A Review of Spiders on Tree Trunks in Europe (Araneae)

European Arachnology 2005 (Deltshev, C. & Stoev, P., eds) Acta zoologica bulgarica, Suppl. No. 1: pp. 221-257.

A review of spiders on tree trunks in Europe (Araneae)

Csaba Szinetár1, Roland Horváth2

Abstract: The present paper provides an overview of the European publications concerning bark-dwelling spiders. A total of 29 works have been investigated during the work: articles, M.Sc. theses and books. Col- lections on which the publications were based upon, demonstrated the presence of a total of 298 spiders on the bark of different tree species. This high species number can be attributed mostly to the species occurring randomly on the tree bark; however, it can unanimously be stated that there are facultative and exclusive bark-dwelling spider species as well. Among the species occurring randomly, a high proportion is constituted by the ground-dwelling linyphiid spiders of small body, which make up a large proportion in the species number, however, their abundance is low. Different authors used different sampling methods and efforts. In spite of this, several conclusions could be drawn concerning the bark-dwelling spider assemblages: i) within the total population in tree bark, the widespread bark-dwelling species were dominant, regardless of tree species and location; ii) in the case of a single tree species, significant differences may be present within a smaller region – even within a town – in the species composition of spider assemblages; iii) the altitude plays an important role in the formation of the bark-dwelling spider assemblages within a given area; iv) the occurrence frequency is characteristically changing with the geographical latitude; v) the composition of the bark-dwelling spider assemblages is seasonally changing even within a single tree species. A significant difference is shown between the summer and winter assemblages.

Key words: bark-dwelling spiders, coniferous trees, deciduous trees, urban habitats, seasonality

Introduction Trees represent well-defined and unique habitats for animals. On the one hand, trees are struc- turally complex and include several microhabitats (foliage, branch, trunks). Thus, they provide ample opportunity for niche-segregation (Lawton 1978). On the other hand, trees provide a stable food source for animals dwelling on them (Southwood 1978). These characters are coupled with a large geographical range in most tree species (Strong 1979). All these factors result in a high species richness and diversity of tree-living animal communities. Tree trunks, as a distinctive microhabitat of the tree, are characterised by numerous unique biotic and abiotic environmental factors, which explains why we can also discern a separate, so-called bark-dwelling arthropod assemblage. Spiders represent a dominant taxon of bark-dwelling predatory arthropods. Well- recognisable adaptation phenomena, which can be observed in bark-dwelling spiders, provide evidence for the evolution of species living on or under tree bark. Adaptations of bark-dwelling spiders can be seen in morphology, phenology, and physiology. Relatively few studies have been published on bark-dwelling-spiders to date, and most of them are of a rather faunistical nature (Koslinska 1967, Albert 1976, Wunderlich 1982, Nicolai 1986, Hansen 1992, Malten 1994). Systematic surveys on bark-dwelling spiders were carried out in Hungary by the mid-nineties (Sebestyén 1996, Horváth, Szinetár 1998, 2002, Bogya et al. 1995, Horváth et al. 2001, 2004, 2005, Szinetár et al. 2002). Up to this date, primarily the black pine (Pinus nigra), Norway spruce (Picea abies), Scots pine (Pinus sylvestris), European beech (Fagus sylvatica), English

1 Berzsenyi College, Department of Zoology, Szombathely, Károlyi Gáspár ter. 4, H-9700, Hungary. E-mail: [email protected] 2 University of Debrecen, Department of Evolutionary Zoology and Human Biology, Debrecen, Hungary 221 European Arachnology 2005 oak (Quercus robur), London planetree (Platanus hybrida) and Apple tree (Pyrus malus) have been investigated for bark-dwelling spider species. In case of the Norway spruce, planetree and apple surveys were carried out in several Hungarian land regions while in the case of the other species, the surveys were carried out in Western Transdanubia only. Simultaneously with the Hungarian surveys, other countries in Europe also conducted similar studies recently (Simon 1995, Weiss 1995, Koponen 1996, 2004, Koponen et al. 1997, Rinne et al. 1998, Pekár 1999, Kubcová, Schlaghamerský 2002). Our overview on European bark-dwelling spider fauna is based on the results of nearly fourteen years of our investigations and foreign literature data.

Overview and evaluation on the survey methods The different authors employ various methods to collect spiders dwelling on the trunk and branches of the trees (parts covered by bark). Beyond the fact that all methods endeavour to capture the species dwelling on bark as efficiently as possible and provide the most comprehensive picture on the assemblages dwelling on bark, these methods resulted in very different samples that can be compared to each other in a very limited manner. The tree eclectors (various stammeclectors, modified pitfall trap, branch traps) of continuous operation, with killing liquid, and mounted on the trunk and the branches are in several aspects similar to the Barber pitfall trap. The animals get into the trap owing to their active movements. A number of similar, automated traps were employed by the authors (Nicolai 1986, Simon 1995, Weiss 1995, Koponen 1996, 2004, Koponen et al. 1997, Rinne et al. 1998, Kubcová, Schlaghamerský 2002). Different preservation liquids were used for the collecting containers e.g. NaCl solution, 4% formaldehyde, ethylene glycol. In this aspect too, the traps are similar to the Barber traps. The other widespread method of trapping from the bark is the trap band, differing significantly from the traps mentioned above. In Hungary and other countries too, one of the often-employed methods is the corrugated cardboard trap band (Bogya et al. 1995, Sebestyén 1996, Horváth, Szinetár 1998, 2002, Pekár 1999, Horváth et al. 2001, 2004, 2005, Kovács 2002). Bands are placed around the trunk, their waved surfaces facing the trunk and covering the whole perimeter of the tree, as if a kind of artificial bark. When sampling, the whole belt is removed from the trunk and placed into a plastic bag. The collection of the animals living on and under the bark is carried out in a laboratory. It can be considered that it is a difficulty (fault) of the method that the animals dwelling between the band and the bark – primarily in the summer period – can quickly escape when the belt is removed, therefore the fast and co-ordinated work of several collectors is needed when removing the trap bands. The vacuum sampler might be an efficient method for collection from the bark N( icolai 1986), and the sifting as well (Sebestyén 1996). Numerous collectors used the manual method, which means surveying the surface of the bark. This, of course is influenced by the structure of the bark (Nicolai 1986, Hansen 1992, Sebe- styén 1996). Scraping the bark off the trunks and branches: the essence of this method is that by disbarking the tree, the spiders dwelling between the bark layers and in the bottom surface of the bark can be collected with little effort and in large number both in terms of specimen and species. In the case of some tree species (e.g. Platanus hybrida), the bark layers can easily be removed, placed in plastic bags, then the surface of the trunk can be brushed and it can be placed along with the bark layers into the bag. The height of collection from the bark also significantly influences the results, which is well demonstrated by the trunk trap captures carried out at different heights (Simon 1995). He was able to establish stratocoenoses or assemblage characteristics to a layer, from the soil to the canopy of Scots pine. The sampling methods, the studied tree species according to sampling sites and the publications by the referenced authors are presented in Table 1. No detailed description is provided on the individual trap types employed by the authors, this can be found in the given publications.

222 Cs. Szinetár & R. Horváth: Spiders on tree trunks

As an addition, it can be mentioned that in case of collection by the beating method often employed for collection from foliage, numerous spider species might be sampled, which primarily dwell on the bark of the branches, instead of the foliage. In the case of these samples, according to the microhabitat choice, it can be established (assumed) whether a species dwells on the bark. The canopy spraying is a similar method. Although it can be assumed that like the beating method, the bark-dwelling species are also under-represented in the samples, in a small number, however, they still might be present (Benfatto et al. 1992).

Table 1. Sampling methods in the individual publications, studied tree species and survey locations (details on methods and locations can be found in the given publications).

Sampling site(s) Publication Sampling method(s) Investigated tree(s) Locality corrugated cardboard Bogya et al. 1999 apple tree, pear tree Nagykovácsi (Hungary) trap band Hansen 1992 hand colletion london planetree Venezia (Italy) Horváth, Szinetár 1998, 2002 and corrugated cardboard Szombathely, Velem, Bozsok, black pine Horváth et al. 2001, trap band Debrecen, (Hungary) 2004, 2005 7 different localities (SW- Koponen 1996 “new” branch trap english oak Finland) 7 different localities (SW- Koponen et al. 1997 “new” branch trap english oak Finland) “new” branch trap and Koponen 2004 english oak Ruissalo (SW-Finland) tree eclector scraping the bark off Koslinska 1967 apple tree Kraków, Lódź (Poland) the trunks and branches english oak, euro- corrugated cardboard 8 different localities in West Kovács 2002 pean beech, scots trap band Hungary pine Kubcová, Schlagham- Lednice, Soutok (Czech tree eclector english oak erský 2002 Republic) Magdeburg Malten 1994 tree eclector white willow (Germany) camperdown elm, hand collection, english oak, europe- Nicolai 1986 vacuum sampler, arbo- an beech, planetree Marburg (Germany) real photo-eclector maple, silver birch, white willow corrugated cardboard Horomĕřice, Doksany (Czech Pekár 1999 apple tree, pear tree trap band Republic) 7 different localities (SW- Rinne et al. 1998 “new” branch trap english oak Finland) corrugated cardboard trap band, 12 different localities (Hun- Sebestyén 1996 london planetree hand collection, gary) bark sifting Grunewald, Eberwalde Simon 1995 tree eclector scots pine (Germany) Nationalpark Bayerischer Weiss 1995 “modified pitfall traps” norway spruce Wald (Germany)

223 European Arachnology 2005

Owing to the differences in the collection methods, the collections are comparable to a certain extent only. Endeavours were made to draw conclusions and make reference to works where the statistical evaluation of standardised samplings was ensured.

Results and Discussion Evaluation and summary of the surveys carried out on the tree trunks and branches According to the results of the publications hereby examined, a total of 298 spider species can be indicated as inhabiting the bark of different tree species in Europe. This accounts for about a quarter of the European spider fauna. This extremely high species number shows that although the majority of the spiders was of accidental occurrence, the spiders are present on the bark for shorter or longer periods, therefore the presence of species characteristic to other microhabitats should also be reckoned with. The more balanced microclimate, diverse bark structure and the high number of hiding places can also be considered an advantageous condition in the case of the trees. The tree trunks provide a willy-nilly landing strip for the ballooning species or those using the aerial dispersal. Certainly, this is the reason for the occurrence of mature forms of small species, dwelling mostly on the ground; however, this presence is presumably of a short duration only. Moreover, the bark is rich in potential prey animals, fulfilling by this the role of an advantageous feeding place in case of a temporary stay (Horváth et al. 2005). The fallen, decayed old trunks have to be mentioned separately. Such works were also examined (Weiss 1995, Kubcová, Schlaghamerský 2002). In such cases the trunks are in close proximity to the ground, their moss and lichen cover might be more developed than in the case of living and upright tree trunks, thence on these trunks the ground-dwelling species settle in a much greater number, which is well demonstrated by the work of Weiss (1995). Classification the bark-dwelling spiders into types according to the strength (closeness) of the relationship

Wunderlich’s (1982) study is still a determinant publication in the European literature. He used the categories on which our study is also based. Real or exclusive bark-dwellers (E): spider species which, as microhabitat specialists, live on or under the bark. They can be found on the bark throughout the year. It is characteristic for them to have mature forms for most of the year or even continuously. Their adaptation to the microhabitat could result in well-distinguishable morphological signs. Facultative bark-dwellers (F1, F2): species which use tree trunks and branches as typical but not exclusive microhabitats with permanent or seasonal character. Rocks, cracks in rocks, artificial walls may also be typical habitats of these species. Spider species dwelling on trees and choosing the bark as a dwelling place for the winter are also classified into this category. This is characteristic primarily for those dwelling in the foliage of deciduous trees. In the case of facultative bark-dwellers we considered it necessary to establish two further sub-categories in order to indicate the frequency of the facultative relationship. F1 is a closer relationship that can be observed more frequently, while F2 is a rareer one, however, it indicates spider-tree species relationships observed by several authors and in several instances. Accidental species (A): The species are listed here regardless of their occurrence frequency. They are species for which other microhabitats mean the typical habitat or such which have no specific habitat preference. In the case of these species it is presumed that the time of stay on the bark is short, however, it is very difficult to study or demonstrate it. These categories can certainly be stated more precisely based on new data gathered on the species. There are rare species for which – owing to the low population number – no reliable knowledge

224 Cs. Szinetár & R. Horváth: Spiders on tree trunks on the microhabitat selection is at our disposal. Apendix 1. includes all species demonstrated from trees together with their short phenological, habitat selection characterisation as well as literature references. Data on 298 spider species were obtained while examining 29 publications, books, and theses. Based on the evaluation of the relationship between the spiders and the bark, the distribu- tion of categories is as follows: 65% of the total species number was constituted by accidental species. In other species, a relationship of different strength and various habitat preferences can be stated. Twenty-four species were classified (8% of the total number of species) into the real bark-dwelling species. Seventy-nine species can be deemed as facultative bark-dwellers. Out of this, 50 species were ranked into the F1 type (17%), while 29 species (10%) were ranked into the F2 type according to the collections, reference literature and own experiences.

