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First record of Africactenus Hyatt, 1954 and redescriptions of two poorly known species of Ctenus Walckenaer, 1805 (Araneae, Ctenidae, Cteninae) from India

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The ctenid genus Africactenus Hyatt, 1954 is recorded for the first time from India. Africactenus unumus sp. nov. is described and illustrated based on male specimens. Detailed redescriptions and illustrations of Ctenus cochinensis Gravely, 1931 (both male and female) and Ctenus indicus Gravely, 1931 (only female) are provided and their distribution in India is updated.
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Accepted by A. Santos: 18 Dec. 2017; published: 5 Mar. 2018
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2018 Magnolia Press
Zootaxa 4388 (3): 395
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http://www.mapress.com/j/zt/
Article
395
https://doi.org/10.11646/zootaxa.4388.3.5
http://zoobank.org/urn:lsid:zoobank.org:pub:9A3DC9D3-0367-4A68-8B75-E96A1D78EE3F
First record of Africactenus Hyatt, 1954 and redescriptions of two poorly known
species of Ctenus Walckenaer, 1805 (Araneae, Ctenidae, Cteninae) from India
PRADEEP M. SANKARAN
1
& POTHALIL A. SEBASTIAN
Division of Arachnology, Department of Zoology, Sacred Heart College, Thevara, Cochin, Kerala 682 013, India.
1
Corresponding author. E-mail: pradeepmspala@rediffmail.com
Abstract
The ctenid genus Africactenus Hyatt, 1954 is recorded for the first time from India. Africactenus unumus sp. nov. is de-
scribed and illustrated based on male specimens. Detailed redescriptions and illustrations of Ctenus cochinensis Gravely,
1931 (both male and female) and Ctenus indicus Gravely, 1931 (only female) are provided and their distribution in India
is updated.
Keywords: distribution, first record, new species, redescription, taxonomy
Introduction
Ctenidae Keyserling, 1877 represents nocturnal and cursorial spiders, frequently found in the tropical forests of
South and Central America, Asia and Africa (Polotow & Jocqué 2015). They can be readily distinguished from the
remaining spider families by their 2-4-2 ocular arrangement (Silva-Dávila 2003). Currently the family has 514
nominal species distributed in 47 genera (World Spider Catalog 2017), but ctenid spiders from India are poorly
investigated. Simon (1897) described the first Indian ctenid spider, the female of Anahita smythiesi (Simon, 1897),
which was originally described under Ctenus Walckenaer, 1805. After Simon, the family received little attention
until Gravely (1931) described four new Ctenus species and recorded the genus Acantheis Thorell, 1891 for the
first time from India. Tikader & Malhotra (1981) revised the Indian Ctenus and recently Jäger (2012) recorded the
genera Anahita Karsch, 1879 and Amauropelma Raven, Stumkat & Gray, 2001 for the first time from India. To
date, only 17 species belonging to four genera have been recorded from India: Acantheis (one species),
Amauropelma (three species), Anahita (one species) and Ctenus (12 species) (World Spider Catalog 2017). In this
paper, we record the African ctenid genus Africactenus Hyatt, 1954 for the first time from India and redescribe two
Ctenus species described by Gravely (1931).
Material and methods
The specimens were studied under a Zeiss Stemi 2000-C stereomicroscope. All measurements are in millimetres
(mm). Lengths of pedipalp/palp and leg segments are given as: total (femur, patella, tibia, metatarsus [except for
pedipalp/palp], tarsus). Drawings were made with the aid of a drawing tube attached to the microscope. The images
of preserved specimens were taken with a Leica DFC295 digital camera attached to a Leica M205C
stereomicroscope with the software package Leica Application Suite (LAS, version 4.3.0) for stacking images
taken at different focal planes. Field photos were taken with Canon EOS 6D with Canon 100 mm Macro photo
lens. The specimens are deposited in a reference collection housed at the Division of Arachnology, Department of
Zoology, Sacred Heart College, Thevara, Cochin, Kerala, India (ADSH).
