider communities were evaluated according to the Bray-Curtis similarity (Fig. 2) from qualitative-quantitative point of view. Hierarchical classification divided the spider communities into two groups. First group represents the spider communities at Fúgelka and Lošonec–lom quarry with eudominant Trochosa terricola. The araneocoenoses in Lindava (with a high dominance of Pardosa lugubris, Trochosa terricola and Coelotes inermis) joined these two communities. The second group is 94

divided into two branches. The spider communities of Katarínka I. and Cajla forest were characterised by the predominance of Pardosa lugubris, Trochosa terricola and Tenuiphantes mengei. Pardosa lugubris and Tenuiphantes mengei predominated in the spider communities at Katarínka II. and Lošonský háj grove.

Fig. 2. Hierarchical cluster analysis (complete linkage) of the studied spider communities in the Malé Karpaty Mts according to Bray-Curtis's index of similarity. Notes: CA – Cajla, FU – Fúgelka, LI – Lindava, LH – Lošonský háj, LL – Lošonec quarry, K1 – Katarínka I., K2 – Katarínka II.

Fig. 3. CCA analysis of the spider communities in seven study sites in the Malé Karpaty Mts and the environmental variables. Notes: E1 – herb stratum, E2 – shrub stratum, E3 – tree stratum. Study sites: CA – Cajla, FU – Fúgelka, LI – Lindava, LH – Lošonský háj, LL – Lošonec lom quarry, K1 – Katarínka I., K2 – Katarínka II.

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The direct gradient analysis (CCA) confirms the influence (or direct correlation) of undergrowth on the spider communities (Fig. 3). The dense herbal stratum (100%) and sparse tree stratum (65%) at the study sites Lošonec–lom quarry and Lindava caused a high trophic offer for phytophagous insects what leads to better nutrition for predators. This food chain caused the rich spider species spectrum and non-balanced communities. The denser undergrowth and canopy (80%) at the study sites Katarínka I., Katarínka II. and Fúgelka forest influenced positively the spider communities, which were balanced. The spider communities in Cajla and Lošonský háj grove were influenced particularly by balanced structure of undergrowth (75%) and canopy. The nominal variable – phragmentation did not influence the spider communities of the study sites. Indirect gradient analysis (PCA) confirms (Fig. 4) the isolation of the spider communities at the study site Lošonec–lom quarry lom and Lindava.

Fig. 4. PCA analysis of the spider communities in seven study sites in the Malé Karpaty Mts. Notes: CA – Cajla, FU – Fúgelka, LI – Lindava, LH – Lošonský háj, LL – Lošonec quarry, K1 – Katarínka I., K2 – Katarínka II.

General characterisation and categorisation of the epigeic spiders in the Malé Karpaty Mts Three spiders – Trochosa terricola, Pardosa lugubris and Tenuiphantes mengei mainly influenced the structure of the spider communities. These three spiders very markedly differentiated the communities. PCA analysis confirms (Fig. 5) the great influence of T. 96

mengei, Pardosa lugubris and Trochosa terricola in the epigeic spider communities, as well as the differences of these communities. Moreover, the species Coelotes inermis, C. terrestris, Dysdera erythrina, Haplodrassus silvestris, Zodarion germanicum, Xysticus luctator, Cercidia prominens, Zora silvestris, Ozyptila simplex, Histopona torpida, Anguliphantes angulipalpis, Panamomops fagei markedly influenced the spider communities in the Malé Karpaty Mts.

Fig. 5. PCA analysis of the species spectrum in the Malé Karpaty Mts.

According to the analysis of the spider communities we can classify spiders into four categories – leaders, typical spiders, specific spiders and tourist species. Leaders Trochosa terricola, Pardosa lugubris and Tenuiphantes mengei are characterised by the highest dominance in the communities and constant presence in the oak-hornbeam forests in the Malé Karpaty Mts. Typical spiders are characterised by a high dominance in the epigeic spider communities. Coexistence of these species is the ground for the spectrum of the spider 97

coenoses in the Malé Karpaty Mts – Anguliphantes angulipalpis, Panamomops fagei, Histopona torpida, Coelotes inermis, C. terrestris, Zodarion germanicum, Haplodrassus silvestris, Zora silvestris and Xysticus luctator. Each type of habitat determines certain number of species, but the similarity of the structure and coexistence of these species is limited. Specific spiders of the araneocoenoses display more or less the mild characteristics of the habitat. This mild characteristic might be the light conditions on the biotope, microhabitats, nourishment, or methods of collecting material. Specific spiders determine each habitat in the Malé Karpaty Mts – Atypus piceus (LI, LL), Drapetisca socialis (KI., FU), Tapinocyba biscissa (all the study sites), Walckenaeria mitrata (LL, KI., LI), Pachygnatha degeeri (LL, KI., FU), Agelena gracilens (KI., LH), Tegenaria ferruginea (KII., LH, LL), Agroeca brunnea (all the study sites), or Ozyptila simplex (LL, FU). Local variability of the species diversity predetermined the structure and productivity of the habitats. Specific spiders have low dominance in the spider communities; however, these spiders create their structure and characterise the habitats as well. Many of the specific spiders signify and specify the habitats or some changes in biotopes. Specific spiders could represent former populations, they can increase in the dominance and their position in the community might be changed. Tourist species often penetrate from the surrounding area. Most of these spiders have different requirements on moisture of habitats. In addition, we found a few tourist species in the Malé Karpaty Mts – Oedothorax retusus, Micaria dives, Pardosa agrestis, or P. monticola. These spiders are accidental species of the communities and only few individuals represent them. However, they could influence the structure of communities by their increase in the dominance. The increase in dominance quickly signals the changes of the habitat conditions.

Discussion The research of the epigeic spider communities in the Malé Karpaty Mts has brought the new information on the fauna and ecological knowledge on spiders. 6,233 spiders belonging to 158 species were collected at seven study sites. The spider communities were balanced and stabilised at the study sites – Katarínka II., Lindava, Cajla, Fúgelka and Lošonský háj grove. These epigeic spider communities correspond to the stable habitat conditions in the oak-hornbeam forest as well. Krumpálová, Szabová (2003, 2005) and Krumpálová, Bartoš (2002) evaluated the spider coenoses as balanced. The exposition of the slope does not seem to be an important factor in these oak-hornbeam forests. The research of the spider community in the oak-hornbeam forest in the adjoining limestone quarry has brought new knowledge on this specific habitat type (Krumpálová, Szabová, 2005). However, the surrounding maize field and brooklet have influenced the spider communities at the Lošonec–lom quarry as well. This influence is connected with 98

the rich herbal stratum that provided better conditions for phytophagous or saprophagous insects based on the high nutritional value. This represents an advantage for predators. The right influence of the lime dust on the spiders was did not confirm by (Krumpálová, Szabová, 2005; Gajdoš, 1992). A high number of subrecedent spiders signalise that many of them penetrate or migrate onto the study site for a short time. Their existence might be short-term, but in the special case they may persist for a longer time. These non-specific spiders (or alien species) can increase in the dominance and their position in the community might be changed. The epigeic spider community at the study site Katarínka I. was affected by various habitat conditions of this young oak-hornbeam forest. The higher value of species diversity reflects varied habitat conditions, diversified herb stratum and hence high nutritional value. In the comparison with pseudoscorpions, the community in the younger oak-hornbeam forests (Christophoryová, Krumpál, 2005) represented the unstable type. The spider communities of the oak-hornbeam forests are rich in the species diversity. Trochosa terricola, Pardosa lugubris and Tenuiphantes mengei may be classified as eudominant. These spiders are characterised by the highest dominance in the communities and constant presence in the oak-hornbeam forests in the Malé Karpaty Mts. PCA analysis confirms the great influence of T. mengei, Pardosa lugubris and Trochosa terricola in the epigeic spider communities, as well as the differences of these communities. The oscillation of the abundance of dominant spiders seems to be the natural fluctuation of certain species in the araneocoenoses. The high dominance of these spiders was noticed in the oak-hornbeam forests in the Malé Karpaty Mts according to Gajdoš (1992) and Krumpálová, Bartoš (2002). The other specialists use different methods (sweeping or quadrate method) to collect spiders. In the oak-hornbeam forests of the southern part of the Malé Karpaty Mts – Devínska Kobyla using the sweeping method Linyphia hortensis, L..triangularis, Mangora.acalypha and Mettelina segmentata were the most dominant (Gajdoš, 1981). Spiders were collected by sweeping method in the oak-hornbeam forest near the city Nitra and Mangora acalypha, Anyphena accentuata and Theridium ovatum dominated in the community (Žitňanská, 1973). The value of species diversity was high at all the study sites. The highest value of species diversity was noticed at the study site Lošonec–lom quarry near the lime stone-pit. High species similarity was found in the comparison throughout the years. The epigeic spider community at Lošonec–lom quarry near the lime stone-pit was different from the other compared spider coenoses as in the papers of the other authors (Krumpálová, Szabová, 2005; Gajdoš, 1981). CCA analysis confirms the influence of undergrowth on the spider communities. The nominal variable – phragmentation did not influence the spider communities. In the contrary to spiders, any environmental variable did not have statistically significant influence on the occurrence of pseudoscorpions (Christophoryová, Krumpál, 2005). Indirect gradient PCA analysis confirms the isolation of the spider communities at the study site Lošonec–lom quarry and Lindava forest. 99

Finally, we can summarise the oak-hornbeam forest near the limestone quarry create the ecological conditions for epigeic araneocoenoses well. We did not notice any negative influence of the lime dust on the spiders, except for high species diversity and immigration of the spiders from the surrounding area. This effect was noticed in the spider community of the young oak-hornbeam forest, as well. Translated by the author Acknowledgement I would like to express my thanks to I. Országh and M. Holecová for their professional consulting on this subject. This work was supported by the Slovak Grant Agency (VEGA, No. 1/0119/03).

References Christophoryová, J., Krumpál, M., 2005: Communities of pseudoscorpions (Pseudoscorpiones, Arachnida) in epigaeon of oak-hornbeam forests in the Malé Karpaty Mts. and Trnavská pahorkatina hills (SW Slovakia). Ekológia (Bratislava), 24, Suppl. 2, p. 76–86. Gajdoš, P., 1981: Arachnocoenoses (Araneae) of the Zobor and Devínska Kobyla hills (in Slovak). Thesis, Faculty of Natural Sciences, Bratislava, 95 pp. Gajdoš, P., Krumpál, M., 1986: Production conditions of spider community (Araneidea) of the vegetation cover of the Zobor and Devínska Kobyla hills - I. Zobor (in Slovak). Biológia, Bratislava, 41, p. 981–991. Gajdoš, P., Krumpál, M., 1988: Production conditions of spider community (Araneidea) of the vegetation cover of the Zobor and Devínska Kobyla hills - II. Devínska Kobyla (in Slovak). Biológia, Bratislava, 43, p..105–114. Gajdoš, P., 1989: Contribution to the knowledge on spiders (Araneae) near Cerová (Malé Karpaty Mts) (in Slovak). Prehľad odbor. prác TOP, Bratislava, 5, p. 207–211. Gajdoš, P., 1992: Arachnocoenoses of soil- and tree photoeclectors in the forest ecosystems in the Malé Karpaty Mts. from a point of view the lime stone pit near Rohožník (in Slovak). Správy Slov. Entomol. Spol. SAV, Bratislava, 4, p. 18–26. Heydemann, B., 1955: Die Frage der topographischen Übereinstimmung des Lebensraumes von Pflanzen- und Tiergesellschaften. Verh. Dtsch. Zool. Ges., Erlangen, p. 444–452. Krumpálová, Z., Bartoš, D., 2002: Lycosid spiders (Araneae) of the oak forests of the Malé Karpaty Mts. near Modra. In Tajovský, K., Balík, V., Pižl, V. (eds): Studies on Soil Fauna in Central Europe, ISB AS CR, České Budějovice, p. 105–111. Krumpálová, Z., Szabová, S., 2003: Epigeic araneocoenoses of oak-hornbeam forest in the Nature Reserve Katarínka – Malé Karpaty Mts (in Slovak). Entomofauna Carpathica, Bratislava, 15, p. 49–55. Krumpálová, Z., 2005: Epigeic spider communities (Araneae) in the natural and human impacted ecosystems in the Malé Karpaty Mts. Abstract Book, 8-th Central European Workshop on Soil Zoology, České Budějovice, p. 41. Krumpálová, Z., Szabová, S., 2005: Spiders (Araneae) of oak-hornbeam forest – influenced by the human activities in Malé Karpaty Mts (in Slovak). Entomofauna Carpathica, Bratislava, 17, p. 55–60. Ludwig, J. A., Reynolds, J. F., 1988: Statistical ecology: a primer of methods and computing. Wiley-Interscience, New York, 337 pp. Mc Cune, J., Mefford, P., 1999: PC-ORD-Multivariate Analysis of Ecological Data. Version 4.0. MjM Softwere design, Gleneden Beach, Oregon, USA, 237 pp. Platnick, N. I., 2005: The world spider catalogue, version 5.0. American Museum of Natural History, online at http://research.amnh.org/entomology/spiders/catalog/index.html.

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Ter Braak, C. J. F., 2003: Program CANOCO, version 4.2. Ordination. In Jongman, R.H.G., Ter Braak, C.J.F., Van Tongeren, O.F.R. (eds): Data analysis in community and landscape ecology. Cambridge University Press, Cambridge, p. 91–173. Tischler, W., 1949: Grundzüge der terrestrischen Tierökologie. Fridrich Vieweg, Braunschweig, 219 pp. Zlinská, J., Šomšák, L., Holecová, M., 2005: Ecological characteristic of studied forest communities of an oak-hornbeam tier in SW Slovakia. Ekológia (Bratislava), 24, Suppl. 2, p. 3–19. Žitňanská, O., 1973: Spinnen des Ökosystems in Báb bei Nitra. Acta Fac. Rer. Nat. Univ. Comen., Zool., 18, p. 31–45. Žitňanská, O., 1979: Arachnofauna of Devínska Kobyla (in Slovak). Final Report, depon. in PriF UK, Bratislava, p. 52–87. Received 30. 7. 2005

Krumpálová Z.: Epigeické pavúky (Araneae) v ekosystémoch dubovo-hrabových lesov Malých Karpát (Slovensko) a ich ekologická kategorizácia. Práca je zameraná na hodnotenie epigeických spoločenstiev pavúkov v ekosystéme dubovo-hrabových lesov Malých Karpát. Pavúky boli odchytávané v rokoch 2000–2002 metódou zemných pascí. Celkovo sme získali 6 233 pavúkov na siedmych študijných plochách. Spoločenstvá pavúkov dubovo-hrabových lesov mali vysokú druhovú diverzitu. Trochosa terricola, Pardosa lugubris a Tenuiphantes mengei mali eudominantné zastúpenie na všetkých sledovaných plochách. CCA analýza potvrdila preukazný vplyv bylinného porastu na spoločenstvá pavúkov. Nominálna premenná – fragmentácia neovplyvnila sledované spoločenstvá pavúkov. Nezaznamenali sme negatívne pôsobenie prachového odpadu z kameňolomu na epigeické pavúky. Zaznamenali sme migráciu pavúkov zo susediacich habitátov, najmä z agrocenóz. Migračný efekt nešpecifických druhov bol zistený aj v spoločenstve pavúkov v mladom dubovo-hrabovom lese. Na základe analýzy zistených druhov epigeických pavúkov a dát z literatúry sme ich rozdelili do štyroch kategórií – charakteristické druhy (lídre), typické druhy, špecifické druhy a zatúlané druhy.

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Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 102–112

SOIL MITES (Acari, Mesostigmata) OF OAK FORESTS IN THE MALÉ KARPATY MTS (W SLOVAKIA) PETER FENĎA, JANA CICEKOVÁ Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská dolina B–1, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected] Abstract Fenďa P., Ciceková J.: Soil mites (Acari, Mesostigmata) of oak forests in the Malé Karpaty Mts (W Slovakia). Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 102–112. During the years 1999–2002 mesostigmatid mites, as important part of soil mesofauna communities, were studied in two oak-hornbeam forests in the Malé Karpaty Mts. 75 species from 4 cohors (Antennophorina, Gamasina, Sejina, and Uropodina) were recorded in these sites. The mite communities in the study site Lošonec–lom quarry regularly impacted by limestone dust from the nearby quarry show greater fluctuations in species richness and abundance of mites during the year. In the study site Lošonský háj grove Nature Reserve there are evident influence of more stable microclimatic conditions in the forest. The comparison of mite communities in consecutive years through hierarchical clustering suggests, that the temperature and air humidity can have stronger influence to mesostigmatid mite assemblages than the type of soil, pH and vegetation. From the faunistical point of view, Pergamasus canestrinii was recorded for the first time in Slovakia. Key words: Acari, mites, soil, oak-hornbeam forests, Malé Karpaty Mts, Slovakia

Introduction The majority of mesostigmatid mites are free-living predators inhabiting the layers of forest litter, where especially speedy predators find the optimal conditions. They hunt in the pore system of the upper layer of the soil (Karg, 1993). Another species play role in the decomposition processes in leaf litter, and a number of species are endoparasites or ectoparasites of mammals, birds, reptiles or insects. The Malé Karpaty (Little Carpathians) Mts form the south–western projection of the Carpathian arch. The eastern and southern slopes of the Malé Karpaty Mts are influenced by the thermophilous Pannonian District. The area is covered by forests, woody steppes and rocky habitats. Forest stands have been under long–term anthropogenic pressure in the cultural land. 102

The mesostigmatid mites of the Malé Karpaty Mts are partially known from fur of small mammals and their nests (Ambros, 1984; Mašán et al., 1994), from birds’ nests (Nosek, Lichard, 1962; Ambros et al., 1992), and also phoretic mites on insects (Mašán, 1999, 2001b; Mašán, Kalúz, 2001). But there is only one paper dealing with soil mites (Kalúz, 2005). Additionally in monographs of cohors Uropodina (Mašán, 2001a), family Macrochelidae (Mašán, 2003), family Zerconidae (Mašán, Fenďa, 2004) and family Ascidae (Kalúz, Fenďa, 2005) were summarized all known data from the Malé Karpaty Mts together with the new findings.

Study area Study sites are situated in the zone of oak-hornbeam forests on southeastern slopes of the Malé Karpaty Mts. Lošonský háj grove Nature Reserve (LH) [48°28' N, 17°24' E, Grid Reference Number of the Databank of the Fauna of Slovakia 7570b] – 80 to 100 year old oak-hornbeam forest (Querco–Carpinetum caricetosum pilosae) at the altitude of 260 m. Under the leaf litter the soil has a lumpy texture being humid in a dry year period too. Soil horizon pH = 4.29, humus content 12.67%. Lošonec–lom quarry (LL) [48°29' N, 17°23' E, 7570a] – 80 to 100 year old oak-hornbeam forest (Querco–Carpinetum melicetosum uniflorae) at the altitude of 340 m. The site is regularly impacted by limestone dust imissions close to the quarry. Soil horizon pH = 6.74, humus content 6.55%. The more detailed pedological and botanical characteristics of the study sites as well as the climatic conditions in the period of 1999–2002 have been included in the paper by Zlinská et al. (2005).

Methods and material The study sites were investigated in the period 1999–2002. At each study plot, at about 1–month intervals from April to November, the material was collected by sifting from the leaf litter and upper part of soil from 1.m2 (16 squares of 25×25 cm). The sifted material was separated using xereclectors of the Moczarski – Winkler’s type (Balogh, 1958), and the animals were deposited in 75% ethyl alcohol. The mites were processed to yield microscopic preparations using chloralhydrate medium Liquid de Swan. The material has been deposited at the Department of Zoology, Faculty of Natural Sciences, Comenius University in Bratislava. The species dominance (in %) was characterized by the scale proposed by Tischler (1949) and the species constancy (in %) was expressed by categories in the sense of Schwerdtfeger (1975). The species abundance relationship after Ludwig and Reynolds (1988) was used to compare the structure of the mite communities [N0 – number of species, N1 – eH' (H' is Shannon’s index), N2 – 1/λ (λ is Simpson’s index), E1 – Pielou evenness index]. In cluster analysis was used Wishart’s index (Podani, 1988).

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Results A total of 8,030 gamasid mites were extracted. Analysis of the material revealed 75 nominated species belonging to 4 cohors (Antennophorina, Gamasina, Sejina, Uropodina) (Table 1). Altogether 64 mite species occurred in LL site and 54 species in LH site. On the other hand, higher abundance was recognized in LH site (Table 1). T a b l e 1. Survey of mesostigmatid mites in study sites during years 1999–2002. Study sites MITES cohors Antennophorina family Celaenopsidae Celaenopsis badius (C. L. K o c h, 1841) cohors Gamasina family Ascidae Aceoseius muricatus (C. L. K o c h, 1839) Arctoseius eremitus (B e r l e s e, 1918) Arctoseius venustulus (B e r l e s e, 1916) Leioseius bicolor (B e r l e s e, 1918) Leioseius minusculus (B e r l e s e, 1905) Zerconopsis remiger (K r a m e r, 1876) family Eviphididae Eviphis ostrinus (C. L. K o c h, 1836) family Laelapidae Eulaelaps stabularis (C. L. K o c h, 1836) Haemogamasus nidi M i c h a e l, 1892 Hypoaspis aculeifer G. C a n e s t r i n i, 1884 Hypoaspis brevipilis H i r s c h m a n n, 1969 Hypoaspis imitata R e i t b l a t, 1963 Hypoaspis oblonga (H a l b e r t, 1915) Hypoaspis praesternalis W i l l m a n n, 1949 Hypoaspis vacua (M i c h a e l, 1891) Hypoaspis spp. family Macrochelidae Geholaspis longispinosus (K r a m e r, 1876) Geholaspis mandibularis (B e r l e s e, 1904) Macrocheles glaber (J. M ü l l e r, 1859) Macrocheles montanus W i l l m a n n, 1951 family Macronyssidae Ornithonyssus sylviarum (C a n e s t r i n i et F a n z a g o, 1877) family Pachylaelapidae Olopachys suecicus S e l l n i c k, 1950 Pachylaelaps magnus (H a l b e r t, 1915) Pachylaelaps resinae K a r g, 1971 Pachylaelaps spp. Pachyseius humeralis B e r l e s e, 1910 family Parasitidae Amblygamasus sp. Eugamasus monticolus B e r l e s e, 1905

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Lošonský Lošonec– háj grove lom quarry N PS N PS 5

4

7

6

1 1 20

1 1 9

4 1 1 1 17

3 1 1 1 12

-

-

35

15

2 1 1 4 10 3

1 1 1 2 7 3

1 6 2 6 1 4 1 33 8

1 1 1 5 1 2 1 15 3

255 2 3 90

32 2 3 23

159 1 4 115

29 1 2 26

1

1

-

-

38 1 23 10 4

15 1 6 8 2

61 1 8 52 23

15 1 6 14 9

33 -

7 -

152 2

27 2

T a b l e 1. (Continued) Study sites MITES Holoparasitus calcaratus (C. L. K o c h, 1839) Holoparasitus sp. Leptogamasus succineus W i t a l i ń s k i, 1973 Leptogamasus spp. Parasitus fimetorum (B e r l e s e, 1903) Pergamasus barbarus B e r l e s e, 1904 Pergamasus brevicornis (B e r l e s e, 1903) Pergamasus canestrinii (B e r l e s e, 1884) Pergamasus crassipes (L i n n a e u s, 1758) sensu B e r l e s e, 1906 Pergamasus mediocris (B e r l e s e, 1904) Pergamasus ruhmi W i l l m a n n, 1938 Poecilochirus carabi G. et R. C a n e s t r i n i, 1882 Porrhostaspis lunulata J. M ü l l e r, 1869 Vulgarogamasus kraepelini (B e r l e s e, 1904) Vulgarogamasus remberti (O u d e m a n s, 1912) family Rhodacaridae Cyrtolaelaps chiropterae K a r g, 1971 Cyrtolaelaps mucronatus (G. et R. C a n e s t r i n i, 1881) Rhodacarus spp. Sessiluncus hungaricus K a r g, 1964 Stylochirus fimetarius (J. M ü l l e r, 1859) sensu M a š á n et K a l ú z, 2001 family Veigaiaidae Veigaia cerva (K r a m e r, 1876) Veigaia exigua (B e r l e s e, 1917) Veigaia kochi (T r ä g å r d h, 1901) Veigaia nemorensis (C. L. K o c h, 1839) Veigaia transisalae (O u d e m a n s, 1902) family Zerconidae Prozercon carpathofimbriatus M a š á n et F e n ď a, 2004 Prozercon tragardhi (H a l b e r t, 1923) Zercon curiosus T r ä g å r d h, 1910 Zercon hungaricus S e l l n i c k, 1958 Zercon peltatus var. peltatus C. L. K o c h, 1836 Zercon vacuus C. L. K o c h, 1839 cohors Sejina family Sejidae Sejus togatus C. L. K o c h, 1836 family Uropodellidae Asternolaelaps sp. cohors Uropodina family Polyaspidae Polyaspinus schweizeri (H u ţ u, 1976) Polyaspis patavinus B e r l e s e, 1881 family Trachytidae Trachytes aegrota (C. L. K o c h, 1841) Trachytes baloghi H i r s c h m a n n et Z i r n g i e b l–N i c o l, 1969 family Trematuridae Trichouropoda elegans (K r a m e r, 1882) Trichouropoda karawaiewi (B e r l e s e, 1904)

Lošonský Lošonec– háj grove lom quarry N PS N PS 420 33 514 34 1 1 3 3 13 4 253 30 293 25 1 1 367 30 74 22 13 5 4 2 21 1 1 1 40 9 74 21 34 12 2 1 26 8 9 2 70 23 1 1 432 31 650 31 8 1 2 1 26 -

2 -

1 3 2 1

1 2 1 1

21 16 2 522 -

8 1 2 29 -

31 508 5

12 29 1

86 5 120 455 3

5 5 20 31 2

5 1 12 262 27 10

3 1 4 29 14 5

20

8

81

21

-

-

2

2

8 1

7 1

3 13

2 9

99 3

6 3

9 7

7 4

1

1

3 -

3 -

105

T a b l e 1. (Continued) Study sites MITES Trichouropoda obscurasimilis H i r s c h m a n n et Z i r n g i e b l–N i c o l, 1961 Trichouropoda orbicularis (C. L. K o c h, 1839) Trichouropoda ovalis (C. L. K o c h, 1839) Trichouropoda penicillata H i r s c h m a n n et Z i r n g i e b l–N i c o l, 1961 family Urodinychidae Dinychus bincheaecarinatus H i r s c h m a n n, W a g r o w s k a–A d a m c z y k et Z i r n g i e b l–N i c o l, 1984 Dinychus perforatus K r a m e r, 1886 Urodiaspis tecta (K r a m e r, 1876) Uroobovella pulchella (B e r l e s e, 1904) family Uropodidae Uropoda misella (B e r l e s e, 1916) Uropoda orbicularis (O. F. M ü l l e r, 1776) Uropoda splendida K r a m e r, 1882 Total individuals / samples H’ N0 N1 N2 E1

Lošonský Lošonec– háj grove lom quarry N PS N PS 49 17 9 7 3 1 336 26 277 29 4 2 9

2

3

3

2 92 -

1 14 -

21 19 1

7 9 1

4 4 267 4325 2.92 54 1.07 1.37 0.72

1 2 28 36

16 8 3705 36 2.84 64 1.04 10.87 0.67

Notes: N – number of specimens, PS – number of positive samples, N0 – number of species, N1 – eH' (H' is Shannon’s index), N2 – 1/λ (λ is Simpson’s index), E1 – Pielou’s evenness index

In addition, in the site Lošonec–lom quarry there were also taken soil samples of wet soil from the bank of the creek inside the forest. Only in these samples were detected species Cheiroseius longipes (W i l l m a n n, 1951), Ch. salicorniae (W i l l m a n n, 1949) and Plesiosejus major (H a l b e r t, 1923), all hygrophilous species from the family Ascidae, Macrocheles carpathicus M a š á n, 2003 and M. tardus (C. L. K o c h, 1841) sensu H y a t t et E m b e r s o n, 1988 from the family Macrochelidae. There was recorded also high number of larval and nymphal stages of big long–legged predators (especially Pergamasus barbarus), that were not recorded by sifting in the drier parts of forest. In the forest soil the eudominant species were Holoparasitus calcaratus (LL 13.9), Veigaia nemorensis (LL 13.7, LH 12.1), Vulgarogamasus kraepelini (LL 17.5), and Zercon peltatus var. peltatus (LH 10.5); the dominant species were Trichouropoda ovalis (LL 7.5, LH 7.8), Zercon hungaricus (LL 7.1), Geholaspis longispinosus (LH 5.9), Holoparasitus calcaratus (LH 9.7), Pergamasus barbarus (LH 8.5), Uropoda splendida (LH 6.2), and Vulgarogamasus kraepelini (LH 9.9). The euconstant species were Geholaspis longispinosus (LL 80.5, LH 88.9), Holoparasitus calcaratus (LL 94.4, LH 91.7), Trichouropoda ovalis (LL 80.5), Veigaia nemorensis (LL 80.5, LH 80.6), Vulgarogamasus kraepelini (LL 86.1, LH 86.1), Zercon hungaricus (LL 80.5), Pergamasus barbarus (LH 83.3), Uropoda splendida (LH 77.8), and Zercon peltatus var. peltatus (LH 86.1); the constant species were Sejus togatus (LL 58.3), Pergamasus barbarus (LL 61.1), Macrocheles montanus (LL 72.2, LH 63.9), Pergamasus mediocris 106

(LH 58.3), Porrhostaspis lunulata (LH 63.9), Trichouropoda ovalis (LH 72.2), and Zercon hungaricus (LH 55.6). All these species are widely distributed in Slovakia.

40

LL LH

Number of species

30

20

10

0 III

IV

V

VI

VII

VIII

IX

X

XI

XII

Month

Fig. 1. Species richness of mites in both study sites during years 1999–2002. Abbreviations of study sites see in chapter “Study area”. Note: Lošonský háj LH, Lošonec–lom LL. 800

LL LH

Abundance

600

400

200

0 III

IV

V

VI

VII

VIII

IX

X

XI

XII

Month

Fig. 2. Abundance of mites in both study sites during years 1999–2002. Abbreviations of study sites see in chapter “Study area”. Note: Lošonský háj LH, Lošonec–lom LL.

107

The abundant species have two patterns of seasonal dynamics. The first pattern has first maximum in spring (April–May) and the second one in summer months (July–August) in LH site and only one maximum in summer (August) in LL site (e.g. species Geholaspis longispinosus, Holoparasitus calcaratus, Pergamasus barbarus, and Veigaia nemorensis). The second pattern has only one spring maximum (April–May) in LH site, and one summer maximum (July–August) in LL site (e.g. species Macrocheles montanus, Vulgarogamasus kraepelini) (Fig. 3). Although the species richness was higher in the LL site, the more balanced community occurred in the LH site (E1 = 0.717, Table 1). That fact is confirmed also by differences of species richness during the years 1999–2002 (Fig. 1) and also by differences of abundance (Fig. 2). The mite communities in the study site Lošonec–lom quarry regularly impacted by limestone dust from the nearby quarry show greater fluctuations in species richness and abundance of mites during the year. In the study site Lošonský háj grove Nature Reserve there is an evident influence of more stable microclimatic conditions in the forest (with maximum abundance of mites in August). 160

LL LH

Abundance

120

80

40

0 III

IV

V

VI

VII

VIII

IX

X

XI

XII

Months

Fig. 3. Abundance of species Vulgarogamasus kraepelini in both study sites during years 1999–2002. Abbreviations of study sites see in chapter “Study area”. Note: Lošonský háj LH, Lošonec–lom LL.

The species Eviphis ostrinus occurred only in Lošonec–lom quarry (dominance 0.9, constancy 41.7). Also species Asternolaelaps sp., Eugamasus monticolus, Leptogamasus succineus, Trichouropoda elegans, T. penicillata, and Veigaia transisalae were recorded only in LL site. Some other species occurred in both sites, but they evidently preferred the conditions in Lošonec–lom quarry: Dinychus perforatus, Hypoaspis vacua, Pachyseius humeralis, Polyaspis patavinus, and Sejus togatus. It is possible, that the above-mentioned species prefer the neutral value (pH = 6.74) of the upper soil horizon in the study site Lošonec–lom quarry, in comparison with pH = 4.29 in Lošonský háj grove Nature Reserve (Zlinská et al., 2005). 108

On the other hand, some species preferred the LH site: Polyaspinus schweizeri, Porrhostaspis lunulata (LH dominance 1.6, constancy 63.9 vs. LL dominance 0.03, constancy 2.8), Trichouropoda obscurasimilis (LH dominance 1.1, constancy 47.2 vs. LL dominance 0.2, constancy 16.4), Veigaia exigua, and V. kochi. The comparison of mite communities in consecutive years through hierarchical clustering (Fig. 4) shows an interesting fact, that the most important factor to form species composition and abundance of soil Mesostigmata seems to be the climatic conditions. The most similarities are between different study sites in the same year (Fig. 4), not between the same study sites. It appears from this that the temperature and air humidity have greater influence on mesostigmatid mite assemblages than the type of soil and vegetation (Zlinská et al., 2005).

Fig. 4. Dendrogram of mite communities in study sites (Wishart’s index). Abbreviations of study sites in chapter “Study area“.

Discussion In Central Europe there are not comparable data from oak or oak-hornbeam forests. The data about human impact on soil mites are known mainly from Poland (Seniczak et al., 1996, 1997, 2002). The concentration of phosphorus and fluorine generally reduced the density of mites and species number of Oribatida and Gamasida (Seniczak et al., 1996), similarly a high concentration of heavy metals greatly reduced the density and species richness of mites. A high concentration of nitrogen pollution reduced slightly the density of mites in all soil horizons (Seniczak et al., 2002). We do not observe shining differences between the study site Lošonec–lom quarry, regularly impacted by limestone dust imissions close to the quarry, and the Nature Reserve Lošonský háj grove. Also the fauna of centipedes (Chilopoda) on both study sites was similar with value 70% (Országh, Országhová, 2005). On the contrary, in Poland a pollution by a cement and lime factory corresponded with lower density and species number of Oribatida and Gamasida (Seniczak et al., 1997). For example in the study sites LL and LH the earwig

109

Chelidura acanthopygia (Dermaptera) responded to the limestone dust caused the neutral value (pH = 6.74) of the upper soil horizon (Országh, 2005). Clearly the age of forest, presence of dead wood and deep layer of leaf litter in the forest, pedological characteristics like pH or content of humus are important factors for mite assemblages. A long–term investigation of the soil fauna in the littoral zone of the Jakubovské rybníky fishponds on a blown sand area in the southwestern Slovakia, where the leaf litter, humus layer or dead wood completely absent, shows comparable fauna (H’ = 2.974, N0 = 106, N1 = 19.57, N2 = 9.901, E1 = 0.634) like in oak forests in the Malé Karpaty Mts (Fenďa, Schniererová, 2005). The fauna of mesostigmatid mites of oak forests in Slovakia is not markedly rich, humidity is probably one of the most important factors for the species richness and abundance of mesostigmatid mites. The study site Lošonský háj grove Nature Reserve has more balanced mite community, there are more stable microclimatic conditions. The deep layer of leaf litter is able to keep the humidity for a long time. The highest diversity of centipedes (Chilopoda) in the Malé Karpaty Mts was observed just in Lošonský háj grove Nature Reserve (Országh, Országhová, 2005). The study site Lošonec–lom quarry is drier, for example the lower abundance of hygrophilous weevils (Coleoptera, Curculionidae) in the Malé Karpaty Mts was detected just in the study site LL (Holecová et al., 2005). From the faunistical point of view, Pergamasus canestrinii was recorded for the first time in Slovakia. The species Sessiluncus hungaricus, Trachytes baloghi, Trichouropoda elegans, T. penicillata, Zercon hungaricus, and probably also Pergamasus ruhmi are typical thermophilic species in Slovakia. Moreover, the species Zercon vacuus is a typical element of forest litter of western Slovakia (Mašán, Fenďa, 2004). Translated by P. Fenďa Acknowledgement Our thanks belong to M. Holecová and I. Országh for collecting material in field. The research was supported by VEGA (Scientific Grant Agency of the Ministry of Education and the Slovak Academy of Sciences), grant No. 1/0119/03.

References Ambros, M., 1984: The fauna of mites (Acari: Mesostigmata) on the small mammals of the Záhorská nížina lowland and the Malé Karpaty Mts (in Slovak). Acta Rerum Naturalium Musei Nationalis Slovaci, 30, p..87–102. Ambros, M., Krištofík, J., Šustek, Z., 1992: The mites (Acari, Mesostigmata) in the birds nests' in Slovakia. Biologia, Bratislava, 47, p. 369–381. Balogh, J., 1958: Lebensgemeinschaften der Landtiere. Akadémiai Kiadó, Akademie–Verlag, Budapest, Berlin, 560 pp.

