QL 453.4 i£ Arachnologische Mitteilungen EUROPEAN CONGRESS OF ARACHNOLOGY E Arachnology 2009 0 16.8. -21.8.2009, Alexandroupoli, Greece P www.european-arachnology.org e a n Volume 40 Nuremberg, January 201 1 ISSN 1018-4171 www.AraGes.de/aramit Arachnologische Mitteilungen Schriftleitung: Theo Blick, Forschungsinstitut Senckenberg, Entomologie III, Projekt Hessische Naturwaldreservate, Senckenberganlage 25, D-60325 Frankfurt/M., E-Mail: theo.blick@senckenberg.de, aramit@theoblick.de Dr. Oliver-David Finch, Carl-von-Ossietzky Universität Oldenburg, Fk 5, Institut für Biologie und Umweltwissenschaften, AG Biodiversität und Evolution der Tiere, D-261 11 Oldenburg, E-Mail: oliver.d.finch@uni-oldenburg.de Guest Editor: Dr. Maria Chatzaki, Lecturer, Dpt. of Molecular Biology and Genetics, Democritos University of Thrace, University Campus, 6th km Alexandroupoli-Komotini, Dragana, 68100 Alexandroupoli, Greece. Tel./Fax: +30-25510-30636, E-Mail: mchatzak@mbg.duth.gr or: maria.chatzaki@gmail.com Redaktion: Theo Blick, Frankfurt/M. Dr. Oliver-David Finch, Oldenburg Dr. Jason Dunlop, Berlin Dr. Ambros Hänggi, Basel Dr. Detlev Cordes, Nürnberg (Layout, E-Mail: bud.cordes@t-online.de) Herausgeber: Arachnologische Gesellschaft e.V. URL: http://www.AraGes.de 9^VTBS0 Nt^ APR 29 2015 ubrar\&z. Wissenschaftlicher Beirat: Dr. Elisabeth Bauchhenß, Schweinfurt (D) Dr. Peter Bliss, Halle (D) Prof. Dr. Jan Buchar, Prag (CZ) Prof. Peter J. van Helsdingen, Leiden (NL) Dr. Peter Jäger, Frankfurt/M. (D) Dr. Christian Komposch, Graz (A) Dr. Volker Mahnert, Douvaine (F) Prof. Dr. Jochen Martens, Mainz (D) Dr. Dieter Martin, Waren (D) Dr. Uwe Riecken, Bonn (D) Dr. Peter Sacher, Abbenrode (D) Prof. Dr. Wojciech Star^ga, Warszawa (PL) Erscheinungsweise: Pro Jahr 2 Hefte. Die Hefte sind laufend durchnummeriert und jeweils abgeschlossen paginiert. Der Umfang je Heft beträgt ca. 50 Seiten. 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Rehbinder (AraGes-Logo) Arachnologische Mitteilungen 40: 1-109 Nürnberg, Januar 2011 25™ EUROPEAN CONGRESS OF ARACHNOLOGY 16-21 AUGUST 2009 ALEXANDROUPOLI, GREECE Organistion commitee Maria Chatzaki maria. chatzaki@gmail. com Katerina Spiridopoulou aikspiridopoulou@gmail.com Iasmi Stathi iasmi@nhmc.uoc.gr Secretariat Stavros Gogolopoulos Kyriakos Karakatsanis Eleni Panayiotou Aggeliki Paspati Emma Shaw DEMODRITUS UNIVERSITY OF THRACE, GREECE DPT. OF MOL. BIOLOGY & GENETICS 6th Km Alexandroupoli — Komotini Univ. Campus, Dragana 68 1 00 Alexandroupoli GREECE Scientific commitee Miquel Arnedo Maria Chatzaki Christo Deltshev Victor Fet Peter Jäger Dimitris Kaltsas Aristeidis Parmakelis Iasmi Stathi Co-organiser & financial support European Society of Arachnology Prefectural Self-Government Prefecture of Evros Prefecture of Rhodopi- Evros Region of Macedonia-Trace NATURAL HISTORY MUSEUM OF CRETE UNIVERSITY OF CRETE PO BOX 2208 71409 Irakleio CRETE GREECE 2 3 Preface The 25th European Congress of Arachnology was held in the city of Alexandroupolis, from 16 to 21 August 2009. It was jointly organised by the Department of Molecular Biology and Genetics of the Democritus University of Thrace and the Natural History Museum of the University of Crete. Arachnologists from all over the world arrived to this little point at the north-eastern border of Greece to attend this meeting, the 25th in a series which started in 1972 in Strasburg. The arachnological meeting has passed through many countries of Europe, but this was the first time that has taken place in Greece. 90 participants, including 30 students, from 28 representative countries from all five conti- nents and 17 accompanying persons attended this meeting. The scientific programme com- prised 51 oral presentations and 34 posters from various thematic fields such as taxonomy and faunistics, phylogenetics, biogeography, phylo- geography, physiology, behaviour, ecology and others. 18 papers were submitted for evaluation for the proceedings volume and of these, 1 1 were selected for publication after peer review by two referees each. During the four days of scientific programme, three keynote presentations were given on special topics by: Miquel Arnedo, Victor Fet and Peter Jäger. Student awards were given to the best three oral presentations and the best three posters on the closing ceremony. Lectures: Holger Frick (Switzerland), Samuel Yu-Lung Hsieh (Germany), and Odile Brug- gisser (Switzerland). Posters: Jin-Nan Huang (Taiwan), Vera Opatova (Czech Republic), and Chueh Hou (Taiwan). During the five-day stay at the congress hall of the Alexander Beach Hotel, participants had the opportunity to enjoy the hospitality of the staff, but also to take advantage of the beautiful doi: 10.5431/aramit4001 beach opposite the hotel and the cosy atmos- phere of Greek taverns. Accompanying persons visited the most important archaeological sites, cities and natural reserves of the area. During the mid-week excursion, all of us had a wonder- ful experience canoeing on the Nestos river or walking by its riverbank and then enjoying the food, wine and nice view of a local tavern in the surroundings of the city of Xanthi, where we spent the rest of the day. An opening ceremony at the pool of the hotel, the standard ‘Russian party’, Thursday’s congress dinner, and finally a beach party at the end of the congress completed the picture of a very pleasant week. The General Assembly of the ESA members closed the meeting of 2009. During this cere- mony we had to say our last goodbyes to the late arachnologists Gershom Levy and Jean-Pierre Maelfait who passed away that year. Never- theless, we also had to congratulate the 70th anniversary of the still very active arachnologist Christo Deltshev. Members of the Society had to vote on the fate and form of the proceedings of the congress, which later in the year lead to a general referendum. Proceedings should thus retain their paper format and be distributed to all members of the Society, either edited by an existing journal or - eventually - by the Society’s official future journal. Scientific contributions and exchange of knowledge harmonised with relaxation and fun due to the commitment of many people who volunteered to offer their help: first of all Iasmi Stathi and Katerina Spyridopoulou did their best for the whole year’s organisation pro- cedure; our students Eleni Panayiotou, Aggeliki Paspati, Kyriakos Karakatsanis and Stavros Gogolopoulous, assisted in various ways during the congress week; Pavlos Georgiadis was the main group leader of the accompanying people’s 4 activities and Emma Shaw assisted in the se- cretariat, as well as other activities; without Peter Jäger there would never have been such a successful last day beach party, since he was the night ’s DJ. Many participants also assisted with the smooth running of the week’s programme, among them session chairs, members of the scientific committee and evaluation teams. I am deeply grateful to the staff of the Alexander Beach Hotel for their patience and kindness, all the facilities they provided and the very well organised service. I also owe my gratitude to Theo Blick and the rest of the Arachnolo- gische Mitteilungen personnel for our excellent cooperation during editing of the proceedings and the numerous reviewers who volunteered to check and decide upon the selected papers of this volume. I finally have to acknowledge our sponsors: the local authorities, Prefectural Self-Government Prefecture of Evros; Prefecture of Rhodo- pi-Evros; Region of Macedonia-Thrace; the Democritus University; the European Society of Arachnology. Special thanks to all secretary members of the above offices who offered their support and help at any moment as well as the members of the ESA council not only for financial support, but also for the psychological and consulting support during the whole year of the organisation of this congress. Maria Chatzaki When citing the whole volume, please use: CHATZAKI M., T. Blick & O.-D. Finch (Eds) (2011): European Arachnology 2009. Proceedings of the 25 th European Congress of Arachnology, Alexandroupoli, 16-21 August 2009. - Arachnologische Mitteilungen 40: 1-109 The contributions to this volume are online available at: 1. http://arages.de/aramit 2 . http :// www. european- arachnology. org/ collo/ collo25 .shtml 3. http://www.mbg.duth.gr/en/index.php?cid=84&st=l The abstract volume (pdf) and the congress photo (jpg) are available at: 1. http://www.mbg.duth.gr/en/index.php?cid=84&st=l 2 . http :/ / www. european- arachnology. org/ collo/ collo25 . shtml Arachnologische Mitteilungen 40:5-14 Nuremberg, January 201 1 Abundant and rare spiders on tree trunks in German forests (Arachnida, Araneae) Theo Blick doi:10.5431/aramit4002 Abstract:The spider fauna active on the bark of trees in forests on eight sites in different regions in Germany was investigated. Trunk eclectors at about 2-4 meters height on living trees were used in different regions of Germany (SW Bavaria, Hesse, Brandenburg) between 1 990 and 2003. In Hesse eclectors were also used on dead beech trees (standing and lying). In this study data, mainly from beech ( Fagussylvatica ) and spruce {Picea abies), from May to Octo- ber are compared - whole year samples (including winter) are only available from Hesse. A total of 334 spider species were recorded with these bark traps, i.e. about one third of the spider species known from Germany. On average, each of the eight regions yielded 1 40.5 (± 26.2) species, each single tree 40.5 (± 1 2.2) species and 502 (± 452) adult spiders per season (i.e. May to Oct.). The 20 most abundant species are listed and characterised in detail. Six of the 20 species were not known to be abundant on bark, three prefer conifers and three beech/broadleaf. Even in winter (December-March) there was a remarkably high activity on the trunks. However, only a few spe- cies occur exclusively or mainly in winter. Finally, the rarity of some bark spider species is discussed and details (all known records in Germany, phenology) of four of them are presented {Clubiona leucaspis,Gongylidiellum edentatum, Kratochviliella bicapitata, Oreonetides quadridentatus).Vr\e diversity and importance of the spider fauna on bark in Central Europe is still underestimated. Key words: bark, common species, distribution, eclectors, Germany, rare species The spider fauna of tree trunks in Germany and Cen- tral Europe is much less well known than the epigeal fauna active on the forest floor. Only the fauna of the tree crowns is more poorly known (SIMON 1995, GOSSNER 2004, RÖDER et al. 2010). Furthermore, the epigeal spider fauna of common forest types - at least in Central Europe — is significantly less inten- sively investigated than that of the epigeal fauna of special habitats, such as those that are extremely dry or wet (HÄNGGI et al. 1995). I estimate that this dis- proportion is much higher, if one compares the epigeal spider fauna in forests with the tree trunk spider fauna. In Central European forests knowledge about the tree trunk fauna reaches an estimated maximum level of 5% compared with the epigeal fauna. Before the research in Strict Forest Reserves in Hesse started, estimates of the species richness of ani- mals (biodiversity) in a beech forest were 1500-2000 species (all animals) (ELLENBERG et al. 1986). Now we know that there are closer to 5000-6000 (DOROW et al. 2004, 2010). In each of the four Strict Forest Reserves investigated until now 162 to 202 spider species were recorded (BLICK 2009). Theo BLICK, Senckenberg Forschungsinstitut und Natur- museum, Projekt Hessische Naturwaldreservate, Sencken- berganlage 25,60325 Frankfurt am Main, Germany, E-Mail: theo.blick@senckenberg.de As the complete spider coenoses and species lists were already published (see below) or will be published elsewhere, the focus here is on the following topics: (a) totals of species and abundances, (b) the 20 most abundant spider species, (c) winter activity, and (d) notes on rarely or very rarely recorded species. Methods Trunk eclectors (Fig. 1, method after BEHRE 1989, see also Braun 1992, DOROW et al. 1992, ENGEL 1999) at heights from 2 to 4 meters on living trees were used in different regions of Germany (SW Bavaria, Hesse, Brandenburg) (Fig. 2). In Strict Forest Reserves in Hesse eclectors were also used on dead beech trees (on standing trees and with an adapted type of trap, also on lying trees) (DOROW et al. 1992). The traps were operated during different time periods in the different regions and projects. The trapping periods from May to October were available for comparison from nearly all sites (except Branden- burg). Hesse (Strict Forest Reserves) was the only area where data were available for the whole year, i.e. two entire years including two winters. Most data sets are from beech - Fagus sylvatica and spruce - Picea abies\ see details below. submitted: 13.1.2010, accepted: 1 5. 4. 20 1 0; online: 10.1.2011 6 T Blick Figure 1 :Trunk eclectors on standing beech trees in northern Hesse, Germany, a - spring, b - winter, c - dead tree Sites The data included in this study came from different sites in Germany (Fig. 2): • Three pairs of pine forests, also part mixed with beech and oak, in Brandenburg (no. 1 in Fig. 2, two forests in the north, one in the south) (unpubl. data from 2000 and 2001, June to September, leg. T. Taeger &c U. Schulz, det. TB). In each of the six forests six trunk eclectors were installed on Scots pine — Pinus sylvestris. A total of 36 trees were investigated. Spiders on tree trunks 7 In four Strict Forest Reserves and their reference areas, where forestry is continued, in Hesse (nos. 2-5 in Fig. 2) trunk eclectors were installed on two living and two dead standing beech trees during two whole years (MALTEN 1999, 2001, MALTEN & BLICK 2007, Blick 2009). Additionally at least two eclectors on lying dead beech trees were installed in each reserve. In total 30 standing and 13 lying dead trees were investigated. Mixed forest (‘Stadtwald’), between the urban area of Frankfurt am Main and its airport. 12 trees (alder - Alnus glutinosa, ash - Fraxinus excelsior , birch - Be- tula pendula, two beeches, douglas fir - Pseudotsuga menziesii , elm - Ulmus laevis , Scots pine, two common oaks - Quercus robur , sessile oak - Quercus petraea) were investigated with trunk eclectors in 2000 (beginning of Fe- bruary/mid-March to beginning of November) (M ALTEN et al. 2003, det. in part byTB) (no. 6 in Fig. 2). Forest near Biburg, east of Augsburg, Bavaria (no. 7 in Fig. 2); eight young beeches, four young spruces (20-40 y), two older beeches, two older spru- ces (about 100 y); beginning of April to end of October 1996 (ENGEL 1999, 2001, det. mainly byTB). Forests near Krumbach and Otto- beuren, Bavaria (no. 8 in Fig. 2); 16 beeches, eight douglas firs, eight common oaks, two northern red oaks - Quercus rubra , two silver firs -Abies alba , 22 spruces (unpubl. data from 1999 to 2003, April to October, with exceptions of 1999 (beginning in June) and of 2000 (until November), leg. K. Engel &c M. Gossner, det. TB). Table 1 Totals of the spiders in the trunk eclectors from the 8 sites region no. species adults 1 123 6174 2 116 7796 3 130 10375 4 149 8399 5 149 6685 6 174 9283 7 106 4904 8 177 44712 Results Totals In total 334 spider species, i.e. one third of the spider fauna of Germany, were recorded with 98328 adult spiders. In the 8 different regions between 106 and 177 spider species were recorded, with an average of 140.5 species (± 26.2 standard deviation) (Tab. 1). Figure 2: Sites with trunk eclectors in Germany, included into this study 1 : forests in Brandenburg (Blick & Schultz unpubl.); 2-5: Strict Forest Reserves in Hesse (2: Hohestein, Malten & Buck 2007; 3: Goldbachs- und Ziebachsrück, Buck 2009; 4: Niddahänge östlich Rudingshain, Malten 1 999; 5: Schönbuche, Malten 2001 ); 6: Stadtwald, Frankfurt am Main, Hesse (Malten et al. 2003); 7: forest near Biburg, east of Augsburg, Bavaria (Engel 1999, 2001); 8: forests near Krumbach and Ottobeuren, Bavaria (Blick et al. unpubl.); see text. 8 T Blick Between 21 and 88 spider species were recorded per tree and season (i.e. May-Oct. - based on 115 living trees/season and a total of 298 species), with an ave- rage (0) of 40.5 species (± 12.2). Altogether 122 to 2696 adult spiders per tree and season were trapped, 0 502 (± 452). Twenty most abundant spider species In Tab. 2 the 20 most abundant spider species are sorted in descending abundance. Regional distribu- tion (focus or exclusivity — if nothing is noted the species was recorded in every one of the 8 regions), the tendency to occur on broadleaves/conifers and other notes are added. To summarise: • 8 of the 20 species (40%) were linyphiids, 10 families were present • only 2 species belong to the same genus (. Xysticus ) • 6 species are not known yet as abundant bark species (compare e.g. WUNDERLICH 1982) • 14 species are abundant in (nearly) all regions investigated, 6 species have a more regional distri- bution • 3 species prefer conifers, 3 species prefer beeches/ broadleaves [3 other species were only or mainly trapped in regions, where all/most of the traps were on beech/broadleaf] • these 20 most abundant species make up 65281 adults, i.e. 66.4 % of the adult spiders from the trunks Most abundant spider species active in winter A similar analysis was made for the winter active spe- cies (trapping months December to March, including long winter trapping periods in the regions which ended at the end of April or even at the beginning of May in some years). This means that mainly data of the regions 2-5 (see Fig. 2) and only some addi- tional data of region 6 could be analysed. All these data came from the federal state Hesse. A total of 140 species, 7356 adults and 15714 juveniles (9833 determinable to species level) were recorded during the winter periods. Even in winter, numerous spiders were found in the trunk eclectors. Only two representatives of the commonly known winter-active spiders (e.g. the linyphiid genera Centromerus, Macrargus and Walcke- naeria; the dictynid Cicurina) are on this list (Tab. 3). The majority of the species are also included in Tab. 2. Compared to the forest floor, fewer species on bark were exclusively active in winter. Only Cicurina cicur and Thyreosthenius parasiticus can strictly be placed in the latter category, but at some sites also Monocephalus castaneipes. The 9 species with more than 100 adult individuals comprise 5930 (81%) of the adult spiders from the trunks in the winter - a much less balanced pattern than in the summer (see above). Rare spider species There are several ‘types’ of rare spider species: (a) widely distributed but rare, (b) at or near the border of their distribution area, (c) with restricted distribu- tion areas (Central European endemics?). Examples of species for these types are listed as follows: species name (family), adults recorded (ind.), and distribu- tion area. PLATNICK (2009), STAUDT (2009) and MIKHAILOV (1997) were used as main sources for the total distribution of each species. (a) widely distributed but rare Araneus saevus (L. Koch, 1872) (Araneidae), 2 ind., Holarctic Carrhotus xanthogramma (Latreille, 1819) (Saltici- dae), 59 ind., Palaearctic Dendryphantes hastatus (Clerck, 1757) (Salticidae), 32 ind., Palaearctic Dipoenatorva (Thorell, 1875) (Theridiidae), 120 ind., Europe to W Siberia Mastigusa arietina (Thorell, 1871) (Dictynidae), 3 ind., Europe Philodromus buchari Kubcovä, 2004 (= P. longipalpis auct. in Central Europe) (Philodromidae), 19 ind., Europe and Turkey Stroemiellus stroemi (Thorell, 1870) (Araneidae), 1 ind., Palaearctic Interestingly there are no records of the very rare tree-living species Philodromus poecilus (Thorell, 1872) (Philodromidae), Palaearctic (see MUSTER 2009) and Xysticus albomaculatus Kulczyriski, 1891 (Thomisidae), Germany to Romania, probably Russia (see JANTSCHER 2001). (b) at or near the border of their distribution Cinetata gradata (Simon, 1881) (Linyphiidae), 76 ind., central and southern Europe, mainly in moun- tainous areas, its northern border of distribution is situated in Germany. Clubiona leucaspis Simon, 1932 (Clubionidae) (Figs. 3a, 4a), 583 ind., southern and central Europe, i.e. the northern border is located in Germany, in Ger- many only in the north-eastern and south-western 9 Spiders on tree trunks Table 2: The twenty most abundant bark spider species (see text) species (family) (bold = not well known as bark species) adults recorded regional focus or exclusivity (nos. see fig. 2) tendency to/focus on tree types notes Hahnia pusilla C.L. Koch, 1841 (Hahniidae) 15308 7 & 8, single specimens in 3 &5 only 58 males, females also on the forest floor, males mainly on the floor Amaurobius fenestralis (Ström, 1768) (Amaurobiidae) 8780 also juveniles were determined (in total 16579), also on the forest floor Pelecopsis elongata (Wider, 1834) (Linyphiidae) 5186 7 & 8 rarely also found on the forest floor or in scree and talus habitats Drapetisca socialis (Sundevall, 1833) (Linyphiidae) 5087 slight preference for beech also 3145 juv. recorded Xysticus audax (Schrank, 1803) (Thomisidae) 4106 preference for conifers Walckenaeria cuspidata Blackwall, 1833 (Linyphiidae) 3076 2 &3, few in 4 [2-4: only traps on beech] also on the forest floor Enoplognatha ovata (Clerck, 1757) (Theridiidae) 2691 Lathys humilis (Blackwall, 1855) (Dictynidae) 2630 mainly 8, not 3 slight preference for conifers Moebelia penicillata (Westring, 1851) (Linyphiidae) 2300 Neon reticulatus (Blackwall, 1853) (Salticidae) 1897 majority in 8 slight preference for conifers only 17 males, both sexes mainly on the forest floor Cryphoeca silvicola (C.L. Koch, 1834) (Hahniidae) 1837 mainly 8, not 1 & 6 also on the forest floor Entelecara erythropus (Westring, 1851) (Linyphiidae) 1574 2-6 (Hesse) [2-6: most traps on beeches/broadleaf] also on the forest floor Coelotes terrestris (Wider, 1834) (Amaurobiidae) 1525 not 1 preference for beech mainly on the forest floor Xysticus lanio (Schrank, 1803) (Thomisidae) 1504 preference for broadleaf Lepthyphantes minutus (Blackwall, 1833) (Linyphiidae) 1491 Philodromus collinus C.L. Koch, 1835 (Philodromidae) 1381 rarer on the forest floor Meioneta innotabilis (O. P. -Cambridge, 1863) (Linyphiidae) 1358 not 3, rare in 2 ,4, 5, 7 Diplocephalus cristatus (Blackwall, 1833) (Linyphiidae) 1288 2&3, singleton in 4 [2-4: only traps on beech] mainly on the forest floor, also in open land habitats Harpactea hombergi (Scopoli, 1763) (Dysderidae) 1170 mainly 6, few in 3 & 8 also on rocks and walls Anyphaena accentuata (Walckenaer, 1802) (Anyphaenidae) 1092 not 4 also juveniles can be determined (in total 10322), rare on the forest floor 10 T Blick Figure 3: Distribution maps for Germany of four of the rare spider species (after Staudt 2009); a Clubiona leucaspis, 1 & 2: Blick & Schultz (unpubl.); 3 & 4: Malten et al. (2003); b Gongylidiellum edentatum, 1 : Blick (2009), 2: Malten (1999); c Kratochviliella bicapitata, 1-4: Blick et al. (unpubl.); d Oreonetides quadridentatus, 1 : Malten (1999), 2: Malten (2001), 3: Blick et al. (unpubl.) Spiders on tree trunks 1 1 Table 3: Most abundant winter active spider species (>100 specimens) species ad./juv. regional focus or winter activity notes (family - only when the species is not exclusivity (if > 50 %) (additional present in Tab. 2) (nos. see fig. 2) to Tab. 2) adults Amaurobius fenestralis (Ström, 1768) 2391 87 % males Walckenaeria cuspidata Blackwall, 1833 1685 2 & 3, few in 4 55% Monocephalus castaneipes (Simon, 1884) (Linyphiidae) 839 only 4 Sc 5 81% also on the forest floor Cicurina cicur (Fabricius, 1793) (Dictynidae) 292 not 6 77% also on the forest floor Moebelia penicillata (Westring, 1851) 156 Thyreosthenius parasiticus (Westring, 1851) (Linyphiidae) 156 not 5 74% rarely on the forest floor Diplocephalus cristatus (Blackwall, 1833) 147 only 2 Sc 3 Drapetisca socialis (Sundevall, 1833) 140 Labulla thoracica (Wider, 1834) (Linyphiidae) 124 not 6 also on the forest floor juveniles Anyphaena accentuata (Walckenaer, 1802) 4555 not 4 59% Amaurobius fenestralis (Ström, 1768) 3105 Diaea dorsata (Fabricius, 1777) (Thomisidae) 905 Clubiona leucaspis Simon, 1932 (Clubionidae) 557 only 6 see section ‘rare spider species’ Cryphoeca silvicola (C.L. Koch, 1834) 199 not 6 75% Lathys humilis (Blackwall, 1855) 111 not 2 Sc 4 84% regions with the highest average temperatures, only lowlands. First record for Germany by MALTEN (1994). Note: some records of C. genevensis L. Koch, 1866 on trees might really represent further records of C. leucaspis. For the typical colouration see: http://spiderling.de/ arages/Fotogalerie/ spe- cies_fg.php?name=Clubiona%201eucaspis (STAUDT 2009). Clubiona marmorata L. Koch, 1866 (Clubionidae), 64 ind., central to south-eastern Europe, its north- western border is situated in Germany. Monocephalus castaneipes (Simon, 1884) (Linyphii- dae), 1035 ind., central to northern, western and south-western Europe, eastern border in Ger- many. Theridion boesenbergi Strand, 1904 (Theridiidae), 105 ind., central and eastern Europe, without northern parts, mainly in mountainous areas, northern border in Germany. (c) with restricted distributions; central European ‘endemics’ (all belong to the Linyphiidae) Gongylidiellum edentatum Miller, 1951 (Figs. 3b, 4b, 10 ind., central Europe (Germany, Czech Rep., Austria), N Italy, SE France, most numerous record (31 specimens) is from a rotting beech stump in Hesse (BLICK 2009), see also below. Kratochviliella bicapitata Miller, 1938, 878 ind., central Europe (Germany, Austria, Czech Rep., Slovakia, Poland), Bulgaria; besides the specimens presented here there is just one other record of a larger number of individuals in Poland (CZAJKA Sc BEDNARZ 1972: “only on the northern surface of the tree trunks. ... We estimated that the whole population consisted of some 6000-7000 specimens at that time”), see also below. Oreonetides quadridentatus (Wunderlich, 1972), 82 ind., central Europe (Germany, Austria, Luxem- bourg) and one record with four specimens from the French Pyrenees (BOSMANS et al. 1986); the 12 T Blick record presented here probably refers to the greatest number of individuals found until now. Pseudocarorita thaleri (Saaristo, 1971), 64 ind., central Europe (Germany, Czech Rep., Austria, Switzer- land, Belgium). More details on four of the rare species mentioned above are presented in Figs. 3 and 4, because very little information has previously been published on them. The phenologies of these four species are presented in Fig. 4. The trapping periods are assigned to the month to which the majority of the days of each trapping period belonged; this may not be the month in which the trap was changed or emptied. For Gongylidiellum edentatum and Kratochviliella bicapitata supplementary data are included, i.e. unpublished data from other sites, or from the sites mentioned in this paper but caught with methods other than trunk eclectors. Discussion TOFT (1976) caught 3195 spiders from the end of April to mid-December with arboreal photoeclectors’ (method after NIELSEN 1974) but gave no species number for the individual methods used (total spe- cies number = 147). ALBERT (1982) was the first to publish detailed data from trunk eclectors (method after FUNKE 1971, i.e. without trapping bottles at the bottom; by contrast the type after BEHRE 1989 uses bottles at the bottom, Fig. l).The eclectors were installed on beeches and spruces and were operated from March/ April to November/December in 1969 and 1971. He gives no total species number for the trunk eclectors, but for the four investigated types: spruce old (52 species, 4025 adults), spruce young (39 sp., 972 ad.), beech old (67 sp., 1525 ad.), beech young (33 sp., 199 ad.). PLATEN (1985, also with the eclector type of FUNKE 1971) recorded 74 species over two whole years, 69 on beech and 37 on spruce. Some other examples: BRAUN (1992), who al- ready used the eclector type of BEHRE 1989, caught 108 spider species (3709 adults) on trunks of Scots pine at 3 different heights above the forest floor (1,4 and 8 meters) from May to Oct. SIMON (1995), also trapped on Scots pine and at different heights (1.5, 5, 10 and 13 meters - species numbers 71, 59, 48, 35, i.e. lower numbers at increasing height), over 3 whole years collecting a total of 103 species (including traps on branches in the crown). GOERTZ (1998) examined 10 trees (5 crack willows — Salix fragilis , 2 almond willows — Salix triandra , 3 black poplars - Populus aff. nigra) from mid-December to end of June and caught a total of 100 species and 20167 determinable -Q — rn ,m~iiIhiI n,d~l a r I I i ,d~L*-nJTi ,-TI H — “i — i— -| — — =i — 'I U -\ 1 1— T T — i — ' T 'i — *— h — — *1 — *1 1 II III IV V VI VII VIII IX X IV V VI VII VIII IX X XI XII Figure 4: Phenology of (a) Clubiona leucaspis, (b) Gongylidiellum edentatum, (c) Kratochviliella bicapitata, (d) Oreonetides quadridentatus (black males, white females, grey juveniles; x-axis: Roman numerals represent the months; y-axis: numbers of trapped specimens); a Clubiona leucaspis (no traps operated from November/XI to January/I), totals 441 3 3, 159$ $, 1231 juv.; b Gongylidiellum edentatum, totals 123 3,45 $ $ (incl.33 3,7$ $ in long winter periods, i.e. Nov./Dec. to end of April/beginning of May - not included in this graph); c Kratochviliella bicapitata, totals 4903 3, 399$ $ ; d Oreonetides quadridentatus, totals 43 3, 83 $ $ (incl.13 in a longer winter period, i.e. mid Nov. to mid of March - not included in this graph) Spiders on tree trunks 13 specimens. Details: 29-55 species (0 41.2 ± 8.8), 580- 3422 spiders (0 2017 ± 918). FINCH (2001) caught from spring to autumn on 6 living trees (3 beeches, 2 oaks, 1 Scots pine - open type after FUNKE 1971) 110 species (0 35.2 ± 15.3), and 56 species on two lying dead trees. All these data fit in the ranges given above for single trees. The eclector type without a bottle traps fewer specimens (and presumably less species) than the type with a bottle. Nevertheless, data from the bottle-less type fit within the range. A possible reason: the range of species and specimen numbers on older trees (with a larger diameter) compared to younger trees is larger than the range resulting from trap type. The data from different regions in this paper already show that it is difficult to compare the published data. The reasons are that different trapping periods, different tree ages and diameters and finally different types of eclectors were used. Nevertheless, important conclusions can be drawn (see also BLICK 2010): • spiders are a species rich group on forests floors, as well as on trees; • to estimate their biodiversity in forests data from both the forest floor and from trees are necessary; • the majority of the dominant species do not prefer conifers or broad-leafed tress; instead they are widely distributed; • in Germany forest spider species that are restric- ted in their distribution to Central Europe (s.l.) occur; • the diversity and importance of the spider fauna on bark in Central Europe was until now underestima- ted compared with the epigeal fauna (even though there is an overlap of both coenoses). Acknowledgements My thanks go: to Andreas Malten (Frankfurt am Main), who determined the spiders of three strict forest reserves and to Ingmar Weiss (Grafenau-Rosenau), who determined a part of the spiders from Frankfurt; to the whole team of the Strict Forest Reserve Project, especially to Wolfgang Dorow (Frankfurt am Main), the old stager of the project; to Aloysius Staudt (Schmelz) for his great work on the German www-distribution maps; toTimTaeger and Ulrich Schultz (Eberswalde) for their field and lab work; to Kerstin Engel, Martin Goßner (formerly Freising), and Andreas Malten for collecting the spiders and for arranging finances; last but not least to Andrew Liston (Müncheberg) for impro- ving the English. Research on the Strict Forest Reserves in Hesse was conducted in cooperation with and financially supported by the ‘Landesbetrieb Hessen-Forst’. References ALBERT R. 