sign in sign up
<<
Home , Amaurobius similis, Anyphaena accentuata, Araniella opisthographa, Bianor, Bolyphantes, Caviphantes

Arachnology (2018) 17 (8), 375–391 375

The (Arachnida: Araneae) of the orientalis Lipsky) and hornbeam (Carpinus caucasica Lagodekhi Reserve, : faunistic results of a Großh.); subalpine forests at 1800–2250 m a.s.l. are transect study and an updated checklist dominated by beech, (Quercus macranthera Fisch. & C.A.Mey. ex Hohen.), maple (Acer trautvetteri Medw.), birch (Betula litwinowii) and rhododendron (Rhododendron Stefan Otto caucasicum Pall.), interspersed with subalpine grasslands GutsMuthsstr. 42, 04177 Leipzig, and meadows (Kvachakidze 1999). According to this email: [email protected] author, the alpine habitats at 2500–3000 m a.s.l. are typi- cally developed as grasslands, rhododendron meadows, George Japoshvili and herb carpets. Subnival vegetation above 3000 m a.s.l. Institute of Entomology, consists of small patches of grass and herbs. Agraricultural University of Georgia, The Lagodekhi Reserve is the best studied single area in Agmashenebeli Alley 13 km, 0159 , Georgia Georgia for arachnology. Before our study, occurrence data and for 169 had been published in 33 studies between Invertebrate Research Center, 1895 and 2015 (see Appendix), indicating this protected Tetrtsklebi, municipality 2200, Georgia area’s signifcance as a refuge for a number of rarely recorded, endemic and specialist species. The occurrence of two species of the rather primitive mygalomomorph family Abstract Nemesiidae, Raveniola zaitzevi (Charitonov, 1948) and R. recki (Mcheidze, 1983), in Lagodekhi Reserve also high- In the largest systematic collection effort in a single lights its importance for the protection of numerous species, location in the Ecoregion, across several altitudinal including Tertiary relicts. zones and throughout the entire vegetation period of one year, In an extensive transect study in the Lagodekhi Reserve 426 juvenile and 359 adult spiders belonging to 69 species from 48 genera and 19 families were collected in the Lagodekhi undertaken in 2014, using Malaise traps and aiming at Reserve, Georgia, in 2014, using Malaise traps on a transect assessing the diversity of insects (e.g. Aslan., Mumladze with seven study plots between 666 and 2559 m a.s.l. in the & Japoshvili 2017; Japoshvili et al. 2017), a signifcant montane, subalpine, and alpine vegetation zones. pilula (Karsch, 1879) is recorded in the Western Palaearctic for number of spiders were collected as bycatch, providing the the frst time. Thirty-eight species are recorded in the Lagodekhi basis for new insights into the spider fauna of the Lagodekhi Reserve for the frst time, raising the number of spider species Reserve. The aim of this paper is to present the arachno- of this reserve to 202, fve additions are frst records for the faunistic results of this study, to note frst or rare species Caucasus Ecoregion and ten species are frst records for Georgia. Diagnostic drawings and analyses are provided for 13 records in the Caucasus Ecoregion, Georgia, or the Lago- species, including Megalepthyphantes pseudocollinus Saaristo, dekhi Reserve, to provide improved information and draw- 1997, Phoroncidia paradoxa (Lucas, 1846), P. pilula (Karsch, ings for poorly known or otherwise interesting species and 1879), and cachetiensis Mcheidze, 1997. An updated to compile a new checklist of the spiders of the Lagodekhi checklist of the spiders in the Lagodekhi Reserve is provided. Reserve.

Keywords: biodiversity • Caucasus • Georgia • new record • redescription Material and methods

This study is based on the material collected in the Lago- Introduction dekhi Reserve, using Malaise traps along an altitudinal transect on the Kudigora Ridge (Fig. 1) during the entire The Lagodekhi Reserve was established in 1912 as the vegetation season of 2014. Malaise traps (obtained from frst protected area in the Caucasus Ecoregion (henceforth BandN Entomological services http://www.entomology. Caucasus, sensu Williams et al. 2006) and nowadays holds org.uk) were set up on bushes at each altitude (one trap at the status of a Strict Nature Reserve. Spanning altitudes of each site) to collect spiders and other invertebrates. Plant approximately 500 to 3500 m a.s.l., the Lagodekhi Reserve associations are named based on Kvachakidze (2009). Trap exhibits a high diversity of natural landscapes and is one sites were: H1: 41.85248°N 46.28777°E, 666 m a.sl., mixed of the world’s best preserved pristine areas. It is located in lower montane forest with dominant beech and hornbeam the northeastern part of Georgia, close to the borders with (Carpineto-Fagetum festucosum), samples N1–N18; H2: and Dagestan (), on the southern slopes 41.85585°N 46.29273°E, 847 m a.s.l., mixed intermediate of the chain. The Lagodekhi Reserve montane forest with dominant beech (Fagetum nudum), consists of the Lagodekhi Nature Reserve (19,749 ha) and samples N19–N36; H3: 41.87147°N 46.31153°E, 1351 the Lagodekhi Managed Reserve (4702 ha) (APA 2016). m a.s.l., mixed intermediate montane forest with domi- The mountain forests, meadows, rivers, and alpine ecosys- nant beech (Fagetum galiosum), samples N37–N54; H4: tems of the Lagodekhi Reserve are very rich in habitat types 41.88273°N 46.32185°E, 1841 m a.s.l., broad-leaved and harbour numerous plant and species, a number upper montane forest with dominant beech and maple with of them endemic to the Caucasus, Georgia, or the reserve rubus underbrush (Fageto-Aceretum rubosum), samples itself (Kikodze et al. 2007). N55–N73; H4/5: 41.88558°N 46.32412°E, 1902 m a.s.l., At 400–1800 m a.s.l., forests near and within the Lago- broad-leaved upper montane forest with dominant maple dekhi Reserve mostly consist of montane beech (Fagus (Aceretum rubosum), samples N74–N83; H5: 41.89805°N 376 Spiders of Lagodekhi Reserve, Georgia

Mikhailov (2013), Otto (2015), Nentwig et al. (2017), and the WSC17. All observations and measurements were made using a Zeiss Stemi SV6 with an Axiocam 105 colour camera and ZEN 2 software. All drawings were made by the frst author based on microscope photographs and using a Wacom Intuos Pro digital drawing tablet with the programs Krita (version 2.9.7) and Gimp (version 2.8.16). Drawings always show the left male palpus; perspective and scale bars are given in the plates and their captions. All measurements are in mm. All altitudes are given in metres above sea level. Abbreviations: Caucasus = Caucasus Ecoregion; TmI range = ratio between a and b, with a being the distance between the proximal end of metatarsus I and the insertion point of the dorsal trichobothrium on this leg segment and b being the length of metatarsus I; WSC17 = World Spider Catalog (2017).

Results

Fig. 1: Map of Lagodekhi Reserve in Georgia showing trap sites H1–H6 on Kudigora Ridge: shaded area = forest cover, contour lines at 600 In total, 426 juvenile and 359 adult spiders (221 males, and 2000 m a.s.l., dashed line = state border of Dagestan and Azer- 138 females) were collected during the sampling period, baijan, dotted line = border of Lagodekhi Reserve. comprising 68 species from 47 genera (plus two females of an unidentifed Agyneta species, ) and 19 fami- 46.33388°E, 2230 m a.s.l., subalpine zone with (Juni- lies (see Appendix). peretum), samples N84–N95; H6: 41.90618°N 46.33340°E, The 38 species listed below are recorded in the Lagodekhi 2559 m a.s.l., alpine grassland (Alchemilletum), samples Reserve for the frst time, among them fve frst records in N96–N110. the Caucasus and ten frst records in Georgia. General collecting started on 2 April 2014 and lasted until 7 November 2014, although in alpine and subalpine areas collecting started later (subalpine 5 May 2014; alpine First records in the Caucasus 25 May 2014) and completed earlier (6 October 2014), due to weather conditions and altitude. Material was sp. () collected every 10 (± 2) days (depending on the weather) 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, and conserved in 96% ethanol and sorted. Identifcation 41.87147°N 46.31153°E, 1351 m a.s.l., mixed montane was done by the frst author using his reference collection forest, Fagetum galiosum, Malaise trap [H3/N37], 12–23 and literature sources on Caucasian spiders (see list in Otto April 2014, leg. Japoshvili & Kirkitadze, Coll. Otto (KVS 2015) as well as Nentwig et al. (2017) and sources listed 452. 1♂, same location, Malaise trap [H3/N38], 23 April–3 therein. May 2014, leg. Japoshvili & Kirkitadze, Coll. Otto (KBS Voucher specimens are deposited in the Entomolog- 258). 1♂, same location, Malaise trap [H3/N39], 5–15 May ical collection of the Agricultural University of Georgia 2014, leg. Japoshvili & Kirkitadze, Coll. Otto (KBS 259). (Tbilisi). and in the comparative collection of the frst The male palpal structure of the collected specimens is author. Synonymy follows the World Spider Catalog (2017) somewhat similar to the structures in A. antipovae Marusik (= WSC17). Distribution information is derived from & Kovblyuk, 2004 which was recently described from the

Phoroncidia pilula Phoroncidia paradoxa (Karsch, 1879) (Lucas, 1846) lacking median indentation with median indentation dorsal tubercle on abdomen (frontal view) (Fig. 13) (Fig. 14) with smooth edge lacking a pronounced step with perpendicular step inner edge of embolus base (Fig. 7b) (Fig. 8b) small apophysis between basal tip of embolus of high triangular shape and with a sharp tip fat and blunt base and tip of apical end of embolus proper (Fig. 7a) (Fig. 8a) 11:10 or 12:10 12:8 length : width ratio of embolus loop (Fig. 2) (Fig. 9) loop shape rather circular longish distal theridiid paracymbium; carapace length:width = 0.68–0.72:0.63–0.65; abdomen length = 1.20–1.28; body length = 1.57–1.67; femur unmarked or distal half darkened (Figs. 4, 10, 15); patella characters shared by males of both species unmarked or partly darkened; tibia with distal third/half darkened; metatarsus unmarked or distal margin darkened; tarsus unmarked

Table 1: Characters differentiating the males of Phoroncidia paradoxa and P. pilula. S. Otto & G. Japoshvili 377 2 4 6

5

3

11 10 12 7 a b

8 a b

9 15 17 16

13 14

Figs. 2–17: Phoroncidia pilula (male from Lagodekhi, KVS 427, 2–7,13). 2 palpus, ventral view; 3 palpus, medioventral view; 4 left leg IV, anterior view; 5 habitus, lateral view; 6 habitus, dorsal view; 7a apophysis near embolus base; 7b embolus base; 13 dorsal tubercle on abdomen, frontal view. Phoroncidia paradoxa (male from , NMB-ARAN-25761, 8–12, 14). 8a apophysis near embolus base; 8b embolus base; 9 palpus, ventral view; 10 left leg IV, anterior view; 11 habitus, lateral view; 12 habitus, dorsal view; 14 dorsal tubercle on abdomen, frontal view. Pho- roncidia paradoxa (female from /Trieste, 15–17). 15 left leg IV, anterior view; 16 habitus, lateral view; 17 habitus, dorsal view.

