Table Of Content2006. The Journal of Arachnology 34:87-97
A REVIEW OF THE LINYPHIID SPIDER GENUS
SOLENYSA (ARANEAE, LINYPHIIDAE)
Lihong Tu and Shuqiang Li^: Institute of Zoology, Chinese Academy of Sciences,
Beijing 100080, China. E-mail: [email protected]
ABSTRACT. The present paper gives a review ofthe Solenysa spiders. Five ofthe six known Solenysa
species were examined, including the types ofS. longqiensis, S. wulingensis and S. circularis. Illustrations
ofthese five species as well as diagnoses and distributional data of all species are provided.
Keywords; Taxonomy, Asia, China, Korea, Japan, spiders, ant mimicry
The linyphiid spider genus Solenysa was tions were made using a drawing tube and
established by Simon (1894) for the sole spe- inked on inkjet plotter paper. Male palps and
cies Solenysa melloteei Simon 1894 from Ja- female epigyna were examined and illustrated
pan. Solenysa remained as a monotypic genus after they were dissected from the spider’s
until five additional species were described bodies. Vulvae of female epigyna were
from China and Korea during the 1990s, in- cleared in boiling KOH solution to dissolve
cluding S. longqiensis Li & Song 1992, S. non-chitinous tissue, and the embolic divi-
wulingensis Li & Song 1992, S. circularis sions of male palps were excised by breaking
Gao et al. 1993, S. protrudens Gao et al. 1993 the column (the membranous connection be-
and S. geumoensis Seo 1996. tween the suprategulum and the radix). For
Members of the genus appear to be ant examination of the genital structures under
mimics and have the posterior part ofthe car- transmitted light microscopy, male palps and
apace drawn into a tubular extension reminis- epigyna were immersed in 75% alcohol so-
cent of the constricted waist of ants. Further, lution, while embolic divisions and vulvae
the epigynum of the females is connected to were mounted in Hoyer’s Solution,
the abdomen by a long, transparent tube or For each species, the synonyms are taken
solenoid. A similar structure is found in some verbatim from Platnick’s spider catalogue
other linyphiids such as Wubanoides Eskov (Platnick 2003). Updated information about
1986 from the Palaearctic region (Tanasevitch the distribution of each species in China is
1996: figs. 7-9) and Metalepthyphantes Lock- provided at the provincial level. The names of
et 1968 from Africa (Locket 1968: figs. 27, localities and distribution data are given ac-
33). cording to current Chinese standard (see Peng
This paper reviews the genus Solenysa. et al. 2003).
Type specimens of S. longqiensis, S. wulin- All measurements are in millimeters. Ter-
gensis, S. circularis and fresh specimens ofS. minology for the somatic morphology and
protrudens and S. melloteei have been exam- genital structures follow Hormiga (2002). The
ined and illustrated. Further diagnoses and abbreviations used as follows:
distributional data ofall six known species are Somatic morphology: AER = anterior eye
provided. row; ALE = anterior lateral eye; AME = an-
METHODS terior median eye; AME-ALE = distance be-
tween AME and ALE; AME-AME = distance
Specimens were examined and measured between AMEs; AMEd = diameter of AME;
using an SZ11-Olympus stereomicroscope. PER = posterior eye row; PLE = posterior
Further details were studied under an Olym- lateral eye; PME = posterior median eye;
pus BX40 compound microscope. All illustra- PMEd = diameter of PME; PME-PLE = dis-
tance between PME and PLE; PME-PME =
' Corresponding author. distance between PMEs.
87
88 THE JOURNAL OF ARACHNOLOGY
—
Map 1. Distribution ofSolenysa spiders. Circles = S. circularis Gao etal.; arrowhead = S. geumoensis
Seo; square = S. longqiensis Li & Song; triangles = S. melloteei Simon; star = S. protrudens Gao et al.;
and ellipse = S. wuUngensis Li & Song.