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Fenďa, P., Schniererová, E., 2005: Mites (Acarina, Gamasida) in littoral zone of Jakubov fishponds (Slovakia). In Tajovský, K., Schlaghamerský, J., Pižl, V. (eds): Contributions to Soil Zoology in Central Europe I. ISB AS CR, České Budějovice, p. 9–14. Holecová, M., Némethová, D., Kúdela, M., 2005: Structure and function of weevil assemblages (Coleoptera, Curculionoidea) in epigeon of oak-hornbeam forests in SW Slovakia. Ekológia (Bratislava), 24, Suppl. 2, p..179–204. Hyatt, K.H., Emberson, R.M., 1988: A review of the Macrochelidae (Acari: Mesostigmata) of the British Isles. Bull. Br. Mus. (Nat. Hist.) Zool., 54, p. 63–125. Kalúz, S., 2005: Soil mites (Acarina) (in Slovak). In Majzlan, O. (ed.): Fauna Devínskej Kobyly. APOP edícia, Bratislava, p. 44–55. Kalúz, S., Fenďa, P., 2005: Mites (Acari, Mesostigmata) of the family Ascidae of Slovakia. Institute of Zoology, Slovak Academy of Sciences, NOI Press, Bratislava, 168 pp. Karg, W., 1993: Acari (Acarina), Milben. Parasitiformes (Anactinochaeta) Cohors Gamasina, Leach, Raubmilben. (Die Tierwelt Deutschlands 59). Gustav Fischer Verlag, Jena–Stuttgart–New York, 523 pp. Ludwig, J.A., Reynolds, J., 1988: Diversity indices. In Statistical Ecology. Whilley–Interscience Public, New York, p. 85–103. Mašán, P., Kalúz, S., Babjaková, A., 1994: Mites (Acarina) from the winter nests of the common mole (Talpa europaea L.) in south Slovakia. Biologia, Bratislava, 49, p. 667–673. Mašán, P., 1999: Mites (Acarina) associated with burying and carrion beetles (Coleoptera, Silphidae) and description of Poecilochirus mrciaki sp.n. (Mesostigmata, Gamasina). Biologia, Bratislava, 54, p. 515–524. Mašán, P., Kalúz, S., 2001: The adult stages of Stylochirus fimetarius (Acari, Mesostigmata) and new systematic status of the genus Iphidosoma. Biologia, Bratislava, 56, p. 483–488. Mašán, P., 2001a: Mites of the cohors Uropodina (Acarina, Mesostigmata) in Slovakia (in Slovak). Annotationes Zoologicae et Botanicae (Bratislava), 223, p. 1–320. Mašán, P., 2001b: First records of mesostigmatid mite species (Acarina) in Slovakia. Biologia (Bratislava), 56, p. 216–218. Mašán, P., 2003: Macrochelid mites of Slovakia (Acari, Mesostigmata, Macrochelidae). Institute of Zoology, Slovak Academy of Sciences, NOI Press, Bratislava, 149 pp. Mašán, P., Fenďa, P., 2004: Zerconid mites of Slovakia (Acari, Mesostigmata, Zerconidae). Institute of Zoology, Slovak Academy of Sciences, NOI Press, Bratislava, 238 pp. Nosek, J., Lichard, M., 1962: Beitrag zur Kenntnis der Vogelnestfauna. Entomologické Problémy (Bratislava), 2, p. 29–51. Országh, I., 2005: Phenology of Chelidura acanthopygia (Dermaptera: Forficulidae) larval instars in oak-hornbeam forests of the Malé Karpaty Mts and Trnavská pahorkatina hills (SW Slovakia). Ekológia (Bratislava), 24, Suppl. 2, p. 152–160. Országh, I., Országhová, Z., 2005: Structure of centipede communities (Antennata: Chilopoda) in oak-hornbeam forests of the Malé Karpaty Mts and Trnavská pahorkatina hills (SW Slovakia). Ekológia (Bratislava), 24, Suppl. 2, p. 124–142. Podani, J. 1988. Syn–tax III user's manual. Abstracta Botanica, Suppl. 1, p. 1–183. Seniczak, S., Dabrowski, J., Klimek, A., Kaczmarek, S., 1996: The mites (Acari) associated with young Scots pine forests polluted by a chemical factory Lubon near Poznan. Bulletin de l'Académie Polonaise des Sciences (Série des Sciences Biologiques), 44, p. 101–104. Seniczak, S., Dabrowski, J., Klimek, A., Kaczmarek, S., 1997: Effect of pollution produced by a cement and lime factory Kujawy on mites (Acari) associated with young scots pine forests in Bielawy, Poland. Bulletin de l'Académie Polonaise des Sciences (Série des Sciences Biologiques), 45, p. 1–6. Seniczak, S., Kaczmarek, S., Klimek, A., Seniczak, A., 2002: The effect of some air pollutants on the vertical distribution of mites (Acari) in soils of young Scots pine forests in Poland. European Journal of Soil Biology, 38, p. 311–314. Schwerdtfeger, F., 1975: Ökologie der Tiere. Band III – Synökologie. Verlag Paul Parey, Hamburg – Berlin, 451 pp. Tischler, W., 1949: Grundzüge der Terrestrischen Tierökologie. Friedrich Vieweg, Braunschweig, 219 pp. Zlinská, J., Šomšák, L., Holecová, M., 2005: Ecological characteristics of studied forest communities of an oak-hornbeam tier in SW Slovakia. Ekológia (Bratislava), 24, Suppl. 2, p. 3–19. Received 30. 7. 2005

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Fenďa P., Ciceková J.: Pôdne roztoče (Acari, Mesostigmata) dubových lesov Malých Karpát (Západné Slovensko). V rokoch 1999–2002 sme uskutočnili výskum mesostigmátnych roztočov v dvoch dubovo–hrabových lesoch Malých Karpát. Zistili sme 75 druhov patriacich do štyroch kohort (Antennophorina, Gamasina, Sejina, Uropodina). Spoločenstvá roztočov na lokalite Lošonec–lom ovplyvnenej vápenným prachom z neďalekého lomu vykazovali väčšie výkyvy v počte druhov a v početnosti počas roka. V prírodnej rezervácii Lošonský háj je zreteľný vplyv stabilnejších mikroklimatických podmienok v lese. Porovnaním spoločenstiev pôdnych roztočov v jednotlivých rokoch na študovaných lokalitách sa ukázalo, že teplota a vzdušná vlhkosť majú na druhovú skladbu a početnosť mesostigmátnych roztočov väčší vplyv ako typ pôdy, pH a vegetácie. Z.faunistického hľadiska sme zaznamenali prvý nález druhu Pergamasus canestrinii pre územie Slovenska.

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Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 113–123

COMMUNITIES OF TERRESTRIAL ISOPODS (Crustacea: Isopoda: Oniscidea) IN EPIGEON OF OAK-HORNBEAM FORESTS OF SW SLOVAKIA IVAN H. TUF, JANA TUFOVÁ Department of Ecology and Environmental Science, Faculty of Science, Palacký University, Svobody 26, 772 00 Olomouc, The Czech Republic, e-mail: [email protected] Abstract Tuf I.H., Tufová J.: Communities of terrestrial isopods (Crustacea: Isopoda: Oniscidea) in epigeon of oak-hornbeam forests of SW Slovakia. Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 113–123. Terrestrial isopods were studied in 10 oak-hornbeam forests in the Malé Karpaty Mts and in their vicinity. In total, seven species were recorded. Individual communities consisted from one to six species, Protracheoniscus politus and Porcellium collicola dominated in most of the studied sites. Beside the age of forest growth, the soil type and pH seems to be important environmental factors affecting the species richness of isopod communities. Key words: Oniscidea, terrestrial isopods, Malé Karpaty Mts, Slovakia

Introduction Terrestrial isopods (Crustacea: Isopoda: Oniscidea) represent one of the main groups of soil macrofauna importantly participated in soil-forming processes. They mechanically destroy dead leaves and others plant rests, they stir anorganic particles with fragments of dead organic matter and enhance thus surfaces of organic matter accessible for soil microorganisms (Hassall et al., 1987; Zimmer et al., 2005). Isopods are decomposers of animal carrions, too (Grassberger, Frank, 2004). On the other hands, together with millipedes play important role in food web as an source of calcium for insectivorous birds and other animals (Graveland, Vangijzen, 1994). In SW Slovakia, the terrestrial isopods were studied by Gulička (1960) in the Svätojurský Šúr within the investigation of ecological consequences of changes of water regime and soil macrofauna. Population ecology and biology of terrestrial isopods were studied in the same site in the seventies by Krumpál (1973, 1976). A detail inventory of terrestrial isopod fauna was done in the Malé Karpaty Protected Landscape Area by Flasarová (Flasarová, 1980, 1986; Flasar, Flasarová, 1989). 113

Study area The Malé Karpaty Mts are situated in the western part of Slovakia approximately between the towns Bratislava and Nové Mesto nad Váhom. It represents the most western part of Carpathian Mountains arch and SW corner of Central Western Carpathians. The highest peak (Záruby) amounts 768 m a.s.l. Fauna and flora in these mountains are enhanced by diffusion of Pannonian biogeographic elements. Within this mountain range, ten forest sites were investigated for terrestrial isopod fauna. Positions of sites on the map, their altitudes, expositions and slopes of uphill and phytocoenological and pedological characteristics are summarized in Zlinská et al. (2005). CA – (Cajla) – forest (Querco-Carpinetum poetosum nemoralis) at the foot of the Malá cajlanská homola hill; VI – (Vinosady) – forest (Querco-Carpinetum poetosum nemoralis) at the foot of the Kamenica hill; FU – (Fúgelka) – forest (Querco-Carpinetum melicetosum uniflorae) near the Dubová village; LI – (Lindava) – forest (Quercetum petrae-cerris) in the Lindava Nature Reserve near the Píla village in vicinity to the Malé Karpaty Mts; HH – (Horný háj grove) – forest (Querco-Carpinetum melicetosum uniflorae) in vicinity to the Malé Karpaty Mts, near the Horné Orešany village, refuge within fields and vineyards; LH – (Lošonský háj grove) – forest (Querco-Carpinetum caricetosum pilosae) in the Lošonský háj Nature Reserve; LL – (Lošonec–lom quarry) – forest (Querco-Carpinetum melicetosum uniflorae) near the Lošonec quarry, litter, trees and herbs strongly covered by calcareous dust; NA – (Naháč–Kukovačník) – small young forest island (Querco-Carpinetum melicetosum uniflorae) surrounded by fields and pastures; NK1 – (Naháč–Katarínka) – young forest (Querco-Carpinetum melicetosum uniflorae) in the Naháč–Katarínka Nature Reserve; NK2 – (Naháč–Katarínka) – old forest growth (Lithospermo-Quercetum virgilianae) in the Naháč–Katarínka NR under monastery ruin.

Material and methods The material of isopods was collected in the years 1999 to 2002 in approximately monthly intervals; eight times per year in 1999–2001 and ten times per year in 2002. Sites LL, LH and HH were investigated in 2000–2002 period only. In each site the litter was sifted from 1 m2 (16 squares of 25 × 25 cm). The sifted material was separated using xereclectors, and the animals were fixed in 75% ethylalcohol. Species were determined by using the key of Frankenberger (1959) and Schmölzer (1965), the used classification is according to Schmalfuss (2003). The computer program JMP (SAS Institute Inc., 1995) was used for cluster analysis of isopod communities (Ward’s method). Species diversity was evaluated according to Odum (1977), all couples of diversity values

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were tested with t-tests (Poole, 1974). Redundant analysis (RDA) was performed for evaluating of the relationships among distribution of species and main environmental factors using the programme CANOCO (ter Braak, Šmilauer, 1998). Species scores were divided by standard deviations, species data were not transformed and were centred by species, using the Monte Carlo Permutation test (500 permutations).

Results In all sites, the occurrence of seven species of terrestrial isopods (Isopoda: Oniscidea) were recorded together (Table 1). The whole material included 3,798 specimens. Generally, the main dominants were Protracheoniscus politus (2,777 individuals) and Porcellium collicola (840 individuals). A total survey of coenoses in studied sites including the values of mean densities and diversity indexes is presented in Table 2, a survey of collected terrestrial isopods in individual years is given in Table 3. T a b l e 1.: Survey of recorded species of terrestrial isopods. Order: Isopoda Suborder: Oniscidea Family: Ligiidae Ligidium hypnorum (C u v i e r, 1792) Family: Trichoniscidae Hyloniscus riparius (C.L. K o c h, 1838) Family: Philosciidae Lepidoniscus minutus (C.L. K o c h, 1838) Family: Agnaridae Orthometopon planum (B u d d e - L u n d, 1885) Protracheoniscus politus (C.L. K o c h, 1841) Family: Trachelipodidae Trachelipus ratzeburgii (B r a n d t, 1833) Porcellium collicola (V e r h o e f f, 1907)

Comparison of data from individual years showed, that total numbers of sampled isopods (in sum from all sites) were very similar in the years 2000–2002 (824, 893, and 833 ind. respectively), beside the year 1999 was more rich for isopods, when 1,248 individuals were obtained from seven sites only. No trends are evident from changes in density at individual sites (Table 3). CA: In this site, only two species were collected. Protracheoniscus politus was more abundant than Porcellium collicola. Moreover, abundance of P. collicola decreased from 25% in 1999 to 0% in 2002 gradually. The highest mean density was in 1999 (8.0 ind.m-2) and the lowest in 2001 (3.3 ind.m-2). VI: During the whole study, only Protracheoniscus politus was collected here. The highest mean density was recorded in 1999 (27.6 ind.m-2) and the lowest in 2000 (4.6 ind.m-2). FU: Three species were collected in this forest site. Protracheoniscus politus was constant and dominant member of the present isopod community. Porcellium collicola was observed only in 2000 and 2002 years in a few specimens, and Ligidium hypnorum 115

T a b l e 2. Isopod communities in individual sites: dominance of species, selected coenological characteristics and t-test of species diversity (Poole, 1974) based on the summarized data from 1999–2002 (sites HH, LH and LL only 2000–2002). Site Ligidium hypnorum Hyloniscus riparius Lepidoniscus minutus Orthometopon planum Protracheoniscus politus Trachelipus ratzeburgii Porcellium collicola Total individual Number of species Mean density (ind.m-2) Pielou’s index of equitability Simpson’s index of dominance Shannon’s index of species diversity CA VI FU LI HH LH LL NA NK1 NK2

CA 86 14 164 2 4.97 0.59 0.76 0.41 8.214*** 4.436*** 0.022ns 5.780*** 4.128*** 6.258*** 2.876** 7.182*** 12.154***

VI 100 454 1 13.76 0 1 0 164 4.146*** 13.275*** 122.469*** 2.911** 14.999*** 7.094*** 1.990* 43.261***

FU 0.7 97.3 2 407 3 12.33 0.13 0.95 0.14 325.02 407 5.805*** 16.125*** 0.174ns 10.970*** 1.998* 2.876** 22.327***

LI 86 14 430 2 13.03 0.58 0.76 0.4 296.17 430 822.76 9.315*** 5.057*** 7.266*** 3.773*** 10.780*** 17.117***

HH LH LL NA NK1 0.5 0.2 0.1 0.1 0.5 48.3 97.1 67 94.3 99.6 2.9 21.8 0.1 51.7 10.1 5.3 145 174 188 511 702 2 2 5 3 4 5.8 6.96 7.52 14.63 20.06 1 0.19 0.55 0.17 0.02 0.5 0.94 0.51 0.89 0.99 0.69 0.13 0.89 0.24 0.03 168.3 332.65 347.67 323.37 200.79 145 174 188 512 702 429.17 376.49 315.11 902.08 597.73 458.57 342.46 291.73 941.98 675.44 179.54 191.43 540.76 824.09 12.459*** 343.29 377.4 221.72 3.282** 10.206*** 312.51 216.98 13.654*** 1.873ns 9.634*** 756.13 38.440*** 2.059* 13.940*** 5.492*** 15.067*** 18.485*** 2.954** 20.361*** 35.164***

Abbreviations of sites – see the text. T-test values are given below the diagonal and degrees of freedom above. Significance levels: *** = P < 0.001; ** = 0.001 < P < 0.01; * = 0.01 < P < 0.05; ns = 0.05 < P (non-significant)

NK2 0.2 18.9 0.2 51.8 1 27.9 623 6 17.8 0.6 0.38 1.08 253.8 623 812.54 915.18 681.13 290.55 258.03 994.61 1089.66 -

was collected in 1999 only. The highest mean density was in 1999 (22.5 ind.m-2) and the lowest in 2000 (6.3 ind.m-2). LI: Only two species were recorded in the litter in this forest. Protracheoniscus P. politus was more abundant than Porcellium collicola, which represented 14% of whole material. The highest mean density was in 2000 (21.4 ind.m-2) and the lowest in 2002 (7.9 ind.m-2). HH: In this site, only two species were collected, too. Protracheoniscus politus was more abundant than Porcellium collicola in the first two years, in the last year P. collicola dominated. But in total evaluation dominances of both species were well-balanced. In this site the highest index of equitability was recorded (Table 2). The highest mean density was in 2002 (7.4 ind.m-2) and the lowest in 2000 (4.3 ind.m-2). LH: In this site, two species were collected. In comparison with other forests, Porcellium collicola was missing here. Protracheoniscus politus was more abundant than Trachelipus ratzeburgii, which represented 3% of the whole sampled material. The highest mean density of the whole community was in 2000 (8.9 ind.m-2) and the lowest in 2001 (5.4 ind.m-2). LL: Relatively rich community of isopods was sampled in this site. Protracheoniscus politus, Porcellium collicola and Trachelipus ratzeburgii were dominant species, and the species Hyloniscus riparius and Orthometopon planum were recorded in 2002, both in one specimen. The highest mean density was in 2001 (9.3 ind.m-2) and the lowest in 2000 (5.1 ind.m-2). NA: In this site, Protracheoniscus politus predominated. Porcellium collicola represented only 5% of the whole sampled material and only 1 specimen of Hyloniscus. riparius was recorded. The highest mean density was in 1999 (20.9 ind.m-2) and the lowest in 2002 (5.8 ind.m-2). NK1: Besides seven hundreds specimens of Protracheoniscus politus, three species Trachelipus ratzeburgii, Hyloniscus riparius and Lepidoniscus minutus were sampled, each in one specimen only. The isopod Porcellium collicola was missing. The highest mean density was in 1999 (39.5 ind.m-2) and the lowest in 2001 (11.5 ind.m-2). NK2: The richest community (six species) inhabited litter in this site. Protracheoniscus politus, Porcellium. collicola and Hyloniscus riparius dominated, Trachelipus ratzeburgii, Orthometopon planum and Ligidium hypnorum were found in a few specimens. It corresponds with the highest index of species diversity (Table 2). The highest mean density was in 1999 and 2001 (23.4 ind.m-2) and the lowest in 2000 (12.8 ind.m-2). Comparison of isopod communities based on the cluster analysis (Fig. 1) divided sites into two groups. One group coupled forests with one to three species, with Protracheoniscus politus and/or Porcellium collicola as dominant. In the second group were sites with other species or (as a subgroup) without P. collicola. The position of the species poor site LH in the diagram relatively closely together with NK1 can explain the highest percentage of P. politus (Table 1). Similar pattern shown set of t-tests where are not significant differences between sites CA and LI (in accord with cluster analysis), and between sites LH – FU and LH – NA, i.e. forests with 2-3 species and dominancy of P. politus more than 90% (Table 2).

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T a b l e 3. Number of collected isopods during individual years in individual sites. Sites were inspected in 1999 to 2001 eight times per year, in 2002 ten inspections. Site Year L. hypnorum H. riparius L. minutus O. planum P. politus T. ratzeburgii P. collicola Total

CA ´99 49 15 64

´00 35 7 42

´01 25 1 26

VI ´02 32 32

´99 221 221

´00 37 37

FU

´01 131 131

´02 65 65

´99 3 177 180

´00 47 2 49

´01 78 78

´02 94 6 100

´99 98 15 113

LI ´00 ´01 - - - - 164 43 - 7 32 171 75

´02 65 6 71

´00 21 13 34

HH ´01 ´02 - - - - 32 17 - 12 50 44 67

´00 67 4 71

LH ´01 42 1 43

´02 60 60

´00 38 3 41

LL ´01 45 21 8 74

´02 1 1 43 17 11 73

´99 1 158 8 167

NA ´00 ´01 135 133 8 10 143 143

´02 57 1 58

´99 1 315 316

NK1 ´00 ´01 - - - 1 - 133 91 1 - 134 92

´02 160 160

´99 1 33 132 21 187

NK2 ´00 ´01 - 8 69 - - 1 58 51 5 31 66 102 187

´02 8 82 1 56 147

Abbreviations of sites – see the text

T a b l e 4. Presence of isopod species during the year based on the summarized data from all sites and all years. Month Ligidium hypnorum Hyloniscus riparius Lepidoniscus minutus Orthometopon planum Protracheoniscus politus Trachelipus ratzeburgii Porcellium collicola

III

IV

V

Trapped individuals VI VII VIII IX

X

XI

III

IV

Proportion of population V VI VII VIII IX

X

XI

ind. total 4 121 1 2 2.777 53 840

The first part of table corresponds with recording probability of these species – increasing intensity of colour corresponds wit h increasing total numbers of isopods in these four categories: white – not recorded, bright grey 1–9 ind., dark grey 10–49 ind., black ≥ 50 ind. The second part of table shows patterns of changes in densities of individual species, with dominancy in these four categories: white – 0 %, bright grey 1–9 %, dark grey 10–49 %, black ≥ 50 %. Numbers of collected individuals are presented in the last column.

CA LI HH NA FU VI LH NK1 LL NK2

4 3 2 1

Fig. 1. Cluster analysis of dissimilarity of isopod assemblages on individuals study plots according to their species presence/absence (JMP, Ward method), for abbreviations of sites see text.

Fig. 2. RDA analysis of community structure and selected environmental factors. Individual axes explain 0.235; 0.077; 0.014; 0.002 of variability, all canonical axes explain 0.328. Abbreviations: exposition – degree of southern exposition of site, %C – content of carbon, %N – content of nitrogen, Ca dust – presence of calcareous dust at surfaces, slope – rate of slope of hillside.

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Redundancy analysis (Fig. 2) showed that the most important environmental factor seemed to be age of forest growth (expressed in the graph by the longest arrow in the third quadrant), although the oldest forests were inhabited on the average by 3.3 species and the youngest ones by 4 species. Nevertheless, in the youngest forests, the species P. politus represented more than 90% of community. The occurrence of individual species during the year is summarised in the Table 4. Lepidoniscus minutus was extracted in June only, two other species, Ligidium hypnorum and Orthometopon planum, were detected in the field in the start and/or in the end of vegetation season (April, October and November). The other species were recorded during all months. Although Protracheoniscus politus, Porcellium collicola and Trachelipus ratzeburgii were numerous during whole year, from the second part of Table.4 is evident, that their densities decreased during months with unfavourable conditions (March, summer, late autumn).

Discussion Generally, the total number of species found in studied sites is low. This result was caused by (1) used method (litter sifting only), (2) apparently also by a low number of studied sites and (3) more or less uniform type of forests, too. Flasarová (Flasarová, 1986; Flasar, Flasarová, 1989) listed for the Malé Karpaty Mts in total 27 species, but she used individual collecting. Thus she sampled isopods from diverse microhabitats, not only from forest litter. She collected isopods under stones, pieces of wood, under the bark of stumps and fallen trunks, under loose plaster of castle masonry, i.e. from microhabitats with usually higher moisture and/or higher content of calcium. In addition, she elaborated material completed during eight years (1978–1985) and collected in about 110 sampling sites. Numbers of species in isopod communities sampled by Flasarová varied from one to 11. Forest community of isopods in her studies consisted from two to three species only. The rich communities with five and more species were sampled in wet sites (shores of brooks and creeks) and/or in synanthropic habitats (quarry, villages and village greens, abandoned yards, churchyards, castle ruins, holiday resorts etc.). Most of terrestrial isopods are hygrophilous, from this reason in more humid sites there form richer communities. For example, the communities in Central Moravian floodplain forests consist from five to eight species (Tuf, 2003). Poor communities (2–3 three species) in studied sites can be determined by relatively dry type of forests in comparison with humid riparian growths. Similarly, pitfall trapping in oak forests, used as another method for sampling of isopods, served 4–7 species per studied site (Farkas et al., 1999). Some isopod species recorded by Flasarová (1986) and missing in this research are frequently associated with man-influenced environments like ruins, quarries, yards etc. (Armadillidium vulgare (L a t r e i l l e, 1804), A. versicolor S t e i n, 1859, Porcelio spinicornis S a y, 1818, Protracheoniscus major (D o l l f u s, 1903), Oniscus asellus L.i.n.n.a e u s, 1758), some of them are more endogeic or associated with deeper soil layers (Haplophthalmus mengei (Z a d d a c h, 1844), H. danicus B u d d e - L u n d, 120

1880, Androniscus roseus (C. K o c h, 1838), Trichoniscus pusillus B r a n d t, 1833, T. pygmaeus S a r s, 1898, Hyloniscus transsilvanicus (V e r h o e f f, 1901)) or they are myrmecophilous (Platyarthrus hoffmannseggii B r a n d t, 1833). Not only moisture, but also availability of calcium is important factor for distribution of terrestrial isopods, too. Calcium is structural element of isopod integument (e.g. Fabritius, Ziegler, 2003). Both studied sites with higher numbers of species (LL – 5 and NK2 – 6) are on the Rendzina soil type with higher content of calcium. In addition, site LL was relatively permanently covered by dust from neighbouring limestone quarry. Both above mentioned sites LL and NK2 were characteristic by pH value higher than 6. Such conditions are preferred by terrestrial isopods as well (Sastrodihardjo, Van Straalen, 1993). Beside these facts, both sites had high values of slope, 8–10° in LL and 45° in NK2 (Holecová, Sukupová, 2002; Zlinská et al., 2005). The higher slope can support higher diversity of microhabitats, for instance in connection with humidity gradient etc. In the site with the higher slope (NK2) was one from dominant species hygrophilous isopod Hyloniscus riparius, stable member of communities from floodplain forests (Tajovský, 1999; Tuf, 2003). The most important factor in RDA analysis is age of forest growth. Beside some irregularities (VI, HH), index of species diversity and index of equitability are positively correlated with age of studied forest. It is in accord with similar trend documented by Tajovský (2001) for terrestrial isopod communities in afforested mine sites and well know for diverse groups of soil invertebrates in primary forest succession (Dunger et al., 2001). Explanation for the highest densities in the year 1999 can be a rainy summer (158 % of mean precipitation for June) and the warmer start and end of the vegetation season – April warmer by 1.8 °C and September warmer by 3.6 °C than mean temperature recorded for these months (Zlinská et al., 2005). Such seasonal conditions can be optimal for development of isopod populations. Otherwise during dry summer they migrate deeper into the soil (Tuf, 2002) and are not detectable by sieving of litter. The records of the hygrophilous species Ligidium hypnorum on two sites only in the spring and autumn months can be caused by higher humidity of litter during these periods (Harding, Sutton, 1985). On the contrary, presence of Orthometopon planum only during could months is surprising and accidental, because this species is originated from Mediterranean region and in Slovakia it prefers warm forests (Frankenberger, 1959). In conclusion, relatively poor but typical terrestrial isopod communities with predominating two species Protracheoniscus politus and Porcellium collicola were described from the several oak-hornbeam forests from the SW Slovakia. Number of species varied among 1 to 6 with mean density from 5 to 20 ind.m-2. The most abundant species was stable during mean year cycle, other more rare species were recorded occasionally during moths with favourable climatic conditions. From the analysis, the main factors affecting structure of communities seems to be age of forest growth, pH and content of Ca (influenced by calcareous dust and soil type). Translated by P. J. Fedor

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Acknowledgement The authors thank to M. Holecová (Comenius University Bratislava) for sampling of material. Her help and help of A. Véle (Palacky University Olomouc) with statistical analyses are highly appreciated. This project was supported by grant of VEGA No. 1/0119/03.

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Tajovský, K., 1999: Impact of inundations on terrestrial arthropod assemblages in southern Moravia floodplain forests, the Czech Republic. Ekológia (Bratislava), 18, Suppl. 1, p. 177–184. Tajovský, K., 2001: Colonization of colliery spoil heaps by millipedes (Diplopoda) and terrestrial isopods (Oniscidea) in the Sokolov region, Czech Republic. Restoration Ecology, 9, p. 365–369. ter Braak, C.J.F., Šmilauer, P., 1998: CANOCO References Manual and User’s Guide to Canoco for Windows: Software for Canonical Community Ordination (verrsion 4). Microcomputer Power, Ithaca, NY, USA, 352 pp. Tuf, I.H., 2002. Contribution to the knowledge of vertical distribution of soil macrofauna (Chilopoda, Oniscidea). In Tajovský, K., Balík, V., Pižl, V. (eds.): Studies on Soil Fauna in Central Europe. ISB AS CR, České Budějovice, p. 241–246. Tuf, I.H., 2003: Development of the community structure of terrestrial isopods (Crustacea, Isopoda, Oniscidea) after a summer flood. In Sfenthourakis, S., de Araujo, P.B., Hornung, E., Schmalfuss, H., Taiti, S., Szlávecz, K. (eds): The biology of terrestrial isopods V. (Crustaceana Monographs, 2). Brill Academic Publisher, Leiden, p. 231–242. Zimmer M., Oliveira R., Rodrigues E., Graca M.A.S., 2005: Degradation of leaf litter phenolics by aquatic and terrestrial isopods. Journal of Chemical Ecology, 31, p. 1933–1952. Zlinská, J., Šomšák, L., Holecová, M. 2005: Ecological charactersitics of studied forest communities of an oak-hornbeam tier in SW Slovakia. Ekológia (Bratislava), 24, Suppl. 2, p. 3–19. Received 30. 7. 2005

Tuf.I.H., Tufová.J.: Společenstva stejnonožců (Crustacea: Isopoda) v epigeonu dubo-habrových lesů JZ Slovenska. Na vybraných deseti lokalitách, reprezentujících dubohabrové porosty v prostoru CHKO Malé Karpaty a blízkého okolí byla zkoumána společenstva suchozemských stejnonožců. Celkem bylo nalezeno sedm druhů, přičemž jednotlivá společenstva byla tvořena jedním až šesti druhy. Protracheoniscus politus a Porcellium collicola byli v těchto společenstvech dominantními druhy, přičemž průměrná abundance na jednotlivých lokalitách se pohybovala mezi 5 až 20 ind.m-2. Z testovaných environmentálních faktorů nejvýznamněji ovlivňovaly druhovou bohatost společenstev stejnonožců stáří porostu, půdní typ (dostatek vápníku) a pH půdy.

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Vol. 24, Supplement 2/2005, p. 152–160

PHENOLOGY OF Chelidurella acanthopygia (Dermaptera: Forficulidae) NYMPHAL INSTARS IN OAK-HORNBEAM FORESTS OF THE MALÉ KARPATY MTS AND TRNAVSKÁ PAHORKATINA HILLS (SW SLOVAKIA) IVAN ORSZÁGH Department of Zoology, Comenius University, Mlynská dolina B-1, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected] Abstract Országh I.: Phenology of Chelidurella acanthopygia (Dermaptera: Forficulidae) nymphal instars in oak-hornbeam forests of the Malé Karpaty Mts and Trnavská pahorkatina hills (SW Slovakia). Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 152–160. In the Malé Karpaty Mts as well as nearby Trnavská pahorkatina hills the phenology of Chelidurella acanthopygia nymphal instars was studied in the period of 1999–2002. Abundance of the species varied between 1–33 ind.m-2 at the study sites. The first specimens, adults of both sexes from the previous year shortly after their overwintering appeared in the samples already in March. The first nymphal instar (N1) was sampled from the second decade of April till the end of the second decade in June. Interval of their existence took approximately 70 days. The second instar (N2) appeared from May (2nd decade) to the end of July in the period of more than 80 days. The third instar (N3) was recorded in the period of more than 75 days from June (beginning of the 2nd half) to the 1st week of September. The fourth instar (N4) was present from the beginning of July to September (1st decade) (period of more than 70 days). The first new generation adults (females and males) appeared at the end of summer (August). After winter they will form a new parental generation for the next year. The obtained data on the instars N1–N4 and length of their existence in the Malé Karpaty Mts declare that the ontogenesis of C. acanthopygia from the 1st nymphal instar to imago takes 100–120 days. Key.words:.Chelidurella acanthopygia, Dermaptera, nymphal instars, phenology, Malé Karpaty Mts, SW Slovakia

Introduction In the Malé Karpaty Mts the earwig Chelidurella acanthopygia (G é n é, 1832) belongs to common inhabitants of fallen leaf litter in oak-hornbeam forests as a part of epigeic macrofauna. It participates on formation of herbal detritus. From the trophic point of view it is classified as a decomposer (Turček, 1974). According to Franke (1985) C. 152

acanthopygia represents an omnivorous species with a preference to animal food. Its ontogenesis includes 4 nymphal instars. In the Malé Karpaty Mts it has one generation annually and overwinters as an adult. The available data declare that its occurrence as well as quantity at a site is determined by temperature, humidity and depth of leaf litter, quantity of decaying wood, presence of bigger-sized stones on substrate. It avoids open landscape and prefers forested areas. There are no detailed studies on correlation between its occurrence and chemical variables of soil as well as undergrowth. It has been recorded in most W and C Europe, some countries of S and SE Europe, Scandinavia and Baltic region. In E Europe its distribution area includes even W Ukraine (Bey-Bienko, 1936; Albouy, Caussanel, 1990). In Slovakia there are some records on this species already from the end of the 19th century (Petrogalli, 1890; Petricskó, 1892). There are just sporadic data published on its distribution in the Malé Karpaty Mts. From the village of Sološnica in the western part of the Malé Karpaty Mts and from the village of Dobrá Voda in the northern part of the mountains the earwig Chelidurella acanthopygia was mentioned by Kočárek, Galvagni (2000). The records from Devínska Kobyla (the southern margin of the Malé Karpaty Mts) were published by Lukáš (2003) and Fedor (2005). Straka (2005) collected Chelidurella acanthopygia in the Biele Karpaty Mts (Javorník – Vršatecké Podhradie; Krivoklát) and Strážovské vrchy Mts (Mitická step – Trenčianske Mitice).

Study area With a total area of 65,504 ha the Malé Karpaty Mts are situated in the western part of Slovakia from Bratislava towards the town of Nové Mesto nad Váhom on NE within the length of 100 km. The altitude varies between 132 m a.s.l. (by the Danube river) and 768 m a.s.l. (Mount Záruby). The main range grows to the altitude of 450–650 m a.s.l. The Malokarpatský kras karst represents a specific area of approximately 180 km2. The Malé Karpaty Mts belong to the gently warm climatic zone with an average annual temperature of 7–9 °C, which declines under 8 °C at the altitudes of 400 m a.s.l. The vegetation period (with the average daily temperature of 10 °C) usually takes 180 days, respectively 160 days at higher altitudes. Precipitation reaches 650–900 mm in the mountains, 600–650 mm in the lowland. The area of the Malé Karpaty Mts belongs to the Subcarpathian district of the deciduous forest province. The eastern side has been influenced by the Steppe province (Pannonian district) as plenty of thermophilous species infiltrate onto S and SE slopes. Dominant forest stands have been considerably replaced by vineyards on the SE and SW slopes, by pastures and farmland on north. From the hypsometric point of view the area is inhabited by colinous and submountainous elements forming the communities in forests, woody steppes and rocky habitats.

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Study sites All the sites are situated on forested area. The plots of Cajla (CA); Vinosady (VI); Fúgelka (FU); Lošonec–lom quarry (LL); Lošonský háj grove (LH); Naháč–Kukovačník (NA); Naháč–Katarínka 1, young forest (NK1) and Naháč–Katarínka 2, old forest (NK2) are located in the zone of oak-hornbeam forests on the SE slopes of the Malé Karpaty Mts. The sites of Lindava (LI) and Horný háj grove (HH) spread out on the boundary of the Malé Karpaty Mts and Trnavská pahorkatina hills. The study sites are situated at the altitude of 240–350 m a.s.l. on acid soils (pH varies between 3.8–5.0) with the only exception of Lošonec–lom quarry (LL) and Naháč–Katarínka 2, old forest (NK2) laying on weakly acid to almost neutral soil (pH 6.5–6.7). The air distance between the first site Cajla (CA) and the most northern one – Naháč–Katarínka 1, young forest (NK1) is measured as approximately 30 km. The map with the study sites and the more detailed pedological and botanical characteristics of the study sites as well as the climatic conditions in the period of 1999–2002 have been included in the paper by Zlinská et al. (2005).

Material and methods The collections of the earwig Chelidurella acanthopygia (Dermaptera) date back to the years of 1999–2002 at 10 study sites in monthly intervals from March to October (resp. to November). In the total of 285 samples 205 were positive on the earwig C. acanthopygia. Generally 1,484 specimens were collected. The Table 1 presents number of nymphs (N1 – N4), and adults obtained at the study sites during the whole period. The material has been deposed at the Department of Zoology, Faculty of Natural Sciences, Comenius University in Bratislava. T a b l e 1. Number of Chelidurella acanthopygia specimens collected during 1999–2002 at the study sites. N1 N2 N3 N4 F M Total PS

CA 0 12 51 22 35 21 141 21

VI 18 42 40 26 10 9 145 21

FU 0 43 20 5 17 13 98 18

LI 4 77 17 37 30 27 192 17

HH 4 49 48 26 29 26 182 20

LH 7 41 20 29 25 30 152 23

LL 0 11 11 5 11 11 49 12

NA 1 49 17 27 37 16 147 23

NK 1 9 64 27 37 47 35 219 27

NK 2 12 35 39 32 21 20 159 23

Total 55 423 290 246 262 208 1484 205

Explications: Abbreviations of study sites (CA → NK 2) see text; N 1 → N 4 – first → fourth nymphal instar; F – female; M – male; PS – Number of positive samples with Chelidurella acanthopygia To sample the square method was used. The leaf litter was sieved on the area of 1 m2 (16 squares of 25x25 cm). The samples were transported in textile bags into a laboratory. The earwigs were extracted in the Moczarski-Winkler’s eclectors (Holdhaus, 1910; Balogh, 1958), and stored in 75% ethyl alcohol. C. acanthopygia was sampled at all the study sites.

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The nymphal instars (N1, N2, N3, N4) were identified according to 5 characters: width of the head in its widest section behind compound eyes; length of the head, distance from the front margin of compound eyes to the rare margin of the head (vertex); width of pronotum; length of pronotum (Fig. 1) and number of antennal segments. All character values and numbers of the measured specimens are presented in the Table 2. T a b l e 2. Width and length of head and pronotum in mm and number of antennal segments of Chelidurella acanthopygia.

WH

LH WP LP AS

1 2 3 1 2 3 1 2 3 1 2 3

N1 0.79 – 0.90 0.88 ± 0.03 n = 39 0.40 – 0.53 0.45 ± 0.04 n = 39 0.69 – 0.79 0.75 ± 0.03 n = 39 0.42 – 0.53 0.45 ± 0.03 n = 39 8

N2 0.98 – 1.14 1.05 ± 0.04 n = 36 0.48 – 0.66 0.57 ± 0.04 n = 36 0.82 – 1.03 0.94 ± 0.04 n = 36 0.53 – 0.71 0.61 ± 0.04 n = 36 10

N3 1.24 – 1.37 1.31 ± 0.03 n = 32 0.69 – 0.85 0.76 ± 0.04 n = 32 1.16 – 1.35 1.24 ± 0.05 n = 32 0.77 – 0.92 0.85 ± 0.05 n = 32 11

N4 1.48 – 1.72 1.58 ± 0.06 n = 35 0.85 – 1.06 0.96 ± 0.05 n = 35 1.48 – 1.69 1.58 ± 0.06 n = 35 0.98 – 1.27 1.15 ± 0.06 n = 35 12

F 1.77 – 2.12 1.97 ± 0.09 n = 25 1.11 – 1.35 1.20 ± 0.06 n = 25 1.74 – 2.22 1.99 ± 0.11 n = 25 1.40 – 1.70 1.54 ± 0.09 n = 25 13

M 1.74 – 2.04 1.91 ± 0.07 n = 36 1.00 –1.35 1.17 ± 0.08 n = 36 1.67 – 2.04 1.85 ± 0.09 n = 36 1.27 – 1.59 1.42 ± 0.07 n = 36 13

Explications: WH – width of head; LH – lenght of head; WP – width of pronotum; LP – length of pronotum; 1 – range (mm); 2 – average ± SD (mm); 3 – number of earwigs measured (n); AS – number of antennal segments; N 1 → N 4 – first → fourth nymphal instar; F – female; M – male All the measurements were realised by the stereomicroscope Stemi 2000 C (Carl Zeiss) with the help of eyepiece micrometer (WPL 16x, Carl Zeiss). The measured earwigs were enlighted by 6-point ringlight (Schott KL electronic 1500, Carl Zeiss), which provides the snap without shadows.

Results and discussion In the research period the earwig Chelidurella acanthopygia usually regularly occurred in the fallen leaf horizon samples in oak-hornbeam forests of the Malé Karpaty Mts and Trnavská pahorkatina hills. At the study sites its abundance varied between 1 and 33 ind.m-2, in most of the samples there were 9–18 specimens recorded from the area of 1 m2. The similar values (3–36 ind.m-2) from the beech forest in N Germany were published by Irmler and Hingst (1993). The study site Lošonec–lom quarry has been regularly impacted by limestone dust from the nearby quarry. This fact has probably reflected in the neutral value (pH = 6.74) of the upper soil horizon. The earwig C. acanthopygia occurred at the site sporadically, its abundance varied between 1 ind.m-2 and 6 ind.m-2. Exceptionally the sample from June 6, 2000 contained 4 ♀ and 6 ♂. In most of the study sites (pH = 3.90–5.00 in the upper soil horizon) the quantitative structure of the earwig was richer.

