Spatial and Temporal Dynamics of Invertebrates Dwelling Karstic Mesovoid Shallow Substratum of Sivec National Nature Reserve (Slovakia), with Emphasis on Coleoptera

Biologia 67/6: 1143—1151, 2012 Section Zoology DOI: 10.2478/s11756-012-0113-y

Spatial and temporal dynamics of invertebrates dwelling karstic mesovoid shallow substratum of Sivec National Nature Reserve (Slovakia), with emphasis on Coleoptera

Michal Rendoš1, Andrej Mock1*&TomášJászay2

1Pavol Jozef Šafárik University, Faculty of Sciences, Institute of Biology and Ecology, Moyzesova 11,SK-04167 Košice, Slovakia; e-mail: [email protected], [email protected] 2The Šariš Museum in Bardejov, Radničné námestie 13,SK-08501 Bardejov, Slovakia; e-mail: [email protected]

Abstract: Interior spaces of the forested rocky debris (MSS) represent a transition zone between the surface and deep underground spaces and a place of animal adaptation to underground life. They serve as a refuge for relict fauna as well. The study was conducted in the limestone scree slopes in Sivec National Nature Reserve (Čierna Hora Mts, Western Carpathians, elevation about 500 m a. s. l.) covered by linden-maple forest from September 2008 to November 2009. The effort was to define the vertical and seasonal aspects of invertebrates and temperature regime. Invertebrates were collected by using subterranean traps (plastic cups with 4% formaldehyde, inserted into the depths 5–95 cm through a plastic tube), which were checked monthly. Almost 26,000 specimens were trapped. Arthropods highly dominated over gastropods and earthworms. Collembola (67.61%) and Acarina (15.55%) were eudominant. Macrofauna was represented mainly by larvae of Holometabola (7.55%) and adult Diptera (5.11%) and Coleoptera (1.13%). All these groups were captured along the total depth gradient. Coleoptera were studied in more details. Among 11 Coleoptera families, Staphylinidae predominated and were captured at all levels. Rather high species diversity was found: 67 spp. excluding common epigeic fauna. Some species supposed to be subterranean, e.g., Bryaxis frivaldszkyi slovenicus, Duvalius bokori valyianus and Omalium validum. Activity of most invertebrate groups decreased significantly with depth (prevalence of surface fauna), but it was not terminated at 1 m under surface; the same was true for beetles, both in activity and diversity. Conspicuous fact is that a mass of subterranean species were traced also close to the surface (35 cm), i.e., probably it is not necessary to put the traps as deep as in this study. Seasonal climate changes affected the activity of invertebrates which was the highest at the end of spring and the lowest during winter, but it was not completely interrupted. Microclimate was characteristic without major temperature fluctuations on the surface. It was stable deeper along with increasing average annual temperature. High diversity and the occurrence of rare faunistic elements as well as specific habitats of MSS are perspective study objects and they merit care; mature design of the next studies considering the effect of season and depth of traps deposition shall do them more effective and less laborious. Key words: mesovoid shallow substratum; invertebrates; Coleoptera; vertical distribution; diversity; temperature regime; seasonal dynamics; limestone scree; Slovakia

Introduction carbonate rocks (Juberthie et al. 1980; Juberthie & De- lay 1981; Camacho 1992). Parameters that affect bio- The concept of Mesovoid Shallow Substratum (origi- diversity in MSS are thickness of soil layer, density of nally in French, milieu souterrain superficiel, MSS) was vegetation cover and especially the flow of organic car- defined by Juberthie et al. (1980). It consists of a sys- bon, which is stronger than in the cave, extension and tem of empty ventilated spaces within the stony debris continuity of subterranean spaces (Gers 1998; Pipan et covered by soil. This formation is in most cases the re- al. 2011). Network of cracks within the bedrock repre- sult of frost weathering in temperature zone. Rock frag- sented a suitable shelter for surviving of invertebrate ments accumulate in several layers on the bedrock. In fauna during the last glaciation. Close contact with the the final phase they are covered by a layer of soil, which surface and stable conditions made the MSS an envi- isolates the MSS from the surface and makes it a sta- ronment that is more attractive and especially more af- ble environment with conditions close to those in caves fordable for fauna and finally more spacious than caves. (Juberthie 2000; Giachino & Vailati 2010). This habi- The adaptation of animals to hypogean environment tat was studied mainly in temperate mountain ranges may also be in progress in this transition zone (Pipan (Culver & Pipan 2009) in both carbonate and non- et al. 2011; Růžička 1993, 1999). There is an assump-

