Coelotes Blackwall, 1841: 618

WANG, XIN-PING, 2002, A Generic-Level Revision Of The Spider Subfamily Coelotinae (Araneae, Amaurobiidae), Bulletin of the American Museum of Natural History 2002 (269), pp. 1-150 : 40-47

Coelotes Blackwall, 1841: 618 . – Roewer, 1954:

54; – Levi and Kraus, 1964: 150; – Lehtinen,

1967: 223; – Brignoli, 1983: 470; – Platnick,

1989: 421; – Platnick, 1993: 569; – Platnick,

1997: 662.

TYPE SPECIES: Clubiona saxatilis Blackwall, 1833 , by original designation, which is a junior synonym of Drassus atropos Walckenaer, 1830 . Male neotype and female paraneotypes of C. atropos designated by de Blauwe (1973), deposited in Collection W. Kulczyn’ski, l’Institut de Zoologie P.A.N. de Varsovie, Poland, not examined.

DIAGNOSIS: Females resemble Eurocoelotes but have lateral epigynal teeth, a small atrium, and short copulatory ducts (figs. 38, 39, 101, 102, 113, 114, 127, 128). Males also resemble Eurocoelotes in having a conductor dorsal apophysis and a short, rounded median apophysis, but have a broad patellar apophysis (figs. 45, 107, 116, 130).

DESCRIPTION: See description of type species in Morphology section (above).

DISTRIBUTION: Europe, East Asia (map 4).

COMPOSITION: Four species groups are recognized within Coelotes : the atropos group, exitialis group, pseudoterrestris group, and charitonovi group, in total about 38 species.

In addition, 85 coelotine species are not well studied and temporarily remain in Coelotes (see appendix).

est, elevation 1675 m, March 31, 1998, 1 male (Hai­Yin Wu, THU, THU­Ar­990026) ; Nantou County, Huei­Sun Experimental Forest , elevation 1550 m, March 31, 1998, 1 female (Hai­Yin Wu, THU, THU­Ar­990027) .

DISTRIBUTION: China (Taiwan).

The atropos Group

DIAGNOSIS: Females have a reduced atrium, reduced copulatory ducts, and a short, less convoluted spermathecae (figs. 38, 39); males generally have a broad patellar apophysis, a small lateral tibial apophysis, a short conductor, and a broad conductor dorsal apophysis (figs. 44, 45).

This species group is well represented in Europe.

There are at least 19 species in the group. Some species, such as C. atropos ( Walckenaer, 1830) and C. pastor Simon, 1875 , show remarkable intraspecific variation, particularly in female genitalia.

1. Coelotes alpinus Polenec, 1972 : male holotype and female paratypes from Alps, Europe, in Prinodoelovni Muzej Slovenije, Ljubljana, Slovenia, not examined. Male and female specimens from Italy, in MZF and MCV, examined.

2. Coelotes atropos ( Walckenaer, 1830) : male neotype and female paraneotype from Europe, in the Collection W. Kulczyn’ski l’Institut de Zoologie P.A.N. de Varsovie, Po­

land, not examined. Male and female specimens from England, France, Denmark, Poland, Switzerland, Russia, in CAS, AMNH, MCZ, NHMB, HEC, and COLL.WALTER, examined.

3. Coelotes atropos anomalus Hull, 1955 : female described from England, type not indicated and depository unknown, not examined. Judged by the illustration, this species may be a junior synonym of C. atropos . Further examination of possible type (s) is need­ ed.

4. Coelotes atropus silvestris Hull, 1955 : female described from England, type not indicated and depository unknown, not examined. Judged by the illustration, this species may be a junior synonym of C. atropos . Further examination of possible type (s) is need­ ed.

5. Coelotes italicus Kritscher, 1956: 1 male paratype from Italy, in MCV, examined. Males and females from Italy, in MCV, MCB, and AMNH, examined.