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First nymphal instar (N1) The first nymphal instar of Chelidurella acanthopygia occurred in the Malé Karpaty Mts from April (2nd decade) to June (end of the 2nd decade) (Table 3). The first specimens appeared in the sample from April 12, 2000 at the site NK1, the last ones in the sample from June 21, 1999 at the sites LH, NK1 and NK2. The first nymphal instar was present in the period of 70 days. Later N1 wasn’t recorded at any of the study sites in the period of 1999–2002. Kočárek (1998) found N1 even at the end of June in Litovelské Pomoraví, Czech Republic. In Germany Franke (1985), recorded N1 on S from Karlsruhe in May, in Denmark Nielsen (1991) app. 25 km on NE from the town of Århus in July (the highest quantity), in June and August (lower quantity). The N1 specimens from the Malé Karpaty Mts are well distinguished in comparison with older nymphs (characters presented in Table 2). Due to their small size (body length 5.5–6.5 mm including cerci) are not visible enough. They are weakly sclerotised, some of the collected specimens were damaged at sieving. T a b l e 3. Occurrence of adults and nymphal instars of Chelidurella acanthopygia in Malé Karpaty Mts and Trnavská pahorkatina hills. F March 8 – June 3 M March 8 – April 26

MAR APR MAY JUN XXX XXXX XXXX X

JUL

AUG

SEP

OCT

NOV

XXX XXXX

N1 April 12 – June 21 N2 May 9 – August 1 N3 June 18 – September 5 N4 July 1 – September 11

XXX XXXX XXX XXX XXXX XXXX X XX XXXX XXXX X XXXX XXXX XX

F August 28 – November 28

X XXXX XXXX XXXX

M August 15 – November 28

XX XXXX XXXX XXXX

Explications: F – female; M – male; N1 → N4 – nymphal instar; the 2nd column includes the 1st and last date of collection; X – projects one quarter on the month; XXXX – projects the whole month

Second nymphal instar (N2) The second nymphal instar occurred from the second decade of May to the end of July, for more than 80 days – the longest period of all the instars. The first N2 specimens appeared in the sample from May 9, 2001 in Vinosady (VI), the last ones were recorded at the site of Fúgelka in August 1, 2001. In the Czech Republic Kočárek (1998) recorded N2 at the end of June, in July and at the beginning of August 1995, in Germany Franke 156

(1985) in June, in Denmark Nielsen (1991) in July and August. The relevant characters are presented in the Table 2. In comparison with N1, N2 has 10 antennal segments. Third nymphal instar (N3) The 3rd nymphal instar is well-sclerotised. Alive specimens are brown to dark brown, being stored in alcohol they are gently pale. The first N3 specimens appeared in the samples in June (at the beginning of the 2nd half), on the 18th of June 2002 at the site NK2, the last ones were recorded in September 5, 2000 at LH. The period of their existence takes more than 75 days. In the Czech Republic N3 nymphs were observed from the end of June to the first decade of September (Kočárek, 1998), in Germany at the end of July and beginning of August (Franke, 1985), in Denmark (Nielsen, 1991) in most of the year, particularly in July, August and September. The relevant characters are presented in the Table 2. Fourth nymphal instar (N4) The 4th instar is formed by specimens with 12-segment antennae, immature cerci and character values included in the Table 2. The first specimens were collected in July 1, 1999 at the site VI, the last ones in September 9, 2001 at the sites CA, VI, FU, LI and HH. The whole existence period of the 4th instar takes more than 70 days. In the Czech Republic the specimens of were recorded in August and September (Kočárek, 1998), in Germany in the second half of August (Franke, 1985), in Denmark in most of the year, particularly in February, August and September (Nielsen, 1991). Adults (females and males) Adults occurred from the first sample at the beginning of the vegetation period (March). Females were recorded till June (1st decade), males till the end of April. Those were the overwintered specimens from the previous year. At the end of summer the first new generation females and males appeared in August. After wintertime they will become the parental generation for the next year. For the whole period of 1999–2002 adults were recorded in July only in 3 cases: July 1, 1999 (♀) in Lindava, July 6, 2000 (♀, ♂) in Lošonec–lom quarry and July 16, 2002 in Naháč–Katarínka l (♀). Most probably these are the specimens from the previous year, which spent wintertime as nymphs. A possibility to overwinter in a nymphal instar (N3, N4) of Chelidurella sp. has been mentioned by Kočárek et al. (2005).

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Conclusion Phenology of Chelidurella acanthopygia nymphal instars was studied at several sites in the Malé Karpaty Mts (Cajla, Vinosady, Fúgelka, Lošonský háj grove, Lošonec–lom quarry, Naháč–Kukovačník, Naháč–Katarínka 1 and Naháč–Katarínka 2) as well as in the Trnavská pahorkatina hills (Lindava and Horný háj) in the zone of oak-hornbeam forests. Being based on the material of 1,484 specimens of nymphs (N1–N4) and adults (♀, ♂) the chronological sequence of the C. acanthopygia ontogenesis was completed (Table 3). The whole development from the 1st instar to imago takes 100–120 days. In the Malé Karpaty Mts the earwig has one generation annually. It spends wintertime in a stage of imago. Few specimens overwinter in nymphal stage. Within the whole study period of 1999–2002 the imagines were recorded only in 3 July samples. The specimens probably survived winter as nymphs.

Fig. 1. Chelidurella acanthopygia – head with pronotum. LH – length of the head; WH – maximum width of the head; LP – length of the pronotum; WP – maximum width of the pronotum.

The paper projects diagnostic characters (Table 2), which allow quite exact identification of nymphal instars. The diagnostic characters include width and length of head and pronotum as well as number of antennal segments (Fig. 1). All the measured specimens (203) with their character values are presented in Table 2. Translated by P. J. Fedor

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Acknowledgement My thanks belong to M. Holecová and Z. Krumpálová for their help in field. The research was supported by the project VEGA No. 1/0119/03.

References Albouy, V., Caussanel, C., 1990: Dermaptčres ou Perce-Oreilles. Faune de France 75. Fédération Française des Sociétés de Sciences Naturelles, Paris, 245 pp. Balogh, J., 1958: Lebensgemeinschaften der Landtiere. Akadémiai Kiadó, Akademie-Verlag, Budapest, Berlin, 560 pp. Bey-Bienko, G. J., 1936: Insectes Dermaptères. Faune de L’URSS. N. S. 5 (in Russian, English summ.). Izdateľstvo Akademii Nauk SSSR, Moskva-Leningrad, 240 pp. Fedor, P.J., 2005: Ucholaky (Dermaptera) (in Slovak). In Majzlan, O. (ed.): Fauna Devínskej Kobyly. APOP, Bratislava, p. 64–65. Franke, U., 1985: Zur Biologie eines Buchenwaldbodens. 7. Der Waldohrwurm Chelidurella acanthopygia. Carolinea, 43, p. 105–112. Holdhaus, K., 1910: Die Siebetechnik zum Aufsammeln der Terricolfauna (nebst Bemerkungen über die Oekologie der im Erdboden lebenden Tierwelt). Z. f. wiss. Insektenbiologie, 6, p. 44–57. Irmler, U., Hingst, R., 1993: Zur Ökologie des Waldohrwurms (Chelidurella acanthopygia) in Schleswig-Holstein (Dermaptera). Faun.-Ökol. Mitt., 9/10, p. 377–390. Kočárek, P., 1998: Life cycles and habitat associations of three earwig (Dermaptera) species in lowland forest and its surroundings. Biologia, Bratislava, 53, p. 205–211. Kočárek, P., Galvagni, A., 2000: Species of Chelidurella (Dermaptera: Forficulidae) in the territory of the Czech Republic and Slovakia. Klapalekiana, 36, p. 89–92. Kočárek, P., Holuša, J., Vidlička, Ľ., 2005: Blattaria, Mantodea, Orthoptera & Dermaptera of the Czech and Slovak Republics. Kabourek, Zlín, 350 pp. Lukáš, J., 2003: Earwings (Dermaptera), snake-flies (Raphidioptera) and scorpion-flies (Mecoptera) of the National Nature Reserve Devínska Kobyla (in Slovak). Biosozologia, 1, p. 39–43. Nielsen, O.B., 1991: Seasonal development of the woodland earwig (Chelidurella acanthopygia Géné) in Denmark (Dermaptera). Ent. Medd., 59, p. 91–98. Petricskó, J., 1892: Fauna of Banská Štiavnica and its environs (in Hungarian). Selmeczbányai Gyógyászati és Természettudományi Egyesület, Selmeczbánya (= Banská Štiavnica), 133 pp. Petrogalli, A., 1890: Excursion to Sitno Mt. (in Hungarian). Jahresheft des Naturwissenschaftlichen Vereines des Trencséner Comitates, 11–12 (1888-1889), p. 132–146. Straka, V., 2005: Blattaria, Mantodea, Orthoptera and Dermaptera collected in Moericke traps in the Trenčiansky kraj region (in Slovak). Naturae Tutela, 9, p. 59–70. Turček, F.J., 1974: Detritus consumption and some ecological remarks of the earwigs Anechura bipunctata (Fabr.) and Chelidurella acanthopygia (Géné) in mat-grass pastures. Biologia, Bratislava, 29, p. 631–635. Zlinská, J., Šomšák, L., Holecová, M., 2005: Ecological characteristics of studied forest communities of an oak-hornbeam tier in SW Slovakia. Ekológia (Bratislava), 24, Suppl. 2, p. 3–19. Received 30. 7. 2005

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Országh I.: Fenológia nymfálnych instarov ucholaka Chelidurella acanthopygia (Dermaptera: Forficulidae) v dubovo-hrabových lesoch Malých Karpát a Trnavskej pahorkatiny (JZ Slovensko). V období rokov 1999–2002 sme v Malých Karpatoch a neďalekej Trnavskej pahorkatine sledovali fenológiu larválnych instarov ucholaka Chelidurella acanthopygia. Abundancia druhu sa na sledovaných plochách pohybovala v rozmedzí hodnôt 1–33 jedincov na m2. Prvé jedince, imága oboch pohlaví, z predchádzajúceho roku sa po prezimovaní objavili vo vzorkách už v marci. Prvé nymfálne instary (N1) boli zbierané od druhej dekády apríla do konca druhej dekády júna. Obdobie ich výskytu trvalo približne 70 dní. Jedince druhého instaru (N2) sa objavovali od mája (2. dekáda) do konca júla s obdobím výskytu viac ako 80 dní. Tretí instar (N3) bol zaznamenaný počas obdobia viac ako 75 dní od júna (začiatok 2. polovice) do 1. týždňa septembra. Štvrtý instar (N4) bol prítomný od začiatku júla do septembra (1. dekáda) s obdobím výskytu viac ako 70 dní. Nová generácia dospelých jedincov (samice a samce) sa objavila na konci leta (august). Táto po prezimovaní utvorí v nasledujúcom roku novú rodičovskú generáciu. Ziskané údaje o instaroch N1 až N4 a dĺžke obdobia ich výskytu v Malých Karpatoch potvrdzujú, že vývin Chelidurella acanthopygia od prvého nymfálneho instaru po imago trvá 100–120 dní.

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Vol. 24, Supplement 2/2005, p. 143–151

MILLIPEDE COMMUNITIES (Diplopoda) OF OAK-HORNBEAM ECOSYSTEMS (THE MALÉ KARPATY MTS, TRNAVSKÁ PAHORKATINA HILLS, SW SLOVAKIA) SLAVOMÍR STAŠIOV Department of Biology and General Ecology, Faculty of Ecology and Environmental Sciences, Technical University, T. G. Masaryka 24, 960 53 Zvolen , The Slovak Republic e-mail: [email protected] Abstract Stašiov S.: Millipedes communities (Diplopoda) of oak-hornbeam ecosystems (the Malé Karpaty Mts, Trnavská pahorkatina hills, SW Slovakia). Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 143–151. Millipede communities were studied in ten oak-hornbeam forest localities in the central and northern part of the Malé Karpaty Mts and the Trnavská pahorkatina hills. The research was conducted during the years 1999–2002. Millipedes were sampled using a dry sieve method from the leaf litter within each locality. In total, 3,654 individuals belonging to 18 millipede species were recorded. The records of Julus curvicornis are of the highest faunistic importance. It is an endemic species of the West Carpathian region. The westernmost locality of its occurrence was found in the Malé Karpaty Mts. Influence of selected environmental factors on the structure of millipedes communities was discussed. Key words: Diplopoda, millipedes, Malé Karpaty Mts, Trnavská pahorkatina hills, Slovakia

Introduction The Malé Karpaty Mts are relatively well-known area of Slovakia concerning the millipede fauna. Data about the millipedes in these mountains were subsequently published by Ortvay (1902), Lang (1933, 1954), Gulička (1955, 1956, 1986), Mišík et al. (1974), Krumpál (1993) and Mock, Janský (2000). Gulička (1986) presented the most comprehensive study with checklist of 30 millipede species from forest communities in the Malé Karpaty Mts Concerning the Trnavská pahorkatina hills, no data about the millipede fauna have been published up to present. This paper presents the results of the millipede research realized at ten oak-hornbeam forest localities situated in the central and the northern part of the Malé Karpaty Mts and the Trnavská pahorkatina hills. 143

Material and methods The research was conducted during four growing seasons (1999–2002) in 10 oak-hornbeam forest localities situated in the central and the northern part of the Malé Karpaty Mts (Cajla, Fúgelka, Lošonský háj grove, Lošonec–lom quarry, Naháč–Katarínka 1, Naháč–Katarínka 2, Naháč–Kukovačník, Vinosady) and in the adjacent Trnavská pahorkatina hills (Horný háj grove, Lindava). This area is situated in western part of Slovakia between the towns Bratislava and Nové Mesto nad Váhom. The altitude varies from 132 m a.s.l. to 768 m a.s.l. These mountains belong to the gently warm climatic zone. From the phytogeographic point of view the mountains belong to the area of West Carpathian flora with dominating forest stands. From the zoogeographic point of view the mountains have been classified into the Subcarpathian district of the deciduous forest province in Eurosiberian subprovince. The studied localities differed in the age of forest growth, anthropic impact (contamination by calcareous dust), fragmentation (isolated forests localities in the cultural landscape and non-fragmented forest complexes) and also in phytocoenological and pedological parameters. The detailed ecological characteristics of studied localities areas (pedobiology, phytocoenology and climatic conditions) are available in the introduction work of this supplement (Zlinská et al., 2005). The following list of studied localities involves their acronyms and selected characteristics: geographic coordinates; altitude; exposition; slope; subassociation of potential vegetation; age of forest growth; species composition of forest. Cajla (CA) { 48°20' N, 17°16' E; 260–280 m a.s.l.; S; 7°; Querco-Carpinetum poetosum nemoralis; 80–100 years; Carpinus betulus, Quercus dalechampii. Vinosady (VI) { 48°19' N, 17°17' E; 280 m a.s.l.; NW; 5–7°; Querco-Carpinetum poetosum nemoralis; 60–80 years; Acer campestre, Quercus cerris, Q. dalechampii. Fúgelka (FU) { 48°22' N, 17°19' E; 350 m a.s.l.; S; 4°; Querco-Carpinetum melicetosum uniflorae; 80–100 years; Quercus dalechampii. Lindava (LI) { 48°22' N, 17°22' E; 240 m a.s.l.; without exposition; 0°; Quercetum petrae-cerris; 80–100 years; Quercus cerris, Q. dalechampii. Horný háj grove (HH) { 48°29' N, 17°27' E; 240 m a.s.l.; W-SW; 5°; Querco-Carpinetum melicetosum uniflorae; 60–80 years; Carpinus betulus, Fraxinus excelsior, Quercus cerris, Quercus dalechampii. Lošonec–lom quarry (LL) { 48°29' N, 17°23' E; 340 m a.s.l.; SW; 8–10°; Querco-Carpinetum caricetosum pilosae; 80–100 years; Quercus dalechampii. Lošonský háj grove (LH) { 48°28' N, 17°24' E; 260 m a.s.l.; N; 1°; Querco-Carpinetum caricetosum pilosae; 80–100 years; Carpinus betulus, Quercus cerris, Q. dalechampii. Naháč–Kukovačník (NA) { 48°32' N, 17°31' E; 300 m a.s.l.; NE; 2–3°; Querco petrae-Carpinetum melicetosum uniflorae; 40–60 years; Carpinus betulus, Quercus cerris, Q. dalechampii. Naháč–Katarínka 1 (NK1) { 48°33' N, 17°33' E; 340 m a.s.l.; NW; 3–5°; Querco petrae-Carpinetum melicetosum uniflorae; 40–60 years; Carpinus betulus, Quercus dalechampii. Naháč–Katarínka 2 (NK2) { 48°33' N, 17°32' E; 320 m a.s.l.; SE; 45°; Lithospermo-Quercetum virgilianae; 80–100 years;, Acer campestre, Quercus cerris, Q. virgiliana, Tilia cordata.

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Material of millipedes was sampled using a dry sieve method (Wallwork, 1970) from leaf litter. At each locality the sample of leaf litter was taken as 16 random quadrates, each of size 25x25 cm. Thus, in total 1m2 of leaf litter was sampled monthly from each study plot. Millipedes were extracted in laboratory from the sieved leaf litter material using a dry extraction in Tullgren xereclectors. The obtained animals were fixed in 70 % ethanol. The material of millipedes is deposed in the collection of author. The similarity of localities and species was evaluated by hierarchical clustering analyses and by Principal Components Analysis (PCA) based on the mean abundance calculated for all years of investigation. Logarithm transformation (log n+1, n = number of captured individuals) was used before data analysis. Dendrograms of cluster analysis were produced using the software STATISTICA for Windows 5.1. (StatSoft, Inc., 1999) with Complete Linkage and Percent Disagreement clustering algorithm. PCA was carried out on log-transformed millipedes abundance of the different study localities using the software Canoco (ter Braak, Šmilauer, 1998). Scaling was focussed on inter-species correlations. Species scores were divided by the standard deviation, and the data were centred by species. The index of species diversity (H´) and the index of species equitability (E) were calculated according to Odum (1971) using ln logarithm.

Results In total 3,654 individuals of millipedes were obtained during the whole research. Among them 3,038 individuals were determined on the species level, the remaining 616 undetermined individuals involved juveniles and females. In total 18 species from 8 families and 5 orders were recorded from the studied localities (Table 1). The highest species number (14) was found in the localities Horný háj grove (HH) and Naháč–Kukovačník 2 (NK2) (Table 1). The locality Fúgelka (FU) with 8 recorded species and the localities Cajla (CA), Lindava (LI), Lošonec–lom quarry (LL), Vinosady (VI), all with 9 species only were the poorest in species richness. The millipedes Glomeris hexasticha, Cylindroiulus boleti and Ommatoiulus sabulosus were the most frequent species occurring at all studied localities. The species Trachysphaera costata (recorded only in one locality), Glomeris connexa and Polyzonium germanicum (both recorded in 2 localities) were the rarest representatives of these invertebrates in the studied area. In total, the most abundant species were Cylindroiulus boleti, Strongylosoma stigmatosum and Ommatoiulus sabulosus with 690, 456 and 424 sampled individuals, respectively. The lowest numbers of individuals were recorded for Trachysphaera costata (1 specimen), Polyzonium germanicum (5 specimens) and Megaphyllum unilineatum (27 individuals). The highest values of species diversity (H) were recorded at the localities Horný háj grove (HH), Cajla (CA) and Lošonec–lom quarry (LL) (Table 1). The lowest value of this index was recorded at the locality Fúgelka (FU). The highest values of equitability index (E) were recorded at the localities Lošonec–lom quarry (LL), Vinosady (VI) and Lindava (LI). The lowest species equitability was recorded at the locality Fúgelka (FU) (Table 1).

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T a b l e 1. Composition of the millipede communities at the studied localities, the summarizing data from 1999–2002: Species spectrum, total number of individual (Σ), Shannon´s index of species diversity (H’), index of species equitability (E). Taxon Glomerida Glomeris connexa C. L. K o c h, 1847 Glomeris hexasticha B r a n d t, 1833 Trachysphaera costata (W a g a, 1857) Polyzoniida Polyzonium germanicum B r a n d t, 1837 Julida Cylindroiulus boleti (C. L. K o c h, 1847) Enantiulus nanus (L a t z e l, 1884) Julus curvicornis V e r h o e f f, 1899 Kryphioiulus occultus (C. L. K o c h, 1847) Leptoiulus proximus (N e m e c, 1896) Megaphyllum projectum (V e r h o e f f, 1894) Megaphyllum unilineatum (C. L. K o c h, 1838) Ommatoiulus sabulosus (L i n n a e u s, 1758) Unciger foetidus (C. L. K o c h, 1838) Unciger transsilvanicus (V e r h o e f f, 1899) Chordeumatida Haploporatia eremita (V e r h o e f f, 1909) Craspedosoma rawlinsii L e a c h, 1815 Polydesmida Strongylosoma stigmatosum (E i c h w a l d, 1830) Polydesmus complanatus (L i n n a e u s, 1761) Indet. spp. Σ Individuals Σ Species H´ E

CA

FU

HH

Locality LH LL

LI

NK1

NK2

NA

Σ

VI

4 -

2 -

2 3 -

35 -

5 -

11 -

12 1

7 -

14 -

6 2 -

8 95 1

62 26 6 1 59 27 89

3 71 1 9 -

98 65 23 155 12 7 117 10 18

63 6 40 82 2 1

81 2 56 1 26 2 1

42 1 26 5 65 6

1 34 49 3 13 -

4 116 2 151 20 11 0 5 1 28 -

124 150 3 4 4

67 10 3 67 38

5 690 69 350 50 395 27 13 424 95 157

19

3 -

8 -

2 -

47 5

14 8

6 -

11 3

13 1

29

104 65

6 299 9 1.78 0.46

3 49 141 8 0.89 0.43

3 1 33 555 14 1.9 0.72

9 90 330 9 1.62 0.74

1 42 269 11 1.33 0.55

43 221 9 1.74 0.79

2 5 72 198 11 1.68 0.7

421 8 127 915 14 1.45 0.55

30 3 140 486 10 1.4 0.61

4 14 240 9 1.69 0.77

456 34 616 3654 18 -

The abbreviations of the localities: CA – Cajla, FU – Fúgelka, HH – Horný háj grove, LI – Lindava, LH – Lošonecký háj grove, LL – Lošonec–lom quarry (LL), NK1 – Naháč–Katarínka 1, NK2 – Naháč–Katarínka 2, NA – Naháč–Kukovačník, VI – Vinosady

Cluster analysis divided localities into two main clusters (Fig. 1). The first of them included the localities Naháč–Katarínka 2 (NK2), Naháč–Kukovačník (NA) and Naháč–Katarínka 1 (NK1). The second comprised the localities Horný háj grove (HH), Lošonský háj grove (LH), Lindava (LI), Fúgelka (FU), Vinosady (VI), Lošonec–lom quarry(LL) and Cajla (CA). Localities from the first cluster were characteristic by relatively low mean height of trees (reaching maximally up to 25 m), whereas within the localities from the second cluster the mean height of trees varied from 25 to 35 m. The most similar localities were Lindava and Fúgelka. In these two localities the lowest values of pH of the leaf litter and the highest values of sorption complex of leaf litter were recorded (Zlinská et al., 2005). PCA divided localities into four quadrates (Fig. 2). Two quadrates included only one locality, Horný háj grove (HH) and Naháč–Katarínka 2 (NK2), respectively. The third quadrate included two localities Naháč–Katarínka 1 (NK1) and Naháč–Kukovačník (NA). The other localities were situated in the fourth quadrate. The locality Horný háj grove (HH) differed from the others by the lowest value of maximum sorption capacity of leaf litter and the highest value of species diversity (Table 1). Typical species for this locality was Enantiulus nanus. The number of individuals of this species collected within this locality represented 94.2% of total catches. The locality Naháč–Katarínka 2 (NK2) was characterized by highest content of nitrogen in leaf litter, the highest values of sorption complex of leaf litter, maximum sorption capacity and maximum degree of sorption complex saturation of soil layers (mineral layer and leaf litter layer), the highest gradient of slope and the highest abundance of individuals. The dominant millipede species from this locality were Strongylosoma stigmatosum and Polyzonium germanicum. Two localities placed in the third quadrate were typical by the lowest content of carbon and humus in leaf litter and by the lowest mean age of the forest growth, lowest average height of trees and lowest diameter of stems measured in the breast height. Dominant millipede species were Trachysphaera costata and Polydesmus complanatus.

Discussion Relatively rich species composition was found out within the set of the studied plots. The number of recorded millipede species represents approximately a quarter of the Slovak millipede fauna. The species spectrum confirms well preserved natural conditions of the studied localities and corresponds with their location at the external border of the West Carpathians. The factors determining relatively rich and unique millipede communities in this area are relatively low elevation (not exceeding 1000 m), presence of large native forest complexes and deep valleys and narrow defiles with inverse microclimate. South-eastern (Unciger transsilvanicus, Strongylosoma stigmatosum), alpine (Haploporatia eremita, Haasea flavescens (L a t z e l, 1884)), Carpathian (Julus curvicornis) and also some common millipedes from the C. Europe occur here (Gulička, 1986). 147

Fig. 1. Cluster analysis of the similarity of localities.

Fig. 2. PCA ordination of millipede species and studied localities. The abbreviations of localities see Table 1. The first two PCA axes λ1 = 0.458, λ2 = 0.192) explained for 65.1% of the variance of the species data.

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The records of the Carpathian endemic species J. curvicornis are of the highest faunistic importance. It has been found in this region for the first time. The former westernmost locality of its occurrence was known from the Veľká Fatra Mts (Stašiov, 2002a). These new findings markedly shifted the known western border of its distribution and may indicate its possible occurrence even in the border mountain range between the Czech Republic and Slovakia. All other recorded species have been already known from the Malé Karpaty Mts, by several authors. The most complex list of the millipede fauna was summarized by Gulička (1986). Author accounted the occurrence of 30 species in this area. Mock, Janský (2000) enriched this list by Glomeris pustulata L a t r e i l l e, 1804. Therefore including Julus curvicornis, in total 32 millipede species are known from the Malé Karpaty Mts. The occurrence of 14 millipede species was recorded from the Trnavská pahorkatina hills (localities Lindava and Horný háj grove). The ascertained species list is therefore the first information about the millipede fauna of this area. Dominant occurrence of Strongylosoma stigomatosum, Polyzonium germanicum, Unciger foetidus and Haploporatia eremita in the locality Naháč–Katarínka 2 may indicate their preference of habitats with higher humus content. Mentioned locality was also featured by highest slope gradient and the occurrence of lime within the stand, which was not recorded from the other localities. Also these factors could have played deciding role in the shaping of millipede community structure of this locality. In contradiction, the species Trachysphaera costata, Polydesmus complanatus, and Glomeris hexasticha seem to be more tolerant to lower humus content in the soil, as indicate their dominant occurrence in the localities Naháč–Katarínka 1 and Naháč–Kukovačník. These species could be considered as more tolerant to humidity. The humidity is one of the main factors influencing the humification of detritus and the soil humus contents. The results of cluster analysis of the similarity of localities showed possible influence of the height of forest growth and also values of pH and sorption complex in leaf litter on the structure of the millipede communities. The tree height could have an indirect effect in the terms of higher litter production (bigger amount of suitable food source for saprophages) by taller trees. Several authors adverted on possible influence of litter properties on the structure of millipede communities. Branquart et al. (1995) studied the influence of environment on terrestrial isopod and millipede communities in oak forest ecosystems in southern Belgium. Authors found out close relationship between investigated communities and type and structure of leaf litter, respectively. Kime (1997) mentioned the potential of millipedes in the biomonitoring of environment properties. He furthermore adverted on the sensitive response of millipedes to the type of soil, form of leaf litter and various climate characteristics. Meyer, Singer (1997) studied millipedes on 6 forest sites in western Austria. They recorded highest epigeic activity of millipedes in mixed forests with mull humus form, in contrast to conifer forests with moder humus form, where the lowest epigeic activity was recorded. Schaefer, Schauermann (1990) investigated the influence of leaf litter on structure and dynamics of soil fauna, including millipedes. They conducted their research in 2 beech forests near Göttingen, Germany. The authors recorded markedly higher biomass of the genus Glomeris within the sites with mull humus from. Stašiov (2002b) studied the 149

influence of the leaf litter pH on millipedes. He conducted his research in 4 beech forest stands in the Kremnické vrchy Mts. He recorded a negative correlation between pH and the epigeic activity of Polyzonium germanicum, Julus curvicornis and Mastigona vihorlatica (A t t e m s, 1899). This correlation may testify the preference of acid detritus by mentioned species. The author has already adverted to positive correlation between values of soil pH and epigeic activity of Polydesmus complanatus. The results from PCA analysis (the allocation of localities and high-affinity species) may indicate possible influence of other investigated factors. Among them the values of sorption complex, maximum sorption capacity of leaf litter, carbon, nitrogen and humus content in the leaf litter, stand composition, age and height of trees and slope inclination, respectively may have important influence on the cenological parameters of millipede communities. The extent of the research did not enable proper evaluation of the effect of mentioned factors on the structure of the millipede communities. The mentioned factors probably have specific effect on different millipede species. It is still unclear whether the preference of particular biotopes by certain species is influenced strictly only by environment conditions evaluated during this research. Assumed effect of these factors should be topical for future research. Translated by the author Acknowledgement Author thanks A..Mock for help, comments and advices regarding the millipede research in investigated region and M..Svitok for help during the analytical data processing. The research was funded by VEGA grant project No. 1/0119/03.

References Branquart, É., Kime, R.,D., Dufrêne, M., Tavernier, J., 1995: Macroarthropod-habitat relationships in oak forest in South Belgium. 1. Environments and communities. Pedobiologia, 39, p. 243–263. Gulička, J., 1955: Two new species of Diplopoda for Czechoslovakia (in Slovak). Biológia, Bratislava, 10, p..367–370. Gulička, J., 1956: Two Alpines species of Diplopoda in Slovakia (in Slovak). Acta Fac. Rerum. Nat. Univ. Comenianae, Zool., 1, p. 79–88. Gulička, J., 1986: Diplopoda of forest communities of the Little Carpathians. In Nosek, J. (ed.): The soil fauna of the Little Carpathians. Results of Research program MAB, Ústav experimentálnej biológie a ekológie SAV, Bratislava, p. 217–224. Kime, R.D., 1997: Biodiversity and land use with regard to diplopods on some west-European sites. Proccedings of the 10th Int. EIS-Coll., 1995, Saarbrücken, p. 75–82. Krumpál, M., 1993: Invertebrates (in Slovak). In Bertová, L. (ed.): Karlova Ves. Vlastivedná monografia. Alfa, Bratislava, p. 57–61. Lang, J., 1933: Contribution to the knowledge of of Czechoslovakian Diplopoda (in Czech). Věst. Král. Spol. Nauk, 2, p. 1–32. Lang, J., 1954: Millipedes – Diplopoda (in Czech). Fauna ČSR 2. Nakladatelství ČSAV, Praha, 187 pp. Meyer, E., Singer, A., 1997: Verteilung, Aktivität und Besiedlungsdichte von Diplopoden in Wäldern Vorarlbergs (Österreich). Ber. Nat.-Med. Verein Innsbruck, 84,: p. 287–306.

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Mišík, M., Gulička, J., Urvichiarová, E., 1974: Devínska Kobyla. Geological conditions, flora and fauna (in Slovak). Obzor, Bratislava, 107 pp. Mock A., Janský V., 2000: Millipedes (Diplopoda) from Slovakia in the collections of the Slovak National Museum in Bratislava (in Slovak). Acta Rer. Natur. Mus. Nat. Slov., 46, p. 3–10. Odum, E.P. 1971. Fundamentals of Ecology, 3rd ed. W. B. Saunders, Philadelphia, 574 pp. Ortvay, T., 1902: Fauna of Bratislava region and of Bratislava, Trnava, Pezinok, Modra, Svätý Jur cities situated in this region (in Hungarian). Kiadja Pozsonyváros anyagi hozzájárulásával Pozsonyvármegye közönsége. Pozsony (= Bratislava), XVI + 648 pp. Schaefer, M., Schauermann, J., 1990: The soil fauna of beech forests: comparison between a mull and moder soil. Pedobiologia, 34, p. 299–314. Stašiov, S., 2002a: Millipedes (Diplopoda) of Blatnická dolina valley (Veľká Fatra NP, Slovakia) (in Slovak). Matthias Belivs Univ Proc, Banská Bystrica, 2, p. 123–133. Stašiov, S., 2002b: Selected groups of epigeic macrofauna (Opilionida, Diplopoda and Chilopoda) as indicators of the status of the top soil layer in submontane beech forest (in Slovak). Scientific Studies. Technická univerzita vo Zvolene, Zvolen, 88 pp. StatSoft, Inc., 1999: Statistica for Windows (Computer program manual). Tulsa. ter Braak, C.J.F., Šmilauer, P., 1998: CANOCO Reference Manual and User’s Guide to Canoco for Windows. Software for Canonical Community Ordination (version 4). Centre of Biometry, Wageningen. Wallwork, J.A., 1970: Ecology of soil animals. McGraw-Hill Publishing Company Ltd., London, 284 pp. Zlinská, J., Šomšák, L., Holecová, M., 2005: Ecological characteristics of studied forest communities of an oak-hornbeam tier in SW Slovakia. Ekológia (Bratislava), 24, Suppl. 2, p. 3–19. Received 30. 7. 2005

Stašiov S.: Mnohonôžky (Diplopoda) v dubovo-hrabových ekosystémoch (Malé Karpaty, Trnavská pahorkatina, JZ Slovensko). Práca prináša výsledky výskumu zameraného na posúdenie vplyvu viacerých sledovaných faktorov a podmienok prostredia na štruktúru taxocenóz mnohonôžok vo vybraných dubovo-hrabových ekosystémoch situovaných v strednej a severnej časti Malých Karpát a v Trnavskej pahorkatine. Výskum sme bol realizovaný v rokoch 1999 až 2002 kvadrátovou metódou, preosievaním listového opadu a následnou extrakciou živočíchov v.Tullgrenových xeroeklektoroch. Celkovo sme na skúmanom území získali 3654 ex. mnohonôžok z.18 druhov z 5 radov. K najvýznamnejším patrí nález západokarpatského endemita Julus curvicornis V e r h o e f f, 1899, u ktorého predstavujú Malé Karpaty územie s doposiaľ najzápadnejším zaznamenaným výskytom v.rámci jeho areálu. Výsledky výskumu poukázali na možný vplyv viacerých faktorov (priemerný vek a výška porastu, priemerná hrúbka stromov, drevinová skladba porastu, subasociácia potenciálnej vegetácie, sklon svahu, pH, hodnota sorpčného komplexu, maximálna sorpčná kapacita, obsah uhlíka, humusu a dusíka v.pokrývkovom humuse, stupňa nasýtenia sorpčného komplexu a obsah humusu v.pokrývkovom humuse i v.minerálnej pôdnej vrstve) na kvalitatívno-kvantitatívnu štruktúru taxocenóz mnohonôžok v podmienkach dubovo-hrabových lesov.

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Vol. 24, Supplement 2/2005, p. 124–142

STRUCTURE OF CENTIPEDE COMMUNITIES (Myriapoda: Chilopoda) IN OAK-HORNBEAM FORESTS OF THE MALÉ KARPATY MTS AND TRNAVSKÁ PAHORKATINA HILLS (SW SLOVAKIA) IVAN ORSZÁGH, ZLATICA ORSZÁGHOVÁ Department of Zoology, Comenius University, Mlynská dolina B-1, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected] Abstract Országh I., Országhová Z.: Structure of centipede communities (Myriapoda: Chilopoda) in oak-hornbeam forests of the Malé Karpaty Mts and Trnavská pahorkatina hills (SW Slovakia). Ekológia (Bratislava), Vol. 24, Suppl. 2/2005, p. 124–142. At 8 sites in the Malé Karpaty Mts and 2 sites in the nearby Trnavská pahorkatina hills the taxocoenoses of centipedes (Chilopoda) were studied in the period of 1999 to 2002. The sites have been covered by 40 to 100 year old oak-hornbeam forests. They are situated at the altitude of 240–350 m a.s.l. The samples have recorded the occurrence of Cryptops anomalans, Henia illyrica, Clinopodes flavidus, Geophilus flavus, Strigamia acuminata, S. transsilvanica, Schendyla nemorensis, Lithobius agilis, L. borealis, L. dentatus, L. erythrocephalus, L. forficatus, L. lapidicola, L. latro, L. lucifugus, L. melanops, L. mutabilis, L. muticus, L. pelidnus, L. piceus, L. aeruginosus, L. austriacus, L. crassipes and L. microps. The highest species richness refers to the site of Lošonský háj grove (17), the lowest diversity was recorded in Vinosady (10). The species diversity index (H') and equitability (e) reached their highest value in the community in Lošonský háj grove. Strigamia acuminata had the abundance (12.53 ind.m-2) at the site of Naháč–Katarínka 2. The most abundant taxa have included Schendyla nemorensis (at Naháč–Katarínka 1–10.16 ind.m-2, Horný háj grove – 9.6 ind.m-2) and Lithobius austriacus (at the site of Naháč–Kukovačník – 8.45 ind.m-2). Schendyla nemorensis and Lithobius muticus may be classified as eudominant species at all the study sites, their dominance was in average 40%. In all the dendrograms of cluster analyses the community at the site of Vinosady is separated from the other ones on the highest level of dissimilarity. The reason lies in the lowest number of determined species, a unique portion of Cryptops anomalans, absence of Strigamia acuminata and the lowest number of obtained specimens per 4 years. As for the species identity the centipede communities at the sites Lošonský háj grove and Naháč–Katarínka 1 may be classified as the most similar on the 87% level of similarity. The communities at Fúgelka and Horný háj grove are close to each other (92% similarity) from the quantitative-qualitative point of view. Most of the recorded species occured throughout the year. Strigamia acuminata and Lithobius muticus were recorded continuously from March to November at most of the study sites. Schendyla nemorensis, present at all the sites, was often excluded from the species richness in the summertime samples. The typical centipede community in the oak-hornbeam forests of the Malé Karpaty Mts consists of the species: Schendyla nemorensis, Strigamia acuminata, Lithobius agilis, L. borealis, L. lapidicola, L.

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mutabilis, L. muticus, L. austriacus; in more southern drier parts of the Malé Karpaty Mts Henia illyrica as well. Key words: Chilopoda, centipedes, species diversity, equitability, dominance, abundance, Malé Karpaty Mts, Trnavská pahorkatina hills, SW Slovakia

Introduction Centipedes play a significant role in stabilization of numerous terrestrial ecosystems. In C. Europe the species of the genus Lithobius have been well analysed from this point of view. Their ontogeny enables them to react promptly onto changes of environmental conditions. This actually originates in their continual egg lying, development during the whole year being proved by a high number of juveniles sampled at the study sites from March to November, slow ontogeny, late sexual maturation under unsuitable ecological conditions as well as an ability to starve for a long time. In C. Europe their portion in energetic flows approximately equals to 13% of predatory arthropods living in a forest fallen leaf horizon (Albert, 1983). Centipedes may be considered as predatory arthropods with a long lifespan. As K-strategists they play an important role in final interactions of trophic chains amongst organisms of oak-hornbeam forest epigeon. They are typical by low average density of species. According to our results density of the species in the Malé Karpaty Mts in 1999–2002 only rarely exceeded a value of 12 ind.m-2. The existing data declare that their distribution is determined by soil texture, depth of a leaf litter layer, ability of soil including a leaf litter layer to keep humidity, quantity of wood litter and presence of bigger stones on the soil surface. An influence of soil chemism on centipedes has not been sufficiently known. In fact it has a more significant effect on hypogeous species (Geophilomorpha) than the species active on the surface (Lithobiomorpha) (Dunger, 1983, 1993). In fact the centipedes of the Malé Karpaty Mts have not been studied in detail yet. The oldest record refers to Attems (1895), who mentioned the species Geophilus longicornis var. austriaca [= Geophilus flavus (D e G e e r, 1778)] from the Kamzík hill (Bratislava, southern part of the Malé Karpaty Mts). Data on Scutigera coleoptrata from the southern part of the Malé Karpaty Mts were published by plenty of authors in the first half of the 20th century, the knowledge on distribution of this species in Slovakia have been summarized by Országh (2001). Dobroruka (1966) published records on Pachymerium tristanicum [? = Geophilus flavus (D e G e e r, 1778)] from Svätý Jur. The species Strigamia acuminata (Zbojnícka jaskyňa cave) and Lithobius forficatus (Čachtická jaskyňa cave) were captured in caves, respectively in the Malé Karpaty Mts (Országh et al. 1994). The more complex information on centipede diversity in the northern part of the Malé Karpaty Mts have appeared in the paper by Országh (2002). Drdul (1997) analysed changing density of soil macrofauna during the vegetation period in the oak forests rounding the Nuclear Power Station Mochovce, on east from the Malé Karpaty Mts.