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Charakterisierung der Lebensräume der häufigsten Spinnenarten Mittel- europas und der mit diesen vergesellschafteten Arten. - Miscellanea Faunistica Helvetiae 4: 1-459 JANTSCHER E. (2001): Revision der Krabbenspinnengat- tung Xysticus C.L. Koch, 1835 (Araneae, Thomisidae) in Zentraleuropa. Thesis, Graz. 328 pp. &81 tables MALTEN A. (1994): Fünf für Deutschland neue Spinnen- arten - Lepthyphantes midas, Neriene furtiva, Hahnia petrobia, Clubiona leucaspis, Diaea pictilis (Araneae: Linyphiidae, Hahniidae, Clubionidae, Thomisidae). - Arachnologische Mitteilungen 8: 58-62 MALTEN A. (1999): Araneae (Spinnen). Naturwaldreserva- te in Hessen, Band 5/2.1. Niddahänge östlich Rudings- hain. Zoologische Untersuchungen 1990-1992, Teil 1. - Mitteilungen der Hessischen Landesforstverwaltung 32/1: 85-197 MALTEN A. (2001): Araneae (Spinnen). Naturwaldreser- vate in Hessen, Band 6/2.1. Schönbuche. 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MÜLLER (2010): Arthropod species richness in the Norway Spruce {Picea abies (L.) Karst.) canopy along an elevation gradient. - Forest Ecology and Management 259: 1513- 1521 - doi: 10.1016/j.foreco.2010.01.027 SIMON U. (1995): Untersuchung der Stratozönosen von Spinnen und Weberknechten (Arachn.: Araneae, Opil- ionida) an der Waldkiefer ( Pinus sylvetris L.). Wissen- schaft & Technik, Berlin. 142 pp. STAUDT A. (2009): Nachweiskarten der Spinnentiere Deutschlands (Arachnida: Araneae, Opiliones, Pseu- doscorpiones). - Internet: http://spiderling.de/arages [accessed 28.XII.2009]. TOFT S. (1976): Life-histories of spiders in a Danish beech wood. - Natura Jutlandica 19: 5-40 WUNDERLICH J. (1982): Mitteleuropäische Spinnen (Araneae) der Baumrinde. - Zeitschrift für angewandte Entomologie 94: 9-21 - doi: 10.1111/j.l439-0418.1982. tb02540.x Arachnologische Mitteilungen 40: 15-22 Nuremberg, January 201 1 On some new or rare spider species from Lesbos, Greece (Araneae: Agelenidae, Amaurobiidae, Corinnidae, Gnaphosidae, Liocranidae) Robert Bosmans doi:10.5431/aramit4003 Abstract: In this paper, three new spider species are described from the Greek Aegean island Lesbos: Tegenaria mael- faiti sp. nov. (Agelenidae), Amaurobiuslesbius sp. nov. (Amaurobiidae) and Agroeca parva sp. nov. (Liocranidae), as well as the unknown male of Arabella pheidoleicomes Bosselaers,2009 (Corinnidae). Diagnostic features and notes on ecology and distribution of these species are given. Two new records for the island are provided, such that currently 300 spider species are known from Lesbos. Key words: Agroeca, Amaurobius, Arabella, description, new species, Tegenaria The spider fauna of Greece is still poorly known compared to other Mediterranean countries. Recently, BOSMANS & ChATZAKI (2005) reported 856 spe- cies known from Greece. In a study of one relatively small region, the Nestos delta, BUCHHOLZ (2007) added 61 spider species to the Greek fauna. The spiders of the island Lesbos were recently surveyed by a group of Belgian arachnologists during several collecting trips (October 2007, March-April 2008, May-June 2008). This resulted in a catalogue of 292 species, some of them new to science. Among these, VAN KEER & BOSMANS (2009) described four new dysderid species. The current paper provides the de- scription of three more new species, belonging to the families Agelenidae, Amaurobiidae and Liocranidae and the description of the unknown male of another liocranid genus. Material and Methods The material treated in this paper was collected by members of the Belgian Arachnological Society ARAB EL during several trips to Lesbos, as detailed in BOSMANS et al. (2009). Specimens were examined and illustrated using a Wild M5 stereomicroscope. Further details were studied using an Olympus CH-2 stereoscopic micro- scope with a drawing tube. Structures of the left palpus are depicted. All morphological measurements are given in millime- Robert BOSMANS, Terrestrial Ecological Unit, Department of Biology, Ghent University, Ledeganckstraat 35, B-9000 Gent, Belgium; E-Mail: robert.bosmans@lne.vlaanderen.be tres. Somatic morphology measurements were taken using a scale reticule in the eyepiece of the stereo microscope. Measurements of the legs are taken from the dorsal side. Spines on leg segments are listed in the following order: dorsal-prolateral-retrolateral-pro- ventral-retroventral. Male palps were detached and transferred to gly- cerol for examination under the microscope. Female genitalia were excised using sharpened needles and then transferred to clove oil for examination under the microscope. Later, palps and epigynes were returned to 70% ethanol. Type material and important reference material is deposited in the Koninklijk Belgisch Instrituut voor Natuurwetenschappen, Brussels, the other material is deposited in the collections of the individuals listed below. The following abbreviations are used in the text: AME: anterior median eyes; CJVK: collection Johan Van Keer; CHDK: collection Herman De Köninck; CRB: collection Robert Bosmans; KBIN: Koninklijk Belgisch Instituut voor Natuur- wetenschappen (Brussels); Legs: Ta: tarsus; Mt: metatarsus; Pa: patella; Fe: femur. Spines: d: dorsal; v: ventral; pi: prolateral; rl: retro- lateral; spines listed between brackets: occurring in pairs or groups at about the same level; *: spines with a terminal position. submitted: 14. 1.2010, accepted: 1 1.5.2010; online: 10.1.2011 16 R. Bosmans Results Family Agelenidae Genus Tegenaria Latreille, 1804 According to BOLZERN et al. (2009), the differences between the genera Tegenaria and Malthonica Simon, 1898 remain unclear. GUSEINOV et al. (2005) distin- guished the two genera by the embolus length but BOLZERN et al. (2008) argued that this character is unsuitable for phylogenetic evaluation. Assigning a new species to a correct genus thus becomes difficult without a comprehensive revision of the two genera (BOLZERN et al., 2009). The species described below has a relatively short embolus and is provisionally placed in the genus Tegenaria. Tegenaria maelfaiti sp. nov. (Figs. 1-5). Type material. Holotype male, 2 paratype males from Greece, Lesbos, Nomos Mandamados: road Klio-Sykami- nia (N 39°22’33” E 26T0’59”), 350 m, litter and stones in small Pinus forest, 27. III. 2008, R. Bosmans leg.; paratype female from Greece, Lesbos, Nomos Agiasos, Agiasos S. (N 39°05’53” E 26°21’57”), 550 m, stones in Castanea forest, 2.IV.2008, L. Baert leg. (KBIN). Deposited in KBIN. Etymology: The species is dedicated to my good friend and excellent ecologist the late Jean-Pierre Maelfait. Diagnosis: Males of the species are readily distingu- ished from other Tegenaria species by the combined characters of the tibial apophyses and the bifid median apophyses. Tegenaria longimana Simon, 1879 from Turkey is a related species, with similar but differently shaped tibial apophysis and distal parts of the median apophysis. Females are recognized by the absence of sclerotisation in the epigyne and the course of the copulatory ducts. Description Measurements (males n=27, females n=4): Male: Total length 3. 8-6.2; prosoma 1.59-2.79 long, 1.46- 2.06 wide. Female: Total length 4. 8-5.0; prosoma 1.87-1.98 long, 1.34-1.43 wide. Leg measurements as in Table 1. Colour: Prosoma yellowish brown to brown, with submarginal stripe, touching at level of fovea, and narrow grey margin; chelicerae yel- lowish brown; sternum grey with median stripe and 3 lateral whitish spots. Legs pale yellowish, Fe, Ti Table 1 : Tegenaria maelfaiti sp. nov. leg measurements (holotype). Fe Pt Ti Mt Ta Palp 1.23 0.42 0.76 - 0.79 I 3.24 0.92 3.29 3.44 1.62 II 2.85 0.78 2.31 2.97 1.37 III 3.23 0.60 1.96 2.57 0.96 IV 2.96 0.76 2.72 3.52 3.51 and Mt greyish at ends, Fe with 2 grey annulations, Ti and Mt with 1 median annulation. Abdomen dorsally greyish black, with anteromedian area and 5-6 chevrons whitish, ventrally whitish with scattered greyish black spots, anastomising in some specimens. Spinnerets whitish, dorsal side of dorsal spinneret and ventral side of ventral spinneret greyish black. Leg spination as in Table 2. Male palp (Figs. 1-3): Tibia elongated, dorsal apophy- sis long, with parallel margins and rounded terminally, retrolateral apophysis a large, rounded lobe. Median apophysis bifid, with shorter, pointed anterior tooth and longer, less pointed posterior tooth. Conductor slightly shorter than alveolus, rounded, folded along its entire retrolateral margin, terminal part pointing in posterior direction. Embolus filiform, semi-circular, arising on mesal side of bulbus. Epigymim and vulva (Figs. 4-5): Epigynal plate slightly protruding over epigastric furrow, posterior margin with two indistinct concavities, with a pair of posteriorly directed teeth but without exterior sclerifications; copulatory ducts wide; spermathecae rounded. Further material examined Agiasos: Oros Olympos (N 39°04’01” E 26°21’30”), 800 m, 1 female, stones in Platanus forest, 2.VI.2008 (CRB); - Kalloni: Anemotia, E. of Moni Voukolon (N 39°04’00” E 26°07’18), 400 m, 1 female, stones in grassland, 29.III.2008 (KBIN); road Anemotia-Skala Kallonis, Karyo (N 39°13’36” E 26°07’50”), 100 m, litter and herbs along rivulet, 29.III.2008 (CRB); - Mandamados: road Klio-Sykaminia (N 39°22’33” E 26°10’59”), 350 m, 24 males, litter and stones in small Pinus forest, 27. III. 2008 (CHDK, CJVK, CRB); - Mithymna: road Argenos-Vafios (N 39°21’06” E Table 2: Tegenaria maelfaiti sp. nov. leg spination (holotype). Fe Pt Ti Mt Ta I (dpl)-d-pl d-d d vrl (plrl) - II d-d-(plrl) d-d d vpl 2v (plrl) - III d-d-(plrl) d-d d (plrl) rl pv (rlplpvrv) - IV d-(pl-rl) d-d (drl) (pvplrl) (dplrl) (pvrv) (plrlrv) (plrlpvrv) - New and rare spiders from Lesbos 17 Figures 1-5: Tegenaria maelfaiti sp. nov. I.Male palp, lateral view; 2. Idem, ventral view; 3. Palpal tibia, dorsal view; 4. Epigyne, ventral view; 5. Vulva, ventral view. C = conductor; CD = copulatory duct; CO = copulatory opening; DTA = dorsal tibial apophysis; E = embolus; ET = epigynal tooth; MA = median apophysis; RTA = retrolateral tibial apophysis; ST = spermatheca. 26°14’29”), 320 m, 3 males, stony grassland and shrubs, 28.III.2008 (CHDK, CJVK, CRB). Ecology: Males were collected in March and April, females in March, April and June. It was mainly found in forest (pine, chestnut and plane forests), but also in shrubs and olive groves and along rivulets. Distribution: Only known from Lesbos. As Tegenaria species tend towards small distribution areas, this species is possibly endemic to the island. Family Amaurobiidae Genus Amaurobius C. L. Koch The genus Amaurobius currently includes 67 species (PLATNICK 2010) and has a mainly Holarctic distri- bution. In Greece, it has been thoroughly studied by Thaler 5c Knoflach (1991, 1993, 1995, 1998, 2002) and WUNDERLICH (1995). Thirteen Amau- robius species have been found in Greece (BOSMANS 5c CHATZAKI 2005). A. candia Thaler 5c Knoflach, 2002, A. cretaensis Wunderlich, 1995 and A. geminus Thaler 5c Knoflach, 2002 are endemics for Crete, A. phaeacus Thaler 5c Knoflach, 1998 is an endemic for Kerkyra, A. deelemanae Thaler 5c Knoflach, 1995 occurs on Crete, the Dodekanisa and Kyklades and A. ausobskyi Thaler 5c Knoflach, A. longipes Thaler 5c Knoflach, 1995,^4. ossa Thaler 5c Knoflach, 1993,^. paon Thaler 5c Knoflach, 1993 and A.pelops Thaler 5c Knoflach, 1991 are endemics occurring in continental Greece. Amaurobius pallidas L. Koch, 1868 and A. strandi Charitonov, 1937 have an eastern Mediter- ranean distribution and A. erberi (Keyserling, 1863) has a circummediterranean distribution. From the Eastern Aegean islands (to which Lesbos belongs) orAy Amaurobius scopoliiTh.ore\\, 1871 has been cited from Chios (STRAND 1917) but this is most prob- ably a misidentification. It is thus not surprising to discover another new Amaurobius species on one of the Aegean Islands. 18 R. Bosmons Amaurobius lesbius sp. nov. (Figs. 6-10) Type material: Holotype male, 1 paratype male two paratype females from Greece, Lesbos, Nomos Kalloni, Skalochori (N 39°15’59” E26° 05’ 30”), 300 m, litter in Quercus forest, 2.X.2007, R. Bosmans & J. Van Keer leg. Deposited in KBIN. Etymology: The species is named after the island of Lesbos. Diagnosis: Amaurobius lesbius n. sp. is closely related to the larger A. erberi (Keyserling), also occurring on Lesbos. Males differ by the reduction of an interme- diate apophysis in the palpal tibia, and the presence of a strongly protruding, rectangular tegular process. Females differ by the closely set spermathecae, sepa- rated by less than Va their diameter in the new species and by their diameter in A . erberi. Description Colour: Prosoma yellowish brown, sometimes with greyish striae, thoracic part paler; chelicerae dark orange brown. Legs yellowish brown. Abdomen in males dark grey to black with anteromedian pale stripe and some subterminal, pale grey chevrons, venter pale to dark grey, in females dorsally pale grey with dark grey pattern. Measurements (males: n=6; females: n= 8): Male: Total length 3. 6-4.6; prosoma 1.81-2.24 long, 1.21- 1.56 wide. Female: Total length 4. 8-6. 8; prosoma 1.99-2.26 long, 1.18-1.54 wide. Leg measurements as in Table 3, leg spination as in Table 4. Male palp (Figs. 6-8): Tibia with small, curved dorsal and larger blunt retrolateral apophyses, the former with a tooth at its base, intermediate apophysis reduced to a short crest. Tegular process strongly protruding, nearly rectangular. Tegular apophysis elongated, terminally hooked and embolus short and bent. Epigyne and vulva (Figs. 9-10): With reversed tra- Table 3: Amaurobius lesbius sp. nov., leg measurements (holotype). Fe Pt Ti Mt Ta Palp 0.72 0.26 0.34 - 0.39 I 1.51 0.76 1.36 1.54 0.91 II 1.42 0.74 1.14 1.08 0.89 III 1.31 0.75 0.99 1.29 0.77 IV 2.02 0.79 1.82 2.28 1.12 Figures 6-10: Amaurobius lesbius sp. nov. 6. Male palp, lateral view; 7. Idem, ventral view; 8. Palpal tibia, dorsal view; 9. Epigyne, ven- tral view; 1 0. Vulva, ventral view. C = conductor; DTA = dorsal tibial apophysis; E = embolus; ITA = intermediate tibial apophy- sis; MP = median plate; RTA = retrolateral tibial apophysis; ST = spermatheca;TA = tegular apophysis;TP = tegular process. New and rare spiders from Lesbos Table 4:Amaurobiuslesbius sp. nov., leg spination (holotype). 19 Fe Pt Ti Mt Ta I pi - 3pl 2rl 3pv 3rv 2pl 2rl 3pv 3rv pdrd* - II P1 - 3pl 2rl 3pv 3rv 3pl 2rl pv 2rv pdrd* - III (pd rd) - 3pl 3pv rv rl 2pl pv 3rv pd rd* - IV rd - pv 3rv rl rl pv 3rv pd pv* - pezoid plate. Spermathecae closely set, separated by less than Va their diameter, basal part covered by epigynal plate. Further material examined Agiasos: Agiasos S. (N 39°05’53” E 26°21’57”), 550 m, 5 males, pitfalls in Castanea forest, X.2007-IV.2008 (CJVK); - Mandamados: road Klio-Sykaminia (N 39°22’33” E 26°10’59”), 350 m, stones in small Pinus forest, 1 subadult male 1 female, 6.X.2007, 6 females, 27. III. 2008 (CHDK, CJVK, CRB). Ecology: All specimens were captured in Castanea , Pinus and Quercus forests. Males were only collected in winter, females in winter and spring. Distribution: At the moment, the species is only known from Lesbos, but as Amaurobius species gen- erally have large distribution areas, it may occur on other East Aegean Islands as well. Family Liocranidae Genus Agroeca C. L. Koch The genus Agroeca currently includes 25 species and has a Holarctic distribution (PLATNICK 2010). In Greece, only Agroeca cuprea Menge, 1873 has been previously reported (BOSMANS Sc CHATZAKI 2005). A new species is described below. Agroeca parva sp. nov. (Figs. 11-14) Type material: Holotype male, 2 male paratypes 1 female paratype from Greece, Lesbos, Polichnitos, Aghios Pavlos E. (N 39°07’38” E 26T4’03”), 10 m, pitfalls in Juncus marsh, 11.X.2007, R. Bosmans leg. Deposited in KBIN. Etymology: The name refers to the small size of the new species. Diagnosis: Agroeca parva sp. nov. is recognised by its small size and its contrasting colour. The male palp and female epigyne suggest the species is closely related to Agroeca annulipes Simon, 1878 and A. maghrebensis Bosmans, 1999, both from the western Mediterranean. Males differ by the truncate palpal tibial apophysis, more pointed in the species from the Maghreb, females by the concave lateral margins of the epigynal depres- sion, straight in the species from the Maghreb. Description Colour: Prosoma yellowish brown with narrow margin and submedian stripe greyish black. Abdomen grey black, median denticulate stripe grey brown, venter yellowish brown with few grey black spots in posterior half. Legs yellowish brown, femora with 2 broad grey annulations, patellae I-III with basal half grey. Measurements (males: n =3, females n = 6): Male: Total length 3. 6-3. 8; prosoma 1.84-2.12 long, 1.44- 1.71 wide. Female: Total length 4. 1-4.5; prosoma 1.83-2.16 long, 1.51-1.64 wide. Leg measurements as in Table 5, leg spination as in Table 6. Figures Agroeca parva sp. nov. 1 1 .Male palp, lateral view; 12. Idem, ventral view; 13. Epigyne, ventral view; 14. Vulva, ventral view. CD = copulatory duct; E = embolus; MA = Median apohysis; RTA = retrolateral tibial apophysis; ST = spermatheca. 20 Table 5:Agroeca parva sp. nov. leg measurements (holotype). Fe Pt Ti Mt Ta Palp 0.72 0.26 0.34 - 0.39 I 1.51 0.76 1.36 1.54 0.91 II 1.42 0.74 1.14 1.08 0.89 III 1.31 0.75 0.99 1.29 0.77 IV 2.02 0.79 1.82 2.28 1.12 Table 6:Agroeca parva sp. nov. leg spination (holotype). Male palp (Figs. 11, 12): Tibia with thick, straight apophysis, terminally truncate at dorsal margin. Median apophysis reversed L-shaped, with distinct anterior concavity and terminally pointed. Embolus gradually narrowing terminally curved towards the median apophysis. R. Bosmans Epigyne and vulva (Figs. 13-14): With large median depression, concave lateral margins and a posteriorly narrowing median septum. Further material examined Agiasos: Megali Limni W., Steni Klidi, (N 39°06’47” E 26°18’30”), 400 m, 2 females, pitfalls in Pinus forest, 3.IV.2008 (CJVK, CRB); - Kalloni: Skala Kalloni (N 39°12’28” E 26°12’01”), 10 m, 1 female, litter in vegetable garden of hotel, 6.X .2007 (CRB); - Polichni- tos: Polichnitos salt works (N 39°06’37” E 26°10’50”), 5 m, stones in Salicornia salt marsh, 2 females, 31.III.2008 (CRB); Vatera (N 39°01’00” E 26°13’59”), 30 m, 1 male, litter in garden of hotel, 9.X.2007 (CRB). Ecology: Males of this species were only collected in October, females in October and March. It was found in salt marshes, pine forests and even in a vegetable garden, so there seem to be no particular habitat preferences. Distribution: Currently, only known from Lesbos, but as all other Agroeca species have large distribution areas, it probably occurs in a much wider area. Fe Pt Ti Mt Ta I 2d lpl - 2 (pv rv) 3 (pv rv) - II 2d lpl - 2 (pv rv) 3 (pv rv) - III 2d lpl Id 2pl 3pv 3rv 2rl 3 pi 3pv 3rv 3rl - IV 2d lpl Id 3pl 3pv 3pl 3rl 3 pi 3pv 3rv 3rl - Family Corinnidae Genus Arabelia Bosselaers, 2009 The genus Arabella was only recently described and includes a single species from the East Mediterranean region. The male is described here for the first time. Arabelia pheidoleicomes Bosselaers, 2009 (Figs. 15-16) Arabella pheidoleicomes Bosselaers, 2009: 51, Figs. 2K, 7 A-G, 8H, 9 (descr. female). Remark The female of this species was recently described by BOSSELAERS (2009), based on material from Rhodes and Lesbos. Later, the corresponding male appeared to be present in the material from Lesbos as well and it is described here for the first time. Somatic morphology is the same as in the female, except as indicated below. Diagnosis The male of this species is recognised by the short, pointed tibial apophysis and the small sperm duct tooth. For a description of the female, see BOSSE- LAERS (2009). Description of the unknown male Colour: As in female, but femora and patella of legs I-II somewhat infuscate, contrasting with paler re- maining segments. Measurements (1 male): Total length 3.5; prosoma 1.36 long, 1.21 wide. Leg measurements as in Table 7, leg spination as in Tab. 8. Male palp (Figs. 15-16): Tibia with small, pointed re- trolateral apophysis. Bulbus ovoid, without apophyses, only with small, anterolateral sperm duct tooth. Material examined Agiasos: Agiasos NW, Voula (N 39°06’26” E 26°21’14”), 350 m, 1 male, 1.IV.2008, dense Pinus forest along rivulet, 1.IV.2008, L. Baert leg.; deposited in KBIN. Distribution Greece (Rhodes, Lesbos) and Cyprus. Ecology Females were always collected together with ants, the only male collected was found in a termite nest. New and rare spiders from Lesbos 21 Figures 1 5-16: Arabella pheidoleicomes Bosselaers, 2009. 1 5. Male palp, lateral view; 1 6. Idem, ventral view. E = embolus. New spider records from Lesbos Fam. Gnaphosidae Drassodes lacertosus O. R- Cambridge, 1872 Identification Levy, 2004: 6, £13-19. Material examined Mithymna: Mithymna (N 39°22’02” E 26°10’30”), 10 m, 1 male in pool of hotel, 28.V.2009, A. Decae leg. (CRB). Distribution: The species is known from Turkey, Is- rael, Syria and Russia (PLATNICK 2010) and is new to Greece. The present record is the most western one. Trachyzelotes barbatus (L. Koch, 1866) Identfication Chatzaki, Thaler 6c Mylonas, 2003: 53, f. 20-21, 26-27. Material examined Mithymna: Mithymna (N 39°22’02” E 26°10’30”), 10 m, 4 males 1 female in pool of hotel, 28.V.2009, A. Decae leg. (CRB). Distribution: Mediterranean to Central Asia, USA. New to Lesbos, but previously recorded in continental Greece and Crete (CHATZAKI et al. 2003). Discussion In their catalogue of the spiders of Lesbos, BOSMANS et al. (2009) list 298 species, some of them previously undescribed. Three Harpactea species and one Stalag- tia species were described in VAN KEER 6c BOSMANS (2009), one Zodarion species in BOSMANS (2009) and three more species are described in the present paper. Additional species belonging to the spider family Linyphiidae will be described in a further paper. Finally, some species were discovered that remain unidentified to species level, for different reasons. Sometimes, only one sex was captured, sometimes the genus is in need of revision or the systematic position is unclear. Further investigations are necessary to solve these problems. The description of these species can only be carried out after revisions of the relevant genera or after the discovery of the corresponding sex. Acknowledgements I wish to thank my friends and colleagues Leon Baert, Jan Bosselaers, Herman De Köninck, Jean-Pierre and Lutgarde Maelfait and Johan Van Keer for joining me on Lesbos and making this journey such a pleasant event and also allow- ing me to examine their material. Arthur Decae is thanked for making available his material collected in Lesbos. The English of the final draft was kindly checked by Anthony Russell-Smith. References BOLZERN A., A. HÄNGGI 6c D. BURCKHARDT (2008): Funnel web spiders from Sardinia: taxonomical notes on some Tegenaria and Malthonica spp. (Araneae: Ageleni- dae). - Revue suisse de Zoologie 115: 759-778 Bolzern A., L. Crespo 6c P. Cardoso (2009): Two new Tegenaria species (Araneae: Agelenidae) from Portugal. - Zootaxa 2068: 47-58 BOSMANS R. (2009): Revision of the genus Zodarion Walckenaer, 1833, part III. South East Europe and Turkey (Araneae: Zodariidae). - Contributions to Natural History 12 (Konrad Thaler Memorial Volume): 211-296 BOSMANS R. 6c M. CHATZAKI (2005): A catalogue of the spiders of Greece. A critical review of all spiders species cited from Greece with their localities. - Nieuwsbrief van de Belgische arachnologische Vereniging 20 (2, suppl.): 1-124 Bosmans R., L. Baert, J. Bosselaers, H. De Köninck, J.-P. Maelfait 6c J. Van Keer (2009): Spiders of Lesbos (Greece). A catalogue with all currently known spider reports from the Eastern Aegean Island of Les- bos. - Nieuwsbrief van de Belgische arachnologische Vereniging 24 (suppl.): 1-71 BOSSELAERS J. (2009): Studies in Liocranidae (Araneae): redescriptions and transfers in Apostenus Westring and Brachyanillus Simon, as well as description of a new genus. - Zootaxa 2141: 37-55 22 R. Bosmans BUCHHOLZ S. (2007): A first contribution to the Arach- nofauna (Arachnida: Araneae) of the Nestos Delta (NE Greece). - Acta Zoologica Bulgarica 59: 241-252 Chatzaki M., K. Thaler & M. Mylonas (2003): Ground spiders (Gnaphosidae; Araneidae) of Crete (Greece). Taxonomy and distribution. III. Zelotes and allied genera. - Revue suisse de Zoologie 110: 45-89 Guseinov E.F., Y.M. Marusik & S. Koponen (2005): Spiders (Arachnida: Aranei) of Azerbaijan 5. Faunistic review of the funnel-web spiders (Agelenidae) with the description of a new genus and species. - Arthropoda S electa 14: 153-177 LEVY G. (2004): Spiders of the genera Drassodes and Hap- lodrassus (Araneae, Gnaphosidae) from Israel. - Israel Journal of Zoology 50: 1-37 - doi: 10.1560/RUQP- 20ML-VDBA-3GKX. PLATNICK N. (2010): The world spider catalog, version 10.5 American Museum of Natural History. - Internet: http://research.amnh.org/ entomology/ spiders/ catalog STRAND E. (1917): Arachnologica varia XIX-XX. - Archiv für Naturgeschichte 82: 158-167 Thaler K. & B. KNOFLACH (1991): Eine neue Amau- robius-Axt aus Griechenland (Arachnida: Araneae, Amaurobiidae). - Mitteilungen der schweizerischen entomologischen Gesellschaft 64: 265-268 THALER K. &B. KNOFLACH (1993): Two new Amaurobius species (Araneae: Amaurobiidae) from Greece. — Bulletin of the British arachnological Society 9: 132-136 Thaler K. & B. KNOFLACH (1995): Über Vorkommen und Verbreitung von Amaurobius- Arten in Peloponnes und Ägäis (Araneida: Amaurobiidae). - Revue suisse de Zoologie 102: 41-60 THALER K. &B. KNOFLACH (1998): Two new species and new records of the genus Amaurobius (Araneae, Amau- robiidae) from Greece. In: SELDEN PA. (Ed.): Proceed- ings of the 17th European Colloquium of Arachnology, Edinburgh 1997, Edinburgh, pp. 107-114 Thaler K. & B. KNOFLACH (2002): A superspecies in the genus Amaurobius on Crete, and additional records from Greece (Araneae: Amaurobiidae). In:TOFT S. 8cN. SCHARFF (Eds): European Arachnology 2000. Procee- dings of the 19th European Colloquium of Arachnology, Aarhus. Univ. Press, Aarhus, pp. 337-344 VAN KEER J. & R. BOSMANS (2009): On some new Harpactea and Stalagtia species from Lesbos, Greece (Araneae: Dysderidae).- Acta Zoologica Bulgarica 61: 2009: 277-285 WUNDERLICH J. (1995): Beschreibung einer bisher un- bekannten Art der Gattung Amaurobius C. L. Koch 1837 von Kreta (Arachnida: Araneae: Amaurobiidae). - Beiträge zur Araneologie 4: 729-730 Arachnologische Mitteilungen 40:23-32 Nuremberg, January 201 1 The faunistic diversity of cave-dwelling spiders (Arachnida, Araneae) of Greece Christo Deltshev doi: 1 0.543 1/aramit4004 Abstract: Until today, from Greek caves a total of 1 09 species of spiders belonging to 25 families are known. One species, the linyphiid Porrhomma convexum (Westring, 1861) was recorded here for the first time in Greece. The 109 species are distributed in caves of different geographic territories as follows:Thrace - 8 species, Macedonia - 1 8, Epirus - 1 , Thessaly - 6, Central Greece - 3, Attiki-Saronic Islands - 24, Peloponnese - 1 5, Evoia-Vories Sporades - 1, Eastern Aegean Islands - 5, Cyclades - 3, Dodecanese - 6, Ionian Islands - 23, Crete - 47. The largest fraction of troglobite species were encountered mainly in the territories of Crete - 15 species (5 of which are anophthalmic), the Ionian Islands - 4, Thrace - 2 (both anophthalmic), the Attiki-Saronic Islands - 2 (both anophthalmic), the Pe- loponnese - 2 (one anophthalmic), and Macedonia, Thessaly, and the Cyclades - each with 2 species. The richness of the troglobitic spiders in these regions strengthens the assumption that they were major centres of speciation and evolution for the species of this group. According to their current distribution, the established 1 09 species can be classified into 12 zoogeograpical categories, grouped into 4 complexes (widely distributed, European, Mediterranean, endemics). The largest number of species belong to the endemic complex (53.2 %) and are also the most characteristic and reflect the local character of the cave-dwelling spiders. Key words: cave-spiders fauna, endemics, troglobites, zoogeography The earliest data on Greek cave-dwelling spiders were presented by SIMON (1885), KULCZYNSKI (1903), ROEWER (1928, 1959), DRENSKY (1936); Kra- TOCHVIL (1937, 1938), HADJISSARANTOS (1940), and FAGE (1945). More recent publications derive from the investigations of BRIGNOLI (1968, 1971a, 1971b, 1972, 1974a, 1974b, 1974c, 1976, 1977, 1978, 1979, 1984), Deeleman-Reinhold (1971, 1977, 1983, 1985, 1989, 1993), DEELEMAN-REINHOLD & DEELEMAN (1988), SENGLET (1971, 2001), Deltshev (1979, 1985, 1999, 2000, 2008), Beron (1985, 1986), Beron & Stoev (2004), Thaler & Knoflach (1995), Wunderlich (1995), Bosse- LAERS (1998), BOSSELAERS &HENDERICKX (2002), GASPARO (2003, 2004a, 2004b, 2005a, 2005b, 2006, 2007, 2008, 2009), CHATZAKI et al 2002, BOSMANS dt CHATZAKI (2005), CHATZAKI &ARNEDO (2006), and PLATNICK (2009). The critical incorporation of all available literature records and the accumulation of new data are now sufficient to allow a critical analysis of the distribution of spiders established in the caves of Greece. Study area and material Greece is a country in south-eastern Europe, situated Christo DELTSHEV, Institute of Biodiversity and Ecosystem Research, 1 Tsar Osvoboditel Blvd; 1 000 Sofia, Bulgaria E-Mail: deltshev@gmail.com submitted: 30. 