Taxonomic notes Several authors draw attention to the great and striking morphological similarity between Clubiona genevensis L. Koch, 1866 and Clubiona leucaspis Simon, 1932, and to the possibil- ity of a misidentification (Malten 1984, Nentwig et al. 2003, Weiss, oral communication). The habitats of the two species are largely different. C. genevenis is a ground-dwelling species of dry sandy lands, whereas C. leucaspis is a typical bark-dwelling species. Following Simon’s (1932) description, Di Franco (1993) published a drawing of C. leucaspis, as well as on the biology of the species from lemon plantations in Sicily. In our opinion the drawing in Figure 12/b on page 56 in Sterghiu (1985), also published by Nentwig et al. (2003), is not of C. genevensis but of C. leucaspis. Likewise, we assume that the occurrence of C. genevensis on trunks and barks is mistakenly reported and stands for the rather similar C. leucaspis, cf. Sterghiu (1985), Heimer, Nentwig (1991), Simon (1995).

Spider assemblages Based on survey results of other tree species and geographical locations, we can formulate the following statements concerning the bark-dwelling spider assemblages: i) In samples from tree trunks, generally irrespective of the tree species and location, the widespread bark-dwelling spider species dominate (Appendix 1). Well-recognisable adaptation phenomena, which can be observed in bark-dwelling spiders, provide evidence for the evolution of species living on or under the bark. This process is a sequential evolution, in which the relationship between the plant and the arthropod living on it is essentially asymmetric, i.e. not of a coevolutionary type, rather, the animals follow the evolution of the host plant without significantly affecting it (Jermy 1987). Adaptations of bark-dwelling spiders can be seen in morphology, phenology, and physiology. One of the morphological adaptations is the typical body shape observable in many species (e.g. Araneidae: Nuctenea umbratica (Clerck, 1757); Philodromidae: Philodro- mus margaritatus (Clerck, 1757), Philodromus fuscomarginatus (De Geer, 1778); Thomisidae: Coriarachne depressa (C. L. Koch, 1837)). The body of these species is characteristically flat- tened, which well reflects an adaptation to modes of living between the bark layers or under the bark. This adaptation can be coupled with colour adaptations. Philodromus fuscomarginatus, for example, blends well with its reddish-brown coloration into the phloem layer of Scots pine (Pinus sylvestris), which is the almost exclusive habitat of this philodromid spider. Philodromus margaritatus with its dark colours blends well into the surface of trees with darker bark, and can thus hide from predators, mainly birds, which feed on it. The most important of the phenological adaptations is that many exclusive bark-dwelling species are eurychron, i.e. adults and juveniles can be found all year-round, or diplochron, i.e., the species has two reproductive periods annu- ally. Reproduction throughout the entire year or throughout a substantial part of the year is made possible by the higher and more even temperature under the bark.

225 European Arachnology 2005

ii) In the case of a given tree species significant differences can be found within a more restricted region – even within a city. In urban sampling areas with strongly polluted air a number of species are missing (e.g. Clubiona pallidula (Clerck, 1757)) or are represented by a smaller population size (e.g. Moebelia penicillata (Westring, 1851)) which, in the garden suburb samples are dominant species on the same tree species (Horváth et al. 2001). iii) The altitude has also an important role in the formation of a bark-dwelling spider assemblage of a given area. In mountain areas some species appear (e.g. Segestria senoculata (Lin- naeus, 1758), Clubiona corticalis (Walckenaer, 1802), Amaurobius fenestralis (Ström, 1768), etc.) which do not occur in urban or lowland biotopes (Horváth, Szinetár 1998, 2002). iv) Occurrence frequency may characteristically change within a genus with geographical latitude. The species Segestria bavarica C. L. Koch, 1843 is primarily Southern European, while Segestria senoculata is more widespread in Central and Northern Europe; the former species occurring northwards only in cities with warmer climate. In Southern Europe the third species of the genus, Segestria florentina (Rossi, 1790) may also appear under the bark (Hansen 1992). v) Summer and winter assemblages show significant differences even in the bark of a single tree species. In winter the foliage-dwelling species migrate in a large number to the bark for wintering. On the other hand, in summer, only the real bark-dwelling and the accidental species can be found on the trunk, resulting characteristic differences (Horváth, Szinetár 2002, Horváth et al. 2001). As a summary, it can be stated that the majority of real bark-dwelling species are not asso- ciated to special tree species nor to a given bark structure (Philodromus margaritatus, Clubiona leucaspis, Segestria senoculata, Marpissa muscosa (Clerck, 1757)). On the other hand, facultative bark-dwelling species show significant differences; the tree species with similar structure (within a given geographical latitude) can be characterised by similar spider assemblages (for example black pine↔scots pine) (Appendix 1). Despite this, with the change of geographical latitude a given tree species exhibits significant differences.

Acknowledgements: The authors wish to express their gratitude to numerous arachnologists colleagues for providing their publications at disposal and for their advices. Special thanks are due to Seppo Koponen, Francesca di Franco, Ingmar Weiss and Jörg Wunderlich. We also wish to thank the Organizing Commit- tee of XXII ECA for supporting our participation in the Colloquium. The participation of Cs.Sz. has been supported by the BDF Scientific Board as well as by the Scientific Board of Vas County General Assembly. Csaba Szinetár was supported by the Bolyai Fellowship of the Hungarian Academy of Sciences.

Received: 09.12.2005 Accepted: 15.04.2006 References

Albert R. 1976. Zusammensetzung und Vertikalverteilung der Spinnenfauna in Buchenwäldern des Solling. - Faunistisch-ökologische Mitteilungen, 5: 65-80. Bogya S., Cs. Szinetár, V. Markó 1999. Species composition of spider (Araneae) assemblages in apple and pear orchards in the Carpathian Basin. - Acta Phytopathologica et Entomologica Hungarica, 34 (1-2): 99-121. Buchar J., V. Ruzicka 2002. Catalogue of spiders of the Czech Republic. Praha, Peres Publishers, 349 p. Benfatto D., F. di Franco, V. Vacante 1992. Spiders of Italian citrus groves. - International Citrus Congress, Acireale, Italy, International Society of Citriculture, 3: 938-942. Di Franco F. 1993. Nuovi dati su Clubiona leucaspis Simon, 1932 (Araneae, Clubionidae). - Animalia, 20 (1/3): 71-75. Dobroruka L. J. 2001. Pseudicius epiblemoides (Araneae: Salticidae) in Central Europe. - Arachnologische Mitteilungen, 22: 50-54. Heimer S., W. Nentwig 1991. Spinnen Mitteleuropas. Berlin und Hamburg, Verlag Paul Parey, 543 p.

226 Cs. Szinetár & R. Horváth: Spiders on tree trunks

Hansen H. 1992. Über die Arachniden-Fauna von urbanen Lebensräumen in Venedig - II. Die Rinde-Be- wohnenden Arten des Stammbereiches von Platanus hybrida. - Bollettino del Museo civico di Storia naturale di Venezia, 41: 91-108. Horváth R., Cs. Szinetár 1998. Study of the bark-dwelling spiders (Araneae) on black pine (Pinus nigra) I. - Miscellanea zoologica hungarica, 12: 77-83. Horváth R., T. Magura, Cs. Szinetár 2001. Effects of immission load on spiders living on black pine. - Biodiversity and Conservation, 10: 1531-1542. Horváth R., Cs. Szinetár 2002. Ecofaunistical study of bark-dwelling spiders (Araneae) on black pine (Pinus nigra) in urban and forest habitats. - Acta Biologica Debrecina, 24: 87-101. Horváth R., Sz. Lengyel, Cs. Szinetár, Sz. Honti 2004. The effect of exposition time and temperature on spiders (Araneae) overwintering on the bark of black pine (Pinus nigra). - In: Samu F., Cs. Szinetár (eds): European Arachnology 2002. Plant Protection Institute and Berzsenyi College, Budapest, 95-102. Horváth R., Sz. Lengyel, Cs. Szinetár, L. Jakab 2005. The effect of prey availability on spider assemblages on European black pine (Pinus nigra) bark: spatial patterns and guild structure. - Canadian Journal of Zoology, 83: 324-335. Jermy T. 1987. Gondolatok a koevulúcióról. [Thoughts on co-evolution]. Budapest, Akadémiai Kiadó, 44 p. Koponen S. 1996. Spiders (Araneae) on trunks and large branches of oak (Quercus robur) in SW Finland. - Revue suisse de Zoologie, vol. hors série: 335-340. Koponen S., V. Rinne, T. Clayhills 1997. Arthropods on oak branches in SW Finland, collected by a new trap type. - Entomologica Fennica, 8: 177-184. Koponen S. 2004. Arthropods from high oak branches – Comparison of two trap types, with a special reference to spiders. - Latvijas Entomologs, 41: 71-75. Koslinska M. 1967. Badania nad fauną zimującą pod korą i w korze jabloni. Część II. Badania nad pajęczakami (Arachnida) [Investigations of fauna overwintering in or under bark of apple trees. Part II. Studies on Arachnida]. - Polskie Pismo Entomologiczne, 37: 587-602. Kubcová L., J. Schlaghamerský 2002. Zur Spinnenfauna der Stammregion stehenden Totholzes in süd- mährischen Auenwäldern. - Arachnologische Mitteilungen, 24: 35-61. Kovács K. 2002. Kéreglakó pókok (Araneae) vizsgálata három fafajon [Study of the bark-dwelling spiders on different trees in West Hungary]. Szakdolgozat (M.Sc. thesis), Department of Zoology, Berzsenyi Dániel College, Szombathely, 34 p. Lawton J. H. 1978. Host-plant influences on insect diversity: the effects of space and time. - In: Mound L. A., N. Waloff (eds): Diversity of Insect Faunas. Symposia of the Royal Entomological Society of London, 9: 105-125. Nicolai V. 1986. The bark of trees: thermal properties, microclimate and fauna. - Oecologia, 69: 148-160. Loksa I. 1969. Pókok I. - Araneae I. - Fauna Hungariae, 97. Budapest, Akadémiai Kiadó, 133 p. Loksa I. 1972. Pókok II. - Araneae II. - Fauna Hungariae, 109. Budapest, Akadémiai Kiadó, 112 p. Malten A. 1994. Fünf für Deutschland neue Spinnenarten - Lepthyphantes midas, Neriene furtiva, Hahnia petrobia, Clubiona leucaspis, Diaea pictilis (Araneae: Linyphiidae, Hahnidae, Clubionidae, Thomis- idae). - Arachnologische Mitteilungen, 8: 58-62. Nentwig W., A. Hänggi, C. Kropf, T. Blick 2003. Central European Spiders - Determination Key. http://www. araneae.unibe.ch/index.html version 8.01.2003. Pekár S. 1999. Some observations on overwintering of spiders (Araneae) in two contrasting orchards in the Czech Republik. - Agriculture, Ecosystems and Environment, 73: 205-210. Platnick N. I. 2005. The World Spider Catalog, Version 6.0. http://research.amnh.org/entomology/spiders/ catalog/INTRO3.html Rinne V., T. Clayhills, S. Koponen 1998. Lounais-Suomen suojeltujen tammilehtojen selkärangattomat eläimet [Invertebrates of protected oak groves in southwestern Finland]. - Metsähallituksen luonnon- suojelujulkaisuja, Sarja A, 90: 1-76. Roberts M. J. 1995. Spiders of Britain and Northern Europe. London, etc. Harper Collins Publishers, 383 p. Sebestyén R. 1996. A közönséges platán (Platanus hybrida) kérgén élõ pókok vizsgálata [Study of the bark- dwelling spiders on London Planetree (Platanus hybrida)]. Szakdolgozat (M.Sc. thesis), Department of Zoology, Berzsenyi Dániel College, Szombathely, 48 p.

