32 O.S. Boikova, N.M. Korovchinsky Arthropoda Selecta 4 (2): 33-48 ©ARTHROPODA SELECrA, 1995 the impact of predators were also emphasized by morphs of the Daplmia carinata King complex I I Limnol. Pijanowska [1990, 1992] but without spe~ifyingthe Oceanogr. Vol.26. ·P.201-218. Erection of infrageneric groupings within the spider genus · relative influence of different predators. · Hanazato T. 1990. Induction of helmet development by a CbaobontS factor in Daplmia ambigua during juvenile stages Clubiona Latreille, 1804 (Aranei Clubionidae): a typological apprpach Certainly, the above is only one of the possible I I J. Plankton Res. Vol.12. P.1287-1294. explanations. Seasonal morphological transformations Have! J.E. 1985. Predation of common ipvertebrate predators on may have more complex roots. According to Havel long- and shortfeathered Dapbnia retrocurva I I Hydro [1985], the enlarged helmet and spine in D. ret biologia. Vol.124. P.141-149. BhiAeAeHue BHYTPHPO,A.OBbiX rpynnupon6K B poAe nayKoB rocurva failed to prevent the rate of their consump Have! J.E. 1987. Predator-induced defences: a review I I Preda tion: direct and indirect impacts on aquatic communities (Ed. Clubiona Latreille, 1804 (Aranei Clubionidae): TitnoAomqecKUH noAXOA tion by Chaoborus and Leptodora. Accessory mar~ W.C. Kerfoot & A. Sih). Hanover, N.H., New England Press. phologiCal structures in zooplankters first of all P.263-278. perform hydrostatic functions, and as such they Kartashov A.J. 1983. [Numbers dynamics of CbaobontS and Kirill G. Mikhailov could have'been acquired as adaptations to life in Leptodora] I I Biotsenozy mezotrofuoP,o ozera Glubokogo. open water [Pijanowska, 1990; Fryer, 1991]. (Ed. N.N. Smimov). Moscow, "Nauka' Pub!. P.94-103 [in K.r. Muxa-H.Aon Russian]. 5. Acknowledgements . Kerfoot W.C. 1980. Perspectives on cyclomorphosis: Separation of · phenotypes and genotypes I I Evolution and ecology of Zoological Museum of the Moscow State University. Herzen Str. 6, K-9 Moscow 103009 Russia. Field studies were carried out at the "Glubokoe zooplankton communities (Ed. W.C. Kerfoot). Hanover, ozero'' Hydrobiological Station. The authors thank N.H., New England Press. P.470-496. 3ooAorwieCI<11H Myaeil Mry, yA. repg:eHa 6, K-9 Moci
Figs 1"11. Structure of male palp in the 'genus Clubiona: 1 - hypothetical primitive construction, 2-11 - morphological types: 2-3 - Paraclubiona: C. corticalis, 3 - C. kurosawai, 4 - ]aponiona: C. japonica, 5 - Bicluona: C. jucunda, 6-11 - Clubiona s.str.: 6 - C. mannorata, 7 - C. reclusa, 8 - C. trivialis, 9 - C. subtilis, 10 - C. basamkini, 11 - C. abboti, 1-3,6,8,9 - lateral view, 4,5,10,11 -ventral view, 7- bulb, ventrally. 1 -after Kraus, 1984, 2,6,9- after Wiehle, 1965, 3- afte~ Ono, 19S6a, 11 - after Dondale & Redner, 1982, Abl;>reviations: Cb- cymbium, E - embolus, EC -conductor of embolar type, EP _: embolar part, H - hematodocha, P - protector, ST - subtegulum, TA - tegular apophysis, TC - conductor of tegular type, TP - tegular part, P11c. 1-11, OrpoeHI1e· IIMhiTbi caMga n poAe Clubiona: 1 -- rnrroTeTM'IeCIGUI rrp11M11TMnHaJI KOHCTPYKIJI1JI, 2-11 - MOp ( 1) include the species with EP either sclerotized Morphotype 1a. Tegulum normal, non~sclero ·.~···~ :~'1 . ; ;.~\· . ( 1a) or non-sclerotized ( 1b; Fig. 2). A considerable tized. TiA varies from very weakly to very strongly .· .. .., .. : :·. enlargement of TP is peculiar in the latter case. In developed. Conductor, if any, of embolar type. . . . . ' S' •., • ,••• , another grouping (2), TP sclerotization is expressed Sometimes membranous protector/ "conductor" . (.. •• • 'tv{.. ••• . •.'ij• only fragmentarily, being provided by a hook- or .structures on tegulum. · · w. ··v hammer-like outgrowth (Figs .4-S); EP is sclerotized · The necessity in merging 1b and 1' groupings is ~ ... ,b;.~.,·'=··• ,, -12 insignificantly and TP enlargement is absent .. confirmed below by the data obtained upon the . Beyond these groupings, there are some species . female genitalia. . · with both TP'enlargenient and a non-sclerotized EP, How well do the above male morphotypes corre combined with a fragmentarily sclerotized TP ( 1', spond to the female ones? Surprisingly, an almost Fig. 3). It is remarkable that the sequence of equal arrangement of the groupings is found in ~plitting in··· tQ groupings firstly by T.P ( 1-2! and then female Clubiona as well. by EP (1a-1b) accepted here is qutte arbttrary and •· Similarly to other spiders, the Clubiona ende can be changed intp the r~verse mode. gyne, i.e. the inner female genitalia, opens by paired Species with strongly sclerotized EP ( 1a) CO of different shape ranging from drcular to differ by. pec11liarities of both E and EP-apophysis fissure-like. Sometimes CO open into a single (Figs 6-7; 11, ·etc.). Apparently, these, details midline groove. CO are disposed at the fore (type 1b represent le.ss generalized characters than sclero + 1', Fig. 12), middle (type 2, Fig. 13), and hind tization-membrimization of either both totally TP (type 1a, Figs 14-22 etcJ part of the genital plate. & EP or TP-enlargement. A considerable buJb Beginning from CO, paired CT direct backwards enlargement of some sp~cies of this grouping (C. ( 1b + 1'), forewards ( 1a), or straight toward the genevensis, C. viridula, etc.) is due to EP, mainly d~pth of the endegyne (2). The only exception is C. to E itselL . · jucunda,, representing a species with the male and As a r1,1le,TiAis well-developed and is important female attributed to d.ifferent morphotypes, 2 (Fig. for species determination in grouping 1a (Figs 8-10 5) and 1a (Fig. 14), respectively. This species etc), whereas in other Clubiona it is either weakly belongs to a mixed morphotype [Mikhailov, 1994]. developed ( 1b + 1 ')or rather uniform, although it Hence, the diagnoses of the morphotypes given can be used for'species identification as \Yell (2). above (except of C. jucunda) must be complement E is functiona:lly associated with a certain struc ed as follows: ' tun! of eithei· tegular or embolar nature, known as . 1b + .1 '. CO in the fore pai't of the epigyne. CT conductor. In some· species, the function of the directed backwards. conductor in COfUlation is obvious, lying in directing · 2. CO at the middle part of the epigyne. CT 16 the embolus movement. Such a conductor with a well directed straight into the depth of the endegyne. expressed role is common in the following groupings: 1a. CO in the hind part or at the hind edge of the •ffiosfspecie'sof 2 (of tegular nature), two rei?resenta epigyne. CT directed strongly forwards .. tives 'oftb +1' (C. pyrifera, C. kurosawm, also of Each of these morphotypes is here allotted the tegular nature), and certaihspecies of 1a (of embolar · subgeneric status. The basic differences between nature - Figs 6, 7). The conductor in grouping 2 is Clubiona subgenera lie in the structure of the bulb weakly sderotized, · but it is peculiar in shape, and the direction of the epigynal tubes, both surrounding E from three sides except ventrally (Fig. apparently more general characters. On the other 4). In many species, a long and thin E of the slipping hand, at the species level such details as the sti·ucture typ'e. (after Wiehle, 1965: Einstoi3-Embolus) lies of TP, EP, and TiA, as well as SPTae and atria are Figs 12-22. Structure of female epigyne and vulva in the genus Clubiona: 12 - Paraclubiona: C. corticalis, 13 - Japoniona: C. calmly, i.e. in a non-erected position, on the tegular more valuable .. According to the principle of the japonica, 14 - Bicluona: C. jucunda, 15-22- Clubiona s.str.: 15 - C. trivialis, 16 - C. abboti, 17 - C. maritima, 18 - C. brrNipes, surface (see Figs 23, 25, 59, 62, etc.). That extent of specific meronomic-taxonomical relation; these lat 19- C. comta, 20- C. genrNensis, 21 -C. terrestris, 22- C. similis. 