12.20 1 0, accepted: 9.4.20 1 0; online: 10.1.2011 on the southern end of the Balkan Peninsula. The country has borders with Albania, the Republic of Macedonia and Bulgaria to the north, and Turkey to the east. The Aegean Sea lies to the east and south of mainland Greece, while the Ionian Sea lies to the west. Both parts of the Eastern Mediterranean basin feature a vast number of islands, islets and rock is- lands (Fig. 1). Two -thirds of the territory of Greece is dominated by limestone, many of which are karstified (CLENDENON 2009). The territory of Greece can be divided into 13 geographical regions (BOSMANS & CHATZAKI 2005; Fig. 1). There are 7 geographical regions on the main- land: Thrace, Macedonia, Epirus, Thessaly, Central Greece, Attica and the Peloponnese. The Ionian Islands are situated on the western border of Greece in the Ionian Sea. There are several island groups in the Aegean Sea on the eastern side of Greece: Evoia and the Sporades, the Saronic Islands (grouped with Attica), the Cyclades, the Eastern Aegean Islands, the Dodecanese and Crete (Fig. 1). Results Species composition The spiders established in the caves of Greece (Main- land and Insular part) are represented by 109 species, included in 52 genera and 25 families: Ctenizidae - 1, Filistatidae - 1, Sicariidae — 1, Scytodidae - 1, Leptonetidae - 9, Pholcidae - 10, Segestriidae - 3, Dysderidae - 12, Oonopidae - 1, Mimetidae - 1, Ere- 24 C. Deltshev LIV1KÖ8 f>ELAQOS Figure 1 : Map of different geographical regions in Greece. sidae - 1, Uloboridae - 1, Nesticidae - 4, Theridiidae - 4, Anapidae - 1, Linyphiidae - 16,Tetragnathidae - 4, Araneidae - 1, Lycosidae - 1, Agelenidae — 21, Amaurobiidae — 4, Gnaphosidae — 6, Philodromidae - 1, Thomisidae - 2, Salticidae - 2 (Table 1). One species is new for the Greek spider fauna: Porrhomma convexum (Westring, 1861) (marked in the list with *), a spider with a Holarctic distribution and widespread in European caves. It is also well represented in the caves of the Balkan Penisula - and not only estab- lished in the caves of Croatia, Romania and Turkey (DELTSHEV 2008). The number of species is high and represents about 13 % of the Greek spiders. This is also evident from a comparison with the number of cave-dwelling spiders recorded from the other coun- tries of the Balkan Peninsula: Bulgaria - 99, Croatia - 63, Serbia - 59, Bosnia and Herzegovina - 52, Macedonia - 44, Montenegro - 44, Slovenia - 43, Al- bania - 10, Turkey - 8, and Romania - 4 (DELTSHEV 2008). The established number of species, however, depends not only on the size of the regions, but also on the degree of exploration. The most characteristic are the families: Leptonetidae (8.2 %), Pholcidae (9.2 %), Dysderidae (11 %), Linyphiidae (14.6 %), and Agelenidae (19.2 %). The families with largest number of anophthalmic species are Leptonetidae (6), Dysderidae (3), Nesticidae (2), and Linyphiidae (1). The genera with the largest number of species are Tegenaria (8), Lepthyphantes (6), Harpactea (5), and Histopona (5). The species are distributed in caves belonging to the geographic territories of Greece as follows: Thrace - 8 species, Macedonia - 18, Epirus - 1, Thessaly - 6, Central Greece - 3, te Attiki- Saronic Islands - 24, the Peloponnese - 15, Evoia-Voroies Sporades - 1, the Eastern Aegean Islands - 5, the Cyclades - 7, the Dodecanese - 6, the Ionian Islands - 23, Crete - 47 (Table 1). We also have to emphasise that the degree of exploration in these territories is not equal: the territories of Evoia- Sporades, the Cyclades, the Dodecanese and Central Greece are less explored. Cave-dwelling spiders can be categorised into four ecological groups (SKET 2008): • troglobites: species limited to a life cycle in caves. Often they show a suite of characters, associated with their adaptation to subterranean life: loss of pigment, loss of eyes and elongation of appenda- ges. • eutroglophiles: species which can live their entire life in caves, but also occur in other environments. • subtroglophiles: species which utilise caves, but must leave the caves to complete their life cycle. • trogloxenes: species which occur underground sporadically. The largest fraction of troglobite species was encoun- tered mainly in the caves of the territories of Crete -15 (5 anophthalmic), the Ionian Islands - 4, Thrace - 2 (2 anophthalmic), the Attiki- Saronic Islands 2 (2 anophthalmic), the Peloponnese -2(1 anophthalmic), and Macedonia, Thessaly and the Cyclades each by 2 species (Table 1). All troglobites are endemics for the territory of Greece or the Balkan Peninsula. Very important is the presence of eutroglophiles (35 species), because together with troglobites (29 species), they can be considered as dependent faunistic elements of caves. The largest number of species is established in the caves of Crete (14 species), the Io- nian Islands (11 species), the Attiki- Saronic Islands (9 species), Macedonia (8 species), and the Peloponnese (8 species). Here, the endemics are represented by 14 species (35 %). A present day example of active subterranean colonisation and cave penetration are the species Lepthyphantes centromeroides and Pallidup- hantes spelaeorum, widespread in the Balkan Peninsula (DEELEMAN-REINHOLD 1978). Here, the species Palliduphantes istrianus should also be included. Cove-dwelling spiders of Greece 25 -C Q. 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J5 "g E £-8 .2 c C LU O I 'S u u O TO Q E 03 03 a= -O 2 c C LU (LI I c < c HI O HI U Distribution:THR - Thrace; MA - Macedonia; EP - Epirus; THE - Thessaly; CGR - Central Greece; ASI - Attiki-Saronic Islands; PE - Peloponnese; ESI - Evia-Sporades; EAI - Eastern Aegean Islands; CY - Cyclades; DO - Dodecanese; 10 - Ionian Islands; CR - Crete. CR X X X X X X X X x ; X X O X X X X X DO X X X X Ö EAI X ESI PE X X X ASI X X X X CGR THE X EP X X MA X X X THR X | O Z ECR £ COS X EPE ECR ECY EAS EAS ETHR ECR EEP EETI s EEAI ECR ECR EKA ECR CAT & stph stph stph tba tba tba tba tba £ tba Cl on ON A u rt q=! 0 0 ""cs H R R R 1 .s g R R s ON On 43 O «! CG O q <18 Ul XI H > ’’'R O R R s Amaurobius strandi Charitonov, 1937 Gnaphosidae Drassodes lapidosus (Walckenaer, 1802) Nomisia ripariensis (Thorell, 1871) Pterotricha lentiginosa (C.L. Koch, 1837) Zelotes clivicola (L. Koch, 1870) Zelotes femellus (L. Koch, 1866) Zelotes oblongus (C.L. Koch, 1833) Sparassidae Eusparassus walckenaeri (Audouin, 1826) Philodromidae Philodromus collinus C.L. Koch, 1835 Thomisidae Synema glob o sum (Fabricius, 1775) oo . An exact provenance for the new fossil specimen is not recorded, but much of the cur- rently available amber derives from the Kaliningrad region on the Baltic coast of Russia. An exact age for amber is difficult to determine objectively, but Baltic amber is traditionally dated at Paleogene (Eocene), or about 45-50 Ma. For comparative purposes, a scanning electron mi- crograph (SEM) photograph of the genital region of a recent cyphophthalmid harvestman is also included here (Fig. 4): 1 female Cyphophthalmus duricorius Joseph, 1868: Slovenia, 4 km NW from Postojna, Pivka Jama camp site, under stones in the old pine forest (N 45°48’18.28" E 14°12T6.01", 550-579 m a.s.l.), 16.IX.1989, leg. and det. P. G. MITOV. The SEM study was made at 10-20 kV with a Philips 515 machine. The specimen was sputter-coated with a 300-400 Ä gold layer. Further SEMs of the American species Siro exilis Hoffman, 1963 were kindly provided by Günther Raspotnik (Figs. 7-8): 1 female Siro exilis Hoffman, 1963: West Virginia, Randolph Co., Monongahela National Forest, nr. Bowden; Otter Creek Wilderness trailhead near Alpena Gap, deep litter of mixed riparian forest (red maple, yellow birch, eastern hemlock, white spruce) with dense understory of rhododendron, 930 m a.s.l., 38° 56.505’ N, 79° 40.084’ W, 13.VI.2006, leg. and det. Roy A. Norton. The specimen was air-dried, mounted on an alu- minium stub and sputtered with gold (AGAR sput- tercoater, Gröpl, Tulin, Austria). The SEM study was made at 20 kV with a Philips XL30 ESEM (Philips/ Siro in Baltic amber 49 FEI, Vienna, Austria) at high vacuum mode. Order Opiliones Sundevall, 1833 Suborder Cyphophthalmi Simon, 1879 Family Sironidae Simon, 1879 Genus Siro Latreille, 1796 Siro balticus sp. nov. Material: Holotype $ and only known specimen, JÖRG Wunderlich collection, F2147/BB/CJW. From Baltic am- ber, exact locality not recorded; Paleogene, Eocene. Diagnosis: Relatively large (length 2.34 mm) fossil Siro species, specifically without the projecting rear end typical of modern European forms, and with body proportions differing from those in the - probably more closely related - extant North American species (see Remarks for details). Derivation of name: From Baltic amber and the Baltic region; the source of this material. Description: Partially complete female specimen preserved in both dorsal (Figs. 1, 5) and ventral view (Figs. 2, 6). All measurements in mm. Body oval; pale brown in colour within the amber, but with darker patches across the body; total length 2.34; maximum prosomal width behind the ozophores 1.33; maximum opisthosomal width 1.34. Lengthrwidth ratio 1.75. Distance between front of scutum (i.e. anterior mar- gin of prosoma) to an imaginary line connecting the anterior (front) bases of ozophores 0.23; total width across (and including) ozophores 1.09. Entire body with pustulate ornament of small, rounded to oval tubercles, generally larger in anterior body regions and smaller posteriorly and on the leg trochanters. Eyes absent. Ozophores conical to slightly pointed and angular, dorsolaterally prominent on the scutum in the type 2 position sensu JUBERTHIE (1970, Fig. 2). Exact position of the ozopore itself - i.e. the opening of the repugnatorial gland - difficult to re- solve, but maybe terminal. Length of ozophores 0.13; width at base 0.22. Slight bulge to body immediately behind the ozophores. Frontal ridge of scutum (i.e. anterior margin of carapace) slightly recurved. Sulcus beginning immediately behind the ozophores curves down towards the midline and defines a posteriorly deeply recurved anterior area of the body; length 0.84 on the midline. Area behind it incorporates the bulge in the body laterally and is also recurved at the midline, length here 0.12. This region followed by eight, quite clearly defined, tergites all with essentially straight posterior margins. Gaps between tergites lack tuberculation. Anterior four tergites longer, lengths c. 0.2; posterior three are notably shorter, lengths 0.13, 0.11 and 0.12 respectively. Posterior end of opistho- soma bluntly rounded to slightly angular in dorsal view. Ventral prosomal complex incomplete. Chelicerae, pedipalps, coxae of legs 1-2 and all walking legs be- yond the trochanter absent. Conceivably, the missing coxae (I— II, or perhaps II only) were free, i.e. not fused to coxae III— IV; which could explain why the breakage point in the fossil lies between the second and third coxae. Third coxae triangular, coming to a point im- mediately above the thoracic complex, but not quite reaching the midline. Third trochanter somewhat Figures 3-4: Details of the thoracic complex region. - 3: Siro balticus sp. nov.; holotype (F2147/BB/CJW).The rounded spiracle (sp) with denticles inside the lumen is a convin- cing character of the genus Siro Latreille, 1796. Coxae numbered. The area between coxal lobes III and IV (i.e. around the coxal pores) has an obtuse, ca. 90°, angle (outlined by black bars: arrowed); another Siro character. Scale bar equals 200 pm. - 4: Same region in a modern cy- phophthalmid, Cyphophthalmus duricorius Joseph, 1 868, SEM. Scale bars equal 1 00 pm. In Cyphophthalmus, by contrast, the area between coxal lobes III and IV has a very acute angle (outlined by black bars: arrowed). 50 J. A. Dunlop & P. G. Mitov Figures 5-6: Interpretative camera lucida drawings of the specimen shown in Figs. 1-2.5: Dorsal view. - 6: Ventral view. Abbrevi- ations: ca = expected site of corona analis, cp = carapace, cx = coxa, oz = ozophore, sp = spiracle, st = sternites 2+3 (subse- quent sternites numbered successively), tc = thoracic complex, tg 1 = tergite 1 (subsequent tergites numbered successively), tr = trochanter. Scale bar equals 1 .0 mm. rounded and cup-shaped, but details equivocal. Fourth coxae much larger than the third and more quadratic in shape. Distinct suture line originates near the an- terolateral corners of the thoracic complex and curves towards the posterior margin of the coxae, bisecting it about a third of the way along its length towards the distal margin. Fourth trochanter emerges from the posterior part of the fourth coxae and is associ- ated with a small, but distinct indent into the coxal margin. Trochanter 4 tubular, longer than wide and with a slightly oval outline, widening distally, length 0.28. Thoracic complex with the outline of an inverted subtriangular structure, dominated centrally by large genital opening (or gonostome) of the female type (Fig. 3). Gonostome anteriorly with semicircular outline; width 0.17, length 0.16; no genital structures visible within this opening. Thoracic complex flanked laterally by sutured-off part of the fourth coxae, pos- teriorly by the second sternite and anteriorly (in part) by the third coxae. More anterior elements missing. Thoracic complex divided anteriorly by a short mid- line sulcus. Either side of this, i.e. on the anterolateral margins of the gonostome, are areas surrounding the coxal pores (= orifices of the coxal glands). Pores themselves clearly visible as small, but distinct holes, diameter 0.0125, lying between coxal lobes III and IV. Sternites two and three apparently fused into a single, large subtriangular plate, projecting anteri- orly between the leg 4 coxae with a procurved and bluntly rounded anterior margin pointing towards the thoracic complex. Tuberculation here heavier, with larger and more oval tubercles. Spiracles present as prominent, round to oval-shaped elements (maxi- mum diameter 0.16) in a lateral position immediately behind the fourth coxae; ring of tiny denticles present within the lumen, expressing - at least on the right side - a distinct invagination towards the centre of the spiracle. No obvious sternal pores, or other structures, located on the midline between the spiracles. Five sternal elements (presumably sternites 4-8) preserved behind the large, spiracle-bearing sclerite. Sternites 4—7 decrease successively in length: i.e. Siro in Baltic amber 51 Figures 7-8: Comparative SEM images of a female of the Recent species Siroexilis Hoffman, 1 963 from the eastern USA. These eastern Siro species are thought to be closer to the European fauna than those of the western USA (cf. Shear 1 980). 7: Overview. Scale bar equals 1 .0 mm. Note the overall similarity to the fossil Siro species in terms of the absence of a projecting rear end (arrowed); a feature usually seen in the (modern) European forms. - 8: Detail of spiracles and thoracic complex region. Scale bar equals 200 pm. 0. 20, 0.19, 0.16 and 0.10. Sternite 8 forms the anterior border of a fairly large oval opening, width 0.32, which in life would probably have contained the corona analis. Posterior margin of opistho- soma somewhat blunt and rectangular in ventral view; 1. e. not smoothly rounded, but details of any specific sclerites here difficult to resolve. In general, tergites slightly wider than sternites and marginal overlap from the overlying tergites can be seen ventrally at the lateral edges of the body. In cyphophthalmids the form of the gonostome dif- fers between the sexes and this specimen clearly has the female type. Since the gonostome is preserved open (Fig. 3), it must be an adult and is thus clearly a mor- tality rather than a moult. Intuitively, the new fossil with its distribution in the Eocene of northern Europe is likely to be a member of Sironidae (see e.g. GlRIBET (2000: Fig. 2) or BOYER et al. (2007: Fig. 1) for distribution maps) because none of the other extant cyphophthalmid families occur in this region today. The fossil is unquestionably modern in appearance and affinities with a number of extant genera need to be considered: i.e. Siro Latreille, 1796, Cyphophthalmus Joseph, 1868 (re-established by BOYER et al. 2005 for a Balkan radiation; see also KARAMAN 2009 and MURIENNE et al. 2010) and the four Iberian genera most recently investigated by MURIENNE 8c GlRIBET (2009). These taxa are mor- phologically rather conservative and appear similar in overall habitus, while a number of taxonomically important details are missing from the new fossil which hinders its unequivocal generic assignment. KARAMAN (2009: table 1) established a series of characters useful in separating Siro from Cyphoph- thalmus. Some of these, such as the number of paired, movable fingers associated with the spermatopositor, the number of anal glands, and the shape of the ad- enostyle of the tarsal gland apophyses, only occur in males and are thus unhelpful in placing this female fossil. An alternative character (cf. KARAMAN 2009: p. 262) which would have been useful is the shape and structure of the prosomal complex (i.e. the coxal lobes of legs II, immediately in front of the genital opening), but unfortunately this feature is equivocal in the fossil. A character discussed by KARAMAN (2009) which is preserved is the form of the spiracles and this does offers useful data about the animal’s affinities. In Cy- phophthalmus the spiracles are semicircular, each with a conical cuticular projection on its posterior margin. This is not seen in the fossil, which by contrast (Fig. 3) seems to share a character seen, so far, only in Siro and which consists of a more rounded spiracle (of circular type, sensu GlRIBET &, BOYER 2002), with denticles inside the lumen (cf. BlVORT &, GlRIBET 2004: Fig. 36c, KARAMAN 2009: Figs. 2A-B). For this reason we are confident in our generic assignment of this fossil to Siro. In support of this hypothesis we also note that the form and width of the area between coxal lobes III and IV is very different between Siro and Cyphophthalmus species. Specifically, the endites (= coxapophysis) of coxae III and IV form either an area (around the coxal pores) with a very acute angle 52 J. A. Dunlop & P. G. Mitov (in Cyphophthalmus’. Fig. 4) or a right/obtuse angle (in Siro: Fig. 3); thus in this context the amber fossil more closely resembles female Siro species. For compara- tive figures of Siro see e.g. RAFALSKI (1958: Fig. 7) for Siro carpaticus Rafalski, 1956, JUBERTHIE (1967: Fig. 7) for Siro rubens Latreille, 1804, and NOVAK Sc GlRIBET (2006: Figs. 5, 7, 12) for Siro crassus Novak Sc Giribet, 2006. For various Cyphophthalmus species we refer also to ElSENBEIS dcWlCHARD (1987: Plate 27, d) (sub Siro duricorius ), MlTOV (1994, Fig. 23) (sub Siro beschkovi) or KARAMAN (2008, 2009). Although incomplete, we feel able to assign this new fossil to its own species. Of particular interest is the posterior end of the body. Females of modern European Siro species typically show a projecting rear end (see e.g. figures in JUBERTHIE 1970), whereas the Baltic amber example has a more smoothly rounded back end which thus resembles, the North American Siro species (e.g. NEWELL 1943, 1947, HOFFMAN 1963, SHEAR 1980; see also Fig. 7). Conceivably our fossil was part of this (formally Laurasian?) lineage (see Discussion). At 2.34 mm in body length the new fossil sironid is somewhat larger than typical American Siro species. Body lengths of females vary from 1.10 mm (in Siro sonoma Shear, 1980) to 2.08 mm (in Siro exilis Hoffman, 1963 - the largest modern American sironid); see also NEWELL (1943, 1947). The largest European sironid is the epigean Siro crassus Novak Sc Giribet, 2006 in which females reach 2.40- 2.61 mm. Direct comparison with the previously described European amber species, Siro platypedibus, is difficult given that the fossils are preserved in completely dif- ferent orientations. The coxo-genital and anal region of the younger Bitterfeld fossil cannot be resolved. The originally proposed diagnosis was based on its flattened legs; a character which cannot be tested in the Baltic form. In any case this has recently been challenged as a possible artefact by KARAMAN (2009). He noted that limb flattening can occur while making preparations of extant material in various mounting media, and that the amber- forming resin as a similarly concentrated viscose medium could have induced comparable effects. Our new fossil and S. platypedibus are, at ca. 2 mm long, similar in overall size. While it may be possible to draw some comparisons based on the profile of the body sculpture in these respective fossils, we currently have little data to argue either for or against the conspecificity of these extinct taxa from the different amber faunas. Discussion As noted above, GlRIBET et al. (2010) inferred a basal divergence time for modern cyphophthalmid lineages perhaps as far back as the Carboniferous (ca. 345 Ma); at which time Europe and North America were part of the single palaeocontinent Laurasia. Indeed, SHEAR (1980, p. 4) commented on the strong simi- larities between some of the (eastern) N. American cyphophthalmids and the European fauna: “... the original divergence took place between the western species and S[iro] exilis (Figs. 7 -8) plus the European forms. The movement of North America away from Europe and Africa, which resulted in the opening of the present Atlantic Ocean, may account for the separation of S. exilis from its European relatives.” The hypothesis that the North American Siro species are not monophyletic was again supported in a recent study by GlRIBET Sc SHEAR (2010), who further re- covered European affinities for some American taxa under some parameters of analysis and reiterated the idea that Siro may be a very ancient genus. What this implies is, first, that there is no fun- damental objection to recovering an American-like Siro species from Baltic amber. This lineage may well have been originally distributed more widely across the Palaearctic, and was present in north-central Europe during the presumably warmer conditions associated with the Baltic amber forest. A possible parallel example of this would be the eupnoid har- vestman genus Caddo Banks, 1892 which was also present in Europe during the early to mid-Paleogene (DUNLOP Sc Mitov 2009, and references therein), but is absent from the modern European fauna and yet still found today in North America. Second, the opening of the Atlantic began during in the Triassic (ca. 200-250 Ma), and this, in turn, offers a minimum divergence time, based on geological evidence, for a common ancestor of these similar-looking American and European Siro species; the latter also including Siro balticus. To reiterate, North America yields Siro (cf. GIRI- BET Sc SHEAR 2010), while both Siro and Cyphoph- thalmus occur in Europe. In recent studies Cyphoph- thalmus was treated as a specifically Balkan radiation (BOYER et al. 2005, KARAMAN 2009); although we should note that Siro has also been reported at least as far south as Slovenia (NOVAK Sc GlRIBET 2006). Recent work by MURIENNE et al. (2010) inferred a late Cretaceous radiation for Cyphophthalmus of ca. Siro in Baltic amber 53 94 Ma and attempted to tie the explosive evolution of its numerous endemic species into the wider geo- logical development of the Balkan Peninsula. Slow rates of evolution for cyphopthalmids in general were postulated by SHEAR (1980) - see also GlRIBET & SHEAR (2010) - and the modern appearance of this amber fossil (cf. Figs. 1-2 and 7-8) compared to extant forms tends to support this supposition: at least for Siro. Unfortunately, we still lack Palaeozoic cyphoph- thalmids which should date from the early phase of their evolution as predicted by molecular analyses. Acknowledgments We thank Jörg Wunderlich (Hirschberg) for making this material available for study and preparing the specimen from a larger block of resin, as well as Ivo Karaman (Novi Sad) and Gonzalo Giribet (Harvard) for valuable comments and to Gonzalo for providing pre-publication copies of relevant papers. Günter Raspotnik (Graz) kindly provided SEM images of Siro exilis material, which in turn was made available by Dr. Roy Norton (SUNY-ESF, Syracuse, New York). Finally, the reviewers are also thanked for helpful remarks. References BANKS N. (1892): A new genus of Phalangiidae. - Pro- ceedings of the Entomological Society of Washington 2 (2): 249-251 BlVORT B.L. DE & G. 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JR (2008): Palaeosiro burmanicum n. gen., n. sp., a fossil Cyphophthalmi (Arachnida: Opiliones: Sironidae) in Early Cretaceous Burmese amber. In: MAKAROV S.E. & R.N. DimITRIJEVIC (eds): Advances in Arachnology and Developmental Biology. Papers dedicated to Prof. Dr. Bozidar Curcic. Inst Zool, Bel- grade; BAS, Sofia; Fac Life Sei, Vienna; SASA, Belgrade & UNESCO MAB Serbia, Vienna, Belgrade, Sofia. Monographs 12. pp. 267-274 RAFALSKI J. (1958): A description of Siro carpaticus sp. n. with remarks on the morphology and systematics of the Cyphophthalmi (Opiliones). - Acta Zoologica Cracoviensia 2: 521-556 SHEAR W.A. (1980): A review of the Cyphophthalmi of the United States and Mexico, with a proposed reclassificati- on of the Suborder (Arachnida, Opiliones). - American Museum Novitates 2705: 1-34 SIMON E. (1879): Les Arachnides de France VII. Con- tenant les ordres des Chernetes, Scorpiones et Opiliones. Roret, Paris. 332 pp. SUNDEVALL J.C. (1833): Conspectus Arachnidium. C. F. Beding, Londini Gothorum. 39 pp. Arachnologische Mitteilungen 40:55-64 Nuremberg, January 201 1 Spiders in a hostile world (Arachnoidea, Araneae) Peter J. van Helsdingen doi:10.5431/aramit4007 Abstract: Spiders are powerful predators, but the threats confronting them are numerous. A survey is presented of the many different arthropods which waylay spiders in various ways. Some food-specialists among spiders feed exclusively on spiders. Kleptoparasites are found among spiders as well as among Mecoptera, Diptera, Lepidoptera, and Heteroptera. Predators are found within spiders' own population (cannibalism), among other spider species (araneophagy), and among different species of Heteroptera, Odonata, and Hymenoptera. Parasitoids are found in the orders Hymenoptera and Diptera. The largest insect order, Coleoptera, comprises a few species among the Carabidae which feed on spiders, but beetles are not represented among the kleptoparasites or parasitoids. Key words: aggressive mimicry, araneophagy, cannibalism, kleptoparasitism, parasitoid SPIDER EATS SPIDER Regular prey Cannibalism Food specialists KLEPTOPARASITES / COMMENSALISM prey of spider eaten by others SPIDER PARASITES /PARASITOIDS Egg parasites Parasitoids proper INSECTS AS PREDATORS ON SPIDERS Figure 1:The spider in its environmental web. Spiders are successful predators with important tools for prey capture, viz, venom, diverse types of silk for snaring and wrapping, and speed. But spiders are prey for other organ- isms as well. This paper presents a survey of all the threats spiders have to face from other arthropods (excluding mites), based on data from the literature and my own observations. Spiders are often defenceless against the attacks of others, just as most spider victims are defenceless against the spiders and their methods of capturing prey. In this article I look at the spider in its environmental context from four angles: when it is preyed on by other spiders; when it is the victim of kleptoparasites (spiders and insects) which steal food from its web; when it is preyed on by other invertebrates (other than spiders); and when the individual spider falls victim to parasitoids. The subjects are dealt with in this order (Fig. 1). The present study is restricted to arthropods. Many of the relationships referred in this article come from subtropical and tropical regions where biodiversity is much higher and food specialization apparently has a better chance to develop. Neverthe- less, the temperate regions contribute to the ecological interactions dealt with here, too. Peter J. van HELSDINGEN, European Invertebrate Survey- Nederland, Leiden, Netherlands. E-Mail: helsdingen@naturalis.nl submitted: 27.11 .2009, accepted: 27.4.20 1 0; online: 10.1.2011 Spiders and their prey The regular prey of spiders consists of insects and other invertebrates, including other spiders. The methods employed are hunting, by sight or other senses, or catching with a web which has sticky threads or is made to entangle the prey. Prey can be wrapped up in silk or held with the legs and chelicerae but as a rule are killed with venom and digested externally by regurgitating digestive fluid over the prey after which the resulting fluid is sucked up. Walking, stalking, waiting, and wrapping are terms which fit. Although the bulk of spider prey consists of insects, preying on spiders is not an exception. A general phenomenon in spiders is cannibal- ism which can already take place inside the egg batch or within the population. SAMU et al. (1999) demonstrated, for Pardosa agrestis (Westring), that when food is scarce preying on individuals in the 56 P.J.v.Helsdingen same population becomes common. Usually not all specimens in a population are of the same age and size as they come from different egg batches and larger individuals then tend to eat the smaller members of the population. Some spiders have distinct food preferences. It is common knowledge that Dysdera species specialize on woodlice (Isopoda), bolas spi- ders (Araneidae: Mastophora , Ordgarius ) attract male moths with chemical compounds which resemble the moth’s pheromones, while Zodarion species feed on ants. Mimetidae and some Salticidae feed exclusively on other spiders, while some Pholcidae hunt other spiders as well (see section “Insects and spiders as predators on spiders”). Kleptoparasites Kleptoparasitism is found in spiders as well as in many insect orders. Prey in spider webs are apparently an easily obtainable source of food once one has devel- oped a method of getting at it without alarming the owner of the web and becoming its prey. Examples are summed up by order. Araneae Kleptoparasites “steal” the prey of the spider from the spider’s web. It is debatable if this might be called commensalism, which is defined as using the food of the host species without causing any harm or negative influence. In many instances, e.g. with web-building species, a spider obtains food by locating the prey in the web, biting it and injecting venom, regurgitating digestive fluid over the prey, and wrapping the victim. All these actions are energy investments made by the host spider and thus are of negative influence on its energy balance, however slight. A kleptoparasite prof- its from the host spider’s energy investment without giving anything in return. True kleptoparasites are able to walk along sticky silk without being trapped. This is not so surprising for kleptoparasitic spiders in which the ability to walk on webs is common in many groups. For other invertebrates this quality must have evolved. Kleptoparasitism occurs in a large number of spider families (Anapidae, Dictynidae, Eresidae, Mys- menidae, Oonopidae, Salticidae, Sparassidae, Sym- phytognathidae, Theridiidae, and Uloboridae) (for a summary and literature references, see AGNARSSON 2002). Ar gyro des species (Theridiidae) are the best- known examples of kleptoparasitism and are found with orb web building Araneidae and Tetragnathidae, and social and subsocial spiders with large communal webs, such as Anelosimus (AGNARSSON 2003 ).Argy- rodes steals the prey and may carry it off to the margin of the web (AGNARSSON 2003). Larger webs, such as those of Nephila , often catch more small prey than the owner needs. Small prey specimens just stick to the spiral threads and are not even bitten or wrapped and Argyrodes often eats from such neglected prey. The habit runs through the whole genus. Argyrodes bryantae Exline Sc Levi was found as kleptoparasite in the webs of Tengella radiata (Kulczynski) (EBERHARD et al. 1993). Argyrodes antipodianus O.P.