Abbreviations used in the text: ALE—anterior lateral eye, AME—anterior median eye, do—dorsal, MA—
median apophysis, pl—prolateral, pld—prolateral-dorsal, PLE—posterior lateral eye, plv—prolateral-ventral,
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PME—posterior median eye, rl—retrolateral, rld—retrolateral-dorsal, rlv—retrolateral-ventral, RTA—retrolateral
tibial apophysis, vt—ventral-terminal, ZSI—Zoological Survey of India, Kolkata, I–IV—1
st
to 4
th
leg.
Taxonomy
Family Ctenidae Keyserling, 1877
Subfamily Cteninae Simon, 1897
Genus Africactenus Hyatt, 1954
Diagnosis. Carapace with deep dorsal indentation (Hyatt 1954: fig. 2; herein Fig. 2A), clypeus high (Steyn et al.
2003: fig. 3; herein Fig. 2B), cheliceral retromargin with four teeth, tibiae I and II usually with six prolateral-
ventral and five retrolateral-ventral spines or rarely six on each row on tibiae I and II, RTA with superior and
inferior denticles (Hyatt 1954: figs 3–4, 6–8; Steyn et al. 2003: figs 7–8, 12–13, 17–18; herein Fig. 3B) and
epigynal plate with M-shaped anterior half (Hyatt 1954: plate XXXII, figs 26–33; Steyn et al. 2003: figs 9, 14, 19).
Type species. Africactenus agilior (Pocock, 1900), by subsequent designation (Hyatt 1954).
Remarks. The genus was erected by Hyatt (1954) for 11 ctenid spiders from Africa as a member of the
subfamily Acantheinae Simon, 1897 and was later transferred to Cteninae Simon, 1897 (Polotow & Brescovit
2014). To date, the genus has 20 nominal species, three are known only from males, eight only from females,
whereas nine are known from both sexes (World Spider Catalog 2017). The genus is currently restricted to Africa
(World Spider Catalog 2017).
Africactenus unumus sp. nov.
Figs 1A, 2A–B, 3, 4
Type material. Holotype, male missing right leg I (ADSH203A) from Areeppara in Edappady, 9
o
42'35.62''N,
76
o
42'48.42''E, Kottayam, Kerala, India, 27 m alt., 28 July 2013, M.S. Pradeep leg., from ground, by hand.
Paratype, male (ADSH203B), same data as holotype, except 20 August 2014.
Additional material examined. INDIA, Kerala: Pathanamthitta, Thiruvalla, Kavumbhagom, 9
o
24'15.60''N,
76
o
32'45.36''E, 3 m alt.,1 male (ADSH203C), Sam Thomas leg., 19 June 2013, from ground, by hand.
Diagnosis. Males of Africactenus unumus sp. nov. seem closely similar to the males of Africactenus decorosus
(Arts, 1912) as both share the wide, transversely oriented apical part of embolus as well as the same position and
orientation of the conductor, but can be separated from the latter by the following combination of characters: RTA
with single inferior denticle (RTA of A. decorosus with three inferior denticles), broad conductor (A. decorosus
with thin conductor), MA lying away from embolus (MA of A. decorosus lying close to embolus), embolus with
embolic apophysis (embolus of A. decorosus lacks embolic apophysis) and embolus narrowing at tip (A. decorosus
with uniformly wide embolus) (Fig. 3A–C, compare with Steyn et al. 2003: figs 7–8).
Description. Male (holotype, Figs 1A, 2A–B). Carapace with deep median indentation (Fig. 2A), brownish,
with faint, blackish, reticulated pattern and mats of greyish hairs along sides, with broad, median, pale brown,
roughly flower vase-like patch, its posterior part just in front of fovea abruptly narrowing towards pedicel, its
apical part with a longitudinal white stripe, which bifurcates posteriorly to continue along lateral margins (Fig. 1A).