227 European Arachnology 2005

Simon E. 1932. Les Arachnides de France. Synopsis géneral et Catalogue des espéces francaises de l’ordre des Araneae, 6 (4): 928-929. Simon U. 1995. Untersuchung der Stratozönosen von Spinnen und Weberknechten (Arachnoidea: Araneae, Opilionida) an der Waldkiefer (Pinus sylvestris). Berlin, Wissenschaft und Technik Verlag, 142 p. Southwood T. R. E. 1978. The components of diversity. - In: Mound L. A., N. Waloff (eds): Diversity of Insect Faunas. Symposia of the Royal Entomological Society of London, 9: 19-40. Sterghiu C. 1985. Arachnida: Fam. Clubionidae. - Fauna Republicii Socialiste Romania. Bucuresti, 5 (4): p. 56. Strong D. R. 1979. Biogeographical dynamics of plant parasites and growth form of their hosts. - The American Naturalist, 114: 1-22. Weiss I. 1995. Spinnen und Weberknechte auf Baumstämmen im Nationalpark Bayerischer Wald. Pro- ceedings of the 15th European Colloquium of Arachnology. Czech Academy of Sciences, Institute of Entomology, České Budějovice, 184-192. Wunderlich J. 1982. Mitteleuropäische Spinnen (Araneae) der Baumrinde. - Zeitschrift für angewandte Entomologie, 94: 9-21.

Обзор върху паяците (Araneae), обитаващи кората на дърветата в Европа

К. Синетар, Р. Хорват

(Резюме) Направен е преглед на информацията за паяците, обитаващи кората на дърветата в Европа. До момента са установени 298 вида под кората на различни видове дървета (Pinus nigra, Picea abies, Fagus sylvatica и др.). Високият брой на видовете се определя от случайно срещащи се, временно пребиваващи и постоянно живеещи под кората видове. По отношение на живеещите под кората паяци могат да се очертаят няколко зависимости: 1) при подкоровите популации доминират широко разпространените паяци, независимо от дървесния вид и местообитание; 2) в случаите на единични дървесни видове, значителни разлики във видовия състав на подкоровите съобщества могат да се наблюдават в по-малки райони, като например територията на един град; 3) надморската височина играе важна роля при формирането на съобществата на подкоровите паяци; 4) честотата на срещане се изменя в зависимост от географската ширина; 5) видовият състав на подкоровите паяци се изменя сезонно, дори и в единични дървесни видове. Значителни разлики са установени между летните и зимните сборове.

228 Cs. Szinetár & R. Horváth: Spiders on tree trunks - - – Exclu – – London E – European – White Willow; 6 Acer pseudoplata Platanus hybrida Salix alba Carpinus betulus Microhabitat preferences Microhabitat under tree bark, in fissures on rock walls under tree bark, holes in walls, in houses forests, rocks, under tree bark, in fissures on rock walls, under stones under tree among leaf litter, bark forests, under tree bark, stones under stones, tree bark, in houses under bark, in houses under bark, in houses – Field Maple; – Connection with bark: with Connection –

– Scots Pine; 2002 e i ss – English Oak; zin e tár – Silver birch; ek ár 2005). ovác s W 1998, , S 1996; ovác s e b es tyén Acer campestre k latnic 1995; 1998, 2002; P zin e tár orváth . 1998; K , S . 2005 Pinus sylvestris i m on e b es tyén 1995; S 5 . 2005 . 2001, 2004, 2005; Quercus Quercus robur 1998, 2002; P et al Betula pendula

zin e tár et al i m on et al et al References , S 1996; H orváth inn e . 2001, 2005; K 1982; S 1982; S zin e tár et al , S orváth orváth opon e n orváth orváth 2002 1992; S 1992 1992 1992 1992 1995; H – Black Pine; Apple – Tree; und e rlich und e rlich an se n orváth an se n an se n orváth ovác s an se n an se n i m on H H W H S H H H H 1995; K W 1998, 2002; R 2002; H 1999; H K 2002; H 1996; H – European Alder; – European

Pinus nigra Pyrus malus 1995). The tree species investigated: 1995). , , Pinus Pinus Pinus Pinus , , , , 4 Platanus s ob e rt Platanus , , Quercus Quercus Pyrus malus Tree Alnus glutinosa – Accidental species. Мicrohabitat preferences based on the quoted literature and own data (cf. data own and literature quoted the on based preferences Мicrohabitat species. Accidental – R , , A Pinus sylvestris , – Pear Tree; – Norway Spruce; Fagus sylvatica Pinus nigra sylvestris hybrida robur Platanus hybrida Fagus sylvatica Picea abies nigra robur Quercus Platanus hybrida Pinus nigra sylvestris Pinus nigra sylvestris hybrida Platanus hybrida Platanus hybrida 1969, 1971, 3 – Horse chesnut; o ks a L Picea abies Season Pyrus communis all year all year all year all year III-IX. V-XI. II-X. all year 1991, 2 C F1 F1 F1 A F1 F1 F1 F1 e ntwig N , , , , , 1916 – Facultative bark-dwelling species, bark-dwelling Facultative – och , 1882) – European Beech; F , 1838) och e i me r , 1758) K . K copoli – Cherry Wild Tree; H och i m on S Aesculus hyppocastaneum al m a s ( S K (L. D ( C. L , 1790) 1 inna e u s 2002, o ss i L (C. L. ( Species R List of spiders sampled on tree trunks, under and on bark of trees in Europe (nomenclature after (nomenclature Europe in trees of bark on and under trunks, tree on sampled spiders of List (

– Camperdown Elm Fagus sylvatica uzic k a Prunus avium R , – Planetree Maple; uchar Segestriidae Segestria bavarica 1843 S. florentina S. senoculata Dysderidae Dasumia canestrini 1867) Harpactea hombergi 1763) H. rubicunda Oonopidae Oonops domesticus inermis Tapinesthis Appendix 1. Appendix species, bark-dwelling sive nus Hornbeam; Planetree; Ulmus glabra B

229 European Arachnology 2005 6 on vegetation in forests and open habitats on vegetation in various forests and open habitats on lower branches of coniferous trees on lower trunks and branches of trees on rock walls, tree trunks in open habitats and in parks bushes and trees edge, park, on bushes in orchards, and under bark in winter period (X-IV.) on herbs, bushes, trees (mainly in coniferous forests) on tree bark and branches among low vegetation and detritus in dry places, rarely on bark

. , ,

, .

et al

2002 2002 1995; i m on , et al orváth

ubcová e i ss . 2005 ubcová zin e tár 1999; W inn e orváth , S orváth et al 1986; S chlagha me r sk ý

ek ár 1999; H 1992; K , S 1995; 5 icolai orváth 1995; R ek ár chlagha me r sk ý chlagha me r sk ý . 1999; H orváth 2002 2002 1996; H an se n i m on , S , S e i ss 2004 ubcová . 2001, 2004; K et al W

1982 1967; N 1996 1996; P 1996; H et al

2002; H 1998, 2002; P ubcová ubcová opon e n ogya 1986; H e b es tyén 1992 1992; S 1995; 1995 1995; K und e rlich an se n an se n icolai o s lin sk a orváth i m on i m on e b es tyén e b es tyén zin e tár chlagha me r sk ý zin e tár chlagha me r sk ý e b es tyén i m on S N S W H S H H S S 1998; B S S S 2002; K 2001; K S 1995; S S K 1998; K ,

Platanus Platanus Pinus Pinus Pinus , , , , , Pyrus 4 , Quercus Quercus Platanus Platanus , , , Pyrus malus Pyrus Pyrus malus , , , Quercus robur Quercus , Picea abies sylvestris robur Picea abies sylvestris hybrida Platanus hybrida robur Quercus Pinus nigra hybrida robur Quercus Pinus sylvestris robur Quercus - Platanus hybrida Pinus sylvestris Pinus nigra hybrida Pinus nigra sylvestris hybrida communis malus 3 summer summer VII-X. early summer V-VII. V-IX. V-VII. V-VIII. summer summer 2 A F1 A F1 F1 F2 F1 F2 E A , , ) och , och K K , 1834) , 1802) l e rc k , 1834) id e r C , 1875) (syn. (C. L. ( W (C. L. ) , 1789) ( , 1875) 1 alc ke na e r hor e ll W ill e r s T lac k wall Continued. ( ( , 1837) Theridion vittatum V B hor e ll ( ( T ( och K Mimetidae Ero aphana Ero E. furcata Uloboridae Hyptiotes paradoxus 1834) Theridiidae Achaearanea lunata 1757) A. riparia A. simulans Theridion simulans Anelosimus vittatus 1836) (syn. Dipoena melanogaster (C. L. D. torva Crustulina guttata Appendix 1.

230 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 on vegetation, in orchards in sandy areas on herb vegetation and bushes in forest habitats on bushes and trees in various forests on low vegetation in open habitats plants in open and on different forest habitats on vegetation in xerothermic habitats on bushes and trees in light forests and their edges in various forests and open habitats, under stones and detritus in mountain spruce forests, moss

. .

, , 1996; et al

et al

. 1999; opon e n inn e opon e n ubcová opon e n ubcová . 1998; . 2001, et al

opon e n 1998, 2002; 1999 chlagha me r sk y et al et al

, S

ogya 1995; K 1995; K ek ár 5 2002; K . 1997; R 2002 1995; K 1995; K inn e zin e tár 2002 2002 1996; K i m on i m on et al e i ss e i ss , S

ubcová orváth ovác s W W zin e tár . 1998; B . 1999; P . 1999 . 1999 1996 al , S 1996; R

orváth opon e n 1986; S e b es tyén 1992; S et al et al et al

e t 1999; K 1999; H 1995; 1995; 1995

e i ss an se n opon e n orváth icolai ogya inn e ogya ogya e b es tyén i m on ek ár ek ár chlagha me r sk ý i m on chlagha me r sk ý P B S H K B N W B S S R 1996; K H 2002 1998 1996; S 1997; H S P 2004, 2005; K S

, , Picea Pinus , , , , Quercus Quercus Quercus Quercus Pinus Pinus , , , , 4 Pyrus Quercus Quercus Quercus Quercus , , , Quercus robur Quercus Pinus nigra , , Pyrus malus Platanus hybrida Picea abies sylvestris malus abies Pinus sylvestris Platanus hybrida Pyrus communis Pyrus malus robur Pinus nigra robur Pyrus malus sylvestris robur Picea abies Fagus sylvatica Fagus Pyrus malus sylvatica Fagus Picea abies sylvestris robur 3 summer summer summer I-VI. V-VII. V-VI. ? IV-V. all year summer 2 A A F2 F1 F2 A A A A A , , , ) , , 1802) ippa inna e u s H ) L , 1833) ( lac k wall ) , 1879) (syn. ) B , 1875) , 1914 ( lac k wall ahn B , 1757) 1 H ( i m on alc ke na e r ( S ( W hor e ll ( ) Continued. ac ks on Theridion Theridion pallens T J l e rc k ( C ( , 1982 Dipoena tristis Theridion tinctum ks ala O Enoplognatha latimana E. mordax E. mordax E. ovata Keijia tincta (syn. Lasaeola tristis (syn. Neottiura bimaculata 1767) (syn. bimaculatum N. suaveolens Theridion suaveolens Paidiscura pallens 1834) (syn. Robertus lividus 1836) R. scoticus Appendix 1.