6. Coelotes mediocris Kulczyn’ski, 1887: type (s), depository unknown, not examined. Males and females from Italy, in NHMB, MCV, MCB, AMNH, IRSNB, COLL.MAURER, examined.

7. Coelotes osellai de Blauwe, 1973 : male holotype, male and female paratypes from Italy, in MCV, examined.

8. Coelotes pabulator Simon, 1875 : male lectotype and female paralectotype from France, in SMNH, examined.

9. Coelotes pastor Simon, 1875 : male lectotype and female paralectotype from France, in MNHN, not examined. Males and females from France and Italy, in MNHN, MCZ, MCV, and AMNH­CU examined.

10. Coelotes pastor carpathensis Ovtchinnikov, 1999 : male holotype, male and female paratypes from Middle Asia, in CSO, not examined.

11. Coelotes pastor cooremani de Blauwe, 1975 : male and female paratypes of C. cooremani de Blauwe from Italy, in MCV, examined. Maurer (1982) reduced the species C. cooremani de Blauwe, 1975 to this subspecies. Male types are C. rudolfi .

12. Coelotes pastor pirinicus Drenski, 1942 : types unknown, not examined.

13. Coelotes pastor pickardi O.P.­ Cambridge, 1873: types unknown, not examined. The species C. pickardi O.P.­ Cambridge, 1873 was reduced to this subspecies by Maurer (1982). Males and females from Italy and Switzerland, in NHMB, COLL.MAURER, HEC, MCB, and AMNH examined.

14. Coelotes pastor tirolensis ( Kulczyn’ski, 1906): types, unknown, not examined. Males and females from Switzerland, Italy, Austria, in NHMB, COLL.MAURER, MCB, and MCV, examined.

The species Coelotes pastor lessinensis Maurer, 1982 , with female holotype and male paratype from Italy, in MCV, examined, is placed as a junior synonym of C. p. tirolensis (Kulczyn’ski) (because of the identical male and female genitalic morphology). NEW SYN­ ONYMY.

15. Coelotes poleneci Wiehle, 1964 : male holotype from Slovenia, in SMF, examined.

16. Coelotes poweri Simon, 1875 : female lectotype and male paralectotype from Italy, in MNHN, examined.

17. Coelotes rudolfi ( Schenkel, 1925) : male and female types from Switzerland, in NHMB, not examined. Male and female specimens from Switzerland, in AMNH and COLL.MAURER, examined.

18. Coelotes solitarius L. Koch, 1868 : male neotype and female paraneotypes from Europe, in Collection W. Kulczyn’ski l’Institut de Zoologie P.A.N. de Varsovie, Poland, not examined. Females from Austria, Italy, Germany, in MCB, MZF, ZMB, and HEC, examined.

19. Coelotes terrestris (Wider, 1834) : male neotype and female paraneotype from Europe, in Collection W. Kulczyn’ski l’Institut de Zoologie P.A.N. de Varsovie, Poland, not examined. Male and female specimens from Europe, in AMNH, COLL.MAURER, NHMB, HEC, MCZ, AMNH­CU, and ZMB, examined.

Coelotes atropos (Walckenaer) Figures 5–54

Drassus atropos Walckenaer, 1830: 170 (male neotype and female paraneotype from Europe, in the Collection W. Kulczyn’ski l’Institut de Zoologie P.A.N. de Varsovie, Poland, not examined). – Walckenaer, 1837: 627.

Drassus trucidator Walckenaer, 1830 (types unknown): 172. – Walckenaer, 1837: 629.

Clubiona saxatilis Blackwall, 1833 (types unknown): 436. – Blackwall, 1834: 332; – Blackwall, 1841: 618, figs. 6–8; – Walckenaer, 1847: 441; – Blackwall, 1861: 169, fig. 109.

Clubiona tigrina Walckenaer, 1842 (types unknown): 483.