Male palp: DSA = distal suprategular developed, divided in two or three branches.
apophysis; E — embolus; EM embolic Epigyne protruding well sclerotizedconnected
membrane; LC lamella characteristica; P = to the abdomen by a long, transversally wrin-
paracybium; PCA = proximal cymbial apoph- kled, membranous solenoid base, which is
ysis; R == radix; SPT = suprategulum; T = packed between the epigynum and abdomen
tegulum; TA = terminal apophysis; TS = ter- in the nonfuncti—onal stage (Fig. 9).
minal sclerite. Description. Small spiders, total length
Epigynum: CD = copulatory duct; CO = of males 1.11-1.61, females 1.27-1.70. Both
copulatory opening; ED = fertilization duct; sexes similar in general appearance with little
S = spermatheca; SL — solenoid. interspecific variation. Caparace chestnut-
The specimens studied here are deposited brown, with numerous impressed, round pits;
in the Institute of Zoology, Chinese Academy fine radial striae little darker in color. Cara-
of Sciences in Beijing (IZCAS), and in Jilin pace in dorsal view elongated oval, slightly
University, Changchun, China (JLU, formerly constricted at level of cervical groove; sides
called Norman Bethune University ofMedical behind cervical groove distinctly crenate, with
Science). four lobes beside coxae; posterior part of car-
TAXONOMY apace drawn into tubular extension (Fig. 22);
AMEs
cephalic region turret-like bearing in
Family Linyphiidae Blackwall 1859 front, PMEs at top, ALEs and PLEs on its
Solenysa Simon 1894 lateral sides (Fig. 1). AMEs black and very
small, their diameter about halfofothers’, rest
Solenysa Simon 1894: 677. AER AME-AME
of eyes subequal; recurved,
—
Type species. Solenysa melloteei Simon almost equal to AMEd and AME-ALE longer,
1894, by mon—otypy. PER almost straight, PME-PME equal to
Diagnosis. Solenysa species can be dis- PMEd and PME-PLE shorter, ALE and PLE
tinguished from other linyphiids by their very juxtaposed. Clypeus broad, concave immedi-
small body size (total length 1.11-1.70) and ately bellow ocular area, sloping in convex
their unique body appearance; especially by line towards frontal margin. Chelicerae slight-
the tubular extension of the posterior end of ly lighter in color than carapace, with some
the carapace and by the many impressed, round pits frontally and some granules antero-
round pits scattered all over the carapace laterally; promargin with 4 and retromargin
(Figs. 1, 2). Cymbium usually with one ortwo with 2-4 teeth; lateral sides with distinct strid-
large proximal apophyses (Fig. 4). Paracym- ulating ridges. Sternum darker in color than
bium L-shaped. Lamella characteristica well carapace, roundish heart-shaped, with many
XU & LI—REVIEW OF SOLENYSA 89
—
Figures 1-11. Solenysa circularis Gao et al. 1. carapace, lateral view; 2. carapace, dorsal view; 3.
right chelicera, anterior view; 4. left male palp, retrolateral view; 5. left male palp, prolateral view; 6. left
male palpal cymbium, dorsal view; 7. right palpal tibia, dorsal view; 8. left male palpal embolus division,
=
dorsal view; 9. epigynum, ventral view; 10. epigyiium, lateral view; 11. vulva, dorsal view. Scale bars
0.1 mm.
90 THE JOURNAL OF ARACHNOLOGY
—
granules, each carrying long hair. Legs long Material examined. CHINA: Zhejiang:
and slender; tibia I-IV and patella I-IV with Mt. Tianmushan (30.4°N, 119.5°E), female
a dorsal spine shorter than diameter of tibia. holotype and 1 male paratype (JLU); Mt. Pu-
Tm I 0.17-0.26. Tm IV absent. tuoshan (30.0°N, 122,4°E), 1 female paratype
Male pedipalp: Tibia with one prolateral (JLU). —
and two retrolateral trichobothria (Fig. 17). Diagnosis. The male of S. circularis is
Cymbium normally with one or two large, easily recognized by the cone-shaped, poste-
proximal apophyses, outstanding in retrolater^ riorly directed cymbial apophysis (Fig. 4) and
al view (Fig. 4), except in S. wulingensis and by the conspicuous, curved spike-like branch
S. geumoensis (Fig. 43). Paracymbium small, of lamella characteristica in prolateral view
L“Shaped, furnished with several long apical (Figs. 5, 8). The epigynum ofthe female (Fig.