* Corresponding author

c 2012 Institute of Zoology, Slovak Academy of Sciences 1144 M. Rendoš et al.

Table 1. Some characters of yearly temperatures at the study plot ( ◦C). The months are marked by Roman numerals.

Depth Max Min Range Average Max 1-day contrast

Surface 20.5 (VII) –4.0 (I) 24.5 7.9 5.5 (IV) 15 cm 18.5 (VII) 1.0 (III) 17.5 8.6 1.0 35 cm 17.5 (VII) 1.5 (II–III) 16.0 7.9 1.0 55 cm 17.5 (VII–VIII) 2.5 (II–III) 15.0 9.1 0.5 95 cm 16.0 (VIII) 3.5 (II–IV) 12.5 9.3 0.5

tion that the obligate cave fauna became semi-extinct tical levels (5, 15, 25, 35, 45, 55, 65, 75, 85 and 95 cm) where in Central Europe during the period of glaciation which plastic cups were fixed as traps and small holes (ø 7 mm) caused noticeable regional and latitudinal differences in were drilled around to pipe to allow the entry of inverte- its distribution (Culver et al. 2006). The subterranean brates into the traps. Top of the pipe was covered with tin spaces with the occurrence of troglobionts are abundant plates. The traps (volume 500 ml) were made of a flexible plastic, fixed on the moving central iron staff and filled with mainly in the Mediterranean region (Gers 1988; Pipan 4% formaldehyde for fixation. Triplicate of traps (or pipes) et al. 2011) while adapted surface fauna with presence were dug in line, at a distance of 30 cm and traps were con- of relics prevails in the northern regions, especially with trolled monthly from October 2008 to November 2009. The cold even permafrost type of MSS (Christian 1987). The obtained material was later fixed in alcohol and gradually type of rock has probably no influence on the special- identified to the level of higher taxa and Coleoptera to the ized subterranean fauna (Giachimo & Vailati 2010). species level using identification keys. The beetle nomencla- Invertebrates dwelling MSS were studied mainly in ture was according to Fauna Europaea (2011). Animals are Europe, especially in France, Romania, Slovenia, Aus- deposed in the authors’ collections. tria and the Czech Republic or in the Canary Islands, Two series of thermodataloggers (iButton DS 1921G focused on some model groups, such as beetles (e.g., #F50 Maxim Dallas, USA) were used for long-term and continuous measurement of temperature (four-hour inter- Gers 1986, 1988; Nitzu et al. 2007; Giachino & Vailati vals). The temperature was measured at five levels: on the 2010), oribatid mites (Arillo et al. 1994), spiders (Nae surface and inside the debris at depths of 15, 35, 55 and & Ilie 2004; Růžička & Zacharda 2010; Laška et al. 95 cm. 2011), centipedes (Ilie 2003a, b), springtails (Gers & Najt 1983; Querner & Krenn 2005) and flies (Gers 1993) or occasionally on the full invertebrate communities Results (e.g., Nitzu et al. 2010). The objectives of this study were to investigate the Temperature regime MSS dwelling invertebrate community in the karstic The largest temperature fluctuations were observed on part of the Čierna hora Mts (the Western Carpathi- the surface. Deeper under the surface, only long-term ans Mts), as the northernmost area of the obligate cave fluctuations were registered, the amplitudes were lower fauna in Central Europe and to monitor the spatial in depth, but synchronised with the climate dynamics and seasonal aspects of both microclimate and inver- on the surface (Table 1, Fig. 1). On the soil surface, the tebrates. Selection of the locality was based on the temperature decreased below zero only for short peri- regional cave fauna inventory (Mock et al. 2009). We ods in contrast to rather strong freezes in the air above presumed that comparisons between communities and the surface (more days with less than –10 ◦C). Deeper well-adapted species in both habitats, caves and scree than 15 cm no frost occurred. Average annual temper- slopes could lead to a more complex understanding of ature increased along the vertical gradient in contrast the subterranean environment functioning. to maximum day temperature.