Amaurobius atropos: Simon, 1864: 139 . – Kulczyn’ski, 1908: 435, figs. 15, 28, 48, 66; – Locket and Millidge, 1953: 19, figs. 14A, 14B; 15A, 15C; – Miller, 1971: 175, pl. XXVIII, figs. 20–22, pl. XXIX, fig. 18.

Amaurobius saxatilis: Simon, 1864: 139 .

Amaurobius trucidator: Simon, 1864: 139 .

Coelotes atropos: Simon, 1875: 32 , fig. 14; – Hansen, 1882: 42, fig. 2; – Becker, 1896: 187, fig. 2; – Chyzer and Kulczyn’ski, 1897: 160, fig. 15; – Bösenberg, 1902: 222, fig. 314. – Dahl, 1931: 22, figs. 34–41; – Reimoser, 1931: 40, fig. 2; – Simon, 1937: 985, 990, 1030, figs. 1511, 1520; – Drensky, 1942: 41, figs. 5c, 6c– Hull, 1955: 51, fig. 2e; – Wiehle, 1963: 289, figs. 107, 110, 112, 117–119; – Lehtinen, 1967: 448, fig. 239; – Loksa, 1969: 108, figs. 73B, 73C, 74B, 75E; – Tyschehenko, 1971: 160, figs. 448, 452; – de Blauwe, 1973: 27, figs. 24–28; – Locket et al., 1974: 43, fig. 23C; – Roberts, 1985: 164, fig. 73a; – Steinberger, 1989: 607, fig. 3; – Heimer and Nentwig, 1991: 356, fig. 927; – Roberts, 1995: 249; – Buchar et al., 1995: 120, figs. 35.12, 35.13.

Coelotes saxatilis: Chrysanthus, 1965: 67 , figs. 1– 3, 8–9.

DIAGNOSIS: This species is similar to C. terrestris but can be distinguished by the different shape of the spermathecae of females (fig. 39) and by the relatively large, rough conductor dorsal apophysis of males (fig. 44).

DESCRIPTION: See descriptions in Morphology section (above) and also by de Blauwe (1973).

MATERIAL EXAMINED: DENMARK: 1 male and 1 female ( CAS). ENGLAND: Oxford Shire , 1 female ; Lake District , 1 male ( AMNH) ; Oxfordshire, Oxford , July 23, 1958, 1 female (H. and L. Levi, MCZ) ; Settle, West Riding Yorkshire, April , 1948, 3 females (B. Percy, AMNH) ; Yorkshire, July , 1948, 1 male (B. Malkin, AMNH) ; West Riding Yorkshire, April , 1948, 2 females (B. Percy, AMNH) ; Yorkshire , 2100 ft, July 9, 1948, 5 females (B. Alkin, AMNH) ; Wales, Lllangoed Castle, Breconshire , July 5, 1959, 3 females (B. Malkin, AMNH). FRANCE: 1 male and 1 female ( MCZ) ; 1 female ( MCZ) ; Paris, 1 male (J. H. Emerton, MCZ). POLAND: Babia Gora , mountain top, 1700 m, 1 female (B. Malkin and A. Kamocki, AMNH) ; Babia Gora , 1450 m, July 25, 1975, 1 male (B. Malkin and A. Kamocki, AMNH) ; Bialewieza, Narewka River , August 26, 1959, 4 males and 1 female (B. Malkin, AMNH) ; Bialewieza , Natl. Pk., September 23–26, 1982, 2 males and 5 females (B. Malkin, AMNH) ; Roztocze Natl. Pk., Bukowa Gora, June 20, 1987, 1 female (B. and H. Malkin, AMNH). RUSSIA: Biolower , 1 female ( Ed. Reimoser , MCZ). SWITZERLAND: Zinal , Val d’Anniviers , unter Steinwall , 1690 m, July 19, 1979, 2 females (J. Walter, COLL. WALTER) ; Mase, Val d’Herens , Fichten­Larchen­Wald, 1900 m, July 16, 1980, 1 male (J. Walter, COLL. WALTER) ; Marchairuz , 1 male and 1 female ( NHMB, 610a) ; 1 male and 1 female ( HEC, B.437, t.45) .