12-15- epigyne, 16-19- vulva, ventrally, 21-22- vulva, TP surface is distinguished by strong membraniza ter five characters .are obviously less gen,eralized inner view (dorsally). 12, 18-22- after Wiehle, 1965, 15,17- after Dondale & Redner, 1982. tion, it has been named earlier also a "conductor", than those of the subgenera. . . P11c. 12-22. OrpoeHHe 31111111Hbi 11 ny,u,nDI CaMI The subgenus Paraclubiona includes only one sis ("tegular apophysis", after Dondale .& Redner, species-group, which formally can be named the 1982) (Fig. 23), slit-like CO combined with a 23 corticalis-group5. This statement is quite prelimi semicircular CT approximating each other, SPTae nary, since numerous species from Southeast Asia ·adjoining each other, and a strongly sclerotized and Australia require a modernTevision. posterior edge of the epigyne (Fig. 17); ]aponiona comprises one.group, whereas Biclu the chabarovi-group: stereospecificity of a char ona is represented by a single species only. acteristic TiA and the epigynal depressions disposed By the conjugation criterion, the following separately_from CO (Figs 52, 63). . groups are distinguishable within Clubiona s.str.: With certain reservations, the obesa-group, (Figs the reclusa-group: a harpoon-like TiA (Figs 60- 32-37, 40-47) is also submitted to this criterion, but 61) and a typical protector oil the male bulb {Fig. the conjugated characters of both male and female 7), c,i considerable distance between the atrium and genitalia display a considerable "infragroup" vari SPT, as well as both direction and shape of the tubes ability. For example, a broad TiA convex ventrally and cavities of the female vulva (Figs 49-50); and concave dorsally is peculiar in the males, but the caerulescens-group: a strong deviatioll. in this apophysis can be either turned around the epigynal structure followed by the male E and TiA longitudinal axis, as in the akagiensis-group, or (Figs 51, 62; seealso Wiehle, 1965, Abb. 106-109)); curved spirally, as in 9. kimyongkii (Fig. 36). the trivi(Jlis-group: a flat TiA combined with a Sinuous CT of the female are typical (Figs 40-42, 44- wide E; a simplestnicture of a circular SPT (Figs 8- 47), but in the sapporensis-subgroup they are almost 10, 15); . . straight (Fig. 43). . the lutescens-grou£: ·a 'bifurcate flat TiA com The·pallidula-group (Figs 48, 53-59) occupies a bined with a wide E (Figs 27-28); an ovoid atrium special position among the others, being distin combined with a curved tubular SPT (Fig. 21); ' guished mainly by the structure of the epigyne, the similis-group: a bifurcate flat TiA of a. namely by the distinctly bifurcate, tubular, rarely different patt~rn combined with a thinE (Fig. 29); ovoid SPTae, as in Fig. 48. The males are character a simple tubular or ovoid non-curved SPT (Fig. 22)'; ized by some polythetical characters (Table 1; cp. the abboti-group: a strongly developed apophy also Figs 53-59) which, combined, outline a typical sis of EP ("tegular apophysis", after Dondale & congregation in the sense of Smirnov [1923, 1924], Redner, 1982), convex ventro-prolaterally and con i.e. a cluster of close species in a multidimensional cave dorso-retrolaterally, combined with a bifur character space. · cate, non-flat TiA (Fig. 11 ); CO paired or fused in Including a single species, the zilla-group (Figs , a groove, a semicircular CT combined with a 31, 39) differs exclusively by the male palp struc wrinkled genital plate projecting beyond the epigas ture. It possesses a large bifurcate EP-apophysis tric furrow (Fig. 16); · ("tegular apophysis:·. after Ono,_ 1986). By the the maritima-group: a big, spur-like EP-apophy- structure of the ep1gyne, accordmg to a recent redescription (Ono, 1986, Figs 6-8), the female of C. zilla is close to C. bakurovi (the obesa-group), 5C. Deeleman-ReLnhold (personal communica displaying no fundamental differences from the set tion) .proposes two species-groups in the Indoneasin of diagnostic features of the latter group. fauna, ofPardclubiona. Thus, the separation of 10-11 species-groups Figs Z3-37. Structure of male palp.in the subgenus Clubiona s.str.: Z3 - C. maritima, Z4 - C. brevipes, Z5 - C. comta, Z6 - C. genevensis, Z7 - C. alpicola, Z8 - C. saxatilis, Z9 - C. ncglecta, 30 - C. japonicola, 31 - C. zilla, 3Z - C. kurilensis, 33 - C. furcata, 34 _:_ C. sapporensis, 35- C. kunasbircmis, 36 - C. kimyongkii, 37- C. irinae. Z3,Z5-Z6,31,34- ventral view, Z4,Z9-30,3Z - lateral vieW, Z7 - upper half ofpalp, ventrally, Z8 - TiA, laterally, 33 ,-- TiA, dorsally, 35 - TiA, dorsolaterally, 37 - TiA, innet view. Z3,33 ~after Dondale & Redner, 198Z, Z4-Z7,Z9- after Wiehle, 1965, Z8- after Thai er, 1981, 31 -after Ono, 1986b, 3Z- after Hayashi, Chikuni, 1984. Abbreviations as in Figs l-11, and "P" -protector-like structure in the group lutescens . . P11c. Z3-37.,CTPOeHHe IIaAbiibi caMlJa BrroApoAe Clubiona s.str.: Z3- C.nuiritim{i, Z4- C. brevipcs; Z5- C. ~omta, Z6- C. gcnevensis, Z7 ,,C. alpicola, Z~ - C. saxatilis, Z9- C. neglecta, 30 - C. japonicola, 31 - C. zilla, 3Z- C. kurilensis, 33 - C. furcata, 34 - C. sapporensis, 35 - C. kunasbircnsis, 36 - C. kimyongkii, J7 - C. irina'c. Z3,Z5-Z6,31,34 - nel!TpaAbHo, Z4,Z9c30,3Z - BY!A c6or 59 E- 57 Figs 51-63. Stmc~re offemale e~igyne (51-52) and ?'~ale _ralp (53-63) in t~e subgenus Gubiona s.str.: 51, 62- C. caemlescens, 52, 63 - C. .cbabarovt, 53 - C. pallidula, 54 - C. sopaikenstS, 55 - C. ezoensts, 56 - C. propinqua, 57,58 - C. mayumiae, 59 - C. odesanensts, 60 - C. subsultans, 61 - C. kulczynskii. 51-52 - epigyne, 53 - TiA, dorsally, 54 - TiA, inner view, 55 - upper part of bulb, ventrolaterally, 56.- upper part of bulb, ventrally, 57,~3 - ventral view, 58 - TiA, ventrolaterally, 59 - upper part of palp, ventrolaterally, 62 - T1A, laterally, 58,61 - ventrolateral VIew. 51,53,60-62 - after Wiehle 1965. Abbreviations as in Figs 1-11. ' P11c. 51-63. CTjX>eHI1e 3nl1l"I1Hbi CaMKI1 (51-52) 11 naAbnbi caMga (53-63) n noApoAe Gubiona s.str.: 51, 62- C. caemlescens, 52, 63 - C. c!Jabarovi, 53 - C. pallidula, 54 - C. S~J!aikensis, 55 - C. ezoensis, _56 - C. propinqua, 57,58 - C. mayumiae, 59 - C. ' Figs 38-50. Structure of female epigyne and vulva in the subgenus Gubiona s.str,: .38 - C. japonicola, 39 - C. zilla, 40 - C. odesanensts, 60- C. subsultans, 61 - C. kulczynskii. 