Caucasus. It is interesting that the spider collection at the The single recorded male has a body length of 1.48, Janashia Museum in Tbilisi contains specimens collected its carapace is dark clay coloured and tinged black with by D. Charitonov and labeled as: “Amaurobius caucasicus, a black line pattern. The abdomen is dark grey. Legs are 25 June, 38, Lagodekhi”, referring to a never formally clay coloured and tinged black with tinging being weakly described species, which is probably conspecifc with our developed or lacking at the proximal and distal ends of the specimens. Due to the lack of data at this point we decided distal leg segments. Habitus and palp are developed as in European specimens (Figs. 18–20). to make this question the topic of another paper (Marusik et This rarely found species has so far been recorded mostly al. in prep.). in Central , with its easternmost records in Cinetata gradata (Simon, 1881) (Linyphiidae) and (Blick & Goßner 2006; Blagoev, Deltshev & 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, Lazarov. 2015; van Helsdingen 2016). Our record of this 41.88558°N 46.32412°E, 1902 m a.s.l., montane forest, species extends its range to the Caucasus. broad-leaved upper montane forest, Aceretum rubosum, Phoroncidia pilula (Karsch, 1879) () Malaise trap [H4–5/N81], 27 September–6 October 2014, 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, leg. Japoshvili & Kirkitadze, Coll. Otto (KVS 464), Figs. 41.85248°N 46.28777°E, 666 m a.s.l., mixed montane 18–20. forest, Fageto-Carpinetum-Festucosum, Malaise trap [H1/ 378 Spiders of Lagodekhi Reserve, Georgia 18 19 20

22 21

23

25

24 26

28 27

Figs. 18–28: Cinetata gradata (male, 18–20). 18 habitus, lateral view; 19 habitus, dorsal view; 20 palpus, retrolateral view. Pocadicnemis juncea (male). 21 palpus, retrolateral view. sp. (females). 22 carapace habitus, dorsal view (KVS 321); 23 , ventral view (KVS 321); 24 vulva, dorsal view (KVS 321); 25 vulva, dorsal view (KBS 328); 26 vulva, dorsal view (KBS 327). Megalepthyphantes cf. pseudocollinus (male from Lagodekhi, KVS460). 27 palpus, retrolateral view. M. pseudocollinus (male from Russia/Voronezh). 28 palpus, retrolateral view (drawing from photo, courtesy of A. Tanasevitch, without scale).

N2], 12–23 April 2014, leg. Japoshvili & Kirkitadze. Coll. 2011, Naturhistorisches Museum Basel, catalogue no. Otto (KVS 427), Figs. 2–7, 13. NMB-ARAN-25761, det. A. Hänggi (see collection details Comparative material: Phoroncidia pilula (= Ulesanis in Hänggi et al. 2014), Figs. 8–12, 14. 1♂, , Rhône minschana Schenkel, 1936; = Phoroncidia borea Logunov Alps, Ardèche, Thueyts, 44.67639°N 4.22139°E, ~500 m & Marusik, 1992) 1♂, 1♀ (syntypes), Naturhistorisches a.s.l., 9–13 October 1986, leg. P. Poot, Coll. Senckenberg Museum Basel, catalogue no. 1467a, label: “Ulesanis Museum Frankfurt, catalogue no. SMF 59538. minschana Schkl, S Kansu, K13, coll. Hummel”. Phoroncidia minuta (Spassky, 1932) is the only species of the , which has been recorded from the Caucasus so Phoroncidia paradoxa (Lucas, 1846) far, and all records were based on females (Spassky 1937; 1♂ (dried specimen), Italy, Garda Mountains, Monte Mcheidze 1964; Logunov & Marusik 1992). We carefully Brione, 45.888056°N 10.873611°E, 250 m a.s.l., 27 examined and compared the single male collected from August 2005, leg. et det. et Coll. B. Knofach. 1♀, Italy, Lagodekhi with the European species P. paradoxa (Lucas, Trieste, Aurisina, 45.75°N 13.67°E, ~200 m a.s.l., 16 1846) and with P. pilula (Karsch, 1879), which was known December 1994, leg. Bertrandi, det. et Coll. B. Knofach, from the Far East only. Despite some similarities with P. Figs. 15–17. 1♂, Switzerland, Ticino, vill. Claro (Al paradoxa, our specimen shared more characters with the Razzei), 46.25117°N 9.03043°E, 412 m a.s.l., 6 September syntype male of Ulesanis minschana Schenkel, 1936, which S. Otto & G. Japoshvili 379 31 29 30

32 33

34 35

36 37 38

Figs. 29–38: Xysticus pseudolanio (H4/KVS 461, male, 29–30). 29 palpus, retrolateral view; 30 palpus, ventral view. tantulus (female, 31–33). 31 habitus, dorsal view (body colour overall black, not depicted here); 32 epigyne, ventral view; 33 vulva, dorsal view. Dictyna ottoi (H3/KBS 265, female, 34, 36). 34 habitus, dorsal view; 36 vulva, dorsal view. Dictyna ottoi (H2/KBS 241, male, 35, 37–38). 35 habitus, dorsal view; 37 palpus, ventral view; 38 palpus, retrolateral view. has since been synonymized with P. pilula (Karsch, 1879). Details of the female habitus of P. paradoxa are also This identifcation is further supported by a second opinion given in Figs. 15–17 to show intraspecifc variability and (Y. Marusik, pers. comm.). Since P. paradoxa has been because published drawings have so far not shown the reported from nearby and characters differentiating habitus in detail. the males of the two species have not yet been published, Phoroncidia pilula is distributed in the southeastern part it seems reasonable to compare these species in more of Russia, northeast , , and (Marusik, detail. Based on comparison of the material listed above, Mikhailov & Omelko 2015; WSC17). Our frst record and literature sources, we derived some, in our opinion, of this species in the Caucasus suggests either a much reliable characters, and present them in Table 1 and Figs. larger distribution range or a new case of Caucasian–Far 2–14. Thaler & Nofatscher (1990: 173) pointed out some East disjunctive range in spiders (Marusik, Mikhailov & palp characteristics typical for P. paradoxa which, in our Guseinov 2006). specimen, were either hardly or not visible, e.g. the guiding ridge of the radix (“Führungsleiste der Radix”), or they Pocadicnemis juncea Locket & Millidge, 1953 (Linyph- differed in the form and proportions heavily depending iidae) on viewing perspective, e.g. the distal median apophysis 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, (“Radixabschnitt mit kräftiger Endkralle”). Perspective was 41.88558°N 46.32412°E, 1902 m a.s.l., broad-leaved upper also important when viewing the embolus base and the more montane forest, Aceretum rubosum, Malaise trap [H4–5/ or less hook-shaped median apophysis (compare Figs. 2–3, N75], 25 June–5 July 2014, leg. Japoshvili & Kirkitadze, 7b with Figs. 8b–9). Coll. Otto (KVS 463), Fig. 21. 380 Spiders of Lagodekhi Reserve, Georgia

The single collected male shows the typical habitus and Coll. Otto. 2♂, same location, Malaise trap [H5/N88], 25 palpus structure (Fig. 21) including the rather long and June–5 July 2014, leg. Japoshvili & Kirkitadze, Coll. Otto. curved median bulbal apophysis of this species (Roberts This species has so far only been recorded from near the 1987; Nentwig et al. 2017). Pocadicnemis juncea is distrib- Caspian Sea in Dagestan (Ponomarev, Khalidov & Aliev uted throughout Europe, with its easternmost records in 2008; Ponomarev et al. 2011a,b; Abdurakhmanov & Alieva Bulgaria (Blagoev, Deltshev & Lazarov. 2015) and the 2011; Ponomarev & Abdurakhmanov 2014). Our records (Mikhailov 2013). Our record of this species are the frst of this species from Georgia. extends its range from Europe to the Caucasus. Dictyna ottoi Marusik & Koponen, 2017 (Dictynidae) Xysticus pseudolanio Wunderlich, 1995 () 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, 41.85248°N 46.28777°E, 666 m a.s.l., mixed montane forest, 41.88273°N 46.32185°E, 1841 m a.s.l., broad-leaved upper Carpineto-Fagetum festucosum, Malaise trap [H1/N6], montane forest, Aceretum rubosum, Malaise trap [H4/N57], 25 May–4 June 2014, leg. Japoshvili & Kirkitadze, Coll. 15–25 May 2014, leg. Japoshvili & Kirkitadze, Coll. Otto Otto. 2♂♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, (KVS 461), Figs. 29–30. 1♂, Georgia, Lagodekhi Reserve, 41.85585°N 46.29273°E, 847 m a.s.l., mixed montane Kudigora Ridge, 41.89805°N 46.33388°E, 2230 m a.s.l., forest, Fagetum nudum, Malaise trap [H2/N21], 5–15 May subalpine zone, Juniperetum, Malaise trap [H5/N86], 4–14 2014, leg. Japoshvili & Kirkitadze, Coll. Otto (KBS 241), June 2014, leg. Japoshvili & Kirkitadze, Coll. Otto. 1♂, Figs. 35, 37–38. 5♂♂, Georgia, Lagodekhi Reserve, Kudi- Georgia, Lagodekhi Reserve, Kudigora Ridge, 41.90618°N gora Ridge, 41.87147°N 46.31153°E, 1351 m a.s.l., mixed 46.33340°E, 2559 m a.s.l., alpine grassland, Alchemilletum, montane forest, Fagetum galiosum, Malaise trap [H3/N40], Malaise trap [H6/N102], 5–15 July 2014, leg. Japoshvili & 15–25 May 2014, leg. Japoshvili & Kirkitadze, Coll. Otto Kirkitadze, Coll. Otto. (3♂), Coll. Agr. Univ. Tbilisi (2♂). 1♂, same location, [H3/ The three collected males clearly differ from related N42], 4–14 June 2014, leg. Japoshvili & Kirkitadze, Coll. species like X. ukrainicus Utochkin, 1968 (cf. Utochkin Otto. 1♀, same location H3, trap N43, 15–25 June 2014, 1968; Mcheidze 1997, 2014) and closely resemble the leg. Japoshvili & Kirkitadze, Coll. Otto. 1♀, same location species description of X. pseudolanio (Figs. 29–30), with H3/N45, 5–15 July 2014, leg. Japoshvili & Kirkitadze, Coll. some minor differences concerning the cymbium, median apophysis and distal embolus (J. Wunderlich, pers. comm.). Otto (KBS 265), Figs. 34, 36. Xysticus pseudolanio has been described from Turkey and Comparative material: 1♂, Georgia, Khevsureti, only its male sex is known (Wunderlich 1995). The type Giorgitsminda, 42.563833°N 45.088472°E, 1700 m a.s.l., locations in northeast and northwest Turkey were situated in montane meadow, under rocks, 24 June 2009, leg. et Coll. forests between 1000 and 1100 m a.s.l., whereas our three Otto (KVS 135). specimens were collected in mountain forest, the subalpine This species is close to D. armata Thorell, 1875 and has zone and on alpine mats between ~1800 and 2600 m. Vege- only recently been described from specimens collected in tation types of lower altitudes are known to reach higher Azerbaijan (Marusik & Koponen 2017). Since the descrip- in the Eastern Caucasus and can even be found in isolated tion only included photographs of this species, we present spots well above their usual upper border, e.g. the so called drawings of the habitus and copulatory organs of both sexes subalpine mountain steppe of the Eastern Caucasus Major, (Figs. 34–38). e.g. in the Lagodekhi Reserve (Kvachakidze 1999; Nakhuts- Our records are the frst of this species in Georgia, but rishvili 2013). Therefore, despite signifcant differences in it is very probable that all earlier records of D. armata in altitudes, environmental conditions of X. pseudolanio in the Georgia actually refer to D. ottoi (cf. Marusik & Koponen Lagodekhi Reserve might be quite similar to the locations 2017). in Turkey. It should be kept in mind, too, that older records braccata (C. L. Koch, 1841) (Theridiidae) of similar looking species in the Caucasus, like records of 2♀♀, Georgia, Lagodekhi Reserve, Kudigora Ridge, X. lanio by T. Mcheidze, might actually belong to X. pseu- 41.85248°N 46.28777°E, 666 m a.s.l., mixed montane dolanio (Marusik, pers. comm.). forest, Carpineto-Fagetum festucosum, Malaise trap [H1/ N8], 15–25 June 2014, leg. Japoshvili & Kirkitadze, Coll. Otto. 2♀♀, Georgia, Lagodekhi Reserve, Kudigora Ridge, First records in Georgia 41.85585°N 46.29273°E, 847 m a.s.l., mixed montane forest, Fagetum nudum, Malaise trap [H2/N26], 25 June–5 Cheiracanthium montanum L. Koch, 1877 (Eutichuridae) July 2014, leg. Japoshvili & Kirkitadze, Coll. Otto. 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, This species is distributed in Europe and the Mediterra- 41.87147°N 46.31153°E, 1351 m a.s.l., mixed montane nean (WSC17). In the Caucasus it has so far been recorded forest, Fagetum galiosum, Malaise trap [H3/N38], 23 April–3 May 2014, leg. Japoshvili & Kirkitadze, Coll. Otto. twice, from Adygea and from North Ossetia-Alania 1♂, same location, Malaise trap [H3/N46], 15–25 July 2014, (Ponomarev & Mikhailov 2007; Ponomarev et al. 2012). leg. Japoshvili & Kirkitadze, Coll. Otto. 1♂, Georgia, Lago- Lathys humilis (Blackwall, 1855) (Dictynidae) dekhi Reserve, Kudigora Ridge, 41.89805°N 46.33388°E, 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, 2230 m a.s.l., subalpine zone, Juniperetum, Malaise trap 41.85248°N 46.28777°E, 666 m a.s.l., mixed montane [H5/N84], 25 May–4 June 2014, leg. Japoshvili & Kirkit- forest, Carpineto-Fagetum festucosum, Malaise trap [H1/ adze, Coll. Otto. 3♂♂, 2♀♀, same location, Malaise trap N2], 12–23 April 2014, leg. Japoshvili & Kirkitadze, Coll. [H5/N86], 4–14 June 2014, leg. Japoshvili & Kirkitadze, Otto. S. Otto & G. Japoshvili 381