231 European Arachnology 2005 6 fields

forests, buildings, under tree bark, in hollow trees, houses under bark, in and around houses, occasionally a field in orchards in and around houses, occasionally a fields in and around houses occasionally a on tree trunks in floodplain forests, on edges, in parks in and near houses, under stones on vegetation in all open habitats, in winter period (I-IV) also on bark in houses and forests, under bark in urban habitats forests, on tree trunks (mainly in coniferous forests)

, ,

e i ss 2002 inn e 2002

zin e tár ogya 1996; 1996; W 1999; , S e i ss ubcová ubcová zin e tár W ek ár 2001, 2004, 1986; . 2001, 2001, , S zin e tár 1995; 1992 1992; 1998; B e b es tyén e b es tyén , S orváth et al. et al.

et al. icolai

i m on 1995; 5 2002; K 2002; K an se n an se n orváth 1998; P 1998, 2002; R et al.

chlagha me r sk ý 2002 2002 orváth i m on , S 1996; S 1996; S 2001 1997; H ovác s orváth ovác s orváth inn e orváth 1982; S 1982; N zin e tár zin e tár 1967; H 1967; H 1996; H 1996 et al. et al. , S , S

opon e n ubcová opon e n 1992 1992; H 1992; S 1999 1999; H . 1998; H und e rlich und e rlich opon e n an se n an se n o s lin sk a orváth orváth o s lin sk a an se n orváth chlagha me r sk ý ek ár e b es tyén e b es tyén chlagha me r sk ý 2004, 2005; K 2004, 2005; K W H H S K K K H 1995; K 1998, 2002; H et al. et al. H 2005; K 1995; K 1998, 2002; R S W P H H S S , , , ,

, Fagus , Platanus Platanus Pyrus Pinus Pinus

, , , , , 4 Platanus Platanus Picea abies , , Quercus Quercus Quercus Pyrus malus , , , , Pinus

, Picea abies nigra sylvestris hybrida robur Quercus Pyrus communis Pyrus malus Platanus hybrida Pinus nigra hybrida Pinus nigra hybrida robur Platanus hybrida Pyrus malus Pinus nigra malus Platanus hybrida Pyrus malus pendula Betula sylvatica Pinus nigra sylvestris hybrida robur 3 all year spring all year autumn V-VII. summer V-VII. early summer V-VII. 2 A F1 F1 A A F2 A F2 E , , 1871 , ) inna e u s , 1870 , 1881 , 1838) O. P.- L 1831 (syn. , 1757) (syn. ( , a m bridg e och och och alc ke na e r C ) 1 K K ahn ( W l e rc k H L.

L. Continued. C ( O. P.- , 1871 C . L K ( a m bridg e Steatoda bipunctata 1758) S. castanea castanea Teutana S. grossa S. triangulosa 1802) Theridion blackwalli familiare T. impressum T. melanurum T. Theridion denticulatum mystaceum T. C Appendix 1.

232 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 on vegetation and rocks on solitary trees and in forests, (mainly on coniferous), in winter period on and under bark on bushes and trees in various habitats, in orchards among leaf litter in different forests among moss und grass in wet habitats among grass and leaf litter in dry and semi-humid forests in various open habitats among leaf litter and detritus in semi-humid forests in wet meadows and various open and forest habitats among vegetation in various semi-humid and humid habitats among vegetation in various open and forest habitats in spruce forests, and various mountain habitats

et al. ,

2002

et al. i m on

1998; 1998; ogya ogya 2002 2002 2002 ubcová orváth et al. et al.

1986; S

chlagha me r sk ý , S inn e inn e 1995; B 1995; B 5 icolai chlagha me r sk ý 1999; K 2002; H e i ss i m on , S ubcová W ek ár 1982; R 1995; R 1999 chlagha me r sk ý chlagha me r sk ý chlagha me r sk ý 1998 1967; N , S , S , S e i ss ubcová 1992; S 1995; et al.

et al. W 1999; K 1995 1995 1995 1995

1999; P e i ss und e rlich an se n o s lin sk a an se n ubcová ubcová ubcová ogya inn e chlagha me r sk ý ek ár i m on i m on i m on B R S K K S W W S 1999; K et al. 2005 2002 K P S 1995; H K H Pinus Pinus Pinus , , , Pyrus Pyrus 4 Platanus Pyrus Pyrus , , , , , Pyrus , Quercus robur Quercus robur Quercus robur Quercus , , , Pyrus malus Pinus nigra sylvestris hybrida communis malus Picea abies sylvestris communis malus robur Quercus Pinus sylvestris robur Quercus robur Quercus Pinus sylvestris Picea abies sylvestris malus Picea abies robur Quercus Pinus sylvestris 3 summer VI-VII. VI-VII. all year summer V-VII. IX-V. all year ? all year all year all year X-IV. 2 A F1 A A A A A A A A A A , ,

, , och , 1873 . K ch e n ke l lac k wall (O. P.- S B (L , 1851) ( , 1851) lac k wall i m on B S ( , 1872 (O. P.- 1 , 1833 och es tring es tring K W Continued. W ahn ( ( , 1863) , 1875) L. , 1851) H es tring a m bridg e a m bridg e W C T. nigrovariegatum nigrovariegatum T. T. pinastri T. varians T. Linyphiidae saltuum Abacoproeces 1872) Agyneta conigera Anguliphantes angulipalpis ( Araeoncus humilis 1841) helveticus Asthenargus 1936 Bathyphantes gracilis 1841) B. nigrinus B. parvulus pabulator Centromerus C Appendix 1.

233 European Arachnology 2005 6 among moss in beech and spruce forests among leaf litter in semi- humid forests among moss and detritus in various open and forest habitats detritus in in moss, leaf litter, forest habitats among detritus in various forests at ground level in various open habitats in very humid mountain habitats among wet leaf litter and in detritus in forests among wet leaf litter and in detritus in forests in leaf litter and detritus forests and in open habitats on tree bark in both decidious and coniferous forests on bushes and trees in various habitats

i m on , 1998; 1996; et al. 2002 2002 ubcová

1986; S chlagha me r sk ý inn e , S icolai opon e n 5 1995; K 1995 2002 e i ss e i ss ubcová 1997; R W W 1982; N 1995; K 1999 chlagha me r sk ý chlagha me r sk ý et al. , S , S

e i ss 1992 et al.

W 1995; 1995 1995 1995; 1995; K 1995 1995 1995 e i ss e i ss e i ss und e rlich an se n ubcová opon e n ubcová ogya i m on chlagha me r sk ý i m on i m on i m on i m on W B S S S H W W K S S S W 1995; 2002 K K , , Fagus Salix , , , , Pinus Pinus , , 4 Quercus Quercus Picea abies , , Ulmus glabra , Picea abies Pyrus malus Picea abies sylvestris robur Pinus sylvestris Pinus sylvestris Platanus hybrida Picea abies Picea abies robur Quercus Picea abies sylvestris Acer pseudoplatanus pendula Betula sylvatica Pinus sylvestris robur Quercus alba Pinus sylvestris robur Quercus 3 ? ? XI-IV. V-VI. ? all year summer all year all year all year autumn summer 2 A A A A A A A A A A E A , , , , 1834) id e r , 1834) , 1841) , 1887) id e r W , 1841) ( und e vall 1894 id e r , 1894 W , S ( a m bridg e ( W , 1869) ( C 1 lac k wall och ulczyńs k i B K lac k wall ( K Continued. , 1833) (O. P.- ( ulczyńs k i B ulczyn sk i ( (L. K K lac k wall B C. silvicola C. similis C. sylvaticus Ceratinella brevis C. major Diplocephalus cristatus ( D. helleri D. latifrons 1863) D. picinus Diplostyla concolor Drapetisca socialis 1833) Entelecara acuminata 1834) Appendix 1.

234 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 on bark on coniferous trees and bushes among moss and grass, on bushes and trees in various habitats, in cultural landscapes wide range of open habitats among grass in wide range of open habitats among moss and detritus in various open and forest habitats among detritus and on vegetation in very humid habitats in mountain spruce forests on herbs and bushes in very humid forests juveniles on tree branches, adults on low vegetation in forests on bushes in forests and their edges , ,

, ubcová ubcová 2002 2002 1998 e b es tyén orváth 1999 , 1999; K et al.

1995; K 5 et al. ek ár

1995; S 1999; H ubcová inn e 2002 2002 2002 e i ss e i ss ogya W et al. W

1982 chlagha me r sk ý chlagha me r sk ý 1998, P 1998 , S , S 1996; R 2002; K ogya 1992; et al. et al. 1995; 1995; B 1995 1995

und e rlich e i ss e i ss e i ss an se n ubcová ubcová opon e n inn e inn e chlagha me r sk ý i m on zin e tár chlagha me r sk ý chlagha me r sk ý W R H K R S W K W K S S 1996; B S S W , Quercus Quercus Pyrus Pinus Platanus , , , , 4 Platanus , Pyrus malus Quercus , , Pinus

, Pyrus communis Picea abies malus Pyrus malus robur Picea abies hybrida robur robur Quercus robur Quercus Picea abies nigra sylvestris hybrida robur Quercus Picea abies robur Quercus Picea abies robur Quercus 3 summer autumn early summer summer all year all year ? autumn - spring summer autumn - spring late autumn and winter early summer 2 E A A A A A A A A A A , , , , , , , id e r W ( , 1833 , 1851) inna e u s lac k wall lac k wall lac k wall a m bridg e , 1834) B ( L B B ( C ( ( und e rlich id e r es tring 1 W ( W lac k wall W ( ( (O. P.- B

Continued. , 1830) und e vall S E. berolinensis E. berolinensis 1969) E. congenera 1879) E. erythropus Erigone atra E. dentipalpis Erigonella hiemalis 1841) Gnathonarium dentatum 1834) Gonatium rubellum 1841) Gongylidium rufipes 1758) Helophora insignis 1841) Hylyphantes graminicola ( Appendix 1.

235 European Arachnology 2005 6 on various vegetation in wetlands, on bushes and trees, on bark among stones and detritus on bark of trees in semi-humid habitats among roots at foot of tree trunks, on rock walls among stones and in cellars on and under tree bark, in hollow trees in forest habitats on vegetation in forest edges and in forests on tree branches on tree trunks in all grassland types

e i ss

i m on , opon e n e i ss , W 2002; 2002 1998; 1996; W ovác s et al. et al.

et al. ubcová 2002 1998; 1998 e b es tyén ubcová 1986; 1986; S ogya 1996; K 1995; opon e n e b es tyén 1998; K inn e et al. et al.

icolai icolai i m on 1995; 5 opon e n 2002; K 1995; S 1998; K et al. chlagha me r sk ý chlagha me r sk ý 1998; B

inn e inn e 2002 2002 i m on , S , S e i ss 1996; K 1996; S 1997; R inn e et al. W et al. 1982; N 1982; S 1982; N

chlagha me r sk ý zin e tár 1967; K et al. , S , S 2004 1996 1996; R 1996; R inn e ubcová opon e n opon e n ubcová inn e 1992; S 1995; 1995 1997; R und e rlich e i ss und e rlich und e rlich o s lin sk a opon e n opon e n an se n opon e n orváth opon e n opon e n ubcová chlagha me r sk ý chlagha me r sk ý i m on K W K W K W H W S K 1997; R S 1995; K 2002; K K H 1995; K K 1996; R et al. 1995 S 1999; K K , , , , Quercus Quercus Pinus Pinus Ulmus Pinus Pinus , , , , , , 4 Quercus Quercus Platanus Platanus Platanus , , , , Quercus Quercus Pyrus malus , , , Salix alba

, Pyrus malus robur Picea abies robur Quercus Picea abies glabra robur Quercus Picea abies sylvestris robur Pinus nigra sylvestris hybrida robur Quercus Acer pseudoplatanus Fagus sylvatica Pinus nigra sylvestris hybrida robur Ulmus glabra Picea abies sylvestris hybrida robur Quercus 3 early summer ? all year all year all year VIII-XI. ? VIII. all year 2 F2 F1 A A E A A E A , , , , , ill e r och hl e rt M , 1834) K ) O , 1968 l e rc k ( lac k wall , 1833) C B id e r ( ( ol m (O. P.- W (C. L. H ( 1 lac k wall Continued. B ( , 1863) Agyneta innotabilis a m bridg e Hypomma cornutum 1833) Kratochviliella bicapitata 1938 Labulla thoracica Lepthyphantes leprosus 1865) L. minutus Linyphia triangularis 1757) boreus Macrargus Meioneta innotabilis (syn. Meioneta rurestris 1836) C Appendix 1.

236 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 in ground layer of subalpine forests on vegetation in wetland habitats on vegetation in various open habitats among leaf litter and detritus in forest habitats on and under bark in hollow trees in moss and lichens coniferous forests in various habitats, under stones on rock steppes on tree trunks, bark on rock walls and tree trunks among leaf litter and detritus in forest habitats, under bark of old trees on shrubs and dry tree branches in forests

1995; 2002; 1998; e b es tyén i m on 1982; orváth ovác s et al.

chlagha me r sk ý chlagha me r sk ý , S , S , inn e 1992; S 1996; S 1997; H 1995 5 und e rlich chlagha me r sk ý W e i ss an se n ubcová ubcová et al. , S

opon e n 2001, 2005; K W 1967; et al.