hairs, deeply curved and with torsion at its 9) is similar to that ofS. protrudens (Fig. 38),
middle. Tegulum triangular in retrolateral but is more rounded and distinctly bulging lat-
view (Fig. 4). Distal suprategular apophysis erally. —
large and well sclerotized. Shape of embolic Description, Total length 1.27. Carapace
membrane variable. Lamella characteristica 0.73 long, 0.44 wide. Abdomen 0.57 long,
conspicuously large, normally with two or 0.43 wide. Tm I 0.26. Tm IV absent. Chelic-
three branches, at least two of them well de-- erae with 4 promargieal and 2 retromarginal
veloped (Fig. 8): the first sword-shaped or teeth (Fig. 3). For further measurements and
spike-like, strongly sclerotized, the second a detailed description of somatic morphology
ribbon-like and the third variable in shape or see Gao, Zhu & Sha (1993).
even missing (Fig. 18). Terminal apophysis Male palp (Figs. 4~8): Lamella character-
strongly sclerotized, usually with 1-3 teeth. istica tripartite (Fig. 8); the first branch big-
'Terminal sclerite’ located in menbranous re- gest, curved spike-like, well sclerotized, sec-
gion between radix and lamella, continuous ond one short and blunt, and third one
with base oflamella, varying in shape and ap- smallest, falciform. Terminal apophysis with
pearance. two small apical processes at its dorsal side:
Epigynum: Protruding and well sclerotized, one truncate, one coniform. Terminal sclerite
variable in shape, connected with abdomen by large, foliaceous, bipartite. Embolus and em-
long, transversally amply wrinkled, membra- bolic membrane shortest among the five spe-
nous solenoid base, which in nonfunctional cies described in this paper, even shorter than
stage is folded between epigynum and abdo- terminal apophysis. Anterior part of radix
men. — drawn into handle-like extension.
Remarks. The members of the genus So- Epigynum (Figs. 9-11): Wider than long,
lenysa seem to be ant mimics and have the sides rounded, posterodorsal part in lateral
posterior part of their carapace drawn into a view turned anteriorly; solenoid base partly
tubular extension resembling the waist ofants. visible (Fig. 9), 3 times as long as the length
Further, the epigynum of the females is con- of epigynum, twisting anteriorly and connect-
nected to the abdomen by a long solenoid ed with its dorsa—lly side.
base. Similar structures are found e.g. in Wub- Distribution. Known only from the orig-
anoides Eskov 1986 (Tanasevitch 1996: figs. inal localities in Zhejiang Province, China
7-9) and Metalepthyphantes Locket 1968
(Map
& 1).
(Locket 1968: figs. 27 33). However, So-
lenysa can be easily distinguished from all Solenysa geumoensis Seo 1996
other linyphiid genera by its unique appear- Solenysa melloteei Namkung 1986: 13, figs. 6-10
ance as described—above. (misidentification).
Distribution. China, Korea and Japan Solenysa geumoensis Seo, 1996: 157; Namkueg
(Map 1). Specimens can be found amongst 2002: 182, figs. 17.35a-b.