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Study area In 1976 the Malé Karpaty Mts were established as a Protected Landscape Area covering a total of 65,504 ha. They are situated in the western part of Slovakia lining from Bratislava towards the town of Nové Mesto nad Váhom on NE at the total length of 100 km. Its maximum broadness (16 km) appears between Lozorno and Sv. Jur near Bratislava, the minimum one (3 km) in the section of Čachtice – Prašník. The altitude ranges from 132 m a.s.l. (near the Danube river) to 768 m a.s.l. (the Záruby Mount). The main range of the mountains grows up to the altitude of 450–650 m a.s.l. the Malokarpatský kras karst is a specific area of these mountains, covering approx. 180 km2. The Malé Karpaty Mts belong to the gently warm climatic zone with an mean annual temperature of 7–9 °C. At the altitudes of over 400 m the temperature declines under 8.°C. In average the vegetation period (average daily temperature 10 °C) takes 180 days, at the higher altitudes under 160 days. The annual precipitations (in the Malé Karpaty Mts) reach 650–900 mm, in lowlands they decline onto 600–650 mm. From the phytogeographic point of view the mountains belong to the area of the West Carpathian flora with dominating forest stands, which have been replaced by vineyards on SE and NW and by bushy pastures, orchards and farmland on N. From the zoogeographic point of view the mountains have been classified into the Subcarpathian district of the deciduous forest province in the Eurosibirian subprovince. The province of steppes of the Pannonian district partially appears rounding the study area. The mountains are inhabited by colinous and submountainous elements, communities of forests, woody steppes and rocky biotopes. Thermophilous species infiltrate from lowlands into the S and SE slopes. The species diversity has been significantly influenced by warm Hungarian lowland.

Study sites The study sites are located in forest stands (Zlinská et al., 2005, Fig. 1). The sites Cajla (CA); Vinosady (VI); Fúgelka (FU); Lošonec–lom quarry (LL); Lošonský háj grove (LH); Naháč–Kukovačník (NA); Naháč–Katarínka 1, young forest (NK1) and Naháč–Katarínka 2, old forest (NK2) are situated in the zone of oak-hornbeam forests on SE slopes of the Malé Karpaty Mts. The sites Lindava (LI) and Horný háj grove (HH) are located in the boundary zone of the Malé Karpaty Mts and Trnavská pahorkatina hills in the zone of oak and oak-hornbeam forests with diverse degree of anthropogenous impact (deforestation, forest tracks, presence of heavy mechanisms, tractors, etc.). The altitude varies between 240 to 350 m a.s.l. The soils may be considered as acid with pH values of 3.8–5.0. However the sites Lošonec–lom quarry (LL) and Naháč–Katarínka 2, old forest (NK2) have a gently acid reaction, or rather neutral (pH 6.47–6.75). The distance between the first site – Cajla (CA) and the last and most northern one – Naháč–Katarínka 1, young forest (NK1) is 30 km (through the air line). The sites are situated in the zone of oak and oak-hornbeam forests with diverse degree of anthropogenous impact. 126

Fig. 1. Hierarchical classification of centipede communities in the forest epigaeon at the study sites according to their species presence/absence similarity (Sørensen’s index, complete linkage). Vertical axis = dissimilarity, abbreviations of study sites see text.

Cajla (CA) [Databank of the Slovak Fauna 7669c grid reference number] – 80 to 100 year old oak-hornbeam forest at the altitude of 260–280 m, with S exposition. Vinosady (VI) [7669d] – 60 to 80 year old oak-hornbeam forest at the altitude of 280 m, with NW exposition. Fúgelka (FU) [7669b] – 80 to 100 year old oak-hornbeam forest at the altitude of 350 m, with S exposition. Lindava (LI) [7670a] – 80 to 100 old oak forest in lowland at the altitude of 240 m. Horný háj grove (HH) [7570b] – 60 to 80 year old oak-hornbeam forest at the altitude of 240 m, with SW exposition. Lošonec–lom quarry (LL) [7570a] – 80 to 100 year old oak-hornbeam forest at the altitude of 340 m, with SW exposition. The site is regularly impacted by limestone dust imissions close to the quarry. Lošonský háj grove (LH) [7570b] – 80 to 100 year old oak-hornbeam forest at the altitude of 260 m, with N exposition and with a 15–20 cm deep fallen leaf horizon at many sites. Under the leaf litter the soil has a lumpy texture being humid in a dry year period too. Naháč–Kukovačník (NA) [7471c] – 40 to 60 year old oak-hornbeam forest at the altitude of 300 m, with NE exposition. The forest is completely surrounded by farmland. Naháč–Katarínka 1, young forest (NK1) [7471a] – 40 to 60 year old oak-hornbeam forest at the altitude of 340 m, with NW exposition. Naháč–Katarínka 2, old forest (NK2) [7471a] – 80 to 100 year old oak forest with lindens and maple trees at the altitude of 320 m, with SE exposition. 127

The map with the study sites and the more detailed pedological and botanical analyses of the study sites as well as the climatic conditions in the period of 1999–2002 are given in the paper Zlinská et al. (2005). The site Naháč–Katarínka 2 (NK2) has been characterized in detail in the paper by Drdulová, Zlatošová (1980).

Material and methods The centipede sampling refers to the years of 1999–2002 at 9 study sites, at the site Horný háj grove (HH) it has included 3 years (2000–2002). At all the sites the samples were collected in the monthly intervals form March till October, resp. November. The January samples 2000–2002 were excluded from the general evaluation. They served to confirm presence of juveniles and imagines in the winter period. In total the material includes 8,303 centipedes. 6,154 individuals of them have been determined (Table 1). The analyses exclude the damaged specimens. 1,843 juveniles and 306 damaged specimens were not determined. The material is deposed at the Department of Zoology, Faculty of Natural Sciences, Comenius University in Bratislava. T a b l e 1. Survey of the centipedes (Chilopoda) recorded. CA a Cryptops anomalans N e w p o r t, 1844 0 Henia illyrica (M e i n e r t, 1870) 0 Clinopodes flavidus C. L. K o c h, 1847 3 0 Geophilus flavus (D e G e e r, 1778) Strigamia acuminata (L e a c h, 1814) 94 Strigamia transsilvanica (V e r h o e f f, 1928) 3 Schendyla nemorensis (C. L. K o c h, 1836) 84 Lithobius agilis C. L. K o c h, 1847 6 Lithobius borealis M e i n e r t, 1868 4 Lithobius dentatus C. L. K o c h, 1844 0 Lithobius erythrocephalus C. L. K o c h, 1847 0 Lithobius forficatus (L i n n a e u s, 1758) 4 Lithobius lapidicola M e i n e r t, 1872 12 Lithobius latro M e i n e r t, 1872 0 Lithobius lucifugus L. K o c h, 1862 2 Lithobius melanops N e w p o r t, 1845 0 Lithobius mutabilis L. K o c h, 1862 10 Lithobius muticus C. L. K o c h, 1847 165 Lithobius pelidnus H a a s e, 1880 1 Lithobius piceus L. K o c h, 1862 0 Lithobius aeruginosus L. K o c h, 1862 0 Lithobius austriacus (V e r h o e f f, 1937) 3 Lithobius crassipes L. K o c h, 1862 3 Lithobius microps M e i n e r t, 1868 0 Lithobius sp. juven. 78 Lithobius sp. damaged 16 Total 488 Species

VI a 3 8 3 0 0 0 82 0 0 0 2 0 1 9 0 0 1 78 0 0 3 0 0 0 34 9 233

FU LI a a 0 0 0 0 2 5 26 2 120 47 2 0 90 130 39 0 8 14 0 0 3 33 6 3 8 6 2 3 0 0 0 0 110 25 121 164 0 0 0 0 0 0 0 202 0 28 0 0 214 336 56 29 807 1027

HH aa 0 0 7 0 22 5 144 17 0 0 0 2 0 0 0 0 25 101 9 0 6 20 12 0 45 23 438

LH a 0 0 1 4 96 4 151 30 8 0 12 27 13 0 0 3 33 114 5 0 0 36 1 1 155 33 727

LL NA a a 0 0 0 0 0 9 34 0 46 165 8 0 223 154 29 40 1 0 0 1 13 15 5 7 14 4 0 4 0 0 0 4 51 91 99 148 1 5 0 0 0 0 0 279 0 5 1 0 117 363 23 47 665 1341

NK 1 a 0 0 5 0 104 2 325 49 3 0 6 18 7 0 0 0 137 246 0 2 0 150 31 2 286 44 1417

NK 2 a 0 9 3 0 401 4 96 102 0 2 1 4 9 0 0 1 72 128 0 0 0 85 2 0 215 26 1160

Abbreviations of study sites (CA → NK 2) see chapter "Study sites"; a – a total of the specimens recorded in 1999–2002; aa – a total number of the specimens recorded in 2000–2002

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For collection of the samples of centipedes as well as mesostigmatid mites (Fenďa, Ciceková, 2005), pseudoscorpiones (Christophoryová, Krumpál, 2005), beetles of Curculionoidea (Holecová at al., 2005) a square method was used. The leaf litter was sieved from a total area of 1 m2 (16 squares of 25x25 cm). The samples were transported in the textile pockets into the laboratory. The centipedes were extracted in the Moczarski-Winkler’s eclectors (Holdhaus, 1910; Balogh, 1958) and have been preserved in 75% ethylalcohol. Generally 285 samples have been obtained (Table 2). Occasionally for a more detailed analysis of diagnostic characters the temporal slides were made in 40 % butyric acid. Five degrees have been used for dominance evaluation: eudominant, dominant, subdominant, recedent, subrecedent (Schwerdtfeger, 1975). Species diversity (diversity of communities) have been calculated according to the Shannon index (Odum, 1971; Spellerberg, Fedor, 2003). All the pairs of diversity values have been compared by the t-test (Poole, 1974). The cluster analysis of the centipede community refers to the NCLAS program (Podani, 1993). The cluster method of complete linkage have been based on Sřrensen’s and Wishart’s indices. T a b l e 2. Number of elaborated samples at individual study sites in 1999–2002. 1999 2000 2001 2002 Total

CA 7 6 7 8 28

VI 4 6 7 6 23

FU 7 8 8 10 33

LI 8 5 8 9 30

HH 0 4 6 5 15

LH 8 8 4 8 28

LL 7 7 9 8 31

NA 8 8 8 9 33

NK 1 8 8 8 8 32

NK 2 8 8 8 8 32

Total 65 68 73 79 285

Abbreviations of study sites (CA → NK 2) see text The values of species data used in the cluster analysis (Wishart similarity ratio) were transformed with log-trans-formation Y' = log(Y+1), which increase the relative rate of rare species that are mostly good indicators.

Results and discussion Species diversity, equitability and species diversity test In the period 1999–2002 the method of leaf litter sieving at 10 sites in the Malé Karpaty Mts led to record 24 centipede species (Table 1) of the families Cryptopidae, Dignathodontidae, Geophilidae, Linotaeniidae, Schendylidae and Lithobiidae. The family of Lithobiidae (17 of the total 24 taxa) may be classified as the most represented. The same fact was published by Stašiov (2002) from the Kremnické vrchy Mts, the Kováčovská dolina valley (15 lithobiid species of the total 27 taxa); by Tajovský (2000) from the Krkonoše Mts, the Czech Republic (in total 15, Lithobiidae 11); by Wytwer (1990) from the Mazovian lowland, Poland (in total 17, Lithobiidae 12); and by Karafiat (1970) from the surroundings of Darmstadt, Germany (in total 15, Lithobiidae 8). In the southern parts of Europe the portion of Lithobiidae appears to be lower; Loksa (1966) presented 13 species (6 of them Lithobiidae) from the Bükk Mts, Hungary; Minelli (1982) recorded 23 species (9 of them Lithobiidae) from northeast Italy, and Zapparoli (1992) 22 taxa (9 of them Lithobiidae) from Central Italy. Mean number of species per one study site in the Malé Karpaty Mts grows up to 13.7. A higher value (16.75) was declared by Stašiov (2002) in the forest community of 129

T a b l e 3. Species diversity test (Poole, 1974) and main coenological variables of the centipede communities at the study sites in 1999–2002. Study site CA VI FU LI HH LH LL NA NK 1 NK 2 No. sp. MA [ind.m-2] e c H'

CA 0.000 2.923** 4.849*** 4.062*** 2.892** 7.071*** 2.794** 5.141*** 5.226*** 2.361* 14 14.1 0.600 0.280 1.583

VI 383.729 0.000 6.995*** 6.384*** 5.404*** 8.715*** 5.211*** 7.231*** 7.298*** 5.103*** 10 8.6 0.567 0.360 1.306

FU 692.378 271.273 0.000 0.979ns 1.973ns 3.175** 1.503ns 0.073ns 0.075ns 3.436*** 13 16.3 0.740 0.179 1.899

LI 723.332 277.205 1192.238 0.000 1.093ns 4.025*** 0.746ns 1.136ns 1.171ns 2.375* 13 22.1 0.721 0.204 1.850

HH 814.925 323.290 1000.489 1072.760 0.000 4.661*** 0.187ns 2.168* 2.218* 0.963ns 12 24.7 0.697 0.242 1.787

LH 789.531 308.669 1042.413 1126.817 1058.675 0.000 3.914*** 3.341** 3.423*** 6.343*** 17 19.3 0.731 0.170 2.072

LL 763.070 382.173 652.737 681.378 774.936 748.594 0.000 1.629ns 1.656ns 1.034ns 13 16.9 0.725 0.242 1.801

Abbreviations of study sites (CA → NK 2) see text; t-test values under the diagonal, degrees of freedom above diagonal. Significance levels: *** = P < 0.001; ** = 0.001 < P < 0.01; * = 0.01 < P < 0.05; ns = 0.05 < P (non-significant); No. sp. – total number of species; MA [ind.m-2] – mean abundance; e – Pielou's index of equitability; c – Simpson's index of species dominance; H’ – Shannon’s index of species diversity

NA 619.603 245.378 1197.151 1380.397 963.152 1034.353 581.452 0.000 0.000ns 3.844*** 15 28.2 0.702 0.186 1.902

NK 1 587.312 236.882 1148.810 1339.283 911.256 980.198 550.880 1968.531 0.000 3.976*** 15 34 0.702 0.188 1.902

NK 2 687.471 263.016 1309.710 1484.267 1073.229 1150.105 645.713 1815.597 1845.931 0.000 15 28.7 0.640 0.248 1.734

Eu-Fagenion. Wytwer (1990) recorded in average 6 species per site in the forest stand Tilio-Carpinetum and 5.33 species in Potentillo albae-Quercetum. Moreover there have been published some more data on this matter, e.g. Karafiat (1970) mentioned 9 species per site in a mixed pine-deciduous forest, Loksa (1966) 12.7 in the community of Ceraso-Quercetum clematidetosum, Zapparoli (1992) on average 6.4 in an oak forest stand (Quercetea ilicis) and 6.16 in a beech community ( Fagetalia sylvaticae). The comparisons mentioned above emphasise a relatively high species diversity of centipedes in the Malé Karpaty Mts, despite different sampling methods as well as forest communities. The highest species richness (17) in Malé Karpaty Mts refers to the site of Lošonský háj grove (LH). This is the 80–100 year old oak-hornbeam forest commonly with a 15–20 cm deep leaf litter horizon. The specimens have been represented with a high value of equitability (0.731) and were involved 3 eudominant, 4 dominant and 2 subdominant species with the lowest value of Simpson’s index of dominance (0.170) (Tables 3, 4). T a b l e 4. Mean dominance (%) of the centipede species at the study sites in 1999–2002. Species CA VI FU LI HH LH LL NA NK 1 Cryptops anomalans 1.58 Henia illyrica 4.21 Clinopodes flavidus 0.76 1.58 0.37 0.76 1.89 0.18 0.97 0.46 Geophilus flavus 4.84 0.30 0.74 6.48 Strigamia acuminata 23.86 22.35 7.12 5.95 17.81 8.76 17.72 9.57 Strigamia transsilvanica 0.76 0.37 1.35 0.74 1.52 0.18 Schendyla nemorensis 21.32 43.16 16.76 19.70 38.92 28.01 42.48 16.54 29.90 Lithobius agilis 1.52 7.62 4.59 5.56 5.52 4.30 4.51 Lithobius borealis 1.01 1.49 2.12 1.48 0.19 0.28 Lithobius dentatus 0.11 Lithobius erythrocephalus 1.05 0.56 5.00 2.23 2.48 1.61 0.55 Lithobius forficatus 1.01 1.12 0.45 0.54 5.01 0.95 0.75 1.66 Lithobius lapidicola 3.05 0.53 1.49 0.91 2.41 2.67 0.43 0.64 Lithobius latro 4.74 0.37 0.45 0.43 Lithobius lucifugus 0.51 Lithobius melanops 0.56 0.43 Lithobius mutabilis 2.54 0.53 20.48 3.79 6.76 6.12 9.71 9.77 12.60 Lithobius muticus 41.88 41.05 22.53 24.85 27.30 21.15 18.86 15.90 22.63 Lithobius pelidnus 0.25 2.43 0.93 0.19 0.54 Lithobius piceus 0.18 Lithobius aeruginosus 1.58 1.62 Lithobius austriacus 0.76 30.61 5.40 6.68 29.97 13.80 Lithobius crassipes 0.76 4.24 3.24 0.18 0.54 2.85 Lithobius microps 0.18 0.19 0.18 Abbreviations of study sites (CA → NK 2) see text

NK 2 0.98 0.33 43.63 0.43 10.45 11.01 0.22 0.11 0.43 0.98 0.11 7.83 13.93

9.25 0.22

On the contrary the maximum value of the dominance index refers to the centipede coenosis at the site of Vinosady (VI) (c = 0.360). The community was represented by 10 species, including two thermophilous centipedes, Cryptops anomalans and Henia illyrica. Low equitability (0.567) in the species structure at Vinosady (VI) hints at the 131

fact, that most of the collected specimens (160) belong to 2 eudominant centipedes, Schendyla nemorensis and Lithobius muticus. Schendyla nemorensis (more than 43%) (Table 4) may be classified as the most dominant species in the taxocoenosis. Apart from the determined species the samples at each site included juveniles and damaged specimens of the genus Lithobius (Table 1) excluded from the analyses. Juveniles occurred in the samples throughout the period of observation (March–November) and their abundance varied between 1 and 40, usually with a value of 5 to 15 specimens per sample. An exceptional abundance of 79 juveniles appeared at the site Naháč–Kukovačník (NA) on July 6, 2000. Generally continual occurrence reflects in egg lying and juvenile ontogeny throughout the year. Regular oscillation in abundance of juveniles throughout the year was not recorded. There were less damaged specimens in the samples. Their abundance varied from 1 to 18, usually 2 to 5 specimens per sample. In the 4-year long research (1999–2002) the juveniles were absent only occasionally, for instance in the sample from November 8, 2002 (site NK1, air temperature 0 to -1 °C). However the sample from November 11, 2000 at the same site (air temperature 4–5 °C) contained 4 juveniles. The t-test values clearly declare that the sites Cajla (CA) and Vinosady (VI) are the most different from the other (Table 3). The species diversity index (H') hints at high species richness at the sites. The highest diversity (2.072) refers to the community at Lošonský háj grove (LH), on the contrary the lowest value (1.306) appears at Vinosady (VI) with 10 species being recorded by the sieving method. The study site Vinosady is warm, however considerably drier during most of the year.

Dominant species in the centipede communities in the Malé Karpaty Mts In accordance with the average values of dominance, only Schendyla nemorensis and Lithobius muticus belonged to the eudominant species at all study sites. Their dominance varied between 10 and 40% (Table 4). Mean abundance of Schendyla nemorensis had the values from 3 to 10 ind.m-2, with the maximum of 10.16 ind.m-2 at the site of Naháč–Katarínka 1 (NK1) (Table 5). The site has been covered by 40–60 year old oak and hornbeam forest with a suitable soil texture and humidity as well as a rich litter horizon, providing suitable shelters. Lithobius muticus, the second dominant species, at all study sites had a dominance 15 to 40%, with an average value of 3 to 8 ind.m-2. The species Strigamia acuminata was eudominant at 5 sites (CA, FU, LH, NA, NK2), Lithobius austriacus at 3 sites (LI, NA, NK1) and L. mutabilis at 2 sites (FU, NK1) only. Similar data on abundance of centipedes were proved by numerous authors from various areas of Slovakia and other European countries. Stašiov (2002) published, that L. mutabilis was the most abundant in the forest community of Eu-Fagenion. In Querceto-Fagetum resp. Fagetum quercino-abietinum, the species Lithobius forficatus was the most common one (Stašiov, Maršalek, 1998). Tajovský (2000) presented the subspecies L. mutabilis sudeticus as the most frequent and most distributed centipede in 132

the Krkonoše National Park, occurring up to the highest altitudes (over 1500 m a.s.l.). Wytwer (1990) considered L. mutabilis as the most abundant species in the forest stands of Potentillo albae-Quercetum and Tilio-Carpinetum. In the Bükk Mts (study site Bélkő) Loksa (1966) recorded Lithobius muticus as the most common with abundance of 17.6 to 43.2 ind.m-2 in summertime. L. calcaratus and Schendyla nemorensis were the most frequent centipedes in the pine-deciduous forests on wind-flown sands in the surroundings of Darmstadt (Karafiat, 1970). In relation to our fauna there are the interesting Zapparoli’s (1992) records of Strigamia acuminata in Central Italy in beech forests only. In the Malé Karpaty this species commonly occurred in oak stands at lower altitudes. T a b l e 5. Mean abundance (ind.1m-2) of the centipede species at the study sites in 1999–2002. Species Cryptops anomalans Henia illyrica Clinopodes flavidus Geophilus flavus Strigamia acuminata Strigamia transsilvanica Schendyla nemorensis Lithobius agilis Lithobius borealis Lithobius dentatus Lithobius erythrocephalus Lithobius forficatus Lithobius lapidicola Lithobius latro Lithobius lucifugus Lithobius melanops Lithobius mutabilis Lithobius muticus Lithobius pelidnus Lithobius piceus Lithobius aeruginosus Lithobius austriacus Lithobius crassipes Lithobius microps

CA

0.11 3.36 0.11 3.00 0.21 0.14

VI 0.13 0.35 0.13

FU

LI

HH

LH

0.06 0.79 3.64 0.06 2.73 1.18 0.24

0.17 0.07 1.57

0.47

0.04 0.14 3.43 0.14 5.39 1.07 0.29

1.10 1.48 0.26 7.19 0.93 0.03

0.43 0.39

0.09 0.18 0.24 0.06

1.10 0.10 0.20 0.10

0.43 0.96 0.46

0.42 0.16 0.45

0.04 3.39

3.33 3.67

0.83 5.47

0.11 1.18 4.07 0.18

1.64 3.19 0.03

3.56

0.09 0.14 0.43

4.33

1.47 0.33 9.60 1.13

0.47 0.13

LL

NA

NK 1 NK 2

0.27

0.16

5.00

3.25 12.53 0.06 0.12 10.16 3.00 1.53 3.18 0.09 0.06 0.19 0.03 0.56 0.12 0.22 0.28

4.67 1.21 0.03 0.45 0.21 0.12 0.12

0.28 0.09

0.07 0.36 5.89 0.04

1.67 6.73 0.60

0.12 2.76 4.48 0.15

4.28 7.69

0.03 2.25 4.00

0.06 0.13 0.11 0.11

6.73 0.93

0.40 1.33 0.80

1.29 0.04 0.04

8.45 0.15 0.03

4.69 0.97 0.06

2.66 0.06

Abbreviations of study sites (CA → NK 2) see text

The biggest portion of subrecendent species with their dominance under 1% was represented at the sites of NA – 8 species, NK1 – 7 species, NK2 – 9 species (Table 4). Generally regular occurrence of the hygrophilous species Lithobius agilis at most of the sites, its eudominant position at NK2, dominant at FU, LH and LL and subdominant at HH, NA and NK1 corresponds with a relatively high degree of humidity in leaf litter during the whole year in central as well as northern part of the Malé Karpaty Mts. A lack of L. agilis at the sites Vinosady (VI) and Lindava (LI) indicates a drier and warmer character of oak-hornbeam forests in the southern part of the Malé Karpaty Mts and Trnavská pahorkatina hills. 133

Seasonal occurrence of centipedes at the study sites in the Malé Karpaty Mts The months of occurrence of the species at 10 study sites are presented in the Table 6. Most of the determined species were recorded throughout the whole sampling period from March to November 1999–2002. Lithobius dentatus and L. aeruginosus are the only species, recorded in several specimens in spring months (March, May) and at the beginning of autumn (September). Some of the species were found at different sites in spring and the autumnal months; Clinopodes flavidus occurred at the sites Lindava (LI) and Lošonský háj grove (LH) just in April and May and at Fúgelka (FU) in October and November only. The other species, Strigamia acuminate and Lithobius muticus were confirmed at most of the sites continuously from March to November. Schendyla nemorensis, the species occurring at all the study sites, was often absent in the samples from the summer months (June, July, August) (CA absence in June; HH, NA in July; LI, LH in August; NK2 in June–July) . The analyses of the centipede assemblages in the Malé Karpaty Mts in the period of 1999–2002 declare, that most of the recorded species occur throughout the year, even in wintertime, as being proved by the background samples from January 2000–2002 (excluded from the analyses).

Hierarchical classification of the centipede communities (Sørensen’s index, complete linkage) The classification of the ten centipede communities according to their species diversity is shown in the dendrogram (Fig. 1). The dendrogram is split out into the community at Vinosady (VI) and a cluster of the other 9 communities on the similarity level of 42%. The reason of this separated position lies in a low number of species (10), the occurrence of Cryptops anomalans, which was not recorded at the other sites and (on the contrary a) the lack of Strigamia acuminata, occurring at all the sites except for VI. The communities of 9 sites are differentiated on the similarity level of 64% into 2 clusters – CA-LL-NA-NK2 and HH-FU-LI-LH-NK1 determined by a different numbers of mutual species. The first cluster (CA-LL-NA-NK2) has 21 species with seven of them mutual: Strigamia acuminata, Schendyla nemorensis, Lithobius agilis, L. forficatus, L. lapidicola, L. mutabilis and L. muticus. The second cluster (HH-FU-LI-LH-NK1) has 20 species and 6 mutual ones: Clinopodes flavidus, Strigamia acuminata, Schendyla nemorensis, Lithobius forficatus, L. mutabilis and L. muticus. Henia illyrica and Lithobius dentatus appear in the first cluster only, L. piceus and L. aeruginosus do just in the second cluster. Of the 18 species mutual for both clusters, there are only Strigamia acuminata, Schendyla nemorensis, Lithobius forficatus, L. mutabilis and L. muticus represented in all 9 centipede communities. On the similarity level of 68% the second cluster is split out with a separate community at the site HH consisting of 12 species. As for the species identity the communities LH and NK1 are the most similar (87%), on the contrary the highest difference appears between the communities at VI and LH-NK1 (46%). 134

T a b l e 6. Occurrence of centipedes in months at the study sites. Species Cryptops anomalans Henia illyrica Clinopodes flavidus Geophilus flavus Strigamia acuminata Strigamia transsilvanica Schendyla nemorensis Lithobius agilis Lithobius borealis Lithobius dentatus Lithobius erythrocephalus Lithobius forficatus Lithobius lapidicola Lithobius latro Lithobius lucifugus Lithobius melanops Lithobius mutabilis Lithobius muticus Lithobius pelidnus Lithobius piceus Lithobius aeruginosus Lithobius austriacus Lithobius crassipes Lithobius microps Lithobius sp. juven. Lithobius sp. damaged

CA

3,8,10 3,5–11 3.7 3–5,7–11 6,8,10–11 7

VI 6–7,9 4–6,8–9 6.8

4–9,11

5 5–6,9 5,7–11

5 4–9,11

FU

LI

HH

LH

10–11 3–11 3–11 11 3–10 4–11 5,7,9

4–5 3–4 4–11

6,8,10

4 4,6,9,11 3–11 4.11 3–7,9–11 3–11 5,8–9,11

4 4,6–7,9 3–7,9 7

5–6,8–11 4–6,10–11 3–7,9–11 4–6,8–11 4–6,8–11 3–8,10–11 3–9,11 5 5.1 10

5

5,8,11 3–11 5–10

LL

4–11 4–11 4–5,7–9 4–11 4,6–11 4

NA

NK 1

NK 2

4–7,10

3,6–8

4–7,10 4,6,10

3–11

3–11 7–8 3–11 3–11 5,8,11

3–6,8–11 4–11

3–11 7–8,10–11 3–5,8–11 3–11

5 4,6–8,11 3,5,9–11 4–5,7,11 5–7,11 4-6,10–11 4–8,11 5,8–10 3.5 4,6-8 5

3.9 8 8-9

8.9 4–11 3–11 3.6

6 3–11 3–11

5.7 3–5,10 3–11 10

4 4–6,8

3–11 4–6,10–11 4–5,8–11 3–8,10–11 4–11 4–11 8–10

7–8 3–11 3–11 4.6

4–11 4–11 9

3–11 3–11 4.6

3

9 3–11 4–6,8–11 4–5,8–11 5, 8,10–11

6 7 4–5,7–11 4–7,9–11

4–8 4–7,11

3–11 4–7,9–11

3–11 4–6,8,10

Abbreviations of study sites (CA → NK 2) see text; 3 → 11 = March → November

4–11 4–5,7–10

3–11 11 9 4–11 3–6,9,11

3–11 3.1 4 4–11 4–5,8–9

3–11 3–11

3–11 3–5,7–11 7 3–11 3–8,10–11

3–11 4.8 3–11 3–9

Oscillation in stability of the species structure at the study sites is shown on the Fig. 2. The communities are classified according to all the years of the study (1999–2002). On the similarity level of 20% the dendrogram is split out again into the community at the site VI and the other coenoses. Low species richness at the site VI in each year (1999, 2001 – 6 species; 2000 – 7 species; 2002 – 5 species) seems to be significant enough to establish a separate cluster. The dendrogram declares higher stability of the communities at the sites VI and LI. Their structure changed with a low significance during the 4 years. The communities at the site VI were on the 54 % level of simiarity, at LI this had the value of 82%. The community at VI was more similar in 1999–2000 (more than 75%) than in the period of 2001–2002 (less than 75%).

Order of the sites: 1–4 CA 1999, 2000, 2001, 2002; 5–8 VI 1999, 2000, 2001, 2002; 9–12 FU 1999, 2000, 2001, 2002; 13–16 LI 1999, 2000, 2001, 2002; 17–19 HH 2000, 2001, 2002; 20–23 LH 1999, 2000, 2001, 2002; 24–27 LL 1999, 2000, 2001, 2002; 28–31 NA 1999, 2000, 2001, 2002; 32–35 NK 1 1999, 2000, 2001, 2002; 36–39 NK 2 1999, 2000, 2001, 2002. Fig. 2. Hierarchical classification of centipede communities in the years 1999–2002 at the study sites according to their presence/absence similarity (Sørensen’s index, complete linkage). Vertical axis = dissimilarity, abbreviations of study sites see text.

The analyses show even higher value of similarity from the communities at LI – in the period of 1999–2000 app. 85%, later in 2001–2002 95%. The sites CA and FU have 136

couples of years which are differentiated by one splitting only. CA has very similar communities just in the period of 2001–2002 (app. 95%). FU likely in 2000, 2002 (app. 95%). The species structure in the communities at the other sites changed more significantly in the years 1999–2002, they are divided by several splittings.

Hierarchical classification of the centipede communities (Wishart’s index, complete linkage) The centipede communities at the study sites in the Malé Karpaty Mts has split out into the coenosis at the site Vinosady (VI) and the cluster of the other sites on the similarity level of 31 %. The community at VI differs from the others with the lowest species richness (10) as well as total quantity of obtained specimens (233, 190 of them identified) (Table 3). On the similarity level of 57% the 9 communities were classified into a couple of communities FU-HH and the cluster with NA, NK1, NK2, LH, CA, LL and LI communities. In the pair of FU-HH the communities are the most similar (92%) from the qualitative-quantitative point of view. They may be defined by low species richness (FU – 13 species, HH – 12 species). The material from these sites includes 17 determined species, with mutual Clinopodes flavidus, Strigamia acuminata, S. transsilvanica, Schendyla nemorensis, Lithobius agilis, L. forficatus, L. mutabilis and L. muticus. The communities at NA, NK1, NK2 and LH being situated in the northern part of the Malé Karpaty Mts and with their similarity of 80% belonged to the richest from the qualitative as well as quantitative point of view. During the period of 4 years the sampling obtained almost 2,500 specimens and 20 determined species at these sites (Table 1). Under the centipede community classification in the period of 1999–2002 (Fig. 4) the community at the site VI has been separated from the other sites on the similarity level of approximately 21% (as in the dendrogram on the Fig. 2). The highest stability (the lowest oscillations in the qualitative and quantitative structures) appears in the coenoses at the site VI (similarity of 68% in 1999–2002), FU (similarity of 75% in 1999–2002), HH (similarity of 78% in 2000–2002). At the site CA the centipede community was stabile only in the years 2000–2002, in 1999 the structure was quite different. The community at NA declared its stability in the period of 1999–2001 however in the last year of studying (2002) the structure significantly changed. The dendrogram (Fig. 4), which explains classification of the communities, takes several variables into account, such as presence or absence of a species and its abundance. The level of stability in the communities during the research period is projected more precisely. The comparison of the community similarity according to the Sørensen’s (Fig. 1) as well as Wishart’s indices (Fig. 3) enables us to conclude the separated position of the community at VI and a close position of the communities at CA-LL. The communities at NA, NK1, NK2 and LH correspond with a high value of similarity according to the Wishart’s index, however they are divided into the couples of NA-NK2 a LH-NK1 on the level of 64% similarity according to the Sørensen’s index. The level of similarity 137

amongst the communities as well as their stability in the period of 1999–2002 is projected on the Figs 2 and 4 more precisely.

Fig. 3. Hierarchical classification of centipede communities in the forest epigaeon at the study sites according to their abundance similarity (Wishart’s index, complete linkage). Vertical axis = dissimilarity, abbreviations of study sites see text.

Conclusion The centipede (Chilopoda) communities in the Malé Karpaty Mts were studied at 9 sites in the period of 1999–2002 and at the site of Horný háj grove in 2000–2002. The communities were sampled in monthly intervals from March to October (resp. November). In total 8,303 specimens were obtained, 6,154 of them were determined. 1,843 juveniles and 306 damaged specimens were excluded from the analyses. In total 24 species have been determined (Table 1). The widest species spectrum (17) refers to the site Lošonský háj grove. This is the centipede community with high equitability (0.731), 3 eudominant, 4 dominant and 2 subdominant species and with the lowest value of the dominance index (0.170). On the contrary the highest value of the dominance index refers to the community at Vinosady (0.360), being formed by 10 species including thermophilous Cryptops anomalans and Henia illyrica. Low equitability (0.567) has been determined by 138

Schendyla nemorensis and Lithobius muticus, which formed the majority (160) of the sampled specimens. The species Schendyla nemorensis reached the highest value of dominance (43%).

Order of the sites: 1–4 CA 1999, 2000, 2001, 2002; 5–8 VI 1999, 2000, 2001, 2002; 9–12 FU 1999, 2000, 2001, 2002; 13–16 LI 1999, 2000, 2001, 2002; 17–19 HH 2000, 2001, 2002; 20–23 LH 1999, 2000, 2001, 2002; 24–27 LL 1999, 2000, 2001, 2002; 28–31 NA 1999, 2000, 2001, 2002; 32–35 NK 1 1999, 2000, 2001, 2002; 36–39 NK 2 1999, 2000, 2001, 2002 Fig. 4. Hierarchical classification of centipede communities in the years 1999–2002 at the study sites according to their abundance similarity (Wishart’s index complete linkage). Vertical axis = dissimilarity, abbreviations of study sites see text.

The t-test declares that the sites of Cajla and Vinosady are the most different from the other sites. The species diversity index hints at high species richness at these sites. The highest value (2.072) appeared in the community from Lošonský háj grove, the lowest one (1.306) at Vinosady with only 10 species obtained by the sieving method. Schendyla nemorensis and Lithobius muticus were eudominant species at all the study sites. The values of their dominance varied between 10 and 40%. The mean abundance of Schendyla nemorensis ranged between 3 and 10 ind.m-2, the highest value (10.16 ind.m-2) refers to the site of Naháč–Katarínka 1. Dominance of Lithobius muticus reached the values between 15 and 40%, its mean abundance varied from 3 to 8 ind.m-2. 139

The research in 1999–2002 enables us to conclude that most of the recorded centipedes occur throughout the year including wintertime. The community similarity dendrograms, based on Sørensen’s and Wishart’s indices declare separated position of the community at Vinosady and close relation between the communities at CA-LL with similarity level of 85% and 77%. The coenoses NA, NK1, NK2 and LH hint at a high degree of similarity (the Wishart’s index dendrogram). The Sørensen’s index of similarity has differentiated the couples of NA-NK2 and LH-NK1 on the 64% level of similarity. Oscillation of the stability in qualitative and quantitative structure of centipedes at the study sites is presented by the dendrograms, which classify the communities from the years 1999–2002. Translated by P. J. Fedor Acknowledgement The authors thank to M. Holecová for her help in the field work as well as in writing the manuscript. The research was supported by the project VEGA No. 1/0119/03.