2004 1998, 2002; R opon e n ubcová 1986; H 1992 1992; K 1992; K 1992; 1999 1995 1995 1995 1995 1995; K 1995 e i ss e i ss e i ss e i ss an se n an se n an se n o s lin sk a icolai orváth opon e n an se n i m on i m on zin e tár ek ár W W K H H S S W H P K N 2002 2002 H S 2002; K W H 1996; K , , , , , Fagus Salix , , Pinus Platanus , , 4 Platanus Picea abies , Pyrus malus , , Picea abies Platanus hybrida robur Quercus Picea abies Acer pseudoplatanus pendula Betula sylvatica Pinus nigra sylvestris hybrida robur Quercus alba Platanus hybrida Picea abies hybrida Pinus sylvestris Pinus sylvestris Picea abies Platanus hybrida robur Quercus Pyrus communis 3 V-X. IV-VI. spring summer all year ? spring autumn all year III-X. all year VI-VIII. 2 A A A A E A A E A F2 A ) , , , , , 1834) , 1842) illidg e es tring M , 1884) (O. P.- id e r ) , 1830) ( W ) W

und e vall , 1884) (syn. lac k wall ( i m on S B ( S ( 1 ( alc ke na e r i m on S W ( und e vall Continued. ( , 1881) (syn. Entelecara penicillata S , 1871) ( ulczyńs k i a m bridg e K C Micrargus georgescuae georgescuae Micrargus 1975 1851) (syn. Microlinyphia impigra Microlinyphia M. pusilla viaria Microneta 1841) Midia midas Lepthyphantes midas Minyriolus pusillus Mioxena blanda Moebelia penicillata Mughiphantes pulcher ( Lepthyphantes pulcher Neriene clathrata 1830) N. emphana Appendix 1.

237 European Arachnology 2005 6 on bushes, tree trunks in floodplain forests on bushes in semi-humid forests on shrubs and herb vegetation in various open and forest habitats, on coniferous trees at ground level in various open habitats among grass in various medows at ground level in various open and forest habitats among moss and detritus in rocky habitats, in pine forests, sometimes on trunks among detritus in various semi- humid forests in detritus various forests and heatlands on branches in the canopy of coniferous trees and on junipers among grass and leaf litter in various open habitats, sporadically also in forests among leaf litter in humid forests in various open and forest habitats, under bark in semi- humid and forests

2002 2002 2002 chlagha me r sk ý , 1998 , S , 1995 5 et al.

ubcová 2002 e i ss ubcová inn e W 1999 chlagha me r sk ý chlagha me r sk ý chlagha me r sk ý 1998 1998 , S , S , S 1996; K 2002; K 2002; 1992; et al.

et al. et al. 1995 1995 1995 1995; R

e i ss opon e n ubcová an se n ubcová ubcová ovác s inn e inn e ogya chlagha me r sk ý i m on i m on i m on K R R B K S H K S W K S K 2002 S , Platanus Quercus Quercus , , 4 Quercus robur Quercus robur Quercus robur Quercus Pyrus malus robur Quercus Pinus sylvestris Picea abies hybrida robur Quercus Pinus sylvestris Picea abies robur robur Quercus Pinus sylvestris Pinus sylvestris robur Quercus 3 IV-VI. IV-VIII. VI-IX. summer autumn V-X. IV-VIII. winter all year all year early summer ? ? ? 2 A F2 A A A A A A A A A A F1 , (O. , , 1834) (C. L. (O. P.- lac k wall id e r B , 1872) , 1913 ( W , 1851) ( , 1757) a m bridg e och , 1834) , 1884) C 1 K l e rc k ac ks on es tring id e r J , 1871) (L. C Continued. i m on ( , 1841) W W S , 1871) ( (O. P.- ( ( , 1836) a m bridg e C lac k wall och a m bridg e K N. montana N. peltata Obscuriphantes obscurus ( B Oedothorax apicatus 1850) O. retusus Palliduphantes pallidus Pelecopsis elongata mengei P. radicicola P. Pityohyphantes phrygianus microphthalmum Porrhomma P.- montanum P. oblitum P. 1871) C Appendix 1.

238 Cs. Szinetár & R. Horváth: Spiders on tree trunks ) Manica 6 at ground level in spruce forests, under stones in scree slopes at ground level, on bushes and herb vegetation in wetlands and floodplain forests among grass in open habitats, in parks among leaf litter and detritus in forests among grass and moss in peat bogs and in marshy forests among detritus in various mainly semi-humid habitats in the nest of ants ( rubida, Formica fusca among moss and detritus in forests among moss and detritus in forests among leaf litter and detritus in forests at ground level in open and forest habitats among leaf litter and detritus and under stones in forests among grass in various open and semi-open habitats, often in disturbed habitats

, 1998 ubcová et al.

1995; chlagha me r sk ý 1998 inn e , S e i ss 1995; K 5 et al. W

2002 i m on ubcová inn e 1997; R 1982; 1998 1998 et al.

1992 1992; S et al. et al. 1995 1995 1995; K 1995 1995 1995; R

, 1995 e i ss und e rlich e i s e i ss an se n opon e n an se n inn e inn e i m on i m on i m on i m on chlagha me r sk ý S R H W K S W R S S H 2002 S W W , , , Quercus Quercus Quercus , , 4

Pinus sylvestris robur Quercus Platanus hybrida Picea abies robur Quercus Picea abies robur Pinus sylvestris Picea abies robur Quercus Pinus sylvestris robur Quercus Pinus sylvestris Picea abies robur Pinus sylvestris Platanus hybrida robur Quercus 3 ? ? autumn - spring autumn - spring all year all year ? ? autumn - spring all year all year III-X. all year 2 A A A A A A A A A A A A A A , , , , , ) , , ) , 1907 ) , 1834) lac k wall ulczyńs k i a m bridg e udouin B , 1887) ( , 1852) (syn. C , 1834) (syn. hor e ll A ) (K esse rt ( lac k wall T , 1913 ( a m bridg e L B id e r 1 C lac k wall (O. P.- W B ( ( ac ks on ulczyńs k i Continued. lac k wall Lepthyphantes J Savignya frontana K B (O. P.- ( (( ) Lepthyphantes mengei P. pallidum P. pygmaeum P. Prinerigone vagans 1826) Saloca diceros 1871) Savignia frontana 1833 (syn. Silometopus reussi 1871) Syedra myrmicarum 1882) affinis Tapinocyba pallens T. 1872) flavipes Tenuiphantes 1854) (syn. flavipes mengei T. (syn. tenebricola T. Lepthyphantes tenebricola tenuis T. Lepthyphantes tenuis Appendix 1.

239 European Arachnology 2005 6 in various habitats and stratums, on scree slopes, walls and often under bark in forest habitats among grass in various grasslands, in orchards and parks on trees, bushes and herb vegetation in forests and open habitats in leaf litter mixed forests, agriculture habitats in orchards, in arable lands among grass in various xerothermic open habitats among grass and moss, on tree trunks in mountain beech and spruce forests among detritus in various open and semi-open habitats at ground level in various open and forest habitats among leaf litter and detritus in forests among moss and detritus in various forest and open habitats among moss and grass in various forests, often in spruce forests

e i ss 1999; W et al.

2002 1995; 1995 chlagha me r sk ý ogya , S e i ss i m on 5 W 1995 e i ss 1996; B ubcová W 1982; S 1982; 1999 chlagha me r sk ý , S opon e n 1992 1992 et al.

1995 1995 1995; K 1995; 1995 1995 und e rlich und e rlich ubcová an se n an se n ogya i m on i m on i m on i m on i m on i m on H W S S S S H S B S 1995; K 2002 K W , Pinus Pinus , , 4 Pyrus , Quercus robur Quercus , Picea abies sylvestris malus Platanus hybrida Picea abies Pinus sylvestris Pinus sylvestris robur Quercus Pinus sylvestris Pinus sylvestris Platanus hybrida Pinus sylvestris Pyrus malus Picea abies sylvestris 3 all year spring - autumn summer ? all year spring - summer all year ? spring - autumn summer spring - autumn 2 F1 A A A A F1 A A A A A , , , id e r W ( i m on

, 1836) S ( , 1834) , 1834) ch e n ke l , 1836 S och

K id e r (O. P.- , 1834) 1 W ( lac k wal B id e r (C. L. Continued. ( lac k wall , 1833 , 1872) W , 1851) B ( es tring a m bridg e lac k wall W C Thyreostenius parasiticus Thyreostenius ( 1947 Tiso vagans Tiso cristatus Trematocephalus 1834) piscator Trichoncoides 1884) cito Trichopterna nasutus Troxochorus acuminata Walckenaeria B antica W. cucullata W. dysderoides W. obtusa W. Appendix 1.

240 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 in dark, humid spaces, entrances of caves among detritus in various open habitats, often in suburban areas agriculture and grasslands among leaf litter and grass in wetlands and forests on vegetation in semi-humid and humid open forest habitats on herb vegetation in partly shaded wetlands, often in floodplain forests on bushes and solitary trees, not only in semi-humid and humid habitats on vegetation in open and forest habitats on herbs and bushes in various open habitats and on forest edges, in orchards on herbs, bushes, trees in various open and forest habitats on shrubs and in canopies of trees on forest edges, often young conifers ,

, 1996; inn e , ubcová ubcová 2004 2002 2002 opon e n e b es tyén ubcová 1995; K opon e n 5 2002; K 1998, 2002; R 1995; K 1995; S i m on 1998; K 2002 2002 2002 e i ss e i ss ovác s W W et al.

chlagha me r sk ý chlagha me r sk ý zin e tár . 1998; K 1967; S al , S , S , S

inn e e t 1999 1995 1995 1995; 1995; 1995

1998; K e i ss o s lin sk a ubcová ubcová orváth inn e chlagha me r sk ý i m on i m on ek ár i m on chlagha me r sk ý i m on chlagha me r sk ý H W K K R K S S P S S 1996; R S et al. S S , , Quercus Quercus Pinus Pinus , , , 4 Platanus , , Quercus Quercus , Pinus , Picea abies Pinus sylvestris Pyrus malus robur robur Quercus robur Quercus robur Quercus Pinus sylvestris Pinus sylvestris Pyrus communis Pyrus malus Picea abies nigra sylvestris robur Quercus Picea abies sylvestris hybrida robur 3 all year all year autumn - spring VI. V-IX. VI-VIII. V-VI. early summer VIII-IX. V-VII. 2 A A A A A A A A A A , , Atea , , 1757 hor e ll und e vall T l e rc k copoli S C S , 1837 ( , 1830 , 1870 , 1874 och 1 , 1831) (syn. och K K , 1802) i m on Continued. ahn S L.

H und e vall ( C. L. S ) alc ke na e r W Tetragnathidae Metellina merianae 1763) Pachygnatha degeeri 1830 listeri P. dearmata Tetragnatha 1873 montana T. obtusa T. pinicola T. Araneidae Aculepeira ceropegia ( Araneus diadematus A. sturmi sturmi Appendix 1.

241 European Arachnology 2005 6 on trees in deciduous forests and on forest edges on trees and bushes in forest and their edges on bushes and trees in all forest and open habitats on bushes and trees forest edges, often on young conifers on bushes and trees in forests, forest edges on herb vegetation in wet meadows, sporadic in semi- humid and forests on vegetation in steppes, in xerothermic forest margins, winter period rarely on bark on bushes and trees xerothermic slopes and in orchards in the canopy of coniferous trees on vegetation in open habitats on buildings and trees near water on shrubs and trees forest edges, near ponds

, orváth opon e n 2005 1999; ubcová 2002 2002 et al.

ek ár 1995; K 5 1999; K orváth 1998, 2002; H i m on 1998; P 2002 ek ár et al.

1999 chlagha me r sk ý chlagha me r sk ý zin e tár 1967; S 1967 1996; H 1996 , S , S , S inn e et al.