51-52- 3nl1r!1Ha, 53- T1A, AOpcaAbHo, 54- TiA, BI1A 113Hyrp11, 55- nepxm111 latericia, 41 ---:- .C. wolcbongsensis, 42 - C. kunasbirensis, 43 - C. sapporensis, 44 --: C. kimyongkii, 45 -. C. cbikunii, 46 - C. bakurovi, '!aCTb 6yAh6yca, neHTjX>AarepaAbHo, 56 - nepxH.fl.fl '!aCTb 6y~6yca, neHrpaAbHO, 57,63 - neHrpaAbHO, 58 - TiA, neHTjX>AarepaAbHO, 47 - C. irinae, 48 - C: pbragmitis, 49 - C. stagnatilis, 50 - C. basbkirica. 38-45,48,49 -vulva, dor~ally, 46,47,50 - epigyne, 38 59 - nepXH.fl.fl '!aCTb nal\hnyca, neHTjX>AarepaAbHO, 62 - T1A, BJ1A c601 degree of morphological generalism of diagnostic capacity for a change. Certainly there is less room for shape. Tegular apophysis of a characteristic shape (hook or medium-sized, straight or slightly arched. Atrium (essential) characters is lower than in species genitalic variability in spider females than in males. or hammer-like), placed in distal part of tegulum. E lori positioned just in front of SPT. SPT unipartite, ovoid or. groups, i.e. there are only little differences between Being surrounded from three sides by abdominal or filiform, directed distad, then retrolaterad. circular. Cover of genital plate thin, usually non-rugose species inside such subgroups. Thus, in the propin organs, the epigyne is actually a structure rather Female. Genital groove either large, sometimes partly (except C. pygmaea). qua~subgroup (pallidula-group) all species includ limited in volume; the shape of its cavities and canals divided,or missing (C. vigil). CO and CT as in Diagnosis. Clubiona (Clubir;ma) trivia/is C:L. Koch, 1843 ed, C. prdpinqua, C. pseudogermanica and C. can be only circular, ovoid, or tubular. Whereas the Atrium either large, vesiculiform, circular, positioned in Clubiona (Clubiona) amurensis Michailov, 1990 hind part of genitalia (C. vigil) or oblong-ovoid and placed Ono, 1992, mayumiae, are almost identical in TiA structure growing capacities of the male bulb are far less Clubiona (Clubiona) aspervida (Fig. 58); such a peculiarity has hitherto remained in fore part. SPT either considerably less in size than atrium Clubiona (Clubiona) baimaensis Song et M. Zhu, 1991 stringent, as well illustrated e.g. by the subgenus and.situated near its back part or almost equal. in size to Clubiona (Clubiona) basarukini Michailov, 1990 unknown in other Clubiona. Differences between Paraclubiona. Not being hollow, the apophyses of the atrium and situated in front of it (C. vigil). Clubiona ( Clubiona) diversa 0. Pickard-Cambridge, 1862 these species lie in body size (with a considerable palp tibia, tegulum, and embolar· part can be consid Clubiona (Japoniona) japonica L.Koch, 1878 Clubiona (Clubiona) duoconcava Zhang et Hu, 1991 intergradation), details of E structure (C. propin erably more slender, acute, branched, etc., that is, Clubiona (Japoniona) flavipes (Saito, 1939) Clubiona (Clubiona) insulana Ono, 1989 qua - all others, in both cases - Figs 56, 57), and much more variable as compared to vulval canals. Clubiona (Japoniona) vigil Karsch, 1879 Clubimia (Clubiona) janae Edwards, 1958 direction of the seminal duct inside EP {C. pseudoger The above statement that the range of variability Clubiona (Clubiona) juvenis Simon, 1878 manica - C. mayumiae). Female distinguishing differs in male and female genitalia has been intro Subgenus Bicluona Michailov, 1994. Clubiona (Clubiona) moesta Banks, 1896 features are also very slight. Similarly, the geneven duced in a discussion of the role of the key-lock Clubiona (Clubiona) pygmaea Banks, 1892 sis-subgroup (comta-group) (Figs 20, 26) can be Type species: Clubiona jucunda (Karsch, 1879). Clubiona (Clubiona) quebecana Dondale et Redner, 1976 (hand-glove) principle in sexual character selection Clubiona (Clubiona) rostrata Paik, 1985 characterized solely by some minor differences· in a [Eberhard, 1985]. Yet this circumstance seems Diagnosis. The male differing by the singular position of E mostly lying between the bulb and the cymbium (Fig. Clubiona (Clubiona) subrostrata Zhang et Hu, 1991 generally uniform structure of the male palp and unimportant, because there are many other reasons Clubiona (Clubiona) subtilis L. Koch, 1866 endegyne. 5). The female differing from that of Paraclubiona and to reject the key-lock principle. j aponiona by CO placed caudally and CT directed Clubiona (Clubiona) subtrivialis Strand, 1906 Besidesthat, a lot of groups have reasonably long forwards (Fig. 14). Clubiona ( Clubiona) transbaicalica Miehailov, 1992 been established within the obesa-group. The latter Characteristics of infrageneric groupings Description. Male. TiA short, npt branched. Tegular ? Clubiona ( Clubiona) yangmingensis Hayashi et Yoshida, 1993 is quite species-rich and differs only by a tendency apophysis rectangular, not unciform. Tegular conductor of variation in some characters (see above), i.e. a Subgenus Paraclubiona (Lohmander, 1945). missing. E filiform. The abboti-group highly dynamic archetype. Separation of some smaller Female. A large genital groove in hind part of epigyne. Diagnosis. Male (Fig. 11): apophysis of EP strongly complexes seems to be useful there from a purely Type species: Clubiona corticalis (Walckenaer, 1802). CT directed forwards, sinuous. SPT large, vesiculiform. practical viewpoint. Besides subgroups encompass Clubiona (Bicluona) jucunda (Karsch, 1879) developed, convex ventro-prolaterally and concave dor Diagnosis. The male differing by the tegular enlarge so-retrolaterally, TiA bifurcate, not flat. Female (Fig, ing two and more species, certain species exist in the ment of the bulb (Figs 2-3), the female by the frontal Subgenus Clubiona sensu stricto. 16): CO either paired or fused into genital groove, CT obesa-group that display highly disjunct and ste position of CO (Fig. 12). semi-circular; genital plate rugose, protruding beyond reospecificallyarrangedgenitalia. Thus, C. bakurovi Description. Male. TiA weakly developed, of relative epigastric furrow. and C. irinae also belong iri species-subgroups of ly simple shape. Bulb large, protruding. Conductor, if Type species: Clubiona pallidula ( Clerck, 1757). Description. Male. TiA as in Diagnosis, with ventral their own. Altogether, no fewer than six subgroups existing, of teguJar nature, weakly sclerotized, complete Diagnosis. The male differing by the conductor of branch always directed parallel to main axis. Base of EP can be established within the Palaearctic obesa ly separated from tegulum, lacking in C. corticalis, C. embolar nature combined with the sclerotized tegular apophysis placed in middle part of bulb, this apophysis cirrosa. Tegular apophysis weakly sclerotized, large (C. apophysis wanting; the bulb is either usually not surpass covering EB. E long, arched around or angled across bulb group: the akagiensis- (Japan), the latericia-, the kurosawai - Fig. 3), small (C. cirrosa) or lacking (C. ing the cymbium or enlarged due to EP. The female sapporensis-, the corrugata-, the bakurovi-, and the· tip; E top lying in proximal half of bulb (except C. corticalis, C. pyrifera). E fili-, spiniform or very short differing by the position of CO either in the hind part or angulata). Bulb surface with a membranous