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Országh I., Országhová Z.: Štruktúra taxocenóz stonožiek (Myriapoda: Chilopoda) dubovo-hrabových lesov Malých Karpát a Trnavskej pahorkatiny (JZ Slovensko). V období rokov 1999–2002 sme na 8 študijných plochách v Malých Karpatoch a 2 plochách na Trnavskej pahorkatine študovali taxocenózy stonožiek (Chilopoda). Plochy boli pokryté 40 až 100 ročným porastom dubovo-hrabového lesa. Situované boli v nadmorskej výške 240–350 m n.m. Vo vzorkách sme zaznamenali výskyt Cryptops anomalans, Henia illyrica, Clinopodes flavidus, Geophilus flavus, Strigamia acuminata, S. transsilvanica, Schendyla nemorensis, Lithobius agilis, L. borealis, L. dentatus, L. erythrocephalus, L. forficatus, L. lapidicola, L. latro, L. lucifugus, L. melanops, L. mutabilis, L. muticus, L. pelidnus, L. piceus, L. aeruginosus, L. austriacus, L. crassipes a L. microps. Najvyššia hodnota druhového bohatstva (17) zodpovedá ploche Lošonský háj, najnižšia diverzita (10) bola zistená vo Vinosadoch. Index druhovej rozmanitosti (H') a ekvitabilita (e) dosiahli najvyššiu hodnotu v spoločenstve v Lošonskom háji. Strigamia acuminata mal druhú najvyššiu hodnotu abundancie 12,53 jedincov na m2 na ploche Naháč–Katarínka 2. K ďalším taxónom s vysokými hodnotami priemernej abundancie patrili Schendyla nemorensis na ploche Naháč–Katarínka 1 (10,16 ind.m-2), na ploche Horný háj (9,6 ind.m-2) a Lithobius austriacus na ploche Naháč–Kukovačník (8,45 ind.m-2). Schendyla nemorensis a Lithobius muticus môžeme klasifikovať ako eudominantné druhy na všetkých

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študijných plochách, ich dominancia dosiahla priemernú hodnotu 40%. Na všetkých dendrogramoch zhlukovej analýzy je spoločenstvo na ploche Vinosady oddelené od ostatných na najvyššej hladine nepodobnosti. Príčinou je najnižší počet determinovaných druhov, unikátne zastúpenie Cryptops anomalans, absencia Strigamia acuminata a najnižší počet nazbieraných exemplárov za 4 roky. Z hľadiska druhovej identity sú na 87% úrovni najpodobnejšie spoločenstvá stonožiek LH – NK 1. Spoločenstvá dvojice FU – HH sú z hľadiska kvalitatívno-kvantitatívneho zloženia najbližšie až s 92% podobnosťou. Väčšina zistených druhov sa vyskytuje počas celého roka. Strigamia acuminata a Lithobius muticus boli na väčšine lokalít zistené súvisle od marca do novembra. Schendyla nemorensis, druh zastúpený na všetkých lokalitách, často absentoval vo vzorkách z letných mesiacov. Charakteristické spoločenstvo stonožiek dubovo-hrabových lesov Malých Karpát je zložené z druhov Schendyla nemorensis, Strigamia acuminata, Lithobius agilis, L. borealis, L. lapidicola, L. mutabilis, L. muticus, L. austriacus, v južnejších suchších častiach Malých Karpát tiež Henia illyrica.

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Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 161–178

BUG ASSEMBLAGES IN EPIGEON OF OAK-HORNBEAM FORESTS IN SW SLOVAKIA KAREL HRADIL State Phytosanitary Administration, Tylova 29, Jičín, The Czech Republic, e-mail: [email protected], [email protected] Abstract Hradil K.: Bug assemblages in epigeon of oak-hornbeam forests in SW Slovakia. Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 161–178. The study contains results of four-year research on epigeic Heteroptera fauna at 8 terrestrial sites of the Malé Karpaty Mts and 2 sites of the Trnavská pahorkatina hills. The bugs were collected during the period of 1999–2002 using a square method and sifting. The 399 obtained specimens belong to 46 species of 17 families. The evaluation of the Heteroptera taxocoenoses was based on their quantitative structure, being expressed by the average abundance values per 1 m2, and on their qualitative aspects with emphasis on the constancy and dominance values. The RDA analysis showed that of the 15 tested variables the E3 cover is the most significant factor. Legnotus limbosus and Eurygaster maura may be classified as eudominant species, Plinthisus brevipennis, Megalonotus sabulicola, M. chiragra, Peritrechus nubilus and Raglius alboacuminatus as dominant and subdominant elements. The occurrence of Myrmedobia exilis is faunistically interesting. Key.words:.bugs, Heteroptera, epigeic assemblages, oak-hornbeam forests, SW ecology, human impacts

Slovakia,

Introduction Bugs live on soil surface, occasionally hypogeically and form a quantitatively as well as qualitatively significant component of zoocoenoses. Predominantly they are phytophagous, feeding on seeds and fruits and often sucking on roots, stalks or lower leaves. Plenty of species specialize on bryophytes, or even lichens on the basis of trunks. Predation or necrophagy may appear less frequently. Some species live in soil or epigeon permanently, the others just temporally. They often survive unsuitable climatic conditions or wintertime in a leaf litter horizon. Species structure and abundance in epigeic assemblages of bugs sensitively reflect changes in humidity, ruderalisation as well as an increase of shadowiness. At open 161

steppelike sites, blown sands, rocky steppes, etc. the epigeic species occur frequently. On the other hand in shadowy habitats, especially in forests, they are less common and diverse, however relatively stabile. In Slovakia the problems on epigeic bug species in the floodplain forest along the Morava river alluvium and along the Danube river in the vicinity of Gabčíkovo dam (including the anthropogenous impact) were analysed by Štepanovičová (1994, 1995, 1997) and Štepanovičová, Degma (1999). Structure and seasonal dynamics of Heteroptera taxocoenoses in the vicinity of the Morava river were studied by Majzlan, Štepanovičová (1998). The papers declare that the epigeic bugs represent good bioindicators to detect hydrological variables in environment. Qualitative – quantitative relations amongst overwintering bugs in the region of the Malé Karpaty Mts were observed by Štepanovičová, Kovačovský (1971), the epigeic Heteroptera in the surroundings of Bratislava were studied by Bianchi (1991). Majzlan et al. (2000) used the sifting method to sample leaf litter in the National Nature Reserve Čachtický hradný vrch in the Protected Landscape Area Malé Karpaty. The other papers on this matter were published by Štepanovičová (2001) from the woody steppe habitats in the Považský Inovec Mts, Štepanovičová, Országh (2002) from the Malé Karpaty Mts, or Štepanovičová, Bianchi (2005) who used sifting and ground traps to study the bug fauna in Devínska Kobyla hill. Rédei, Hufnagel (2003a, b) analysed epigeic Heteroptera in Hungary by sifting, Bakonyi et al. (2002) used ground traps. In Bulgaria Strawinski (1960) dealt with bugs living under stones. In the Czech Republic epigeic bugs were studied in the floodplain forest in Moravian Lednice (Křístek, 1985, 1991; Křístek, Dobšík, 1985), or in Central Bohemia (Karlštejnsko) by Verner (1959) in his complex study on the arthropods inhabiting soil of oak-hornbeam forests.

Study area In the period of 1999–2002 the epigeic bug fauna was studied in SW Slovakia in the Malé Karpaty Mts and Trnavská pahorkatina hills. The regular samples come from the following sites: Cajla (CA) forest community of Querco-Carpinetum poetosum nemoralis on SE slopes of Malá cajlanská homola hill at the altitude of 260–280 m a.s.l. Vinosady (VI) forest community of Querco-Carpinetum poetosum nemoralis on the basis of Kamenica hill at the altitude of 260–280 m a.s.l. Fúgelka (FU) forest community of Querco-Carpinetum melicetosum uniflorae with Rubus fruticosus a R. hirtus (oceanic species), 3 km NW from the village of Dubová. The altitude of 350 m a.s.l. Lindava (LI) forest community of Quercetum petrae-cerris (on acid psedocley) situated app. 1 km E from the village of Píla (Trnavská pahorkatina hills). Nature Reserve. Altitude of 240 m a.s.l.

162

Horný háj grove (HH) forest community of Querco-Carpinetum melicetosum uniflorae within the cadastral area of Horné Orešany (Trnavská pahorkatina hills) at the altitude of 240 m a.s.l. It forms a larger complex of island forest surrounded by vineyards and farmland. Lošonec–lom quarry (LL) forest community of Querco-Carpinetum caricetosum pilosae at the altitude of 340 m a.s.l. Lošonský háj grove (LH) being situated close to the village of Lošonec (in the vicinity of Smolenice), in 1984 established as the Nature Reserve. The forest community of Querco-Carpinetum caricetosum pilosae in the deluvium under the slopes. The altitude of 260 m a.s.l. Naháč–Kukovačník (NA) young stand of Querco petrae-Carpinetum melicetosum uniflorae forming a small island amongst farmland at the altitude of 300 m a.s.l. Naháč–Katarínka 1 (NK1) (Nature Reserve) young oak-hornbeam sprout forest of Querco petrae-Carpinetum melicetosum uniflorae at the altitude of 340 m a.s.l. Naháč–Katarínka 2 (NK2) (Nature Reserve) old woody stand of Lithospermo-Quercetum virgilianae (on limestone) under the ruins of monastery. The forest is 80–100 years old and situated at the altitude of 320 m a.s.l. The detailed analysis on the study sites has been accompanied in the paper by Zlinská et al. (2005).

Methods The soil macrofauna was collected by M. Holecová (Faculty of Natural Sciences, Comenius University, Bratislava) by the square method combined with sifting. In each study plot, at about 1-month intervals from March to November, the material was collected from the leaf litter and the upper part of soil from 16 squares. Each square consisted of the area of 25x25 cm, i.e. altogether an area of 1 m2 was sifted, representing one sample. The samples were extracted using xereclectors of the Moczarski’s type. The material has been deposed at the collection of Karel Hradil in Miletín (Czech Republic). The species dominance is characterised by the scale proposed by Tischler (1949) and completed by Heydemann (1955): ED = eudominant, D = dominant, SD = subdominant, R = recedent, SR = subrecedent. Constancy is defined by the following symbols: EU = euconstant (75–100%), C = constant (50–75%), AS = accessory (25–50%), AD = accidental species (0–25%) (Pelikán, 1993). The cluster analysis of the communities was done using the computer program NCLAS (Podani, 1993). The clustering method complete linkage in combination with Sørensen’s index and Wishart’s similarity ratio was used. Species diversity was evaluated according to Odum (1977). All the couples of diversity values were compared with a t-test, to find out if they are significantly different (Poole, 1974). Principal component analysis (PCA) was performed to assess the relationship among the Heteroptera communities at 10 study sites using the program Canoco (ter Braak, Šmilauer, 1998). For PCA a canonical ordination (RDA) with forward selection of environmental variables was used to find out environmental variables influencing the community. At all the study sites the following environmental variables were measured (range of values of gradient variables or categories of categorical ones are in brackets): pedological and chemical characteristics of leaf litter such as total organic carbon (3.80–14.40%); total nitrogen content (0.35–1.1%); content of exchange bases (Ca, Mg, Na, K) (12.60–49.40 mval/100g); total acidity (1.8–14.9 mval/100g); pH in H2O (3.90–6.74); humus content (6.55–24.83%); age of a forest stand (40–100 years); cover of E3: (65–80%); cover of E2 (1–50%); cover of E1 (40–100%); lying and standing dead wood (2–20%); forest

163

fragmentation (0–50%), exposition of study plots (E, W, S, NE, NW, SW, SE), slope (0–45°); pollution by calcareous dust (present, absent). The zoogeographical distribution accords to Josifov (1986). Habitat and trophic categories, humidity preferences and interactions towards soil were used according to Stehlík (1983, 1984, 1985, 1986, 2002), Stehlík, Heiss (2000), Stehlík, Vavřínová (1993, 1995, 1996, 1998, 1999) and Štepanovičová, Bianchi (1998, 2003).

Results In the period of 1999–2002 totally 399 bug specimens of 46 species and 17 families were recorded. Their average abundance at all the study sites reached a value of 2.44 specimens per 1 m2. A survey of the collected specimens with the values of their abundance, dominance, constancy, diversity indices, equitability is presented in the Table 1. Phenology, zoogeographical distribution, habitat and trophic categories, humidity preferences and interactions to soil are included in the Table 2. Holopalaearctic (23.26%) and Western Palaearctic species (20.93%) may be classified as eudominant. The group of dominant elements includes Holomediterranean (9.3%), Euro-Siberian species (9.3%) and Western Euro-Siberian species (6.98%). The other species were subdominant (Table 2). The analysis on trophic relations showed significant dominance of phytophagous species (83%). The rest was formed by zoophagous (15.21%) and mycetophagous bugs (2.17%). An interesting fact lies in comparison of species abundance in accordance with their interactions with soil. The samples consisted of dominating geophilous (58.7%), relatively well-represented geoxene (37.21%) – accidental in this habitat and finally geobiont (6.52%) species. According to the humidity preferences the most common were euryhygric (69.77%) and then xerophilous species (27.91%) in the samples. Only one species belonged to the category of hygrophilous insects. The data correspond with drier ecological conditions at the sites. The hierachical classification of the Heteroptera assemblages at all the study sites and according to the species identity (Sørensen’s index) as well as abundance similarity (Wishart’s index) is projected on 2 dendrograms (Fig.1, 2). The analysis on qualitative similarity in the taxocoenoses hints at two clusters. The first cluster includes the sites of VI, NA, FU, NK1 and HH, the second one CA, LI, NK2, LH and LL. The dendrogram declares that the sites are not significantly similar however there are not any significant differences amongst the communities. The sites of VI (9 mutual species of 12 total) and NA (9 of 21 total species) are relatively the most similar (less than 60%). The fact corresponds with the constancy index values, as amongst 46 recorded species there are only 5 euconstant and 1 (up to 3) constant species. The other bugs were accessory or accidental. Approximately the same E3 cover (70–75%), cover of E2 and E1, similar pH values (4.18–4.48) as well as humus content in soil were mutual factors at all the sites. The couples of FU – NK1, LI – NK2 and LH – LL show a similar species spectrum of bugs. 164

T a b l e 1. Survey of bug species being recorded in oak-hornbeam forest epigeon in the Malé Karpaty Mts and Trnavská pahorkatina hills. No. Species CA VI FU LI HH LH LL NA NK1 NK2 % Do Con. 3.54 3.54 SD AD 1 Acalypta musci (S c h r a n k, 1781) 1.18 1.18 R AD 2 Acalypta spp. nymfy 3 Aelia acuminata (L i n n a e u s, 0.29 0.29 1.18 0.29 0.59 0.59 0.29 0.29 3.83 SD EC 1758) 4 Aneurus (Aneurus) l. laevis 0.59 0.29 1.47 2.36 SD AS (F.a.b.r.i.c i u s, 1775) 0.29 0.29 0.59 SR AD 5 Arma custos (F a b r i c i u s, 1794) 6 Campylosteira verna (F a l l é n, 0.29 3.83 4.13 SD AD 1826) 7 Carpocoris purpureipennis (D e 0.29 0.29 SR AD G.e.e.r, 1773) 8 Ceraleptus gracilicornis 0.29 0.29 SR AD (H.e.r.r.i.c.h-S.c.h.a.e.f.f.e.r, 1835) 9 Deraeocoris (K.) lutescens 0.29 0.29 SR AD (S.c.h.i.l.l.i.n.g, 1837) 10 Drymus b. brunneus (R. F. 0.59 0.59 SR AD S.a.h.l.b.e.r.g, 1848) 11 Drymus sylvaticus (F a b r i c i u s, 0.59 0.29 0.29 0.59 1.77 R AS 1775) 0.29 0.59 0.88 SR AD 12 Elasmucha grisea grisea (L.i.n.n.a.e.u.s, 1758) 13 Emblethis verbasci (F.a.b.r.i.c.u.s, 0.29 0.29 SR AD 1803) 14 Eremocoris plebejus (F.a.l.l.é.n, 0.29 0.29 SR AD 1807) 15 Eremocoris podagricus 0.29 0.29 SR AD (F.a.b.r.i.c.i.u.s, 1775) 16 Eurydema oleracea (L.i.n.n.a.e.u.s, 0.29 0.59 0.88 SR AD 1758) 17 Eurygaster a. austriaca 0.29 0.88 0.29 0.29 1.77 R AS (S.c.h.r.a.n.k, 1778) 18 Eurygaster maura (L.i.n n a e u s, 2.06 1.77 2.06 1.77 0.29 1.47 0.59 1.47 0.59 0.88 12.98 ED EC 1758) 0.59 0.88 7.67 2.06 4.72 5.01 0.88 1.77 23.6 ED EC 19 Legnotus limbosus (G.e.o f f r o y, 1785) 20 Loricula exilis (F a l l é n, 1807) 1.47 0.59 2.06 SD AD 21 Lygus rugulipennis P o p p i u s, 0.29 0.59 0.29 0.29 0.88 2.36 SD AS-C 1911 22 Megalonotus chiragra 0.59 0.59 0.29 0.29 0.29 0.29 0.29 0.29 0.59 3.54 SD EC (F.a.b.r.i.c.i.u.s, 1794) 23 Megalonotus sabulicola 1.77 0.59 0.29 0.29 0.29 0.29 0.29 0.59 0.29 0.29 5.01 D EC (T.h.o.m.s.o.n, 1870) 24 Metopoplax origani (K.o.l.e.n.a.t.i, 0.29 0.29 SR AD 1845) 25 Microporus nigritus (F a b r i c i u s, 0.29 0.29 SR AD 1794) 26 Nabis brevis brevis S c h o l z, 1847 0.29 0.29 SR AD 27 Nabis p. pseudoferus R.e.m.a.n.e, 0.29 0.29 SR AD 1949

165

T a b l e 1. (Continued) No. Species 28 Nabis rugosus (L i n n a e u s, 1758) 29 Neottiglossa pusilla (G.m.e.l.i.n, 1789) 30 Orius sp. 31 Palomena prasina (L.i.n.n.a.e.u.s, 1761) 32 Peritrechus gracilicornis P u t o n, 1877 33 Peritrechus nubilus (F a l l é n, 1807) 34 Piesma capitatum (W o l f f, 1804) 35 Piesma maculatum (L a p o r t e, 1833) 36 Plinthisus brevipennis (L.a.t.r.e.i.l.l.e, 1807) 37 Pyrrhocoris apterus (L i n n a e u s, 1758) 38 Raglius alboacuminatus (G o e z e, 1778) 39 Rhoplaus (R.) parumpunctatus (S.c.h.i.l.l.i.n.g, 1829) 40 Taphropeltus hamulatus (T.h.o.m.s.o.n, 1870) 41 Thyreocoris scarabaeoides (L.i.n.n.a.e.u.s, 1758) 42 Trapezonotus a. arenarius (L.i.n.n.a.e.u.s, 1758) 43 Tritomegas bicolor (L.i.n.n.a.e.u.s, 1758) 44 Tritomegas sexmaculatus (R.a.m.b.u.r, 1839) 45 Tropistethus holosericus (S.c.h.o.l.z, 1846) 46 Tropistethus sp. Number of species

CA VI FU

LI HH LH LL NA NK1 NK2 % Do Con. 0.29 0.29 0.59 SR AD 0.29 0.29 SR AD 0.59 SR AD 0.29 0.59 SR AD

0.59 0.29 0.29

0.29 SR AD 0.29

0.29

2.65 0.29 1.18 4.72 SD AS-C

0.29 0.29

1.18 0.29 1.77 R AS 2.06 0.29 2.65 SD AS 5.31

0.59

0.29 0.29 0.29 0.29

5.9 D

AD

1.18 1.47 R

AD

0.88 0.88 0.29

2.95 SD

0.29

C

0.29 SR AD 0.29 0.29 SR AD

0.29

0.29 SR AD

0.29

0.29 SR AD 0.29

0.29 0.29 12

12

11

11

0.29

0.59 SR AD

0.59

0.88 SR AD

0.29 0.59 0.29 9 7 11 21

0.59 SR AD 0.88 SR AD 14

16

Abbreviations: Study site: CA – Cajla, VI – Vinosady, FU – Fúgelka, LI – Lindava, HH – Horný háj grove, LL – Lošonec–lom quarry, LH – Lošonský háj grove, NA – Naháč–Kukovačník, NK1 – Naháč–Katarínka 1, NK2 – Naháč–Katarínka 2; % dominance; Do – category of dominance: ED – eudominant, D – dominant, SD – subdominant, R – recedent, SR – subrecent; Co – category of constancy: EK – euconstant, C – constant, AS – accessory, AD – accidental

The analysis on similarity of the bug assemblage abundances at all the study sites (Fig. 2) hints at two clusters on the low level of site similarity (10–60 %). The first cluster is represented by the CA, VI, LH, FU and NK1 sites, the second one by LI, NA, HH, LL and NK2. The highest similarity appears between CA and VI as well as HH and LL, the lowest values refer to CA and NK2. The highest abundance of bugs refers to the site NA (4.53 ex.m-2), the lowest one to FU (1.63 ex.m-2), however there isn’t a significant difference between these values. 166

T a b l e 2. Survey of bug species - phenology, zoogeographical distribution, habitat and trophic categories, humidity preferences, interactions to soil. Species Tingidae Acalypta musci Acalypta spp. nymfy Campylosteira verna Microphysidae Loricula exilis Miridae Lygus rugulipennis Deraeocoris lutescens Nabidae Nabis brevis brevis Nabis p. pseudoferus Nabis rugosus Anthocoridae Orius sp. Aradidae Aneurus l. laevis Oxycarenidae Metopoplax origani Rhyparochromidae Drymus b. brunneus Drymus sylvaticus Emblethis verbasci Eremocoris plebejus Eremocoris podagricus Megalonotus chiragra Megalonotus sabulicola Peritrechus gracilicornis Peritrechus nubilus Plinthisus brevipennis Raglius alboacuminatus Taphropeltus hamulatus Trapezonotus a. arenarius Tropistethus holosericus Tropistethus sp. Piesmatidae Piesma capitatum

CA

VI

FU

LI

HH

LH

LL

NA

NK1

NK2

Zg

Hab.

Tr

H

Sub.

-

-

-

-

10

-

-

3, 10

-

5, 7 3, 7 -

E E

st st

F F F

e sx

Gb Gf Gb

-

-

7

-

9

-

-

-

-

-

WES

st

Z

e

Gf

11 -

-

-

10, 11 -

-

-

3

11 -

11 -

11 -

HP HM (1)

et et

F Z

e e

Gx Gx

-

-

-

-

-

-

-

10 9

9 -

4

WES (BMo) WP WES

et et et

Z Z Z

e e e

Gf Gf Gf

-

-

-

-

-

-

11

-

-

-

-

-

Z

-

Gx

-

-

5, 7

-

-

-

-

4

3, 7

-

WES

st

M

e

Gx

-

-

-

-

-

-

-

-

5

-

HM (1)

et

F

e

Gx

8 9 4, 9, 10 11 9 -

4, 5 4 4 7 4 -

6 4, 8 10 11 4 -

11 11 9 4-7 9 -

10 10 10 10

10 9 -

4 7 4, 9 5 4

4, 11 4 3 4 4, 11 4 - 11 3, 5 10 -

et et st et et et st st et st et et st st -

F F F, N F F F F F F F F F F F F

sh e sx e e e sx sx e sx e e sx sx -

Gf Gf Gf Gf Gf Gf Gf Gf Gf Gf Gf Gf Gf Gf Gf

-

11

-

-

-

-

-

4, 9, 10

et

F

e

Gf

ES HP HM (1)+MA WP MoM 11 9, 10 ES 4 10 WP NM (1)+MA 11 3, 9, 10 WP WP 3 WP 4 WP HA WP -

11

ES

T a b l e 2. (Continued) Species Piesma maculatum Pyrrhocoridae Pyrrhocoris apterus Coreidae Ceraleptus gracilicornis Rhopalidae Rhopalus parumpunctatus Cydnidae Legnotus limbosus Microporus nigritus Tritomegas bicolor Tritomegas sexmaculatus Thyreocoridae Thyreocoris scarabaeoides Scutelleridae Eurygaster a. austriaca Eurygaster maura Pentatomidae Aelia acuminata Arma custos Carpocoris purpureipennis Eurydema oleracea Neottiglossa pusilla Palomena prasina Acanthosomatidae Elasmucha grisea grisea

CA -

VI 4

FU -

LI -

HH -

LH -

LL -

9

-

-

-

-

-

-

-

11

-

-

-

-

-

-

-

-

-

-

-

-

-

5 -

5 5 7

-

-

-

-

4 - 7 5, 7, 8 3 -

8 8, 9, 11 4, 7 - 9 8, 9, 11 7 - 9

-

NA NK1 3, 4, 9, 11 11

NK2 -

Zg HP

Hab. et

Tr F

H e

Sub. Gf

-

11

HA

et

F, N

sx

Gf

-

-

-

HM (1)

st

F

sx

Gx

-

4

-

HP

et

F

e

Gx

4, 5 -

HM (1)+MA HP HP NM (1)

st st et et

F F F F

e sx e e

Gf Gb Gf Gf

-

WP

st

F

sx

Gf

HM (1) HP

et et

F F

e e

Gx Gx

4, 5, 7, 8 3-7, 9, 10 5, 11 10 4 -

-

-

-

9

10

8, 11 5, 7, 10, 11

7 4, 10

9 - 11

-

9 8, 9 8, 9, 11

3 -

9 -

4, 8-10 4 8

4 9 -

-

10 11 10 -

3 -

4, 9 -

9 -

3 10, 11 11

HP SES ES (BMo) HP ES HP

et st et et et et

F Z F F F F

e e e e sx e

Gx Gx Gx Gx Gx Gx

-

-

-

3

-

-

-

-

-

3

PM

et

F

e

Gx

Zoogeographical distribution (Zg): E – western European-Siberian species occurring in Europe only, HP – Holopalearctic species, WES – western Euro-Siberian species, SES – southern Euro-Siberian species, ES – Euro-Siberian species, Bmo – Boreo-montane species, HM (1) – Holomediterranean species, PM – Ponto-Mediterranean species, MA – Central Asian species, WP – western Palaearctic species, MoM – Montanmediterranean, NM (1) – northern Mediterranean species reaching Central Europe in the north, HA – Holarctic. Habitat (Hab.): et – eurytopic, st – stenotopic. Trophic interactions (Tr): F – phytophagous species, M – mycetophagous species, N – necrophagous species, Z – zoophagous species. Ecological valency (V): e – euryhygric, sh – hygrophilous, sx xerophilous. Habitat (Hab.): et – eurytopic, st – stenotopic . Topic preferences (Sub.): Gb – geobiont, Gf – geophilous, Gx – geoxenous. 3 – 11: month of occurrence. Abbreviations of study sites see in Table 1.

The analyses on the epigeic bug taxocoenosis structure includes indices of diversity as well as equitability. Species diversity expresses ratio between number of species and number of specimens. Equitability hints at a degree of balanced dominance of the present species. At all the study sites the diversity indices varied between relatively low values 1.554 (LI) – 2.542 (NA), what corresponds with several dominant species and hence low diversity. The relatively high values of equitability at the sites CA (0.924) and LI (0.911) declare a balanced structure of specimens amongst the recorded species. At the other sites the equitability varied between 0.648 – 0.889. The highest value of diversity (Table. 3) refers to the site NA (moreover with the highest abundance as well – 4.53 ex.m-2). The value was high significantly different from the diversity at LI (t = 5.24) and significantly different from the sites LH (t = 3.334), LL (t = 3.155), HH (t = 2.524) and FU (t = 2.122). High significant differences appeared in values of Heteroptera taxocoenosis diversity between the sites LI – NK1 (t = 3.646), NK2 (t = 4.609), and significant differences originate between LI – CA (t = 2.368), VI (t = 3.016), FU (2.447); NK1 – LH (t = 2.464), LL (t = 2.378) and the sites NK2 – HH (t = 2.157), LH (t = 2.911) a LL (t = 2.767).

Fig. 1. Hierarchical classification of bug assemblages at the study sites according to their species presence/absence similarity (Sørensen’s index, complete linkage). Vertical axis = dissimilarity, for abbreviations of study sites see Table 1.

The PCA analysis (Fig. 3, for species numbers see Table 1) confirmed the similarity of 2 sites – LL and LI, which are parallelly in negative correlation with NK. The same fact is expressed in the dendrogram. According to the Wishart´s index the group of 169

similar sites includes VI, LH, CA, FU, NK1. The couples of NA – NK2; NA – LL, LI; or NA – FU, NK1, VI, HH, LH, CA, are dissimilar what actually contradicts with the Wishart’s index. The first two PCA axes account for 74.7% of the variance of the species data. The eigenvalues of the two first axes are λ1 = 0.566, λ2 = 0.182 (Fig. 3). For codes of taxa and codes of localities see Table 1. A detailed study on factors influencing occurrence of bugs in taxocoenoses at the sites, being based on RDA ordination (Fig. 4) has shown that just E3 played a significant role. The closest affinity to this factor has appeared at Rhopalus parumpunctatus, Metopoplax origani, Loricula exilis, Nabis pseudoferus, Aelia acuminata, Aneurus laevis and Carpocoris purpureipennis. Of these species only Loricula exilis and Nabis pseudoferus may permanently inhabit shadowy habitats. The others are classified as accidental or they use leaf litter horizon for overwintering. The group of heliophilous species preferring sparse canopy layer includes the species such as Tropistethus holosericeus, Legnotus limbosus, Plinthisus brevipennis, Raglius alboacuminatus, Trapezonotus arenarius or Microporus nigritus.

Fig. 2. Hierarchical classification of bug assemblages at the study sites according to their abundance similarity (Wishart’s index, complete linkage). Vertical axis = dissimilarity, for abbreviations of study sites see Table 1.

The results of RDA ordination analysis are is given in Fig 4. Eigenvalues of the two first axes of RDA are λ1 = 0.294 and λ2 = 0.322. The first two axes account for 61.7% of the total variance of the species data. 170

T a b l e 3. Species diversity test (Poole, 1974) and basic coenological characteristics (Begon et al., 1997) of the bug assemblages at the study sites in 1999–2002.

CA VI FU LI HH LH LL NA NK1 NK2

Study site Σ spp. MA [ind.m-2] e c H’ 2.133 2.264 2.105 1.554 1.869 1.730 1.720 2.542 2.438 2.455

CA 12 1.67 0.924 0.162 2.133 0.488ns 0.110ns 2.368* 0.855ns 1.394ns 1.360ns 1.801ns 1.115ns 1.383ns

VI 12 1.57 0.886 0.132 2.264 47.000 0.636ns 3.016** 1.308ns 1.897ns 1.835ns 1.283ns 0.659ns 0.859ns

FU 11 1.63 0.858 0.154 2.105 49.783 46.823 2.447* 0.803ns 1.373ns 1.335ns 2.122* 1.300ns 1.649ns

LI 11 3.47 0.911 0.301 1.554 54.391 51.815 64.080 1.117ns 0.676ns 0.602ns 5.244*** 3.646*** 4.609***

HH 9 2.13 0.878 0.218 1.869 36.139 33.722 32.907 31.058 0.433ns 0.445ns 2.524* 1.853ns 2.157*

LH 7 1.56 0.648 0.214 1.730 33.835 31.323 30.657 28.986 30.990 0.030ns 3.334** 2.464* 2.911**

LL 11 2.73 0.851 0.311 1.720 54.599 51.641 53.075 54.526 42.665 40.383 3.155** 2.378* 2.767**

NA 21 4.53 0.889 0.112 2.542 44.393 41.660 52.533 120.014 25.726 23.251 45.914 0.461ns 0.500ns

NK1 14 2.33 0.717 0.111 2.438 45.745 42.770 44.600 47.222 34.390 31.902 50.992 37.976

NK2 16 2.82 0.835 0.115 2.455 46.760 44.026 54.490 106.842 27.308 24.929 48.175 112.564 40.305

0.074ns

Explications: Σ spp. – total number of species, MA [ind.m-2] – mean abundance, H’ – Shannon’s index of species diversity, e – Pielou’s index of equitability, c – Simpson’ index of dominance. T-test values are under the diagonal and degrees of freedom are above it. Significance levels: *** P < 0.001; ** 0.001 < P < 0.01; * 0.01 < P < 0.05; ns 0.05 < P (non-significant). Abbreviations of study sites see Table 1.

Discussion The papers dealing with quantitative and qualitative analyses on epigeic bug fauna declare that the most frequent families include (Table 4): Rhyparochromidae, Pentatomidae (eudominant), Tingidae, Miridae, Cydnidae, Nabidae, Anthocoridae and Oxycarenidae (dominant). The qualitative analyses on epigeic bug fauna in forests show that species richness does not reach high values (21–28 species). The following species may be considered as the typical elements: Rhyparochromidae – Drymus brunneus, D. ryeii, Scolopostethus affinis, S. thomsoni, Rhyparochromus alboacuminatus, Cydnidae – Legnotus limbosus. The geoxenous species such as Aelia acuminata or Eurygaster maura (Křístek, Dobšík, 1985; Křístek, 1991; Štepanovičová, Degma, 1999) are common as well.

Fig. 3. Principal component analyses of bug taxocoenoses in forest epigeon at the study sites (for abbreviations of study plots and species codes see Table 1).

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Hence the species diversity in these habitats does not seem to be high what may hint at the fact that these conditions are not optimal for bugs. Just a few species have been adapted to inhabit the sites permanently with a high value of dominance. Apart from these characteristic species the samples from sifting contain plenty of accidental Heteroptera, which primarily inhabit the different habitats and they use the leaf litter horizon of oak-hornbeam forests for overwintering particularly. They are often dominant.

Fig. 4. RDA canonical ordination of bug taxocoenoses and environmental characteristics of the localities (for abbreviations of study plots and species codes see Table 1).

According to many references humidity is one of the limiting factors for a species structure, what has been actually proved by dominance of hygrophilous species in more humid localities as well as mesophilous to xerophilous species under drier conditions in epigeon of deciduous forests. In the Malé Karpaty Mts and Trnavská pahorkatina hills the four year long research of epigeic bugs at ten study sites has recorded 46 species in total. The average abundance varied between 1.56–4.53 (in average 2.44) ex.m-2, what may be considered as a low degree of abundance in comparison with the similar research on epigeic bugs in 173

T a b l e 4. A survey of bugs recorded at the research of epigeic fauna (%). Authors 1 2 3 4 5 6 7 8 9 10 11 12 13 D Family/Country CZ-B CZ-M SK SK SK SK SK SK SK SK HU HU SK Ceratocombidae 1.19 0.64 0.9 0.91 Dipsocoridae 4.35 1.27 0.9 2.17 Naucoridae 1.19 1.19 Pleidae 0.64 0.64 Hebridae 1.27 1.8 1.54 Hydrometridae 1.19 0.64 0.92 Veliidae 0.64 0.64 Gerridae 1.19 1.27 0.9 1.12 Saldidae 4.35 1.19 1.91 0.03 1.87 Tingidae 4.35 8.7 5.2 9.52 8.33 9.52 7.14 15.29 10.81 6.52 8.54 Microphysidae 4.35 2.55 2.7 2.17 2.94 Miridae 8.7 26.09 3.4 4.76 3.57 6.37 6.31 4.35 7.94 Nabidae 8.7 13.04 6.8 4.55 4.76 3.57 2.38 3.18 7.21 6.52 6.07 Anthocoridae 8.7 2.17 5.44 Reduviidae 7.14 0.64 3.89 Phymatidae 2.38 2.38 2.38 2.55 1.8 2.17 2.95 Aradidae 3.39 4.76 3.57 Piesmatidae 4.35 8.7 1.7 1.69 4.76 2.38 3.82 1.8 4.35 3.73 Berytidae 1.7 9.09 2.55 2.7 4.01 Lygaeidae 4.76 4.76 1.91 1.8 3.31 Cymidae 1.7 1.69 1.19 1.27 0.9 1.35 Blissidae 8.69 1.27 4.98 Geocoridae 2.38 0.64 0.9 1.31 Artheneidae 1.69 1.19 1.44 Heterogastridae 1.69 1.19 0.64 1.17 Oxycarenidae 1.7 3.39 2.38 19.04 3.82 0.9 2.17 4.77 Rhyparochromidae 30.43 21.74 34.48 45.5 56.52 45.76 57.14 35.71 47.62 30.95 24.84 27.93 32.61 37.79 Pyrrhocoridae 1.7 4.55 1.69 1.19 1.27 1.8 2.17 2.05 Stenocephalidae 2.38 0.9 1.64 Coreidae 13.04 5.2 4.55 6.78 4.76 2.38 3.18 4.51 2.17 5.17 Alydidae 2.38 2.38 Rhopalidae 4.35 3.4 9.52 1.27 1.8 2.17 3.75 Plataspidae 1.69 1.19 4.76 2.55 Cydnidae 4.35 8.8 9.09 8.69 5.08 4.76 5.95 4.76 7.14 3.18 4.51 8.7 6.25 Thyreocoridae 4.55 1.69 1.19 4.76 2.38 0.64 0.9 2.17 2.29 Scutelleridae 5.2 4.55 4.35 6.78 4.76 4.76 7.14 1.91 2.7 2.17 4.43 Pentatomidae 26.1 17.3 13.64 4.35 16.95 4.76 11.9 4.76 7.14 8.28 11.71 13.04 11.66 Acanthosomatidae 1.7 0.64 2.17 1.5 Authors: 1 – Verner, 1959, 2 – Křístek, 1991, 3 – Štepanovičová, Kovačovský, 1971, 4 – Bianchi, 1991, 5 – Štepanovičová, 1994, 6 – Štepanovičová, 1995, 7 – Majzlan, Štepanovičová, 1998, 8 – Štepanovičová, Degma, 1999, 9 – Majzlan et al., 2000, 10 – Štepanovičová, 2001, 11 – Rédei, Hufnagel, 2003a, 12 – Rédei, Hufnagel, 2003b, 13 – This work. Abbreviations: CZ-B – Czech Republic, Bohemia, CZ-M – Czech Republic, Moravia, SK – Slovakia, HU – Hungary, D.–.dominance

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floodplain forests along the Danube river with the doubled values (Štepanovičová, 1995). Low numbers (1–2 ex.) are also presented by Verner (1959), Olechowicz (2004) recorded the abundance of 0.9–3.3 (ex.m-2) in her research on soil fauna in shelter belts. An eudominant status was recorded at the mesophilous Legnotus limbosus, a typical epigeic species, being evaluated as euconstant bug at all the study sites except for LH. Eurygaster maura, an eudominant species, trophically connected with quite different habitats – grassland including cereals. However it uses the fallen leaf litter horizon to survive winter. The group of significantly common species includes also Plinthisus brevipennis, Megalonotus sabulicola, M. chiragra, Peritrechus nubilus or Raglius alboacuminatus. During researchs of epigeic bugs in the floodplain forest along the Dyje river in Moravia (Czech Republic) species Drymus brunneus and Scolopostethus thomsoni were dominated (Křístek, Dobšík, 1985), in floodplain forests of the Morava river Drymus brunneus, D. ryeii were dominated and in drier forests there was mainly Legnotus limbosus (Štepanovičová, 1994). The analysis has proved an occurrence of some faunistically interesting species such as Acalypta musci (Schrank, 1781) – European species, which occurs primarily in different mosses, in relatively humid places, woodland habitats but may be also found on fungi growing on tree trunks (Rédei et al., 2004). Ceraleptus gracilicornis (Herrich-Schaeffer, 1835), Holomediterranean species, trophically bound to Fabaceae, which shows a tendency towards necrophagy. Occurrence of this thermophilous species is probably limited to the southern part of Slovakia (Stehlík, Vavřínová, 1995). Loricula exilis (Fallén, 1807) – western Euro-Siberian species, predator inhabiting leaf litter or as arboricole on bark of coniferous trees covered by bryophytes or lichens, occasionally on Quercus or Fagus (Štepanovičová, Országh, 2002). This represents the 4th locality in the Malé Karpaty Mts and the 3rd published record in Slovakia. The comparison of bug species diversity at the sites shows, that the highest values appear at the site of Naháč–Kukovačník (NA) (moreover with the highest average abundance of bugs per 1m2) and Naháč–Katarínka (NK1, NK2). In the Nature Reserve Naháč–Katarínka (NK1) the lowest value of the dominance index was recorded what hints at equitable structure of dominance amongst more species in the assemblage and hence a suitable stage of the site. The same results refer to the research on the Nabidae taxocoenosis structure from this area published by Bulánková, Holecová (1998). The study site Lošonec–lom quarry has been influenced by limestone dust. The community had the 4th highest value in abundance of bugs (per 1 m2) in spite of that also the highest dominance index (disbalanced structure in dominance), a relatively low species richness (11), the second lowest index of diversity and relatively low equitability in the comparison with the other study sites. The values may hint at some degree of negative impact from the quarry on epigeic bugs, although due to dominating phytophagous species the effect is not as remarkable as in the case of zoophagous species of Nabidae, which decreases in their abundance because of a decline in their food (caterpillars) on leaves covered by immissions Bulánková, Holecová (1998), Holecová, Bulánková (1999, 2000). Apart from the variables mentioned above there were 15 environmental factors observed at the sites, which may influence a structure in the Heteroptera communities. Amongst all of them the E3 cover was the only significant. The RDA ordination shows 175

that the species Loricula exilis and Nabis pseudoferus more frequently occur in shadowy habitats while Tropistethus holosericeus, Legnotus limbosus, Plinthisus brevipennis, Raglius alboacuminatus, Trapezonotus arenarius or Microporus nigritus prefer sunny sites. Translated by P. J. Fedor Acknowledgement I am grateful to M. Holecová for granting me the Heteroptera material obtained in the forests of the Malé Karpaty Mts and of the Trnavská pahorkatina hills by the sifting and square method. This study was by the Slovak Scientific Grant Agency VEGA as a Project No. 1/0119/03.