1995 1995; P 1995 1995 1995 2004 e i ss e i ss e i ss ubcová o s lin sk a ubcová o s lin sk a orváth ogya i m on i m on chlagha me r sk ý e b es tyén e b es tyén K K W K S W K H S W B S S et al. S 1996; R , , , , Quercus Quercus Quercus Platanus , , , 4 Quercus robur Quercus Picea abies Pinus sylvestris Pyrus communis Pyrus malus robur Pinus sylvestris Picea abies Pyrus malus Pinus nigra Pinus sylvestris Pyrus communis Pyrus malus robur Picea abies Pyrus malus Pinus nigra hybrida Platanus hybrida 3 V-VI. VI-VIII. V-VIII. V-VII. IV-VI. IV-VII. IV-VI. V-VI. IV-VI. V-VI. VII-X. all year 2 A A A A A A A A A A A A , , , 1869) , 1802) , 1802) und e vall , 1775) ) , 1772) hor e ll , 1757) S och T ( , 1757) , 1870) ( K , 1847) alla s l e rc k 1 (L. P l e rc k abriciu s ( e ntz C ( alc ke na e r C F alc ke na e r H hor e ll ( ( , 1802) ( Continued. W T W ( ( ( Araneus triguttata alc ke na e r W A. triguttatus (syn. Araniella alpica A. cucurbitina A. displicata Cyclosa conica C. oculata Gibbaranea bituberculata ( G. gibbosa G. omodea Hypsosinga pygmaea 1831) Larinioides ixobolus 1873) L. patagiatus Appendix 1.

242 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 on vegetation in all open habitats and in forest edges on conifers peat bogs forests, buildings, under tree bark, on wooden constructions on shrubs and lower branches of trees in subalpine forests on tree trunks and rocky walls on bushes and lower braches in light forests on shrubs and solitary trees, various natural and artificial vertical surfaces on wall of houses, tree trunks and on rocky walls on fallen tree trunks, in forests clearings in the mountains in open habitats, early stages of succession in mountain spruce forests and meadows on ground layer in various forests and their edges

i m on orváth zin e tár 1996; 2002; opon e n , S 1997; 1998 1992; S ovác s et al. 1999

. 1998; H et al. e b es tyén orváth

1996; K an se n 1998 5 et al.

2002 et al. inn e

opon e n et al.

1996; S opon e n ogya 1997; H inn e 1982; H zin e tár inn e 1996; R et al. , S 2004 1996; K opon e n 1992 1995 1995; K 1995; B 1995 1995 1995 1995 1995 2001, 2004, 2005; K 1997; R e i ss und e rlich e i ss e i ss e i ss e i ss orváth opon e n opon e n opon e n an se n e b es tyén i m on i m on i m on H W H S W W K S S W W W K K et al. al. 1998, 2002; R 1995; K S , , , Pinus , 4 Platanus , Quercus Quercus , Pinus , Pinus nigra Picea abies nigra sylvestris hybrida robur Platanus hybrida Pinus sylvestris Pyrus malus Picea abies Picea abies Platanus hybrida robur Quercus Pinus sylvestris Pinus sylvestris robur Quercus Picea abies Picea abies Picea abies 3 V-VII. IV-VIII. all year females VI-VIII. males summer V-VI. IV-V. summer summer- autumn summer III-IX. VII-IX. III-IX. 2 A E F1 A F1 A A F2 F2 A A A , , , , och , ) K och ) , 1861) , 1845) K l e rc k , 1802) , 1802) C ( och hor e ll alc ke na e r (C. L. (C. , 1870) , 1757) K T ( (C. L. es tring W , 1757) ( och W ( K 1 l e rc k alc ke na e r (C. L. C alc ke na e r l e rc k (L. W ( Continued. C ( W Zygiella montana Zygiella stroemi ( ( Mangora acalypha 1802) Nuctenea silvicutrix 1835) N. umbratica montana Parazygiella 1839) (syn. stroemi Stroemiellus 1872) (syn. Zilla dioida Zygiella atrica Z. x-notata Lycosidae Acantholycosa lignaria 1758) agrestis Pardosa ferruginea P. lugubris P. Appendix 1.

243 European Arachnology 2005 6 in underlogged meadows and on marshy pond margins on wet meadows and forest clearings in humid habitats, often floodplain and in marshy forests in ground and on herb layer various open and partly shaded habitats on bushes in various habitats among herb vegetation on various forest edges among stones, roots, in hollow trees mainly in ground layer various dry and open habitats among branches of bushes in forests bark, tree under trees, hollow in in buildings in houses, dry open habitats, on tree trunks in houses, dry open habitats on rock walls, in houses, dry open habitats under stones in scree slopes semi-humid and humid forests

1998, zin e tár , S 2002 2002 2002 , 1999 1999 zin e tár , S orváth 2001 1995 5 ubcová et al. et al.

2002 i m on et al.

orváth ogya ogya 2002 1999; H chlagha me r sk ý chlagha me r sk ý chlagha me r sk ý 1998; K 1996 , S , S , S ovác s orváth 1992 1992 1992; S et al.

et al. 1995; B 1995; B 1995; H 1995 1995

e i ss e i ss ubcová ubcová an se n an se n an se n ubcová inn e ogya chlagha me r sk ý i m on e b es tyén i m on i m on R W S S H H K K S W S K 2002; K 2002; H S H B

, , , Pyrus Pinus , , 4 Quercus robur Quercus , Quercus robur Quercus Picea abies Pinus sylvestris Pyrus malus Pinus nigra malus Pinus nigra sylvestris Platanus hybrida Platanus hybrida robur Quercus Quercus robur Quercus Platanus hybrida Picea abies Pinus sylvestris Pyrus malus robur Quercus Pinus sylvestris Platanus hybrida 3 IV-IX. V-VII. V-IX. VII-VIII. summer VII-X. IV-XI. summer- autumn V-IX. II-XI. ? ? all year II-X. 2 A A A A A A A A A F2 A A A A , , , , 1872 och , 1757) och K , 1834 K , 1757) , 1785) alc ke na e r , 1833) , 1870) , 1840 , 1804) och hor e ll l e rc k , 1872 W T ( (C. L. C K , 1916

C. L. och och ( l e rc k och 1 K och C K K anz e r ( K ourcroy P i m on C. L. F ( S Continued. (L. ( L. C. L. (C. L. P. prativaga P. P. riparia P. Pirata hygrophilus Pisauridae Pisaura mirabilis Agelenidae Agelena gracilens 1841 A. labyrinthica Histopona torpida 1834) agrestis Tegenaria 1802) campestris T. ferruginea T. nemorosa T. pagana T. parietina T. silvestris T. Appendix 1.

244 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 under stones in scree slopes and under tree bark on forest edges under stones, in wet decaying trunks and tree stumps in shaded forests under tree bark, among detritus and under stones in mountain forests under tree bark among moss, leaf litter and detritus in forest habitats under tree bark and stones among grass, leaf litter and under stones in open habitats in leaf litter and under stones in forests and open habitats with high humidity on vegetation in various open habitats ,

1996 ubcová et al.

1998 e b es tyén 1995 5 et al.

1998; K opon e n 2002 i m on inn e et al. 1982; S 1982

1967 1996; K inn e 1992 1992; S 1995 1995 1995; R e i ss e i ss und e rlich und e rlich opon e n an se n o s lin sk a an se n i m on chlagha me r sk ý S W W W K H W K H S 1997; R , , , , Quercus Quercus , 4 Pinus sylvestris Picea abies Picea abies robur Acer campestre Aesculus hyppocastaneum Platanus hybrida Pyrus communis robur Quercus Platanus hybrida Platanus hybrida Pyrus malus Pinus sylvestris Platanus hybrida 3 summer VIII. autumn - spring III-VI. III-XII. autumn summer -autumn spring - autumn V-VI. 2 F1 E A F1 A A F1 A A , , , och , 1789) K och , 1793) , 1897 K

inna e u s L L. livi e r ( , 1841 (C. L. O ( (O.P.- och abriciu s (O. P.- 1 K F ulczyńs k i ( K Continued. , 1863) , 1861) C. L. a m bridg e a m bridg e C Textrix denticulata Textrix Cybaeidae Cybaeus angustiarum 1868 Hahnidae Cryphoeca silvicola 1834) Hahnia picta H. pusilla maerens Tuberta Dictynidae subnigra Argenna Cicurina cicur Dictyna arundinacea 1758) C Appendix 1.

245 European Arachnology 2005 6 on various sun-exposed vertical surfaces, on walls of houses, tree trunks in towns on bushes in humid mountain habitats, in peat bogs on tree branches in various forests and their edges on tree branches forest edges, in orchards and gardens on trees (mainly coniferous) in forests, and parks winter period on and under bark in the nest of ants on xerothermic slopes and forest under bark margins, on foliage mainly in oak forests, on shrubs forest edges on bushes on foliage of trees and bushes, on walls of houses, in winter period on and under bark under stones ,

ek ár , ubcová orváth e b es tyén zin e tár ubcová 1996; P 2001 1992; , S 2002; K 1995; S et al.

an se n 5 2002; K orváth 2002 1998, 2002; H i m on e b es tyén zin e tár 2002 2002 , S orváth 2002 ovác s 1982; H 1982 zin e tár zin e tár 1967; S 1967; S 1996; H , S , S ovác s orváth 1992 1995 1995 1995 2001; K und e rlich und e rlich orváth o s lin sk a o s lin sk a orváth an se n i m on chlagha me r sk ý chlagha me r sk ý i m on i m on e b es tyén 1996; K 1999; H K H S W W S S S H H S S 1998, 2002; H et al. K

Platanus Pinus Pinus , , , , Pyrus 4 Platanus Platanus , , , Pyrus malus Pyrus Quercus , , , Quercus robur Quercus , Pinus nigra Pinus sylvestris Picea abies sylvestris hybrida Pinus nigra hybrida communis malus Pinus nigra sylvestris hybrida robur robur Quercus Pinus sylvestris Pinus sylvestris Pinus nigra Platanus hybrida Platanus hybrida 3 VI-XI. early summer spring - summer V-VIII. IV-VI. all year V. IV-VIII. VI-X. autumn - winter 2 A A A A F1 F2 A A F2 A , , , , 1855) , 1951) e y se rling hor e ll alc ke na e r , 1856 T K ( ( , 1856 W o e w r ( lac k wall 1 , 1870) , 1850) R , 1869 B ( ( hor e ll Continued. T hor e ll i m on uca s e ng

S T L ( M ( D. civica D. major D. pusilla D. uncinata Lathys humilis Mastigusa arietina 1871) Nigma flavescens 1830) N. puella N. walckenaeri Amaurobidae erberi Amaurobius 1863) Appendix 1.

246 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 under tree bark, in hollow trees, hollow in bark, tree under in fissures on rock walls under stones in forest steppes, rock steppes and their edges, on tree trunk under stones in spruce, beech and scree forests at gruond level in forests at gruond level in forests under stones on rock steppes, on sand dunes on trees, and in buildings and in various xerotherm open habitats, orchards, in winter under bark on herb vegetation rock steppes and other xerothermic habitats on leaves and branches of trees in deciduous and mixed forests, winter on and under bark

, ,

2005 ovác s et al.

et al.

1999; orváth 2002; orváth 1998 et al. et

e b es tyén inn e 1982; et al.

et al. orváth

1998, 2002; 1995; H 1996; H orváth ogya inn e H 1996; S e i ss 1997; R 5 und e rlich W 2002; H chlagha me r sk ý zin e tár W e b es tyén et al. , S , S

opon e n 2001, 2004, 2005; K 2001 1997; R 1982; 1999 zin e tár 1967; et al. et al. et al. , S 2004 1998, 2002; 1998, 1998, 2002; B opon e n orváth ubcová 1992; S et al.

1995; K 1995 1995 1995 und e rlich e i ss e i ss e i ss orváth an se n orváth opon e n o s lin sk a orváth opon e n ogya zin e tár zin e tár i m on 1998; H W H W W W B K H H S K S 1996; K 2005 K 2002; K H S

, , , Picea , , , Pyrus , Platanus Pinus , , 4 Pyrus malus , Quercus robur Quercus Pinus nigra , , Picea abies nigra Pinus nigra Picea abies Picea abies Picea abies Pyrus malus Pinus nigra hybrida robur Quercus Alnus glutinosa Carpinus betulus sylvatica Fagus abies Pinus sylvestris Platanus hybrida Prunus avium malus 3 IV-XI. VI-IV. autumn - spring autumn - spring all year VI-XI. V-VI. V. V-VIII. 2 F1 A A A A A F1 A F1 , , , ) och 1872 och K , 1834) ahn , 1871 K H och L. ( id e r K (L. , 1768) , 1868 W L. ( hor e ll T 1 och trö m K S , 1802) ( Continued. Coelotes inermis L. alc ke na e r W A. fenestralis A. jugorum Callobius claustrarius 1833) Coelotes terrestris inermis Eurocoelotes 1855) (syn. Titanoecidae schineri Titanoeca Miturgidae Cheiracanthium mildei 1864 C. oncognathum Anyphaenidae Anyphaena accentuata ( Appendix 1.