—
grass, leaf-litter and other detritus, but are Material examined, None.
very rare in museum collections. Diagnosis.—Judging from the illustrations
Solenysa circularis Gao et al. 1993 by Namkung (1986, 2002), the male ofS. geu-
Figs. 1-11 moensis is close to that of S. wulingensis as
Solenysa circularis Gao et al. 1993: 66, figs. 7-10; both have cymbium without outstanding prox-
Song et al. 1999: 204, figs. 1 16H-I, N-O. imal apophysis (Namkung 1986, fig. 9) and in
TU & LI—REVIEW OF SOLENYSA 91
prolateral view basically similar lamella char- apophysis with twisted bases in dorsal view
acteristica (Namkimg 1986, fig. 8). They dif- (Fig. 17). Embolic division (Fig. 18) conspic-
fer in that one branch (exactly which one is uously large. Lamella characteristica with two
not clear) of the lamella characteristica of S. well-developed, ribbon-like branches, apex of
geumoensis has a forked tip (Namkung 1986, first one with a strongly sclerotized claw, sec-
fig. 7) while S. wulingensis has all branches ond one with truncate, somewhat serrate end.
entire (Fig. 46), Furthermore, the apical por- Embolus slightly longer than lamella charac-
tion of the paracymbium of S. geumoensis is teristica. Embolic membrane petale-shaped.
longer than that of S. wulingensis (Namkung Terminal apophysis fist-like. Radix quadran-
1986, fig. 7). Epigynumhalfround shape, pos- gular.
terior part wider than anterior part, the trans- Epigynum (Figs. 19-20): Apple-shaped,
parent vulval structures convergent anteriorly widest close to its anterior edge, about 1.5
(Namkung 1986, fig. 10). times wider than long. Solenoid base about
Description,—See Namkung (1986, 2002). three times as long as the length ofepigynum,
Distribution.—Korea (Map 1). covered with triangularextension ofabdomen.
Remarks,— geumoensis is Distribution.^—Known only from the type
thought to be written by Seo in 1996, but the locality in Fujian Province, China (Map 1).
original description on this species is not easy
to obtain. A further study on this species will Solenysa melloteei Simon 1894
be necessary in future, in case that we can get Figs. 21^30
the types or fresh material. Solenysa mellottei [sic] Simon 1894: 677; Oi, 1960:
153, figs. 52^54.
Solenysa longqiensis Li & Song 1992 Solenysa mellotteei [sic] Simon: Yaginuma 1986:
Figs. 12-20 78, fig. 42.2; Me & Saito 1987: 23, fig. 21; Chi-
Solenysa longqiensis Li & Song, 1992: 6, figs. lA- kuni 1989: 56, fig. 48.
—
G; Song et al. 1993: 861, figs. 17A-G; Li et al. Material examieed* JAPAN: no detailed
1994: 80, figs. 18, 19; Song etal. 1999: 204, figs. data, 1 male —and 1 female (IZCAS).
116J, K, Q, R.
Diagnosis.' ^The male of this species can
Material examined.—CHINA: Fujian: Ji- be distinguished from all otherSolenysa males
angle County (26.7°N, 117.4°E), Mt. Longqi, by the broad, coiled embolus and the hook-
Yujiapieg Town, 10 August 1991, male ho- like cymbial apophysis (Fig. 24). Epigynum
lotype, 1 male and 6 female paratypes (IZ- (Fig. 28) somewhat apple-shaped, like that of
CAS). — S. longqiensis (Fig. 19), but apically narrower
Diagnosis. The male of this species can and transparent vulval structures of different
be distinguishedfrom all otherSolenysa males shape. Solenoid base without cover.
by the bipartite lamella characteristica (Fig. Description.—Total length 1.27. Carapace
18) and the peculiarly twisted cymbial apoph- 0.77 long, 0.45 wide. Abdomen 0.50 long,
ysis (Figs. 15-17). The female has a some- 0.43 wide. Tm I 0.17. Tm IV absent. Chelic-
what apple-shaped epigynum (Fig. 19), like erae with 4 promargirial and 2 retromarginal
that of S. melloteei (Fig. 29), but distinctly teeth (Fig. 23). For further measurements and
broader apically and the transparent vulval a detailed description of somatic morphology
structures are different. Further the solenoid see Oi (1960).