conductor. irinae-subgroups (for the geographical distribution and almost missing (C. pJJrifera), directed straightly at the rear edge of the epigyne as well as by CT directed of the last five of these, see Mikhailov, 1992). Female. CO large (only in C. angulata small), either distad or spiralling at tip (C. cirrosa). forwards. separated (in most species) or f~sed in~o genital groove, A total subdivision of all Clubiona species-groups Female. CO in fore part of epigyne, removed far away Description. Male. TiA usually well-developed and either partly (C. opeongo, C. ltttoralts) or totally (C. seems impossible due to morphologicalr~easons. from epigastric furrow. CT directed caudally. Atrium sclerotized, often of a complex shape. Bulb usually catawba). CT arched or sinuous (C. angulata). Other Establishment of three levels of infrageneric vesiculiform, poorly visible, positioned in hind part of elongated. Embolar conductor not always expressed. E of features as in Diagnosis. groupings (subgenus - species-group - species genitalia. SPT tubular, sometimes circular, considerably variable shape, from aciculi- and aculeiform to oblong Clubiona (Clubiona) abboti L. Koch, 1866 subgroup) is undertaken here for ordination both of less in size than atrium. , loriform, directed retrolaterad or proximad, suffi<;iently Clubiona (Clubiona) adjacens Gertsch et Davies, 1936 Clubiona (Paraclubiona) corticalis (Walckcnaer, 1802) long coiling around tegular tip. Clubiona (Clubiona) angulata Dondale et Redner, 1976 generalism arid the extent of morphological differ Clubiona( Paraclubiona) chakrabartei Majumder et Tikadcr, Female. CT more often long. Atrium sderotized, ences. Nonetheless, at each level the groupings Clubiona (Clubiona) bishopi Edwards, 1958 1991 usually placed in fore part of genitalia. SPT ovoid, Clubiona (Clubiona) catawba Gertsch, 1941 themselves differ considerably by the extent of Clubiona (Paraclubiona) cirrosa Ono, 1989 circular or tubular, sometimes bipartite and strongly morphological similarity between species included. Clubiona (Clubiona) dyasia Gertsch, 1941 Clubiona (Paraclubiona) concirma (Thorell, 1887) curved sagittally. Clubiona (Clubiona) estes Edwards, 1958 Thus, at the subgeneric level, the greatest degree of ? Clubiona (Paraclubiona) drassodes 0. Pickard-Cam Clubiona (Clubiona) gertschi Edwards, 1958 bridge, 1874 congeneric similarity seems to occur amongst]aponio The trivia/is-group Clubiona (Clubiona) johnsoni Gertsch, 1941 na species, while at the species-group one among the Clubiona ( Paraclubiona) kurosawai Ono, 1986 Clubiona (Clubiona) kagani Gertsch, 1941 reclusa-grouf members. This further supports the Clubiona ( Paraclubiona) lyriformis Song et M. Zhu, 1991 Diagnosis. The male differing by the not branched, Clubiona (Clubiona) kastoni Gertsch, 1941 Clubiona (Paraclubiona) melanosticta Thorell, 1890 specificity o the meronomic-taxonomical relation, wide, flat or almost flat TiA, weakly angled or dentate, Clubiona (Clubiona) kiowa Gertsch, 1941 Clubiona (Paraclubiona) mora/is Song et M. Zhu, 1991 rarely attenuating (Figs 8-10). The female differing by Clubiona (Clubiona) /ittoralis Banks, 1895 for a considerably more widely distributed feature is Clubiona (Paraclubiona) pyrifera Schenkel, 1936 the simple structure of circular or ovoid SPTae combined Clubiona (Clubiona) marna Roddy, 1966 known to be a general meronomic-taxonomical Clubiona (Paraclubiona) shillongensis Majumder et Tika- with CO brought close together or fused into a single Clubiona (Clubiona) mutata Gertsch, 1941 inconformity [Lyubarsky, 1991]. der, 1991 groove, and CT straight or almost straight (Fig. 15). Clubiona (Clubiona) newmani lvie et Barrows, 1935 The female spider genitalia have traditionally been Clubiona (Paraclubiona) yaginwnai Hayashi, 1989 Description. Male. TiA as in Diagnosis, tending to Clubiona (Clubiona) nicholsi Gertsch, 1941 viewed as less variable than the male ones [Eberhard, enlarge in C. rostrata and C. transbaicalica. EP with a Clubiona (Clubiona) odelli Edwards, 1958 1985: 30]. This trend is especially well-pronounced in Subgenus ]aponiona Michailov, 1990. small apophysis near EB, sometimes bearing 1-3 teeth. EB Clubiona (Clubiona) opeongo Edwards, 1958 araneoid spiders, such as the Linyphiidae, to a lesser covered by EP-apophysis (except C. insulana). E placed Clubiona (Clubiona) oteroana Gertsch, 1941 extent also in the Theridiidae. The same phenomenon Type species: Clubiona japonica L.Koch, 1878. totally in apical part of bulb, arched around or angled Clubiona (Clubiona) pikei Gertsch, 1941 is characteristic of Clubiona as well. Diagnosis. The male differing by the characteristic across tegular tip, sometimes stretching proximad along - Clubiona (Clubiona) plumbi Gertsch, 1941 Expressing it more exactly, at the species level shape of the tegular conductor (Fig. 4). The genital a membranous, slightly depressed "conductor" up to C/ubiona (Clubiona) pomoa Gertsch, 1941 the degree of generalism of differences (differential groove or CO placed in the middle of the epigyne. CT midway of bulb (C. trivia/is, C. rostrata). Seminal auct C/ubiona (Clubiona) procteri Gertsch, 1941 characters, see Nelson & Platnick, 1981: 160-164) in directed downwards, either retracted in the depth of the relatively short. Clubiona (Clubiona) rhododendri Barrows, 1945 epigyne or shifted a little forwards (Fig. 13). Female. Hind edge of epigyne usually projecting a Clubiona (Clubiona) sa/titans Emerton, 1919 Clubiona female genitalia can be stated to be lower Description. Male. TiA strongly sderotized, not little beyond epigastric furrow (except C. moesta). CO than that in the male genitalia. branched, little, thin or strongly attenuating closer to small, placed near hind edge of epigyne, fused at latter's The marmorata-group Yet, tr·aited generally, as accepted for all spiders, top. Bulb non-protruding. Conductor tegular, consider midline, rarely separated from, but situated very closet?, this statement ignores certain differences in the ably larger than E, weakly sclerotized, of a characteristic each other, as in C. quebecana and C. pygmaea. CT thm Diagnosis. The male differing both by the strongly Infrageneric groupings within the spider genus Clubiona 45 44· K.G. Mikhailov The similis-group small. Atrium tubular, SPT circular, its diameter not developed, flat, wide TiA and embolar conductor (Fig. or. almost parallel .to each other in the beginning (C. exceeding of epigynal width. 