247 European Arachnology 2005 6 on trees in detritus various forests, at gruond level, on shrubs and tree trunks on sun-exposed scree slopes, in buildings, on vertical surfaces, under bark on sun-exposed scree slopes, under bark on tree bark, bushes in sun- exposed forests and edges on bushes in semi-humid forests on trees (mainly in deciduous forests), among moss, under stones, under bark in winter period on bark under tree bark, on peat bogs and in semi-humid humid mountain forests

, ,

, orváth ubcová orváth 1998, zin e tár opon e n e b es tyén ubcová , S 1998; H chlagha me r sk ý zin e tár 2002; K 1998; H , S , S 1996; K orváth 2005 1995; S 5 2002; K et al. zin e tár

ovác s 2002 2002 i m on et al. , S ubcová opon e n orváth ovác s inn e 1982; H 1982 2002; K orváth orváth 1992 1992; S 1995; K 1995; K 1995; H 1995 2001; K 1997; R und e rlich und e rlich e i ss an se n an se n i m on chlagha me r sk ý i m on zin e tár chlagha me r sk ý i m on H S W W H W S S S S 2002; H 2002 1998 et al. et al. S 1996; H , , , Pinus Pinus , , , Quercus Quercus 4 Platanus , , Quercus Quercus , Pinus , Platanus hybrida Pinus sylvestris robur Quercus Pinus nigra - Fagus sylvatica Pinus nigra sylvestris hybrida robur Picea abies Carpinus betulus Picea abies nigra sylvestris robur Pinus nigra Pinus sylvestris 3 ? all year III-XII. VI-IX. IV-VII. V-IX. IV-VII. IV-IX. 2 A F2 F2 F2 F1 A F2 E1 , , , , 1802) , 1867 alc ke na e r W och ( lac k wall lac k wall , 1875) B B K , 1866

, 1839 ( 1 L. alc ke na e r och och W hor e ll ( K Continued. K T ( L. C. A. sabina Liocranidae brunnea Agroeca 1833) Liocranum rupicola 1830) L. rutilans Clubionidae Clubiona brevipes 1841 C. caerulescens C. comta C. corticalis Appendix 1.

248 Cs. Szinetár & R. Horváth: Spiders on tree trunks is a 6 C. genevenis Notes: ground-dwelling species of dry sandy lands. See details in text. forests, edges, under tree bark, on trunks (mainly on coniferous) in winter period on bark on tree bark and branches in deciduous forests and on solitary trees on herb vegetation and bushes in various open and forest habitats on tree trunks (mainly in deciduous forests) often in nests, in winter period under bark at ground level and on herb vegetation in wetlands and forest edges on herb vegetation in wetlands

, 2002; 1999; ovác s et al.

orváth 2002 et al.

inn e , 2001, 2004, 1996; 1994; K ogya 2004 ovác s et al.

2005 1995; H 5 ubcová e n alt 1998; R chlagha me r sk ý 2002 e b es tyén opon e n , S i m on et al.

2001; K orváth 1999 zin e tár 1998; K 1993; M et al. , S 1996; S 1998, 2002; B ubcová orváth 2002; K 1992; S et al.

et al. 1995 1995

ranco F e i ss an se n orváth i opon e n orváth ovác s ogya inn e zin e tár chlagha me r sk ý i m on H D B R K W S K S 2005; K 2002; H H H 1998, 2002; H S , , , , Picea , , , , , , Pinus Pinus , , 4 Platanus Quercus , , Pyrus , Prunus avium Pinus nigra , , Alnus glutinosa Fagus sylvatica Pinus nigra sylvestris hybrida malus robur Quercus Pinus nigra sylvestris robur Pyrus malus robur Quercus Alnus glutinosa Carpinus betulus sylvatica Fagus abies Pinus sylvestris Platanus hybrida Prunus avium robur Quercus Picea abies Pinus sylvestris 3 IV-VI. III-VII. ? V-VIII. IV-IX. female all male year, IV-VIII. V-X. 2 A E F1 F1 A A

, 1863 , 1897 , 1866 , 1866 , 1851 , 1757) C. leucaspis och och , 1932 K a m bridg e (1992) and K 1 C ulczyńs k i L. l e rc k es tring L. i m on K C S Continued. W an se n ( O. P.- H (1995) – according to self- , 1932. i m on i m on S S C. genevensis* In case of revision all data are C. leucaspis C. marmorata C. lutescens C. pallidula C. reclusa C. stagnatilis Appendix 1.

249 European Arachnology 2005 6 under tree bark and among detritus in forest habitats among leaf litter and detritus, under tree bark in various forest habitats on bushes in wide range of open habitats on and under bark among grass and detritus in various open and forest habitats, sometimes on tree trunks ? under stones, on various sun- exposed vertical surfaces in xerothermic habitats at ground level in various open habitats under bark of fallen trunks and tree stumps in mountain forests under tree bark on solitary trees and forest edges, in hollow trees ?

, 1995; et al.

e i ss ubcová 2002; W inn e 1998; et al.

1999; 1995; 5 2002 2002; K 1995; R inn e 2002 i m on ek ár e i ss W 1982 1999 1999 chlagha me r sk ý zin e tár zin e tár , S , S , S 1996; R 2004 1992; S 1992; P 1992 1992 et al. et al.

1995; 1995 1995 e i ss und e rlich e i ss an se n orváth an se n orváth an se n opon e n ubcová opon e n an se n ogya ogya i m on chlagha me r sk ý S H W W H H B B W K H K S K H 1998 H , Platanus Pinus Pinus , , , 4 Quercus Quercus Platanus , , Pyrus malus , Pinus , Picea abies sylvestris robur Picea abies nigra sylvestris hybrida Picea abies Pyrus communis Pinus nigra hybrida robur Quercus Platanus hybrida Pyrus malus Pyrus malus Picea abies robur Quercus Platanus hybrida 3 IV-X. IV-X. all year V-IX. V-VIII. IV-X. ? IV-X. IV-XI. IV-X. III-X. ? 2 F1 F2 A E F2 A A A F1 E ? , , , , och

och es tring K , 1964 K och W , 1983 ( K , 1875 , 1843 , 1851 i e hl an es trini (C. L. (L. W (C. L. C och ( rignoli 1 K B hor e ll es tring T , 1802) Continued. W

C. L. alc ke na e r W C. subsultans C. terrestris C. trivialis Corinnidae Cetonana laticeps 1868) festivus Phrurolithus 1835) Zodariidae Zodarion hamatum Gnaphosidae Drassodes lapidosus ( Drassyllus pusillus 1833) Gnaphosa montana 1866) cognatus Haplodrassus 1861) Micaria cyrnea Appendix 1.

250 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 among grass, moss in various open habitats on solitary trees, and under bark in various open and forest habitats under tree bark, in hollow trees in beech, oak and floodplain forests, in houses under tree bark in forests, parks, in buildings and houses ? ? in various open and forest habitats, sometimes on tree trunks on shrubs and trees forest steppes, in oak forests and orchards on shrubs and trees in forest edges and in orchards

, , ,

ovác s 1999; orváth 1998; 1995; 2004 e b es tyén e i ss orváth ubcová et al. ubcová zin e tár ,

et al.

2002 W 2001, 1986; 1992; chlagha me r sk ý , S opon e n 1995; H , S , inn e ogya 1996; S et al.

icolai ubcová an se n 1997; H i m on 1995; 5 2002; K orváth 2002; K 2002 2002 opon e n ubcová i m on et al.

2001, 2005; K 2001, 2004, 2005; K orváth ovác s 1982; N 1982; K 1982; H 1982; S chlagha me r sk ý 1996; H et al. et al. , S

1998, 2002; R 1998, 2002; B opon e n 1992; K 1992 1992 1992; S K 1992; 1999 und e rlich und e rlich und e rlich und e rlich an se n orváth an se n an se n an se n ubcová an se n orváth ek ár zin e tár chlagha me r sk ý chlagha me r sk ý e b es tyén chlagha me r sk ý zin e tár P W W W H H H H S 1998, 2002; H 1996; K 2002 2002 2004, 2005; K K S H S S S H W H S , , , Platanus Pinus Pinus Pinus , , , , Platanus 4 Pyrus Platanus , , , , Pyrus Quercus Quercus , , , Quercus robur Quercus , Pyrus communis Fagus sylvatica Pinus nigra hybrida communis robur Quercus robur Quercus Pinus nigra sylvestris hybrida robur Platanus hybrida Platanus hybrida Picea abies sylvestris hybrida robur Platanus hybrida robur Quercus Fagus sylvatica Pinus nigra sylvestris malus 3 spring - early autumn IV-X. II-X. all year ? ? ? early summer IV-VII. 2 A E F1 F1 ? ? A F1 F1 , , 1833) , 1831) 1861 , 1866) , ulczyńs k i , 1866) K , 1757) och und e vall och K , 1846) 1 K (S es tring und e vall (L. l e rc k W (L. (S

Continued. uca s C ( L ( , 1758) inna e u s L M. pulicaria M. subopaca Scotophaeus quadripunctatus ( S. scutulatus S. validus Zelotes tenuis Zoridae Zora spinimana Philodromidae albidus Philodromus 1911 aureolus P. Appendix 1.

251 European Arachnology 2005 Pinus 6 on shrubs and trees in forest edges and in orchards on shrubs and trees in forests, mainly on coniferous trees on vegetation in forests and forest edges on tree trunks in coniferous forests on tree trunks in pine forests, on and under bark of sylvestris on trees (mainly coniferous), shrubs and herbs in forests, forest edges, parks on tree trunks and branches in forests and on forest edges on herb vegetation and trees and shrubs (mainly on coniferous) in forests, on forest edges, in parks

et al. i m on et al.

ovác s 2002 1999 e b es tyén zin e tár zin e tár orváth 2002 orváth ovác s 1992; S ek ár 2002 , S , S 1995; 1995; K 1999 e i ss an se n i m on 1995; S ovác s 5 et al.

W orváth orváth 1998; H 1998, 2002; 1999; H 1999; P 1999; K i m on ogya 2002 ek ár et al. et al.

1982; H 1982; 1982; S 1995 chlagha me r sk ý zin e tár zin e tár 1996; H 1996; H , S , S , S e i ss ovác s ogya ogya 1992; S W 1995; B und e rlich und e rlich und e rlich an se n orváth orváth ubcová i m on e b es tyén e b es tyén 2001; K H W H S W W H 2001, 2004, 2005; K S S 1998; B 1998, 2002; P 2002 1995; 1996; B K , , , , Picea , , , , , , , Quercus Quercus Pinus Pinus Quercus Pinus Pinus , , , , , , 4 Platanus Quercus Platanus , , , Pyrus malus , Pinus nigra , Pinus sylvestris Platanus hybrida Pyrus communis Pyrus malus Picea abies sylvestris hybrida Pinus nigra sylvestris robur Pinus sylvestris Pyrus malus Picea abies sylvestris Pinus nigra sylvestris hybrida Alnus glutinosa Carpinus betulus sylvatica Fagus abies Pinus sylvestris Platanus hybrida Pyrus communis Pyrus malus robur Pinus nigra robur 3 spring - summer early summer IV-VII. early summer VI-VIII. IV-VII. V-VIII. IV-VII. 2 F1 E F1 F1 F1 F2 E E , , ee r , 1802) , 1803) , 1757) G , 1826 e , 1835 D , 1870 a m bridg e ( C l e rc k och chran k 1 C K S ( i m on ( alc ke na e r S O. P.- W Continued. ( alc ke na e r C. L. W P. cespitum P. collinus P. dispar P. emarginatus P. fuscomarginatus P. 1778) longipalpis P. margaritatus P. praedatus P. 1871 Appendix 1.