base of S. longqiensis is twisted under a tri- Male palp (Figs. 24—27): Tibia twice as
angular cover while that ofS. melloteei is ex- long as patella (Fig. 24), with a dorsal apoph-
posed. ysis bearing two long bristles (Fig. 26). Cym-
Description.—Total length 1.47, Carapace bium with a forward pointing, hook-like prox-
0.80 long, 0.50 wide. Abdomen 0.76 long, imal apophysis (Fig. 24). Lamella tripartite,
0.45 wide. Tm I 0.22. Tm IV absent. Chelic- the first branch short, spike-like and well
erae with 4 promarginal and 3 retromarginal sclerotized; second one ribbon-like, long and
teeth (Fig. 14). For further measurements and slender, tapering off distally; third one small
a detailed description of somatic morphology and thin (Fig. 27). Embolus plate-like, half
see Li & Song (1992), spiral with a hooked tip (Fig. 24). Embolic
Malepalp (Figs. 15-18): Tibia slightly lon- membrane indiscernible. Terminal apophysis
ger than patella. Two proximal cymbial truncate, with a sickle-shaped dorsal process
92 THE JOURNAL OF ARACHNOLOGY
—
Figures 12-20. Solenysa longqiensis Li & Song: 12. carapace, lateral view; 13. carapace, dorsal view;
14. left chelicera, anterior view; 15. left male palp, retrolateral view; 16. left male palp, prolateral view;
17. left male pedipalp, dorsal view; 18. left male pedipalp embolus division, dorsal view; 19. epigyeum,
ventral view; 20. vulva, dorsal view. Scale bars = 0.1 mm.
TU & LI—REVIEW OF SOLENYSA 93
—
Figures 21—30. Solenysa melloteei Simon: 21. carapace, lateral view; 22. carapace, dorsal view; 23.
left chelicera, anterior view; 24. left male palp, retrolateral view; 25. left male palp, prolateral view; 26.
left palpal tibia, dorsal view; 27. left male pedipalp embolus division, dorsal view; 28. epigynum, ventral
view; 29. epigynum, lateral view, lifted; 30. epigynum, lateral view, expanded. Scale bars = 0.1 mm.
94 THE JOURNAL OF ARACHNOLOGY
(Figs. 24, 27). Terminal sclerite triangular, Distribution.—Known from China (Jiang-
blunt tipped. Radix short, wide and bipartite. su, Taiwan) (Map 1).
Epigynum (Figs. 28-30): Apple-shaped, &
Solenysa wulingensis Li Song 1992
widest at its middle, about 1.8 times wider Figs. 40-46
than long. Solenoid base exposed, short, about
twice as long as the length of epigynum, and Solenysa wulingensis Li & Song 1992: 7, figs, 2A-
can lift epigynum inversely in nonfunctional E; Song & Li 1997: 404, figs. 6A-E; Song et al.
1999: 204, figs. 117A, B.
stage. —
Distribution, Apparently restricted to Ja- Material examined.—-CHINA: Hunan:
pan (Map 1). Zhang]iajie National Nature Reserve (29.1°N,
llCr-'l°E), Dayong City, 13 August 1990, male
Solenysa protrudens Gao et al. 1993 holotype and 2 male paratypes (IZCAS).
Figs. 31-^39
Diagnosis,—The male of this species can
Solenysa protudens Gao et al. 1993: 65, figs. 1—6; be easily distinguished from all otherSolenysa
Song et al. 1999: 204, figs. 116L, M, P, S. males by the rudimentary cymbial apophysis
Material examined.-—CHINA: Taiwan: (Fig. 45). The female is unknown.