-Cambridge shows a transition to araneophagy (WHITEHOUSE 1986). Social Uloboridae have been observed as solitary kleptoparasites in the webs of other spiders. Philoponella republicana (Simon) is known to occur in webs of Cyrtophora nympha Simon (ROBINSON 1977) and also in Anelosimus webs in French Guiana (LOPEZ 1987). P. tingena (Chamberlin Sc Ivie) has been recorded from webs of Nephila clavipes L. and “Achaearanea spec.” (OPELL 1979). Two species of Mysmenopsis (Mysmenidae) lead a kleptoparasitic life in webs of Tengella radiata (Kulczynski) (Tengel- lidae) (EBERHARD et al. 1993). Both have a broad host spectrum. M. tegellacompta Platnick is found in webs of Tengella radiata as well as in a diplurid web and an agelenid web (species not established), while M. dipluramigo Platnick & Shadab has been found in webs of T radiata , a ctenid web and a pisaurid web (Eberhard et al. 1993). Mecoptera Scorpionflies (Mecoptera) of the family Panorpidae have been observed to land directly onto a spider web or walk into it from the surrounding vegetation and eat from the prey they find there. When the owner of the web approaches the scorpion fly it may ward the spider off by hitting it with the thick end of its abdomen. Scorpionflies have been found in webs of Agelenidae, Tetragnathidae, Theridiidae, and Araneidae (THORNHILL 1975). Diptera Diptera also have their kleptoparasitic species. The gall midge Didactylomyia longimana (Nematocera, Cecidomyiidae) was detected as a very common kleptoparasite in orb webs of Nephila clavipes (Tet- ragnathidae), Argiope aurantia Lucas, Mastophora bisaccata (Emerton), Eriophora ravilla (C.L. Koch), and Scoloderus cordatus (Taczanowski) (all Araneidae) (SlVINSKI Sc STOWE 1980). The females were found on the prey of the spider, while the males were hang- ing inactively in the web. Among the biting midges 57 Spiders in a hostile world (Ceratopogonidae) a number of species associated with spider webs have also been found (SlVINSKI &, Stowe 1980). Among the suborder Brachycera there are several families which comprise species with kleptoparasitic behaviour. Examples of species of Chloropidae and Milichiidae (both of acalyptrate fly families) are listed by NENTWIG (1985). Desmometopa species (Milichi- idae) were observed to feed on the prey (honey bee, Apis mellifera) of a lynx spider (Oxyopidae, probably Oxyopes heterophthalmus (Latreille)) and also of other spiders (RICHARDS 1953; ROBINSON & ROBINSON 1977): Phyllomyza spec. (Milichiidae) on prey of Nephila clavipes (L.), Conioscinella spec, in the web of Argiope argentata (Fabricius). Best known, relatively, are the members of the genus Microphor (Micropho- ridae) with eight species and eight further species of related genera in Europe (PAPE 2010). Most of the Microphor species, if not all, are seen in association with spiders and their prey. They usually sit on the prey item while the spider is sucking on it. CHVALA (1986) stressed that MACQUART (1827) previously noted an association with spiders. Megaselia scalaris Loew (Phoridae) was found on the webs of Tengella radiata (Kulczynski) (EBERHARD et al. 1993). Lepidoptera Caterpillars of some lepidopteran families are known to feed on spider prey. POCOCK (1903) states that the larvae of Batrachedra stegodyphobius Walsingham (Batrachedridae) live in the communal web of a Ste- godyphus species (Eresidae) in South Africa. Accord- ing to Pocock pupation of the noctuid moth occurs in the spider web and the adult moths are seen fluttering about the web. ROBINSON (1977) reported the larvae of Neopalthis madates Druce (Noctuidae) living in the communal web of Anelosimus eximius Simon in Panama. The caterpillars of Tallula watsoni Barnes &c McDunnough (Lepidoptera, Pyralidae) seem to live exclusively in the webs of Anelosimus studiosus (Hentz), where they eat dead and living leaves from the sup- porting tree or shrubs and attack or eat the spiders (DEYRUP et al. 2004). All these species are not only kleptoparasites but also inquilines which live in the web permanently. Heteroptera Among the bugs (Hemiptera-Heteroptera) there exist kleptoparasitic specialists in several families. Arachnocoris (Nabidae) is a genus which occurs with nine species (2-5 mm) in the Neotropical Region (Lopez-Moncet 1997). An upside-down position in the spider web is typical for this taxon. The different species were found in webs of Araneidae {Micrathena) (with sticky silk),Theridiidae ( Tidarren fordum (Key- serling) (= Tidarren sisyphoides (Wzlckenzer)), Anelosi- mus eximius (Keyserling)) .(sticky) as well as those of Pholcidae ( Physocyclus sp.) (non-sticky silk). Strangely, specimens of Arachnocoris trinitatis Bergroth, one of the best studied species of the genus, are usually found in empty webs of the pholcid Mesobolivar aurantiacus (Mello-Leitao). It is hypothesized that the bug uses the web for catching prey and finding a mate (SEW- LAL &c STARR 2008). It is not clear whether the bug emptied the web by capturing and devouring the spider or by chasing it away. The genus Ranzovius (Miridae) comprises at least four species which are associated with spiders (WHEELER ScMcCAFFREY 1984). All specimens in this genus are very small (2-2.5 mm) and are found in orb webs as well as in sheet webs. R.fennahi Carvalho lives in large webs of the social Anelosimus eximius (Keyserling) while R. contubernalis Henry occurs in the communal webs of the social Anelosimus studiosus (Hentz). In the large spatial webs of the latter a lot of prey remnants are scattered throughout the web which attract pyralid larvae, cockroaches and ants which behave as scavengers. R. californicus (Van Duzee) consumes prey in the webs of Hololena curt a (McCook) (Agelenidae). R. agelenopsis Henry can be found in high numbers in the webs of the com- mon Agelenopsis pennsylvanica (C.L. Koch) where R. contubernalis can be found as well. Agelenopsis species are often common in shrubs and hedges. The webs of other common spider species in the same habitat, such as the linyphiid Frontinella pyramitela (Walckenaer) and various araneids (probably Zygiella species) were checked for the presence of Ranzovius but none were found (WHEELER &McCAFFREY 1984). Apparently Ranzovius prefers Anelosimus and Agelenopsis for its kleptoparasitic practises. In the Reduviidae the relatively common spe- cies Reduvius personatus L. has been found in webs of “house spiders”, a name used in the U.S.A. for Parasteatoda tepidariorum (C.L. Koch) (AMYOT & SERVILLE 1843). There are a number of striking examples of kleptoparasites within the subfamily Emesinae (Reduviidae), viz. the genera Eugubinus , Ploiaria , Emesa , Empicoris , and Stenolemus . They all feed on the prey of the spiders the webs of which they invade. In the case of Eugubinus araneus Distant this was a theridiid (in Bombay) (DISTANT 1904), while 58 P.J. v.Helsdingen E. intrudans Distant and E. reticolus Distant were seen in webs of Cyrtophora cicatrosa (Stoliczka) (India) (DISTANT 1915). Stenolemus represents a transition to araneophagy. More web-invading heteropteran species can be found in the Anthocoridae, viz., Cardiastethus inqui- linus China & Myers in South Australia, in the web of a gregarious oxyopid (CHINA & MYERS 1929). Species of the Plokiophilidae, with the genera Plokiophila , Plokiophiloides, Lipokophila , and Embi- ophila are found in the webs of Dipluridae andTengel- lidae in the southern hemisphere (McGAVIN 1993). The very small Plokiophila cubana (China &c Myers) occurs on the webs of Diplura macrura (C.L. Koch) (Dipluridae) in Cuba. Lipokophila eberhardi Schuh and L. tengella Schuh were found on the webs of Tengella radiata (Kulczynski) (Tengellidae) (EBERHARD et al. 1993). According to CARAYON (1974) Plokiophilidae spend their whole life in the webs of spiders. They live there from egg stage to death. The egg is deposited on a thread in the spider’s web and the young bug hatches immediately. Plokiophiloides asolen Carayon lives in webs of the social Agelena consociata Denis, while P balachowskyi Carayon lives in webs of the social Agelena republicana Darchen. P biforis Carayon was collected from webs of Lathrothele catamita (Simon) (Dipluridae). The reduviids Themonocoris bambesanus Carayon and two Anthocoridae ( Cardiastethus affinis Poppius and C. lateralis Poppius) live there too. When reduviid bugs live in a spider web these are free of kleptoparasitic spiders (LOPEZ-MONCET 1997). Possibly bugs live in spider webs because they are safe there from ants which are everywhere but hardly ever enter spider webs (LOPEZ-MONCET 1997). Insects and spiders as predators on spiders Araneae For spiders any other spider is potential prey when it falls within the limits of its range of possibilities (size, danger, risk, defence of prey, etc.). Some spiders have made a habit of eating spiders of other species, a habit called “araneophagy”. For cannibalism (occasional eat- ing of specimens of the own species), see above. Mimetidae are specialized predators on other spiders which they attack in the web of the prey by producing signals resembling those of an entangled insect or a potential mate wanting to pair, so-called “aggressive mimicry” (JACKSON & WHITEHOUSE 1986). From observations made by Bristowe (1958) it is clear that Mimetidae have very strong, paraly- zing venom. Salticidae of the subfamily Spartaeninae are specialized in capturing spiders in their webs by stealthy approach combined with aggressive mimicry. The genus Portia is the best known genus (five species) which exploits this type of prey capture, but there are three other genera which show this type of behaviour as well, viz. Brettus (two species), Gelotia (one species), and Cyrba (two species) (WANLESS 1984). All these salticids share the characters of good vision with the ability to walk over sticky and non-sticky webs (JACK- SON 1986). The pholcid Pholcus phalangioidesvz ntures into the webs of other spiders and overwhelms the owner (JACKSON & BRASSINGTON 1987). Some Palpimanidae invade the web and lure the host out (HENSCHEL et al. 1992). There are many examples of insects which are predators of spiders. The following examples are listed by order. Heteroptera Stenolemus species (Reduviidae, Emesinae) can be found in the surroundings of the spider webs which they penetrate to prey on the spider. Stenolemus are large, up to 1 cm overall body length, with long, thin legs. The 1990 catalogue of the Reduviidae of the world (Maldonado Capriles 1990) listed 78 species, four of which are known to be predators on spiders. S. tfrar^m^^gz^Maldonado-Capriles &Van Doesburg from Dutch Guiana (Surinam) was found in the communal web of Anelosimus rupununi Levi. They have peculiarly modified antennae which may be an adaptation to their habit of walking through webs (MALDONADO-CAPRILES & VAN DOESBURG 1966). S. lanipes Wygodzynski has been observed to eat juveniles of Achaearanea tepidariorum (C.L. Koch) (= Parasteatoda t.) (HODGE 1984). S. giraffa Wygodzynski (Australia) has a striking, elongate pro thorax, hence its name. S. edwardsi Bergroth has been recorded as preying on young specimens of Badumna (Desidae) in Australia ( Wignall & Taylor 2008). Stenolemus bituberus Stal was found in the webs of — and seen actually feeding on — spiders of the fami- lies Desidae, Pholcidae,Theridiidae, and Uloboridae. Most likely araneophagy will be found subsequently among the many other Stenolemus species known. Neuroptera The subfamily Mantispinae of the Mantispidae are predators of spider eggs. The front legs resemble those of the praying mantis, hence the name mantispid flies. They are fairly long, up to 5 cm. The stalked eggs are Spiders in o hostile world 59 Odonata All Pseudostigmatidae (“Helicopter damsel flies”) in which the adult feeding habits are known prey exclusively on web-building spiders. Gifted with very good vision they aim di- rectly at the spider. Species showing this behaviour are Mecistogaster linearis (Fabricius), M. modesta Selys, M. ornata Rambur, Megaloprepus coerulatus (Drury), and Pseudostigma accedens Selys (CORBET 1999). M. coerulatus was seen preying on small Argyrodes spec. (Theridiidae) at a Nephila web (YOUNG 1980). M. modesta was seen at work near orb webs, as well as at the lampshade-shaped webs of pholcids. deposited on the substratum. Different strategies are employed to reach the spider’s eggs (REDBORG 1998). The larvae of one group of mantispids, the “boarders”, attach themselves to a passing bee, beetle, or spider. The larva then rides along on the spider, usually curled around the pedicel, feeding itself with haemolymph fluid from the spider, acting as a leech. Their final destination is the egg cocoon or egg batch. When a mantispid larva has settled on a young spider it has to get on the newly emerging next instar of the spider when it moults. It may seek refuge temporarily in a book lung during the moulting process. They wait for the construction of the egg cocoon, slip into it, feed on the eggs and pupate in the cocoon or egg batch. In the other strategy, that of the “borers”, the larva is attract- ed by spider silk and thus finds an egg sac and bores into it to feed on the eggs. Spiders which suffer from mantispid egg predation by spider boarders belong to a wide range of families of web builders as well as active hunters (REDBORG 1998), while the indepen- dent egg sac penetrators all feed exclusively on the eggs of hunting spiders. local spider fauna must be considerable. Vespa crabro acts as a regular kleptoparasite as well as predator on Argiope bruennichi (Scopoli) (Figs. 2-3). Ants are about the largest and ever present group of predator insects, often occur in very high numbers in certain habitats and are known to bring all types of prey to their nests, among which spiders do not fail. I have not found any literature on the relative importance of spiders in the ants’ diet. Coleoptera Carabid beetles are known to feed on spiders on agri- cultural fields, but no quantitative data are available. Hymenoptera Species from the Vespoidae, such as the hornet ( Vespa crabro L.), and Vespula species capture spiders as food for their brood. Because the colonies of these social living insects are often very large the impact on the Figures 2, 3: Vespa crabro in web of Argiope bruennichi steeling the spiders prey (2) (Photo Jeanette Hoek), and with remnant of Argiope bruennichi (3) (Photo Marcel Wasscher). 60 PJ.v.Helsdingen Figure 4: Tromatobia ornata on egg cocoon of Argiope bruennichi. Photo Gerben Winkel. Parasites and Parasitoids Egg parasitoids Some Pimplinae (Hymenoptera, Ichneumonidae) are predators of spider eggs ( Gelis , Hemiteles , Tromatobia, Zaglyptus, some Scelionidae) (RICHARDS 1977). The pimpline larva eats from the eggs in the spiders egg sac. Species of Tromatobia parasitize the egg sacs and adults of spiders. Tromatobia is a species-rich genus (FlTTON et al. 1988). A striking example is Argiope bruennichi (Scopoli), a species which has spread relatively quickly under its own power and of which the egg cocoons are parasitized by Tromatobia ornata Gravenhorst (Fig. 4). The parasite may have travelled along with the spider when it spread over the Netherlands over the last 25 years. True parasitoids Among Hymenoptera, the Ichneumonidae are also parasitoids of adult insects and spiders. They paralyze their prey, place an egg and after hatching the larva feeds on it while it remains in a stable, paralysed condition (endoparasitoids); or they place an egg on the victim which then continues its normal life until it succumbs because it is slowly weakened by its un- invited ecto-parasitic guest. Spider-wasps (Hymenoptera, Pompilidae) are specialized parasites of spiders. Their search for and capture of spider specimens is followed by a paralys- ing sting. The spider is then brought to a suitable place where it is burrowed, an egg is put on the spider, and the burrow is closed. The pompilid larva when full-grown pupates in the burrow. Most pompilid waps are polyphagous and hunt for spiders in general or special- ize on webspiders. Some are monophagous, at least regionally, such as Homono- tus sanguinolentus Fabricius which exclusively hunts for Cheiracanthium erraticum (Walckenaer) (Miturgidae) which is then left in its own silken nest (NIELSEN 1936). There are exceptions in this sequence. Eoferreola rhom- bica (Christ) parasitizes on Eresus sandaliatus (Martini 8c Goeze) (Eresidae). This spider lives in a burrow with a cribellate web above the entrance. Having located the spider in its burrow the wasp enters, paralyzes the spider, places its egg on the animal and leaves the burrow without closing it. It does not make a burrow of its own (HAUPT 1927). Neither does Aporus unicolor Spinola, which locates Atypus (Atypidae) in its burrow and leaves it there af- ter having paralyzed it and provided an egg. Ceropales species (Pompilidae) are known as kleptoparasites of other Pompilidae in that they follow other pompilid wasps with prey and put an egg on the prey just before the prey is buried by the true hunter (OEHLKE 8c Wolf 1987). Within the Ichneumonidae the Pimplinae com- prise the spider-ectoparasitoids of the Polysphincta group of genera of which we often see the larva externally on the abdomen (RICHARDS 1977). Spe- cies of the Polysphincta genus -group of the Pimplinae attack spiders. They first immobilize the spider, then put an egg on the spider, usually on the abdomen. The spider regains consciousness and leads a normal life until the larva is full-grown and pupates in the body of the dead spider. Many genera are distinguished, such as Dreisbachia , Schyzopyga, Polysphincta, Acrodac- tyla, Synarachna, and Zatypota (FlTTON et al. 1988; GAULD et al. 2006). A curious phenomenon in this respect is a proce- dure which is called “manipulation of the host behav- iour”. When the larva of Hymenoepimecis argyraphaga Spiders in o hostile world 61 Gauld (Ichneumonidae) is ready to pupate it stimu- lates the host spider Plesiometa argyra (Walckenaer) (= Leucauge argyra (Walckenaer)) (Tetragnathidae) just before it will die to produce a hub for a new web and repeat this over and over again, thus fabricating a “cocoon web” for its own parasitoid wasp (EBER- HARD 2000, 2001), this process probably stimulated by chemicals brought into the host’s body. I am not aware of any other cases of manipulation of behaviour by parasitoids of spiders. Within the Apoidea, the Crabronidae or digger wasps, comprise species which specialize on spiders and show a behaviour equivalent to that of the Pom- pilidae. Species of the genus Miscophus hunt for small spiders, and Trypoxylon species capture larger spiders, which they put in a cell and close off with mud. Diptera When the larvae of Acroceridae (Cyrtidae, Oncodi- dae) hatch they try to find a spider and climb on it, enter the book lungs and develop inside the abdomen. The spider dies when the parasitoid is full-grown and pupates. More than 500 species are known world- wide, mostly in the tropics. Recent additional data of acrocerid infestations in the Nearctic Region are given by LARRIVEE & BORKENT (2009). Discussion Spiders are strongly armed, well-equipped preda- tors: they have fangs to inject their venom, different types of silk for their webs - sticky or cribellate - and methods to wrap up their victims very quickly, and they have very short reaction time. For spiders the transition from hunting to invading a strange web, to kleptoparasitism, to becoming predator of a non- specific spider species, or to cannibalism, is nearly a continuum. The driving force in all instances is the search for food to meet the requirements of the individual’s energy balance. Food shortage will force spiders to eat individuals of their own kind. Other spiders, being live objects, are always on the menu. Once it is possible for an individual spider to enter a strange web unobserved or with misleading be- haviour the intruder can benefit from available food (kleptoparasitism), protection from other organisms which cannot enter the web (such as ants), or capture the owner by surprise (araneophagy). There are many examples of insects that follow the same strategies. There are also many invertebrates that have found ways and means to master spiders: they have won the arms race. It is clear that spider kleptoparasites benefit from using the prey collected by others. They do not have to invest in silk for webs, venom, or energy needed for hunting, jumping and overwhelming, while silk production for wrapping can be omitted. One may expect that exclusively kleptoparasitic spider species even have lost their capacity to produce venom, while the glands for silk production may have undergone reduction. The kleptoparasitic spider also gains pro- tection from other predators, such as the ever present ants, which, however, have been observed only in a few spider species to enter the web. For the host spider the stealing of food means loss of invested energy but for a larger host {Nephifo communal Anelosimus) this may be negligible, also because the kleptoparasites often eats from smaller prey for which the web owner has no interest. Aggression towards kleptoparasites by the web owner is nearly always negligible too, partly because of size differences (small kleptoparasites in the webs of larger species), partly because of subdued aggres- sion (spiders in communal webs). Kleptoparasitism appears to be not so rare a phenomenon, although the number of spider species which employ this feeding behaviour is relatively restricted as far as we know now. The number of insects that play a kleptoparasitic role in relation with spiders probably is larger than we know now. It may be expected that observations of spiders in their natural environment may reveal more kleptoparasitic relationships, especially in the acalyptrate Diptera and reduviid Heteroptera. Spiders are not defenceless against predators. They can defend themselves with their chelicerae and fight back, but against most stinging Hymenoptera they seem to hardly have a chance, although we do not know how many attempts by Ichneumonidae, Cra- bronidae, and Pompilidae meet with failure. Spiders can drop from the web, change colour when hitting the ground, run away, hide in self spun cells, or flee to the other side of the web, putting the web between himself and predator (JACKSON et al. 1993). How- ever, their chances of defence against predators which successfully deploy “aggressive mimicry” seem to be very slight. In the described cases the victim spiders were lured within striking distance of the predators “on perfidious pretexts” and the victim had very slight chances to escape. Apparently the “behavioral” arms race has been won by the predators, although we do not register where such methods are developing right now and have a lower percentage of successful attempts. It is difficult to detect evolution at work and 62 P.J. v.Helsdingen understand the direction the selective forces might move into. If we look at the orders of invertebrates (other than spiders) which have developed scavengers, klep- toparasites, or predators, and parasitoids of spiders we must conclude that the largest order, the Coleoptera with 359,891 described species, has hardly developed any (carabid beetles can feed on spiders on arable land if no other food is available), that the Diptera (152,244) have some kleptoparasites among them but are under-represented as to predators (none) and parasitoids (only one family). Lepidoptera (156,793 species) are represented with noctuid kleptoparasites, but this feature seems to be rather exceptional in the order. The Odonata are a small order (5,680 species) of which only few genera have developed into spider predators. By far the most kleptoparasites are found among the Hemiptera (100,428 species), while some have become predators. The largest number of general predators, parasites and parasitoids are found in the Hymenoptera (144,695 species), which all possess poison glands and thus are able to overwhelm and/or parasitize spiders. They have developed a weapon of their own and are clearly ahead in the arms race. (All data on species numbers after ADLER & FOOTTIT 2008.) Reflections Interactions as brought together and discussed in this paper are of importance for understanding the biology of the species, of spiders as well as of the many insects involved. Discovering new relation- ships and interactions will help us to understand the many interesting behavioural patterns and food chains which exist in the invertebrate world. They illuminate an important aspect of the “web of life” and demonstrate the intricacies of food chains. It is clear that this asks for collecting observations in the field more than collecting specimens. Pitfalls and canopy fogging yield specimens and give insight in the composition of the fauna but they do not help us to find patterns of behaviour, parasitic relationships or food chains. This paper is meant to stimulate the observing type of invertebratologists who sits down amidst the invertebrates at work and tries to discover the patterns of interactions between organisms, the way organisms react to each other. References Adler P.H. & R.G. FOOTTIT (2008): Introduction. In: FOOTTIT R.G. &P.H. ADLER (eds.): Insect biodiversity, science and society. Wiley-Blackwell, Oxford. 632 pp. AGNARSSON I. (2002): Sharing a web - on the relation of sociality and kleptoparasitism in theridiid spiders (Theridiidae, Araneae). - Journal of Arachnology 30: 181-188 - doi: 10.1636/0161-8202(2002)030[0181: SAWOTR]2.0.CO;2 AGNARSSON I. (2003): Spiderwebs as habitat patches. 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Altogether two endemic harvestmen (Nemastoma bidentatum relictum, Nemastoma schuelleri) and 8 endemic spiders {Abacoproeces molestus,Collinsia (caliginosa) nemen- ziana, Mughiphantes severus, Mughiphantes styriacus, Pelecopsis alpica, Scotophaeus nanus, Troglohyphantes novicordis, Troglohyphantes tauriscus), beside 9 subendemic harvestman and 38 subendemic spider species have been recorded from Austria. Hot-spots of endemism in the Eastern Alps are the north-eastern (Ennstaler Alps) and southern Calcareous Alps (Karawanken, Karnische Alps) and the Central Alps (HoheTauern, Gurktaler Alps, Ötztaler and Stubaier Alps). Most of the endemic arachnid species occur from the nival down to the montane zone. Important habitats are rocky areas, caves and woodlands. High absolute numbers and percentages of endemics can be found within the har- vestman families Cladonychiidae, Ischyropsalididae and Nemastomatidae and in the spider genera Lepthyphantes s. Land Troglohyphantes. Jhe conservation status of these highly endangered taxa - 85 % of the spider species and 100 % of the harvestman taxa are endangered in Austria - is poor. Key words: conservation, Eastern Alps, endangering, endemism, ice age, massifs de refuge, nunataks, red list, subendemics, protection, vertical distribution Zoogeography appears to be one of the most amusing and stimulating of the natural sciences: every few years its fundamental concepts change and one can begin anew. Paolo Marcello BRIGNOLI (1983: 181) The so-called Eastern Alps belong to the 30-35 mil- lion year old European Alpine system, and are largely contained within the national borders of Austria. Despite intensive research efforts by several Austrian zoologists in the past, like Rudolf Heberdey, Karl Holdhaus, Herbert Franz and Heinz Janetschek or the German Gustaf de Lattin as well as more recently renowned „Alpine- arachnologists“ like Konrad Thaler and Jürgen Gruber, a comprehensive faunal catalogue for the region is lacking. A recent study, co-ordinated by the Environment Agency Austria (Umweltbundesamt), aimed to fill this deficit. A comprehensive overview of plant, fungus and animal species, whose range lies entirely (endemics) or predominantly (subendemics) within the political borders of Austria, has now been com- pleted (RABITSCH & ESSL 2009). Altogether 748 (sub)endemic animal and plant species have been identified. Among the 548 animal species in Aus- tria 10 pseudoscorpion (7 endemic, 3 subendemic) Christian KOMPOSCH, ÖKOTEAM - Institute for Animal Ecology and Landscape Planning, Bergmanngasse 22, 801 0 Graz, Austria. E-Mail: c.komposch@oekoteam.at submitted: 1 9. 1 .20 1 0, accepted: 20.4.20 7 0; online: 10.1.2011 (MAHNERT 2009), 11 harvestman (KOMPOSCH 2009a) and 46 spider species (KOMPOSCH 2009b) can be found (Fig. 1,2). Scorpions (KOMPOSCH 2009c) and palpigrades (CHRISTIAN 2009) include no real (sub)endemic species of Austria, whereas 10 oribatid mites (SCHATZ SCHUSTER 2009) are classified as endemic and subendemic; many more oribatids are pseudoendemics, i. e. the current taxonomy of faunis- tic knowledge is not sufficient and a wider distribution is assumed. So why is it important to publish these data in addition to the endemics book? This is done for the following reasons: first the endemics book is of re- gional distribution, mainly spread in Austria, and due to the small print run it will soon be sold out (compare BLICK 2009). Second it is written in German, and for the relevant chapters even an English abstract is miss- ing. The present publication closes this gap for the arachnological section. Furthermore it expands and updates the knowledge of the two treated arachnid groups. Material and Methods This paper deals with species and subspecies, whose ranges lie entirely (endemics) or predominantly (subendemics) within the political borders of Austria. 66 C. Komposch Figure 1 : Habitus of endemic and subendemic harvestmen of Austria. From left to right and from the top downwards: Holo- scotolemon unicolor, Paranemastoma bicuspidatum, Ischyropsalis kollari, Megabunus armatus, Leiobunum roseum, Leiobunum subalpinum. Subendemics are defined in this context as taxa with more than 75 % of their total range within Austria or if they are local or regional endemics with less than 10 known localities or if their area is restricted to less than 1000 km2 respectively with less than 75 % of their total range in Austria (RABITSCH & ESSL 2009). All available, published and unpublished, data on endemic and subendemic harvestmen and spiders have been collected, reviewed, geographically located, digitised and stored in a private database (data base ÖKOTEAM). Altogether more than 2250 data-sets were analysed (Araneae: 1050, Opiliones: 1200). The distribution maps (see appendix) - showing the natural zones as well as the political borders of the federal countries - have been printed by the Environ- ment Agency Austria (UBA). All photos were taken by the author. Results The geographical localisation and digitalisation of all available data facilitated the present drawings of distribution maps (appendix 1, 2) and - for the first time - the clear identification of centres and hotspots of faunal endemism within the Eastern Alps. A complete list of all endemic and subendemic harvestman and spider taxa is given in table 1, a se- lection of habitus photographs is presented in figure 1. Altogether two endemic ( Nemastoma bidentatum Endemie harvestmen and spiders in Austria 67 Figure 2: Habitus of endemic and subendemic spiders of Austria. From left to right and from the top downwards: Mughiphantes variabilis,Xysticus secedens, Troglohyphantes tauriscus, Troglohyphantes noricus. relictum, Nemastoma schnellen ) and nine subendemic harvestmen are recorded from Austria (table 1, ap- pendix 1). The spider fauna shows eight endemic (. Abacoproeces molestus , Collinsia ( caliginosa ) nemen- ziana , Mughiphantes sever us, Mughiphantes styriacus, Pelecopsis alpica, Scotophaeus nanus, Troglohyphantes novicordis, Troglohyphantes tauriscus) and 38 suben- demic species (table 1, appendix 2). Therefore a total of 57 subendemic and endemic harvestman (11) and spider (46) species can be found within the Austrian Republic. Regarding the main vertical distribution of Opiliones and Araneae in the Eastern Alps the colline zone (115 up to ca. 250-400/500 m) harbours 2 harvestmen and 4 spider species, the submontane (up to ca. 350-500/700 m) 3 / 9, the montane (up to ca. 1500-2000 m) 10 / 21, the subalpine (up to ca. 1800-2100 m) 8 / 23, the alpine (up to ca. 2500-2800 m) 3 / 18, and the nivale zone (up to 3798 m) 1 / 6 species. Of the (sub)endemic taxa, 39 spider species (85 %) are “Critically Endangered” up to “Vulnerable” and two species (4 %) are placed in the category “Data deficient” (KOMPOSCH in press b). All 11 harvest- man taxa are vulnerable or endangered in Austria (KOMPOSCH 2009d). The horizontal distribution of arachnids shows major differences between the different natural areas in Austria. Rich in endemic spiders and harvestmen are the mountainous federal states of Styria (St), Carinthia (C), Tyrol (T) and Salzburg (S). MAURER 5c HÄNGGI (1990) pointed out, that endemics of the Northern Alps mostly show bigger areas than south-alpine ones. Concerning the fauna of the whole Alpine arc this is definitively correct. With respect to the distribution maps of Austrian (sub)endemics it could hardly be verified, as the political borders of Austria comprise a much bigger area of the Northern, as opposed to the Southern, Alps. Discussion The number of endemic and subendemic arachnids within Austria is remarkably high and differs widely in the nine federal states (Bundesländer). Centres of arachnological and zoological diversity and endemism are, within the entire Alps, the Southern Calcare- ous Alps with their peak in the south-western parts (Mercantour National Park and Alpi Marittime NP) (Maurer 5c Thaler 1988). The hotspot of arachnological endemism in Austria is situated in the Eastern Alps. Following the definitions of subende- mism of RABITSCH 5cESSL (2009) the arachnological hotspots in Austria are the central Hohe Tauern (C, S, T) and the Gurktaler Alps (C, S, St), but also the 68 C Komposch Table 1 : List of endemic (E), subendemic (S), and pseudoendemic (P) harvestmen (Opiliones) and spider (Araneae) species of Austria, related to the altitudinal zones of their main occurrence in the area (colline to nivale). RLA = Red List of endangered harvestmen and spiders of Austria (Komposch 2009d, Komposch in press). Categories of endangerment used: LC = Least Concern, NT = Near Threatened, DD = Data Deficient, VU = Vulnerable, EN = Endangered, CR = Critically Endangered.