6). The female differing by CO widely separated from nnnor, C. pseudomznor), then curved ventrad in fore part 1/5 each other combined with CT almost straight and the of endegyne to return backwards parallel to each other . · Diagn~sis. The. male differing by the weakly convex, Clubiona (Clubiona) zilla Dllnitz 'et Strand, 1906. epigyne strongly sclerotized. . (except C. pseudominor), but quickly arching and di bifurcate TiA wtth the ventral branch more thin, narrow and Description. Male. TiA attaining ea. 0.4-0.5 of cym ver~ing laterad. Atrium bordering SPT, both simple, attenuating, combined with a thinE (Fig. 29) and the absence The obesa-group biallength. Conductor slightly protruding beyond cym ovmd. of a conductor. The female differing by CT long, strongly bium and covering latter's base in ventral view. E coiled Clubiona (Clubiona) comta C.L. Koch, 1839 curved in distal half, combined with the simple ovoid or Diagnosis. The male differing by the wide TiA usually around tegular. tip, not stretching proximally. Seminal Clubiona (Clubiona) decora Blackwall, 1859 tubular SPT not curved in the depth of the vulva (Fig. 22). convex ven~rally ~nd concave dorsally (Figs 32-33, 35- duct relatively short. Clubiona (Clubiona) diniensis Sirnon, 1878 Oescription. Male. TiA bifurcate, with the ventral 36). Sometlmes TIA turned around its axis or strongly Female (known only in C. marmorata). CO small. Clubiona (Clubiona) genevensis L. Koch, 1866 branch either thin (C. similis, C. neglecta, C. aducta, C. developed. E thin, usually supplied with a membraneous CTslightly arched. Atrium bordering SPT, circular. SPT Clubiona ( Clubiona) leucaspis Sirnon, 1932 pseudosimi!~ - Fig. 29) or attenuating (C. germanica, "co~du~tor" on tegula~ surface. Female differing by unipartite, cylindrical, slightly narrowed in middle part, Clubiona (Clubiona) minor Wunderlich, 1987 C. congenttlts). EB apparently fused with EP-apofhysis. cavity-l.Ike CO usually .situat~d at the posterior angles of stretching along epigynal midline. Epigyne strongly Clubiona (Clubiona) paral/e/a Hu et Li, 1988 E thin, arising from the middle or distal part o bulb the gemtal plate, combmed With unparallel CT (Figs 40- sclerotized in hind part, with two depressions in the Clubiona (Clubiona) pseudominor Wunderlich, 1987 directed prolaterodistad, then arched around or angularly 47). middle presumably destined for TiA fixation during Clubiona ( Clubiona) tenerifensis Wunderlich, 1991 across tegular tip, moving on proximad, arching on Description. Male. TiA see Diagnosis, it slightly copulation. Clubiona (Clubiona) vegeta Sirnon, 1878 ventral bulb surface (i_n the upper half in most species, torsate, especially so in the akagiensis-subgroup. TiA can Clubiona (Clubiona) marmorata L. Koch, 1866 Clubiona (Clubiona) wunderlichi Michailov, 1992 over all surface returnmg to EB in C. neglecta). b.e divid~d iQto two weak branches or strongly developed, Clubiona (Clubiona) hyrcanica Michailov, 1990 Clubiona ( Clubiona) zhangmuensis Hu et Li, 1988 Female. CO opening mostly at hind edge of epigyne, eith~r SI~ple (~: bakurovi) or strongly branched (C. sometimes in groove in epigynal hind part (C. geimanica, kurtlensts, C. mnae - Figs 32, 37). EP-apophysis of The maritima-group The genevensis-subgroup C. congentilis). CT wide, directed parallel forward or different shape, being close to EB (except C. obesa and slightly diverging (C. similis, C. aducta - Fig. 22), then C. mixta). ,E beginning from bulb prolaterally, arched coiling several times in middle part of vulva, converging Diagnosis. Male: EP with a large spur-like apophysis Diagnosis. The male differing by the weakly devel around its distal top (except the latericia-subgroup), toward midline and finally a little diverging. Atrium thin, obtuse apically in C. obesa and C. mixta. Conductor protruding beyond cymbium (Fig. 23). Female: CO slit oped and non-furcate TiA as well as by a dark spot of thick circular and ovoid, placed in front of SPT and then more like, CT sinuous, SPTae adjointed, hind edge of epigyne hairs on the pro lateral side of the cymbium (fig. 26). The lacking only in the latericia-subgroup. Seminal duct of ~at~rally. SPT ~imple, ovoid, s~me.times considerably less medium length. · strongly sclerotized (Fig. 17). female differing by the large genital groove characteris m size than atnum (C. germamca) or completely circular Description. Male. TiA with three branches, each not tically semi-circular or inverted cordiform (Fig. 20). Female. CO cavity-like, positioned near posterior lateral (C. pseudosimilis), rarely tubular (C. similis). Epigynal ~les o~ genital pl.ate, only in C. spiralis slit-like and situated longer than 1/3 of cymbiallength. EP-apophysis cover Description of both male and female coincides with surface smooth. ing EB. E long, arched around bulb tip, then stretching the group description including the subgroup diagnosis. m the middle of epigyne. CT from narrow to moderately wide, Clubiona (Clubiona) similis L. Koch, 1866 no~ parallel; if almo~t parallel, as in the sapporensis-subgroup proximad along a membranous "conductor". C. genevensis, C. decora, C. diniensis, C. /eucaspis, C. Clubiona (Clubiona) aducta Sirnon, 1932 Female. Main features see in Diagnosis. 'CT thin, minor, C. paral/ela, C. pseudominor, C. tenerifensis, C. vegeta, (FJg. 43), convergmg to the epigyne median line. Atrium Clubiona (Clubiona) congentilis Kulczyii.ski, 1913 circular or ovoid, bordering SPT. SPT either unipartite long, first diverged, then converged. C. wunderlichi, C. zhangmuensis. Clubiona (Clubiona) gennanica Thorell, 1871 Clubiona (Clubiona) maritima L.Koch, 1866 circular, ovoid, tubular, or tends to be bipartite (C. kurilensis, Clubiona(Clubiona) neg/ecta 0. Pickard-Carnbridge, 1862 C. praematura, especially the sapporensis-subgroup). Geni The lutescens-group Clubiona (Clubiona) pseudoneglecta Wunderlich, 1994 tal plate.wi?e and smooth, slightly wrinkled in C. kurilensis, The brevipes-group Clubiona (Clubiona) pseudosimilis Michailov, 1990 usually mctsed posteriorly. Diagnosis. The male differing by the bifurcate convex Clubiona (Clubiona) obesa Hentz, 1847 Diagnosis. Male differing by the characteristically TiA (Fig. 28) combined with the wide E and missing The japonicola-group Clubiona ( Clubiona) aciformis Zhang et Hu, 1991 bifurcate EP-apophysis (Fig. 24), female by CT diverging embolar conductor (Fig. 27). The female differing by the Clubiona (Clubiona) akagiensis Hayashi, 1985 from the median groove combined with the simple straight or almost straight CT curved at the end corn bined Clubiona (Clubiona) bakurovi Michailov, 1990 circular SPTae (Fig. 18). with the ovoid atrium and tubular, unipartite SPT curved Diagnosis. By the structure of the male genitalia, the group is close to the lutescens-group differing by the thin Clubiona (Clubiona) bryantae Gertsch, 1941 Description. Male. TiA flat, either subdivided at from the depth of the vulva to its surface (Fig. 21). Clubiona ( Clubiona) charitonovi Michailov, 1990 tip ,(C. brevipes) or attenuating and curved (C. Description. Male. TiA with two convex branch E or more thin TiA (Fig. 30); by the structure of the female genitalia it is close to the similis-group, distin C/ubiona (Clubiona) chechtsirica Michailov, 1995 alexeevi). EP-apophysis bifurcate, in lateral view es of an almost equal length (in C. pseudosaxatilis, guished by the circular SPTae (Fig. 38). Clubiona (Clubiona) chikunii Hayashi, 1986 directed (sub)rectangularly to bulb surface, not cover dorsal one a little larger). EP-apophysis fused with Clubiona (Clubiona) chippewa Gertsch, 1941 ing EB. E arched around tegular tip. Seminal duct EB, so latter considerably wide; only in C. fruteto Description. Male. TiA with two