Activity and seasonal dynamics of invertebrates along Material and methods the vertical gradient The research was conducted at one of the MSS sites (ge- Almost 26,000 individuals of 24 higher invertebrate ographic co-ordinates 48◦5031.27 Nand21◦0633.97 E) taxa were collected, and more than 99.8% of individu- in the National Nature Reserve (NNR) Sivec, the Western als belonged to Arthropoda; some earthworms and gas- Carpathians Mts (eastern Slovakia). A massif of Mesozoic tropods were collected besides the major groups. Meso- limestone with steep debris at its base is the typical fea- fauna, especially Collembola, was highly predominant ture of the reserve territory. The study plot is located at over macrofauna; more than 83% of all individuals be- 530 m a. s. l., a few metres under the top of the slope on longed to it. Adult Diptera and Coleoptera and the lar- the foothill of rock faces. The north-east scarred slope is val stages of several groups of holometabolic insects be- close to the valley and it is covered by the linden-maple forest (Tilio-Acerion). During the winter season the area is longed to the dominant groups, exceeding 5% (Table 2). usually covered by snow. Majority of the individuals of all groups (60%) was Invertebrates were collected by subterranean traps de- trapped at 5 cm under the surface. Only eight higher signed based on Schlick-Steiner & Schlick (2000) and Laška taxa were distributed along the entire vertical gradient et al. (2008): 110 cm long PVC pipe (ø 110 mm) with 10 ver- and 14 taxa were found in a depth of 95 cm. Besides Invertebrates dwelling karstic mesovoid shallow substratum 1145

Fig. 1. Daily average ﬂuctuations of air temperature within MSS at diﬀerent depths.

Table 2. Overall distribution of higher taxa of invertebrates along the vertical gradient. Depth (cm)/Taxon Gastropoda Lumbricidae Pseudoscorpiones Araneae Opiliones Acarina Isopoda Symphyla Pauropoda Chilopoda Diplopoda Collembola Diplura Plecoptera Psocoptera Thysanoptera Heteroptera Auchenorrhyncha Aphidinae Coleoptera Hymenoptera Formicoidea Mecoptera Diptera Larvae Total

5 17 5 17 96 18 2752 21 0 0 42 43 11454 33 0 19 0 0 9 22 237 41 56 1 247 427 15557 1531530361002492635301013192414101131093311 2520130165100566021000019511003072914 350010014100110275000000115003888552 451002092100214019021015260057177760 5510030142211333582000106140096151774 652023010410031455960000011817001752781182 7501030920003338480000094310175221907 8510010950004123972000000577002152461010 951003095000264961310000446101821921106

Total 28 7 26 117 18 4039 26 1 4 69 97 17561 95 1 22 1 2 14 91 293 114 59 1 1328 1961 25973

D % 0.11 0.03 0.10 0.45 0.07 15.55 0.09 0.00 0.02 0.27 0.37 67.61 0.37 0.00 0.08 0.00 0.01 0.05 0.35 1.13 0.44 0.23 0.00 5.11 7.55 100.00