* Tapinocyba affinis is endemic to the alpine mountainous system (Thaler 1999). The subspecies orientalis, although not clearly shown up to now, seems to have a wider distribution in Slovakia, Czech Republic and Germany (Miludge 1 979, Komposch 2009b, Blick in litt.); in this case it would lose the subendemic-status in Austria. family taxon Opiliones (harvestmen) 1 Cladonychiidae Holoscotolemon unicolor Roewer, 1915 2 Nemastomatidae Mitostoma alpinum (Hadzi, 1931) 3 Nemastoma bidentatum relictum Gruber & Martens, 1968 4 Nemastoma schnellen Gruber & Martens, 1968 5 Paranemastoma bicuspidatum (C. L. Koch, 1835) 6 Ischyropsalididae Ischyropsalis hadzii Roewer, 1950 7 Ischyropsalis kollari C. L. Koch, 1839 8 Phalangiidae Megabunus armatus (Kulczynski, 1887) 9 Megabunus lesserti Schenkel, 1927 10 Sclerosomatidae Leiobunum roseum C. L. Koch, 1839 11 Leiobunum subalpinum Komposch, 1998 Araneae (spiders) 1 Linyphiidae Abacoproeces molestus Thaler, 1973 E CR 1 2 Centrophantes roeweri (Wiehle, 1961) S EN 1 1 3 Collinsia (caliginosa) nemenziana Thaler, 1980 E VU 1 4 Diplocephalus rostratus Schenkel, 1934 S EN 5 Incestophantes kotulai (Kulczynski, 1904) s VU 1 6 Meioneta alpica (Tanasevitch, 2000) s DD 1 7 Meioneta ressli Wunderlich, 1973 s VU 1 8 Metopobactrus nodicornis Schenkel, 1927 s EN 1 9 Micrargus alpinus Relys & Weiss, 1997 s VU 1 10 Mughiphantes armatus (Kulczynski, 1905) s VU 11 Mughiphantes rupium (Thaler, 1984) s CR 1 12 Mughiphantes severus (Thaler, 1990) E CR 1 13 Mughiphantes styriacus (Thaler, 1984) E CR 1 14 Mughiphantes triglavensis (Miller & Polenec, 1975) s EN 1 1 15 Mughiphantes variabilis (Kulczynski, 1887) s NT 1 1 16 Palliduphantes montanus (Kulczynski, 1898) s LC 1 1 17 Pelecopsis alpica Thaler, 1991 E CR 1 18 Scotinotylus clavatus (Schenkel, 1927) s EN 1 1 19 Silometopus rosemariae Wunderlich, 1969 s VU 1 1 20 Styloctetor austerus (L. Koch, 1884) s VU 1 21 Syedra apetlonensis Wunderlich, 1992 s CR 1 22 Tapinocyba affinis orientalis Millidge, 1979 s* NT 1 1 23 Tenuiphantes jacksonoides (Van Helsdingen, 1977) s NT 1 1 1 24 Troglohyphantes fagei Roewer, 1931 s VU 1 1 S EN 1 S EN E EN E EN S EN S EN S VU 1 S EN S VU S EN S VU 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1111 1 1 1 1 1 1 E/S RLA g c .5 a g c g .2 T3 ~ ■§ U CA Endemie harvestmen and spiders in Austria 69 family taxon E/S RL A colline submontane montane subalpine alpine nival 25 Troglohyphantes karawankorum Deeleman-Reinhold, 1978 S CR 1 1 26 Troglohyphantes latzeli Thaler, 1986 S CR 1 1 27 Troglohyphantes noricus (Thaler 8c Polenec, 1974) S VU 1 1 1 28 Troglohyphantes novicordis Thaler, 1978 E CR 1 29 Troglohyphantes subalpinus Thaler, 1967 S VU 1 1 1 30 Troglohyphantes tauriscus Thaler, 1982 E EN 1 1 31 Troglohyphantes thaleri Miller 8c Polenec, 1975 S VU 1 1 1 32 Troglohyphantes typhlonetiformis Absolon 8c Kratochvil, 1932 S CR 1 1 33 Tetragnathidae Pachygnatha terilis Thaler, 1991 s EN 1 1 34 Lycosidae Arctosa renidescens Buchar & Thaler, 1995 s EN 1 1 35 Pardosa giebeli (Pavesi, 1873) s VU 1 1 36 Pardosa saturatior Simon, 1937 s VU 1 1 1 37 Agelenidae Tegenaria mirifica Thaler, 1987 s CR 1 38 Hahniidae Cryphoeca lichenum lichenum L. Koch, 1876 s NT 1 1 39 Cryphoeca lichenum. nigerrima Thaler, 1978 s EN 1 1 40 Gnaphosidae Haplodrassus aenus Thaler, 1984 s EN 1 1 41 Haplodrassus bohemicus Miller 8c Buchar, 1977 s CR 1 42 Scotophaeus nanus Wunderlich, 1995 E? EN 1 43 Zelotes zellensis Grimm, 1982 s VU 1 1 44 Philodromidae Philodromus depriesteri Braun, 1965 s DD 1 45 Thanatus firmetorum Muster 8c Thaler, 2003 s VU 1 1 1 46 Thomisidae Xysticus secendens L. Koch, 1876 s VU 1 1 1 Ötztaler and Stubaier Alps (T) as well as the Koralpe (C, St) (all Central Alps), the Gesäuse National Park (Ennstaler Alps, Northern Calcareous Alps, St) and in particular the Karawanken (Southern Calcareous Alps, C) with their massifs de refuge, marking the margin of the Wiirm-ice-shields (Fig. 3). Regions outside the Alps are poor in endemics. For spiders and harvestmen a maximum of 12 taxa per grid-cell was found in Austria. Concerning all animals, the maximum was 46 taxa in a grid cell in the Gesäuse NP (St). The Hochobir (Fig. 4) in the Karawanken (Southern Calcareous Alps, C) came second with 41 endemic taxa. A comparison between the arachno- logical hotspots (Fig. 5) and these of the whole fauna (RABITSCH 8cESSL 2009: 882) reveals a quite similar picture. Differences can be found in the araneological hotspot in the Ötztaler and Stubaier Alps, intensively explored by Konrad Thaler and his students, and - probably a methodical artefact — the comparatively low frequency in the eastern parts of the Northern Calcareous Alps (Styria, Upper and Lower Austria). Regarding a wider definition of subendemism the Austrian number one would be the Southeastern Calcareous Alps with the Karawanken and Steiner Alps at the borderline between Carinthia/ Austria and Slovenia, followed by the Karnische Alps, marking the border between Carinthia/ Austria and Italy. Without attempting to explain the origin of these en- demics in detail, I would like to point out three main reasons for the richness of the small-scale distributed species in the Alps: 1) The recent climate history with large-scale expansi- on of the last ice-shields (Fig. 3 is of importance to understand today's distribution ranges. Like other zoogeographers, DE LATTIN (1967) previously pointed out that the geological event with the strongest influence on the Holarctic faunas was irruption by the Pleistocene ice-ages. HOLDHAUS 70 C. Komposch (1954) wrote about the “rettungslose Vernichtung” [desperate devastation] of the Alpine fauna by the glaciations. 2) The Alpine inhabitants were pushed towards refu- gia by the glaciations (e.g. BRIGNOLI 1983, WEISS 6e FERRAND 2007), then underwent evolutionary processes and post-glacial re-immigration over both short and long distances (HOLDHAUS 1954, JANETSCHEK 1956, THALER 2003). 3) Survival was possible in massifs de refuge, i.e. un- glaciated massifs at the periphery of the ice shields (e.g. Fig. 3), in caves and - quite rarely — on inner Alpine nunataks (MAURER 6c HÄNGGI 1990, Holderegger 6cThiel-Egenter 2009). Nu- nataks are steep mountain peaks free of ice, which appear through the ice crust. Due to the particular forms of glaciation (Fig. 3), survival and re-wandering processes in the Eastern Alps the diversity of endemic and subendemic spiders and harvestmen shows a South- North and an East-West decline (cf. Muster 2001, 2002). Most of the endemic arachnid species occur from the alpine down to the montane zone (table l).The most important habitats are rocky areas, caves and woodlands. High absolute numbers and percentages of endemics can be found within the soil-inhabiting harvestman-families Cladonychiidae, Ischyropsalididae and Nemastomatidae and the spi- der-family Linyphiidae ( Lepthyph - antes spp. s. 1. and Troglohyphantes spp.). Beyond these linyphiid taxa Maurer 6c Hänggi (1990) list Coelotes , Cryphoeca, Cybaeus , Nesti- cus , dysderids and leptonetids. The threat status of endemic spider- and harvestman-species in Austria is in general high. The major threats are caused by fore- stry (old-growth forests are being replaced by common spruce mo- nocultures), hydraulic engineering (large Alpine valleys are flooded and changed to hostile reservoirs), agriculture (intensification and bi- ocide use) and tourism (devastating high Alpine and mountainous areas for building and expanding ski-regions; a big problem is the enormous need for water for ski cannons, which leads to the construction of water reservoirs, water diversion and the destruction of spring communities, brooklets and brooks). Furthermore climate warming leads to habitat loss and endangers cold stenothermic species inhabiting rather low peripheral mountain chains of the Alps, which were not glaciated during the Pleistocene; DlRNBÖCK et al. in press). Despite these various hazardous impacts on sensitive habitats and their species, endemic arachnids and insects are so far not protected by Austrian laws. The coverage of the distribution of endemics by nature reserves is rather poor. Firstly none of the Austrian nature conservation areas or National Parks has been established up to now in response to a high or even outstanding diversity among endemic invertebrates. Not more than 10 % of Figure 3:The eastern Alps at maximum glaciation during the last Ice Age (Würm): orography and political borders of Austria (modified after van Husen 1 987, Rabitsch & Essl 2009) Figure 4: The Hochobir, an impressive peak and massif de refuge within the Kara- wanken (Southern Alps, Carinthia), is a well known hot spot of endemic arachnids. Endemie harvestmen and spiders in Austria 71 10° 11° 12° 13° 14° 15° 16° 17° 9ft’ 3(Y 40' fin’ 10’ 90’ 30“ 40’ ace 10* ?0’ 3(T 40’ scr 10’ 20’ 30* 40’ 50* 10’ 90’ 30’ 40’ 50’ 10’ 90’ 30’ 40’ 50’ 10’ 20’ 30* 40' ace 10’ 20’ 30" 40’ ace 10’ 20’ 30' 40’ 50* 10’ 20’ 30’ 40’ 50* 10’ 20’ 30’ 40’ 50* 10’ 20’ 30’ 40’ 50’ 10’ 20’ 30’ 40’ 50’ 10’ 20' 30' 40’ 50’ 10’ 20’ 30* 40’ 501 10* 20’ 3CT 40’ 50“ 10' 10° 11° 12° 13° 14° 15° 16° 17° Figure 5: Cumulative distribution map showing all endemic and subendemic spiders and harvestmen in Austria (modi- fied after Komposch 2009b). The main hotspots of arachnological endemism are encircled by rings and ovals. the 16 quadrants with the highest number of endemic animals in Austria are covered by nature conservation areas. Secondly the coverage of large parts of the country by Natura-2000-areas rules out protection of the endemic species inside these areas as they are not regarded as conservation objects by the European Union. It is quite astonishing that comparative data on endemic arachnids from other central European countries are missing or weak. Therefore a comparison to the Austrian results is hardly possible. This highly unsatisfactory situation is in need of change. Available data refer to Romania: TATOLE (2006) carried out a zoogeographical analysis of the 961 hitherto registered spider species and identifies about 6 % as endemic. The spider fauna of Bulgaria is represented by 975 species, including “76 species (10 %) established in Bulgaria (35 species) and other territories of the Balkan Peninsula (41 species)” (DELTSHEV 2005: 309). Following this author, this phenomenon can be attributed to the relative isola- tion of the mountains compared with the lowlands in the context of paleo-environmental changes since the Pliocene. MAURER &, HÄNGGI (1990) disclose in their catalogue of Swiss spiders a value of 7 % of merely alpine species - i.e. just partly endemic or subendemic taxa of Switzerland. The spider fauna of the Caucasus, comprising Armenia, Azerbaijan and Georgia as well as parts of the Iran, Turkey and Russia, is poorly known; the value of endemic spiders is 22 % (> 226 endemics from a total of 1022 species) (MARUSIK et al. 2006). Conclusions The presented results provide a valuable basis for both zoogeographical inferences involving glacial refugia and postglacial re-colonisation of the fauna of the Alps as well as conservation planning in Austria. Although known from other animal groups, the high diversity of endemic spiders and harvestmen from the Gurktaler Alps and the Gesäuse National Park (KOMPOSCH 2010) was surprising. Further faunistic and taxonomic work especially in these areas should bring forward new and undescribed species. As pointed out before, the status of subendemism strongly relates to its definition. A second step towards the recording and documentation of the subendemic fauna of Austria - and the endemic fauna of the Alps - is urgently needed. From the conservation perspec- tive it is not acceptable to pay attention merely to spe- cies with a 75 % area-quotient within national borders. Taking into account the fact that several rare, small scale-distributed and endangered species share their area equally within two or three countries (e.g. within 72 C Komposch the south-eastern Alps: Slovenia, Italy and Austria), only a percentage of about 20 clearly leads to a main distribution within each country and consequently to a main responsibility for each country. The need for action is great, and the time pressure due to habitat destruction and climate warming is high. The following four steps should be rapidly undertaken: 1) Clarification of targets and general conditions of a national and international protection concept, including basic research, politics, protection measures, university training and public relations (Komposch in press a). 2) Identification and prioritisation of endemic taxa (local, regional and endangered endemics) and regions (hotspots of endemism) - with continuative and representative mappings - with a high need for protection. 3) Drafting of specific protection measures, showing deficits of knowledge and priorities of research. Stopping the loss of organismic biologists and specialists able to identify a species. 4) Implementation of the protection concept (e.g. KLAUSNITZER 2010) (anchoring the protection of endemics in federal conservation laws as well as in the FFH-directive of the European Union), accompanied by long-term monitoring and read- justment of measures. Conservation efforts must focus on these unique treasure of our Adpine fauna. Ackowledgements For arachnological data and help I am deeply grateful to Konrad Thaler (f), Jürgen Gruber and Albert Ausobsky. Thanks for data, literature, fieldwork and intensive dis- cussions to Franz Essl, Thomas Frieß, Werner Holzinger, Barbara Knoflach, Brigitte Komposch, Jochen Martens, Christoph Muster, Tone and Ljuba Novak, Wolfgang Paill, Axel Schönhofer and Reinhart Schuster. Financial support came from the Umweltbundesamt Vienna (project-sup- port), the Österreichische Forschungsgemeinschaft and the Styrian Government (both support of the congress partici- pation in Alexandroupolis 2009) and technical support from Wolfgang Rabitsch. Big thanks go to the two anonymous reviewers, Jason Dunlop for improving the English, Theo Blick and Oliver-D. Finch for discussion und valuable comments and to Maria Chatzaki for her patience with the manuscript. References BLICK T. (2009): Wolfgang Rabitsch & Franz Essl (2009): Endemiten. Kostbarkeiten in Österreichs Pflanzen- und Tierwelt. 924 S. Mit zahlreichen Einzelbeiträgen ver- schiedener Autoren. - Arachnologische Mitteilungen 37: 39-40 - doi: 10.543 l/aramit3708 BRIGNOLI P.M. (1983): Dispersion, dispersal and spiders. - Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg, 26: 181-186 CHRISTIAN E. (2009) Palpigradi (Tasterläufer). In: RA- BITSCH W. &.F. ESSL (Red.): Endemiten. Kostbarkeiten in Österreichs Tier- und Pflanzenwelt. Naturwissen- schaftlicher Verlag für Kärnten und Umweltbundesamt, Wien. pp. 406-407 DELTSHEV C. (2005): Fauna and zoogeography of spiders (Araneae) in Bulgaria. - Journal of Arachnology 33: 306-312 - doi: 10.1636/CH05-6.1 DIRNBÖCKT, F. Essl &W. RABITSCH (in press): Dispro- portional risk for habitat loss of high-altitude endemic species under climate change. - Global Change Biology - doi: 10.1111/j.l365-2486.2010.02266.x HUSEN D. van (1987): Die Ostalpen in den Eiszeiten. Populärwissenschaftliche Veröffentlichungen der Ge- ologischen Bundesanstalt, Wien. 24 pp. & 1 map De LATTIN G. (1967): Grundriß der Zoogeographie. G. Fischer, Stuttgart. 602 pp. Holderegger R. & C. Thiel-Egenter (2009): A discussion of different types of glacial refiigia used in mountain biogeography and phylogeography. - Journal of Biogeography 36: 476-480 - doi: 10.111 1/j. 1365- 2699.2008.02027.x HOLDHAUS K. (1954): Die Spuren der Eiszeit in der Tierwelt Europas. - Abhandlungen der zoologisch- botanischen Gesellschaft in Wien 18: 1-493, Taf. 1-52 JANETSCHEK H. (1956): Das Problem der inneralpinen Eiszeitüberdauerung durch Tiere (Ein Beitrag zur Ge- schichte der Nivalfauna). - Österreichische Zoologische Zeitschrift 6: 421-506 KLAUSNITZER B. (2010): Entomologie - quo vadis? - Nachrichtenblatt der Bayerischen Entomologen 59: 99-111 KOMPOSCH C. (2009a): Weberknechte (Opiliones). In: RA- BITSCH W. &F. ESSL (Red.): Endemiten. Kostbarkeiten in Österreichs Tier- und Pflanzenwelt. Naturwissen- schaftlicher Verlag für Kärnten und Umweltbundesamt, Wien, pp 476-496 KOMPOSCH C. (2009b): Spinnen (Araneae). In: RABITSCH W. & F. ESSL (Red.): Endemiten. Kostbarkeiten in Österreichs Tier- und Pflanzenwelt. Naturwissen- schaftlicher Verlag für Kärnten und Umweltbundesamt, Wien. pp. 408-463 Endemie harvestmen and spiders in Austria 73 KOMPOSCH C. (2009c): Skorpione (Scorpiones). In: RABI- TSCH W. 6c F. ESSL (Red.): Endemiten. Kostbarkeiten in Österreichs Tier- und Pflanzenwelt. Naturwissen- schaftlicher Verlag Air Kärnten und Umweltbundesamt, Wien. pp. 496-500. KOMPOSCH C. (2009d): Rote Liste der Weberknechte (Opiliones) Österreichs. In: ZULKA P. (Red.): Rote Listen gefährdeter Tiere Österreichs. Checklisten, Ge- fährdungsanalysen, Handlungsbedarf. - Grüne Reihe des Lebensministeriums 14/3: 397-483 KOMPOSCH C. (2010): Alpine treasures - Austrian endemic arachnids in the Gesäuse National Park. - eco.mont 2(2): 21-28 - doi: 10.1553/eco.mont-2-2s21 KOMPOSCH C. (in press): Rote Liste der Spinnen Öster- reichs (Arachnida: Araneae). In: ZULKA P (Red.): Rote Listen gefährdeter Tiere Österreichs. Checklisten, Ge- fährdungsanalysen, Handlungsbedarf. - Grüne Reihe des Lebensministeriums 14/4 MAHNERT V. (2009): Pseudoscorpiones (Pseudoskorpi- one). In: RABITSCH W. 6c F. ESSL (Red.): Endemiten. Kostbarkeiten in Österreichs Tier- und Pflanzenwelt. Naturwissenschaftlicher Verlag für Kärnten und Umweltbundesamt, Wien. pp. 501-508 Marusik Y.M., K.G. Mikhailov 6c E.F. Guseinov (2006): Advance in the study of biodiversity of Cauca- sian spiders (Araneae). In: DELTSHEV C. 6c P. STOEV (Eds.): European Arachnology 2005. - Acta zoologica bulgarica, Suppl. 1: 259-268 MAURER R. 6c A. HäNGGI (1990): Katalog der schweizeri- schen Spinnen. - Documenta Faunistica Helvetiae 12: unpaginated MAURER R. 6c K. Thaler (1988): Über bemerkenswerte Spinnen des Pare National du Mercantour (F) und seiner Umgebung (Arachnida: Araneae). - Revue suisse de Zoologie 95: 329-352 MlLLIDGE A.F. (1979): Some erigonine spiders from southern Europe. - Bulletin of the British arachnological Society 4: 316-328 MUSTER C. (2001): Biogeographie von Spinnentieren der mittleren Nordalpen (Arachnida: Araneae, Opiliones, Pseudoscorpiones). - Verhandlungen des Naturwis- senschaftlichen Vereins in Hamburg 39: 5-196 MUSTER C. (2002): Substitution patterns in congeneric arachnid species in the northern Alps. - Diversity and Distribution 8: 107-121 - doi: 10.1046/j.l472- 4642.2002.00131.x RABITSCH W. 6c F. ESSL (Red.) (2009): Endemiten - Kostbarkeiten in Österreichs Pflanzen- und Tierwelt. Naturwissenschaftlicher Verlag für Kärnten und Um- weltbundesamt, Wien, 924 pp. SCHATZ H. 6c R. Schuster (2009): Oribatida (Hornmil- ben). In: RABITSCH W. 6c F. ESSL (Red.): Endemiten. Kostbarkeiten in Österreichs Tier- und Pflanzenwelt. Naturwissenschaftlicher Verlag für Kärnten und Um- weltbundesamt, Wien. pp. 464-475 TATOLE A. (2006): On the biogeography of Romanian spiders (Araneae). In: DELTSHEV C. 6c P. STOEV (Eds.): European Arachnology 2005. - Acta zoologica bulgarica, Suppl. 1: 281-285 THALER K. (1999): Beiträge zur Spinnenfauna von Nord- tirol - 6. Linyphiidae 2: Erigoninae (sensu Wiehle) (Arachnida: Araneae). - Veröffentlichungen des Tiroler Landesmuseums Ferdinandeum 79: 215-264 THALER K. (2003): The diversity of high altitude arachnids (Araneae, Opiliones, Pseudoscorpiones) in the Alps. In: Nagy L., G. Grabherr, C. Körner 6c D.B.A THOMPSON (eds.): Alpine biodiversity in Europe. - Ecological Studies 167: 281-296 WEISS S. 6c N. FERRAND (2007): Phylogeography of southern European refugia: evolutionary perspectives on the origins and conservation of European biodiversity. Springer, Dordrecht. 377 pp. - doi: 10.1007/1-4020- 4904-8 Appendix 1 : Distribution patterns for endemic and subendemic harvestmen (Opiliones) in Austria - 1 1 maps. 74 C. Komposch ! Nemastoma schuelleri vor 1900 «> 1900-1950 • nach 1950 A Fundzeitpunkt unbekannt Ischyropsalis hadzii Leiobunum roseum 75 Endemie harvestmen and spiders in Austria Appendix 2: Distribution patterns for endemic and subendemic spiders (Araneae) in Austria - 46 maps. Centrophantes roeweri 76 C Komposch Mughiphantes rupium vor 1900 C 1900-1950 • nach 1950 A Fundzeitpunkt unbekannt Mughiphantes styriacus vor 1900 4) 1900-1950 • nach 1950 a Fundzeitpunkt unbekannt Mughiphantes variabilis Palliduphantes montanus O vor 1900 4 1900-1950 • nach 1950 i ! Endemie harvestmen and spiders in Austria 11 78 C. Komposch Endemie harvestmen and spiders in Austria 79 Scotophaeus nanus Thanatus firmetorum Haplodrassus bohemicus Xysticus secedens Philodromus depriesteri Arachnologische Mitteilungen 40:80-84 Nuremberg, January 201 1 Ground-living spiders (Araneae) at polluted sites in the Subarctic Seppo Koponen doi:10.5431/aramit4009 Abstract: Spiders were studied around the Pechenganikel smelter combine, Kola Peninsula, north-western Rus- sia.The average spider density was 6-fold greater and the density of Linyphiidae specimens 1 1.5-fold higher at slightly polluted sites, compared with heavily polluted sites. Altogether, 18 species from 10 families were found at heavily polluted sites, the theridiid Robertus scoticus clearly dominating (23.3 % of identifiable specimens), also Neon reticulatus (9.6 %), Thanatus formicinus (9.6 %) and Xysticus audax (8.2 %) were abundant. The most numerous among 58 species found at slightly polluted sites were Tapinocyba pollens (18.5 %), Robertus scoticus (13.7 %),Masosundevalli (9.5 %) and Alopecosa aculeata (8.2 %).The family Linyphiidae dominated at slightly polluted sites, 64 % of species and 60 % of individuals; compared with heavily polluted sites, 23 % and 38 % respectively. Key words: density, diversity, Kola Peninsula, smelter In the late 1980's, reports of heavy pollution loads from the Russian smelters in the Kola Peninsula and their possible effect on needle losses among pine in northern Finland prompted active studies on effects of pollution on forest ecosystems in northern Fin- land, Russia and Norway (see KOZLOV et al. 1993, Tikkanen &c Niemelä 1995). Spiders, like some other predatory arthropod groups, have been observed at heavily polluted sites near smelters in northern Europe (BENGTSSON & Rudgren 1984, Koneva 1993, Koponen & Niemelä 1994, Koponen & Koneva 2006). Gen- erally, markedly high concentrations of heavy metals have been found in spiders near pollution sources (Bengtsson & Rundgren 1984, Koponen & Niemelä 1995, Maelfait & Hendrickx 1998), and spiders have often been used as indicators in monitoring the effects of pollution (e.g. CLAUSEN 1986). In the present paper, information will be given on spider assemblages near the Pechenganikel smelter complex, in the subarctic Kola Peninsula, NW Rus- sia. For general data on the nature and degree of pol- lution in the area, see KOZLOV et al. (1993, 2009) and Norwegian Pollution Control Authority (2002). Spider densities near the Pechenganikel smelter have been briefly dealt with by KONEVA & KOPONEN (1993). For the spider fauna of natural forests in the subarctic and northern boreal taiga zones in Fennoscandia, see e.g. KOPONEN (1977, 1999) and RYBALOV (2003). Seppo KOPONEN, Zoological Museum, University of Turku, FI-20014Turku, Finland. E-Mail: sepkopo@utu.fi submitted: 77. 1.2010, accepted: 17.3.2010; online: 10.1.201 1 Figure 1 : Pechenganikel (the study area) and Monchegorsk smelters in Kola Peninsula. Material and Methods The present study area, the Pechenganikel smelter combine (Nikel and Zapolyarny), ca 69°30’N, 30°20’E, is situated in a subarctic pine forest area, where there are also birch woods, bogs and treeless fells (Fig. 1). The altitude of sites varies between 100 and 200 m asl. The smelter is very close to Norway (only about 10 km from the border) and to Finland (40 km from the border), and the distance to the Arctic Ocean is about 50 km. The yearly mean temperature is about -1°C, and permanent snow cover lasts 6-7 months. Yearly emission in 1990, just before the spider sampling, was about 250 000 tons S02, ca. 200 tons Cu, 300 tons Ni and 10 tons Co (see KOZLOV et al. 2009). This means that sulphur load was the same as in Monchegorsk smelter, central Kola Peninsula, but copper, nickel and cobalt loads were only 10 % of 81 Spiders at polluted sites in the subarctic Table 1: Composition of the spider fauna in the studied heavily (5 sites) and slightly pollut- ed areas (9 sites), Pechenganikel. Heavily polluted (H) Slightly polluted (S) S/H Species 18 58 Species/ site 3.6 6.4 1.8 Families 10 13 Individuals 101 1083 Ind./m2 20.2 120.3 6.0 range 3-32 71-266 Linyphiidae ind./m2 7.6 87.7 11.5 % of Linyphiidae (of spec.) 22.2 63.8 2.9 % of Linyphiidae (of ind.) 37.6 59.8 1.6 % of Lycosidae (of ind.) 10.9 11.7 1.1 % of the most dominant species 23.3 18.5 that in Monchegorsk. The area of complete damage to forests was 45 km2 and severe damage 300 km2 around the Pechenganikel complex; in these areas S02 concentration was over 80 g/m3, and annual dry sulphur deposition ca 4 g/m2(GYTARSKY et al. 1997, Norwegian Pollution Control Authority 2002). Spiders were collected at five study sites in heavily polluted areas, 2.5 — 7 km from the smelter, and at nine sites in slightly (or moderately) polluted areas, 6-55 km from the smelter. The average cover of ground and field layer vegeta- tion indicates the degree of pollution: 30 % coverage at heavily and 80 % at slightly polluted sites. Thus heavily polluted sites are more open and because the ground is often black, their microclimatic conditions in summer are more extreme, i.e. they are much drier and warmer than at slightly polluted sites. Ground- and soil-living spiders were collected, in 1991-92 from 25 x 25 cm squares by hand-sorting. The field work was done by Dr. Galina G. Koneva (Moscow) and identification was by the author. The material consists of ca 620 identifiable spider speci- mens (of the total 1185 individuals) and is deposited in the Zoological Museum, University of Turku, Finland. Results Great differences were found (Table 1) in the den- sity of spiders in heavily and slightly polluted areas, the averages being 20 and 120 ind./m2 (ranges 3-32 and 72-266 ind./m2) respectively. The average spider density was 6-fold greater and the density of Linyphiidae specimens even 11. 