processes separated !rom ~heir ~ases. E~-apophy~is passes on directly into E, Clubiona ( Clubiona) corrugata B!lsenberg et Strand, 1906 relatively short. :um the .border bet~een them is visible. E beginning Clubiona (Clubiona) eskovi Michailov, 1995 Female (known only in C. brevipes). CO fused into m proximal or middle part of tegulum, directed m C. ;apomcola a httle covenng the latter's base. E in the upper half of the bulb, directed retrolaterad, then Clubiona (Clubiona) furcata Emerton, 1919 genital groove of a characteristic multiangular shape. CT prolaterodistad, then arched around tegular tip and Clubiona (Clubiona) fusoidea Zhang, 1993 diverged laterad, then curved forward. Atrium a little moving on retrolaterad along -alveolar wall, some prolaterodistad, exceeding forward the alveola's limits. In C. yasudai a membraneous "conductor" expressed as Clubiona (Clubiona) fuzhouensis Gong, 1985 bigger than SPT, bordering it. SPT unipartite, circular. times coiling at bulb tip; often with a thin crest. Clubiona (Clubiona) ikedai Ono, 1993 Clubiona (Clubiona) brevipes Blackwall, 1841 Membraneous tegular "conductor" expressed as a a tegular apophysis. Seminal duct relatively short. .F.emale: Cs> separated from .each other, very large, Clubiona (Clubiona) inaensis Hayashi, 1989 Clubiona (Ciubiona) a/exeeui Michailov, 1990 protector-like tegular apophysis ("P" in Fig. 28). Clubiona (Clubiona) irinae Michailov, 1991 Seminal duct long. positioned m hmd part of the gem tal plate. CT wide, first directed forward, then in the middle part of the vulva Clubiona (Clubiona) kimyongkii Paik, 1990 Female. W~ll-visible CO placed in hind part of The comta-group rectangularly laterad outward. Atrium circular, placed in Clubiona (Clubiona) kunashirensis Michailov, 1990 genital plate. CT wide, directed foreward and. a little Clubiona (Clubiona) kurenshikovi Michailov, 1995 laterad. SPT tubular, thin or thick, curved outwards from f~ont of SP!, closer to epigynal midline. SPT simple, Diagnosis. The male (Figs 25-28) differing by the circular, a httle larger than atrium. Clubiona ( Clubiona) kurilensis Bllsenberg et Strand, 1906 total or almost total fusion of EP-apophysis with E depth of vulva, thus looking as if bipartite. Atrium Clubiona (Clubiona) latericia Kulczynii.ski, 1926 circular, positioned in front of SPT, more median in C. Clubiona ( Clubiona) japonica/a Bllsenberg et Strand, 1906. itself combined with the embolar tip directed proxi Clubiona (Clubiona) yasudai Ono, 1991. ? Clubiona ( Clubiona) /ena Bllsenberg et Strand mad and reaching the base of the bulb. The female lutescens and C. golovatchi, more lateral in others. Atrio Clubiona (Clubiona) logunovi Michailov, 1990 (Figs 19-20) differing by the total or almost total spermathecal tube short and wide. Genital plate smooth, The zilla-group Clubiona (Clubiona) manshanensis M.Zhu et An, 1988 fusion of CO combined with CT bordering each other usually without groove (except C. riparia; poorly ex Clubiona (Clubiona) marusiki Michailov, 1990 and parallel in the beginning, then strongly curved. pressed in C. lutescens and C. saxatilis). Clubiona (Clubiona) maya Hayashi et Yoshida, ·1991 A considerable extent of CT corresponding to the Clubiona (Clubiona) lutescens Wcstring, 1851 Diagnosis. The male differing by the large branched Clubiona ( Clubiona) microsapporensis Michailov, 1990 length of the male E. Clubiona (Clubiona) alpico/a Kulczyii.ski, 1882 EP-apophysis being wider than bulb (Fig. 31). Female • Clubiona (Clubiona) mimula Charnberlin, 1928 Description. Male. TiA flat, curved, attenuating in C. Clubiona (Clubiona) andreinii Caporiacco, 1936 differing by the presence of a transverse furrow with slit Clubiona (Clubiona) mi:rta Ernerton, 1980 comta (Fig. 25), simple and short in the. genevensis Clubiona (Clubiona) frutetorum L. Koch, 1866 like CO at its corners (Fig. 39). Clubiona (Clubiona) nenilini Michailov, 1995 subgroup (Fig. 26)'. EP-apophysis well-visible only in C. Clubiona (Clubiona) golovatchi Michailov, 1990 Description. Male. TiA wide, short, thick. EP-apo Clubiona (Clubiona) paiki:Michailov, 1991 comta at EB. EB solid, sometimes overgrowing prolat Clubiona (Clubiona) hilaris Sirnon, 1878 physis large, with 2 branches in the upper third, the lower ? Clubiona (Clubiona) paralena l\1ichailov, 1995 erad, a little beyond cymbium (C. comta, C. genevensis). Clubiona (Clubiona) proszynskii Michailov, 1995 branch trifurcate. E moderately thick, arising prolateral Clubiona (Clubiona) phragmitoidesSchenkel, 1963 E long, curved around tegular tip and directed proximad. Clubiona (Clubiona) pseudosaxatilis Michailov, 1992 ly, then arched around the bulb top and ending at its Clubiona (Clubiona) praematura Ernerton,'1909 Seminal duct relatively short. Clubiona (Clubiona) riparia L. Koch, 1866 · lower third. Membraneous "conductor" present. Clubiona (Clubiona) rileyi Gertsch, 1941 Female. Genital groove either small (C. comta, Fig. Clubiona (Clubiona) saxatilis I,.. Koch, 1867 Female. CO slit-like. CT straight, converging to the Clubiona (Clubiona) sapporensis Hayashi, 1986 19) or large (C. genevensis etc. - Fig. 20). CT parallel Clubiona (Clubiona) terrestris Westring, 1851 middle of epigyne, more thin near atria. Atrium and SPT Clubiona (Clubiona) spiralis Ernerton, 1909 Infrageneric groupings within the spider genus Clubiona 46 K.G. Mikhailov 47 Clubiona (Clubiona) subborea./is Michailov, 1992 phragmitoides). CT narrow, directed laterad, then epigastric furrow, combined with CT being wide, con Clubiona (Clubiona) zyuzini Michailov, 1995 Clubiona (Clubiona) tsumsakii Hayashi, 1987 converging and becoming parallel to each other. SPT verging to the middle of the epigyne and then almost The hummeli-subgroup Clubiona (Clubiona) uenoi Orio, 1986 and atrium placed on sides of vulva or in its middle (C. parallel. Clubiona (Clubiona) wo!clwngsensis Paik, 1990 corrugata). SPT irregularly ovoid (C. corrugata), Description. Male. TiA large, carrying one external Diagnosis. Having a typical epigyne, the species of Clubiona (Clubiona) yoshidai Hayashi, 1989 tubular, either of medium length (C. kimyongkii, C. and two internal branches. EP-apophysis covt;ring EB. E this subgroup differ only by male characters: E long, EP kurenshikovi) or strongly elongate (C. phragmitoides). long, occupying 2/3 of bulb length. Membraneous apophysis with a double ridge, membraneous "conductor" "conductor" present. . . The latericia-subgroup Atrium circular, bordering SPT, larger than it (C. present (Fig. 59). kimyongkii, C. kurenshikovi) or smaller (C. corruga Female. CO rounded, with curved furrows in g·enital Description. Male. TiA with a cluster of branches, a plate flowing into them from lateral sides. Direction of Diagnosis. The male differing by the short E, not ta). By CT structure, the species of this group resemble little similar to the obesa-group (especially C. flexa). EP C. kurilensis. CT see in Diagnosis. SPT complex, curved, tubular, covering EB. wider than 2/3 of bulb, combined with TiA typical in the bordering circular atrium. 