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Štepanovičová, O., 2001: Bugs (Heteroptera) in the woody steppes habitats of the Nature Rezerve Kňaží vrch in Tematínske kopce (Považský Inovec Mts) (in Slovak). Folia Faunistica Slovaca, 6, p. 75–79. Štepanovičová, O., Országh, I., 2002: To the occurrence of bug Myrmedobia exilis (Heteroptera, Microphysidae) in Slovakia (in Slovak). Folia Faunistica Slovaca, 7, p. 35–38. Štepanovičová, O., Bianchi, Z., 2003: On the fauna of bugs (Heteroptera) of blown sands in the Podunajská nížina lowland (in Slovak). Entomofauna Carpathica, 15, p. 35–40. Štepanovičová, O., Bianchi, Z., 2005: Bugs (Heteroptera) (in Slovak). In Majzlan (ed.): Fauna Devínskej Kobyly. APOP, Bratislava, p. 75–82. Ter Braa, C.J.F., Šmilauer, P., 1998: CANOCO References Manual and User’s Guide to Canoco for Windows: Software for Canonical Community Ordination (version 4). Microcomputer Power (Ithaca, NY, USA), 352.pp. Tischler, W., 1949: Grundzüge der terrestrischen Tierökologie. Friedrich Vieweg, Braunschweig, 219 pp. Verner, P.H., 1959: Study on arthropods living in soil of oak-hornbeam forest in environment of Karštejn (in Czech). Bohemia Centralis, A, 1, (7), p. 345–408. Wishart, D., 1969: An algorithm for hierarchical classifications. Biometrics, 22, p. 165–170. Zlinská, J., Šomšák, L., Holecová, M., 2005: Ecological characteristics of studied forest communities of an oak-hornbeam tier in SW Slovakia. Ekológia (Bratislava), 24, Suppl. 2, p. 3–19. Received 30. 7. 2005

Hradil K.: Společenstva ploštic v epigeonu dubo-habrových lesů v oblasti JZ Slovenska. Rozborem sběrů epigeické fauny ploštic na osmi lokalitách v Malých Karpatech a dvou lokalitách v Trnavské pahorkatině v letech 1999–2002 bylo zjištěno celkem 46 druhů náležejících do 17 čeledí. Ploštice byly sbírány kvadrátovou metodou a prosevy. Průměrná abundance byla 2,44 jedinců na 1 m2. U jednotlivých druhů byla stanovena jejich dominance a konstance. Na základě výsledků RDA analýzy z 15 testovaných proměnných byl potvrzen signifikantní vliv zápoje korun (E3) na složení společenstev ploštic. S eudominantním zastoupením byli zjištěny druhy Legnotus limbosus a Eurygaster maura, dominantní a subdominantní byly druhy Plinthisus brevipennis, Megalonotus sabulicola, M. chiragra, Peritrechus nubilus nebo Raglius alboacuminatus. Faunisticky zajímavý byl nález druhu Myrmedobia exilis.

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Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 179–204

STRUCTURE AND FUNCTION OF WEEVIL ASSEMBLAGES (Coleoptera, Curculionoidea) IN EPIGEON OF OAK-HORNBEAM FORESTS IN SW SLOVAKIA MILADA HOLECOVÁ1, DANKA NÉMETHOVÁ2, MATÚŠ KÚDELA1 .Department of Zoology, Comenius University, Mlynská dolina B–1, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected], [email protected] 2 .Centre for Biostatistics & Analyses, Faculty of Medicine and Faculty of Natural Sciences, Masaryk University, Kamenice 3, 625 00 Brno, The Czech Republic, e-mail: [email protected] 1

Abstract Holecová M., Némethová D., Kúdela M.: Structure and function of weevil assemblages (Coleoptera, Curculionoidea) in epigeon of oak-hornbeam forests in SW Slovakia. Ekológia (Bratislava), Vol. 24, Suppl.2/2005, p. 179–204. In 1999–2002, we studied the weevil assemblages in the epigeon of oak-hornbeam forests in SW Slovakia, with regard to assemblage structure, species richness, seasonal occurrence and ecological requirements of individual species. The investigation was performed on 10 study plots (forest stands of the age from 40 to 100 years) in the Malé Karpaty Mts and Trnavská pahorkatina hills, SW Slovakia. Each site was affected, to some extent by human activities of which forest fragmentation and pollution with calcareous dust were the most important. The soil macrofauna was collected by sifting the litter, surface layer of soil and remnants of twigs. Altogether 4,090 individuals of weevils classified into 78 species, 43 genera and 4 families were recorded. The number of species found at the particular sites ranged from 22 to 31 species. The mean abundance of weevils varied from 2.09 to 49.71 ind.m-2. Geophilous species Ceutorhynchus pallidactylus, geobionts Acalles fallax, Barypeithes chevrolati, Barypeithes mollicomus, Brachysomus hirtus, Trachodes hispidus and geoxenes Ceutorhynchus obstrictus and Sitona macularius predominated in the material from the oak-hornbeam forest epigeon. The canopy architecture and content of exchange bases in the leaf litter from amongst 13 gradient and 2 categorial variables analysed had direct and significant influence on the structure of epigeic weevil assemblages. The forest fragmentation caused increase the abundance of herbicolous, euryhygric and ubiquitous species in the epigeon of island woods. The weevil community was less abundant and unstable in the forest site polluted with calcareous dust. Key.words:.weevils, Curculionoidea, Coleoptera, epigeic assemblages, oak-hornbeam forests, SW Slovakia, ecology, human impacts

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Introduction The oak-hornbeam forests in Slovakia used to be the most frequent forest climatic zone formation at lower altitudes. In the past they covered continuous and large areas, especially in plains and lowlands from the altitude of approximately 100 m above sea level, in hilly and submountainous regions up to 600 m above sea level and in all the Inner–Carpathian hollows (Michalko, 1986). These forest stands in Slovakia as well as in the other countries of Europe have been under long-term human impact. There are just some fragments under considerable anthropogenic pressure in the present agricultural land. In the cultivated land they represent a refuge for many animal species. Weevils belong to predominant beetle groups in the ecosystem of deciduous forests in Europe (Funke, 1971; Grimm, 1976; Schauermann, 1973, 1976; Nielsen, 1978 a, b, c; Holecová, 1991b, 1992; Dajoz, 2000, etc.). Geobiont species associated with forest floor and leaf litter are important phyllophages, mycophages but also decomposers of dead and decaying wood. They have an influence on soil and vegetation, both directly and indirectly (Wallwork, 1976; Speight, 1989; Dajoz, 2000). Due to their low mobility and inability to take flight they often live in isolated populations. They are sensitive indicators of negative human impacts such as forest fragmentation, clear cutting, pollution, disturbance, changes of soil moisture and vegetation (Holecová, 1986). Soil beetles in the forests of the oak-hornbeam vegetation tier in various parts of Slovakia were studied by Drdul (1973, 1977, 1997), Kožíšek, Drdul (1991), Majzlan (1986, 1991). Majzlan, Hošták (1996), Holecová (1991a, 1995), Holecová, Sukupová (2000, 2002), Holecová et al. (2002) focused on epigeic weevils. Our research originated in the grant being concentrated on animal communities in oak-hornbeam forests in SW Slovakia. Several studies of various groups of soil microfauna (ciliates, naked amoebae) and arthropods (ants, spiders, bugs) in this territory have been already published (Holecová et al., 2005; Krumpálová, Bartoš, 2002; Krumpálová, Szabová, 2003; Mrva, Matis, 2000; Mrva, 2003; Štepanovičová, Országh, 2002; Tirjaková, Bartošová, 2004; Tirjaková, Mrva, 2005; Tirjaková et al., 2002). The present study gives the results of four-year investigation (1999–2002) on weevil assemblages (Coleoptera, Curculionoidea) in epigeon of oak-hornbeam forests in SW Slovakia. The aims of the study were the following: { to characterise structure, dynamics and seasonal occurrence of weevil communities; { to analyse representation of topic, trophic and ecological–bionomic groups; { to find out biotic, abiotic and anthropic factors with an influence on weevil epigeic assemblages.

Study area The investigation refers to 10 study plots (the age of forest stands varies from 40 to 100 years) in Malé Karpaty Mts and Trnavská pahorkatina hills (SW Slovakia). All the study plots are affected to a certain degree by human activities. 180

Cajla (CA), 48°20' N, 17°16' E, GRN (Grid Reference Number of the Databank of the Fauna of Slovakia) 7669c, 260–280 m a.s.l.: an 80–100 year old forest at the foot of the Malá cajlanská homola hill, oriented onto S and neighbouring meadows and vineyards on S and E, from N and W closed forest complexes. Quercus dalechampii and Carpinus betulus predominate in the tree layer. Vinosady (VI), 48°19' N, 17°17' E, GRN 7669d, 280 m a.s.l.: a 60–80 year old forest at the foot of the Kamenica hill, oriented onto NW, W neighbouring drier subxerophilous meadows and shrub complexes. Besides Quercus dalechampii, the tree stratum consists of Q. cerris and Acer campestre. Fúgelka (FU), 48°22' N, 17°19' E, GRN 7669b, 350 m a.s.l.: an 80–100 year old forest near the Dubová village, oriented onto S. Besides Quercus dalechampii, the tree stratum consists of Acer pseudoplatanus. Lindava (LI) (Nature Reserve), 48°22' N, 17°22' E, GRN 7670a, 240 m a.s.l.: an 80–100 (120) year old forest near the village of Píla. Quercus dalechampii and Q. cerris predominate in the tree layer. Horný háj (HH), 48°29' N, 17°27' E, GRN 7570b, 240 m a.s.l.: a larger complex of an island forest 60–80 years old near the village of Horné Orešany, surrounded by fields and vineyards, oriented onto W and SW. Quercus cerris, Q. dalechampii, Carpinus betulus and Fraxinus excelsior predominate in the tree layer. Lošonec–lom quarry (LL), 48°29' N, 17°23' E, GRN 7570b, 340 m a.s.l.: an 80–100 year old forest oriented onto SW, neighbouring mesophilous meadows and pastures. The tree layer consists of Quercus dalechampii, Q. cerris and Carpinus betulus. The leaf litter, herbage undergrowth and trees are heavily covered with calcareous dust from a nearby quarry. Lošonský háj grove (LH) (Nature Reserve), 48°28' N, 17°24' E, GRN 7570b, 260 m a.s.l.: an 80–100 year old oak-hornbeam forest oriented onto NE, surrounded by closed forest complexes. Quercus dalechampii, Q. cerris and Carpinus betulus predominate in the tree stratum. Naháč–Kukovačník (NA), 48°32' N, 17°31' E, GRN 7471c, 300 m a.s.l.: a small forest island, approximately 40–60 year old surrounded by fields and pastures, oriented onto NE. Quercus dalechampii, Q. cerris and Carpinus betulus predominate in the tree layer. Naháč – Katarínka 1 (NK1) (Nature Reserve), 48°33' N, 17°33' E, GRN 7471a, 340 m a. s. l.: a 40–60 year old forest oriented onto NW, surrounded by closed forest ecosystems. Quercus dalechampii and Carpinus betulus predominate in the canopy. Naháč–Katarínka 2 (NK2), (Nature Reserve), 48°33' N, 17°32' E, GRN 7471a, 300 m a.s.l.: an 80–100 year old forest below the monastery ruins. Quercus virgiliana, Q. cerris and Tilia cordata predominate in the tree stratum. The study plots LI and HH are situated in the Trnavská pahorkatina hills, the other ones in Malé Karpaty Mts. Managed oak-hornbeam forests cover the study area. According to the Geobotanic Map of Slovakia (Michalko, 1986) Carpathian mesophilous oak-hornbeam woods (Carpinion betuli I s s l e r 1931 em. M a y e r 1937) (the study plots CA, VI, FU, LL, LH, NA, NK1), subxerothermophilous forests with 181

European turkey oak (Quercion confertae–cerris H o r v a t 1954) (the study plot LI), and Submediterranean xerothermophilous woods (Quercion pubescentis–petrae B r.–B l. 1931) (NK2) are considered as typical natural vegetation in this territory. The map, pedological and phytocoenological characteristics of the investigated area are given in detail by Zlinská et al. (2005).

Material and methods The soil macrofauna was collected by the square method combined with sifting. At each study site, in about one-month intervals from March to November, the material was collected from the leaf litter and upper part of soil from 16 squares. Each square comprised 25x25 cm, i.e. altogether an area of 1m2 was sifted, representing one sample. The samples were extracted using xereclectors of the Moczarski’s type. The material is deposited at the Department of Zoology, Comenius University in Bratislava. In total, 310 samples were used for statistical analysis. The species dominance is characterised by the scale proposed by Tischler (1949) and completed by Heydemann (1955): ED = eudominant, D = dominant, SD = subdominant, R = recedent, SR = subrecedent. The species constancy was expressed by categories according to Tischler (1949) and Schwerdtfeger (1975): EC = euconstant, C = constant, As = accessoric and A = accidental. The indices of Shannon–Wiener (H'), Pielou (e) and Simpson (c) were used as the alpha diversity indices (Odum, 1977; Spellerberg, Fedor, 2003). All the couples of Shannon–Wiener’s diversity indices were compared with a t–test (Poole, 1974). The trophic groups of weevil adults were established according to Brown, Hyman (1986). Four basic groups we distinguished: S1 = monophages, S2 = narrow oligophages, S3 = wider oligophages; and G = polyphages (S1 – S3 = specialists and G = generalists). Other ecological characteristics of weevil adults (habitat preference, humidity preference, topic groups, bionomic groups, relationship between weevil imagoes and subsoil) are given according to Koch (1992). The cluster analysis of weevil communities was done using the computer program NCLAS (Podani, 1993). The clustering method complete linkage in combination with Wishart’s similarity ratio was used (Wishart, 1969). Effects of environmental variables on weevil community composition were analysed using the redundancy analysis (RDA) ordination technique by CANOCO software program (ter Braak, Šmilauer, 1998). The values of species data used in the cluster analysis and RDA were transformed with log–transformation Y' = log (Y + 1). At all the study plots the following environmental variables were measured (range of values of gradient variables or categories of categorial ones are in brackets). Gradient variables: pedological and chemical characteristics of leaf litter such as total organic carbon (3.80–14.40%); total nitrogen content (0.35–1.1%); content of exchange bases (Ca2+, Mg2+, Na+, K+) (12.60–49.40 mval/100g); total acidity (1.8–14.9 mval/100g); pH of litter in H2O (3.90–6.74); humus content (6.55–24.83%); age of a forest stand (40 – 100 years); cover of E3: (65–80%); cover of E2 (1–50%); cover of E1 (40–100%); lying and standing dead wood (2–20%); forest fragmentation (0–50%); slope (0–45°). Categorial variables: exposition of study plots (E, W, S, NE, NW, SW, SE); pollution by calcareous dust (present, absent). Mean abundance and number of species among the localities were compared using single factor analysis of variance and Tukey’s HSD test at significance level 0.05. The tests were performed with SPSS 11.0 for Windows. Nomenclature of weevils was assumed according to Alonso–Zarazaga, Lyal (1999).

182

Results and discussion Community structure A total of 4,090 individuals of 78 species, 43 genera and 4 families (Anthribidae, Apionidae, Erirhinidae, Curculionidae) were examined during the four-year research (Table 1). At the study sites 22 to 31 species were recorded. The mean abundance of weevils varied from 2.09 (the study plot LL) to 49.71 ind.m-2 (the study plot HH) (Table 2). The geobiont Acalles fallax, Barypeithes mollicomus and geophilous Ceutorhynchus pallidactylus occurred as eudominants, geoxenous C. obstrictus as dominant, geobiont Barypeithes chevrolati, Brachysomus hirtus, Trachodes hispidus and geoxenous Sitona macularius as subdominants. The characteristic species spectrum is represented by three groups of species: 1. highly dominant species (eudominant, dominant, subdominant) with a high constancy (euconstant or constant) – geobionts Acalles fallax, Barypeithes mollicomus, Trachodes hispidus, geophilous species Ceutorhynchus pallidactylus and geoxenous species C. obstrictus, Sitona macularius; 2. euconstant or constant, non–dominant species – saproxylic geobionts Acalles camelus, Ruteria hypocrita and geoxenous species Ceutorhynchus alliariae, C..scrobicollis (herbicoles associated with Alliaria petiolata), Coeliodes trifasciatus, Curculio glandium, Furcipus rectirostris, Phyllobius argentatus (arboricolous species associated with tree canopy); 3. differential species – saproxylic geobionts Acalles echinatus, Acallocrates colonnellii, phyllophagous geobionts Barypeithes albinae, Brachysomus dispar, B.rachysomus setiger, and Coeliodes proximus, C. rana, C. transverseoalbofasciatus (arboricolous species associated with oaks but aestivating in leaf litter). The first two groups of species have a wider ecological amplitude. The third group is represented by stenovalent species with only a low dominance and constancy and associated exclusively or namely with forests of the oak-hornbeam vegetation tier. Seasonal occurrence and dynamics Abundance of weevils in soil reflected two peaks: vernal (May – the 1st half of June) and serotinal to autumnal (August–September). The spring peaks were determined by typical geobionts of the Barypeithes and Brachysomus genera. The peaks in serotinal and autumnal were defined by some xylodetriticoles such as the species of Acalles, Ruteria genera, Trachodes hispidus and geophilous Ceutorhynchus pallidactylus. Seasonal dynamics in species richness corresponds with abundance dynamics as well. The peak appeared in vernal aspect (May–June) with species richness of 15 geobionts and 5 (in May), resp. 2 geophiles (in June). The second peak in species richness hints at serotinal (August) with a frequent occurrence of saproxylic geobionts ( Acalles spp., Ruteria 183

T a b l e 1. List of weevil species and numbers of individuals found in the epigeon of oak-hornbeam forests in SW Slovakia during 1999–2002. Family, species / study plot CA VI FU LI HH LH Anthribidae Anthribus albinus (L i n n a e u s, 1758) 1 0 0 0 0 0 Apionidae Catapion seniculus (K i r b y, 1808) 0 0 0 0 0 0 Ceratapion gibbirostre (G y l l e n h a l, 1813) 1 0 0 0 0 1 Cyanapion columbinum (G e r m a r, 1817) 0 0 0 0 0 0 Diplapion confluens (K i r b y, 1808) 0 0 0 0 2 0 Eutrichapion punctigerum (P a y k u l l, 1792) 1 0 0 0 0 0 Holotrichapion aestimatum (F a u s t, 1890) 0 0 0 0 0 0 0 0 Holotrichapion pisi (F a b r i c i u s, 1801) 0 0 0 0 Ischnopterapion virens (H e r b s t, 1797) 1 0 0 0 0 0 Kalcapion pallipes (K i r b y, 1808) 0 0 0 0 2 0 Omphalapion hookerorum (K i r b y, 1808) 0 0 0 0 2 0 Protapion apricans (H e r b s t, 1797) 0 0 0 0 0 0 Protapion fulvipes (G e o f f r o y, 1785) 4 2 1 0 0 1 Protapion trifolii (L i n n a e u s, 1768) 0 0 0 0 0 0 Protapion nigritarse (K i r b y, 1898) 0 1 0 0 0 0 Pseudapion rufirostre (F a b r i c i u s, 1775) 1 0 0 0 0 0 Synapion ebeninum (K i r b y, 1808) 0 0 0 0 2 0 Trichopterapion holosericeum (G y l l e n h a l, 2 0 0 0 0 1 1833) Erirhinidae Tanysphyrus lemnae (P a y k u l l, 1792) 0 0 0 0 0 0 Curculionidae Acalles camelus (F a b r i c i u s, 1792) 0 0 1 1 0 18 Acalles fallax B o h e m a n 1844 0 0 58 0 40 62 Acalles echinatus (G e r m a r, 1824) 16 1 0 20 0 6 Acallocrates colonnellii B a h r, 2003 0 0 0 2 6 0 Barypeithes albinae F o r m a n e k, 1903 0 0 8 69 1 0 Barypeithes chevrolati (B o h e m a n, 1843) 50 30 5 0 0 28 Barypeithes mollicomus (A h r e n s, 1812) 0 0 0 0 455 7 Bradybatus creutzeri G e r m a r, 1824 0 1 0 0 0 0 Bradybatus fallax G e r s t a e c k e r, 1860 0 0 1 0 0 0 Bradybatus kellneri B a c h, 1854 0 2 0 0 0 0

LL NA NK1 NK2 total

%

CD

Co

CC Geo Top Tro Bio-i Hab Hum

0

0

1

0

2

0.05 SR 20.00 A GB T

G

XF

ST HY

1 0 0 0 0 0 0 1 0 0 1 1 2 0 0 0 0

1 0 1 0 0 1 7 0 0 0 0 2 0 0 0 0 0

0 1 0 0 0 0 1 0 0 0 0 2 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0

2 3 1 2 1 1 8 2 2 2 1 14 2 1 1 2 3

0.05 0.07 0.02 0.05 0.02 0.02 0.20 0.05 0.05 0.05 0.02 0.34 0.05 0.02 0.02 0.05 0.07

S2 S3 S2 S3 S2 S2 S2 S2 S1 S3 S1 S2 S2 S2 S2 S3 S1

FF FF FF FF FF FF FF FF FF FF FF FF FF FF FF FF FF

EU EU ST EU EU ST EU EU ST EU EU UB EU EU ST EU ST

0

1

0

0

1

0.02 SR 10.00 A GX H S2

2 4 0 0 0 0 1 0 0 0

14 4 32 72 1.76 R 70.00 C 74 69 98 405 9.90 ED 70.00 C 0 0 0 43 1.05 R 40.00 As 0 0 6 14 0.34 SR 30.00 As 0 0 0 78 1.91 R 30.00 As 0 0 0 113 2.76 SD 40.00 As 9 191 118 781 19.10 ED 60.00 C 0 0 0 1 0.02 SR 10.00 A 0 0 0 1 0.02 SR 10.00 A 0 1 0 3 0.07 SR 20.00 A

SR SR SR SR SR SR SR SR SR SR SR SR SR SR SR SR SR

20.00 30.00 10.00 10.00 10.00 10.00 20.00 20.00 20.00 20.00 10.00 80.00 10.00 10.00 10.00 10.00 20.00

A As A A A A A A A A A EC A A A A A

GX GX GX GX GX GX GX GX GX GX GX GX GX GX GX GX GX

GB GB GB GB GB GB GB GX GX GX

H H H H H H H H H H H H H H H H A

T T T T T T T A A A

G G G S3 ? ? G S2 S2 S2

EH XE XE XE XE XE EH HY XE EH EH XE XE XE EH EH

FF ST HY XF ST HY XF ST HY XF ST HY XF ST HY FF ST HY FF ST HY FF EU HY FF ST XE FF ST EH FF ST EH

T a b l e 1. (Continued) Family, species / study plot Brachysomus echinatus (B o n s d o r f f, 1785) Brachysomus dispar P e n e c k e, 1910 Brachysomus hirtus (B o h e m a n, 1845) Brachysomus setiger (G y l l e n h a l, 1840) Calosirus apicalis (G y l l e n h a l, 1727) Ceutorhynchus alliariae C h. B r i s o u t, 1860 Ceutorhynchus chalibeus G e r m a r, 1824 Ceutorhynchus erysimi (F a b r i c i u s, 1787) Ceutorhynchus minutus (R e i c h, 1797) Ceutorhynchus obstrictus (M a r s h a m, 1802) Ceutorhynchus pallidactylus (M a r s h a m, 1802) Ceutorhynchus rhenanus S c h u l t z e, 1895 Ceutorhynchus scrobicollis N e r e s et W.a.g.n.e.r, 1924 Ceutorhynchus typhae (H e r b s t, 1795) Coeliodes proximus S c h u l t z e, 1895 Coeliodes rana (Fabricius, 1787) Coeliodes transverseoalbofasciatus G o e z e, 1777 Coeliodes trifasciatus B a c h, 1854 Curculio glandium (M a r s h a m, 1802) Curculio pellitus (B o h e m a n, 1843) Curculio venosus (G r a v e n h o r s t, 1807) Furcipus rectirostris (L i n n a e u s, 1758) Hypera nigrirostris (F a b r i c i u s, 1775) Hypera postica (G y l l e n h a l, 1813) Kyklioacalles suturatus (D i e c k m a n n, 1983) Leiosoma cribrum (G y l l e n h a l, 1834) Microplontus campestris (G y l l e n h a l, 1837) Nedyus quadrimaculatus (L i n n a e u s, 1758) Ophrohinus suturalis (F a b r i c i u s, 1775) Otiorhynchus ovatus (L i n n a e u s, 1758)

CA 0 0 0 0 0 0 0 4 0 3 58

VI FU LI HH LH 3 1 0 0 0 0 0 0 3 23 0 0 1 3 0 7 0 0 0 0 1 0 0 0 0 1 0 0 12 0 0 0 0 3 0 1 8 3 7 4 1 0 0 0 1 5 16 48 30 2 13 27 36 839 84

LL 5 0 0 0 0 1 4 0 0 0 5

NA NK1 NK2 0 0 20 0 0 0 16 0 134 0 0 0 0 0 0 2 1 0 0 2 0 2 8 2 0 0 0 84 100 16 94 151 39

total 29 26 154 7 1 17 9 39 2 304 1346

% 0.71 0.64 3.77 0.17 0.02 0.42 0.22 0.95 0.05 7.43 32.91

CD Co SR 40.00 SR 30.00 SD 40.00 SR 10.00 SR 10.00 SR 50.00 SR 30.00 SR 90.00 SR 20.00 D 90.00 ED 100.00

CC Geo Top As GB T As GB T As GB T A GB T A GX H C GX H As GX H EC GX H A GX H EC GX H EC GF H

Tro Bio-i G FF ? FF ? FF G FF S3 FF S1 FF S3 FF S3 FF S3 FF S3 FF S3 FF

Hab EU ST ST ST ST ST ST UB UB UB UB

Hum EH XE XE XE HY HY XE EH EH EH EH

0 0

2 2

0 0

0 1

0 2

0 0

0 3

0 0

0 4

0 39

2 51

0.05 SR 10.00 A GX H S2 1.25 R 60.00 C GX H S1

FF FF

ST XE ST HY

4 0 1 0

2 0 0 0

2 0 0 0

3 0 2 2

10 0 3 0

2 0 0 0

1 0 0 0

3 0 0 0

11 0 8 1

5 4 0 0

43 4 14 3

1.05 R 10.00 A 0.10 SR 10.00 A 0.34 SR 40.00 As 0.07 SR 20.00 A

GX GX GX GX

H A A A

S3 S2 S2 S2

FF FF FF FF

UB ST ST ST

EH XE XE XE

3 7 2 0 1 0 0 4 0 0 0 0 0

1 4 0 0 0 0 0 3 0 0 0 0 0

1 3 0 0 0 3 0 4 0 0 0 0 0

5 5 0 0 0 0 0 0 0 1 0 0 0

11 2 0 0 0 0 2 0 1 0 1 1 0

1 2 0 4 0 0 0 2 1 0 0 0 1

1 1 0 0 3 1 0 0 0 0 0 0 1

3 1 0 0 3 2 1 0 0 0 0 0 0

4 2 0 0 1 0 0 0 0 0 0 0 0

0 0 0 0 1 0 0 0 0 0 1 0 0

30 27 2 4 9 6 3 13 2 1 2 1 2

0.73 0.66 0.05 0.10 0.22 0.15 0.07 0.32 0.05 0.02 0.05 0.02 0.05

GX GX GX GX GX GX GX GB GF GX GX GX GF

A A A A A H H T H H H H HA

S2 S2 S2 S2 S2 S2 S3 G S2 S1 S1 S2 G

FF FF FF FF FF FF FF XF FF FF FF FF FF

ST EU ST ST ST EU EU ST ST ST UB ST EU

XE EH XE EH HY EH EH HY HY XE EH XE XE

SR SR SR SR SR SR SR SR SR SR SR SR SR

90.00 90.00 10.00 10.00 50.00 30.00 20.00 40.00 20.00 10.00 20.00 10.00 20.00

EC EC A A C As A As A A A A A

T a b l e 1. (Continued) Family, species / study plot Otiorhynchus raucus (F a b r i c i u s, 1777) Phyllobius argentatus (L i n n a e u s, 1758) Phyllobius maculicornis G e r m a r, 1824 Polydrusus marginatus S t e p h e n s, 1831 Polydrusus viridicinctus G y l l e n h a l, 1834 Rhinoncus bruchoides (H e r b s t, 1784) Rhinoncus perpendicularis (R e i c h, 1797) Rhynchaenus pilosus (F a b r i c i u s, 1781) Ruteria hypocrita (B o h e m a n, 1837) Sciaphilus asperatus (B o n s d o r f f, 1785) Simo variegatus (B o h e m a n, 1843) Sitona humeralis S t e p h e n s, 1831 Sitona lineatus (L i n n a e u s, 1758) Sitona macularius (M a r s h a m, 1802) Stenocarus cardui (H e r b s t, 1784) Strophosoma melanogrammum (F o r s t e r, 1771) Trachodes hispidus (L i n n a e u s, 1758) Trachyphloeus bifoveolatus (B e c k, 1817) Tychius picirostris (F a b r i c i u s, 1787) Total

CA 0 3 0 1 0 0 0 0 4 0 0 0 0 1 0 1

VI FU LI HH LH LL 0 0 0 0 0 0 1 3 2 9 0 3 0 0 1 0 0 0 0 1 8 3 0 3 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 2 0 1 5 1 4 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 0 0 0 0 0 27 1 0 5 6 3 40 1 9 0 0 0 0 0 0 4 5 0 0 0 2

2 0 0 176

0 24 6 22 0 0 0 0 0 1 0 0 93 184 221 1550

9 0 0 269

NA NK1 NK2 1 5 9 7 10 4 0 0 0 0 0 3 0 3 2 1 0 0 1 0 0 0 0 0 4 1 12 0 0 1 0 0 1 0 0 0 0 0 0 10 3 6 0 2 0 0 0 0

total 15 42 1 19 6 1 1 3 37 1 1 4 28 84 2 12

% 0.37 1.03 0.02 0.46 0.15 0.02 0.02 0.07 0.90 0.02 0.02 0.10 0.68 2.05 0.05 0.29

CD SR R SR SR SR SR SR SR SR SR SR SR SR SD SR SR

Co 30.00 90 10 60.00 30.00 10.00 10.00 20.00 90.00 10.00 10.00 10.00 20.00 100.00 10.00 40.00

CC Geo Top As GF TH EC GX A A GX A C GX A As GX A A GX H A GX H A GX A EC GB T A GF H A GX A A GX H A GX H EC GX H A GX H As GX A

Tro Bio-i G FF G FF G FF G FF S2 FF S2 FF S2 FF S2 FF G XF G FF G FF S2 FF S2 FF S2 FF S2 FF G FF

4 12 27 3 109 2.67 SD 90.00 EC GB T G 7 0 0 0 7 0.17 SR 10.00 A GB T G 0 0 0 1 2 0.05 SR 20.00 A GX H S2 68 357 614 558 4090 100.00

Hab EU EU EU EU ST ST EU ST ST EU ST EU EU EU ST EU

Hum EH EH EH XE XE XE EH EH HY HY XE EH EH XE XE EH

XF EU EH FF EU XE FF EU NE

Symbols and abbreviations: % – dominance; CD – category of dominance: ED – eudominant, D – dominant, SD – subdominant, R – recedent, SR – subrecedent; Co – constancy in %; CC – category of constancy: EC – euconstant, C – constant, As – accessoric, A – accidental; Geo – interaction of weevils to subsoil: GB – geobiont, GF – geophilous, GX – geoxenous; Top – topic group: T – terricolous, TH – terriherbicolous, H – herbicolous, HA – herbiarboricolous; A – arboricolous; Tro – tropic group: S1 – monophages, S2 – narrow S2 – narrow oligophages, S3 – wider oligophages, G – generalists (polyphages); Bio-i - bionomical group of imagoes: XF – xylophagous (wood-eating), FF – phyllophagous (leaf-eating); Hab – habitat preference: ST – stenotopic, EU – eurytopic, UB – ubiquitous; Hum – humidity preference: HY – hygrophilous, XE – xerophilous, EH – euryhygric. Abbreviations of study plots see in Material and methods.

T a b l e 2. Species diversity test and basic coenological characteristics of weevil assemblages at study plots in 1999–2002. Study plot Σ spp. Σ GB spp. Σ GF spp. Σ GX spp. MA±SE MAGB±SE MAGF±SE MAGX±SE e c H' CA VI FU LI HH LH LL NA NK1 NK2

CA 25 6 1 18 5.7±1.23 2.5±0.82 1.9±0.79 1.2±0.28 0.67 0.2 2.154 2.077* 1.339ns 0.306ns 6.435*** 0.283ns 5.600*** 0.213ns 1.459ns 0.600ns

VI 23 7 1 15 3.0±0.80 1.6±0.75 0.4±0.18 1.0±0.21 0.8 0.14 2.493 211.390 1.047ns 2.536* 8.002*** 2.064* 2.911** 2.199* 3.754*** 2.023*

FU 22 8 1 13 5.8±0.88 3.4±0.52 0.9±0.31 1.5±0.34 0.76 0.15 2.334 343.528 178.880 1.874* 9.576*** 1.263ns 4.779*** 1.422ns 3.538*** 1.155ns

LI 22 6 1 15 7.0±1.10 3.2±0.91 1.1±0.52 2.7±0.47 0.68 0.18 2.114 351.344 173.556 396.939 7.519*** 0.693ns 6.596*** 0.636ns 1.345ns 1.165ns

HH 31 8 2 21 49.7±16.60 17.2±7.35 27.1±14.25 5.4±0.95 0.42 0.38 1.444 224.119 110.143 260.706 323.340 8.885*** 14.271*** 9.679*** 8.823*** 13.157***

LH 25 9 3 13 8.2±1.25 5.0±0.88 2.7±0.94 0.7±0.22 0.68 0.18 2.191 346.788 164.405 411.999 476.095 413.721 6.166*** 0.100ns 2.270* 0.361ns

LL 25 6 2 17 2.2±0.48 0.7±0.25 0.0±0.03 1.2±0.29 0.92 0.06 2.959 201.697 159.250 168.322 163.459 87.224 153.328 6.518*** 8.780*** 6.732***

NA 26 6 2 18 10.5±1.51 4.0±0.63 2.9±0.85 4.0±0.97 0.66 0.18 2.181 322.251 148.993 399.314 482.724 605.977 583.636 134.887 2.352* 0.519ns

NK1 26 6 2 18 17.2±3.22 9.4±2.37 4.9±2.04 5.0±1.36 0.6 0.2 1.987 258.944 122.857 315.153 392.947 1358.224 504.455 102.679 725.889

NK2 26 8 3 15 18.1±2.92 13.6±2.80 1.5±0.52 2.8±0.43 0.7 0.15 2.222 248.298 119.077 298.008 370.527 1321.477 473.986 98.031 679.726 1165.871

3.514***

Symbols and abbreviations: Σ spp. – total number of species, Σ GB spp. – total number of geobiont species, Σ GF spp. – total number of geophilous species, Σ GX spp. – total number of geoxenous species, MA [ind.m-2] – mean abundance of weevils, MAGB [ind.m-2] – mean abundance of geobionts, MAGF [ind.m-2] – mean abundance of geophilous species, MAGX [ind.m-2] – mean abundance of geoxenes, H' – Shannon’s index of species diversity, e – Pielou’s index of equitability, c – Simpson’ index of dominance. T-test values are under the diagonal and degrees of freedom are above it. Significance levels: *** = P < 0.001; ** = 0.001 < P < 0.01; * = 0.01 < P < 0.05; ns = 0.05 < P (non-significant). Abbreviations of the study plots see in Material and methods.

hypocrita, Trachodes hispidus), geophiles (Otiorhynchus raucus, Ceutorhynchus pallidactylus). However, a rapid increase of number of geoxenous species (Fig. 1) is significant as well. The first group is formed by geoxenes, which migrate into the forest ecosystem from the nearby open habitats in the period of mild or severe drought and later to hibernate. Geoxenous species from the upper forest strata create the second group, which includes particularly arboricolous species, often aestivating in forest soil (e.g. the genera Coeliodes and Curculio from oaks, Furcipus rectirostris from wild cherries, an arboricolous generalist Strophosoma melanogrammum, etc.). The Table 3 presents the seasonal occurrence of the species.

Fig. 1. Cumulative numbers of weevil species recorded in individual months of the field study (1999–2002). Explanations: T – total number of weevil species, G – geobionts, GF – geophiles, GX – geoxenes.

Ecological requirements of the weevil assemblages According to the relationship between the weevil adults and subsoil, the curculionids were classified into three groups: geobionts living permanently in leaf litter, geophilous species living in other forest strata but occurring regularly in soil and geoxenous species occurring in leaf litter accidentally. The geoxenous species predominated qualitatively (by species number), but the geophilous species predominated quantitatively (by number of individuals). The quantitative proportion of geobiont and geophilous individuals is balanced (Fig. 2).