This species has so far been recorded from a number of 41.85585°N 46.29273°E, 847 m a.s.l., mixed montane locations in the North Caucasus and Azerbaijan (e.g. Dunin forest, Fagetum nudum, Malaise trap [H2/N22], 15–25 May 1988; Guseinov 1999; Ponomarev & Volkova 2013; see 2014, leg. Japoshvili & Kirkitadze, Coll. Otto. further references in Otto 2015). It seems to be wide-spread This species has so far only been recorded from a single and abundant in the Caucasus. Our record is the frst of this location in Adygea in the North Caucasus (Ponomarev et al. species from Georgia. 2012). Our records are the second example of this species in Megalepthyphantes cf. pseudocollinus Saaristo, 1997 the Caucasus and the frst from Georgia. (Linyphiidae) Sibianor tantulus (Simon, 1868) (Salticidae) 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, 1♀, Georgia, Lagodekhi Reserve, Kudigora Ridge, 41.87147°N 46.31153°E, 1351 m a.s.l., mixed montane 41.90618°N 46.33340°E, 2559 m a.s.l., alpine grassland, forest, Fagetum galiosum, Malaise trap [H3/N53], 27 Alchemilletum, Malaise trap [H6/N102], 5–15 July 2014, September–6 October 2014, leg. Japoshvili & Kirkitadze, leg. Japoshvili & Kirkitadze, Coll. Otto (KVS 468), Figs. Coll. Otto (KVS 460), Fig. 27. 31–33. Comparative material: 1♂, Russia, Voronezh Oblast, The single collected female resembles S. tantulus very Savalskoe Lesnichestvo near Ternovka railway station, well, especially in the structure of the epigyne and vulva 51.67383°N 41.54173°E, 150 m a.s.l., forest, 1980, leg. K. (Figs. 32–33), e.g. the insemination ducts lacking the frst Mikhailov, Coll. A. Tanasevitch, Fig. 28. loop seen in related species (Logunov & Marusik 1991; Megalepthyphantes pseudocollinus has been recorded Logunov 2001). However, according to these studies, leg twice in the Caucasus Ecoregion, from Krasnodar Kray colouration of our specimen is more similar to S. auro- (Russia), near the Crimea (Ponomarev & Mikhailov 2007) cinctus, especially the dark femur and tibia I with contrasting and from North Ossetia-Alania (Ponomarev & Komarov brighter patella (but femur I antero-laterally bright, cf. Fig. 2013). 31). The single collected male shows close affnity to M. This species is distributed in the Palaearctic (Logunov pseudocollinus Saaristo, 1997, despite having a rather 2001; Mikhailov 2013; WSC17) and has been recorded broad lamella characteristica (Fig. 27) instead of the usual from the Caucasus in North Ossetia-Alania (Logunov 2001; slender one (cf. Saaristo 1997). Variation seems to be quite Ponomarev & Komarov 2013). Our record is the frst of this large within this species as shown in a specimen from Voro- species from Georgia. nesh, Russia (Fig. 28) and there are more yet undiscovered The records in North Ossetia-Alania are from the steppe, species to be expected in (and living close to) the Caucasus, steppe meadows, and young oak forests. The occurrence of as indicated by a number of single females collected, which S. tantulus at such a comparably high altitude in our study have so far not been attributed to any of the extant species in might be attributed to climatic conditions and elements the genus Megalepthyphantes (Coll. Otto; A. Tanasevitch, of steppe vegetation in the alpine zone of the Lagodekhi pers. comm.). We think it is possible that our collected male Reserve (see comments for Xysticus pseudolanio above). actually belongs to one of these yet undescribed species, but teberdaensis Tanasevitch, 2010 (Linyph- this question can only be answered based on newly collected iidae) pairs of males and females as well as a thorough study of the 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, (few!) specimens of M. pseudocollinus already existing in 41.88273°N 46.32185°E, 1841 m a.s.l., upper montane private and institutional collections. forest, Fageto-Aceretum rubosum, Malaise trap [H4/N72], Nigma favescens (Walckenaer, 1830) (Dictynidae) 27 September–6 October 2014, leg. Japoshvili & Kirkit- 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, adze, Coll. Otto. 41.85585°N 46.29273°E, 847 m a.s.l., mixed montane In the Caucasus, T. teberdaensis has so far been recorded forest, Fagetum nudum, Malaise trap [H2/N19], 12–23 April fve times from Adygea and Karachay-Cherkessia (Tana- 2014, leg. Japoshvili & Kirkitadze, Coll. Otto. 1♂, same sevitch 2010; Ponomarev et al. 2012; Martynovchenko & location, Malaise trap [H2/N20], 23 April–4 May 2014, leg. Mikhailov 2014; Ponomarev & Chumachenko 2014). Our Japoshvili & Kirkitadze, Coll. Otto. 4♂♂, same location, record is the frst of this species from Georgia. Malaise trap [H2/N21], 5–15 May 2014, leg. Japoshvili & Xysticus kulczynskii Wierzbicki, 1902 (Thomisidae) Kirkitadze, Coll. Otto. 2♂♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, This species has been recorded from nine locations in 41.85248°N 46.28777°E, 666 m a.s.l., mixed montane the North Caucasus, , and Azerbaijan (Kulczyński forest, Carpineto-Fagetum festucosum, Malaise trap [H1/ 1895; Dunin 1988, 1989; Guseinov 1999; Ponomarev & N4], 5–15 May 2014, leg. Japoshvili & Kirkitadze, Coll. Khatshikov 2009; Ponomarev & Volkova 2013). Our record Otto. 2♂♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, is the frst of this species from Georgia. 41.87147°N 46.31153°E, 1351 m a.s.l., mixed montane albidus Kulczyński, 1911 () forest, Fagetum galiosum, Malaise trap [H3/N39], 5–15 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, May 2014, leg. Japoshvili & Kirkitadze, Coll. Agr. Univ. 41.85248°N 46.28777°E, 666 m a.s.l., mixed montane Tbilisi. 2♀♀, same location, Malaise trap [H3/N41], 25 forest, Carpineto-Fagetum festucosum, Malaise trap [H1/ May–4 June 2014, leg. Japoshvili & Kirkitadze, Coll. Otto. N4], 5–15 May 2014, leg. Japoshvili & Kirkitadze, Coll. 1♀, same location, Malaise trap [H3/N42], 4–14 June 2014, Otto. 1♀, same location, Malaise trap [H1/N8], 15–25 leg. Japoshvili & Kirkitadze, Coll. Otto. June 2014, leg. Japoshvili & Kirkitadze, Coll. Otto (KVS Genital characters of both males and females clearly 437). 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, distinguish this species from similar taxa such as X. 382 Spiders of Lagodekhi Reserve, Georgia ferrugineus Menge, 1876 or X. spasskyi Utochkin, 1986. X. show close affnity to (Blackwall, kulczynskii has so far only been recorded from a few loca- 1833) when comparing habitus, body size and TmI range. tions in the Eastern parts of Azerbaijan (Logunov, Marusik However, the epigyne and vulva differ somewhat from the & Koponen 2002). Our record is the frst of this species characters in this species (Figs. 23–26) as well as from the from Georgia. much smaller Centromerita concinna (Thorell, 1875). We and a colleague (T. Blick, pers. comm.) therefore judge our specimens probably not to be conspecifc with these two Further additions to the checklist of Lagodekhi Reserve species. As no males have been found in this study and the likely C. bicolor has not yet been recorded in the Caucasus, The following 23 species have previously been recorded we decided to present drawings of the specimens collected from Georgia and are now recorded from the Lagodekhi and leave the question of our material’s species determina- Reserve for the frst time. Detailed collection data can be tion open until more material, including males, has been found in the Appendix. found in the Caucasus. This should be more likely during 1. opisthographa (Kulczyński, 1905) (Araneidae) winter months because other species in this genus, including 2. Blackwall, 1841 (Clubionidae) the two mentioned above, are winter active (T. Blick, pers. 3. (Walckenaer, 1802) (Clubionidae) comm.). 4. Cheiracanthium mildei L. Koch, 1864 (Eutichuridae) Clubiona caucasica Mikhailov & Otto, 2017 (Clubio- 5. Drassodes lapidosus (Walckenaer, 1802) (Gnaphosidae) nidae) 6. Scotophaeus scutulatus (L. Koch, 1866) (Gnaphosidae) Twenty-fve males and 20 females of a species vaguely 7. Pelecopsis crassipes Tanasevitch, 1987 (Linyphiidae) resembling C. caerulescens L. Koch, 1867 were found 8. Trichoncus affnis Kulczyński, 1894 (Linyphiidae) in the samples. Further analysis revealed it to be a new 9. (Walckenaer, 1802) (Philodro- species which was subsequently described in Mikhailov et midae) al. (2017). All records of C. caerulescens in Lagodekhi by 10. Philodromus dispar Walckenaer, 1826 (Philodromidae) Mikhailov (1990) thus refer to this species. 11. Pisaura novicia (L. Koch, 1878) (Pisauridae) 12. dubius C. L. Koch, 1835 (Salticidae) Trochosa cachetiensis Mcheidze, 1997 (Lycosidae) 13. Pseudicius encarpatus (Walckenaer, 1802) (Salticidae) 1♂, Georgia, Lagodekhi Reserve, Kudigora Ridge, 14. Segestria bavarica C. L. Koch, 1843 (Segestriidae) 41.87147°N 46.31153°E, 1351 m a.s.l., mixed montane 15. (Ohlert, 1865) (Salticidae) forest, Fagetum galiosum, Malaise trap [H3/N39], 5–15 May The single collected female exhibits the conspicious 2014, leg. Japoshvili & Kirkitadze, deposited at Naturhistor- ”frst loop” of the insemination ducts and colouration of isches Museum Basel, catalogue no. NMB 27262 (ex Coll. leg I, typical for S. aurocinctus (cf. Logunov 2001); but as Otto, KVS 454, Figs. 39, 40, 48, 54, 55). 1♀, same location, females of S. aurocinctus and S. turkestanicus Logunov, Malaise trap [H3/N40], 15–25 May 2014, leg. Japoshvili & 2001 are barely separable, males from the Lagodekhi reserve Kirkitadze, Naturhistorisches Museum Basel, catalogue no. are needed in order to verify (or correct) our determination. NMB 27263 (ex Coll. Otto, KVS 455, Figs. 41, 52, 53). 1♂, 16. Sittipub relictarius (Logunov, 1998) (Salticidae) same location, Malaise trap [H3/N42], 4–14 June 2014, leg. 17. Anelosimus vittatus (C. L. Koch, 1836) (Theridiidae) Japoshvili & Kirkitadze, Naturhistorisches Museum Basel, 18. Neottiura bimaculata (Linnaeus, 1767) (Theridiidae) catalogue no. NMB 27264 (ex Coll. Otto, KBS 262). 19. Theridion varians Hahn, 1833 (Theridiidae) Comparative material: Trochosa robusta: ♂, , Stei- 20. livens Simon, 1876 (Thomisidae) ermark, Zöbing, Pollhammer, 15°44′25″E 47°01′18″N, 20 21. Xysticus gallicus Simon, 1875 (Thomisidae) May–2 June 2008, leg. Frieß, Coll. C. Muster (Figs. 42, 49), 22. Trachelas minor O. Pickard-Cambridge, 1872 (Trachel- ♀, Austria, Burgenland, Seewinkel, Kleine Neubruchlacke, idae) 16°49′57″E 47°46′47″N, 20–30 April.1994, ex Coll. N. 23. Zora spinimana (Sundevall, 1833) (Zoridae) Milasowszky, KNL74C2 (Fig. 43); Trochosa ruricola: ♂,♀, Germany, Hannover Vahrenheide, Kugelfangtrift, 15 April 2005, leg. P. Sprick, Coll. Otto, Lyc16VS (Figs. 44–45, 50); Notes on other rarely collected or poorly known species: Trochosa hispanica: ♂,♀, Switzerland, Tessin, Bissone, 21 April–7 June 1984, leg. A. Pedroli, Naturhistorisches Centromerita sp. (Linyphiidae) Museum Basel, catalogue no. NMB 2373r (Figs. 46–47, 1♀, Georgia, Lagodekhi Reserve, Kudigora Ridge, 51). 41.90618°N 46.33340°E, 2559 m a.s.l., alpine grassland, Despite having been recorded in the Caucasus more than Alchemilletum, Malaise trap [H6/N107], 15–25 July I 2014, twenty times, this species remains poorly known because leg. Japoshvili & Kirkitadze, Coll. Otto (KBS 327), Fig. 26. the diagnosis and drawings of Trochosa cachetiensis in 1♀, same location, Malaise trap [H6/N108], 25 August–4 Mcheidze (1997, 2017) are not detailed, and the type mate- September 2014, leg. Japoshvili & Kirkitadze, Coll. Otto rial from the Janashia Museum is not available for study. In (KBS 328), Fig. 25. 1♀, Georgia, Tusheti region, near vill. order to facilitate further taxonomic studies on this species, Diklo, 42.39701°N 45.69386°E, 2100 m a.s.l., subalpine we provide a short description of the somatic and genital pasture, 4 September 2009, leg. F. Walther, Coll. Otto (KVS characters together with diagnostic drawings as well as a 321), Figs. 23–24. discussion of differential characters concerning related The two females from our study and a conspecifc female species. Terminology of epigyne and vulva structure follows from a collection in a neighboring Georgian mountain region Hepner & Milasowszky (2006). S. Otto & G. Japoshvili 383 39 40 41