the regular occurrence of some invertebrate groups, Seasonal climate changes affected the activity of Symphyla, Plecoptera, Thysanoptera, Mecoptera and invertebrates. Fluctuations of the locomotion activity scorpionflies were sporadically present, and harvestmen (=presence in traps) during the year were observed were caught only at the surface. Both high taxa diver- in all dominant groups and at all depths (Fig. 3). In sity and specimen number declined strongly within the general, the activity culminated twice, during spring zone between –5 and –35 cm, then the decrease of both and autumn (summer and beginning of winter in Aca- parameters stopped and other horizons showed equal rina and adult Diptera, respectively). These fluctua- characteristics, sometimes even with moderate increase tions were observed along the depth gradient within (Fig. 3). Deeper, the structure of invertebrate communi- the talus, sometimes with moderate short-time de- ties changed, springtails highly predominated but their creases at deeper levels. The activity of some arthro- ratio slightly decreased with depth (from 74% to 51%), pods (Coleoptera, Diptera) was interrupted during the while the relative abundance of adult dipterans and in- winter and summer period, in winter for longer pe- sect larvae increased. riod, nevertheless the activity of the other invertebrates 1146 M. Rendoš et al.

Fig. 2. Monthly ﬂuctuations of the activity of dominant groups of invertebrates.

Fig. 3. Overall activity (number of individuals) and the number of higher invertebrate taxa along the vertical gradient.

(e.g., Collembola and Acarina) dropped only slightly robium fulvipenne (Gravenhorst, 1806) (Table 3). In (Fig. 4). general, the overall activity, species and family richness decreased with the depth from –5 to –35 cm. Species diversity and seasonal dynamics of activity of 58 spp. (85.3%) were recorded at 5 cm under the Coleoptera surface, while only 14, 4 and 1 species were found A total of 67 species of Coleoptera were captured (Ta- at –15 cm, –25 cm and –35 cm, respectively. Thus, ble 3). Staphylinidae was the most diversiﬁed among more than 95% of species occurred between the depths the 14 families (36 spp., 53.7%) and inhabited the of –5 to –35 cm. Deeper under the surface all men- entire vertical gradient (Table 4). Five staphylinid tioned parameters showed low values, their changes species predominated over the rest, each of them par- had no linear character and at the bottom of the ticipated with more than 5% in the community abun- vertical gradient they even rose slightly. Only 9 spp. dance: Liogluta microptera (Thomson, 1867), Omal- (12.2%) dwelled deeper than –35 cm, three of them ium rivulare (Paykull, 1789), O. rugatum Mulsant & (4.4%) were found exclusively there: Anaspis sp., Du- Ray, 1880, Bryaxis nigripennis (Aubé, 1844) and Lath- valius bokori valyianus (Bokor, 1922) and Eusphalerum Invertebrates dwelling karstic mesovoid shallow substratum 1147

Table 3. Diversity and spatio-temporal distribution of Coleoptera. Taxon Family Ind. D (%) Depth (cm) Months