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T a b l e 3. Seasonal occurrence of individual weevil species in epigeon of studied oak-hornbeam forests in SW Slovakia. Family, species / month of occurrence 3 4 5 6 7 8 9 10 11 Anthribidae + Anthribus albinus (L i n n a e u s, 1758) + Apionidae Catapion seniculus (K i r b y, 1808) + Ceratapion gibbirostre (G y l l e n h a l, 1813) + + + Cyanapion columbinum (G e r m a r, 1817) + Diplapion confluens.luens (K i r b y, 1808) + + Eutrichapion punctigerum (P a y k u l l, 1792) + Holotrichapion aestimatum (F a u s t, 1890) + Holotrichapion pisi (F a b r i c i u s, 1801) + + + + + Ischnopterapion virens (H e r b s t, 1797) + + + Kalcapion pallipes (K i r b y, 1808) + Omphalapion hookerorum (K i r b y, 1808) + + Protapion apricans (H e r b s t, 1797) + Protapion fulvipes (G e o f f r o y, 1785) + + + + + + + Protapion trifolii (L i n n a e u s, 1768) Protapion nigritarse (K i r b y, 1898) + Pseudapion rufirostre (F a b r i c i u s, 1775) + Synapion ebeninum (K i r b y, 1808) + Trichopterapion holosericeum (G y l l e n h a l, 1833) + + + Erirhinidae Tanysphyrus lemnae (P a y k u l l, 1792) + Curculionidae Acalles camelus (F a b r i c i u s, 1782) + + + + + + + + + Acalles fallax B o h e m a n, 1844 + + + + + + + + + Acalles echinatus (G e r m a r, 1824) + + + + + + + + Acallocrates colonnellii (B a h r, 2003) + + + + + + Barypeithes albinae F o r m a n e k, 1903 + + + + + Barypeithes chevrolati (B o h e m a n, 1843) + + + + + + Barypeithes mollicomus (A h r e n s, 1812) + + + + + + Bradybatus creutzeri G e r m a r, 1824 + Bradybatus fallax G e r s t a e c k e r, 186 + Bradybatus kellneri B a c h, 1854 + + Brachysomus echinatus (B o n s d o r f f, 1785) + + + + Brachysomus dispar P e n e c k e, 1910 + + + + + Brachysomus hirtus (B o h e m a n, 1845) + + + + + + Brachysomus setiger (G y l l e n h a l, 1840) + + + Calosirus apicalis (G y l l e n h a l, 1727) + Ceutorhynchus alliariae C h. B r i s o u t, 1860 + + + + + Ceutorhynchus chalibeus G e rm a r, 1824 + + + + Ceutorhynchus erysimi (F a b r i c i u s, 1787) + + + + + + + + Ceutorhynchus minutus (R e i c h, 1797) + + + Ceutorhynchus obstrictus (M a r s h a m, 1802) + + + + + + + + + Ceutorhynchus pallidactylus (M a r s h a m, 1802) + + + + + + + + + Ceutorhynchus rhenanus S c h u l t z e, 1895 + + Ceutorhynchus scrobicollis N e r e s h e i m e r et W.a.g.n.e.r, 1924 + + + + + + + + Ceutorhynchus typhae (H e r b s t, 1795) + + + + + + + Coeliodes proximus S c h u l t z e, 1895 + + + + Coeliodes rana (F a b r i c i u s, 1787) + + + + + + +

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T a b l e 3. (Continued) Family, species / month of occurrence Coeliodes transverseoalbofasciatus G o e z e, 1777 Coeliodes trifasciatus B a c h, 1854 Curculio glandium (M a r s h a m, 1802) Curculio pellitus (B o h e m a n, 1843) Curculio venosus (G r a v e n h o r s t, 1807) Furcipus rectirostris (L i n n a e u s, 1758) Hypera nigrirostris (F a b r i c i u s, 1775) Hypera postica (G y l l e n h a l, 1813) Kyklioacalles suturatus (D i e c k m a n n, 1983) Leiosoma cribrum (G y l l e n h a l, 1834) Microplontus campestris (G y l l e n h a l, 1837) Nedyus quadrimaculatus (L i n n a e u s, 1758) Ophrohinus suturalis (F a b r i c i u s, 1775) Otiorhynchus ovatus (L i n n a e u s, 1758) Otiorhynchus raucus (F a b r i c i u s, 1777) Phyllobius argentatus (L i n n a e u s, 1758) Phyllobius maculicornis G e r m a r, 1824 Polydrusus marginatus S t e p h e n s, 1831 Polydrusus viridicinctus G y l l e n h a l, 1834 Rhinoncus bruchoides (H e r b s t, 1784) Rhinoncus perpendicularis (R e i c h, 1797) Rhynchaenus pilosus (F a b r i c i u s, 1781) Ruteria hypocrita (B o h e m a n, 1837) Sciaphilus asperatus (B o n s d o r f f, 1785) Simo variegatus (B o h e m a n, 1843) Sitona humeralis S t e p h e n s, 1831 Sitona lineatus (L i n n a e u s, 1758) Sitona macularius (M a r s h a m, 1802) Stenocarus cardui (H e r b s t, 1784) Strophosoma melanogrammum (F o r s t e r, 1771) Trachodes hispidus (L i n n a e u s, 1758) Trachyphloeus bifoveolatus (B e c k, 1817) Tychius picirostris (F a b r i c i u s, 1787)

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With regard to habitat preference, the weevil beetles were classified into four groups: terricoles (16 spp.) living in the leaf litter; terriherbicoles (1 sp.) associated with the litter and herbage undergrowth; herbicoles preferring the herbage stratum (41 spp.); herbiarboricoles associated with both herbaceous and woody plants (1 sp.) and arboricoles associated with woody plants (19 spp.). In total, the herbicoles predominated by number of species and individuals (Fig. 3). The terricoles and arboricoles predominated quantitatively in the spring aspect. The herbicoles occurred more numerously during both the summer and autumnal aspects (e.g. the herbicolous species Ceutorhynchus palidactylus aestivated here, other species were possibly searching for their hibernation place or occurred accidentally in the forest epigeon). Considering the trophic requirements of weevil adults, four groups were distinguished: monophages (S1) associated with one plant species (7 spp.); narrow oligophages (S2) associated with one plant genus (35 spp.); wider oligophages (S3) living on more genera from one plant family or relative families (13 spp.), unknown 190

trophics (4 spp.), and polyphages or generalists (G) (19 spp.). Trophic specialists (S1–S3) predominated both by number of species and individuals (Fig. 4).

21%

6%

73%

Fig. 2. Percentages of weevil groups according to their relationship to subsoil. Explanations: GB – geobionts, GF – geophiles, GX – geoxenes.

Fig. 4. Percentages of weevil trophic groups. Explanations: S1 – monophages, S2 – narrow oligophages, S3 – wider oligophages, G – generalists, ? – unknown.

Fig. 3. Percentages of weevil topic groups. Explanations: T – terricoles, TH – terriherbicoles, HA – herbiarboricoles, H – herbicoles, A – arbo- ricoles.

Fig. 5. Percentages of weevil bionomic groups. Explanations: XF – xylophages (wood-eating), FF – phyllophages (leaf-eating).

191

Two bionomical groups of weevil adults are distinguishable in the examined material: phyllophages (leaf-eating curculionids) (70 spp.) and xylophages (wood-eating, namely saproxylic weevils) (8 spp.). Phyllophages predominated both by number of species and individuals (Fig. 5). The weevil adults differ in their habitat preference. Stenotopic species predominated qualitatively but a quantitative proportion between stenotopic a eurytopic is balanced (Fig. 6).

Fig. 6. Percentages of weevil groups according to their habitat preference. Explanations: ST – stenotopic, EU – eurytopic, UB – ubiquitous.

Fig. 7. Percentages of weevil groups according to their moisture preference. Explanations: HY – hygrophilous, XE – xerophilous, EH – euryhygric.

With regard to humidity preference, euryhygric and hygrophilous species predominate quantitatively (by number of individuals) (Fig. 7). However, amongst the stands compared there are significant differences in a qualitative-quantitative structure of the species according to their humidity preferences. The lowest quantity of the hygrophilous species appeared in the forest fragments (HH, NA) and in the stand polluted by limestone dust (LL) (35, 31, resp. 27%). Comparison of weevil assemblages Hierarchical classification Based on a qualitative–quantitative similarity (Wishart’s similarity ratio, complete linkage) the hierarchical classification divided the weevil taxocoenoses into two separate clusters connected on the low level of their similarity (Figs 8, 9).

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Fig. 8. Hierarchical classification of weevil assemblages in the forest epigeon of individual study plots according to their qualitative-quantitative similarity (Wishart’s similarity ratio, complete linkage). Vertical axis designates dissimilarity. The dendrogram based on qualitative–quantitative representation of all recorded weevil species.

The first cluster is built up of the coenoses VI and CA communicating with LI on the low level of their similarity. The study sites VI and CA are situated close to each other from the geographical point of view and are in contact with open landscape. In such light and drier forests with 100% undergrowth the communities are typical by quantitative dominance of phyllophagous geobionts, especially Barypeithes chevrolati (CA, VI) and B. albinae (LI) (Fig. 10). Acalles echinatus, Ruteria hypocrita (CA, VI, LI) Kyklioacalles suturatus (CA, VI), Trachodes hispidus (CA, LI) appear mutually as xylophagous geobionts. The second cluster with its three divisions (1. FU+LH, 2. HH+NA+NK1+NK2, 3. LL) includes the other assemblages. The first division contains the communities from the sites FU and LH, representing old closed forest stands (80–100 years of age) with significant canopy (80%) and lower density of undergrowth (40, resp. 75%) with sufficient amount of decaying wood. There is no significant difference between the compared assemblages (Table 3). The communities are typical by quantitative dominance of xylophagous geobionts, particularly Acalles fallax (A. camelus, Kyklioacalles suturatus, Ruteria hypocrita, Trachodes hispidus as well). Barypeithes chevrolati is the only phyllophagous geobiont mutual for both the compared coenoses (Table 1). The second division may be determined by the communities at the sites HH, NA, HK1 and NK2 being situated in the northern part of the study area. The compared coenoses are abundantly formed by geobiont Barypeithes mollicomus (a phyllophagous 193

species) and Acalles fallax (a xylophagous species). Ruteria hypocrita and Trachodes hispidus may be classified as the mutual xylophagous geobionts. The lowest diversity refers to the community HH with a gradation and massive occurrence of two species – Barypeithes mollicomus and Ceutorhynchus pallidactylus. There is a highly significant difference (P < 0.001) between the diversity of this assemblage (H' = 1.444) and the other coenoses of the second division (NA, NK1, NK2). The dominance of the HH assemblage has been concentrated onto low species richness, what actually appears in a higher value of the dominance index (c = 0.382). On the other hand the communities NA, NK1 and NK2 are typical by higher richness of codominant species with a consequence of lower values of the dominance index (NA: c = 0.175, NK1: c = 0.200, NK2: c = 0.153). The quantitatively poor assemblage LL represents a separate line of the second cluster on the low level of similarity towards the first two divisions. With a high diversity (H' = 1.959) the species appear in low quantities and hence the differences amongst their dominance are supposed to be minimal (c = 0.064). There is a high significant difference (P < 0.001) (Table 2) in diversity between this coenosis and the other studied assemblages.

Fig. 9. Hierarchical classification of weevil assemblages based on species associated with forest floor according to their qualitative-quantitative similarity (Wishart’s similarity ratio, complete linkage). Vertical axis designates dissimilarity.

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Fig. 10. Qualitative and quantitative proportion of bionomic groups within geobiont species. Explanations: XF – xylophagous (wood-eating), FF – phyllophagous (leaf-eating), horizontal axis (study plots, abbreviations see in Material and methods).

Relationship between weevil assemblages and environmental variables Twenty weevil species associated with the forest floor (geobionts and geophiles) were selected for the redundancy analysis. The cover of tree stratum and content of exchange bases in leaf litter from amongst 13 gradient and 2 categorial variables were significant and explanatory in the analysis (P value of the Monte Carlo permutation test was lower than 0.05). The result of redundancy analysis is shown in Fig. 11. Eigenvalues of the two first canonical axes are λ1 = 0.316 and λ2 = 0.114. The first two canonical axes 195

account for 43% of the total variance of the species data and 100% of the species–environment relation. The scatter of the sites and species forms three groups.

Fig. 11. RDA ordination diagram of weevil geobiont and geophilous species, study plots and environmental factors. Symbols: 1 – Anthribus albinus, 2 – Acalles camelus, 3 – Acalles fallax, 4 – Acalles echinatus, 5 – Acallocrates colonnellii, 6 – Barypeithes albinae, 7 – Barypeithes chevrolati, 8 – Barypeithes mollicomus, 9 – Brachysomus echinatus, 10 – Brachysomus dispar, 11 – Brachysomus hirtus, 12 – Brachysomus setiger, 13 – Kyklioacalles suturatus, 14 – Ruteria hypocrita, 15 – Trachodes hispidus, 16 – Trachyphloeus bifoveolatus, 17 – Ceutorhynchus pallidactylus, 18 – Leiosoma cribrum, 19 – Otiorhynchus ovatus, 20 – Otiorhynchus raucus, 21 – Sciaphilus asperatus. Abbreviation of study plots see in Material and methods.

The species Brachysomus echinatus, Acallocrates colonnellii, Brachysomus hirtus (the upper right quadrant of the ordination diagram) prefer the litter with a higher content of exchange bases (49.40 mval/100g) and lower acidity (NK2: pH in H2O = 6.74; the soil type is Rendzic Leptosols). The second group involves sites and species situated in the central part of RDA diagram (the upper left quadrant of the scatter). The forest sites LL, LI, VI have lower tree canopy (65–70%) and/or are influenced by open, non-forest habitats in their close vicinity. This group is formed by xerophilous species living in forest habitats, forest 196

ecotons, shrub formations but also in grassland (Otiorhynchus ovatus, Trachyphloeus bifoveolatus, Brachysomus setiger). The third group (the lower left and right quadrant of the diagram) comprises namely hygrophilous species preferring shady forest sites (CA, NA, FU, HH, LH, NK1) with higher cover of tree stratum (75–90%). Twelve forest species (Acalles camelus, A. echinatus, A. fallax, Ruteria hypocrita, Kyklioacalles suturatus, Anthribus albinus, Trachodes hispidus, Barypeithes albinae, B. chevrolati, B. mollicomus, Leiosoma cribrum, Brachysomus dispar) and one ubiquitous species (Ceutorhynchus pallidactylus) can be included in this most numerously represented group. Otiorhynchus raucus is an euryhygric species living in open habitats, forests with lower canopy as well as shady and more humid forest stands. Direct and significant influence of canopy architecture was observed also in epigeic spiders (Krumpálová, 2005), bugs (Hradil, 2005) and ants (Holecová et al., 2005).

Fig. 12. Number of weevil species recorded in the epigeon of the study plots. Explanations: median (median horizontal line), arithmetic mean (…) ± standard deviation (box), 1st and 99th percentile (x) and range (vertical lines) are displayed; for study plots abbreviations see Material and methods.

Number of species and abundance Qualitative and quantitative data were compared using single-factor analysis of variance (ANOVA). Mean number of species was significantly different amongst the study plots (F = 14.2; P < 0.001). Five partially overlapping homogenous subsets were recognised. Mean number of the species in LL was significantly lower than in FU, NA, HH, NK1, 197

NK2. Mean number of the species in CA and VI was significantly lower than in HH, NK1, NK2. On the contrary, the mean number of the species in NK2 was significantly higher than that in LL, VI, CA, LI, LH, FU (Fig. 12). Significant differences were found in mean number of geobiont species among the study plots (F = 10.2; P < 0.001). Three, partially overlapping homogenous subsets were recognised. Mean number of geobiont species in NK2 was significantly higher than that in all the other study plots and mean number of geobiont species LL was significantly lower than in FU, LH, NA, NK1, NK2 (Fig. 13).

Fig. 13. Number of geobiont weevil species in the epigeon of the study plots. Explanations: median (median horizontal line), arithmetic mean (…) ± standard deviation (box), 1st and 99th percentile (x) and range (vertical lines) are displayed; for study plots abbreviations see Material and methods.

Mean abundance of weevils was significantly different among the study plots (F = 6.5; P < 0.001). Mean abundance in the study plot HH was significantly higher than that in all other localities, while no significant differences were found among the remaining study plots (Fig. 14). Similarly, significant differences were found in the mean abundance of geobiont species among the localities (F = 4.3; P < 0.001). Three, partially overlapping homogenous subsets were recognized: the first consists of LL, VI, CA, LI, FU, NA, LH and NK1 (P = 0.402); the second of CA, LI, FU, NA, LH, NK1 and NK2 (P = 0.094); the third of NK1, NK2 and HH (P = 0.549). Mean abundance of geobionts at the sites LL and VI was significantly lower than in HH and NK2 (Fig. 15).

198

Fig. 14. Abundance of weevils recorded in the epigeon of the study plots (ind.m-2). Explanations: median (median horizontal line), arithmetic mean (…) ± standard deviation (box), 1st and 99th percentile (x) and range (vertical lines) are displayed; for study plots abbreviations see Material and methods.

Fig. 15. Abundance of geobiont weevil species recorded in the epigeon of the study plots (ind.m-2). Explanations: median (median horizontal line), arithmetic mean (…) ± standard deviation (box), 1st and 99th percentile (x) and range (vertical lines) are displayed; for study plots abbreviations see Material and methods.

199

Human impact All the study stands represent managed and, in some way, human-modified forests. The most serious anthropogenic impact appears in a stand fragmentation (a lack of originally continuous forest habitat and its contact with nearby open landscape) as well as in particulate pollution with calcareous dust from the nearby quarry at the site LL. Although the forest stand fragmentation did not declare any significant and explainable impact within a complex analysis of the weevil communities (Fig. 11) the separate evaluation of all the sites hints at increased proportion of herbicoles (HH – 62% of the total specimens), NA – 60%), euryhygric species (HH – 62%, NA – 61%) and ubiquists (HH – 58%, NA – 51%) in the fragmented forest complex. The calcareous dust pollution from the nearby quarry was evident at the site LL, what was actually considerable during the low precipitation years (2000, 2001). Humus content in litter (LL: 6.55%) was measured as low in the comparison with the other stands (CA – 11.21%, VI – 13.79%, FU – 24.83%, LI – 14.48%, HH – 16.55%, LH – 12.76%, NA – 12.41%, NK1 –12.76% and NK2 – 9.31%). The weevil coenosis in LL behaves as unstable and from the qualitative–quantitative point of view it seems poor in geobionts (Figs. 13, 15). Of the 25 species recorded during a four–year period there is only one xerophilous geoxene (Sitona macularius) being present in epigeon during three seasons (1999, 2000 and 2001, respectively). Only 3 species were recorded within two seasons (two geobionts living also in non–forest habitats: Brachysomus echinatus, Trachyphloeus bifoveolatus and one geoxene: Strophosoma melanogrammum). The typical forest geobionts preferring dead wood (Acalles fallax, A. camelus and Trachodes hispidus) were observed just in 1999, when the maximum precipitation referred to the spring and summer months. Therefore the calcareous dust was repeatedly flowed out from the forest floor and its vegetation. Occurrence of the remaining 21 weevil species was confirmed only in one season. Unlike the other forest stands, the weevil species structure at this anthropically affected site was different in each year (apart from the 4 above mentioned species) (Table 4). Fourteen species were recorded in 1999 – Acalles camelus, A. fallax, Brachysomus echinatus, Ceutorhynchus pallidactylus, C. scrobicollis, Coeliodes trifasciatus, Curculio glandium, Furcipus rectirostris, Phyllobius argentatus, Polydrusus marginatus, P. viridicinctus, Sitona macularius, Trachodes hispidus, Trachyphloeus bifoveolatus. In the vegetation period 2000 there were only four species observed – Barypeithes chevrolati, Brachysomus echinatus, Sitona macularius, Strophosoma melanogrammum. In 2001 we recorded five species – Ceutorhynchus alliariae, C. chalibeus, Sitona macularius, Strophosoma melanogrammum, Trachyphloeus bifoveolatus and in 2002 nine species – Catapion seniculus, Ischnopterapion virens, Protapion apricans, P. trifolii, Ceutorhynchus chalibeus, C. typhae, Hypera nigrirostris, Otiorhynchus ovatus, Polydrusus marginatus. Mean abundance of all the weevils (MA) as well as mean abundance of the geobiont weevil species (MAGB) reached low values in all the seasons (arithmetic mean ± SD, MA: 1999 = 5,57 ± 2,149 ind.m-2, 2000 = 1.00 ± 2,138 ind.m-2, 2001 = 0.88 ± 0,835 ind.m-2, 2002 = 1.63 ± 2.326 ind.m-2; MAGB: 1999 = 2,71 ± 1,704 ind.m-2, 2000 = 0.25 ± 0,463 ind.m-2, 2001 = 0.25 ± 0.707 ind.m-2, 2002 – geobiont species were absent). 200

T a b l e 4. Number of weevil species recorded in individual study plots from one to four years of the research. Study plot CA VI FU LI HH LH LL NA NK1 NK2 1 year 14 8 9 8 16 15 21 11 9 8 2 years 5 7 6 6 6 2 3 5 7 5 3 years 2 4 4 3 3 3 1 2 3 7 4 years 4 4 3 5 6 5 0 8 7 6 Abbreviation of the study plots see in Material and methods.

Epigeic weevil assemblages are likely to be sensitive to mechanical disturbance of habitats, particulate, chemical pollution and also to forest cutting. Majority of typical weevil geobionts are apterous or brachypterous. They have low mobility and are not able to survive unfavourable conditions (Holecová, 1986). On the contrary, the woodlands affected by imissions are to a higher extent attacked by leaf-feeding insects (mainly moth caterpillars, beetles, aphids, etc.). This fact was observed at the Lošonec–quarry by Bulánková and Holecová (1998, 2000) as well as by other authors in various parts of Slovakia (Bulánková, 1990; Cicák et al., 1999; Kulfan et al., 2002, 2004, etc.). Only a few species (in general mostly known as “pests”) can thrive in areas with pollution impacts. Such the gradations often lead into an abundance increase of their natural predators and parasitoids. Bulánková and Holecová (1998, 2000) hinted at considerably higher abundance of Nabidae predators in the forest habitat polluted by calcareous dust (LL) in the comparison with the background without pollution impact. The effects of air pollutants on insects have been reviewed by several authors (Alstaed et al., 1982, Dopcherty et al., 1997, ect.). It is evident that any polluted area has its own special features (pollutants and other anthropogenic impacts, abiotic conditions, flora, fauna, ect.). Also, differences in response of particular insect species to pollution are apparent. The factors act in many combinations and interferences (Cicák et al., 1999; Führer, 1985; Kulfan, 1988; Kulfan et al., 2002, 2004; Zelinková et al., 2004). Translated by M. Holecová and P. J. Fedor Acknowledgement We would like to thank I. Országh and Z. Krumpálová for their assistance in the field and laboratory work. This research was supported by VEGA (Scientific Grant Agency of the Ministry of Education and the Slovak Academy of Sciences), grant number 1/0119/03.

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Holecová M., Némethová D., Kúdela M.: Štruktúra a funkcia spoločenstiev nosáčikov (Coleoptera, Curculionoidea) v epigeóne dubovo-hrabových lesov JZ Slovenska. V rokoch 1999–2002 sme študovali štruktúru a dynamiku taxocenóz nosáčikov (Coleoptera, Curculionoidea) v epigeóne dubovo-hrabových lesov. Výskum sme uskutočnili na 10 študijných plochách, v lesných porastoch veku 40–100 rokov patriacich do zväzov Carpinion betuli, Quercion confertae-cerris, Q. pubescentis-petrae. Študované lesné porasty sa nachádzajú v orografických celkoch Malé Karpaty a Trnavská pahorkatina. Pôdnu makrofaunu sme zbierali metódou presevov v mesačných intervaloch. Materiál sme extrahovali v xereklektoroch Moczarskeho typu. Celkove sme zozbierali 4 090 jedincov, ktoré patrili do 78 druhov, 43 rodov a 4 čeľadí. Na jednotlivých študijných plochách sme zistili od 22 do 31 druhov. Priemerná abundancia nosáčikov dosahovala hodnoty od 2.1 do 49.7 ex.m-2 Charakteristické druhové spektrum bolo zastúpené 3 skupinami druhov: (1) druhmi s vysokou hodnotou dominancie a konštantnosti – Acalles fallax, Barypeithes mollicomus, Trachodes hispidus, Ceutorhynchus pallidactylus, C. obstrictus, Sitona macularius; (2) druhmi s vysokou konštantnosťou, ktoré nedominujú – Acalles camelus, Ruteria hypocrita, Ceutorhynchus alliariae, C. scrobicollis, Coeliodes trifasciatus, Curculio glandium, Furcipus rectirostris, Phyllobius argentatus; (3) diferenčnými druhmi – Acalles echinatus, Acallocrates colonnellii, Barypeithes albinae, Brachysomus dispar, B. setiger, Coeliodes proximus, C. rana, C. transverseoalbofasciatus. RDA ordinácia ukázala, že z.vybraných 13 gradientových a 2 kategoriálnych premenných má na štruktúru spoločenstiev Curculionoidea priamy a signifikantný vplyv zápoj stromovej etáže a obsah výmenných báz (Ca2+, Mg2+, K+, Na+) v pôde. Fragmentácia porastov mala za následok vzrast početnosti herbikolných, euryhygrických, ubikvistických druhov. Nestabilné a kvantitatívne chudobné bolo spoločenstvo porastu zaprášeného vápenatým prachom z.blízkeho lomu.

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Ekológia (Bratislava)

Vol. 24, Supplement 2/2005, p. 205–222

BIODIVERSITY OF SELECTED INVERTEBRATE GROUPS IN OAK-HORNBEAM FOREST ECOSYSTEM IN SW SLOVAKIA MILADA HOLECOVÁ1, MIROSLAV KRUMPÁL1, IVAN ORSZÁGH1, ZUZANA KRUMPÁLOVÁ2, SLAVOMÍR STAŠIOV3, PETER FEDOR4 Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská dolina B-1, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected], [email protected], [email protected] 2 Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, 845 06 Bratislava, The Slovak Republic, e-mail: [email protected] 3 Department of Biology and General Ecology, Faculty of Ecology and Environmental Sciences, Technical University, T.G. Masaryka 24, 960 53 Zvolen, The Slovak Republic, e-mail: [email protected] 4 Department of Ecosozology, Faculty of Natural Sciences, Comenius University, 842 15 Bratislava, The Slovak Republic, e-mail: [email protected] 1

Abstract Holecová M., Krumpál M., Országh I., Krumpálová Z., Fedor P.: Biodiversity of selected invertebrate groups in oak-hornbeam forest ecosystem in SW Slovakia. Ekológia (Bratislava), Vol. 24, Supplement 2/2005, p. 205–222. The paper summarizes analyses of 4–year long coenological research on micro- and macrofauna in oak-hornbeam forest ecosystems in SW Slovakia. The studied forest ecosystems, 40–100 years of age, are situated in the orographic units of the Malé Karpaty Mts. and Trnavská pahorkatina hills and may be classified into 3 vegetation types: Carpinion betuli, Quercion confertae-cerris and Quercion pubescentis-petraeae. In total we determined 39,987 invertebrates (except for Protozoa) and thus recorded 575 species of 4 phyla (Ciliophora, Rhizopoda, Tardigrada, Arthropoda). Twelve taxocoenoses of ciliates, naked amoebae, water bears, pseudoscorpions, spiders, mesostigmatid mites, terrestrial isopods, centipedes, millipedes, earwigs, bugs, weevils were analysed more in detail. Apart from forest epigeon we were focused on some other microhabitats, such as decaying wood mater, mosses and dendrotelmae. Of the studied microfauna just the community from decaying wood possessed affinity to soil. In the other microhabitats (dendrotelmae and mosses) species are predominantly interacted with aquatic environment. There are stronger bonds onto soil at arthropods, represented particularly by epigeic, partially by typically edaphic species. Of the 15 analysed variables just age of a stand, depth of leaf litter, undergrowth coverness of canopy and sporadically pollution (dust from the quarry) appeared as significantly influencing the studied arthropod communities. Key.words:.invertebrates, coenoses, oak-hornbeam forest, epigeon, mosses, decaying wood, tree-holes, SW Slovakia tree-holes, SW Slovakia

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Introduction Oak-hornbeam forests used to be the most frequent forest climatically zone formation at lower altitudes in Slovakia. In past they continually covered large areas particularly in lowlands from the altitude of 100 m a.s.l. In hills they spread up to 600 m a.s.l. and occurred in all Carpathian basins. For a long time these forest stands have been under intensive anthropogenous impact in Slovakia as well as in other European regions. Nowadays they have survived in fragments attacked and impacted by humans. However in cultural land they provide refugium for many animal species. Animals in such the forest type have not yet been intensively studied. Relevant papers dealing with fauna of oak-hornbeam forests are usually focused on pests (e.g. Patočka et al., 1999), or are limited by partial taxocoenoses of certain invertebrates (for example naked amoebae: Mrva, Matis, 2000; Mrva, 2003; ciliates: Tirjaková, 2002; Tirjaková et al., 2002; water bears: Nelson et al., 1979; Guoth, 1986; Dastych, 1988; Degma et al., 2004, 2005a; terrestrial isopods: Gulička, 1960; Krumpál, 1973, 1976; Flasarová, 1980, 1986; Flasar, Flasarová, 1989; millipedes: Gulička, 1986; Branquart et al., 1995; Korsós, 1997; David et al., 1999; centipedes: Wytwer, 1990; Tajovský, 2001; spiders: Žitňanská, 1981; Gajdoš, 1992; Gajdoš, Krumpál, 1986, 1988; Jedličková, 1988; Noflatscher, 1991; Esjunin et al., 1994; Krumpálová, Bartoš, 2002; Krumpálová, Szabová, 2003, 2005; mites: Kalúz, 1981, 2005; Ambros, Kalúz, 1985, 1987; Mašán et al., 1994; Kalúz, Fenďa, 2005; bugs: Štepanovičová, Kovačovský, 1971; Bianchi, 1991; Rédei, Hufnagel, 2003a, b; Bakonyi et al., 2002; beetles: Korbel, 1966, 1973; Drdul, 1973; Czechowski, 1989; Majzlan, 1986, 1991; Majzlan, Hošták, 1996; Majzlan et al., 2000; Holecová, Sukupová, 2002; Holecová et al., 2002 etc.). General and complex view on arthropod fauna in Central-European oak-hornbeam forest has been included in just a few of papers (e.g. Balogh, Loksa, 1948; Verner, 1959; Loksa, 1966, 1968; partially Nosek, 1986). This study has been focused to analyse the structure and biodiversity of soil microand macrofauna in oak-hornbeam forest ecosystem more complexly. The research hinted at stands of different age and anthropogenous impact. Impacts of forest fragmentation as well as some ecological and environmental factors have been taken into account. The research was realised during 4 vegetation seasons (1999–2002) at 10 regularly studied sites in central and northern part of the Malé Karpaty Mts and nearby Trnavská pahorkatina hills. Apart from intensive analyses of soil fauna we were focused on some other microhabitats, such as mosses, decayed wood mater (in various decay degree) and dendrotelmae, which have not been intensively studied yet, particularly from the microfauna point of view.

Material and methods To study epigeic and soil macrofauna we applied almost all the available collecting methods, such as sieving the leaf litter and upper part of soil, direct sampling of soil (leaf litter, soil) as well as formaldehyde ground traps. From samples of mosses, wood and tree-holes the microfauna was studied by direct analyses of sampled

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material and modifications of non-flooded Petri-dish method. Material was collected in regular monthly intervals at the same study plots and periods.

Results and discussion General characterization of oak-hornbeam invertebrate assemblages seems to be very disputable due to the rich material of various systematic groups, often from many types of microhabitats. However this paper will project a certain synthesis. In total we determined 39,987 invertebrates (except for Protozoa) of 575 species from 4 phyla (Ciliophora, Rhizopoda, Tardigrada, Arthropoda). Generally the study presents detailed ecological analysis of 12 systematic groups (ciliates, naked amoebae, water bears, pseudoscorpions, spiders, mesostigmatid mites, terrestrial isopods, centipedes, millipedes, earwigs, bugs, weevils) being provided by the authors. In this part we summarize the obtained data, analyse epigeic and soil animal communities from oak-hornbeam forests in the Malé Karpaty Mts. Moreover environmental impact on the studied animals was taken into account. Protozoa were studied in decaying wood matter dendrotelmae and mosses – in markedly different microhabitats in some degree interacted with soil. Decaying wood matter often represents refugium for various soil animals and is inhabited by specific communities. The structure of ciliate assemblages with a majority of soil species has proved this fact. Interactions towards a certain tree or shrub species do not appear as significant (Bartošová, Tirjaková, 2005). In contradiction to decaying wood matter tree-holes represent specific habitats being isolated from soil, what has been actually proved by the communities of Ciliophora. Four indication communities have been established. They are particularly affected by tree and shrub diversity, size and age of a telma as well as by presence of Rotifera and other Metazoa. Moreover time as an important factor has to be taken into account. The communities were represented by aquatic, terrestric, limnetic and eurypotent species in diverse proportion and were influenced by many factors mentioned above. Low frequency in occurrence of species may hint at diversity and significant impact of environment. Interactions with soil have not been proved (Tirjaková, Vďačný, 2005). However mosses possess stronger interactions to soil than dendrotelmae do. Therefore the fact that 32 recorded taxa (23 species) of naked amoebae with significant bonds to aquatic environment appears as very disputable. The community of naked amoebae seems to be very similar to aquatic assemblages (Mrva, 2005). Twenty-one species of Tardigrada obtained from mosses have not shown any significant interactions to soil. Some expressive differences in species diversity refer to individual sites with high mutual similarity at the least-diverse communities. Mostly the differences correspond with accidental records of various species. Distribution and dispersal process at Tardigrada in various strata is supposed to be passive and accidental with insignificant interactions with specificity of environment (Degma et al., 2005b). In the study area we recorded 7 epigeic terrestrial isopod species. Specific communities were usually formed by 1–6 of them with relatively low average 207

abundance. Protracheoniscus politus and Porcellium collicola may be classified as indicative species. The communities were influenced by age of a stand, soil type, content of Ca and pH of soil (Tuf, Tufová, 2005). Of ten recorded pseudoscorpion species just Neobisium muscorum was recorded at all the study sites. Four communities being classified for the study area were not significantly impacted by 15 measured environmental variables. The highest stability refers to the assemblage in 80–100 year old oak-hornbeam forest stands, the lowest values hints at more arid and younger forest (60–80 years) with markedly reduced leaf horizon (Christophoryová, Krumpál, 2005). One hundred and fifty-eight species and 24 families of spiders were recorded in the study area. Their communities in oak-hornbeam forests were considerably diverse with the indication species of Trochosa terricola, Pardosa lugubris and Tenuiphantes mengei. Of the measured environmental variables the undergrowth appears as the only factor significantly influencing the communities. The other variables seem to be with no important impact on taxocoenoses of spiders (Krumpálová, 2005). The studied microhabitats were inhabited by 75 mesostigmatid mites. Holoparasitus calcaratus, Veigaia nemorensis, Vulgarogamasus kraepelini and Zercon peltatus var. peltatus were the eudominant species in forest soil. Temperature, air humidity, soil type, pH and vegetation may be classified as the most significant factors influencing structure and frequency in the communities of mites (Fenďa, Ciceková, 2005). The centipede (Chilopoda) communities were formed by 10–17 species of the total richness of 24 species recorded in the study area. Schendyla nemorensis and Lithobius muticus belonged to the category of eudominant species at all the sites. The typical centipede community in the oak-hornbeam forests of the Malé Karpaty Mts consists of the following species: Schendyla nemorensis, Strigamia acuminata, Lithobius agilis, L. borealis, L. lapidicola, L. mutabilis, L. muticus, L. austriacus; in more southern drier parts of the Malé Karpaty Mts Henia illyrica as well. The highest diversity (17 species) refers to the 80–100 year old oak-hornbeam forest, the lowest one to the more arid and younger (60–80 years old) stand with minimal leaf litter. Most species occurred during the whole year, what was actually proved by additional samples from January 2000–2002 being excluded from the paper (Országh, Országhová, 2005). Eighteen millipede species were recorded in the studied oak-hornbeam forests. Individual communities were formed by 8–14 species. Cylindroiulus boleti, Strongylosoma stigmatosum and Ommatoiulus sabulosus may be classified as indication species. The results of cluster analysis of the similarity of localities showed possible influence of the height of forest growth and also values of pH and sorption complex in leaf litter on the structure of the millipede communities. The tree height could have an indirect effect in the terms of higher litter production (bigger amount of suitable food source for saprophages) by taller trees (Stašiov, 2005). Fourty-six species of Heteroptera were recorded in the oak-hornbeam forests. The communities at the study sites were formed by 21–28 species, including eudominant Legnotus limbosus and Eurygaster maura. The following species may be considered as the typical elements: Drymus brunneus, D. ryeii, Scolopostethus affinis, S. thomsoni, Rhyparochromus alboacuminatus, Legnotus limbosus. The geoxenous species such as Aelia acuminata or Eurygaster maura are common as well. The research has proved the 208

impact of canopy coverness (E3) on structure of bug communities. Tropistethus holosericeus, Legnotus limbosus, Plinthisus brevipennis, Raglius alboacuminatus, Trapezonotus arenarius or Microporus nigritus may be classified as more heliophilous species with preference to scarce cover. Moreover the nearby quarry is supposed to negatively influencing (air pollution) the community of epigeic Heteroptera in oak-hornbeam forests of the Malé Karpaty Mts (Hradil, 2005). In epigeon of oak-hornbeam forests we recorded 78 species of weevils. Their communities were usually formed by 22–31 species. Acalles fallax, Barypeithes mollicomus, Trachodes hispidus, Ceutorhynchus pallidactylus, C. obstrictus and Sitona macularius may be classified as species with high value of dominance and frequency. Of 13 gradient and 2 categorial variables being analysed in the research there are just several: coverness of canopy and content of exchangeable bases (Ca2+, Mg2+, K+, Na+) in soil with a significant impact on communities of weevils. Fragmentation of stands leaded to increase in abundance of herbicolous, euryhygric and ubiquitous species. The community at the site being influenced by calcareous dust from the nearby quarry may appear as unstable and quantitatively poor (Holecová et al., 2005). Apart from the groups studied and mentioned above we include notes on thrip communities (Thysanoptera), which undisputedly form an important part of forest ecosystem and may indicate character and dynamics of ecological conditions. However the thrips being sampled from epigeon predominantly have arboricolous origin and many of them possess strong interactions with soil in some ontogenetical stage. The taxocoenoses consist of approximatelly 15 species and are significantly equitable in older, more stable forests. For instance in Cajla (site 1) Hoplandrothrips williamsianus P.r i e s n e r, 1923 may be considered as an indication species of vital oak-hornbeam forests, followed by zoophagous Aeolothrips versicolor U z e l, 1895. In more opened forest stands with a real possibility for infiltration of thrips from the nearby ecosystems, such as Horný háj grove, the communities are enriched in several more heliophilous species such as Limothrips denticornis H a l i d a y, 1836 and Frankliniella tenuicornis (U z e l, 1895). Of the studied Protozoa and Tardigrada there was only one group in decaying wood matter with affinity towards soil. The other communities possessed interactions to aquatic environment. The observed arthropods were more significantly interacted with soil. The communities were predominantly represented by epigeic species, partially by typical edaphic species. Of the 15 analysed variables just age of a stand, depth of leaf litter, coverness of herbage undergrowth, canopy architecture and sporadically pollution (dust from the quarry) appeared as significantly influencing the studied arthropod communities. Translated by P. J. Fedor Acknowledgement The work has been supported by the Slovak Grant Agency VEGA, Grant No. 1/0116/03.

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Holecová M., Krumpál M., Országh I., Krumpálová Z., Stašiov S., Fedor P.: Biodiverzita vybraných skupín bezstavovcov v ekosystéme dubovo-hrabového lesa v oblasti JZ Slovenska. Práca sumarizuje výsledky štvorročného cenologického výskumu mikro- a makrofauny lesného ekosystému dubovo-hrabového vegetačného stupňa v oblasti JZ Slovenska. Študované lesné porasty veku 40–100 rokov sa nachádzajú v orografických celkoch Malé Karpaty a Trnavská pahorkatina. Jedná sa o 3 typy dubovo-hrabových (Carpinion betuli), dubovo-cerových (Quercion confertae-cerris) a xerotermofilných dubových lesov (Quercion pubescentis-petraeae). Za celé obdobie výskumu bolo spolu determinovaných 39 987 exemplárov bezstavovcov (s výnimkou Protozoa). Zistili sme 575 živočíšnych druhov, patriacich do 4 kmeňov (Ciliophora, Rhizopoda, Tardigrada, Arthropoda). Celkove boli spracované synúzie 12 systematických skupín (nálevníky, nahé meňavky, pomalky, štúriky, pavúky, mesostigmátne roztoče, suchozemské rovnakonôžky, stonôžky, mnohonôžky, ucholaky, bzdochy, nosáčiky). Okrem lesného epigeónu, sme venovali pozornosť štúdiu ďalších mikrohabitatov, akými sú rozkladajúca sa drevná hmota, machy a dendrotelmy. Z mikrofauny javila afinitu k lesnej pôde iba skupina študovaná v odumretej drevnej hmote. Pre ostatné mikrohabitaty (dendrotelmy a machy) je charakteristický výskyt druhov viazaných tiež na vodné prostredie. U študovaných skupín Arthropoda je už väzba na pôdu oveľa výraznejšia. Prevládali epigeické druhy a len menej boli zastúpené typicky edafické druhy. Z 15 sledovaných environmentálnych premenných javili vplyv na niektoré spoločenstvá článkonožcov iba vek porastu, hrúbka vrstvy hrabanky, pokryvnosť bylinnej etáže a korunový zápoj, u niektorých skupín tiež znečistenie vápenatým prachom z lomu.