42 43

44 45

46 47

48 50 49 51

52 54 55

53

Figs. 39–55: 39 Trochosa cachetiensis male habitus, dorsal view. 40–47 right leg, prolateral view, of Trochosa spp. 40 T. cachetiensis male; 41 T. cache- tiensis female; 42 T. robusta male; 43 T. robusta female; 44 T. ruricola male; 45 T. ruricola female; 46 T. hispanica male; 47 T. hispanica female. 48–51 bulb median apophysis, apical view, of Trochosa spp. 48 T. cachetiensis; 49 T. robusta; 50 T. ruricola; 51 T. hispanica. 52–55 T. cachetiensis. 52 epigyne, ventral view; 53 vulva, dorsal view; 54 left bulb, ventral view; 55 left bulb, retrolateral view.

Males (n = 2) Body length 8.60/8.20, carapace length paired brown spots. Fovea thin, black. Chelicera dark 4.30/4.55, width 3.20/3.42, yellow to clay coloured, with brown, fang smooth, brown-red, lacking a hump, inner and median band bordered by broad brown lateral bands; cara- outer row of cheliceral furrow each with 3 teeth; medium pace edge with irregular brown fringe. Brown fringe and tooth in each row largest, apical tooth smallest. Sternum brown lateral bands separated from each other by bright bright yellow (in one specimen fringed brown), labium and irregular thin band (Fig. 39). Clypeus thin. Eyes with black gnathocoxae brown. Legs light brown with pale femora fringe on brown background connected to the brown lateral and coxae. Annulation weak and better visible on femora. bands. Bright median band behind ocular region with faint Opistho soma dorsum with dark brown pattern on clay 384 Spiders of Lagodekhi Reserve, Georgia coloured background, heart region with lanceolate bright Discussion mark and pairwise arranged bright spots behind, each with small dark spot inside. Sides with dark brown irregular Despite not having been initially planned as an arach- markings. Venter bright, laterally bordered by a row of dark nological study per se and with a number of other studies brown spots. Cymbium and palpal tibia brown, darker than having contributed a larger total sum of specimens and patella and femur. Cymbium with apical claw. Embolus tip species, this initially entomological study with a focus hook-like (Figs. 48, 54, 55), median apophysis rather thick on hymenopterans and beetles turned out to be the most (Fig. 48). systematic, industrious and thorough arachnological feld Female (n = 1) Body length 12.61, carapace length 5.31, study ever undertaken in the Caucasus as well, comprising width 4.05. Habitus as in male, but fovea rather thick, short, seven trap sites at altitudes between 666 and 2559 m a.s.l. black. Sternum bright yellow with broad brown fringe. and a collection period between April and November with Pedipalpus tip with claw. Epigyne with rounded helmet- a fortnightly sampling scheme. Not surprisingly, the results shaped broadenings and copulatory ducts visible through refect this systematic and thorough collection effort: 69 cuticula (Fig. 52). Vulva with large and bulging copulatory collected species, 38 species new to the Lagodekhi Reserve, ducts lacking small apical appendices (Fig. 53). fve species new to the Caucasus Ecoregion, 10 species new Despite the small number of specimens collected, their to Georgia, at least one new species to science and a fair similarity with the description of Trochosa cachetiensis amount of faunistic as well as aut- and synecological data Mcheidze, 1997 and the marked differences from the rarely collected in this quantity and quality in the Caucasus other species in question are striking. The habitus given in before; all of these forming the basis for this paper and Mcheidze (1997) shows the same reduced pairwise carapace a number of arachnological follow-up publications to marks as all three newly collected specimens, which only this transect study, which had not even been specifcally rarely occurs in this reduced variation in the other European designed as a spider study. These results exemplify the importance of systematic species. The large and bulging basal parts of the copula- and long-term collections of invertebrates for a thorough tory ducts of the female specimen (Fig. 53) are markedly assessment of species diversity in medium to highly diverse different from the other relevant species and very similar to taxa such as spiders and insects. Traditionally, and espe- the drawing given in Mcheidze (1997). This is also the case cially in the Caucasus, collections are made by hand during with the helmet-shaped markings, which appear rounded, the daytime or more rarely using pitfall traps by collectors when looked at through the epigynal cuticula (Fig. 52). No visiting regions of interest for a couple of weeks, only. specifc characters link the male specimens to the descrip- Systematic collections and transect studies using ground- tion of T. cachetiensis because the published drawing is too and arboricolous traps are almost non-existent in the arach- schematic and no relevant information was given in the text. nological study history of the Caucasus Ecoregion (but see A tarsal claw is absent in Mcheidze‘s drawing. The hook- Mikhailov & Mikhailova 2002). As seen in this paper, a like embolus is characteristic for T. robusta (Simon, 1876), single transect with Malaise traps, spanning several altitu- but the specimens differ from this species in the shape of the dinal zones and lasting the whole vegetation season in the median apophysis (Figs. 48–49) and the tarsus of the frst leg Caucasus Ecoregion can easily extend the species lists of (Fig. 40), which is stout and thick in male T. robusta (Fig. a study area, a country, the Caucasus Ecoregion and even 42). The frst leg of T. ruricola (De Geer, 1778) (Fig. 44) add new species to science—not to mention the signifcant shows no marked differences compared with the new speci- amount of never-before collected data on local species abun- mens, but they differ in the shapes of the median apophysis dance, altitudinal and habitat preferences of spider species, and embolus (Fig. 50) Males of T. hispanica Simon, 1870 phenology data etc.—even if it is the arachnologically best differ in embolus shape, median apophysis and patterns studied area in Georgia. of the frst leg (Figs. 46, 51). In females, the legs show no The admittedly huge effort of trap placement, fortnightly characteristics valuable in distinguishing the species (Figs. maintenance and post-collection sorting is easily mitigated 41, 43, 45, 47). The specimens also differ from the recently by the amount and quality of data collected for several taxa– described Caucasian species T. hirsuta Ponomarev, 2009 in one combined effort of systematic collecting! The 2014 (not studied), both in characteristics of the male bulb struc- transect study in the Lagodekhi Reserve has so far not only ture, relative length of the tarsus of the frst leg and epigynal yielded this faunistic overview on the spider fauna of the structure. Lagodekhi Reserve and initial to answers to other ecolog- Our collected specimens from the type locality of T. ical and taxonomic questions concerning the spider fauna of cachetiensis differ distinctly from all other species in the Caucasus (Otto & Japoshvili, in prep.) but also studies Europe and the Caucasus (Hepner & Milasowszky 2006; of the obtained insect specimens are starting to be published M. Hepner and A. Hänggi, pers. comm.), exhibit some of (Aslan, Mumladze & Japoshvili 2017; Japoshvili & Kost- the more distinct characters given in the species description jukov 2016; Japoshvili, Kostjukov & Kosheleva 2016; by Mcheidze (1997), and a second opinion (A. Ponomarev, Japoshvili et al. 2017). pers. comm.) further strengthens our conclusion that the Therefore, we stress the importance of systematic and specimens from Lagodekhi indeed belong to T. cachetiensis. long-term collections using traps