1. Abax ovalis (Duftschmid, 1812) C 10 3.41 5, 25 V, VII, IX, XI 2. Abax parallelepipedus (Piller et Mitterpacher, 1783) C 5 1.71 5, 25 V, VI 3. Acrotona troglodytes (Motschulsky, 1858) S 3 1.02 5, 35 VI, X 4. Acrotrichis intermedia (Gillmeister, 1845) P 2 0.68 5 V, VII 5. Adexius scrobipennis (Gyllenhal, 1827) Cu 2 0.68 5, 15 VIII, XI 6. Agathidium laevigatum laevigatum Erichson, 1845 L 1 0.34 5 VIII 7. Agriotes pilosellus (Schönherr, 1817) E 2 0.68 5 VII 8. Amphichroum canaliculatum (Erichson, 1840) S 4 1.37 5 V,VI 9. Anaspis sp. Scr 3 1.02 85 VII, VIII, IX 10. Anthobium atrocephalum (Gyllenhal, 1827) S 9 3.07 5 V, VI, VIII, XI 11. Anthobium melanocephalum (Illiger, 1794) S 1 0.34 5 V 12. Anthophagus bicornis (Block, 1799) S 3 1.02 5, 15 V–VII 13. Atheta fungi (Gravenhorst, 1806) S 4 1.37 5 V–VII 14. Atheta putrida (Kraatz, 1856) S 5 1.71 5 XI 15. Atomaria sp. Cr 1 0.34 5 XI 16. Bryaxis frivaldszkyi slovenicus (Machulka, 1926) S 4 1.37 5, 65, 85, 95 V–VII, XI 17. Bryaxis nigripennis (Aubé, 1844) S 20 6.83 5, 15 V–VII, IX, X 18. Catops fuliginosus fuliginosus Erichson, 1837 L 1 0.34 5 XI Catops sp. L 1 0.34 5 X 19. Catops subfuscus subfuscus Kellner, 1846 L 1 0.34 5 V 20. Cephenium majus Reitter, 1881 Sc 2 0.68 5 V 21. Colon affine Sturm, 1839 L 1 0.34 5 IX 22. Cryptophagus sp. Cr 4 1.37 5 VI, VII, IX 23. Denticollis linearis (L., 1758) E 1 0.34 5 VII 24. Dienerella elongata (Curtis, 1830) L 1 0,34 15 XI 25. Duvalius bokori valyianus (Bokor, 1922) C 1 0.34 65 V 26. Euconnus (Cladoconnus)denticornis (Müller et Kunze, 1822) Sc 1 0.34 5 V 27. Euconnus transsilvanicus (Saulcy,1876) Sc 2 0.68 5 V, VI 28. Eusphalerum longipenne (Erichson, 1839) S 3 1.02 75 VI, VII 29. Eusphalerum rectangulum (Baudi di Selve, 1870) S 4 1.37 5 VI, VII 30. Eusphalerum semicoleoptratum (Panzer, 1795) S 1 0.34 5 VII 31. Gymnetron sp. Cu 1 0.34 15 VIII 32. Habrocerus capillaricornis (Gravenhorst, 1806) S 8 2.73 5, 15 VI, VII, XI 33. Chrysomelidae sp. Ch 2 0.68 15, 25 VIII,X 34. Ischnosoma longicorne (Mäklin, 1847) S 1 0.34 5 VI 35. Lathrobium fulvipenne (Gravenhorst, 1806) S 17 5.80 5–25, 45–95 VI-VIII 36. Leiodes sp. L 1 0.34 5 VII 37. Liogluta granigera (Kiesenwetter, 1850) S 11 3.75 5 X 38. Liogluta microptera Thomson, 1867 S 29 9.90 5, 15, 85 V-VII,X,XI 39. Medon fusculus (Mannerheim, 1830) S 2 0.68 15, 75 IX 40. Meligethes aeneus (F., 1775) N 2 0.68 5, 95 V,VII 41. Molops piceus piceus (Panzer, 1793) C 14 4.78 5, 15 V-VII,IX-XI 42. Nargus velox (Spence, 1815) L 1 0.34 5 IX 43. Neuraphes elongatulus (Müller et Kunze, 1822) Sc 1 0.34 5 V 44. Ocalea badia Erichson, 1837 S 4 1.37 5, 15 X, XI 45. Ocypus macrocephalus (Gravenhorst, 1802) S 1 0.34 5 V 46. Omalium rivulare (Paykull, 1789) S 27 9.22 5 V–VII, XI 47. Omalium rugatum Mulsant et Rey, 1880 S 20 6.83 5 V–VII 48. Omalium validum Kraatz, 1857 S 1 0.34 15 X 49. Othius punctulatus (Goeze, 1777) S 6 2.05 5, 15 I, II, IV, V, XII 50. Oxypoda acuminata (Stephens, 1832) S 2 0.68 5 XI 51. Oxypoda alternans (Gravenhorst, 1802) S 1 0.34 5 X 52. Oxypoda formosa Kraatz, 1856 S 4 1.37 5, 15 X, XI 53. Oxypoda vittata Märkel, 1842 S 3 1.02 5 XI 54. Parabolitobius inclinans (Gravenhorst, 1806) S 1 0.34 5 IV 55. Proteinus brachypterus (F., 1792) S 1 0.34 5 V 56. Proteinus crenulatus Pandellé, 1867 S 3 1.02 5, 45 XI 57. Pterostichus burmeisteri Heer, 1841 C 13 4.44 5, 15 IV–VIII, X, XI 58. Pterostichus foveolatus (Duftschmid, 1812) C 1 0.34 5 V 59. Pterostichus oblongopunctatus (F., 1787) C 1 0.34 5 V 60. Ptomaphagus variicornis (Rosenhauer, 1847) L 1 0.34 5 VII 61. Quedius fumatus (Stephens, 1833) S 1 0.34 5 VII 62. Quedius mesomelinus mesomelinus (Marsham 1802) S 1 0.34 15 V Quedius sp. S 1 0.34 5 V 63. Quedius suturalis Kiesenwetter, 1845 S 2 0.68 5 V, XII 64. Rhizophagus dispar (Paykull, 1800) M 1 0.34 5 IV 65. Scaphisoma assimile Ericson, 1845 Sca 1 0.34 5 X 66. Tachinus laticollis Gravenhorst 1802 S 1 0.34 5 VIII 67. Trimium carpathicum Saulcy, 1875 S 2 0.68 5 VI Total 293 100.00 Explanations: Abbreviated names of familes: C – Carabidae, Ch – Chrysomelidae, Cr – Cryptophagidae, Cu – Curculionidae, E – Elateridae, La – Latridiidae, L – Leiodidae, M – Monotomidae, N – Nitidulidae, P – Ptiliidae, S – Staphylinidae, Sc – Scydmaenidae, Sca – Scaphidiidae, Scr – Scraptiidae. 1148 M. Rendoš et al.