6 group. The female differing by the strong sclerotization C. corrugata, C. kimyongkii, C. kurenshikovi, C. phragmi Female. See Description of the group . toides Hind part of genital plate strongly sclerotized corre 'C. hwmneli, C. flexa, C. odesanensis of the inner structure (Fig. 40) and the SPTae ovoid or spondingly to the strong male TiA. tubular (C. kunashirensis - Fig. 42), positioned in the C. irinae fore part of the genitalia. The akagiensis-subgroup The propinqua-stibgtoup Description. Male. TiA bifurcate from base (C. latericia, C. bryantae) or with the ventral branch more Diagnosis. The male differing by the tip of TiA curved The pallidula-group Diagnosis. The male differing by the semicircular strongly developed (C. kunashirensis - Fig. 35). EP downwards, combined with E thin and attenuating, ventral branch of TiA directed v~ntrad, then after apophysis covering EB, sometimes small (C. kunashiren directed retrolaterad or retrolatero-apicad, as well as by Diagnosis. The male differing by the thick, sometimes curvature, dorsad (Fig. 58). Female is indistinguishable. sis, C. maya). E usually short, not longer than 1 I 2 of the considerable EP-apophysis. The female differing by weakly-divided TiA (Figs 53-54, 58) and short E; when Description. Male. EP-apophysis and "conductor" bulb width (2/3 in C. tsurusakii), straight, acute. CT semicircular (Fig. 45), directed backwards in C. E long, EP-apophysis carrying a double ridge (Fig. 59). lacking. E relatively short, not longer than 2/3 of bulb Conductor lacking (except C. tsurusakii). chikunii and C. ikedai, then laterad, foreward and to the The female differing by the SPTae distinctly bipartite, widt~, dagger-like, sometimes with an uneven edge (C. Female. CO consist of wide depressions (C. latericia midline of the epigyne. tubular, rarely ovoid (Fig. 48). propmqua). - Fig. 40, C. bryantae) or small grooves (C. tsurusakii, Description. Male. Curvature of TiA tip mostly Description. Male. TiA thick, with a cluster of short Female. See Description of the group. C. kunashirensis, C. wolchongsensis - Fig. 42), some expressed in C. akagiensis and C. chikunii. E directed branches depressed from inwards (C. hummeli, in C. C. propinqua, C. mayumiae, C. pseudogennanica times positioned beyond the epigastric furrow on a small retrolaterad in C. akagiensis, almost retrolaterad in C. pallidula there are two branches, but one of them is semi epigynal outgrowth (C. kunashirensis, weaker in C. chikunii, and retrolatero-apicad in C. ikedai, its length transparent, thin and poorly visible - Fig. 53), without The reclusa-group maya). CT ·sinuous, more rarely almost straight (C. not exceeding bulb width. EP-apophysis directed retro depressions (C. odesanensis), or with two distinct branches (C. phragmitis, C. zacharovi, the propinqua-subgroup kunashirensis). Mutual disposition of atrium and SPT laterad in C. akagiensis, retrolatero-apicad in C. chiku Diagnosis. The male differing by the characteristic obscure due to the absence of sufficient material. nii, and apicad in C. ikedai. - Fig. 58). EP with a thick apophysis carrying a tooth (C. pallidula C. ezoensis - Fig. SS). EP-apophysis harpoon-like ventral branch of TiA (Figs 60-61) and the C. latericia, C. bryantae, C. kunashirensis, C. maya, C. Female. CO small, rounded (C. chikunii, C. ikedai), presence of a tegular conductor named here protector tsurusakii, C. wo/chongsensis positioned in the middle (C. ikedai), hind (C. chikunii) covering EB (c. pallidula, the hummeli-subgroup). E short, thick, curved, in C. phragmitis, C. sopaikensis (Fig. 7). The female differing by a short diverging CT part of epigyne or at its hind edge (C. akagiensis), which and the considerable distance between atrium and SPT is inserted in two latter species. Atrium small in C. and the propinqua-subgroup more or less dagger-like, The sapporensis-subgroo.p directed retrolaterad and occupying 1/2-2/3 of bulb (Figs 49-50). chikunii and C. ikedai, in C. akagiensis its disposition Description. Male. TiA divided into 2-3 branches, remains obscure due to the absence of material available width, with a free top (Fig. 57); if E long, as in the Diagnosis. The male differing by the presence of one hummeli-subgroup, it is directed proximad at 1/3 with a harpoon-like ventral one, in C. bashkirica the (ventral) branc:;h of TiA turned typically for the total for study. SPT ovoid (C. akagiensis) or tubular. typical contour is visible only in lateral view. As a rule, C. akagiensis, C. chikunii, C. dkedai length of bulb and combined with a double ridge on EP group, combined with the short E, approximately equal apophysis (Fig. 59). "Conductor" largely lacking, being EP carrying an apophysis with 1-2 acute teeth at top or to the bulb ~idth (Fig. 34). The female differing by the on true conductor (lacking in C. bashkirica). E thin and The bakurovi-subgroup present only in the hummeli-subgroup. Seminal duct complex structure of SPT supplied with diverse out- long, sinuous. curved, EB covered by EP (except C. bashkirica). Tip growths (Fig. 43). · Female. CO ovoid, not deep, positioned laterally near of E positioned near tip of protector at the distal edge Description. Male. TiA see Diagnosis. EP-apophysis Diagnosis. The male differing by the strong indivisi hind edge of genital plate, rarely at the edge, as in C. of alveolum, sometimes protruding beyond i.ts limits. In covering EB partly or completely. E short, directed ble TiA, the large tooth on the long EP-apophysis, and by langei and C. zyuzini, or removed from the edge, as in C. C. bashkirica, protector similar to the corresponding retrolaterad, then proximad. Membraneous "conduCtor" the absence of a "conductor". The female (Fig. 46) ezoensis and C. mayumiae, directed laterad and back flat membraneous structure in the lutescem-group. present. differing by the strongly sclerotized ridge (septum) ward. CT arched laterad. SPT bipartite, first part small, Female. CO slit-like, vague, positioned at the hind Female. Ovoid CO positioned near hind edge of positioned along the midline in the hind part of the genital elongate, lateral to the second one, large, ovoid in plane, edge or in the hind part (C. bashkirica - Fig. SO) of genital plate. CT sometimes ·almost parallel, a little plate, combined with the depression in the plate serving but more often tubular and arched in lateral view, genital plate, rarely rounded or ovoid, situated in the converging. Tubular SPT with an outgrowth, strongly for fixation of the male palp during copulation. positioned closer to midline of epigyne (Fig. 48). Atrium middle part of genital plate (C. maracandica, C. rybini). curved from the depth of· vulva outward, so in a Description. Male. TiA strong, convex retrolaterally circular or tubular, by shape similar to the smaller part CT short, directed laterad. SPT consisting of two reni transverse plane it seems bipartite. Atrium larger than and concave prolaterally. E positioned in distal part of of SPT. Genital plate smooth, in C. komissarovi protrud form parts (a different shape in C. bashkirica). Atrium SPT, ovoid, positioned more laterally, as in the palli- bulb, curved behind tegulum. EP-apophysis large and ing a·little beyond epigastric furrow. removed from SPT, located near hind edge of genital plate dula-group. . · . long, covering EB, with a large tooth. Clubiona ( Clubiona) pallidula Clerck, 1757 (Figs 49-50). Atrio-spermathecal tube long, directed to C. sapporensis, C. charitonovi, C. eskovi, C. fuzhouensis, · Femalt;. CO positioned near midline of epigyne, Clubiona (Clubiona) ezoensis Hayashi, 1987 the median line of epigyne. Genital plate mostly wrin C. manshanensis, C. microsapp6rensis, C. nenilini divided by a strongly sclerotized longitudinal ridge (Fig. Clubiona (Clubiona) flexa Zhang et Chen, 1993 kled, usually covered by hairs. 