5 -fold at slightly polluted sites. Also spe- cies number/site was higher at slightly polluted sites: 1.8-fold. Altogether, 18 and 58 species of spiders were caught in heavily (5 sites) and slightly (9 sites) polluted areas, respectively. In- terestingly, as many as 10 fami- lies but only 18 species were found at heavily polluted sites, and at slightly polluted sites the material was much more species-rich, but represented no more than 13 families. The theridiid Robertus scoticus clearly dominated at the heavily polluted sites (Table 2: 23.3 % of identifiable specimens), and was the second most abundant at the slightly polluted sites (Table 3: 13.5 %). Other abundant species at heavily polluted sites were Neon reticulatus (9 .6 %), Thanatus formicinus (9.6 %), Xysticus audax (8.2 %), Agyneta gulosa (5.5 %), and Alopecosa aculeata (5.5 %). Of these, especially^. Table 2:Total number of spider specimens found at five most polluted and damaged sites (2.5 - 7 km from the smelter), Pechenganikel. Robertus scoticus 17 23.3 % of identifiable Neon reticulatus 7 9.6 Thanatus formicinus 7 9.6 Xysticus audax 6 8.2 Agyneta gulosa 4 5.5 Alopecosa aculeata 4 5.5 Haplodrassus sp. 4 5.5 Dictyna uncinata 3 4.1 Evarcha falcata 3 4.1 Heliophanus sp. 3 4.1 Micaria alpina 3 4.1 Scandichrestus tenuis 3 4.1 Xysticus obscurus 3 4.1 Tapinocyba p aliens 2 Pardosa palustris 1 Tetragnatha sp. 1 Tibellus maritimus 1 Walckenaeria antica 1 Number of species 18 Number of identifiable specimens 73 82 S. Koponen Table 3:Total number of specimens of the most abundant spiders found at nine slightly polluted sites (6 - 55 km from the smelter), Pechenganikel. Tapinocyba pallens 101 18.5 % of identifiable Robertus scoticus 75 13.7 Maso sundevalli 52 9.5 Alopecosa aculeata 45 8.2 Micrargus herbigradus 29 5.3 Hahnia ononidum 26 4.8 Macrargus rufus 18 3.3 Centromerus arcanus 16 2.9 Pardosa hyperborea 15 2.7 Evarcha falcata 12 2.2 Agyneta gulosa 11 2.0 Palliduphantes antroniensis 11 2.0 Xysticus obscurus 11 2.0 Minyriolus pusillus 10 Microcentria pusilla 7 Neon reticulatus 7 Sisicus apertus 7 Macrargus multesimus 6 Porrhomma pallidum 6 Xysticus audax 6 Cnephalocotes obscurus 5 Walckenaeria karpinskii 5 Gnaphosa microps 4 Micaria alpina 4 Pardosa palustris 4 Pocadicnemis pumila 4 Scotinotylus alpigena 4 Scandichrestus tenuis 4 Haplodrassus signifer 3 Hilaira herniosa 3 Microneta viaria 3 Tenuiphantes mengei 3 Zorn nemoralis 3 Bolyphantes luteolus 2 Cercidia prominens 2 Clubiona trivialis 2 Gnaphosa sticta 2 Gonatium rubens 2 Pelecopsis mengei 2 Robertus lyrifer 2 Semljicola faustus 2 Tenuiphantes tenebricola 2 Thanatus formicinus 2 Agnyphantes expunctus 1 Agyneta conigera 1 Agyneta subtilis 1 Bolehthyphantes index 1 Decipiphantes decipiens 1 Dictyna sp. 1 Diplocentria bidentata 1 Gonatium rubellum 1 Oreonetides vaginatus 1 Tetragnatha sp. 1 Tibellus maritimus 1 Walckenaeria dysderoides 1 Walckenaeria lepida 1 Walckenaeria nudipalpis 1 Zornella cultrigera 1 Number of species 58 Number of identifiable specimens 547 aculeata and also Agyneta gulosa were found in good numbers at slightly polluted sites. Altogether, four linyphiid species (i.e., Agyneta gulosa , Scandichrestus tenuis, Tapinocyba pallens, and Walckenaeria antica) were found among the total 18 species (22 %) at heavily polluted sites. Only one linyphiid was among the six most abundant spider species (Table 2 ). The percentage of Lycosidae specimens was about 1 1 % (Table 1). The most abundant species at slightly polluted sites (Table 3) were Tapinocyba pallens (18.5 %), Robertus scoticus (13.7 %), Maso sundevalli (9.5 %), Alopecosa aculeata (8.2 %), Micrargus herbigradus (5.3 %), Hahnia ononidum (4.8 %), and Macrargus rufus (3.3 %). The dominant species of the slightly polluted sites, Tapinocyba pallens , was found at the heavily polluted sites but only in small numbers. About 64 % of all species and 60 % of specimens at slightly polluted areas belonged to the family Linyphiidae. Three linyphiids were among the six most abundant species. The percentage of Lycosidae was 12 % of all individuals caught (Table 1). Discussion Leg deformations or other injuries caused by pollu- tion, could not be found in spiders collected at the polluted sites. Deformations have been observed previously in, e.g., oribatid mites near a smelter (EEVA & PENTTINEN 2009). The average spider density (about 120 ind./m2) at slightly polluted sites was of the same magnitude as found previously in natural northern boreal or sub- arctic forests (e.g. KOPONEN 1977). Also the species composition and the great proportion of Linyphiidae among species and specimens in slightly polluted ar- eas resembled the situation in natural forest habitats. The family Linyphiidae is known to dominate in Spiders at polluted sites in the subarctic 83 the ground layer of northern areas (e.g. KOPONEN 1993). Proportions of Lycosidae specimens were almost equal and rather low at heavily and slightly polluted sites, i.e. 11 % and 12 %, respectively In southern Finland, Xerolycosa and Alopecosa species dominated at the most polluted sites near a smelter (KOPONEN 8cNlEMELÄ 1994). On the other hand, lycosids were not found in the eroded industrial barren area close to the Monchegorsk smelter, central Kola Peninsula (KOPONEN 8c KONEVA 2006). However, different collecting methods, pitfall traps in southern Fin- land and hand-sorting both in Pechenganikel and Monchegorsk, must be borne in mind. When spiders along a pollution gradient were studied by KOPONEN 8c KONEVA (2006) near the Monchegorsk smelter complex, Robertus scoticus and Agyneta gulosawere also found there in the most pol- luted areas (black, dead barren sites 2.5-5 km from the smelter). In addition to these two species, only Steatoda phalerata occurred at the most polluted sites. S. phalerata was not found in the Pechenganikel area at all; which may have conditions too harsh for this thermophilous species. Of the present species found at heavily polluted sites, Micaria alpina , Alopecosa aculeata , Evarcha falcata and Tapinocyba pallens were caught in good numbers at rather heavily polluted sites in Monchegorsk (10 km from the smelter). The spider densities near the Monchegorsk smelter were lower than in the present study area (KOPONEN Sc KONEVA 2006), probably indicating the effect of much greater copper and nickel emissions at Monchegorsk (see Material and Methods). Emissions of heavy metals and especially sulphur dioxide have decreased greatly since the field work of this paper. According to KOZLOV et al. (2009), the decrease in emissions from 1985 (heavy pollution period) to 2005 was for S02 about 70 %, Co 75 %, Cu 20 %, and Ni 15 %. However, pollution in the study area has continued all the time, and emissions at the beginning of this century were still above the critical level for sensitive biota (NORWEGIAN POLLUTION CONTROL Authority 2002). Thus the environ- ment for the ground-living fauna is nowadays not better than in the early 1990s. Acknowledgements I wish to thank Dr. Galina G. Koneva (Moscow) for col- lecting and sorting the spider material, and Dr. Mikhail V. Kozlov (Turku) for organizing a trip to the Pechenganikel area in 2006. References BENGTSSON G. 8c S. RUNDGREN (1984): Ground-living invertebrates in metal-polluted forest soils. - Ambio 13: 29-33 CLAUSEN I.H.S. (1986): The use of spiders (Araneae) as ecological indicators. - Bulletin of the British arachno- logical Society 7: 83-86 EeVAT. Sc R. PENTTINEN (2009): Leg deformities of ori- batid mites as an indicator of environmental pollution. - Science of the Total Environment 407: 4771-4776 - doi: 10.1016/j.scitotenv.2009.05.013 GytarskyM.L., R.T. Karaban 8cI.M. Nazarov (1997) On the assessment of sulphur deposition on forests growing over the areas of industrial impact. - Environ- mental Monitoring and Assessment 48: 125-137 - doi: 10.1023/A1005744301889 KONEVA G.G. (1993): Changes in soil macrofauna around ’’Severonikel” smelter complex. In: KOZLOV M.V., E. HAUKIOJA Sc V.T. YARMISHKO (Eds): Aerial pollution in Kola Peninsula. Kola Scientific Center, Apity. pp. 362-364 KONEVA G.G. &S. KOPONEN (1993): Density of ground- living spiders (Araneae) near smelter in Kola Peninsula. In: Kozlov M.V., E. Haukioja Sc V.T. Yarmishko (Eds): Aerial pollution in Kola Peninsula. Kola Scientific Center, Apity. p. 365 KOPONEN S. (1977): Spider fauna (Araneae) of Kevo area, northernmost Finland. - Reports from the Kevo Subarctic Research Station 13: 48-62 KOPONEN S. (1993): On the biogeography and faunistics of European spiders: latitude, altitude and insularity. - Bulletin de la Societe Neuchäteloise des Sciences Naturelles 116 (1): 141-152 KOPONEN S. (1999): Common ground-living spiders in old taiga forests of Finland. - The Journal of Arachnology 27: 201-204 KOPONEN S. 8cG.G. KONEVA (2006): Spiders along a pol- lution gradient (Araneae). - Acta Zoologica Bulgarica, Suppl. 1: 131-136 KOPONEN S. Sc P. NIEMELÄ (1994): Ground-living spi- ders in a polluted pine forest, SW Finland. - Bollettino della Accademia Gioenia di scienze naturali 26 (345): 221-226 KOPONEN S. Sc P. NIEMELÄ (1995): Ground-living ar- thropods along pollution gradient in boreal pine forest. — Entomologica Fennica 6: 27-131 Kozlov M.V., E. Haukioja 8c V.T. Yarmishko (Eds) (1993): Aerial pollution in Kola Peninsula. Kola Scien- tific Centre, Apatity. 419 pp. Kozlov M.V., E.L. Zvereva 8c V.E. Zverev (2009): Impacts of point polluters on terrestrial biota. Environ- mental Pollution 15. Springer, Dordrecht, Heidelberg, London, New York. 466 pp. 5. Koponen 84 MAELFAIT J.-P. & F. HENDRICKX (1998); Spiders as bio-indicators of anthropogenic stress in natural and semi-natural habitats in Flanders (Belgium): some recent developments. In; SELDEN P.A. (Ed): Proceedings of the 17th European Colloquium of Arachnology, Edin- burgh 1997. British arachnological Society, Dorcester. pp. 293-300 Norwegian Pollution Control Authority (SFT) (2002): Air pollution effects in the Norwegian-Russian border area. International cooperation on environmental issues and environmental protection in the polar areas, TA-1860, Oslo. 34 pp. http://www.sft.no/publikasjoner/ internasjonalt/1860/tal860.pdf [accessed 1.VII.2009] RYBALOV L. (2003): Population of soil-dwelling inver- tebrates of the old-growth spruce forest of the Nature Reserve Friendship. In: HEIKKILÄ R. &T. LlNDHOLM (Eds): Biodiversity and conservation of boreal nature. -The Finnish Environment 485: 206-211 TlKKANEN E. & I. NlEMELÄ (Eds) (1995): Kola Penin- sula pollutants and forest ecosystems in Lapland: final report of the Lapland Forest Damage Project. Ministry of Agriculture and Forestry & Finnish Forest Research Institute, Helsinki. 82 pp. Arachnologische Mitteilungen 40:85-93 Nuremberg, January 201 1 Ground spider communities in experimentally disturbed Mediterranean wood- land habitats Yael Lubin, Noa Angel & Nirit Assaf doi:10.5431/aramit4010 Abstract:The protected Mediterranean woodland habitats in Israel are undergoing tree encroachment, resulting in loss of open patches with herbaceous vegetation. We suggested that this process results in a ground spider community dominated by shade-loving species. At three Mediterranean woodland sites located along a rainfall gradient, we examined the effects on the ground-spider community of experimental removal of the woody vege- tation in 1000 m2 plots by cutting and overall plant biomass reduction by grazing and browsing by livestock. Pitfall traps were placed in replicated plots of four treatments (control, cutting, grazing/browsing, and cutting together with grazing/browsing) and in two different habitat patch types (open, woody). ANOVA and multivariate analyses were performed on family abundance by treatment and habitat patch type.Tree-cutting reduced the number of families in plots at two of the three sites. Grazing did not have a significant effect on the number of families or on the ground spider community composition. The spider community of cut-woody patches was more similar to that of open patches than to that of uncut woody patches. Most spider families separated along an axis of open versus woody patches, with woody habitat families predominating at all sites. Families typical of open habitats were positively associated with cut-woody patches as well. The overall effect on ground spider diversity of such manipulations may depend on the scale of habitat changes. Key words: diversity, family composition, ordination, patchiness, similarity Patterns of diversity and community structure are the consequence of a large number of factors operating at small and large scales. These effects of different factors may be deduced by investigating the naturally occur- ring patterns over a range of sites (e.g. OXBROUGH et al. 2005, De MAS et al. 2009). However, this is a static approach and differences among sites may result from historical as well as current factors. Large-scale habitat manipulation experiments provide an opportunity to examine changes in communities over time at multiple levels of organization and in different groups of or- ganisms (BROWN et al. 2001). Large-scale controlled manipulations will change habitat features in such a way as to provide insight into how specific changes affect communities (SHACHAK et al. 2008). In the present study, we use such an experimental approach to examine effects of a manipulation at the landscape scale on the composition of spider communities at the habitat scale in a Mediterranean ecosystem. Mediterranean landscapes are patchy, with patches consisting mostly of two habitat types: woody patches that are occupied by shrubs or trees, and contrasting Yael LUBIN, Noa ANGEL & Nirit ASSAF, Mitrani Department of Desert Ecology, Blaustein Institutes for Desert Research, Ben- Gurion University of the Negev, Sede Boqer Campus, 84990 Israel, e-mail: lubin@bgu.ac.il submitted: 19. 1.2010, accepted: 24.3.2010; online: 10.1.2011 open patches that are devoid of woody plants (NAVEH 1982, GABAY et al. 2008). In protected Mediterra- nean woodland areas in Israel, the proportion of area covered by woody patches increases along a climatic gradient from south to north with increasing annual precipitation and productivity (KADMON &c DANIN 1997). Open patches are mostly covered by herbace- ous vegetation that is green in the winter and spring months (January-April) and otherwise dry. Medi- terranean woodlands in Israel have been exposed to regimes of grazing and forest cutting for thousands of years (NAVEH &lDAN 1973). The combined effect of disturbance from intensive grazing and browsing by livestock, and removal of woody vegetation for fire- wood, has maintained the mosaic of open and woody patches in the Mediterranean landscape. With the establishment of nature reserves in the Mediterranean region of Israel, grazing and tree-cutting were largely eliminated; consequently the woodland became more closed and with reduced cover of grasses and annual plants (PEREVOLOTSKY 2006). Such changes could have positive and negative effects on biodiversity of different groups of organisms owing to a number of different processes that may act in different ways. For example, the increased structural diversity of trees may create more niches for arboreal arthropods, while increased shade may lower annual plant productivity and reduce arthropod abundance in the understory. 86 Y. Lu bin, N. Angel & N. Assaf The effect of encroachment of the woodland and subsequent re-opening by cutting and grazing on the diversity of a range of organisms was the aim of a multi-year investigation of biological diversity in three Mediterranean woodland sites along a preci- pitation gradient in Israel (SHACHAK et al. 2008). In this study we tracked the changes in community composition of ground-active spiders in woody and open patches following two large-scale manipulations of the landscape: 1) tree cutting and removal of the woody vegetation canopy and 2) grazing and brow- sing. The structure of the habitat is well-known to in- fluence habitat selection by spiders (e.g., UETZ 1991, Langellotto ScDenno 2004, Mallis &Hurd 2005) and thus was expected to affect community composition following the manipulation. We report on the changes following the manipulations, concen- trating on effects of cutting and woody vegetation removal on the spider community found in the two habitat patch types. Our working hypothesis was that creating large open areas in the landscape by removal of woody canopy will change the spider community composition from predominantly shade-loving taxa to a community dominated by taxa typical of open habitats, and that this change will be accompanied by a loss of shade-loving taxa. Methods Research sites and experimental design The study was conducted in three Long-term Eco- logical Research sites in the Mediterranean region of Israel, listed from south to north: Adulam - in the Adulam Nature Reserve near Bet Guvrin (31°40’ N, 34°50’ E, 200 m elevation, ca. 400 mm annual rainfall), Ramat Hanadiv - in the southern Carmel mountains near Zichron Ya’acov (32°30’ N, 34°55’ E, 120 m elevation, ca. 600 mm annual rainfall), and Meron - on the slopes of Mt. Meron in the Upper Galilee (33°15’ N, 35°25’ E, 850 m elevation, ca. 900 mm annual rainfall). All of these sites have a canopy of small trees (3-5 m height) and woody shrubs, do- minated by Quercus calliprinos and Pistacia palaestina at Meron (ZOHARY 1973), Phillyrea media , Pistacia lentiscus and Sarcopoterium spinosum at Ramat Hanadiv (BAR MASSADA et al. 2008) and Phillyrea media and Pistacia lentiscus at Adulam (MlLMAN 2007). At each site, 5-6 blocks were marked and within each block we established four plots of 1000 m2 each. Each plot within a block received a different treatment, such that the treatments were replicated across the blocks. The treatments were: 1) control (no manipulation of the habitat), 2) grazing and browsing, 3) cutting and removal of the woody vegetation from the plot, and 4) cutting and woody plant removal together with grazing and browsing. Grazing and browsing by livestock was introduced into the plots in the late spring (May-June) after the arthropod sampling. Livestock used were typical of the region, sheep at Adulam, goats at Ramat Hanadiv and cows at Meron. Grazing pressure was moderate and typical of the area, with the exception of Ramat Hanadiv where grazing was more intensive. Cutting and removal of the woody vegetation was done in the late autumn. The trees and woody shrubs were cut at the base and the above-ground parts were removed from the plot. Each site was cut twice during the four-year study, once in the first year and repeated in the second year, with the exception of Adulam, where cutting was repeated after a 2-year interval. We measured ground spider activity-density using pitfall traps, which were set for 5 days in the late spring of 2006 (April-May), the season in the Mediterranean ecosystem when many plants are flowering and there is new growth of herbaceous vegetation. Flowering in spring begins in the south and progresses northward, such that the peak of flowering at the northernmost site (Meron) is about one month later than at the southern site (Adulam). Pitfall trapping was timed at each site to the local flowering period. At each site, dry pitfall traps consisting of two plastic cups (10 cm diameter at the top x 10 cm height) placed one inside the other were dug into the ground such that the rim was flush with the surface. In each plot there were 18 traps: three traps in each of three patches beneath trees or woody shrubs and, similarly, three traps in each of three open patches with herbaceous vegetation cover. In cut plots, traps were placed in former woody patches; these were easily distinguished by the remaining base of the tree or shrub and an area of leaf litter surrounding it. The samples from the three traps in each patch were combined. Thus, the samples in all analyses represent three replicates per patch type per plot. The traps were positioned and kept closed 1-2 weeks before the trapping began to allow the ground to settle around the traps. The traps were checked every morning for 5 days and each day all spiders were collected and transferred to 70 % alcohol for later identification. All spiders were identified to family level (indi- viduals of unknown family were removed from the data set), and for some families adults were identified to species. The analyses below were performed at the Ground spider communities in Mediterranean woodlands family level in order to take advantage of the full data set. Classification by family is somewhat arbitrary, as species may be shifted from one family to another. Nevertheless, we justify this approach here because 1) species identification was possible for only some of the families, 2) juveniles constituted 16-40 % of the indi- viduals and could not be reliably identified to species, and 3) creating morpho-species, especially with large numbers of juveniles, would artificially augment the number of species. Species in a family often belong to a single foraging guild and thus analysis by family can provide an indication of composition in relation to the diversity of habitat use and foraging strategies. Finally, we are cautious about inferring species-level processes from those observed at the family level. CARDOSO et al. (2004), for example, showed a significant cor- relation between numbers of species and families at sites in Portugal, but nonetheless found that species richness could not be inferred from family richness. Statistical analysis Factorial AN OVA was performed on the total number of families at each site, with grazing and tree removal as factors and plots as sample replicates. The effect of the different blocks was not significant in any of the analyses and thus this was not entered as a factor. In order to test for differences in the spider com- munity composition we used multivariate analyses. The analyses were performed on the matrix of family abundance (fourth-root transformed) with plots as the sample replicates and block as a covariable. The environmental variables were the treatments (control, grazing, cutting, cutting and grazing combined) and habitat patch type (open and woody patches). Analysis of similarity (AN O SIM) was performed on a Bray-Curtis dissimilarity matrix of family ab- undance with Primer V5.2.2 (CLARKE 8c GORLEY 2006). ANOSIM was used first to compare family composition in plots of the different treatments (cut- ting, grazing) and then to compare between specific a priori combinations of treatment and habitat patch type. To further explore the relationship between the spider family abundance and environmental variables of treatments and patch types we performed constrai- ned ordinations. Detrended correspondence analysis (DCA) was performed to determine which ordination method to use. DCA showed that the length of the gradient was below 3 and therefore the axes were relatively short, implying that the response curves of the family scores were linear. Therefore, principal 87 components analysis (PCA) and redundancy analy- sis (RDA) are appropriate for these data. We used CANOCO software version 4.53 (TER BRAAK 8c SMILAUER 2004). The points in the two-dimensional PCA graphs represent sample scores of patch types per plot for the first two axes. Thus, the number of samples is equal to the number of plots in which the patch type appeared. In the PCA graphs, the closer the points are to each other, the more similar they are in their family composition. Redundancy analysis (RDA), also performed in CANOCO (TER BRAAK 8c SMILAUER 2004), was used to determine the environmental variables (treatments and patch type) that best explain the family composition and to determine which families correlate best with the different treatments and patch types. RDA is a form of direct ordination in which axes are constrained by environmental variables and like PCA, it is based on linear correlation responses of the family scores curves (LEPS 8c SMILAUER 2003). We used reciprocal averaging of family abundances and site scores (representing the treatment and patch type variables) and manual forward selection with 499 Monte-Carlo permutations, to choose the set of environmental variables to include in the model, selecting only the significant variables. In order to compare among the three sites, we limited the RDA to the 10 most abundant families in each site (11 in Meron, due to a tie between the abundances; see Table 1). These families represent about 90 % of the total abundance in each site. Finally, we quantified the contribution of the most abundant families to the differences in family composition between treatment and habitat patch type combinations, using the same Bray-Curtis dissimilarity matrix from the ANOSIM (see above). A similarity percentage analysis (SIMPER) was per- formed in PRIMER (v5.2.2) to calculate the average Bray-Curtis dissimilarity for each family, which is expressed as the average contribution of the family to the overall dissimilarity between pairs of treatment and patch-type combinations. Results Sample sizes were small (Table 1), reflecting the relatively low productivity of a seasonal, semi-arid landscape and the short sampling period. The do- minant families in all three sites were ground spiders (Gnaphosidae), jumping spiders (Salticidae), wolf spi- ders (Lycosidae) and ant-eating spiders (Zodariidae). These four families together constituted 78 %, 80 % 88 Y. Lubin, N. Angel & N. Assaf Table 1 :Total numbers of individuals of the different families collected by pitfall trapping in Mediterranean woodland habitats at three sites in Israel (listed from south to north according to increasing rainfall). The abundances of the ten families used at each site (1 1 at Meron) for RDA are shown in bold. Family Adulam Ramat Hanadiv Meron Agelenidae 13 42 10 Araneidae 4 6 3 Cithaeronidae 2 Clubionidae 3 7 Corinnidae 4 3 Dictynidae 1 1 Dysderidae 13 25 40 Eresidae 2 1 Filistatidae 2 16 3 Gnaphosidae 187 190 254 Hahniide 1 1 Hersiliidae 7 4 Linyphiidae 29 25 54 Liocranidae 2 3 Lycosidae 145 307 118 Miturgidae 4 1 22 Nemesiidae 1 Oecobiidae 20 2 Oonopidae 4 1 Oxyopidae 3 2 Palpimanidae 2 2 Philodromidae 6 5 Pholcidae 3 Pisauridae 4 2 6 Salticidae 124 66 91 Sicariidae 2 1 Scytodidae 4 14 13 Segestriidae 2 Theraphosidae 1 Theridiidae 6 10 14 Thomisidae 23 3 10 Zodariidae 92 65 134 Zoridae 1 Totals 700 786 808 and 74 % of the total abundance in Adulam, Ramat Hanadiv and Meron respectively. Tree cutting and removal reduced the number of families in Adulam (F120=13.67, p=0.0014) and in Meron (F1 16=26.45, p=0.006), but not in Ramat Hanadiv (Fj 16=5.0, p>0.5).The effect of grazing and the interaction between grazing and cutting were not significant at any of the sites (Fig. 1). The full AN OVA models (cutting, grazing and the interaction between them) explained 41 % of the variance in number of families in Adulam (F1 20=4.63, p=0.012) and 38.5 % in Meron (Fj 16=3.34, p=0.046), but only 17 % in Ramat Hanadiv (Fj 16= 1.1, ns). Analysis of similarity (AN O SIM) based on Bray- Curtis dissimilarity indices showed that family com- position differed significantly between cut and uncut plots at all sites (Adulam, global R=0.09, p<0.005; Ramat Handiv, global R=0.06, p=0.051; Meron, glo- bal R=0.08, p<0.05) while grazing had no significant effect. When we segregated the cutting treatment according to open and woody patch types, ANOSIM showed that open patches did not differ from one another in family composition, regardless of cutting treatment (Table 2). As cut and uncut open patches did not differ from one another, for the principal components analysis we segregated the samples into three groups: open patches (cut and uncut combined), uncut-woody and cut-woody patches. In the principal components analysis (PCA) there was no segregation along the first four axes according to grazing, and thus grazing is not shown in the PCA graphs (Fig. 2A, 2C, 2E; see RDA results below). The first principal components axis accounted for 27 % of the spider family variance at Adulam, 34 % at Ramat Hanadiv and 38 % at Meron (Appendix). In Adulam (Fig. 2 A) and Meron (Fig. 2E), the first axis separated the samples from woody and open patches, with the cut-woody samples falling closer to the open patches. At these two sites, the variance in the second axis (12 % and 10 %, respectively) was not clearly associated with any treatment or patch- type gradient in the samples. At Ramat Hanadiv, the separation of woody and open samples was along the second axis, which accounted for 16 % of the variance (Fig. 2C). As in Adulam and Meron, the cut-woody samples were closer to the open samples, and indeed, intermixed with them along the second axis. The first axis could not be attributed to either cutting or to the patch type. Adding the grazing treatment did not show a clear pattern along this axis and thus, the first axis represents variation from other unmeasured factors. In the redundancy analysis (RDA), the first two axes were correlated with the environmental variables of treatment and patch type (Figs. 2B, 2D, 2F; correlation coefficients: Adulam: 0.75 and 0.49 for axes 1 and 2 respectively, Ramat Hanadiv: 0.74 and 0.31, Meron: 0.78 and 0.62). However, the total variance in family composition explained by Ground spider communities in Mediterranean woodlands 89 Table 2: Analysis of similarity (ANOSIM) tables showing R (similarity index) for the comparison between pairs of treatment and patch type combinations at the three sites. The comparisons are arranged in decreasing order of R values. A high index of similarity (p<0.05; shown in italics) indicates that there was a significant difference in family composition between the two treatment-habitat types. Habitat patches in plots that were not cut are noted as 'woody' or 'open', while those in cut plots are noted as 'cut- woody' and 'cut-open'. A. Adulam Treatment -patch type R Significance cut-open vs. woody 0.474 0.001 cut-woody vs. woody 0.397 0.001 open vs. woody 0.391 0.001 cut-open vs. cut-woody 0.098 0.03 cut-open vs. open 0.031 0.2 cut-woody vs. open -0.034 0.8 B. Ramat Hanadiv Treatment -patch type R Significance open vs. woody 0.382 0.001 cut-open vs. woody 0.31 0.001 cut-woody vs. woody 0.293 0.001 cut- woody vs. open 0.006 0.4 cut-open vs. open -0.04 0.7 cut-open vs. cut-woody -0.06 0.85 C. Meron Treatment -patch type R Significance open vs. woody 0.7 0.001 cut-open vs. woody 0.564 0.001 cut-woody vs. woody 0.386 0.001 cut-woody vs. open 0.141 0.01 cut-open vs. cut-woody 0.005 0.4 cut-open vs. open -0.014 0.5 treatment and patch type was low in all three sites (Adulam: 22 %, Ramat Hanadiv: 13 %, Meron: 29 %). Grazing was not a significant variable in any of the sites (p>0.5). Both patch type and cutting were significant in Adulam (patch type: F=6.71, p=0.002; cutting: F=3.84, p=0.002) and in Meron (patch type: F=11.21, p=0.002; cutting: F=3.77, p=0.004). In Ramat Hanadiv patch type was the only significant explanatory variable (F=2.62, p=0.036); cutting was not significant (F=1.94, p=0.096), but was included in the analysis as it appears to influence the distribution of some families. Most of the spider families at the three sites were segregated along the axis of patch type, showing an Figure 1: Mean and 95 % Cl of the number of families in plots under two landscape manipulations, cutting and gra- zing, at three Mediterranean woodland sites (from south to north): Adulam (31 °40' N, 34°50' E, 200 m elevation, c. 400 mm annual rainfall), Ramat Hanadiv (32°30' N, 34°55' E, 120 m elevation, c. 600 mm annual rainfall) and Mt. Meron (32°00' N, 35° 20'E, 900 m elevation, c. 800 mm annual rainfall). affinity either to woody patches or to open patches. At Adulam and Ramat Hanadiv, the two southern sites, there were more families associated with woody patches than with open patches (Figs. 2B and 2D, respectively). Families with strong woody-patch affi- nity (indicated by the length of the arrows) included Axis 2 90 SAMPLES HI Woody open FI Woody - Cut Y Lubin, N. Angel & N. Assaf -1.0 1.0 -1.0 1.0 Ground spider communities in Mediterranean woodlands one of the four dominant families, namely the wolf spiders (Lycosidae), as well as sheetweb spiders (Li- nyphiidae), cobweb spiders (Theridiidae, mainly ju- veniles of Crustulina , Enoplognatha and Euryopis ), and woodlouse hunters (Dysderidae, mainly unidentified species of Harp acted). The families associated with the open, herbaceous patches were the remaining three numerically dominant families: the ground spiders (Gnaphosidae, mainly Pterotricha levantina Levy, 1995 and Zelotes scrutatus (O. P.-Cambridge, 1872) at Meron and Adulam and Pterotricha cambridgei (L. Koch, 1872) and Z. scrutatus at Ramat Hanadiv), jumping spiders (Salticidae, unidentified species of Aelurillus at all sites) and ant-eating spiders (Zodari- idae, e.g., Ranops expers (O. P.-Cambridge, 1876) and Zodarion lutipes (O. P.-Cambridge, 1872) at Meron, Palaestina expolita O. P.