Description.—Total length 1.40. Carapace
Taidong (22.7°N, 121.1°E), Lanyu, August 0.80 long, 0.50 wide. Abdomen 0.67 long,
2000-February 2001, 3 males and 2 females 0.45 wide. Tm I 0.19. Tm IV absent. Chelic-
(IZCAS). The type material was deposited in
erae with 4 promargieal and 3 retromarginal
JLU but could not be found despite thorough
teeth (Fig. 42). For further measurements and
checking of the collection by the authors.
a detailed description of somatic morphology
Diagnosis.—The male of this species can see Li & Song (1992).
be distinguished from all otherSolenysa males Malepalp (Figs. 43^6): Tibia slightly lon-
by the stout, erect cymbial apophysis (Fig. ger than patella. Cymbial apophysis small,
34), petal-shaped embolic membrane and nearly inconspicuous (Figs. 43, 45), Paracym-
well-developed branches of lamella character- bium strongly curved at its middle, proximal
istica (Figs. 34, 35). Epigynum similar to that end with small lateral curvature. Lamella
of S. circularis, but more angular with nearly characteristica tripartite (Fig. 46), first branch
straight sides (F—ig. 38). spike-like, well sclerotized, second one larg-
Description. Total length 1,67. Carapace est, ribbon-like, tapering offdistally, third one
0.83 long, 0.50 wide. Abdomen 0.90 long, small, thin. Embolus (Fig. 46) short, embolic
0.50 wide. Tm I 0.22. Tm IV absent, Chelic- membrane flower-shaped. Terminal apophysis
erae with 4 promarginal and 4 retromarginal with two apical and one median process. Ter-
teeth (Fig. 33). For further measurements and minal sclerite foliaceous. Radix elongated,
a detailed description of somatic morphology blunt tipped.
see Gao et al. (1993). Female: Unknown.
Malepalp (Figs. 34-37): Tibia nearly twice Distribution.—Known only from the type
as long as patella, with small dorsal process locality in Hunan Province, China (Map 1).
bearing two long bristles (Fig. 34). Cymbium
ACKNOWLEDGMENTS
with a stout and erect proximal apophysis,
pointing dorsally. Lamella characteristica tri- We are grateful to Dr. Michael L Saaristo
partite (Fig, 37), the first and third branches (University of Turku, Finland) and Dr. Xiep-
sword-shaped, robust, welLsclerotized, the ing Wang (Illinois Natural History Survey,
latter longer than the former; the second rib- USA) for their support during our study on
bon-like, with denticles on its distal end. Em- the Chinese linyphiid spiders. The present
bolus slightly longer than lamella character- study was supported by the National Natural
istica. Embolic membrane petale-shaped, with Sciences Foundation of China (NSFC-
two processes. Terminal apophysis small, tri- 30270183, 30370263), by the National Sci-
angular (Fig. 37). ence Fund for Fostering Talents in Basic Re-
Epigynum (Figs. 38, 39): Rectangular, search (NSFC-J0030092), and was also partly
about three times as wide as long. Solenoid supported by the Kadoorie Farm and Botanic
base about three times as long as the length Garden, Hong Kong Special Administrative
of epigynum. Region, China.
TU & LI—REVIEW OF SOLENYSA 95
—
Figures 31-39. Solenysa protrudens: 31. carapace, lateral view; 32. carapace, dorsal view; 33. left
chelicera, frontal view; 34. left male palp, retrolateral view; 35. left male palp, prolateral view; 36. left
male palp, dorsal view; 37. left male palp embolus division, dorsal view; 38. epigyne, ventral view; 39.
vulva, dorsal view. Scale bars = 0.1 mm.
96 THE JOURNAL OF ARACHNOLOGY
—
Figures 40-46. Solenysa wulingensis Li & Song: 40, carapace, lateral view; 41, carapace, dorsalview;
4452.. lleefftt cmhaelleicepraal,p,andtoerrsiaolrvviieeww;; 4463.. lleefftt mmaallee ppaeldpi,parleptroelmatbeorlalusvideiwv;isi4o4n.,ldefotrsmaalleviepawl.p,Scparloelatbearrasl=vie0w.1;
mm.