46). CT directed laterad, then forward, widely separated Clubiona (Clubiona) haeinsensis Paik, 1990 Clubiona (Clubiona) reclusa O.Pickard-Cambridge, 1863 The. corrugata~subgroup from each other, almost parallel. SPT simple, 'arched, Clubiona (Clubiona) hummeli Schenkel, 1936 Clubiona (Clubiona) bashkirica Michailov, 1992 atrium bordering it. Clubiona (Clubiona) hwanghakensis Paik, 1990 Clubiona (Clubiona) californica Fox, 1938 Diagnosis. The male differing by the long simple TiA A strongly chitinized hind edge of epigyne as much Clubiona ( Clubiona) komissarovi Michailov, 1992 Clubiona (Clubiona) canadensis Emerton, 1890 sometimes reaching midlength of the cymbium, combined as two strongly sclerotized grooves near hind edge of Clubiona (Clubiona) kumadaomm Ono, 1993 Clubiona ( Clubiona) evoronensis Michailov, 1995 with the relatively shortE (Fig. 36). The female differing genital plate obviously serving as a support for huge Clubiona (Clubiona) langei Michailov, 1991 Clubiona (Clubiona) interjecta L.Koch, 1879 by the thin and long CT, small atria and SPTae (Fig. 44). male TiA during copulation. Inner membrane of female Clubiona (Clubiona) mandschurica Schenkel, 1953 Clubiona ( Clubiona) kulczynskii Lessert, 1905 Description. Male. TiAlong, convex retrolaterally, genitalia, almost transparentin most Clubiona species, Clubiona (Clubiona) mayumiae Ono, 1993 Clubiona (Clubiona) maracandica Kroneberg, 1875 concave pro]aterally, sometimes spiralling (C. kimyong also strongly sclerotized here. Epigyne projecting Clubiona (Clubiona) odesanensis Paik, 1990 Clubiona (Clubiona) norvegica Strand, 1900 kii). E short, lying free above tegulum in alveolum (C. beyond epigastric furrow, weakening the total con Clubiona (Clubiona) oligerae Michailov, 1995 Clubiona (Clubiona) pacifica Banks, 1896 kimyongkii), or curved around tegul urn (C. corrugata); struction. Clubiona (Clubiona) orientalis Michailov, 1995 Clubiona ( Clubiona) rosserae Locket et Millidge, 1953 in the last case membraneous conductor is weakly C. bakurovi Clubiona (Clubiona) phragmitis C.L.Koch, 1843 Clubiona (Clubiona) rybini Michailov, 1992 developed. In C. kimyongkii, tip of E curved proximad, C/ubiona (Clubiona) propinqua L.Koch, 1879 Clubiona (Clubiona) saurica Michailov, 1992 as in the japonica/a-group. EPcapophysis covering EB The irinae-subgroup Clubiona ( Clubiona) pseudogemwnica Schenkel, 1936 Clubiona ( Clubiona) stagnatilis Kulczynski, 1897 (C. kimyongkii) or not, carrying two teeth (C. corruga-. Clubiona (Clubiona) sopaikensis Paik, 1990 Clubiona (Clubiona) subsultans Thorell, 1875 ta). Seminal duct long (C. corrugata) or of medium Diagnosis. The male differing by ·the strong and Clubiona (Clubiona) tateyamensis Ono, 1989 length (C. kimyongkii). strongly subdivided TiA (Fig. 37) and long E. The Clubiona (Clubiona) venusta Paik, 1985 6The epigyne of C. flexa is very similar to C. wolchong Female. CO cavity-like, opening at hind edge of female differing by the strong sclerotization of the hind Clubiona (Clubiona) zacharovi Michailov, 1991 sensis in the latericia-subgroup of the obesa-group. genital plate or circular, positioned in its hind part (C. edge of the genital plate (Fig. 47) not hanging over the ©ARTHROPODA SELECfA, 1995 48 K.G. Mikhailov Arthropoda Selecta 4 (2): 49-71 The caerulescens-group Kraus, 0. 1984. Male spider genitalia: evolutio11ary changes in Remarks on the Ural spider fauna, 5. New records of spider species of st.ructure and function I I Verh. naturwiss. Ver. Hamburg. Diagnosis. The male differing by the strong develop N.F. Bd.27. S.373-382. the family Linyphiidae from the Urals. (Arachnida Aranei) ment of divided TiA and the free E without any conductor Lange, A. B. 1969. [Order Spiders (Aranei)] I I Zhizn zhivot nykh. Moscow, Prosveshchenie Pub!. Vo).3. P.56-88 [in and EP~apophysis (Fig. 62). The.female differing by the Russian]. · · ·. epigyne hanging far over the epigastric furrow and Lohmander, H. 1945. Vorlllufige Spinnennotizen I I Ark. Zoo!. 3a.MeTIUt no <}>ayHe nayKoB Ypa.Aa,. 5. Ho:BI>Ie HaXOAKH nayRon ce.M.eUCTB~ strongly sclerotized at its hind edge (Fig. 51). Bd.35A. Nr.16. P.1-21. . Description. Male. TiA strong, with three branches, Lyubarsky, G.Y. 1991. [Objectivation of the taxonomical rank Linyphiidae Ha Ypa.Ae (Arachnida Aranei) the middle one bifurcate at top. E beginning in proximal category] I I Zhurn. obshch. bioi. Vol. 52. No.5. P.613-616 half of bulb, very long, directed distad, then arched [in Russian, with English summary]. around tegulum and moving on!roximo-laterodistad. Mayr, E. 1963. Animal Species and Evolution. Cambridge, Mass.: S.L. Esyunin*, V.E. Efi.mik*, A.B. Polyanin** Female. CT directed forwar , being parallel to each Belknap Press of Harvard Univ. Press. 797 pp. other at midway. SPT simple, bordering an ovoid atrium. Mayr, E. 1969. Principles of Systematic Zoology. New York etc.: C.A. EcroHuH*, B. E. E<}>H.Mu.K*, A.B. lloMIHu.H** Ca. 2/5 of epigyne length located beyond epigastric McGraw-Hill Book Company. 428 pp. furrow. Meyen, S.V. 1978. [Main aspects of organism typology] I I Sclerotization of the hind part of the genital plate Zhurn. obshch. bioi. Vol.39. No.4. P.495-508 [in Russian, * Department of Zoology, Perm State University, Bukireva Street 15, Perm 614600 Russia. seems to correspond to the large size of the male TiA. with English summary]. *Ka<}leApa 300AOrl111 6ecrro3BOHO'lHbiX >KHBOTHbiX, DepMCKHH rocyAapcTBeHHbiH yHunepcuTeT, YA· ByKupena 15, DepMb 614600 PoccWI. Clubiona (Clubiona) caernlescens L.Koch, 1866 Mikhailov, K.G. 1990. The spider genus Clubiona Latreille 1804 in the Soviet Far East, 1 (Arachnida, Aranei, Clubionidae) - ** Department of Dog-breeding, Perm Higher Military School, Perm 108, Russia. The chabarovi-group Korean Arachnology I I Vol.5. No.2. P.139-175. ** Ka<}leApa KHHOAOrl1l1, DepMcKoe BbiCIIIee noeHHOe KOMaHAHO-TbiAonoe )"l11Al1llje MBA Poccul1, DepMD 108, Poccl1JI. Mikhailov, K.G. 1991. The spider 'genus Gubiona Latreille 1~04 in the Soviet Far East, 2 (Arachnida, Aranei, Clubionidae) I Diagnosis. The male differing by TiA strong but I Korean Arachnology. Vol.6. No.2. P.207-235. KEY WORDS: spiders, fauna, faunistics, Urals. weakly sclerotized (Fig. 63), the female by the weakly Mikhailov, K.G. 1992a. [Systematics and fauna of the spider sclerotized epigyne hanging over the epigastric furrow, genus Clubiona Latreille, 1804 (Aranei, Clubionidae) of KJIIQqEBbiE CJIOBA: rray1