-Cambridge, 1872 and Z. lu- tipes at Ramat Hanadiv and Zodarion judaeorum Levy, 1992 at Adulam). Cutting had a positive influence on funnel-web spiders (Agelenidae, mainly juveniles of Agelena and Maimund) and wolf spiders (Lycosidae) at Ramat Hanadiv (Fig. ID). Analysis of similarity (ANOSIM) results (Table 2) showed that there were significant differences in family composition between cut-woody patches and woody patches (uncut) at all sites. We conducted a percentage similarity analysis (SIMPER) of family dissimilarity between these two patch types in order to determine which families contributed most to this difference. The results support the segregation of families along the axis of cut- woody versus woody patches as suggested in the RDA. Among the families that contributed the most to the community dissimila- rity between these two habitats at Adulam and Meron were Gnaphosidae and Salticidae (together, 36 % and 21 % contribution to dissimilarity at Adulam and Me- ron, respectively), with higher average abundance of both families in cut-woody patches and Linyphiidae Figure 2: Multivariate analyses of ground spider families at three sites Adulam (A, B), Ramat Hanadiv (C, D) and Meron (E, F). Figures A, C, and E are show the spider family composition per plot along the first (x) and second (y) principal component axes, coded by patch type (woody, open and cut-woody). Figures B, D, and F show plots of the spider families along the first two RDA axes repre- senting the environmental variables of habitat type (axis 1 , woody vs. open) and cutting (axis 2, cut vs. uncut). The spider family codes are: AGE=Agelenidae, DYS=Dysderidae, GNA=Gnaphosidae, HER=Hersiliidae, LIN=Linyphiidae, LYC=Lycosidae, MIT=Miturgidae, OEC=Oecobiidae, SAL=Salticidae, SCY=Scytodidae,THE=Theridiidae, THO=Thomisidae, ZOD=Zodariidae 91 (12.5 % and 11 % contribution, respectively) found only in woody patches. At Ramat Hanadiv, Lycosidae and Gnaphosidae together contributed 49 % of the dissimilarity between cut and uncut woody patches and both had higher average abundance in cut-woody patches. Discussion At two of the three sites, plots where trees had been cut and removed (with or without grazing) had fewer spider families on average than did control plots. The ground spider community of cut woody patches dif- fered significantly from that of uncut woody patches and was more similar to that of open patches. Grazing had no significant effect on family richness, nor did it influence the family composition at any of the sites. Intensive grazing and browsing by goats at Ramat Hanadiv reduced the cover of woody vegetation (BAR MASSADA et al. 2008), and yet we found no effect on spider family composition in our analysis. Grazing alone also had little effect on the species richness of herbaceous plants at all three sites (AGRA &Ne’EMAN 2009; Y. Lubin, G. Ne’eman & A. Per- evolotsky unpubl.). There may be several reasons for the lack of a strong grazing effect. First, the Mediter- ranean flora and fauna has a long history of grazing and organisms are well- adapted to it (NOY-MEIR et al. 1989, WARD 2005); second, grazing may have been insufficient in duration and intensity to produ- ce a change; and third, the sampling of herbaceous vegetation and of spiders took place several months after the livestock were removed from the plots and there may have been sufficient time for the new annual vegetation to emerge following winter rains and for spiders to re-colonise following the disturbance. At all sites, cutting and removal of the woody vegetation had a marked effect on the spider family composition at the patch level. Analysis of similarity (ANOSIM) gave low, but significant values of R for the difference between cut and uncut plots. The reason for the low global R values is that the open patches in both cut and uncut plots were similar in family composition, reducing the overall strength of the difference between the two treatments. The PCA diagrams show that the spiders from former woody patches (cut-woody) were more similar in family composition to those from open patches than to those from uncut woody patches. Thus, community composition shifted from families typically found beneath trees and shrubs toward families typical of open patches with herbaceous vegetation. The 92 Y. Lu bin, N. Angel & N. Assaf shift was not complete, however, as some cut-woody samples overlapped with the woody samples at all three sites, and there were significant differences in family composition between cut-woody and open habitats at two of the three sites. One reason for the incomplete shift may be that the cut trees re-sprout rapidly following cutting and at the time of sampling most cut trees had new growth to a height of 0.3-1 m, thus providing conditions that may allow shade-loving species to persist or to recolonise. Another option is that, owing to modification of soil characteristics around woody plants (SEGOLI et al. 2008), the litter and soil moisture conditions surrounding the cut tree change only slowly and thus spiders still perceive this habitat as a woody patch. The distinct family preferences for either woody or open patches are seen in the RDA, as this was the main axis of separation at all three sites. Most families were associated with the tree habitat, an association possibly related to a more favorable microclimate and higher prey abundance. Gnaphosids and salticids were notable exceptions being clearly associated with open patches. Most of the gnaphosid species at these sites are nocturnal (personal observation), while the jumping spiders are all diurnal, which suggests that these two taxa temporally partition the terrestrial niche of open, herbaceous vegetation patches. Differences in family composition between cut and uncut plots (ANOSIM) largely reflect dif- ferences between cut-woody patches and woody (uncut) patches. In the percentage similarity analysis (SIMPER), families that showed strong associations with either open or woody patches in the RDA also contributed the most to the dissimilarity between the cut-woody and woody patches. This again supports the view that open habitat species benefitted from tree cutting. Two families in particular, Agelenidae and Lycosidae, showed an association with cutting at Ramat Hanadiv (RDA) and also contributed the most to the dissimilarity between cut-woody and woody patches at this site (SIMPER). The funnelweb spiders (Agelenidae) at Ramat Hanadiv mostly belonged to a single species, Agelescape livida (Simon, 1875). These spiders construct their webs on shrubs and were able to take advantage of an abundance of websites on the regenerating trees (personal observation). A large proportion of agelenids and lycosids trapped at Ramat Hanadiv were juveniles (90 % and 77 %, respectively). Juvenile lycosids and agelenids are highly mobile (e.g., WALKER et al. 1999) and young individuals searching for suitable sites may have moved preferentially into the newly accessible cut-woody patches. At the other two sites, both of these families were associated with the woody habitat. However, overall there were lo- wer proportions of juveniles at Adulam (20 %) and Meron (16 %) than at Ramat Hanadiv (42 %), and in particular, the proportion of juvenile lycosids was lower at these two sites (39 % and 19 %, respectively) than at Ramat Hanadiv. Thus the shift from woody patches into the cut woody patches at Ramat Hanadiv may have been a consequence of the preponderance of mobile juveniles in these two families. In conclusion, opening the woodland by tree- cutting had a small, but significant effect on family richness, and a pronounced effect on spider family composition. In general, there was a shift of family composition from predominately woody habitat taxa to more open-habitat taxa. The patterns observed here at a family level should be verified at the species level once species identifications are available. The extent to which such changes will increase or decrease overall diversity at the landscape scale will depend greatly on the spatial scale of removal of the woody vegetation. Acknowledgments This study was part of a national project on patterns and me- chanisms of biodiversity in water-limited ecosystems, fun- ded by the Israel Science Foundation (grant #1077/03), the Israel Ministry of Infrastructure (Eshkol grant #3-2539), the Israel Nature and Parks Authority and Yad Hanadiv Foundation. N. Angel was supported by a fellowship from the Albert Katz School for Desert Studies, Ben-Gurion University of the Negev. N. Assaf was supported by a post- doctoral fellowship of the Blaustein Center for Scientific Cooperation and the Council for Higher Education. We thank especially I. Musli, E. Groner, A. Perevolotsky, G. Ne’eman and the many other participants in the project. This is publication no. 719 of the Mitrani Department of Desert Ecology. References AGRA H. 6c G. 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PEREVOLOTSKY (Eds.): Biodi- versity in drylands: towards a unified framework. Oxford University Press, Cambridge/UK. pp. 233-249 ZOHARY M. (1973): Geobotanical foundations of the Mid- dle East and adjacent areas. Fischer Verlag, Stuttgart. 739 pp. Arachnologische Mitteilungen 40:94-104 Nuremberg, January 201 1 Spatial and temporal structure of the spider community in the clay semi-desert of western Kazakhstan Tatyana V. Piterkina Abstract: The spatial and temporal structure of spider communities was studied in the clay semi-desert of the north-western Caspian Lowland, western Kazakhstan (49°23' N, 46°47' E).The soils and vegetation are complex, being composed of a mosaic of desert and steppe plant communities. Besides the native associations, there are plantations of different tree species.The ground-dwelling spider assemblages in the native habitats are the most diverse. The number of species inhabiting forest plantations is three times as small. Gnaphosidae is the leading family in the ground layer.They show high abundance and diversity levels during the whole season. Thomisidae, Lycosidae, Philodromidae, and Salticidae are abundant as well. The species diversity of herbage-dwelling spiders in different open native habitats is very similar. The spectrum of dominant families (Thomisidae, Oxyopidae, Araneidae, and Salticidae) and the seasonal dynamics of their ratio in desert and steppe associations have much in common. Spider assemblages of native and artificial habitats are characterised by change from multispecies polydominant spring-summer communities to impoverished imbalanced autumn ones. Seasonal changes in the species structure of mature spider groupings in native habitats are well pronounced, while the impact of seasonal conditions is even stronger than between-habitat differences. Complexes of typical species with different levels of habitat preference are revealed. Key words: Araneae, ecology, habitat preference, seasonal dynamics Caspian Lowland, a semi-desert zone (MlLKOV Sc Gvozdetsky 1986). Study area, material and methods The Dzhanybek plain is the most arid area in the Ciscaspian semi-desert due to both internal drainage and soil salinity, despite its northernmost location. The climate of the territory is characterised by high atmospheric drought and aridity. Hot summers and severe winters are typical: the summer temperatures exceed 40°C, the winter temperatures sink lower than -35°C. The average annual air temperature (for 1951-2003) is 7.3°C; 18°C during the warm period and -3.5°C during the cold period. The average annual precipitation (for 1951-2003) is 295 mm, ranging from 44 (in 1984) to 354 mm (in 1993) (SAPANOV 2006). The sharp disparity of heat and moisture causes the very low humidity of the territory. The evaporative power reaches 1000 mm, which is 3 times the total rainfall. In addition, the meteorological conditions of the region are characterised by long-term fluctua- tions with regular cyclic reiterations of drought and moist periods (RODE 1959, LlNDEMAN et al. 2005, SAPANOV 2006). Another characteristic feature of the study area is a well pronounced complex pattern of soils and vegetation, with a combination of typical desert, semi- desert and steppe habitats. The co-existence of such Spiders of steppe and semi-desert regions of the Palaearctic, unlike those of the temperate zone, are still poorly studied. There is some faunistic infor- mation (e.g. Ponomarev 1981, 1988, 2005, 2008, Ponomarev &, Tsvetkova 2003, Ponomarev &TSVETKOV 2004a, 2004b, POLCHANINOVA 1992, 1995, 2002, KOVBLYUK 2006, EFIMIK et. al. 1997, ESYUNIN Sc EFIMIK 1998, ESYUNIN et al. 2007, TU- NEVA Sc ESYUNIN 2003), but very little attention has been paid to such ecological aspects as the structure of populations, their dynamics, and the mechanisms of community function in arid and semi-arid conditions (ESYUNIN 2009). This paper is focused on studying the spatial and temporal structure of spider assemblages in the clay semi-desert in the Volga and Ural rivers’ interfluve. The research was carried out in the environs of the Dzhanybek Research Station of the Russian Acade- my of Sciences (49°23'N, 46°47'E), located on the border between the Western Kazakhstan Province of the Republic of Kazakhstan and the Volgograd Province of the Russian Federation. The area studied is a flat, nearly undrained plain in the north-western Tatyana V. PITERKINA, Laboratory of Synecology, Institute of Ecology and Evolution, Russian Academy of Sciences, Leninskii Prospect 33, Moscow 1 19071, Russia, piterkina@yandex.ru submitted: 1 9. 1.20 1 0, accepted: 25.4.20 1 0; online: 10.1.2011 Spider community in a semi-desert of W -Kazakhstan contrasting biotopes is caused by pronounced micro- relief and, consequently, differences in moisture, soil substrates and their properties (RODE & POLSKIKH 1961). Microelevations are occupied by plant communi- ties of the desert type, with Kochia prostrata, Artemisia pauctflora , and Salsola laricina on saline soils. The groundwater is saline. Forb-grass vegetation ( Stipa spp., Festuca valesiaca , Agropyron cristatum , etc.) on dark chestnut and meadow chestnut soils with fresh groundwater occupies microdepressions (down to 0.4 m deep); they represent steppe habitats. This mosaic of elements constitutes most of the territory. Large depressions (down to 1-1.5 m deep, area of 1-100 hectares) with steppe plant communities take up about 10-15 % of the area. These large depressions are best supplied with water, due to runoff from the surrounding area. Besides these mentioned native associations, there are 50-year-old plantations com- posed of different tree species. Material for this work was collected by the author (April-October 2004-2005) and Dr. K.G. Mikhailov (June- September 1984) in three native habitats (de- sert associations of microelevations, and steppe asso- ciations of large depressions, and microdepressions) and three artificial ones: oak ( Quercus robur) forest belts, oak patch in a park, and elm (Ulmus pumila) forest belts. The collections in the latter habitat took place only in 1984. In recent years, the vitality of the forest-belt has become very poor; the trees are very sparse so the conditions in it have approached those of an open habitat. Traditional collecting methods were used: pitfall trapping (one transect — 10 traps), hand-sorting of soil and litter samples (0.25 x 0.25 m, 10 samples) and sweeping (one sample - 4 x 25 sweeps, 3 times a day, at 00:00, 8:00 and 16:00). Sampling was carried out every 7-10 days. Pitfall traps were set in microeleva- tions, microdepressions and woody plantations. Soil and litter samples were taken in all studied habitats. As the plantations had a rather poor and scattered herbaceous layer, sweeping was undertaken only in native habitats. The material includes a total of 15000 pitfall days, 570 soil and litter samples, and 268 sweeping samples. One of the most important features of the spider population in the clay semi-desert is its strongly pronounced seasonality and vertical stratification. Thus, I analysed the structure of spider complexes separately by layer, i.e. ground and herbaceous layers, 95 and seasons, i.e. spring, summer and autumn. When calculating the ratio of families, I considered spiders of all instars. With respect to the seasonal changes in species compositions I used mature spiders only, although I suggest that the differences revealed might reflect certain phenological trends. Taxa with a relative abundance of >5 % were considered predominat. The habitat preference of species was calculated using Pesenko’s coefficient (if.) (PESENKO 1982), which represents a mathematical transformation of the share of a species in a single biotope to its share in all other biotopes: R. = (n../N - n./N)/(n../N. + n./N), y ij j i y j 1 where n.— number of specimens of /-species in samp- les from y-biotope with total volume A.; n. — number of specimens of /-species in all other biotopes with total volume N. Single records of species were omitted from the calculation. The choice of this coefficient was based on the variety of the collecting methods used, which caused the heterogeneity of the data obtained and the diffi- culties in their unification. Using relative indices (not absolute ones) simplifies the interpretation of data and makes miscellaneous information comparable. The value of the coefficient ranges from -1 (absolute avoidance) to +1 (absolute preference). Statistical data analysis was performed using Statistica 6.0. Results About 20000 spider specimens were captured and studied, with about 7000 of these spiders being ma- ture. Altogether, 172 species from 88 genera and 21 families were recovered. Taking into account the scant information published previously, the spider fauna of the Dzhanybek Station amounts to 184 species from 93 genera and 22 families. A checklist and the distribution of species between the study habitats has been made available elsewhere (PlTERKINA 2009, PlTERKINA & MIKHAILOV 2009). Since the time of these mentioned papers some taxonomical changes have taken place or some identifications were refined, thus some species names may not coincide. Namely, Ero sp. turned out to be Ermetus inopinabilis Pono- marev, 2008, Theridion cf. uhligi Martin, 1974 - T. uhligi , Thanatus constellatus Charitonov, 1946 - T. oblongius cuius (Lucas, 1846), and Eresus cinnaberinus (Olivier, 1789) - E. kollari Rossi, 1846. 96 T. V. Piterkina Species structure of spider communities and its seasonal dynamics The communities of ground-dwelling spiders in the native habitats - microelevations and microdepres- sions - are the most diverse (about 90 species). The number of species inhabiting forest plantations is three times as small (about 30 species) (Tab. 1). The activity of spiders in the open habitats fluc- tuates from 20 to 70 ind. / 100 pitfall-days, with the highest numbers in spring and summer. The amp- litude of its fluctuation is much higher in the forest plantations (from 3-4 to 100 ind./100 pitfall-days). The density of the spider population, based on soil and litter samples, reaches its highest values in autumn (up to 117 ind. /m2). Gnaphosidae is the dominant family in the native associations. They exhibit high abundance and diver- sity levels (about 50 %) during the whole vegetation season, this being quite typical of arid and semi-arid landscapes. The proportions of Lycosidae and Saltici- dae are less, but also stable. Linyphiidae predominate in spring and autumn, Oxyopidae in summer, Titanoe- cidae in spring and summer, Thomisidae in summer and autumn. The dominant complex of the tree plan- tations is less diverse. The proportion of Gnaphosidae is significantly lower than in native habitats (about 20-30 %), while the abundance of Thomisidae is high and stable during the entire vegetation season (about A 1 -Pearson r Complete Linkage 30-50 %). Pisauridae show a peak in their abundance in spring and autumn, whereas Liocranidae peak in summer. Seasonal change in species dominance is well pro- nounced and the species set is relatively stable across different years (Tab. 1). For example, in the desert habitats, T. veter anica , Haplodrassus cf. soerenseni , E. eltonica , D. ros trains, Z. orenburgensis predominate in spring populations in both years of study. The stable summer dominants are P. braccatus , H. horridus , Oxy- opes c£ xinjiangensis , D. rostratus and Z. orenburgensis. The autumn populations are rather imbalanced. Cher- acteristic is a high level of predominance of 1-2 species that can change in different years (Z. orenburgensis , X. marmoratus or D. rostratus). The dominant complexes of oak plantations have much in common and include several species abundant during the whole vegetation season (Z. gallicus, O. pratkola, X. luctator) (Tab. 1). The species diversity of herbage-dwelling spiders in the open native habitats is very similar: about 50 species (Tab. 2). The abundance ofhortobiotic spiders fluctuates with a high amplitude, reaching its maxi- mum in summer (about 100 ind. / 100 sweeps). The spectrum of predominating families and the seasonal dynamics of their proportion in desert and steppe associations have much in common. Uloboridae and Linyphiidae are abundant in spring, Araneidae and Oxyopidae in spring and autumn, Salticidae in sum- B 1 -Pearson r Complete Linkage Figure 1 : Clustering the mature spider complexes for separate seasons: A - ground-dwelling spiders, B - herbage-dwelling spi- ders. Habitats: 1 : microelevations, 2: microdepressions, 3: large depressions, 4: elm shelter-belt, 5: oak shelter-belt, 6: oak patch in a park. Seasons: spr - spring, sum - summer, aut - autumn. Spider community in a semi-desert of W -Kazakhstan 97 mer and autumn. Philodromidae, Clu- bionidae and Miturgidae are numerous during the whole vegetative period. The seasonal change of the pre- dominant complexes of species is also well-pronounced (Tab. 2). In spring and, especially, summer, the sets of abundant species are not stable in different years. On the contrary, the autumn populations of all habitats are very similar. They are mainly formed by two species, Xysticus marmoratus and X. striatipes. Co-domi- nance of Cheiracanthium cf. virescens adds originality to the autumn assemblages of microelevations, E. michailovi to those of microdepressions, and H. lineiventris to those of large depressions (Tab. 2). Clustering the mature spider com- plexes for separate seasons (Fig. 1) yielded interesting results. Two large clusters were revealed among ground- dwelling spiders: assemblages of native biotopes and of forest plantations (Fig. 1 A). Within them, the populations were not united by habitat, as one would expect, but by season. The cluster of open habitats includes populations of microelevations and microdepressions during spring, summer and autumn. Microclimatic conditions in woody plantations were presumably compara- tively smoother, even though no direct abiotic measurements were taken. The cluster of artificial forests appears to be less differentiated. The same tendency is also obvious when clustering the herbage-dwelling spider complexes: three pronounced clusters united spring, summer and autumn assemblages of microelevations, microdepressions and large depressions respectively (Fig. IB). Habitat preferences of species Spider assemblages of desert associa- tions are the most specific. The share of species collected only in microelevations is highest (24 %), whereas it is half this in the other biotopes. Most of unique species, with few exceptions, exhibit low abundance levels and hardly play coenotic roles (Tab. 3). "O c D 0 01 cu OJ 'u cu Q. 1/1 Oh c n CJ § 00 .■«H a -c go £ a c -a r JR N N Q O 1 1 «S § R ^ e? 3; O <3 ^ On -id N Q CO b R

an q a" a" r ^oq ÜSNÜÜ< \0 — ' g CN g .Sq. ‘'I | -S ' • S a ?> 5 3 J3 " 73 a -S . pR ^ ^ N r S ju pR- cq Si g 8 § «Sä § ■«s ’S “i S Is a” ^ -cr-S Or R '5T £ ^ S -S g 3 a 1*8 51| § R 5S -R -<5 OR Se a -9 or ar -gs Si cq N ^ Ü ^ O cj '■Ö ’u OJ fcv 0 ^ R ?s ts'1 X ■§ ^ gf? N^^°0 R vO «3 R II S « JH'* ■v s s 5 f?-£ ^ ^ O 03 Qq N R d- R -1 R ^ S R R -3 £ ’S R » JS N Ö ^ Q c? R R S$ ^ ^ js -R g s ■VS 'S 55 ■i ru <3 iS 00 rs ^ ^ X o ^ R3 '■§ I I g ^ *X). •«V, .3 •§ s s "§ ^ >0 R R R N*^s -S > R VO N ^ Q Ü Table 2: Species structure of populations of mature herbage-dwelling spiders. Numbers in brackets shows relative abundance in %. Microelevations (desert habitats) Spider community in o semi-desert of W -Kazakhstan 99 Oh CD S .1 XÖ X>? < s .« | § •CX, 5- S 8 §?.§ I X Q Ö E3 m ^ 'I * | | § * 1 1! | ^ a X a ^ § I 1? si Q S ^ .b l^-s hß ^ ^<3 .-Sh- a S 00 3 i f. E-H o I 'U ■^LO 3 s 3 « 'S r r a e r a a a -h a «s 'k, -<) ?ü U 0 1 § 1| 5 5 S K ja £ a “'R « £ § 5 R I C§ & O ~R S 3 1 ll g I R“ lit l si a 'O "a X "S Ö5 A a X S £ vo 1 f § f s Ö ^ rX q ?X ^ ChTN V (Nj nTs a hO O ^ *o x X x I X 8 -b -9 00 X_ |o § § p a no § -S -2 'tJ ON Eg X Sä *»o a S X R^-S R § T— 1 1 r ‘S -§ -3 ulepeira a, '\ctyna late cyopes cf. : ?. liophanus lanatus ob leiracanth ) ^ Q Ö ^ X Ü X R c? S ‘§xoo & X I x c+h & Se -S VO o S ^ S -ö o H ■S ~§ .3 <5 Cx) 'S 'S 'S a R R R g a a ?s p g 'S a ^ . x X . X CJ bJj fH ih^N *- r> t-* 'X? rvTN 0 X^ 0 § Vh Oh X CO CO ^ X 3 R g s x ^ X ^ S £ a o ON 3 T3 C^c- § b S Njh ' X R Xi at U c-\ Q o 0.7) and relatively low (0.3>/y>0.7) levels of habitat preference, was revealed for eac ii habitat (Tab. 4). Discussion It is well known that the denser the vegetation the greater is density of spiders, and the greater the diversity of vegetation the greater the spider species diversity (DUFFEY 1962). But the spider assemblages of both the ground and herbaceous layers of open na- tive habitats (microelevations, microdepressions and large depressions) are very similar not only in species diversity but also in density. This was rather surpris- ing as the low, sparse and rather poor desert plant communities look much more miserable compared to the dense forb-grass vegetation of steppe habitats. This reveals a complex of species well adapted to the extreme conditions of desert associations. On the contrary, the communities of forest plantations ap- pear to be significantly impoverished. The poorness of soil fauna under Dzhanybek plantations was dem- onstrated for other arthropods as well (CHERNOVA 1971, KRIVOLUTSKII 1971, etc.). Calculating the level of habitat preference (if .) revealed complexes of typical species for each habitat (Tab. 4). In spite of mosaic structure and a compara- tively small size of desert and steppe elements (some tens of square meters) in complex Northern Caspian semi-desert, the spider groupings formed on them are rather specific and contain sets of species asso- ciated with the particularities of the substrate (soil) and vegetation of those elements. The complexes of typical species of native habitats - microelevations and microdepressions - are the richest (35-40 species). Most of the typical species in desert associations are dwellers of arid and semi-arid landscapes: these are steppe (D. rostratus , Z. orenburgensis , G. steppica, etc.), semi-desert (S. crassipedis , T. mikhailovi , W. stepposa) and steppe-desert species (H. horridus , O. lugubris)-, with some participation of nemoral-steppe and ne- moral ones. The share of steppe species ( B . cinerea , G. leporina , H. isaevi, etc.) decreases significantly in associations of microdepressions and large depressi- ons, while nemoral-steppe ( E . michailovi , Z. electus , T. arenarius , etc.) and nemoral-subtropical species (P. chrysops , P.fasciata,A. lobata , etc.) prevail. Most of the typical species are quite abundant and predominate in these biotopes. In addition, there is a complex of species which can inhabit several types of native habitats with simi- lar probability levels (except for woody plantations). These are Trichoncoides cf. pis cat or, G. bituberculata, A. v-insignitus, A. cursor , P. histrio, Z. segrex, etc. Complexes of typical species of woody plantations are poor and include 12-15 species, although the level of habitat preference is very high (Tab. 4). Most of them are nemoral species. Populations in the plan- tations are very likely composed of highly eurytopic species ( D.pusillus , Z.gallicus , P mirabilis ) and typical dwellers of intrazonal associations ( S . zimmermanni , T. schineri ) with a small participation of forest species (O. praticola) which could be introduced with plant material. On the other hand, the structure of spider as- semblages is heavily determined by macroclimatic conditions and their seasonal changes. The analysis of seasonal features of population structure shows that the spring and summer spider assemblages of both ground and herbaceous layers are characterised by high species diversity levels and a relatively high number of predominating species, as opposed to the impoverished, imbalanced autumn populations (Tab. 1-2). The same pattern was recovered by ESYUNIN (2009) for spiders of steppe and steppe-like habitats in the Ural Mountains. Clustering the spider complexes for separate seasons confirmed the prevailing role of seasonal differences in species proportions for mature spider groupings of native habitats when comparing be- tween-habitat differences (Fig. 1). The populations of native associations were not united by habitats, but by seasononality. A similar trend has been also shown by ESYUNIN (2009) for the spider populations of steppe-like habitats in the Ural Mountains. It is interesting to note that such a tendency was Spider community in o semi-desert of W -Kazakhstan 101 Table 4: Pesenko's coefficient of a habitat preference {Fij) of spiders. Species are grouped according to their preference to a certain habitat. Within the groups species are ranked in order of decreasing the values of Fij. Grey background: high level of habitat preference (0.7 < F..< 1.00); bold: relatively low level of habitat preference (0.3 < F..