Fig. 4. Monthly ﬂuctuations of activity of the main invertebrate groups observed along the vertical gradient (logarithmic transforma- tion).

Table 4. Structure of the complete beetle assemblage at the family level.

Family Number of species D (%) inds D (%)

1. Carabidae 7 10.45 45 15.36 2. Chrysomelidae 1 1.49 2 0.68 3. Cryptophagidae 2 2.99 5 1.71 4. Curculionidae 2 2.99 3 1.02 5. Elateridae 2 2.99 3 1.02 6. Latridiidae 1 1.49 1 0.34 7. Leiodidae 7 10.45 8 2.73 8. Monotomidae 1 1.49 1 0.34 9. Nitidulidae 1 1.49 2 0.68 10. Ptiliidae 1 1.49 2 0.68 11. Staphylinidae 36 53.73 211 72.01 12. Scydmaenidae 4 5.97 6 2.05 13. Scaphidiidae 1 1.49 1 0.34 14. Scraptiidae 1 1.49 3 1.02

Total 67 100.00 293 100.00

longipenne (Erichson, 1839), belonging to diﬀerent fam- Nitidulidae, Scraptiidae and Staphylinidae, with the ilies. predominance of members of the latest family (66.7% On the family level, the highest diversity was con- of species) (Fig. 5). During the year, clear maximum centrated at surface of the debris (–5 to –35 cm), only of the total beetle activity was measured in the trap four of them were present at deeper levels: Carabidae, controls from May to July (e.g., during the months be- Invertebrates dwelling karstic mesovoid shallow substratum 1149