becoming the norm rather We hope that this question can be studied more thoroughly, than remaining the exception in studying and monitoring when more specimens are collected and Tamara Mcheidze‘s invertebrate species diversity in the Caucasus Ecoregion. type material becomes available for study in the Janashia Only by such elaborate collecting efforts can biodiversity— Museum in Tbilisi. which is essentially the diversity of invertebrate taxa—in S. Otto & G. Japoshvili 385 the Caucasus Ecoregion be adequately assessed and efforts DUNIN, P. M. 1992: The spider family Dysderidae of the Caucasian fauna in nature and environmental protection be based on mean- (Arachnida Aranei Haplogynae). Arthropoda Selecta 1: 35–76. GUSEINOV, E. F. 1999: Spiders of Lenkoran Natural Region and ingful and exhaustive data. Absheron Peninsula in Azerbaijan (Autoreferat). Ph.D. thesis, Zoological Institute of the Azerbaijanian Academy of Sciences Baku. [in Russian] Acknowledgements HÄNGGI, A., HEER, X., TRIVELLONE, V., PALTRINIERI, P. & MORETTI, M. 2014: Eleven new spider species (Arachnida: Araneae) for Switzerland discovered in vineyards in Ticino– A number of colleagues greatly helped in the process of What are possible reasons?. Mitteilungen der Schweizerischen species determination, either by discussing issues, cross- Entomologischen Gesellschaft 87: 215–228. checking determinations or loaning specimens to the frst HEPNER, M. & MILASOWSZKY, N. 2006: The separation of Central author. For this invaluably help we are indebted to T. Blick European Trochosa females (Araneae: Lycosidae). Arachnologische Mitteilungen 31: 1–7. (Hummeltal, Germany), A. Hänggi (Basel, Switzerland), JAPOSHVILI, G. & KOSTJUKOV, V. 2016: The frst Record of Pronotalia M. Hepner (Vienna, Austria), P. Jäger (Frankfurt/Main, Gradwell, 1957 (Hymenoptera, Eulophidae) Wasps from Georgia. Germany), B. Knofach-Thaler (Innsbruck, Austria), M. Entomological Review 95: 855–859. Kovblyuk (Simferopol, Ukraine), C. Kropf (Bern, Swit- JAPOSHVILI, G., KOSTJUKOV, V. & KOSHELEVA, O. 2016: zerland), Y. Marusik (Magadan, Russia), K. Mikhailov New record of Tamarixia (Hymenoptera: Eulophidae) from Transcaucasia with some taxonomic and biogeographical notes. (Moscow, Russia), N. Milasowszky (Vienna, Austria), Journal of Entomological Research Society 18: 113–120. C. Muster (Putbus, Germany), A. Nadolny (Sevastopol, JAPOSHVILI, G., SALAKAIA, M., KIRKITADZE, G. & Ukraine), S. Ponomarev (Rostov-on-Don, Russia), A. BATSANKALASHVILI, M. 2017: List of encyrtids (Hymenoptera: Tanasevitch (Moscow, Russia), J. Wunderlich (Hirsch- Encyrtidae) from subalpine and alpine zones of Lagodekhi protected areas with new records from Georgia and Transcaucasia. Turkish berg, Germany). F. Walther (Hamburg, Germany) kindly Journal of Zoology 41: 342–344. collected spiders for the frst author on his travels in Georgia KIKODZE, A., GOKHELASHVILI, R., TAGAGARI, I., in 2009. S. Shetekauri (Tbilisi, Georgia) and H–J. Zündorf KAZALIKASHVILI, I. & KIKODZE, D. 2007: Protected areas of (Jena, Germany) patiently explained botanical concepts to a Georgia. In A. Kikodze & R. Gokhelashvili (eds.), Protected areas of Georgia. Tbilisi: Tsignis Sakhelosno. zoologist (frst author). J. Dunlop (Berlin, Germany) kindly KOVBLYUK, M. M., KASTRYGINA, Z. A. & MARUSIK, Y. M. 2013: had a gracious look at our English text. We also thank the The spider genus Pireneitega Kishida, 1955 in the Caucasus Rustaveli National Science Foundation for their fnancial (Aranei: : Coelotinae). Arthropoda Selecta 22: 59–73. supports under ref: FR/221/7-110/13. Finally, we express KULCZYŃSKI, L. 1895: Araneae a Dre G. Horvath in Bessarabia, Chersoneso Taurico, Transcaucasia et Armenia Russica collectae. our gratitude to Ms Tamar Chunashvili for her kind help in Természtrajzi Füzetek 18: 3–38. sorting material. KVACHAKIDZE, R. 1999: Vegetation of the Lagodekhi State Nature This study was supported by the Shota Rustaveli National Reserve. Tbilisi: Institute of Botany of the Georgian Academy of Science Foundation (grant FR/221/7-110/13). Sciences. [in Georgian] KVACHAKIDZE, R. 2009: Vegetation of Georgia. Tbilisi: Botanical Garden and Institute of Botany of the Georgian Academy of Sciences. [in Georgian] References LOGUNOV, D. V. 1996: A review of the genus Phlegra Simon, 1876 in the fauna of Russia and adjacent countries (Araneae: Salticidae: ABDURAKHMANOV, G. M. & ALIEVA, S. V. 2011: Results of a study Aelurillinae). Genus 7: 533–567. on the spider fauna (Aranei) of Dagestan. Southern Russia: Ecology, LOGUNOV, D. V. 1998: New species and new records of Palaearctic Development 1: 44–66. Sitticus (Araneae: Salticidae). Zoosystematica Rossica 7: 77–83. AGENCY OF PROTECTED AREAS 2016: Agency of Protected Areas. LOGUNOV, D. V. 1999: Redefnition of the genus Habrocestoides Tbilisi: Ministry of Environment and Natural Resources Protection Prószyński, 1992, with establishment of a new genus, Chinattus of Georgia, online at: http://apa.gov.ge/en/protected-areas/reserve gen n. (Araneae: Salticidae). Bulletin of the British Arachnological ASLAN, E. G., MUMLADZE, L. & JAPOSHVILI, G. 2017: List of leaf Society 11: 139–149. beetles (Coleoptera: Chrysomelidae) from Lagodekhi reserve with LOGUNOV, D. V. 2001: A redefnition of the genera Peckham new records for Transcaucasia and Georgia. Zootaxa 4277: 86–98. & Peckham, 1885 and Simon, 1885, with the BLAGOEV, G., DELTSHEV, C. & LAZAROV, S. 2015: The spiders establishment of a new genus Sibianor gen. n. (Araneae: Salticidae). (Araneae) of Bulgaria. Sofa: National Museum of Natural History, Arthropoda Selecta 9: 221–286. Bulgarian Academy of Sciences, online at: http://www.nmnhs.com/ LOGUNOV, D. V. 2015: Taxonomic-faunistic notes on the jumping spiders spiders-bulgaria/ of the Mediterranean (Aranei: Salticidae). Arthropoda Selecta 24: BLICK, T. & GOSSNER, M. 2006: Spinnen aus Baumkronen- 33–85. Klopfproben (Arachnida: Araneae), mit Anmerkungen zu Cinetata LOGUNOV, D. V. & GUSEINOV, E. F. 2002: Faunistic review of the gradata (Linyphiidae) und Theridion boesenbergi (Theridiidae). jumping spiders of Azerbaijan (Aranei: Salticidae), with additional Arachnologische Mitteilungen 31: 23–39. faunistic records from neighbouring Caucasian countries. CHARITONOV, D. E. 1948: A new species of Brachythele in Georgia. Arthropoda Selecta 10: 243–260. Soobshcheniia Akademii Nauk Gruzinskoi SSR 9: 135–139. [in LOGUNOV, D. V. & DEMIR, H. 2006: Further faunistic notes on Cozyptila Russian] and Xysticus from Turkey (Araneae, Thomisidae). Arachnologische CHARITONOV, D. E. 1956: Overview of the spider family Dysderidae Mitteilungen 31: 40–45. in the fauna of the USSR.. Uchenye Zapiski, Molotovskii LOGUNOV, D. V. & MARUSIK, Y. M. 1991: Redescriptions and Gosudarstvennyj Universitet Imeni A. M. Gorkogo 10: 17–39. [in morphological differences of Bianor aurocinctus (Ohlert) and B. Russian] aemulus (Gertsch) (Aranei, Salticidae). Sibirskii biologicheskii DUNIN, P. M. 1988: Cribellate spiders (Aranei, Cribellatae) of Azerbaijan. Zhurnal 2: 39–47. [in Russian] Entomologicheskoye obozreniye 67: 190–203. [in Russian] LOGUNOV, D. V. & MARUSIK, Y. M. 1992: The spider genus DUNIN, P. M. 1989: Fauna and altitudinal distribution of spiders Phoroncidia (Aranei, Theridiidae) in the USSR. Proceedings ot the (Arachnida, Aranei) of the Azerbaijan part of the southern Zoological Institute, Leningrad 226: 91–97. [in Russian] macroslope of the Caucasus Major. In A. B. Lange (ed.), Fauna LOGUNOV, D. V. & MARUSIK, Y. M. 1998: A brief review of the genus i Ekologiy Paukov i Skorpionov: Arakhnologicheskii Sbornik. Chalcoscirtus Bertkau, 1880 in the faunas of Central and the Moscow: Akademia Nauk SSSR: 31–39 [in Russian] Caucasus (Aranei: Salticidae). Arthropoda Selecta 7: 205–226. 386 Spiders of Lagodekhi Reserve, Georgia