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A P P E N D I X 1. Review of taxa found in studied oak-hornbeam forest of SW Slovakia in 1999–2002

Phylum: RHIZOPODA Class: LOBOSEA Subclass: GYMNAMOEBIA Order: EUAMOEBIDA Family: Amoebidae Deuteramoeba algonquinensis (B a l d o c k, R.o.g.e.r.s o n et B e r g e r, 1983) Family: Hartmannellidae Hartmannella cantabrigiensis P a g e, 1974 Hartmannella vermiformis P a g e, 1967 Saccamoeba limax (D u j a r d i n, 1841) Saccamoeba stagnicola P a g e, 1974 Family: Thecamoebidae Dermamoeba granifera (G r e e f f, 1866) P a g e et B l a k e y, 1979 Dermamoeba minor (P u s s a r d, A.l.a.b.o.u.v.e.t.t.e et P o n s, 1979) Paradermamoeba levis S m i r n o v et G.o.o.d.k.o.v, 1994 Paradermamoeba valamo S m i r n o v et G.o.o.d.k.o.v, 1993 Sappinia diploidea (H a r t m a n n et N ä g l e r, 1908) Thecamoeba quadrilineata (C a r t e r, 1856) Thecamoeba sphaeronucleolus (G r e e f f, 1891) Thecamoeba striata (P e n a r d, 1890) Thecamoeba terricola (G r e e f f, 1866) Family: Vannellidae Platyamoeba stenopodia P a g e, 1969 Vannella sp. Vannella lata P a g e, 1988 Vannella platypodia (G l ä s e r, 1912) Family: Paramoebidae Korotnevella bulla (S c h a e f f e r, 1926) Korotnevella diskophora S m i r n o v, 1999 Korotnevella stella (S c h a e f f e r, 1926) Mayorella penardi P a g e, 1972 Mayorella vespertilioides P a g e, 1983 Order: LEPTOMYXIDA Family: Flabellulidae Flamella sp. 1 Flamella sp. 2 Family: Leptomyxidae Leptomyxa reticulata G o o d e y, 1914 Rhizamoeba sp. Unidentified leptomyxid amoeba

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Class: ACANTHOPODIDA Family: Acanthamoebidae Acanthamoeba sp.1 Acanthamoeba sp.2 Acanthamoeba sp.3 INCERTAE SEDIS Stygamoeba sp. Phylum: CILIOPHORA Class: HETEROTRICHEA Order: HETEROTRICHIDA Family: Blepharismidae Blepharisma hyalinum P e r t y, 1849 Blepharisma sp. Order: ARMOPHORIDA Family: Metopidae Metopus hasei S o n d h e i m, 1929 Metopus minor K a h l, 1927 Class: SPIROTRICHEA Subclass: OLIGOTRICHIA Order: HALTERIIDA Family: Halteriidae Halteria grandinella (O. F. M u e l l e r, 1773) Subclass: HYPOTRICHIA Order: EUPLOTIDA Family: Euplotidae Euplotes affinis (D u j a r d i n, 1841) Euplotes muscicola K a h l, 1932 Order: UROSTYLIDA Family: Urostylidae Bakuella pampinaria E i g n e r et F o i s s n e r, 1992 Hemicycliostyla sphagni S t o k e s, 1886 Hemisincirra gellerti (F o i s s n e r, 1982) Hemisincirra interrupta (F o i s s n e r, 1982) Holosticha muscorum (K a h l, 1932) Holosticha sp. Holosticha tetracirrata B u i t k a m p et W.i.l.b.e.r.t, 1974 Paraurostyla macrostoma F o i s s n e r, 1982 Order: SPORADOTRICHIDA Family: Oxytrichidae Cyrtohymena candens K a h l, 1932 Cyrtohymena muscorum (K a h l, 1932) Cyrtohymena quadrinucleata (D r a g e s c o et N.j.i.n.é, 1971) Gastrostyla steini E n g e l m a n n, 1862 Gonostomum affine (S t e i n, 1859)

Appendix 1. (Continued) Gonostomum kuehnelti F o i s s n e r, 1987 Histriculus vorax (S t o k e s, 1891) Oxytricha granulifera quadricirrata B l a t t e r e r et F.o.i.s.s.n.e.r, 1988 Oxytricha setigera S t o k e s, 1891 Oxytricha similis E n g e l m a n n, 1862 Oxytricha sp. Steinia platystoma (E h r e n b e r g, 1831) Sterkiella histriomuscorum (F o i s s n e r, B.l.a.t.t.e.r.e.r, B e r g e r et K o h m a n n, 1991) Stylonychia pustulata (O. F. M u e l l e r, 1786) Tachysoma pellionellum (O. F. M u e l l e r, 1773) Urosomoida agiliformis F o i s s n e r, 1982 Class: PROSTOMATEA Order: PRORODONTIDA Family: Plagiocampidae Plagiocampa metabolica (K a h l, 1926) Family: Urotrichidae Urotricha globosa S c h e w i a k o f f, 1892 Family: Colepidae Coleps hirtus (O. F. M u e l l e r, 1786) Class: LITOSTOMATEA Subclass: HAPTORIA Order: HAPTORIDA Family: Enchelyidae Enchelys gasterosteus K a h l 1926 Enchelys polynucleata (F o i s s n e r, 1984) Family: Acropisthiidae Acropisthium mutabile P e r t y, 1852 Family: Tracheliidae Dileptus breviproboscis F o i s s n e r, 1981 Order: SPATHIDIIDA Family: Spathidiidae Apospathidium atypicum (B u i t k a m p et W.i.l.b.e.r.t, 1974) Arcuospathidium australe F o i s s n e r, 1988 Arcuospathidium japonicum F o i s s n e r, 1988 Epispathidium amphoriforme (G r e e f f, 1888) Epispathidium sp. Protospathidium vermiforme F o i s s n e r, A.g.a.t.h.a et B e r g e r, 2002 Spathidium bavariense K a h l, 1930 Spathidium muscicola K a h l, 1930 Spathidium spathula (M ü l l e r, 1773) Spathidium turgitorum F o i s s n e r, A g a t h a et B e r g e r, 2002 Order: PSEDOHOLOPHRYIDA Family: Pseudoholophryidae Pseudoholophrya terricola B e r g e r, F o i s s n e r et A d a m, 1984

Order: PLEUROSTOMATIDA Family: Litonotidae Acineria uncinata T u c o l e s c o, 1962 Litonotus muscorum (K a h l, 1931) Class: PHYLLOPHARYNGEA Subclass: PHYLLOPHARYNGIA Order: CHLAMYDODONTIDA Family: Chilodonellidae Chilodonella uncinata (E h r e n b e r g, 1838) Odontochlamys gouraudi C e r t e s, 1891 Pseudochilodonopsis mutabilis F o i s s n e r, 1981 Pseudochilodonopsis sp. Subclass: SUCTORIA Order: EXOGENIDA Family: Podophryidae Sphaerophrya terricola F o i s s n e r, 1986 Class: NASSOPHOREA Order: SYNHYMENIIDA Family: Scaphiodontidae Chilodontopsis muscorum K a h l, 1931 Order: MICROTHORACIDA Family: Microthoracidae Drepanomonas dentate F r e s e n i u s, 1858 Drepanomonas exigua P e n a r d, 1922 Drepanomonas obtusa P e n a r d, 1922 Drepanomonas pauciciliata F o i s s n e r, 1987 Drepanomonas revoluta P e n a r d, 1922 Drepanomonas sphagni K a h l, 1931 Leptopharynx costatus M e r m o d, 1914 Leptopharynx eurystomus (K a h l, 1931) Family: Pseudomicrothoracidae Pseudomicrothorax agilis M e r m o d, 1914 Class: OLIGOHYMENOPHOREA Subclass: PENICULIA Order: PENICULIDA Family: Frontoniidae Frontonia angusta K a h l, 1931 Frontonia depressa (S t o k e s, 1886) Frontonia leucas (E h r e n b e r g, 1833) Family: Parameciidae Paramecium caudatum E h r e n b e r g, 1833 Subclass: SCUTICOLITIA Order: PHILASTERIDA Family: Philasteridae Philasterides armatus (K a h l, 1926) Family: Cinetochilidae Cinetochilum margaritaceum (E h r e n b e r g, 1831) Sathrophilus mobilis (K a h l, 1926) Sathrophilus muscorum (K a h l, 1931) Family: Cohnilembidae Kahlilembus attenuatus (S m i t h, 1897)

215

Appendix 1. (Continued) Family: Pseudocohnilembidae Pseudocohnilembus pusillus (Q u e n n e r s t e d t, 1869) Family: Uronematidae Homalogastra setosa K a h l, 1926 Order: PLEURONEMATIDA Family: Cyclidiidae Cyclidium elongatum (S c h e w i a k o f f, 1889) Cyclidium glaucoma O. F. M u e l l e r, 1773 Cyclidium muscicola K a h l, 1931 Subclass: HYMENOSTOMATIA Order: HYMENOSTOMATIDA Family Glaucomidae Glaucoma scintillans E h r e n b e r g, 1830 Pseudoglaucoma muscorum K a h l, 1931 Family Ophryoglenidae Ophryoglena flava E h r e n b e r g, 1833 Ophryoglena oblonga G a j e v s k a j a, 1927 Ophryoglena sp. Family Tetrahymenidae Tetrahymena edaphoni F o i s s n e r, 1986 Tetrahymena pyriformis complex Tetrahymena rostrata (K a h l, 1926) Family Turaniellidae Colpidium colpoda (L o s a n a, 1829) Dexiostoma campylum (S t o k e s, 1886) INCERTAE SEDIS Dexiotrichides centralis (S t o k e s, 1885) Hexotricha caudata L a c k e y, 1925 Pithothorax processus K a h l, 1926 Subclass: PERITRICHEA Order: SESSILIDA Family: Opisthonectidae Opisthonecta henneguyi F a u r e-F r e m i e t, 1906 Telotrochidium cylindricum F o i s s n e r, 1978 Telotrochidium sp. Family: Operculariidae Opercularia arboricolum (B i e g e l, 1954) Propyxidium spp. Family: Scyphidiidae Scyphidia spp. Family: Epistylidae Epistylis entzii S t i l l e r, 1935 Epistylis sp. Rhabdostyla pyriformis P e r t y, 1852 Family: Vorticellidae Vorticella aquadulcis complex Vorticella astyliformis F o i s s n e r, 1981 Vorticella infusionum D u j a r d i n, 1841 Vorticella microstoma E h r e n b e r g, 1830 Vorticella spp. Peritrichia gen. sp.

216

Class: COLPODEA Order: COLPODIDA Family: Colpodidae Bresslaua vorax K a h l, 1931 Colpoda aspera K a h l, 1926 Colpoda cavicola K a h l, 1935 Colpoda cucullus (O. F. M u e l l e r, 1773) Colpoda edaphoni F o i s s n e r, 1980 Colpoda henneguyi F a b r e-D o m e r g u e, 1889 Colpoda inflata (S t o k e s, 1884) Colpoda lucida G r e e f f, 1888 Colpoda maupasi E n r i q u e z, 1908 Colpoda reniformis K a h l, 1931 Colpoda steinii M a u p a s, 1883 Family: Hausmanniellidae Anictostoma terricola F o i s s n e r, 1993 Hausmanniella patella (K a h l, 1931) Family: Marynidae Ilsiella elegans F o i s s n e r, A g a t h a et B.e.r.g.e.r, 2002 Order: CYRTOLOPHOSIDIDA Family: Cyrtolophidae Cyrtolophosis acuta K a h l, 1926 Cyrtolophosis elongata (S c h e w i a k o f f, 1892) Cyrtolophosis mucicola S t o k e s, 1885 Plesiocaryon elongatum (S c h e w i a k o f f, 1892) Family: Platyophryidae Platyophrya macrostoma F o i s s n e r, 1980 Platyophrya spumacola K a h l, 1927 Platyophrya vorax K a h l, 1926 Phylum: TARDIGRADA Class: EUTARDIGRADA Family: Hypsibiidae Hypsibius convergens (U r b a n o w i c z, 1925) Hypsibius dujardini (D o y e r e, 1840) Hypsibius cf. morikawai I t o, 1995 Hypsibius pallidus T h u l i n, 1911 Isohypsibius lunulatus (I h a r o s, 1966) Isohypsibius prosostomus T h u l i n, 1928 Diphascon (D.) brevipes (M a r c u s, 1936) Diphascon (D.) pingue (M a r c u s, 1936) Diphascon (A.) belgicae R i c h t e r s, 1911 Diphascon (A.) prorsirostre T h u l i n, 1928 Diphascon (A.) scoticum M u r r a y, 1905 Astatumen bartosi (W e g l a r s k a, 1959) Astatumen trinacriae (A r c i d i a c o n o, 1962) Family: Macrobiotidae Macrobiotus cf. harmsworthi M u r r a y, 1907 Macrobiotus hufelandi C.A.S. S c h u l t z e, 1834 Macrobiotus pallarii M a u c c i, 1954 Macrobiotus cf. seychellensis B i s e r o v, 1994

Appendix 1. (Continued) Macrobiotus cf. vanescens P i l a t o et al., 1991 Macrobiotus sp.1 Minibiotus intermedius ( P l a t e, 1889) Minibiotus sp. 1 Phylum: ARTHROPODA Subphylum: CHELICERATA Order: PSEUDOSCORPIONES Family: Chthoniidae Chthonius (Ephippiochthonius) boldorii B e i e r, 1934 Chthonius (Ephippiochthonius) fuscimanus S.i.m.o.n, 1900 Chthonius (Ephippiochthonius) sp. 1 Chthonius (Ephippiochthonius) sp. 2 Family: Neobisiidae Neobisium (Neobisium) erythrodactylum (C. L. K.o.c.h, 1873) Neobisium (Neobisium) muscorum (L e a c h, 1817) Roncus lubricus C. L. K o c h, 1873 Family: Chernetidae Chernes similis C. L. K o c h, 1873 Pselaphochernes scorpioides (H e r m a n n, 1804) Chernetidae gen. sp. Order: ARANEAE Family: Atypidae Atypus piceus (S c h u l z e r,1776) Family: Segestriidae Segestria senoculata (L i n n a e u s, 1758) Family: Dysderidae Dysdera erythrina (W a l c k e n a e r, 1802) Harpactea rubicunda (C. L. K o c h, 1838) Family: Mimetidae Ero furcata (V i l l e r s, 1789) Family: Theridiidae Enoplognatha oelandica (T h o r e l l, 1875) Enoplognatha ovata (C l e r c k, 1757) Enoplognatha thoracica (H a h n, 1833) Enoplognatha sp. Episinus angulatus (B l a c k w a l l, 1836) Euryopis flavomaculata (C. L. K o c h, 1836) Robertus lividus (B l a c k w a l l, 1836) Robertus sp. Steatoda albomaculata (D e G e e r, 1778) Steatoda bipunctata (L i n n a e u s, 1758) Steatoda phalerata (P a n z e r, 1801) Steatoda sp. Theridion tinctum (W a l c k e n a e r, 1802) Family: Linyphiidae Abacoproeces saltuum (L. K o c h, 1872) Anguliphantes angulipalpis (W e s t r i n g, 1851) Asthenargus paganus (S i m o n, 1884)

Bathyphantes nigrinus (W e s t r i n g, 1851) Centromerus arcanus (O. P. C a m b r i d g e, 1873) Centromerus brevivulvatus D a h l, 1912 Centromerus incilium (L. K o c h, 1881) Centromerus sellarius (S i m o n, 1884) Centromerus sylvaticus (B l a c k w a l l, 1841) Centromerus sp. Ceratinella brevis (W i d e r, 1834) Ceratinella major K u l c z y ń s k i, 1894 Ceratinela scabrosa (O. P. C a m b r i d g e, 1871) Diplocephalus latifrons (O. P. C a m b r i d g e, 1863) Diplocephalus picinus (B l a c k w a l l, 1841) Diplostyla concolor (W i d e r, 1834) Drapetisca socialis (S u n d e v a l l, 1833) Moebelia penicillata (W e s t r i n g, 1851) Lasiargus hirsutus (M e n g e, 1869) Lepthyphantes minutus (B l a c k w a l l, 1833) Linyphia hortensis S u n d e v a l l, 1830 Linyphia triangularis (C l e r c k, 1757) Linyphia sp. Macrargus rufus (W i d e r, 1834) Megalepthyphantes collinus (L. K o c h, 1872) Meioneta affinis (K u l c z y n s k i, 1898) Micrargus herbigradus (B l a c k w a l l, 1854) Micrargus sp. Microlinyphia pussila (S u n d e v a l l, 1830) Microneta varia (B l a c k w a l l, 1841) Minicia marginella (W i d e r, 1834) Neriene clatrata (S u n d e v a l l, 1830) Oedothorax retusus (W e s t r i n g, 1851) Palliduphantes alutacius (S i m o n, 1884) Palliduphantes pallidus (O. P. C a m b r i d g e, 1871) Panamomops fagei M i l l e r et K r a t o c h v í l, 1939 Pocadicnemis juncea L o c k e t et M i l l i d g e, 1953 Pocadicnemis pumila (B l a c k w a l l, 1841) Sintula corniger (B l a c k w a l l, 1856) Tapinocyba biscissa (O. P. C a m b r i d g e, 1872) Tapinocyba insecta (L. K o c h, 1869) Tenuiphantes mengei (K u l c z y ń s k i, 1887) Tenuiphantes tenbricola (W i d e r, 1834) Tenuiphantes flavipes (B l a c k w a l l, 1854) Tenuiphantes tenuis (B l a c k w a l l, 1852) Tenuiphantes zimmermani (B e r t k a u, 1890) Tenuiphantes sp. Tapinopa longidens (W i d e r, 1834) Thyreostenius biovatus (O. P. C a m b r i d g e, 1875) Trichoncus affinis K u l c z y n s k i, 1894 Walckenaeria antica (W i d e r, 1834)

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Appendix 1. (Continued) Walckenaeria atrotibialis (O. P. C a m b r i d g e, 1878) Walckenaeria corniculans (O. P. C a m b r i d g e, 1875) Walckenaeria cucullata (C. L. K o c h, 1836) Walckenaeria cuspidata B l a c k w a l l, 1833 Walckenaeria dysderoides (W i d e r, 1834) Walckenaeria furcillata (M e n g e, 1869) Walckenaeria incisa (O. P. C a m b r i d g e, 1871) Walckenaeria mitrata (M e n g e, 1868) Walckenaeria monoceros (W i d e r, 1834) Walckenaeria nudipalpis (W e s t r i n g, 1851) Walckenaeria obtusa B l a c k w a l l, 1836 Walckenaeria vigilax (B l ac k w a l l, 1853) Walckenaeria sp. Family: Tetragnathidae Pachygnatha degeeri S u n d e v a l l, 1830 Pachygnatha listeri S u n d e v a l l, 1830 Family: Araneidae Araneus diadematus C l e r c k, 1757 Araneus quadratus C l e r c k, 1757 Araneus sp. Cercidia prominens (W e s t r i n g, 1851) Family: Lycosidae Alopecosa aculeata (C l e r c k, 1757) Alopecosa pulverulenta (C l e r c k, 1757) Arctosa lutetiana (S i m o n, 1876) Pardosa agrestis (W e s t r i n g, 1861) Pardosa amentata (C l e r c k, 1757) Pardosa hortensis (T h o r e l l, 1872) Pardosa lugubris (W a l c k e n a e r, 1802) Pardosa monticola (C l e r c k, 1757) Pardosa sp. Trochosa ruricola (D e G e e r, 1778) Trochosa terricola T h o r e l l, 1856 Trochosa sp. Family: Pisauridae Pisaura mirabilis (C l e r c k, 1757) Family: Zoridae Zora nemoralis (B l a c k w a l l, 1861) Zora silvestris K u l c z y ń s k i, 1897 Zora spinimana (S u n d e v a l l, 1833) Family: Agelenidae Agelena gracilens C. L. K o c h, 1841 Agelena sp. Histopona torpida (C. L. K o c h, 1837) Tegenaria campestris C. L. K o c h, 1834 Tegenaria ferruginea (P a n z er, 1804) Tegenaria silvestris L. Koch, 1872 Tegenaria sp. Textrix denticulata (O l i v i e r, 1789) Family: Cybaeidae

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Cybaeus angustiarum L. K o c h, 1868 Family: Hahnidae Hahnia helveola S i m o n, 1875 Hahnia ononidum S i m o n, 1875 Family: Dictynidae Cicurina cicur (F a b r i c i u s, 1793) Dictyna uncinata T h o r e l l, 1856 Family: Amaurobiidae Coelotes inermis (L. K o c h, 1855) Coelotes terrestris (W i d e r, 1834) Coelotes sp. Family: Liocranidae Agroeca brunnea (B l a c k w a l l, 1833) Apostenus fuscus W e s t r i n g, 1851 Scotina celans (B l a c k w a l l, 1841) Family: Corinnidae Cetona laticeps (C a n e s t r i n i, 1868) Phrurolithus festivus (C. L. K o c h, 1835) Family: Zodariidae Zodarion germanicum (C. L. K o c h, 1837) Family: Clubionidae Clubiona caerulescens L. K o c h, 1867 Clubiona corticalis (Walckenaer, 1802) Clubiona comta C. L. Koch, 1839 Clubiona terrestris Westring, 1851 Clubiona sp. Family: Gnaphosidae Drassyllus praeficus (L. K o c h, 1866) Drassyllus villicus (T h o r e l l, 1875) Haplodrassus silvestris (B l a c k w a l l, 1833) Micaria dives (L u c a s, 1846) Trachyzelotes pedestris (C. L. K o c h, 1837) Zelotes apricorum (L. K o c h, 1876) Zelotes aurantiacus M i l l e r, 1967 Zelotes erebeus (T h o r e l l, 1871) Zelotes subterraneus (C. L. K o c h, 1833) Zelotes sp. Family: Philodromidae Philodromus dispar W a l c k e n a e r, 1826 Philodromus sp. Thanatus formicinus (C l e r c k, 1757) Family: Thomisidae Diaea dorsata (F a b r i c i u s, 1777) Ozyptila praticola (C. L. K o c h, 1837) Ozyptila simplex (O. P. C a m b r i d g e, 1862) Xysticus cristatus (C l e r c k, 1757) Xysticus lanio C. L. K o c h, 1835 Xysticus luctator C. L. K o c h, 1870 Xysticus sp. Family: Salticidae Ballus chalybeius (W a l c k e n a e r, 1802)

Appendix 1. (Continued) Carrhotus xanthogramma (L a t r e i l l e, 1819) Euophrys frontalis (W a l c k e n a e r, 1802) Euophrys sp. Evarcha falcata (C l e r c k, 1757) Marpisa sp. Neon reticulatus (B l a c k w a l l, 1853) Order: ACARINA Family: Celaenopsidae Celaenopsis badius (C. L. K o c h, 1841) Family: Ascidae Aceoseius muricatus (C. L. K o c h, 1839) Arctoseius eremitus (B e r l e s e, 1918) Arctoseius venustulus (B e r l e s e, 1916) Leioseius bicolor (B e r l e s e, 1918) Leioseius minusculus (B e r l e s e, 1905) Zerconopsis remiger (K r a m e r, 1876) Family: Eviphididae Eviphis ostrinus (C. L. K o c h, 1836) Family: Laelapidae Eulaelaps stabularis (C. L. K o c h, 1836) Haemogamasus nidi M i c h a e l, 1892 Hypoaspis aculeifer G. C a n e s t r i n i, 1884 Hypoaspis brevipilis H i r s c h m a n n, 1969 Hypoaspis imitata R e i t b l a t, 1963 Hypoaspis oblonga (H a l b e r t, 1915) Hypoaspis praesternalis W i l l m a n n, 1949 Hypoaspis vacua (M i c h a e l, 1891) Hypoaspis spp. Family: Macrochelidae Geholaspis longispinosus (K r a m e r, 1876) Geholaspis mandibularis (B e r l e s e, 1904) Macrocheles glaber (J. M ü l l e r, 1859) Macrocheles montanus W i l l m a n n, 1951 Family: Macronyssidae Ornithonyssus sylviarum (C a n e s t r i n i et F.a.n.z.a.g.o, 1877) Family: Pachylaelapidae Olopachys suecicus S e l l n i c k, 1950 Pachylaelaps magnus (H a l b e r t, 1915) Pachylaelaps resinae K a r g, 1971 Pachylaelaps spp. Pachyseius humeralis B e r l e s e, 1910 Family: Parasitidae Amblygamasus sp. Eugamasus monticolus B e r l e s e, 1905 Holoparasitus calcaratus (C. L. K o c h, 1839) Holoparasitus sp. Leptogamasus succineus W i t a l i ń s k i, 1973 Leptogamasus spp. Parasitus fimetorum (B e r l e s e, 1903) Pergamasus barbarus B e r l e s e, 1904 Pergamasus brevicornis (B e r l e s e, 1903) Pergamasus canestrinii (B e r l e s e, 1884)

Pergamasus crassipes (L i n n a e u s, 1758) sensu B e r l e s e, 1906 Pergamasus mediocris (B e r l e s e, 1904) Pergamasus ruhmi W i l l m a n n, 1938 Poecilochirus carabi G.et R.C a n e s t r i n i, 1882 Porrhostaspis lunulata J. M ü l l e r, 1869 Vulgarogamasus kraepelini (B e r l e s e, 1904) Vulgarogamasus remberti (O u d e m a n s, 1912) Family: Rhodacaridae Cyrtolaelaps chiropterae K a r g, 1971 Cyrtolaelaps mucronatus (G. et R. C a n e s t r i n i, 1881) Rhodacarus spp. Sessiluncus hungaricus K a r g, 1964 Stylochirus fimetarius (J. M ü l l e r, 1859) sensu M.a.š.á.n et K a l ú z, 2001 Family: Veigaiaidae Veigaia cerva (K r a m e r, 1876) Veigaia exigua (B e r l e s e, 1917) Veigaia kochi (T r ä g å r d h, 1901) Veigaia nemorensis (C. L. K o c h, 1839) Veigaia transisalae (O u d e m a n s, 1902) Family: Zerconidae Prozercon carpathofimbriatus M a š á n et F.e.n.ď.a, 2004 Prozercon tragardhi (H a l b e r t, 1923) Zercon curiosus T r ä g å r d h, 1910 Zercon hungaricus S e l l n i c k, 1958 Zercon peltatus var. peltatus C. L. K o c h, 1836 Zercon vacuus C. L. K o c h, 1839 Family: Sejidae Sejus togatus C. L. K o c h, 1836 Family: Uropodellidae Asternolaelaps sp. Family: Polyaspidae Polyaspinus schweizeri (H u ţ u, 1976) Polyaspis patavinus B e r l e s e, 1881 Family: Trachytidae Trachytes aegrota (C. L. K o c h, 1841) Trachytes baloghi H i r s c h m a n n et Z.i.r.n.g.i.e.b.l–N.i.c.o l, 1969 Family: Trematuridae Trichouropoda elegans (K r a m e r, 1882) Trichouropoda karawaiewi (B e r l e s e, 1904) Trichouropoda obscurasimilis H i r s c h m a n n et Z i r n g i e b l–N i c o l, 1961 Trichouropoda orbicularis (C. L. K o c h, 1839) Trichouropoda ovalis (C. L. K o c h, 1839) Trichouropoda penicillata H i r s c h m a n n et Z.i.r.n.g.i e b l–N i c o l, 1961 Family: Urodinychidae Dinychus bincheaecarinatus H i r s c h m a n n,

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Appendix 1. (Continued) W.a.g.r.o.w.s.k.a–A d a m c z y k et Z.i.r.n.g.i.e.b.l–N i c o l, 1984 Dinychus perforatus K r a m e r, 1886 Urodiaspis tecta (K r a m e r, 1876) Uroobovella pulchella (B e r l e s e, 1904) Family: Uropodidae Uropoda misella (B e r l e s e, 1916) Uropoda orbicularis (O. F. M ü l l e r, 1776) Uropoda splendida K r a m e r, 1882 Subphylum: BRANCHIATA Order: ISOPODA Suborder: Oniscidea Family: Ligiidae Ligidium hypnorum (C u v i e r, 1792) Family: Trichoniscidae Hyloniscus riparius (C. L. K o c h, 1838) Family: Philosciidae Lepidoniscus minutus (C. L. K o c h, 1838) Family: Agnaridae Orthometopon planum (B u d d e–L u n d, 1885) Protracheoniscus politus (C. L. K o c h, 1841) Family: Trachelipodidae Trachelipus ratzeburgii (B r a n d t, 1833) Porcellium collicola (V e r h o e f f, 1907) Subphylum: MYRIAPODA Class: CHILOPODA Order: SCOLOPENDROMORPHA Family: Cryptopidae Cryptops anomalans Newport, 1844 Order: GEOPHILOMORPHA Family: Dignathodontidae Henia illyrica (M e i n e r t, 1870) Family: Geophilidae Clinopodes flavidus C. L. K o c h, 1847 Geophilus flavus (D e G e e r, 1778) Family: Linotaeniidae Strigamia acuminata (L e a c h, 1814) Strigamia transsilvanica (V e r h o e f f, 1928 Family: Schendylidae Schendyla nemorensis (C. L. K o c h, 1836) Order: LITHOBIOMORPHA Family: Lithobiidae Lithobius agilis C. L. K o c h, 1847 Lithobius borealis M e i n e r t, 1868 Lithobius dentatus C. L. K o c h, 1844 Lithobius erythrocephalus C. L. K o c h, 1847 Lithobius forficatus (L i n n a e u s, 1758) Lithobius lapidicola M e i n e r t, 1872 Lithobius latro M e i n e r t, 1872 Lithobius lucifugus L. K o c h, 1862 Lithobius melanops N e w p o r t, 1845 Lithobius mutabilis L. K o c h, 1862

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Lithobius muticus C. L. K o c h, 1847 Lithobius pelidnus H a a s e, 1880 Lithobius piceus L. K o c h, 1862 Lithobius aeruginosus L. K o c h, 1862 Lithobius austriacus (V e r h o e f f, 1937) Lithobius crassipes L. K o c h, 1862 Lithobius microps M e i n e r t, 1868 Class: DIPLOPODA Order: GLOMERIDA Family: Glomeridae Glomeris connexa C. L. K o c h, 1847 Glomeris hexasticha B r a n d t, 1833 Family: Trachysphaeridae Trachysphaera costata (W a g a, 1857) Order: POLYZONIIDA Family: 48)zoniidae Polyzonium germanicum B r a n d t, 1837 Order: JULIDA Family: Julidae Cylindroiulus boleti (C. L. K o c h, 1847) Enantiulus nanus (L a t z e l, 1884) Julus curvicornis V e r h o e f f, 1899 Kryphioiulus occultus (C. L. K o c h, 1847) Leptoiulus proximus (N e m e c, 1896) Megaphyllum projectum (V e r h o e f f, 1894) Megaphyllum unilineatum (C. L. K o c h, 1838) Ommatoiulus sabulosus (L i n n a e u s, 1758) Unciger foetidus (C. L. K o c h, 1838) Unciger transsilvanicus (V e r h o e f f, 1899) Order: CHORDEUMATIDA Family: Mastigophorophyllidae Haploporatia eremita (V e r h o e f f, 1909) Family: Craspedosomatidae Craspedosoma rawlinsii L e a c h, 1815 Order: POLYDESMIDA Family: Paradoxosomatidae Strongylosoma stigmatosum (E i c h w a l d, 1830) Family: Polydesmidae Polydesmus complanatus (L i n n a e u s, 1761) Subphylum: HEXAPODA Class: INSECTA Order: DERMAPTERA Family : Forficulidae Chelidurella acanthopygia (G é n é, 1832) Order: HEMIPTERA Suborder: HETEROPTERA Family: Tingidae Acalypta musci (S c h r a n k, 1871) Acalypta spp. nymphae Campylosteira verna (F a l l é n, 1826) Family: Microphysidae Loricula exilis (F a l l é n, 1807)

Appendix 1. (Continued) Family: Miridae Lygus rugulipennis P o p p i u s, 1911 Deraeocoris lutescens (S c h i l l i n g, 1837) Family: Nabidae Nabis b. brevis S c h o l z, 1847 Nabis p. pseudoferus R e m a n e, 1949 Nabis rugosus (L i n n a e u s, 1758) Family: Anthocoridae Orius sp. Family: Aradidae Aneurus l. laevis (F a b r i c i u s, 1775) Family: Oxycarenidae Metopoplax origani (K o l e n a t i, 1845) Family: Rhyparochromidae Drymus b. brunneus (R. F. Sahlberg, 1848) Drymus sylvaticus (F a b r i c i u s, 1775) Emblethis verbasci (F a b r i c I u s, 1803) Eremocoris plebejus (F a l l é n, 1807) Eremocoris podagricus (F a b r i c i u s, 1775) Megalonotus chiragra (F a b r i c i u s, 1794) Megalonotus sabulicola (T h o m s o n, 1870) Peritrechus gracilicornis P u t o n, 1877 Peritrechus nubilus (F a l l é n, 1807) Plinthisus brevipennis (L a t r e i l l e, 1807) Raglius alboacuminatus (G o e z e, 1778) Taphropeltus hamulatus (T h o m s o n, 1870) Trapezonotus a. arenarius (L i n n a e u s, 1758) Tropistethus holosericus (S c h o l z, 1846) Tropistethus sp. Family: Piesmatidae Piesma capitatum (W o l f f, 1804) Piesma maculatum (L a p o r t e, 1833) Family: Pyrrhocoridae Pyrrhocoris apterus (L i n n a e u s, 1758) Family: Coreidae Ceraleptus gracilicornis (H e r r i c h – S.c.h.a.e.f.f.e.r, 1835) Family: Rhopalidae Rhopalus parumpunctatus (S c h i l l i n g, 1829) Family: Cydnidae Legnotus limbosus (G e o f f r o y, 1785) Microporus nigritus (F a b r i c i u s, 1794) Tritomegas bicolor (L i n n a e u s, 1758) Tritomegas sexmaculatus (R a m b u r, 1839) Order: COLEOPTERA Suprafamily: CURCULIONOIDEA Family: Anthribidae Anthribus albinus (L i n n a e u s, 1758) Family: Apionidae Catapion seniculus (K i r b y, 1808) Ceratapion gibbirostre (G y l l e n h a l, 1813) Cyanapion columbinum (G e r m a r, 1817)

Diplapion confluens (K i r b y, 1808) Eutrichapion punctigerum (P a y k u l l, 1792) Holotrichapion aestimatum (F a u s t, 1890) Holotrichapion pisi (F a b r i c i u s, 1801) Ischnopterapion virens (H e r b s t, 1797) Kalcapion pallipes (K i r b y, 1808) Omphalapion hookerorum (K i r b y, 1808) Protapion apricans (H e r b s t, 1797) Protapion fulvipes (G e o f f r o y, 1785) Protapion trifolii (L i n n a e u s, 1768) Protapion nigritarse (K i r b y, 1898) Pseudapion rufirostre (F a b r i c i u s, 1775) Synapion ebeninum (K i r b y, 1808) Trichopterapion holosericeum (G y l l e n h a l, 1833) Family: Erirhinidae Tanysphyrus lemnae (P a y k u l l, 1792) Family: Curculionidae Acalles camelus (F a b r i c i u s, 1782) Acalles fallax B o h e m a n 1844 Acalles echinatus (G e r m a r, 1824) Acallocrates colonnellii (B a h r, 2003) Barypeithes albinae F o r m a n e k, 1903 Barypeithes chevrolati (B o h e m a n, 1843) Barypeithes mollicomus (A h r e n s, 1812) Bradybatus creutzeri G e r ma r, 1824 Bradybatus fallax G e r s t a e c k e r, 1860 Bradybatus kellneri B a c h, 1854 Brachysomus echinatus (B o n s d o r f f, 1785) Brachysomus dispar P e n e c k e, 1910 Brachysomus hirtus (B o h e m a n, 1845) Brachysomus setiger (G y l l e n h a l, 1840) Calosirus apicalis (G y l l e n h a l, 1727) Ceutorhynchus alliariae Ch. B r i s o u t, 1860 Ceutorhynchus chalibeus G e r m a r, 1824 Ceutorhynchus erysimi (F a b r i c i u s, 1787) Ceutorhynchus minutus (R e i c h, 1797) Ceutorhynchus obstrictus (M a r s h a m, 1802) Ceutorhynchus pallidactylus (M a r s h a m, 1802) Ceutorhynchus rhenanus S c h u l t z e, 1895 Ceutorhynchus scrobicollis N e r e s h e i m e r et W a g n e r, 1924 Ceutorhynchus typhae (H e r b s t, 1795) Coeliodes proximus S c h u l t z e, 1895 Coeliodes rana (F a b r i c i u s, 1787) Coeliodes transverseoalbofasciatus G o e z e, 1777 Coeliodes trifasciatus B a c h, 1854 Curculio glandium (M a r s h a m, 1802) Curculio pellitus (B o h e m a n, 1843) Curculio venosus (G r a v e n h o r s t, 1807) Furcipus rectirostris (L i n n a e u s, 1758) Hypera nigrirostris (F a b r i c i u s, 1775)

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Appendix 1. (Continued) Hypera postica (G y l l e n h a l, 1813) Kyklioacalles suturatus (D i e c k m a n n, 1983) Leiosoma cribrum (G y l l e n h a l, 1834) Microplontus campestris (G y l l e n h a l, 1837) Nedyus quadrimaculatus (L i n n a e u s, 1758) Ophrohinus suturalis (F a b r i c i u s, 1775) Otiorhynchus ovatus (L i n n a e u s, 1758) Otiorhynchus raucus (F a b r i c i u s, 1777) Phyllobius argentatus (L i n n a e u s, 1758) Phyllobius maculicornis G e r m a r, 1824 Polydrusus marginatus S t e p h e n s, 1831 Polydrusus viridicinctus G y l l e n h a l, 1834 Rhinoncus bruchoides (H e r b s t, 1784)

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Rhinoncus perpendicularis (R e i c h, 1797) Rhynchaenus pilosus (F a b r i c i u s, 1781) Ruteria hypocrita (B o h e m a n, 1837) Sciaphilus asperatus (B o n s d o r f f, 1785) Simo variegatus (B o h e m a n, 1843) Sitona humeralis S t e p h e n s, 1831 Sitona lineatus (L i n n a e u s, 1758) Sitona macularius (M a r s h a m, 1802) Stenocarus cardui (H e r b s t, 1784) Strophosoma melanogrammum (F o r s t e r, 1771) Trachodes hispidus (L i n n a e u s, 1758) Trachyphloeus bifoveolatus (B e c k, 1817) Tychius picirostris (F a b r i c i u s, 1787)