< 0.7). Habitats as in Fig. 1. Species Number of specimens Habitats 1 2 3 4 5 6 Chalcoscirtus nigritus 17 1.00 -1.00 -1.00 -1.00 -1.00 -1.00 Heriaeus horridus 54 1.00 -1.00 -1.00 -1.00 -1.00 -1.00 Lepthyphantes spasskyi 7 1.00 -1.00 -1.00 -1.00 -1.00 -1.00 Micaria guttulata 7 1.00 -1.00 -1.00 -1.00 -1.00 Nomisia ausser eri 4 1.00 -1.00 -1.00 -1.00 -1.00 Robertus arundineti 5 1.00 -1.00 -1.00 -1.00 -1.00 -1.00 Urozelotes sp. 4 1.00 -1.00 -1.00 -1.00 -1.00 Evippa eltonica 189 0.98 -0.94 -1.00 -1.00 -1.00 Titanoeca veteranica 115 0.96 -0.90 -1.00 -1.00 -1.00 Aelotes orenburgensis 204 0.93 -0.82 -1.00 -0.94 -1.00 Drassodes rostratus 153 0.91 -0.78 -1.00 -1.00 -0.92 -1.00 Lasaeola tristis 23 0.91 -0.75 -0.85 Phaeocedus braccatus 47 0.91 -0.77 -1.00 -1.00 -1.00 Micaria pallipes 56 0.89 -0.71 -1.00 -1.00 -1.00 -1.00 Oxyopes cf. xinjiangensis 114 0.82 -0.92 -0.36 -1.00 -1.00 -1.00 Thanatus mikhailovi 22 0.80 -0.51 -1.00 -1.00 -1.00 -1.00 Microlinyphia pusilla 16 0.71 -0.26 -0.76 -1.00 -1.00 -1.00 Silometopus crassipedis 23 0.71 -0.35 -0.59 -1.00 -1.00 -1.00 Talanites mikhailovi 10 0.71 -0.35 -1.00 -1.00 -1.00 Trachyzelotes adriaticus 5 0.71 -0.35 -1.00 -1.00 -1.00 Talanites strandi 14 0.69 -0.31 -1.00 -1.00 -1.00 Xysticus marmoratus 278 0.66 -0.44 -0.41 -1.00 -1.00 -1.00 Gnaphosa lucifuga 79 0.65 -0.23 -1.00 -1.00 -1.00 -1.00 Theridion uhligi 6 0.64 -0.21 -1.00 -1.00 -1.00 Ozyptila lugubris 19 0.62 -0.18 -1.00 -1.00 -1.00 Theridion innocuum 8 0.62 -0.26 -0.54 -1.00 -1.00 -1.00 Drassyllus sur 33 0.60 -0.16 -1.00 -1.00 -1.00 Nurscia albomaculata 36 0.59 -0.15 -1.00 -1.00 -1.00 Ozyptila pullata 22 0.58 -0.13 -1.00 -1.00 -1.00 -1.00 Pellenes albopilosus 26 0.57 -0.11 -0.63 -1.00 -1.00 -1.00 Archaeodictyna consecuta 18 0.53 -0.41 -0.18 Ceratinella brevis 3 0.50 -0.02 -1.00 -1.00 -1.00 Euophrys frontalis 3 0.50 -0.02 -1.00 -1.00 -1.00 Gnaphosa steppica 72 0.50 -0.08 -1.00 -0.83 -0.77 Walckenaeria stepposa 3 0.50 -0.02 -1.00 -1.00 -1.00 Haplodrassus cf. soerenseni 102 0.42 0.05 -1.00 -0.88 -0.83 Aelurillus m-nigrum 5 0.39 0.13 -1.00 -1.00 -1.00 Cheiracanthium cf. virescens 99 0.36 -0.15 -0.20 -1.00 -1.00 -1.00 Phlegra bicognata 24 0.36 0.16 -1.00 -1.00 -1.00 Uloborus walckenaerius 26 0.35 -0.22 -0.12 -1.00 -1.00 -1.00 Aelotes caucasius 39 0.32 0.10 -0.08 -0.71 -1.00 Improphantes contus 7 -1.00 1.00 -1.00 -1.00 -1.00 Heliophanus flavipes 4 -1.00 1.00 -1.00 Phlegra fasciata 10 -1.00 1.00 -0.05 -1.00 -1.00 -1.00 Walckenaeria alticeps 4 -1.00 1.00 -1.00 -1.00 -1.00 -1.00 Haplodrassus kulczynskii 48 -0.94 0.98 -1.00 -1.00 -1.00 Trichopterna cito 169 -0.98 0.96 -0.43 -0.19 -0.91 -1.00 Berlandina cinerea 139 -0.83 0.92 -1.00 -1.00 -0.76 Cercidia levii 37 -1.00 0.92 -0.80 -1.00 -1.00 -1.00 Trichoncus villius 28 -0.79 0.92 -1.00 -1.00 -1.00 -1.00 Thanatus arenarius 140 -0.76 0.90 -0.54 -1.00 -0.91 -1.00 Zelotes electus 73 -0.96 0.90 0.31 -0.83 -0.77 Haplodrassus isaevi 37 -0.69 0.88 -1.00 -1.00 -1.00 102 T V. Piterkina Species Number of specimens Habitats 1 2 3 4 5 6 Gnaphosa leporina 24 -1.00 0.81 -1.00 -1.00 0.30 Thanatus atratus 45 -0.52 0.81 0.40 -1.00 -1.00 -1.00 Zelotes longipes 60 -0.62 0.79 -0.29 -0.63 -1.00 Evarcha michailovi 77 -0.92 0.77 -0.52 -1.00 -1.00 -1.00 Heliophanus koktas 19 -1.00 0.77 -0.50 Pardosa plumipes 5 -0.45 0.77 -1.00 -1.00 -1.00 Drassodes lapidosus 4 -0.33 0.71 -1.00 -1.00 -1.00 Clubiona genevensis 27 -0.26 0.66 -0.74 -1.00 -1.00 -1.00 Drassodes villosus 7 -0.25 0.66 -1.00 -1.00 -1.00 Alopecosa schmidti 31 -0.24 0.65 -1.00 -1.00 -1.00 Thanatus pictus 96 -0.11 0.57 -0.16 -1.00 -1.00 -1.00 Thanatus sp. 8 0.00 0.49 -0.05 -1.00 -1.00 -1.00 Alopecosa taeniopus 41 -0.65 0.47 0.25 0.30 -0.63 Agyneta saaristoi 30 -0.05 0.43 -0.54 -1.00 -0.39 -1.00 Haplodrassus signifer 45 0.05 0.42 -1.00 -1.00 -0.65 Eresus kollari 13 0.13 0.39 -1.00 -1.00 -1.00 Gnaphosa taurica 135 -0.31 0.38 0.35 -0.36 -0.10 Xysticus striatipes 426 -0.60 0.35 -0.03 -0.29 -0.80 -1.00 Agroeca maculata 61 0.15 0.33 -1.00 -1.00 -0.73 Philaeus chrysops 5 0.20 0.32 -1.00 -1.00 -1.00 -1.00 Simitidion simile 9 0.12 0.32 -0.54 -1.00 -1.00 -1.00 Scotargus pilosus 4 0.20 0.32 -1.00 -1.00 -1.00 Xysticus cristatus 43 0.20 0.32 0.22 -1.00 -1.00 -1.00 Trichoncoides cf. piscator 11 0.41 0.32 -1.00 -1.00 -1.00 -1.00 Zelotes segrex 15 0.26 0.26 -1.00 -1.00 -1.00 Aelurillus v-insignitus 30 0.20 0.13 -1.00 -1.00 0.01 Alopecosa cursor 34 0.26 0.27 -1.00 -1.00 -1.00 Gibbaranea bituberculata 82 0.09 0.02 -0.10 -1.00 -1.00 -0.90 Philodromus histrio 18 -0.01 -0.11 0.12 -1.00 -1.00 -0.90 Oxyopes lineatus 43 -1.00 -0.62 0.84 -1.00 -1.00 -1.00 Neoscona adianta 23 -1.00 -0.42 0.73 Argiope lob at a 4 0.03 -1.00 0.69 Thanatus oblongiusculus 99 -0.51 -0.53 0.67 Aculepeira armida 53 -0.49 -0.53 0.66 -1.00 -1.00 -1.00 Oxyopes heterophthalmus 29 -0.60 -0.43 0.64 -1.00 -1.00 -1.00 Thomisus onustus 20 -0.71 -0.23 0.54 Dictyna latens 45 0.02 -0.63 0.50 Agyneta spp. ( $ ) 29 -0.01 -0.38 0.34 -1.00 -0.25 -0.09 Heliophanus lineiventris 75 -0.38 0.01 0.22 -1.00 -1.00 -1.00 Heriaeus melloteei 10 -0.04 -0.15 0.18 -1.00 -1.00 -1.00 Pardosa xinjiangensis 6 -1.00 -1.00 1.00 -1.00 -1.00 Micaria rossica 7 -0.25 -0.13 0.92 -1.00 -1.00 Pseudeuophrys obsoleta 5 -1.00 -1.00 0.91 0.63 0.40 Ermetus inopinabilis 8 -0.33 -1.00 0.90 0.59 -1.00 Titanoeca quadriguttata 3 -1.00 -0.02 0.88 0.53 -1.00 Zelotes atrocaeruleus 6 -0.14 -0.02 0.88 -1.00 -1.00 Xysticus ninnii 85 -0.79 0.24 0.22 0.86 0.04 -0.42 Tibiaster djanybekensis 24 0.56 -1.00 -1.00 0.83 -0.49 -1.00 Zelotes gallicus 96 -1.00 -1.00 0.83 0.75 0.47 Mangora acalypha 5 0.23 0.49 -1.00 0.69 -1.00 -1.00 Cheiracanthium pennyi 24 -1.00 0.10 0.10 -1.00 1.00 -1.00 Pis aura mirabilis 105 -1.00 -1.00 0.13 0.91 0.37 Zora pardalis 56 -1.00 -0.68 0.10 0.88 0.28 Lathy s stigmatisata 93 -0.80 -0.25 -0.25 0.21 0.70 0.37 Titanoeca schineri 73 -1.00 -1.00 0.55 0.87 0.44 Xysticus luctator 198 -1.00 -0.98 -1.00 0.73 0.80 Spider community in a semi-desert of W -Kazakhstan 1 03 Species Number of specimens Habitats 1 2 3 4 5 6 Drassyllus pusillus 90 -1.00 -1.00 0.40 0.63 0.83 Sitticus zimmermanni 29 -1.00 -0.87 0.42 0.55 0.84 Ozyptila praticola 155 -1.00 -1.00 -1.00 -1.00 0.60 0.88 Xysticus robustus 10 -1.00 -1.00 -1.00 -1.00 0.68 0.84 Zelotes subterraneus 3 -1.00 -1.00 -1.00 -1.00 1.00 Philodromus cespitum 5 -1.00 0.10 0.10 -1.00 -1.00 1.00 Agroeca cuprea 4 -1.00 -1.00 -1.00 -1.00 1.00 not revealed for snout-beetles (Coleoptera, Curcu- lionoidae) investigated at the Dzhanybek Station during the same period. These phytophagous insects showed that the influence of between-habitat differ- entiation on the structure of their populations — which was determined by their close links with the plants on which they forage (KHRULEVA et al. in press) - was much stronger than seasonal changes. Spiders being a group of mobile generalist predators are more likely to be influenced by abiotic factors. Acknowledgements I would like to thank the managers of the Dzhanybek Research Station for the opportunity to work there. I am also thankful to Kirill G. Mikhailov for the material he collected in 1984 and for his valuable advice, as well as to the following arachnologist colleagues, Galina N. Azarkina, Alexander V. Gromov, Dmitri V. Logunov, Yuri M. Maru- sik, Vladimir I. Ovtsharenko, and Andrei V. Tanasevitch for their help in identifying some of the spider taxa. I am deeply indebted to all staff of the Laboratory of Synecology for their constant help and encouragement. Sergei I. Golovatch kindly checked the English of an advanced draft. The study was supported by the Russian Foundation for Basic Research, the Program “The Origin and Evolution of the Biosphere”, the Program for the Support of the Leading Academic Schools and Young Scientists. References CHERNOVA N.M. (1971): [Springtails of plantations in the Northern Caspian clayey semi-desert] In: RODE A.A. (Ed.): Zhivotnye iskusstvennykh lesnykh nasazhdenii v glinistoi polupustyne. Nauka, Moscow, pp. 24-33. [in Russian] DUFFLEY E. (1962): A population study of spiders in limestone grassland. The field-layer fauna. - Journal of Animal Ecology 31: 571-599. Efimik V.E., S.L. Esyunin & S.F. Kuznetsov (1997): Remarks on the Urals spider fauna, 7. New data on the fauna of the Orenburg Region (Arachnida, Aranei). - Arthropoda Selecta 6: 85-90 ESYUNIN S.L. (2009): Geographical variation in spider as- semblages (Arachnida: Aranei) of steppe and steppe-like habitats of the Urals, Russia. In: GOLOVATCH S.L, O.L. Makarova, A.B. Babenko & L.D. Penev (Eds.): Species and communities in extreme environments. Festschrift and a Laudatio in Honour of Academician Yuri Ivanovich Chernov. Pensoft Publishers & KMK Scientific Press, Sofia & Moscow, pp. 403-418 ESYUNIN S.L. & V.E. Efimik (1998): Remarks on the Urals spider fauna, 8. New and unidentified species from steppe landscapes of the South Urals. - Arthropoda Selecta 7: 145-152 Esyunin S.L.,T.K.Tuneva &G.S. Farzalieva (2007): The remarks on the Ural spider fauna (Arachnida, Ara- nei), 12. Spiders of the steppe zone of Orenburg Region. - Arthropoda Selecta 16(1): 43-63 Khruleva O.A., B.A. Korotyaev &T.V. Piterkina (in press): [Stratification and seasonal dynamics of the weevil (Coleoptera, Curculionoidae) assambleges in the Northern Caspian semi-desert]. - Zoologicheskii Zhurnal 90 [in Russian] KOVBLYUK M.M. (2006): [Gnaphosidae spiders (Arach- nida: Aranei) in Crimean fauna]. PhD Theses, Institute of Zoology, Ukranian Academy of Sciences, Kiev. 18 pp. [in Ukrainian] KRIVOLUTSKII D.A. (1971): [The population of oribatid mites in the soils of the Northern Caspian semi-desert and their changes under the influence of afforestation.] In: RODE A.A. (Ed.): Zhivotnye iskusstvennykh lesnykh nasazhdenii v glinistoi polupustyne. Nauka, Moscow, pp. 13-23 [in Russian] Lindeman G.V., B.D. Abaturov, A.V. Bykov & V. A. LOPUSHKOV (2005): [Dynamics of the vertebrate animal population in semidesert of the area east of the Volga river]. Institute of Forestry, Nauka, Moscow. 252 pp. [in Russian] MlLKOV F.N. &N.A. GVOZDETSKY (1986): [Physical geo- graphy of the USSR]. Vysshaya Shkola Pubis., Moscow. 512 pp. [in Russian] PESENKO Y.A. (1982): [Principles and methods of quanti- tative analysis in faunistical researches]. Nauka, Moscow. 288 pp. [in Russian] PITERKINA T.V. (2009): Spiders (Arachnida, Araneae) of the Dzhanybek Research Station, West Kazakhstan: a local fauna in a biogeographical aspect. In: GOLOVATCH S.I., O.L. Makarova, A.B. Babenko & L.D. Penev (Eds.): Species and communities in extreme environ- 104 T V. Piterkina merits. Festschrift and a Laudatio in Honour of Acad- emician Yuri Ivanovich Chernov. Pensoft Publishers & KMK Scientific Press Sofia, Moscow, pp. 335-356 Piterkina T.V. & KG. Mikhailov (2009): [Annotated check-list of spiders (Aranei) of Dzhanybek Station.] In: TlSHKOV A. A. (Ed.): Zhivotnyie glinistoi polu- pustyni Zavolzhya (konspekty faun i ekologicheskiye kharakteristiki). KMK Scientific Press, Moscow, pp. 62-88 [in Russian] POLCHANINOVA N.Y. (1992): [Spiders of Proval’skaya Steppe]. In: Fauna i ekologiya paukov, skorpionov i lozhnoskorpionov SSSR. - Trudy ZIN AN SSSR 226 (for 1990): 98-104 [in Russian] POLCHANINOVA N.Y. (1995): [Spiders (Arachnida, Aranei) of the “Askania-Nova” Reserve]. In: Fauna i ekologiya paukov SSSR: Mezhvuzovskii sbornik nauchnikh tru- dov. Perm University, Perm (for 1994). pp. 89-98 [in Russian] POLCHANINOVA N.Y. (2002): [To the spider fauna of Ka- zatskii district of the Central-Chernozyem Reserve]. In: ALEKHINA V.V.: Izuchene i okhrana prirody lesostepi. Materialy nauchno-prakticheskoi konferentsii, posv- yashchennoi 120-letiyu so dnya rozhdeniya Kurskaya Province, Zapovednoe, January 7, 2002. Tula. pp. 111- 112 [in Russian] PONOMAREV A.V. (1981): [To the fauna and ecology of spider family Gnaphosidae in the semi-desert zone of European part of the USSR]. In: Fauna i ekologiya nasekomykh. Perm University, Perm. pp. 54-68 [in Russian] PONOMAREV A.V. (1988): [Characteristics of the spider fauna of the semi-desert zone of the USSR European part.] In: Fauna i ekologiya paukoobraznykh. Mezh- vuzovskii sbornik nauchnykh trudov. Perm University, Perm. pp. 51-61 [in Russian] PONOMAREV A.V. (2005): [Spiders (Aranei) of Ros- tovskaya Province: fauna, landscape-zonal distribution. PhD Theses, Stavropol State University, Stavropol. 22 pp. [in Russian] PONOMAREV A.V. (2008): Additions to the fauna of spiders (Aranei) of the south of Russia and Western Kazakhstan: new taxa and finds. - Caucasian Entomological Bulletin 4: 49-61 Ponomarev A.V. &A.S. Tsvetkov (2004a): [General- ized data on spiders (Aranei) of the “Rostovskii” Reserve] - Trudy Gosudarstvennogo prirodnogo zapovednika “Rostovskii”. Donskoi izdatel’skii dom, Rostov-on-Don 4: 84-104 [in Russian] Ponomarev A.V. &A.S. Tsvetkov (2004b): [Spiders]. In: SHOLOKHOVA M.A.: Flora, fauna i mikobita Go- sudarstvennogo muzeya-zapovednika. Gosudarstvenyi muzei-zapovednik. Rostov-on-Don. pp. 81-87 [in Russian] Ponomarev A.V. & Y.A. Tsvetkova (2003): [Spiders (Aranei) of the territory of the Razdorskii Museum and Nature Reserve.] - Istoriko-kul’turnye i prirodnye issle- dovaniya na territorii Razdorskogo mezeya-zapovednika. Izdatel’stvo Rostovskogo universiteta, Rostov-on-Don 1: 167-208 [in Russian] RODE A. A. (1959): [Climatic conditions of the Dzh- anybek Station region]. - Soobshcheniya Laboratorii lesovedeniya AN SSSR 1: 5-78 [in Russian] Rode A.A. & M.N. Polskikh (1961): [Soils of the Dzhanybek Station, their morphological structure, mechanical and chemical composition, and physical properties.] In: Pochvy polupustyni severo-zapadnogo Prikaspiya i ikh melioratsiya. AN SSSR Pubis., Moscow, pp. 3-294 [in Russian] SAPANOV M.K. (2006): [Conditions for protective af- forestation growing in the semi-desert of the northern Caspian Region in view of climate changes in the second half of the XX century]. - Lesovedenie 6: 45-51 [in Russian] TUNEVA T.K. & S.L. ESYUNIN (2003): A review of the family Gnaphosidae in the fauna of the Urals (Aranei), 3. New species and new records, chiefly from the South Urals. - Arthropoda Selecta 11: 223-234 Arachnologische Mitteilungen 40: 105-109 Nuremberg, January 201 1 Regional variations in agrobiont composition and agrobiont life history of spiders (Araneae) within Hungary Ferenc Samu, Csaba Szinetär, Eva Szita, Kinga Fetykö & Dora Neidert doi:10.5431/aramit4012 Abstract: Agrobiont spider species are well adapted to arable systems, which have fairly uniform vegetation structure and pest assemblages over continent-wide areas. We wanted to study, whether agrobiont spider sub- assemblages and the life history of the most prominent agrobiont, Pardosa agrestis, show any regional variation within Hungary, where only modest climatic differences exist between the NW and SE parts of the country. We studied agrobiont species of spider assemblages in 27 alfalfa and 21 cereal fields with suction sampling and pitfalls.The similarity structure of these agrobiont sub-assemblages (Sorensen distance measure) was congruent with the geographic distance matrices (Eucledian distance), as tested by Mantel tests. However, if we considered sub-assemblages consisting of the non-agrobiont species, this congruency was always higher. Thus, agrobionts responded only moderately to geographical variation if we compare them to non-agrobiont species. We studied the generation numbers and the occurrence of the first adult individuals in P. agrestis; the most common agrobiont spider in Hungary.This comparison involved comparing fields along a NW - SE gradient during 6 sampling years in pairwise comparisons, where in each year a northern and a southern population was compared with a minimum distance of 1 26 km in between. In generation numbers there was no difference; we found two generations across Hungary. In contrast, the first occurrence of adult individuals was on average 1 5 days earlier in both generations in the more southern populations. Thus, it can be concluded that agrobionts show a fairly stable and relatively low magnitude response over country-sized geographical ranges. Key words: alfalfa, biological control, cereal, climatic gradient, natural enemy, Pardosa agrestis, regionality Agrobiont spider species are well adapted to ara- ble systems (SAMU 5c SZINETÄR 2002, KAJAK 5c OLESZCZUK 2004). Agricultural crops have a fairly uniform vegetation structure and pest assemblages over continent-wide areas (WECHSUNG et al. 1995). It is interesting to see how this large scale homogeni- sation of habitat structure affects spiders. On the one hand, the same crop might attract different spiders from the local fauna, whose species might contribute differently to the biological control potential of the natural enemy complex (BATÄRY et al. 2008, SAMU et al. 2008). On the other hand, uniform and low diversity spider assemblages of large crop areas might negatively affect nearby local natural assemblages (JEANNERET et al. 2003). Thus, information about the regional variation of crop spider assemblages is important both from biological control and nature conservation perspectives. Previous data indicates that spider assemblages even on the same crop plants vary Ferenc SAMU, Eva SZITA, Kinga FETYKÖ, Dora NEIDERT, Plant Protection Institute, Hungarian Academy of Sciences, P.O. Box 1 02 Budapest, H-1 525, Hungary, E-Mail: samu@julia-nki.hu Csaba SZINETÄR, University of West Hungary, P.O. Box 1 70, Szombathely, H-9701 , Hungary, E-Mail: szcsaba@ttmk.nyme.hu submitted: 1 7.3.201 0, accepted: 20.4.201 0; online early: 10.1.201 1 within Europe (HÄNGGI et al. 1995, FINCH et al. 2008). However, the extent and scale of this variation is largely unknown. Pardosa agrestis is the most common (c. 40 % dominance in arable fields) agrobiont spider in Hungary (SAMU 5c SZINETÄR 2002). However, this species is not an agrobiont in England and Western Continental Europe. It gains an agrobiont status along a NW-SE gradient in Central Europe (BLICK et al. 2000). The species has a second generation in Hungary (SAMU et al. 1998), while more northern populations are known to have a single generation (S. Toft pers. comm.). Although geographical details in the generation number shift have not yet been studied in detail, the fact that one generation populations are non- agrobionts or less dominant agrobionts (OBERG et al. 2008) suggests a relationship between generation number and the agrobiont status of the species. Since dominance increases from NW towards SE, and the existence of a second generation is known only from a starting point in Austria (ZULKA et al. 1997), we can infer the generation number to be an adaptation to climatic conditions. Since in Middle-Hungary the second generation was only facultative, i.e. part of the population had a one generation life-cycle, while the rest had two generations (KISS 5c SAMU 2005), 106 we suspected Hungary to be in a transition zone. Therefore we wanted to know if populations in NW Hungary and SE Hungary have different generation numbers. We decided to conduct observations over a modest geographical gradient within Hungary. We wanted to study responses to climatic differences existing between the NW and SE parts of the country. The objectives were (a) agrobiont sub-assemblages (only the agrobiont species out of all spider species) - where we studied the species composition - and (b) the life history (generation number and timing of genera- tions) of the most prominent agrobiont, Pardosa agrestis (Westring, 1861). Material and Methods Regional differences in agrobiont sub-assemblages We studied the agrobiont sub-assemblages in 27 al- falfa and 21 cereal fields located in various regions of Hungary. Datasets with comparable sampling efforts F Samu, C. Szinetär, E. Szito, K. Fetykö & D. Neidert and at least one cropping season sampling were selec- ted from the Hungarian spider database (SAMU 2000). The sampling method was either suction sampling or pitfall trapping, or both in parallel (Tab. 1). Standard parametric and multivariate community analyses (McCUNE & GRACE 2002) were run separately both by method and by the type of crop using PC-ORD, version 5.1. Following SAMU & SZINETÄR (2002) we regarded a species as an agrobiont if its average dominance was higher than 1 % in arable fields and it occurred on at least 75 % of the fields. Thus the following species were arable agrobionts: Pardosa agrestis (Westring, 1861), Meioneta rurestris (C. L. Koch, 1836), Oedo- thorax apicatus (Blackwall, 1850), Pachygnatha degeeri Sundevall, 1830, Erigone dentipalpis (Wider, 1834), Xysticus kochi Thorell, 1872, and Tibellus oblongus (Walckenaer, 1802). The seven agrobiont species represented on average 75 % of the spider individuals in the studied fields (mean ± S.D. = 75.5 ± 16.05, Table 1 : Locations of fields considered in the study. Fields that belonged to the same settlement considered to have the same location. Fields by and large represent a NW-SW gradient, and pairings were made to maximise distance in this direction (see text for details). # of fields sampled # of individuals (species) caught alfalfa cereal P. agrestis study Settlement Latitude (N) Longitude (E) pitfall D-vac pitfall D-vac Bank 47° 53' 39" 19° 08' 50" 1 533 (18) Decs 46° 16' 41" 18° 46' 43" 2 155 (13) 277 (12) 317(10) 203 (16) 80 Diösjeno 47° 55' 30" 19° 03' 01" 1 1961 (41) Felsönäna 46° 27' 46" 18° 32' 44" 6 4145 (61) 6348 (35) 576 (24) 660 (20) 2826 Fülöpszälläs 46° 48' 39" 19° 11' 07" 3 210 (19) 2504 (42) 200 (17) 561 (26) Gencsapäti 47° 16' 54" 16° 35' 33" 1 586 Hatvan 47° 41' 17" 19° 36' 36" 1 3713 (96) Iclod 40° 59’ 21” 23° 48’ 52” 1 1741 (40) Kartal 47° 39' 20" 19° 32' 06" 2 2700 (77) Kirälyhegyes 46° 17' 03" 20° 41' 12" 3 763 (40) 244 (15) 161 (18) 111 (8) 538 Kunpeszer 47° 03' 33" 19° 17' 32" 4 55 (8) 81 (11) 118 (18) Kunszentmiklös 47° 00' 53" 19° 09' 17" 2 217 (10) 484 (16) Livada 47° 00’ 18” 23° 50’ 24” 1 1489 (33) Nagykoväcsi 47° 32’ 52" 18° 56’ 03" 4 2867 (63) 8888 (76) 1176 (40) 1815 (39) 5556 Päsztö 47° 55' 12" 19° 44 07" 1 146 (9) Päty 47° 32’ 11" 18° 50' 49" 4 1259 (26) 313 (19) Retsäg 47° 54 34" 19° 05' 54" 1 333 (24) Romhany 47° 54' 36" 19° 12' 57" 1 1793 (41) Szekszärd 46° 19' 27" 18° 49' 23" 2 132 (10) 301 (16) 184 (13) Szombathely 47° 14' 58" 16° 38' 26" 2 4179 (37) Tevel 46° 22' 45" 18° 30' 28" 5 392 (27) 1474 (24) 152 (16) 96 (10) 319 Variations in agrobiont spiders in Hungary 107 N = 48 fields), while the remaining 25 % of the spi- ders belonged to 246 non-agrobiont species. The rare end of any community is always the most difficult to sample, and most vulnerable to undersampling bias. Therefore, when statistically considering the non-agrobiont species of spider assemblages (= non- agrobiont sub-assemblage), we only included those species into these analyses for which the whole data set had a cumulated individual count >50. Applying the inclusion rule, for the possible crop/method combinations we achieved the following number of non-agrobiont species: 29 spp. in alfalfa/pitfall; 28 spp. in alfalfa/D-vac; 32 spp. in cereal/pitfall and 25 spp. in cereal/D-vac. We tested whether spider assemblages changed with geographical location by performing Mantel tests, with Monte Carlo simulation to derive signi- ficance (MCCUNE &c MEFFORD 2010). This tested for congruence between the geographical distance (Eucledian distance) of the study plots and the dis- tance structure of the spider assemblages (Sorensen distance measure). Agrobionts and non-agrobionts were tested separately in the study plots. We analysed data sets gained by the two sampling methods and in the two crops separately, and also treated the agrobiont and the non-agrobiont sub- assemblages separately. Regional differences in Pardosa agrestis life history We studied the generation numbers and the occur- rence of first adult individuals in the common agro- biont wolf spider Pardosa agrestis. This comparison involved comparing fields along a NW — SE gradient during 6 sampling years (in the period 1993-2003) in pairwise comparisons, where in each year one north- ern and one southern population was compared, with a minimum distance of 126 km in between (see Tab. 1 for the sampling locations). Population samples were collected by pitfalls, emptied at weekly intervals. Results a) Regional differences in agrobiont sub-assem- blages The similarity structure of the analysed spider sub-as- semblages was congruent with the geographic distance matrices, except for the alfalfa agrobiont sub-assem- blage sampled by pitfalls. Relative to each other, the similarity between geographically close non-agrobiont sub-assemblages was always higher than it was bet- ween agrobiont sub-assemblages (Fig. 1). Although average agrobiont similarity among fields was a moderate 37 % (Sorensen similarity, abundance data) it was nearly twice as high as the 20 % similarity among fields if non-agrobiont species were considered (a significant difference: paired t-test comparing average similarity - based on all possible field pairings - of the agrobiont and non-agrobiont sub-assemblages for each crop and sample method combination: t = 7.37, d.f. = 3, P = 0.005). Figure 1: Standardised Mantel statistics of relationships between distances in the physical and in the community space separately analysed by (i) sub-assemblage type (agrobionts or non-agrobionts), (ii) crop type and (iii) col- lecting method. Asterisks denote the significance of the Mantel statistics at the P=0.05 level, obtained by Monte Carlo simulation. 240 220 200 180 160 140 120 100 80 Figure 2: First occurrence of adult P. agrestis individuals in days counted from 1 st January. Each year a new pair of northern and a southern populations (N = 6 pairs) were compared with a minimum distance of 1 26 km in between. 108 b) Regional differences in Pardosa agrestis life history Comparing the northern and southern populations we found two generations in each year. Thus the basic pattern of P. agrestis life-cycle seems to be uniform across the country The first adult occurrence of P. agrestis was significantly earlier in the more southern populations, on average by 16 days in the first, and by 14 days in the second generation (Fig. 2). The multi-factor AN OVA revealed that the effect of years was marginal, while interestingly, males did not occur significantly earlier than females in the samples (Tab. 2). Table 2: ANOVA analysis result between first occurrences of adult P. agrestis individuals.The six studied years were taken as random factors. (* indicated by asterisk). Source SS d.f. F P Year* 1273.77 5 1.7706 0.1503 North-South 2466.78 1 17.1442 0.0003 Generation 60704.8 1 421.9015 0.0001 Sex 0.44 1 0.003 0.9565 Discussion Agricultural habitats have very specific traits (such as special repeated pattern of disturbance (e.g. ploughing, harvest), monotypic, uniform vegetation structure, dominance of open soil surface, etc.), and the majority of the individuals - those belonging to agrobiont species - are adapted to these conditions (WlSSINGER 1997). Since crops have the same traits over large geographic areas, we expected the agrobi- onts to show little variation. Indeed, in the present study we found that similarity between agrobiont sub-assemblages was less determined by regionality (geographical proximity) than was the case for the non-agrobiont sub-assemblages. Between-field similarity of agrobionts was much higher than that of non-agrobonts. We interpret these results to mean that while both agrobiont species composition and non-agrobiont species composition change geographically, the agrobiont composition is much more conservative and stable in this respect. Moulting to the adult stage occurred ca. two weeks earlier in the southern populations of P. agrestis in the present study. Although the facultative nature of the two generations could not be studied by simple pitfall trapping, it can be speculated that more southern populations had more time to successfully complete a second generation and become juveniles that are large- enough for successful overwintering (KISS Sc SAMU F. Samu, C. Szinetdr, E. Szita , K. Fetykö & D. Neidert 2002). Based on this finding, it can be hypothesised that P agrestis populations in the south of Hungary are likely to have two obligate generations per year. To summarise, the effect of regionality on agrobi- ont assemblages seems to be scalable down to relatively small geographical areas, such as Hungary. However, at smaller scales the effects become more subtle. Both in the structure of agrobiont spider assemblages and in the life history of a prominent agrobiont species, small but consistent regional differences could be observed within the country. Agrobiont assemblages were less affected than non-agrobiont species, which indicates that agroecosystems host fairly stable animal assemblages, and in those assemblages the agrobiont component is mostly affected by large-scale factors, such as climate. As opposed to agrobionts, some 25 % of the assemblages in the studied fields consisted of a high number of non- agrobiont species, which showed much stronger regional variability; presumably they were more influenced by neighbouring habitats. We believe that both the agrobiont and non-agrobiont com- ponents are important in the relationship between agricultural and natural areas, because natural areas can give shelter and increase the number of both types of species, while large and uniform arable land will homogenise and reduce regional species pools, which may negatively influence nearby natural habitats. Acknowledgements We are grateful for funding by the grant: OTKA T048434. References Batäry R, A. BAldi, F. Samu, T. Szüts Sc S. Erdos (2008): Are spiders reacting to local or landscape scale effects in Hungarian pastures? - Biological Conservation 141: 2062-2070 - doi: 10.1016/j.biocon.2008.06.002 Blick T, L. Pfiffner Sc H. Luka (2000): Epigäische Spinnen auf Ackern der Nordwest-Schweiz im mittel- europäischen Vergleich (Arachnida: Araneae). - Mit- teilungen der Deutschen Gesellschaft für allgemeine angewandte Entomologie 12: 267-276 Finch 0.-D.,T. BLICK& A. SCHULDT (2008): Macroeco- logical patterns of spider species richness across Europe. - Biodiversity and Conservation 17: 2849-2868 - doi: 10.1007/sl0531-008-9400-x HÄNGGI A., E. Stöckli &W. NentWIG (1995): Habitats of Central European spiders. — Miscellanea Faunistica Helvetiae 4: 1-460 JEANNERET R, B. SCHUPBACH &H. LUKA (2003): Quan- tifying the impact of landscape and habitat features on Variations in agrobiont spiders in Hungary 109 biodiversity in cultivated landscapes. - Agriculture, Ecosystems 6c Environment 98: 311-320 - doi: 10.1016/ S0167-8809(03)00091-4 Kajak A. 6c M. OLESZCZUK (2004): Effect of shelter- belts on adjoining cultivated fields: patrolling intensity of carabid beetles (Carabidae) and spiders (Araneae). - Polish Journal of Ecology 52: 155-172 KISS B. 6c F. SAMU (2002): Comparison of autumn and winter development of two wolf spider spe- cies ( Pardosa , Lycosidae, Araneae) having different life history patterns. - Journal of Arachnology 30: 409-415 - doi: 10.1636/0161-8202(2002)030[0409: COAAWD]2.0.CO;2 KISS B. & F. SAMU (2005): Life history adaptation to changeable agricultural habitats: Developmental plas- ticity leads to cohort splitting in an agrobiont wolf spider. - Environmental Entomology 34: 619-626 - doi: 10.1603/0046-225X-34.3.619 McCUNE B. 6cJ.B. GRACE (2002): Analysis of ecological communities. MjM Software Design, Gleneden Beach. 300 pp. McCUNE B. 6cM.J. MEFFORD (2010): PC-ORD. Mul- tivariate analysis of ecological data. Version 5.1. MjM Software Design, Glenden Beach. OBERG S., S. Mayr 6c J. Dauber (2008): Landscape effects on recolonisation patterns of spiders in arable fields. - Agriculture, Ecosystems 6c Environment 123: 211-218 - doi: 10.1016/j.agee.2007.06.005 SAMU F. (2000): A general data model for databases in ex- perimental animal ecology. - Acta Zoologica Academiae Scientiarum Hungaricae 45: 273-292 Samu F, J. Nemeth, F. Töth, E. Szita, B. Kiss 6c C. SZINETÄR (1998): Are two cohorts responsible for bimodal life history pattern in the wolf spider Pardosa agrestis in Hungary? In: SELDEN PA. (Ed.): Proceed- ings of the 17th European Colloquium of Arachnology. Edinburgh, 14-18 July, 1997. British Arachnological Society, Burnham Beeches, Bucks, pp. 215-221 SAMU F. 6c C. SZINETÄR (2002): On the nature of agrobiont spiders. - Journal of Arachnology 30: 389-402 - doi: 10.1636/0161-8202(2002)030[0389: OTNOAS]2.0.CO;2 Samu F, A. Horväth, E. Szita, B. Bernäth, E. Botos 6c K. FETYKÖ (2008): The effect of source habitats on arable spider communities: is proximity the most im- portant? - IOBC/wprs Bulletin 34: 89-92 Wechsung G., F. Wechsung F, G.W. Wall, F.J. Adamsen, B.A. Kimball, R.L. Garcia, P.J. Pinter 6c T. KARTSCHALL (1995): Biomass and growth rate of a spring wheat root system grown in free-air C02 enrichment (FACE) and ample soil moisture. - Journal of Biogeography 22: 623-634 WlSSINGER S. (1997): Cyclic colonization in predictably ephemeral habitats: a template for biological control in annual crop systems. - Biological Control 10: 4-15 - doi: 10. 1006/bcon. 1997.0543 ZULKA K.P, N. MlLASOWSZKY 6cC. LETHMAYER (1997): Spider biodiversity potential of an ungrazed and grazed inland salt meadow in the National Park ‘Neusiedler See-Seewinkef (Austria): implications for management (Arachnida: Araneae). - Biodiversity and Conservation 6: 75-88 - doi: 10.1023/AT018375615960 Arachnologische Instructions to Authors The journal "Arachnologische Mitteilungen" publishes scientific papers about W-Palaearctic arachnids (excluding mites and ticks) in German or English. 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However, in such cases authors should consult the editors with respect to file formats, etc. Separata will be made available to the authors in a digital form (PDF-format, preferably sent by e-mail). ISSN 1018-4171 www.AraGes.de Arachnologische SMITHSONIAN LIBRARIES mi 11 il inn CO 90 B8 01 811 9 073 Maria Chatzaki: 25 th EC A Alexandroupolis, Greece: Preface 1-4 Theo Blick: Abundant and rare spiders on tree trunks in German forests (Arachnida: Araneae). . . 5-14 Robert Bosmans: On some new or rare spiders from Lesbos, Greece (Araneae: Agelenidae, Amaurobiidae, Corinnidae, Gnaphosidae, Liocranidae) 15-22 Christo Deltshev: The faunistic diversity of cave-dwelling spiders (Arachnida: Araneae) of Greece 23-32 Christo Deltshev, Stoyan Lazarov, Maria Naumova &c Pavel Stoev: A survey of spiders (Araneae) inhabiting the euedaphic soil stratum and the superficial underground compartment in Bulgaria 33-46 Jason A. Dunlop &, Plamen G. Mitov: The first fossil cyphophthalmid harvestman from Baltic amber 47-54 Peter J. van Helsdingen: Spiders in a hostile world (Arachnoidea: Arachnida) 55-64 Christian Komposch: Endemic harvestmen and spiders of Austria (Arachnida: Opiliones, Araneae) 65-79 Seppo Koponen: Ground-living spiders (Araneae) at polluted sites in the Subarctic 80-84 Jael Lubin, Noa Angel & Nirit Assaf: Ground spider communities in experimentally disturbed Mediterranean woodland habitats 85-93 Tatyana V. Piterkina: Spatial and temporal structure of the spider community in the clay semi-desert of western Kazakhstan 94-104 Ferenc Samu, Csaba Szinetär, Eva Szita, Kinga Fetykö & Dora Neidert: Regional variations in agrobiont composition and agrobiont life history of spiders (Araneae) within Hungary 105-109 ISSN 1018-4171 www.AraGes.de