LOGUNOV, D. V., MARUSIK, Y. M. & KOPONEN, S. 2002: Redescription OTTO, S. 2015: Commented checklist of the spider species (Araneae) in the of a poorly known spider species, Xysticus kulczynskii Wierzbicki Caucasus Ecoregion. Leipzig: Caucasian Spiders Database, online 1902 (Araneae: Thomisidae). Acta Arachnologica 51: 99–104. at: http://caucasus-spiders.info/wp-content/uploads/2015/10/2015_ LOGUNOV, D. V. & RAKOV, S. Y. 1998: Miscellaneous notes on Middle Otto_ChecklistCaucasus.pdf Asian jumping spiders (Aranei: Salticidae). Arthropoda Selecta 7: OVTSHARENKO, V. I., PLATNICK, N. I. & SONG, D. X. 1992: A review 117–144. of the North Asian ground spiders of the genus Gnaphosa (Araneae, MARTYNOVCHENKO, F. A. & MIKHAILOV, K. G. 2014: Spiders Gnaphosidae). Bulletin of the American Museum of Natural History (Aranei) of Teberda State Reserve: fauna and biotopic distribution. 212: 1–88. Euroasian Entomological Journal 13: 355–371. PONOMAREV, A. & ABDURAKHMANOV, G. 2014: Spiders (Aranei) MARUSIK, Y. M. 1987a: Comparative study of the webs of the orb- of North Caspian Coast and Islands. The South of Russia: Ecology, weaving spiders (Aranei, Araneidae, Tetragnathidae, ) Development 1: 76–121. [in Russian] from Lagodekhi reserve. Vestnik Zoologii 3: 83–86. PONOMAREV, A., ABDURAKHMANOV, G., ALIEVA, S. & MARUSIK, Y. M. 1987b: Three new species of the family Nesticidae DVADNENKO, K. 2011a: Spiders (Arachnida: Aranei) of the (Aranei) from the fauna of the USSR. Zoological Journal Moscow coasts and islands in northern Dagestan. Southern Russia: Ecology, 66: 461–463. Development 4: 126–143. [in Russian] MARUSIK, Y. M. 1989: New data on the fauna and synonyms of the PONOMAREV, A. V., ALIEV, M. A., KHALIDOV, A. K. & SHAVLUKOV, spiders of the USSR (Arachnida, Aranei). In A. B. Lange (ed.), Z. A. 2011b: Supplementary data on the spider fauna (Aranei) of Fauna i Ekologiy Paukov i Skorpionov: Arakhnologicheskii Dagestan. In Contemporary Pproblems in biology and ecology: Sbornik. Moscow: Akademia Nauk SSSR: 39–52 material reports at the international applied-science conference MARUSIK, Y. M. 2008: Synopsis of the rauda-group (Araneae, Makhachkala. Makhachkala: Dagestan State Pedagogic University: Thomisidae), with revalidation of Ozyptila balkarica Ovtsharenko, 77–82. [in Russian] 1979. Zootaxa 1909: 52–64. PONOMAREV, A. V. & CHUMACHENKO, Y. A. 2014: Spiders (Aranei) MARUSIK, Y. M. & KOPONEN, S. 2017: On two sibling species of in herpetobiont mesofauna of the Northwest Caucasus. The South of Dictyna (Araneae: Dictynidae) from Ukraine and Caucasus. Russia: Ecology, Development 2: 95–101. [in Russian] Entomologica Fennica 28: 41–48. PONOMAREV, A. V., KHALIDOV, A. K. & ALIEV, M. A. 2008: Additions MARUSIK, Y. M. & KOVBLYUK, M. M. 2004: New and interesting to the spider fauna (Aranei) of Dagestan. Communications of the cribellate spiders from (Aranei: Amaurobiidae, Dagestan State Pedagogic University (Biosciences) 3: 60–71. [in Zoropsidae). Arthropoda Selecta 13: 55–61. Russian] MARUSIK, Y. M., MIKHAILOV, K. G. & GUSEINOV, E. F. 2006: PONOMAREV, A. V. & KHATSHIKOV, E. A. 2009: Preliminary data on Advance in the study of biodiversity of Caucasian spiders (Araneae). the fauna of spiders (Aranei) of ‘Bolshoi Utrish’ (Russia, Krasnodar In C. Deltshev & P. Stoev (eds.), European Arachnology 2005. Acta Kray). Zhivotniy Mir Gornykh Territoriy: 109–113. [in Russian] zoologica bulgarica, Suppl. No. 1: 259–268. PONOMAREV, A. & KOMAROV, Y. 2013: Preliminary compilation of MARUSIK, Y. M., MIKHAILOV, K. G. & OMELKO, M. M. 2015: material on the spider fauna (Aranei) of North Ossetia-Alania. Trudy Taxonomic notes on spiders (Arachnida: Aranei) of the Russian Far Severo-Ossetinskogo Gosudarstvennogo Prirodnogo Zapovednika East. Arthropoda Selecta 24: 117–124. 2: 76–111. [in Russian] MCHEIDZE, T. 1964: Spiders (Araneina). Zhivotnyi mir Gruzii (AN Gruz. PONOMAREV, A. V., KOVBLYUK, N. M., CHUMACHENKO, Y. SSR Publ.) 2: 48–116. [in Russian] A. & VOLKOVA, D. D. 2012: Preliminary data on the fauna of MCHEIDZE, T. 1968: A study of arachnoids of the district. Trudy spiders (Aranei) of the Republic of Adygea. In G. G. Matisho & Tbiliskogo Gosudarstvennogo Universiteta 123: 213–223. [in R. D. Khunagov (eds.), Social-humane and ecological problems Georgian] of development of contemporary Adygea. Collection of scientifc MCHEIDZE, T. 1979: New species of spiders of the genus Dysdera papers. Rostov-on-Don: SSC RAS Publishers: 447–481 [in Latr. (Dysderidae) from Georgia. Soobshcheniya Akademii Nauk Russian] Gruzinskoi SSR 93: 421–468. [in Georgian] PONOMAREV, A. V. & MIKHAILOV, K. G. 2007: Addition to fauna of MCHEIDZE, T. 1983: New species of spider Brachythele recki sp. n. from spiders (Aranei) of the Russian Caucasus. In G. G. Matishov (ed.), Georgia. Fauna i ekologia bespozvonochnykh Gruzii, Tbilisi: 155– Studies of the southern scientifc center of the Russian Academy 159. [in Georgian] of Sciences. Rostov-on-Don: SSC RAS Publishing: 130–150 [in MCHEIDZE, T. 1997: Spiders of Georgia: systematics, ecology, Russian] zoogeographic review. Tbilisi: Tbilisi University. [in Georgian] PONOMAREV, A. V. & VOLKOVA, D. D. 2013: The frst results of the MCHEIDZE, T. 2014: Georgian spiders – systematics, ecology and study of spiders (Aranei) fauna of the Abrau Peninsula. In S. M. zoogeographic analysis. In S. Otto (ed.), Virtual library of Malkhazova & S. V. Velitshko (eds.), Biodiversity of State National biology. Frankfurt am Main: vifabioDOC, online at: http://dx.doi. Reserve “Utrish”. Anapa: Scientifc-technical Council of the State org/10.5431/mcheidze2014 online at http://nbn-resolving.de/urn/ Reserve “Utrish”: 228–247. [in Russian] resolver.pl?urn:nbn:de:hebis:30:3-325366 RAKOV, S. Y. & LOGUNOV, D. V. 1996: A critical review of the genus MIKHAILOV, K. G. 1987: Contribution to the spider fauna of the genus Heliophanus C. L. Koch, 1833 of middle Asia and the Caucasus Micaria Westring, 1851 of the USSR. I (Aranei, Gnaphosidae). (Aranei, Salticidae). Arthropoda Selecta 5: 67–104. Spixiana 10: 319–334. ROBERTS, M. J. 1987: The spiders of Great Britain and Ireland. Volume MIKHAILOV, K. G. 1990: The spider genus Clubiona LATREILLE 2: Linyphiidae and checklist. Colchester: Harley Books. 1804 in the Caucasus, USSR (Arachnida: Araneae: Clubionidae). SAARISTO, M. I. 1997: Description of Megalepthyphantes pseudocollinus Senckenbergiana biologica 70: 299–322. n. sp. (Araneae: Linyphiidae: Micronetinae). Bulletin of the British MIKHAILOV, K. G. 2013: The spiders (Arachnida: Aranei) of Russia and Arachnological Society 10: 257–259. adjacent countries: a non-annotated checklist. Arthropoda Selecta SCHMIDT, P. 1895: Beitrag zur Kenntniss der Laufspinnen (Araneae Supplement No. 3. Moscow: KMK Scientifc Press. Citigradae Thor.) Russlands. Zoologische Jahrbücher (Systematik) MIKHAILOV, K. G. & MIKHAILOVA, E. A. 2002: Altitudinal and 8: 439–484. biotopic distribution of the spider family Gnaphosidae in North SPASSKY, S. A. 1937: Some facts on the spiders fauna of the Black Sea coast. Ossetia (Caucasus major). In S. Toft & N. Scharff (eds.), European Sbornik Trudov Azovo-Chernomorskogo Selksokhozyaistvennogo Arachnology 2000. Proceedings of the 19th European Colloquium Instituta 5: 131–138. [in Russian] of Arachnology. Aarhus: Aarhus University Press: 261–265. TANASEVITCH, A. 1986: New and little-known species of Lepthyphantes MIKHAILOV, K. G., OTTO, S. & JAPOSHVILI, G. 2017: A new species Menge 1866 from the Soviet Union (Arachnida: Araneae: from the Clubiona caerulescens group from the Caucasus (Araneae: Linyphiidae). Senckenbergiana biologica 67: 137–172. Clubionidae). Zoology of the Middle East 63: 362–368. TANASEVITCH, A. V. 1987: The linyphiid spiders of the Caucasus, USSR NAKHUTSRISHVILI, G. 2013: The vegetation of Georgia (South (Arachnida: Araneae: Linyphiidae). Senckenbergiana biologica 67: Caucasus). Heidelberg: Springer. 297–383. NENTWIG, W., BLICK, T., GLOOR, D., HÄNGGI, A. & KROPF, C. TANASEVITCH, A. V. 1990: The spider family Linyphiidae in the fauna 2017: Spiders of Europe, version 2.2017. Bern: University of Bern, of the Caucasus (Arachnida, Aranei). Fauna of the terrestrial online at: www.araneae.unibe.ch invertebrates of the Caucasus. Moscow: Nauka: 5–114 S. Otto & G. Japoshvili 387

TANASEVITCH, A. 2010: A new Tenuiphantes Saaristo et Tanasevitch, WORLD SPIDER CATALOG 2017: World spider catalog, version 17.5. 1996 from the Caucasus, Russia (Arachnida: Aranei: Micronetinae). Bern: Natural History Museum, online at: http://wsc.nmbe.ch Arthropoda Selecta 19: 37–39. WUNDERLICH, J. 1995: Zur Kenntnis west-paläarktischer Arten der THALER, K. & NOFLATSCHER, M.-T. 1990: Neue und bemerkenswerte Gattungen Psammitis Menge 1875, Xysticus C. L. Koch 1835 und Spinnenfunde in Südtirol (Arachnida: Aranei). Veröffentlichungen Ozyptila Simon 1864 (Arachnida: Araneae: Thomisidae). Beiträge des Museums Ferdinandeum in Innsbruck 69: 169–190. zur Araneologie 4: 749–774. UTOCHKIN, A. S. 1968: Spiders of the genus Xysticus of the USSR fauna ZAITZEV, F. A. 1948: Overview on the cave fauna of Georgia. Trudy (Opredelitel’). Perm State University: 1–73. [in Russian] Zoologicheskogo Instituta AN SSSR 8(15): 1–173. VAN HELSDINGEN, P. J. 2016: Araneae, version 2016. Fauna Europea ZAWADSKI, A. M. 1902: Material on the fauna and biology of the Database, online at: www.european-arachnology.org WERSHBITSKY, S. 1902: O paukach Kavkaskovo kraja (De spiders (Araneina) of the Caucasus. Dnevnik Zoologicheskogo Araneis regionis Caucasicae). Zapiski Kiewskogo obshchestva Otdeleniya Obshchestva Lyubitelei Yestestvoznaniya, Antropologii Estestvoznaniya 17: 461–504. [in Russian] i Etnographii 3(3): 1–5. [in Russian] WILLIAMS, L., ZAZANASHVILI, N., SANADIRADZE, G. & ZYUZIN, A. A. & LOGUNOV, D. V. 2000: New and little-known species of KANDAUROV, A. 2006: An ecoregional conservation plan for the the Lycosidae from Azerbaijan, the Caucasus (Araneae, Lycosidae). Caucasus. Tbilisi: WWF Caucasus Programme Offce. Bulletin of the British Arachnological Society 11: 305–319.