252 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 on vegetation in various habitats, edges in orchards, winter period mainly under bark among grass in various open habitats, also in orchards under tree bark in pine forests on leaves of bushes and trees in forest habitats, in winter period on and under bark on branches of solitary trees (mainly oak) and on forest edges, in winter period on and under bark on flowers in medows, other open habitats on herbs and bushes, lower branches of trees in semi-open and partly shaded habitats, often in orchards, in winter period on bark in various forest habitats, at ground level and also on tree trunks

2002 , , ovác s , 2002; 1998, e b es tyén , ovác s 1999; 1999; 2004

ubcová orváth ubcová 1995; K 1999 zin e tár zin e tár et al. et al. et al.

1996; S ek ár ubcová , S , S 2001; K i m on ek ár 5 S 2002; K ogya ogya 1995; H

1998; K 2002 2002 2002 et al. orváth opon e n

e i ss orváth orváth 2002; K W et al. 1982;

1999; P 1999 zin e tár 1996; B 1996; B , S 1998, 2002; P orváth ovác s inn e 2002; H 1992; K et al. et al.

1999; H 1995; H 1995; und e rlich orváth ovác s an se n ogya ogya i m on i m on zin e tár chlagha me r sk ý e b es tyén chlagha me r sk ý e b es tyén ek ár chlagha me r sk ý H P S B W S S B H 1999; K S 2002; H 2002 S 1996; R K S S S , , , , Picea Pinus , , , , Platanus Platanus Pinus , , , Quercus Quercus Pyrus 4 Quercus Quercus Pyrus , , , , Pyrus malus Pyrus , , Quercus robur Quercus Pinus Pinus nigra , , , Carpinus betulus sylvatica Fagus abies Pinus sylvestris robur Quercus Pyrus communis Pyrus malus sylvatica Fagus sylvestris robur Picea abies nigra sylvestris communis robur Pinus nigra hybrida robur Quercus Pyrus malus Pinus nigra hybrida communis malus Platanus hybrida robur Quercus 3 V-VI. early summer autumn V-IX. IV-VI. IV-VI. V-VII. V-VIII. III-XII. 2 F1 F1 F1 A F2 A E F2 , , och , 1777) , 1757) K (C. L. alc ke na e r , 1826 W l e rc k ) (C. L. ( C abriciu s 1 ( F , 1876 ( Continued. i m on alc ke na e r , 1775) S W Diaea pictilis , 1837) abriciu s och F P. rufus P. oblongus Tibellus 1802) Thomisidae Coriarachne depressa Diaea dorsata D. livens (syn. Misumena vatia Misumenops tricuspidatus ( Ozyptila praticola 1837) K Appendix 1.

253 European Arachnology 2005 6 among grass in dry habitats on bushes and lower branches of trees, also on tree trunks forest steppes and edges on vegetation (mainly flowers) in open habitats and on edges on herb vegetation and bushes on forest steppes, light forests and their edges on tree trunk in floodplain forests and in orchards on flowers in medows and other open habitats among grass on xerothermic habitats on and under bark on bushes and herb vegetation in various open and forest habitats among grass in medows and other open habitats, also in fields and in orchards among grass and on vegetation in various open habitats, also fields, orchards and gardens 1996;

, , 2002 opon e n 2002; zin e tár 1999 , S ubcová ovác s ubcová 5 et al.

1995; K 2002 2002 orváth e i ss ogya 2002; K W 1982 1982 1999; K 1999; K 1999 chlagha me r sk ý 1998 1996 , S ovác s 1992 1992; H et al. et al. et al.

et al. 1999 1995; 1995; B

und e rlich und e rlich an se n ubcová an se n ogya ogya ogya inn e chlagha me r sk ý chlagha me r sk ý ek ár i m on i m on e b es tyén R H B H W B P B W S S S 2002; K S K S , , , Picea Picea , , , , Quercus Quercus Pinus , , 4 Quercus Quercus , Pyrus malus Pinus nigra , , Platanus hybrida sylvatica Fagus abies robur Quercus sylvatica Fagus abies Pinus sylvestris Platanus hybrida robur Quercus ? Pyrus malus robur Pyrus communis Pyrus malus Pyrus communis Picea abies sylvestris robur Pinus sylvestris Pyrus malus Platanus hybrida 3 ? summer V-VII. J: autumn, spring early summer ? V-VIII. IV-V. spring - autumn spring - early summer IV-VI. summer 2 F1 A F1 E F2 A A A F2 A A , , , , , 1802) , 1872 , 1772) a m bridg e abriciu s alc ke na e r C F , 1757) ulczyńs k i alla s ( , 1803) hor e ll W P K ( T alc ke na e r , 1872 1

, 1875 W l e rc k (O. P.- ( Continued. ( C chran k i m on S hor e ll ( T S O. sanctuaria 1871) Pistius truncatus Synaema globosum 1775) Tmarus piger stellio T. Thomisus onustus 1805 Xysticus acerbus X. albomaculatus 1891 X. audax X. cristatus X. kochi Appendix 1.

254 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 on shrubs and herb vegetation in forests and their edges, on tree trunks on vegetation in wet meadows and on marshy pond margins on soil surface in xerothermic open habitats on bushes and among leaf litter in forests, also canopies and on bark (mainly oak) on bushes and trees in rock steppes and in orchards on branches in coniferous in forests and their margins, winter period on and under bark among moss, leaf litter and under stones in forest habitats at ground level and on herbs in various open and forest habitats at ground level and on herbs in various open and forest habitats on bushes rock steppes, in pine forests and on xerothermic forest edges

2002; 2004 2002; ubcová opon e n inn e 2005 e b es tyén 2002 ovác s 1999 opon e n zin e tár et al.

1996 , S et al. 1999; K

1997; R 1995; K 1995; S 5 1999; K 2002; K et al. orváth ogya i m on

i m on et al.

orváth e b es tyén et al.

1999 chlagha me r sk ý 1998 ogya , S opon e n ogya 2002; H 1986; S 1992; S 1992; B et al.

et al. 1995; S 1995; H 1995 1995 1995

1998; B e i ss e i ss icolai an se n ubcová ovác s an se n ogya inn e chlagha me r sk ý i m on i m on i m on K N S B W H R W H S S S 1996; K K al. 1996; B , , , , , , , , Quercus Quercus Quercus Quercus Pinus , , , 4 Quercus Quercus , Betula pendula Fagus sylvatica Pinus sylvestris Pyrus malus robur Pinus sylvestris Platanus hybrida Pyrus malus Picea abies Platanus hybrida Pyrus malus robur Quercus Picea abies Carpinus betulus Pinus sylvestris Platanus hybrida Pyrus malus robur Pinus nigra sylvestris robur Pinus sylvestris 3 spring - early summer VI-VII. spring - summer summer summer V-VIII. spring early summer V-VII. spring - summer V-VI. 2 F2 A A F1 A F1 A A A A , , , , ) l e rc k , 1757) und e vall C ( S alc ke na e r ( , 1835 alc ke na e r 1835 W , ( W ( l e rc k och 1 C K och , 1831) K , 1802) Continued. Ballus depressus , 1819) ahn C. L. H C. L. ( alc ke na e r e ill atr L W X. lanio X. ulmi Salticidae Aelurillus v-insignitus 1757) Ballus chalybeius 1802) (syn. xanthogramma Carrhotus ( Dendryphantes rudis 1833) Euophrys frontalis 1802) falcata Evarcha Heliophanus cupreus ( H. dubius Appendix 1.

255 European Arachnology 2005 6 on vegetation various xerothermic open habitats on vegetation various xerothermic open habitats on vegetation various xerothermic open habitats, on meadows and on forest edges ? on branches of trees (mainly coniferous) in xerothermic habitats, in winter period on and under bark under tree bark, on the trunks in various open and partly shaded habitats on bushes, trees, under bark and on surface of rocks in various open and forest habitats ? under bark of solitary trees and in forests orchards, parks, on tree bark on trunks of oak

, , et

2002 1998; orváth et al. 1999; 1999; ogya

orváth orváth 1996; ovác s 2002; et al. et al.

2002 zin e tár orváth 2002; H 1996; H 1996; H , S ogya ogya ovác s e b es tyén 5 ovác s 1998, 2002; B 2002; H orváth e b es tyén e b es tyén 2001, 2005; K 2001; K 1982; S 1999 ovác s chlagha me r sk ý , 2001 zin e tár 1998; K et al. et al. , S , S 1996; H 1998, 2002; B 1998, 2002; B 1992 1992 1992 1992; S 1992 1992; S et al.

et al.

2005 1999; K und e rlich an se n an se n an se n an se n orváth orváth opon e n an se n an se n orváth ubcová obroru k a ogya inn e zin e tár zin e tár B H H H H K H H D H R al. 2004, 2005 et al. K H H W S S , , , , Picea , , , Quercus Quercus Pinus Pinus , , , , Pyrus 4 Platanus Platanus , , , Pyrus Pyrus malus , , Quercus robur Quercus Pinus nigra , , Pyrus malus Platanus hybrida Platanus hybrida Platanus hybrida sylvatica Fagus abies Pinus sylvestris Platanus hybrida Pyrus malus robur Carpinus betulus Fagus sylvatica Pinus nigra sylvestris hybrida communis malus Pinus nigra robur Quercus Platanus hybrida Pinus nigra sylvestris hybrida robur Quercus robur Quercus 3 spring - summer ? V-VI. summer IV-X. summer V-VIII. ? 2 A A A F1 E F1 A E A E , 1757) , 1846) , och l e rc k K , 1867 (syn. C , 1868 ( ) hyz e r 1 , 1832 C ) , 1868 i m on (C. L. , 1802) , 1826) (syn. , 1802) S ahn Continued. , 1891 H lac k wall

i m on B S ( alc ke na e r alc ke na e r alc ke na e r ulczyńs k i W W W K H. flavipes H. kochi H. tribulosus Icius hamatus nidicolens Macaroeris ( Marpissa muscosa Pseudeuophrys erratica ( Euophrys erratica vafra P. Euophrys vafra Pseudicius encarpatus ( epiblemoides P. Appendix 1.

256 Cs. Szinetár & R. Horváth: Spiders on tree trunks 6 on tree bark of solitary trees, rarely on walls of houses ? on rock walls, the surface of scree slopes and on walls houses, on tree trunks ? on the trunks and lower branches of trees in various open and forest habitats among short vegetation and stones on rock steppes xerothermic habitats among grass on sand dunes, spoil heaps on various sun-exposed vertical surfaces, on rock walls, walls of houses, on tree trunks in various habitats, on edges, on rock walls among grass on rock steppes, on bark on vegetation in various open habitats, rarely on bark at ground level, in various open and dry habitats, under stones

1999; i m on , et al.

opon e n et al. opon e n

ubcová 1992; S orváth 1999 ogya . 1998; K 1996; K an se n al ek ár

5 2002; K e t

2002; H 1996; B opon e n inn e 1982; P 1982; H 1982 1982 1999 1999 zin e tár , S e b es tyén 1992 1992 et al. et al.

1995 1995 1995; K 1995 1997; R e i ss und e rlich und e rlich e i ss und e rlich und e rlich an se n an se n orváth ogya ogya chlagha me r sk ý i m on i m on W H W H B W W S B W W S S H et al. 2004 2005 1995; S , Pinus Quercus Quercus , , 4 Platanus , Pyrus malus , Picea abies robur Platanus hybrida Pyrus malus Platanus hybrida Pyrus malus – Pinus nigra sylvestris hybrida robur Quercus Pinus sylvestris Pyrus malus Picea abies Pyrus communis Pinus sylvestris 3 summer ? spring -autumn V-VI. spring - autumn summer all year ? ? spring - autumn ? 2 F1 ? F2 E A A A A F2 F2 A , , ) ) , , och och , 1797) K K , 1837) i m on hl e rt , 1775) , 1846) S ( O anz e r och ( P (C. L. , 1757) K och (C. L. ( , 1846 K , 1836) , 1868) 1 abriciu s F l e rc k uca s ( uca s i m on Continued. (C. L. L C Bianor aurocinctus Euophrys petrensis

( S L (C. L. ( ( Salticus cingulatus S. mutabilis S. scenicus S. unciger S. zebraneus Sibianor aurocinctus 1865) (syn. Sitticus distinguendus 1868) S. pubescens S. saxicola Synageles hilarulus 1846) S. venator petrensis Talavera 1837) (syn. Appendix 1.

257