The exitialis Group

DIAGNOSIS: Females have lateral atrial margins, broad copulatory ducts, and long, convoluted spermathecae (figs. 101–104); males generally have a well­developed lateral tibial apophysis, a long conductor, and a slender conductor dorsal apophysis (figs. 107–109).

This species group is well represented in Japan.

Nine species are here included in the group.

1. Coelotes antri (Komatsu, 1961) : female holotype from Japan, depository unknown, not examined. Male and female from Shikoku, Japan, in AMNH, CAS, and SMF, examined.

2. Coelotes eharai Arita, 1976 : male holotype, male and female paratypes from Mt. Daisen, Tottori, Japan, in the Ohtemon­Gak­ uin University and in the Tottori Museum, Japan, not examined. A female from Shikoku, Japan, in AMNH, examined.

3. Coelotes exitialis L. Koch, 1878 : male and female types from Japan, depository unknown, not examined. Male and female from Japan, in AMNH, CAS, and MCZ, examined.

4. Coelotes hiratsukai Arita, 1976 : male holotype, male and female paratypes from Funaoka­cho, Tottori, Japan, in the Ohtemon­Gakuin University and in the Tottori Tottori Museum, Japan, not examined. One female from Shikoku, Japan, in AMNH, examined.

5. Coelotes kitazawai Yaginuma, 1972 : female holotype from Fuji­san, Japan, depository unknown, not examined.

6. Coelotes micado Strand, 1907 : types from Japan, depository unknown, not examined.

7. Coelotes musashiensis Nishikawa, 1989 : female holotype, male and female paratypes from Tokyo, Japan, in the Arachnological Society of Japan and in NSMT, not examined. Two females from Japan, in ZMB, examined.

8. Coelotes yaginumai Nishikawa, 1972 : female holotype and male paratype from Ibaraki, Osaka, Japan, in the Arachnological Society of Japan, not examined.

9. Coelotes yodoensis Nishikawa, 1977 : male holotype, male and female paratypes from Minoo, Osaka, Japan, in the Arachnological Society of Japan and in the Osaka Natural History Museum, Japan, not examined.

Coelotes exitialis L. Koch

Figures 101–112

Coelotes exitialis L. Koch 1878: 755 , figs.17–19

(types from Japan, depository unknown, not ex­

amined). – Uyemura, 1936: 10, figs. unnum­ bered; – Yaginuma, 1960: 93, figs. 82.1, 82.2;

– Yaginuma, 1971: 93, figs. 82.1, 82.2; – Ya­

ginuma, 1973: 25, figs. 5, 6; – Nishikawa, 1974:

175, figs. 1, 2; – Nishikawa, 1975: 173, figs. 2–

51; – Paik, 1978: 342, figs. 152.1, 152.2; – Ya­

ginuma, 1986:147, fig. 79; – Chikuni, 1989:

100, fig. 11.

DIAGNOSIS: Females of this species can be recognized by the broad epigynal teeth and the anteriorly situated copulatory ducts (figs. 101, 102), and males by the broad, slightly bifid patellar apophysis and the long, slender, slightly spiral conductor (figs. 107–109).

DESCRIPTION: See also description of Koch (1878). Total length 8.00–15.0. From front, anterior eye row slightly procurved, posterior row procurved; eye sizes and arrangements: AME smallest, ALE largest, PLE slightly smaller than ALE, PME subequal to PLE; AME­AME, AME­ALE, and ALE­PLE sub­ equal or AME­ALE slightly longer, PLE­ PME wider than PLE diameter, PME­PME slightly wider than AME­AME. Clypeal height slightly more than twice AME diameter; chilum divided, elongated (fig. 105). Chelicerae with three promarginal and four retromarginal teeth (fig. 106). Labium slightly longer than wide. Length of female 1st leg patella + tibia shorter than carapace length.