Appendix: Checklist of the spiders in Lagodekhi Reserve

Previous References of records in Lagodekhi Synonyms and No. Species Author Caucasus Distribution Reserve and comments misspellings records Agelenidae 1 Eratigena agrestis (Walckenaer, 1802) Mcheidze (1997) Tegenaria agrestis 10 Holarctic 2 Tegenaria domestica (Clerck, 1757) Mcheidze (1964) 82 Cosmopolitan Amaurobiidae 3 Amaurobius pallidus L. Koch, 1868 Mcheidze (1997) 8 Europe–Caucasus 4 Amaurobius similis (Blackwall, 1861) Mcheidze (1997) 9 Holarctic 5 Amaurobius sp. this study (frst record Caucasus) – 6 Pireneitega spasskyi (Charitonov, 1946) Mcheidze (1964, 1997), Kovblyuk, Coelotes spasskyi 68 Caucasus Kastrygina & Marusik (2013), this study 7 accentuata (Walckenaer, 1802) Mcheidze (1997), this study 19 Palaearctic Araneidae 8 Aculepeira ceropegia (Walckenaer, 1802) Marusik (1987a), Mcheidze (1964, Araneus ceropegius 50 Palaearctic 1997) 9 Araneus diadematus Clerck, 1757 Mcheidze (1964), this study 143 Holarctic 10 Araneus grossus (C. L. Koch, 1844) Mcheidze (1997) 31 Palaearctic 11 Araneus marmoreus Clerck, 1757 Mcheidze (1964, 1997) 53 Holarctic 12 Araneus quadratus Clerck, 1757 Marusik (1987a), Mcheidze (1997), 33 Palaearctic this study 13 Araneus svanetiensis Mcheidze, 1997 Mcheidze (1997) 5 Georgia 14 Araniella alpica (L. Koch, 1869) Mcheidze (1997) Araneus alpicus 3 Europe–Caucasus 15 (Clerck, 1757) Mcheidze (1997) Araneus cucurbitinus 85 Palaearctic 16 Araniella inconspicua (Simon, 1874) Mcheidze (1997) Araneus incaspicuus 5 Palaearctic 17 Araniella opisthographa (Kulczyński, 1905) this study (frst record in Lagodekhi 13 Palaearctic Reserve) 18 Argiope bruennichi (Scopoli, 1772) Mcheidze (1997) 82 Palaearctic 19 Larinioides cornutus (Clerck, 1757) Mcheidze (1997) Araneus cornutus 28 Holarctic 20 Larinioides suspicax (O. Pickard-Cambridge, 1876) Wershbitsky (1902), Mcheidze (1997) Araneus folium 17 Palaearctic 21 Mangora acalypha (Walckenaer, 1802) Marusik (1987a), this study 106 Palaearctic 22 Neoscona adianta (Walckenaer, 1802) Mcheidze (1964, 1997) Araneus adiantus, A. 102 Palaearctic bisantinus 23 Neoscona subfusca (C. L. Koch, 1837) Mcheidze (1997) Araneus dalmaticue 23 Palaearctic 24 Nuctenea umbratica (Clerck, 1757) Mcheidze (1997) Araneus umbraticus 56 Europe–Caucasus 25 Parazygiella montana (C. L. Koch, 1834) Mcheidze (1997) Zigiella montana 25 Palaearctic 26 Singa hamata (Clerck, 1757) Mcheidze (1997) 12 Palaearctic 27 Singa lucina (Audouin, 1826) Mcheidze (1997) 8 Palaearctic 28 Singa nitidula C. L. Koch, 1844 Mcheidze (1997) 19 Palaearctic 29 Zygiella x-notata (Clerck, 1757) Mcheidze (1997) Zigiella x-notata 9 Holarctic Clubionidae 30 Kulczyński, 1882 Mikhailov (1990) 23 Palaearctic 31 Clubiona brevipes Blackwall, 1841 this study (frst record in Lagodekhi 8 Palaearctic Reserve) 32 Clubiona caucasica Mikhailov & Otto, 2017 Mikhailov (1990), this study formerly misidenti- 24 Caucasus, Turkey fed as C. caerules- cens L. Koch, 1867 388 Spiders of Lagodekhi Reserve, Georgia

Previous References of records in Lagodekhi Synonyms and No. Species Author Caucasus Distribution Reserve and comments misspellings records Clubionidae (cont.) 33 Clubiona corticalis (Walckenaer, 1802) this study (frst record in Lagodekhi 11 Palaearctic Reserve) 34 Clubiona golovatchi Mikhailov, 1990 Mikhailov (1990), this study 33 Caucasus 35 Westring, 1851 Mikhailov (1990, 1997), this study 63 Holarctic 36 Clubiona neglecta O. Pickard-Cambridge, 1862 Mcheidze (1997) 49 Palaearctic 37 Clubiona pseudosimilis Mikhailov, 1990 Mikhailov (1990), this study 21 Europe–Caucasus 38 Clubiona similis L. Koch, 1867 Mcheidze (1997) 28 Europe Dictynidae 39 Dictyna ottoi Marusik & Koponen 2017 this study (frst record in Georgia) 4 Europe–Caucasus 40 Lathys humilis (Blackwall, 1855) this study (frst record in Georgia) 17 Palaearctic 41 Nigma favescens (Walckenaer, 1830) this study (frst record in Georgia) 9 Europe Dysderidae 42 Cryptoparachtes fedotovi (Charitonov, 1956) Zaitzev (1948), Charitonov (1956), Harpactocrates 10 Caucasus Dunin (1992), Mcheidze (1997) fedotovi 43 Dysdera bogatschevi Dunin, 1990 Dunin (1992) 10 Caucasus 44 Dysdera crocata C. L. Koch, 1838 Mcheidze (1964, 1968, 1997) 50 Cosmopolitan 45 Dysdera hungarica Kulczyński, 1897 Dunin (1992), Mcheidze (1979, 1997) 30 Europe 46 Dysdera richteri Charitonov, 1956 Mcheidze (1997) 9 Caucasus 47 Harpactea zaitzevi Charitonov, 1956 Zaitzev (1948), Charitonov (1956), 9 Caucasus Dunin (1992) Eutichuridae 48 Cheiracanthium (Walckenaer, 1802) Mcheidze (1997) Chiracanthium[sic] 28 Palaearctic erraticum erraticum 49 Cheiracanthium mildei L. Koch, 1864 this study (frst record in Lagodekhi 32 Holarctic Reserve) 50 Cheiracanthium L. Koch, 1877 this study (frst record in Georgia) 6 Palaearctic montanum Gnaphosidae 51 Drassodes albicans (Simon, 1878) Mcheidze (1997) 2 Europe 52 Drassodes lapidosus (Walckenaer, 1802) this study (frst record in Lagodekhi 89 Palaearctic Reserve) 53 Gnaphosa dolosa Herman, 1879 Ovtsharenko, Platnick & Song (1992) 13 Palaearctic 54 Gnaphosa mcheidzeae Mikhailov, 1998 Mcheidze (1997) Gnaphosa caucasica 1 Georgia 55 Gnaphosa modestior Kulczyński, 1897 Ovtsharenko, Platnick & Song (1992) 6 Europe–Caucasus 56 Micaria formicaria (Sundevall, 1831) Mikhailov (1987) 16 Palaearctic 57 Micaria silesiaca L. Koch, 1875 Mikhailov (1987) 6 Palaearctic 58 Scotophaeus scutulatus (L. Koch, 1866) this study (frst record in Lagodekhi 20 Palaearctic Reserve) 59 Zelotes longipes (L. Koch, 1866) Mcheidze (1997) Zelotes serotinus 26 Palaearctic 60 Zelotes subterraneus (C. L. Koch, 1833) Mcheidze (1997) 23 Palaearctic Linyphiidae 61 Allomengea scopigera (Grube, 1859) Tanasevitch (1987) 1 Holarctic 62 Bisetifer cephalotus Tanasevitch, 1987 Tanasevitch (1987) 11 Europe–Caucasus 63 Bolyphantes alticeps (Sundevall, 1833) Mcheidze (1997), this study 14 Palaearctic 64 Bolyphantes lago- (Tanasevitch, 1990) Tanasevitch (1990) 2 Georgia dekhensis 65 Caviphantes dobrogicus (Dumitrescu & Miller, 1962) Tanasevitch (1987) Caviphantes 5 Europe–Caucasus dobrogica 66 Centromerita sp. Dahl, 1912 (Centromerita) this study – 67 Ceratinella scabrosa (O. P.-Cambridge, 1871) Tanasevitch (1987) 18 Palaearctic 68 Cinetata gradata (Simon, 1881) this study (frst record in the Caucasus) – Europe 69 Erigone dentipalpis (Wider, 1834) Mcheidze (1997) 36 Holarctic 70 Erigone longipalpis (Sundevall, 1830) Mcheidze (1997) 5 Palaearctic 71 Frontinellina frutetorum (C. L. Koch, 1834) Tanasevitch (1987), this study 86 Palaearctic 72 Incestophantes amotus (Tanasevitch, 1990) Tanasevitch (1990) Lepthyphantes 11 amotus 73 Linyphia hortensis Sundevall, 1830 Mcheidze (1997), this study 30 Palaearctic 74 Linyphia triangularis (Clerck, 1757) Mcheidze (1964, 1997), this study Liniphia triangularis 63 Holarctic 75 Megalepthyphantes cf. Saaristo, 1997 this study (frst record in Georgia) 2 Palaearctic pseudocollinus 76 Microlinyphia pusilla (Sundevall, 1830) Mcheidze (1997) Linyphia pusilla 34 Holarctic S. Otto & G. Japoshvili 389

Previous References of records in Lagodekhi Synonyms and No. Species Author Caucasus Distribution Reserve and comments misspellings records Linyphiidae (cont.) 77 Microneta viaria (Blackwall, 1841) Tanasevitch (1987), this study 90 Holarctic 78 Minicia alticola Tanasevitch, 1990 Tanasevitch (1990) 1 Georgia 79 emphana (Walckenaer, 1841) Mcheidze (1997), this study Linyphia emphana 51 Palaearctic 80 Neriene peltata (Wider, 1834) Tanasevitch (1987), Mcheidze (1997) 37 Palaearctic 81 Neriene radiata (Walckenaer, 1841) Mcheidze (1997), this study Linyphia marginata 13 Holarctic 82 Pelecopsis crassipes Tanasevitch, 1987 this study (frst record in Lagodekhi 12 Central Asia Reserve) 83 Pocadicnemis juncea Locket & Millidge, 1953 this study (frst record in the Caucasus) – Europe 84 Tenuiphantes contortus (Tanasevitch, 1986) Tanasevitch (1986), Mcheidze (1997) Leptyphantes [sic] 21 Caucasus lagodechensis), Lept- hyphantes contortus 85 Tenuiphantes mengei (Kulczyński, 1887) Tanasevitch (1987), this study Lepthyphantes 91 Palaearctic mengei

86 Tenuiphantes teberdaensisTanasevitch, 2010 this study (frst record in Georgia) 5 Caucasus 87 Tenuiphantes tenuis (Blackwall, 1852) Tanasevitch (1987) Lepthyphantes tenuis 29 Palaearctic 88 Trichoncus affnis Kulczyński, 1894 this study (frst record in Lagodekhi 1 Europe Reserve) Lycosidae 89 Alopecosa albofasciata (Brullé, 1832) Schmidt (1895) Lycosa albofasciata 14 Palaearctic 90 Alopecosa cuneata (Clerck, 1757) Mcheidze (1997) 16 Palaearctic 91 Alopecosa pulverulenta (Clerck, 1757) Mcheidze (1964, 1997) Tarentula pulveru- 58 Palaearctic lenta 92 Arctosa cinerea (Fabricius, 1777) Schmidt (1895), Wershbitsky (1902) Lycosa cinerea 17 Palaearctic 93 Arctosa lagodechiensis Mcheidze, 1997 Mcheidze (1997) 1 Georgia 94 Geolycosa charitonovi (Mcheidze, 1997) Mcheidze (1997) Alopecosa chari- 27 Caucasus tonovi 95 Hogna radiata (Latreille, 1817) Mcheidze (1964) Lycosa radiata, 71 Palaearctic Tarentula radiata 96 Lycosa piochardi Simon, 1876 Mcheidze (1997) 4 Central Asia 97 Lycosa praegrandis C. L. Koch, 1836 Mcheidze (1997) Lycosa nordmanni 11 Europe–Caucasus 98 Mustelicosa dimidiata (Thorell, 1875) Mcheidze (1997) Trochosa dimidiata 24 Palaearctic 99 Pardosa agrestis (Westring, 1861) Mcheidze (1964) Pardosa arenicola 83 Palaearctic pseudomonticola 100 Pardosa agricola (Thorell, 1856) Mcheidze (1997) Pardosa arenicola 17 Palaearctic 101 Pardosa amentata (Clerck, 1757) Mcheidze (1964) 30 Palaearctic 102 Pardosa bifasciata (C. L. Koch, 1834) Mcheidze (1997) Pardosa calida 30 Palaearctic 103 Pardosa hortensis (Thorell, 1872) Mcheidze (1964) 46 Palaearctic 104 Pardosa lugubris (Walckenaer, 1802) Mcheidze (1964) Pardosa chelata 45 Palaearctic 105 Pardosa ninigoriensis Mcheidze, 1997 Mcheidze (1997) 1 Georgia 106 Pardosa plumipes (Thorell, 1875) Mcheidze (1997) 2 Palaearctic 107 Pardosa pontica (Thorell, 1875) Zyuzin & Logunov (2000) 45 Europe–Caucasus 108 Pardosa wagleri (Hahn, 1822) Mcheidze (1997) 4 Palaearctic 109 Piratula knorri (Scopoli, 1763) Mcheidze (1997) Pirata knorri 6 Europe 110 Trochosa cachetiensis Mcheidze, 1997 Mcheidze (1997), this study 23 Caucasus 111 Trochosa ruricola (De Geer, 1778) Mcheidze (1964) 61 Holarctic 112 Thorell, 1856 Schmidt (1895), Mcheidze (1997) Lycosa terricola 46 Holarctic Nemesiidae 113 Raveniola recki (Mcheidze, 1983) Mcheidze (1983, 1997) B. zaitzewi 4 Georgia 114 Raveniola zaitzevi (Charitonov, 1948) Charitonov (1948), Zaitzev (1948), 4 Caucasus Mcheidze (1964) Nesticidae 115 Carpathonesticus Marusik, 1987 Marusik (1987b) 1 Georgia eriashvilii 116 Carpathonesticus Marusik, 1987 Marusik (1987b) 1 Georgia mamajevae Oxyopidae 117 Oxyopes heterophthalmus (Latreille, 1804) Mcheidze (1997) O. heteroptalmus 26 Palaearctic [sic] 118 Oxyopes lineatus Latreille, 1806 Mcheidze (1964), this study 76 Palaearctic 119 Oxyopes ramosus (Martini & Goeze, 1778) Mcheidze (1997) 6 Palaearctic 390 Spiders of Lagodekhi Reserve, Georgia