Fig. 5. Relative abundance, species and family richness of Coleoptera along the vertical gradient. fore the true controls). Among 47 spp. (29 spp. ex- face. On the contrary, bases of scree slopes can be per- clusively), captured in this period, also some of spe- manently cooled even as permafrost in such cases and cific representatives of the underground were collected these are typical climatic anomalies of slope deposits (e.g., D. bokori valyianus, Bryaxis frivaldszkyi sloveni- and their close surroundings. The climatic characteris- cus (Machulka, 1926)), but the other 20 species were tics of such habitats are equal to those that are typical active only out of this period (Table 3). of other altitudes or latitudes than the site under investigation, thus it is a potential refuge for organisms Discussion (Zacharda et al. 2007). Favourable microclimate without extreme temperatures enabled uninterrupted activ- Specific climate characteristics within the scree slopes ity of invertebrates during the whole year. Warmed un- have been studied by several authors (e.g., Nitzu et derground up to the surface probably caused that no al. 2007; Zacharda et al. 2007; Pipan et al. 2011). vertical migration of fauna was observed. In addition, The basic characteristics of the year-round tempera- no retardation of seasonal activity at any of the inves- ture regime is transient there compared to static cave tigated levels was observed. There is no general agree- spaces and surface (soil); it is more stable than on the ment about the depth of border between soil and MSS. surface but still clearly influenced by seasonal fluctua- For example, on the Canary Islands traps with 75 cm tions. Such conditions can be considered as suitable for long plastic tubes were used and troglomorphic inverte- adaptive processes leading to the obligate underground brates were recorded (López & Oromí 2010). Querner & strategy of life (like subterranean species) and mod- Greben-Krenn (2005) used only 50 cm tubes and they erate climate within the slope can be used as refuge also recorded some troglophilous springtails at various by relict fauna, locally supported by the presence of depths. At the site investigated in this study the max- such representatives of millipedes and springtails in the imum thickness of organic soil and rhizosphaera run scree (Rendoš et al., unpublished). In general, this phe- to the depth of 35–45 cm and a mixture of stones and nomenon was confirmed mainly in forested slopes or mineralised soil followed, except for solitary roots which in the MSS covered by a huge soil layer (Pipan et overgrew to a depth of several metres. al. 2011). Even the type of slope debris that was in- The distribution pattern with typical sudden de- vestigated in this study strongly impacts the surface cline of activity of majority of invertebrates (and species climate in de-escalation of extreme values of temper- diversity) along the depth gradient is probably caused ature in summer or winter, considering the fact that by rapid limitation of food sources and open spaces. its surrounding has also peculiar conditions (the slope Gers (1993, 1998) observed that the distribution of is “breathing”). In deeper horizons more stable and in- fauna in MSS was induced by accessibility of food (dead creased temperature were observed. The study plot was plant material), which was concentrated at the surface situated at the upper part of the slope, which is under and during rainfall it was distributed deeper under the the influence of warmer air deposited within the scree surface. The occurrence of some species is aggregated, (Růžička & Růžička 1990; Růžička 1993). The lower e.g., flies and their larvae (Gers 1993). We did not winter temperatures were only slightly below zero, al- recognize aggregation of any taxa, except of the gen- though during January the air temperature was under eral concentration of majority of the specimens close –10 ◦C and rather huge layer of snow covered the sur- to the surface. The distribution and activity of insect 1150 M. Rendoš et al. larvae was vertically continuous; more surprising was Coleoptera is the first model group we studied at a rather continuous vertical distribution of adult flies species level. Their relative activity was rather low, but and hymenopterans (mainly small body forms without based on the partial unpublished data on other inver- reduced wings). Deep penetration of plant louses can tebrate groups, beetles are the group with the highest be explained by deposition of food sources (long roots species richness dominating other groups of macrofauna of trees). Dry periods prevented water transport of or- that occurred within the scree. High species richness ganic material deeper to the underground. Availability (alpha diversity) was recognized and high value of di- of food, which is not continuous and permanent over versity index (Shannon’s index = 3.56, Simpson’s index the year, can essentially influence activity and repro- 1–D = 0.96) have premised total local diversity will duction of animals in the MSS (Gers 1998). Cone-like be higher in reality. Nevertheless such high biodiversity distribution pattern is characteristic by a strong drop with absence of obligate subterranean forms has shown in the