Tarsal organ situated close to distal end of tarsus, slightly anterior of distalmost trichobothrium. Only female spinnerets examined: apex of ALS with 2 major ampullate gland spigots (MAP), 45–50 piriform gland spigots; PMS with about 47 aciniform gland spigots, 2 cylindrical gland spigots (CY), 1 or 2 minor ampullate gland spigots (mAP); PLS with 56 aciniform gland spigots and 2 cylindrical gland spigots (figs. 110–112).

Female epigynum with epigynal teeth broad, may vary in shape and position ( Nishikawa, 1974); atrium reduced, with more or less apparent lateral atrial margins; copulatory ducts broad; spermathecal heads situated anteriorly; spermathecae elongated, convoluted (figs. 101–104). Male palp with patellar apophysis large, slightly bifid, may vary within species ( Nishikawa, 1974); RTA long, with distal end sharply extended beyond tibia; lateral tibial apophysis present; cymbial furrow short; conductor long, more or less spiral, slender, its shape may vary within species ( Nishikawa, 1974); conductor dorsal apophysis long; conductor lamella moderately developed; embolus basal origin, moderately long; median apophysis spoonlike, rounded, with distal end smooth (figs. 107–109).

MATERIAL EXAMINED: JAPAN: Kobu ( Kobe ?), February 1909, 1 female (together with a male P. luctuosus ) (S. C. Thompson, CAS) ; Shizuoka Pref., Izu Penninsula , 100– 400 m, August 10. 1962, 1 male (K. Sekiguchi and A. Ogura, MCZ) ; Fuji Mt., SE slope of Mt. Fuji , February 18, 1955, 2300 m, floor mat., 1 female (D. C. Lowrie, AMNH) .

DISTRIBUTION: Japan.

Figures

Fig. 5. Coelotes atropos (Walckenaer), female body, dorsal view.

Figs. 6–11. Coelotes atropos (Walckenaer), female. 6. Eyes and chelicerae, front view. 7. Chelicera, lateral view. 8. Chelicera, ventral view. 9. Labium, endite, and palp, ventral view. 10. Palp, dorsal view. 11. Sternum.

Figs. 12–19. Coelotes atropos (Walckenaer), female. 12. Chelicera, ventral view. 13. Cheliceral glands. 14. Cheliceral gland, close view. 15. Inner, distal long setae of chelicera. 16. Proximal short setae of chelicera. 17. Promargin setae of chelicera. 18. Median depression of cheliceral fang. 19. Apex of cheliceral fang.

Figs. 20–27. Coelotes atropos (Walckenaer), female, legs. 20. First leg femur, right. 21–24. First to fourth legs, left. 25, 26. Fourth leg, showing trichobothria distribution. 27. Fourth leg tibia, showing trichobothria distribution.

Figs. 28–30. Coelotes atropos (Walckenaer). 28. Tarsus, showing position of tarsal organ (double arrows) and the distalmost trichobothrium (single arrow). 29. Trichobothrium. 30. Tarsal organ.

Fig. 31. Coelotes atropos (Walckenaer). Trachea and spinnerets.

Figs. 32–34. Coelotes atropos (Walckenaer), female spinnerets. 32. ALS, left. 33. PMS, right. 34. PLS, right. Abbreviations are spelled out on p. 6.

Figs. 101, 102. Coelotes exitialis Koch, female. 101. Epigynum. 102. Vulva.

Figs. 103–107. Coelotes exitialis Koch. 103. Female, epigynum. 104. Female, vulva. 105. Carapace, front view. 106. Chelicera, ventral view. 107. Male palp, retrolateral view.

Figs. 108, 109. Coelotes exitialis Koch, male palp. 108. Ventral view. 109. Retrolateral view, showing conductor dorsal apophysis.

Figs. 110–112. Coelotes exitialis Koch, female spinnerets, ventral view, right. 110. ALS. 111. PMS. 112. PLS. Abbreviations are spelled out on p. 6.