Previous References of records in Lagodekhi Synonyms and No. Species Author Caucasus Distribution Reserve and comments misspellings records Philodromidae 120 Philodromus albidus Kulczyński, 1911 this study (frst record in Georgia) 1 Europe 121 Philodromus aureolus (Clerck, 1757) Mcheidze (1997), this study 46 Europe 122 Philodromus cespitum (Walckenaer, 1802) this study (frst record in Lagodekhi 21 Holarctic Reserve) 123 Philodromus dilutus Thorell, 1875 Mcheidze (1997) 6 Europe–Caucasus 124 Philodromus dispar Walckenaer, 1826 this study (frst record in Lagodekhi 26 Holarctic Reserve) 125 Philodromus rufus Walckenaer, 1826 Mcheidze (1997), this study 10 Holarctic 126 Rhysodromus histrio (Latreille, 1819) Mcheidze (1997) Philodromus histrio 11 Holarctic 127 macellus Simon, 1875 Mcheidze (1997) 22 Palaearctic Pholcidae 128 Pholcus phalangioides (Fuesslin, 1775) Mcheidze (1964) 70 Cosmopolitan Pisauridae 129 Pisaura mirabilis (Clerck, 1757) Mcheidze (1997) 92 Palaearctic 130 Pisaura novicia (L. Koch, 1878) this study (frst record in Lagodekhi 27 Palaearctic Reserve) Salticidae 131 Attulus goricus (Ovtsharenko, 1978) Logunov (1998), this study Sitticus goricus 8 Caucasus 132 Attulus penicillatus (Simon, 1875) Logunov & Guseinov (2002), this Sitticus penicillatus 5 Palaearctic study 133 Ballus chalybeius (Walckenaer, 1802) Logunov & Rakov (1998), this study 37 Palaearctic 134 Chalcoscirtus pseudoin- Ovtsharenko, 1978 Logunov & Marusik (1998) 8 Europe fmus 135 Chinattus caucasicus Logunov, 1999 Logunov (1999) 45 Central Asia 136 Euophrys frontalis (Walckenaer, 1802) Mcheidze (1997) Evophrys [sic] 45 Palaearctic frontalis 137 Evarcha arcuata (Clerck, 1757) Logunov & Guseinov (2002), Logunov 91 Palaearctic (2015) 138 Heliophanus cupreus (Walckenaer, 1802) Rakov & Logunov (1996), this study 80 Palaearctic 139 Heliophanus dubius C. L. Koch, 1835 this study (frst record in Lagodekhi 9 Palaearctic Reserve) 140 Heliophanus patagiatus Thorell, 1875 Mcheidze (1997) 4 Palaearctic 141 Marpissa zaitzevi Mcheidze, 1997 Mcheidze (1997) 2 Georgia 142 Myrmarachne formicaria (De Geer, 1778) Logunov & Guseinov (2002) 32 Holarctic 143 Philaeus chrysops (Poda, 1761) Logunov & Rakov (1998), Logunov & 102 Palaearctic Guseinov (2002) 144 Phintella castriesiana (Grube, 1861) Logunov & Guseinov (2002) 34 Palaearctic 145 (Hahn, 1826) Logunov (1996) 26 Palaearctic 146 Pseudicius encarpatus (Walckenaer, 1802) this study (frst record in Lagodekhi 14 Palaearctic Reserve) 147 Sibianor aurocinctus (Ohlert, 1865) this study (frst record in Lagodekhi 4 Palaearctic Reserve) 148 Sibianor tantulus (Simon, 1868) this study (frst record in Georgia) – Palaearctic 149 Sittipub pubescens (Fabricius, 1775) Mcheidze (1997) Sitticus pubescens 1 Holarctic 150 Sittipub relictarius Logunov, 1998 this study (frst record in Lagodekhi Sitticus relictarius 15 Palaearctic Reserve) Segestriidae 151 Segestria bavarica C. L. Koch, 1843 this study (frst record in Lagodekhi 39 Europe Reserve) Tetragnathidae 152 Metellina mengei (Blackwall, 1869) Marusik (1987a) 5 Palaearctic 153 Metellina merianae (Scopoli, 1763) Marusik (1987a) 28 Palaearctic 154 Metellina segmentata (Clerck, 1757) Mcheidze (1964, 1997), this study Meta segmentata 62 Holarctic 155 Pachygnatha clercki Sundevall, 1823 Mcheidze (1997) Pachygnatha clarki 10 Holarctic 156 Pachygnatha listeri Sundevall, 1830 Mcheidze (1997) 13 Palaearctic 157 Tetragnatha montana Simon, 1874 Mcheidze (1997) 36 Palaearctic 158 Tetragnatha pinicola L. Koch, 1870 Mcheidze (1997) 13 Palaearctic Theridiidae 159 Anelosimus vittatus (C. L. Koch, 1836) this study (frst record in Lagodekhi 9 Palaearctic Reserve) 160 Dipoena braccata (C. L. Koch, 1841) this study (frst record in Georgia) 2 Europe S. Otto & G. Japoshvili 391

Previous References of records in Lagodekhi Synonyms and No. Species Author Caucasus Distribution Reserve and comments misspellings records Theridiidae (cont.) 161 (Clerck, 1757) Mcheidze (1997), this study 60 Holarctic 162 favomaculata (C. L. Koch, 1836) Mcheidze (1997) 5 Palaearctic 163 Heterotheridion nigro- (Simon, 1873) Mcheidze (1997), this study Theridion tubercu- 17 Palaearctic variegatum latum 164 Neottiura bimaculata (Linnaeus, 1767) this study (frst record in Lagodekhi 13 Holarctic Reserve) 165 Parasteatoda lunata (Clerck, 1757) Mcheidze (1997) Theridium lunatum 44 Palaearctic 166 Phoroncidia pilula (Karsch, 1879) this study (frst record in the Caucasus) – Palaearctic 167 impressa (L. Koch, 1881) Mcheidze (1997) Theridium impressum 68 Holarctic 168 Phylloneta sisyphia (Clerck, 1757) Mcheidze (1997) Theridium sisiphium 4 Palaearctic 169 tincta (Walckenaer, 1802) Mcheidze (1964, 1997) Theridium tinctum 30 Holarctic 170 Robertus scoticus Jackson, 1914 Mcheidze (1997) 1 Palaearctic 171 simile (C. L. Koch, 1836) Mcheidze (1997) Theridium simile 10 Holarctic 172 bipunctata (Linnaeus, 1758) Mcheidze (1997), this study 28 Holarctic 173 Steatoda castanea (Clerck, 1757) Mcheidze (1964, 1997) Teutana castanea 74 Holarctic 174 Steatoda triangulosa (Walckenaer, 1802) Mcheidze (1997) Teutana triangulosa 63 Cosmopolitan 175 Theridion albipes L. Koch, 1878 Mcheidze (1997) Theridium albipes 12 Caucasus 176 Theridion melanurum Hahn, 1831 Mcheidze (1997) Theridium denticu- 14 Holarctic latum 177 Theridion varians Hahn, 1833 this study (frst record in Lagodekhi 18 Holarctic Reserve) Thomisidae 178 (Fabricius, 1777) Mcheidze (1997) 13 Palaearctic 179 Simon, 1876 this study (frst record in Lagodekhi 2 Holarctic Reserve) 180 tricuspidata (Fabricius, 1775) Mcheidze (1997) tricus- 74 Palaearctic pidatus 181 (Clerck, 1757) Mcheidze (1964, 1997) 90 Holarctic 182 Ozyptila conostyla Hippa, Koponen & Oksala, Marusik (1989) Oxyptila conostyla 5 Palaearctic 1986 183 Ozyptila orientalis Ovtsharenko, 1979 Marusik (2008) 13 Caucasus balkarica 184 (C. L. Koch, 1837) Mcheidze (1997) Oxyptila praticola 11 Holarctic 185 (Blackwall, 1846) Mcheidze (1997) Oxyptila trux 14 Holarctic 186 Runcinia grammica (C. L. Koch, 1837) Mcheidze (1997) Runcinia lateralis 47 Palaearctic 187 Synema caucasicum Utochkin, 1960 Mcheidze (1997) Synaema caucasicum 10 Georgia 188 Tmarus punctatissimus (Simon, 1870) Mcheidze (1997) T. horvathi 6 Palaearctic 189 Xysticus acerbus Thorell, 1872 Mcheidze (1997) 12 Palaearctic 190 Xysticus bacurianensis Mkheidze, 1971 Logunov & Demir (2006) 31 Europe–Caucasus 191 Xysticus gallicus Simon, 1875 this study (frst record in Lagodekhi 33 Palaearctic Reserve) 192 Xysticus kochi Thorell, 1872 Mcheidze (1997) 96 Palaearctic 193 Xysticus kulczynskii Wierzbicki, 1902 this study (frst record in Georgia) 4 Central Asia 194 Xysticus lanio C. L. Koch, 1835 Mcheidze (1997) 21 Palaearctic 195 Xysticus nubilus Simon, 1875 Mcheidze (1997) 2 Europe 196 Xysticus pseudolanio Wunderlich, 1995 this study (frst record in the Caucasus) – Europe Titanoecidae 197 Titanoeca nivalis Simon, 1874 Mcheidze (1997) 26 Holarctic Trachelidae 198 Trachelas minor O. P.-Cambridge, 1872 this study (frst record in Lagodekhi 9 Palaearctic Reserve) Uloboridae 199 paradoxus (C. L. Koch, 1834) Zawadski (1902) 19 Palaearctic 200 Octonoba yesoensis (Saito, 1934) Marusik (1987a), Mcheidze (1997) Uloborus georgicus 5 Palaearctic 201 Uloborus plumipes Lucas, 1846 Mcheidze (1997) 20 Cosmopolitan Zoridae 202 Zora spinimana (Sundevall, 1833) this study (frst record in Lagodekhi 20 Palaearctic Reserve)