number of trapped individuals along the vertical the upper horizons of scree habitat could be supposed gradient, especially at two uppermost levels. This trend as surface-subterranean ecotone. Deeper from –15 cm extends to 35 cm under the surface. Deeper, the activity it is continuously converted into true MSS. In compar- of animals is rather constant. In some of them the total ison with 20 caves and cavities in the same orographic vertical distribution is more like the shape of a sand- unit several collectors used a combination of collect- glass, most markedly in Diptera or Diplura. Expres- ing methods during multi-year investigation and they sive dominance of mesofauna (mites and springtails) found only 50 species (15 families), although diversity was expected, however, compared with similar studies of families was higher in the caves (Mock et al. 2009). In the peculiarities of particular sites are distinguished. both types of underground, caves and under the surface In some cases deep and continuous penetration of big- of scree slope the family Staphylinidae predominated, sized arthropods as well as lower relative abundance of followed by small-sized representatives of other families mesofauna were observed (Laška et al. 2011). The dif- (except of cave entrances with large-sized epigeic bee- ferences are probably due to peculiarities of the regional tles). Beetles belong especially to the “surface” species fauna and its potential to colonise the underground (Pi- without specific adaptations to life underground, how- pan et al. 2011) and differences in topography, latitude, ever, there were no large epigeic forms (Anoplotrupes, altitude, vegetation, exposition of the slope, but some Carabus spp. et al.), and even if no specific “MSS forms” contrasts can be subjective (e.g., differences in method- or troglobite was found, some troglophilous species were ology). present, e.g., the common Quedius mesomelinus or the Seasonal dynamics of invertebrate activity within infrequent Bryaxis frivaldszkyi slovenicus, B. nigripen- the slope followed changes in temperature, and there nis, Omalium validum and Duvalius bokori valyianus. is no evidence of occurrence of abundant species inde- But there is probably no potential for the occurrence of pendent of oscillation of climatic variables, which is in obligate troglobitic beetles in the whole West Carpathi- contrast with cave inhabitants which lost the seasonal ans (Košel 2009). In comparison with the local cave rhythm (Culver & Pipan 2009). However, in this phase fauna, in the scree slope there is an absence of some im- of investigation we are not able to exclude the occur- portant species (e.g., Atheta spelaea or Duvalius hun- rence of subterranean forms with other temporal model garicus), but their distribution pattern is discontinu- of activity, but based only on Coleoptera. These species ous, they were recorded in different parts of the Čierna are rare and can be caught only sporadically, being im- hora Mts (Mock et al. 2009) and there is no evidence possible to rate. of any troglobitic beetles in the Western Carpathians. Slight differences among the seasonal activity of From the faunistic point of view, the presence of some single groups of invertebrates were observed in this species is remarkable. B. frivaldszkyi slovenicus and D. study. The activity of dominant groups culminated bokori valyianus are endemic for the neighbouring part rather steeply during late spring months (Collembola, of the West Carpathians and they prefer subterranean Coleoptera), summer months (Acarina) to September habitats. A bulk of the species showed simple pattern (Diptera) and then it strongly decreased, probably due of seasonal activity similar to other groups of terrestrial to the dry period. Seasonal fluctuations of activity in arthropods in temperate forests in general: two peaks adult dipterans are probably caused by the end of yearly, especially the spring-summer one is massive. It the larval development during the maximum precipi- can be a good tool for faunistic short-time research: tation period (June–July) and consecutive incubation most of the beetles can be collected in the “fruitful” of adults. Close to the surface, both increase and de- late spring period close to surface. But there are many crease in seasonal activity of invertebrates were more scattered species dwelling deeper levels and occurring sudden and shorter in time in comparison with those as adults in different seasons. at deeper levels, except for Diptera, where the aestival peaks (at –65 cm, –85 cm, –95 cm) were the high- Acknowledgements est. Winter decline was longer than the aestival one (up to 4 months), but the activity of some inverte- The study was supported by the grant Vega 1/0139/09. Au- brates was continuous during all seasons, completely thors express great thanks to Dr. P. Ľuptáčik, J. Rendoš, interrupted only in some cases and levels for a short M. Goga, E. Miková, S. Kuchár (Košice) for the help during period. field works, Dr. P. Ľuptáčik also for checking of tempera- Invertebrates dwelling karstic mesovoid shallow substratum 1151 ture data, but mainly to Dr. I.H. Tuf and MSc J. Mikula Juberthie C., Delay B. & Bouillon M. 1980. 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