Fovea long (1.39), longitudinal, black. Eye field, clypeus, chelicerae, labium, endites, sternum, spinnerets, and legs
brownish. Clypeus very high (Fig. 2B). Chelicerae dorsally with greyish-brown stiff hairs; promargin with a thick
layer of long, brownish, inwardly curved hairs, with one teeth and two denticles, retromargin with four teeth and
two small denticles. Fangs black. Sternum clothed with black hairs. Opisthosoma elongated oval, olive-brown,
with irregularly scattered black spots and striae; dorsum medially with a broad longitudinal, straw-coloured patch,
anterior 2/3
rd
of which medially possesses a pale yellowish band; dorsum with scattered clusters of white, stiff hairs
(Fig. 1A); venter medially with a broad yellowish patch, the middle of which bears a longitudinal olive-brown
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FIGURE 1. Habitus of Indian ctenid spiders. A. Africactenus unumus sp. nov.: male (holotype), dorsal. B, D–E. Ctenus
cochinensis Gravely, 1931: B. Female (live), dorsal (ADSH20196); D. Male (ADSH20195), dorsal; E. Female (ADSH20196),
dorsal. C, F. Ctenus indicus Gravely, 1931: C. Female (live), fronto-dorsal (ADSH201037); F. Same, dorsal. Scale bars: A, D–
F 2 mm. Photo credit B & C Jimmy Paul.
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FIGURE 2. Africactenus unumus sp. nov. (A–B) and Ctenus indicus Gravely, 1931 (C–E). A. Male prosoma, lateral; B. Same,
frontal. C. Left femur IV retrolateral view showing humps; D–E. Close up of humps on femur IV. Scale bars: A, C 2 mm; B 1
mm; D–E 0.2 mm.
stripe. Scopulae on tarsi entire; scopulae on metatarsi I–II entire, 2/3
rd
of metatarsus III with reduced scopula,
metatarsus IV lacks scopula. Body length 14.99. Carapace length 7.98, width 6.71. Opisthosoma length 7.01, width
3.98. Eye diameters: ALE 0.24. AME 0.36. PLE 0.53. PME 0.44. Eye interdistances: ALE–ALE 1.46. ALE–PME
0.19. AME–ALE 0.30. AME–AME 0.25. PLE–PLE 1.72. PME–PME 0.36. Clypeus height at AMEs 0.71, at ALEs
1.29. Chelicerae length 3.79. Measurements of pedipalp and legs: pedipalp 12.97 [5.05, 1.85, 2.87, 3.20], I 51.44
[13.82, 4.17, 15.39, 14.25, 3.81], II 45.77 [12.38, 4.11, 13.19, 12.65, 3.44], III 36.71 [10.22, 3.22, 10.12, 10.41,
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INDIAN CTENID SPIDERS
2.74], IV 53.19 [14.83, 3.34, 13.47, 18.03, 3.52]. Leg formula: 4123. Spination of pedipalp: femur pld 1 do 4 rld 1,
patella pl 1 rl 1, tibia pl 1 plv 1 rl 1 rld 1, tarsus/cymbium spineless; legs: femur I pl 2 pld 2 plv 1 do 1 rld 4, II pl 1
pld 3 do 3 rld 4, III–IV pld 4 do 3 rld 4; patellae I–IV pl 1 rl 1; tibiae I–II pl 1 plv 6 do 4 rl 2 rlv 6, III pl 2 plv 3 do
3 rl 1 rld 1 rlv 3, IV pl 2 plv 3 do 4 rl 2 rlv 3; metatarsus I pl 1 plv 2 rl 2 rlv 3, II pl 1 plv 3 rl 2 rlv 3, III pl 1 pld 3
plv 3 rl 1 rld 3 rlv 3, IV pl 1 pld 4 plv 4 rl 2 rld 3 rlv 4; tarsi I–IV spineless. Pedipalp (Fig. 3A–C): palpal segments
brownish with faint black annulations on femur, patella and tibia. RTA short with broad base and one tiny, sharp
inferior denticle (Fig. 3B, arrow). Ventro-lateral tibial apophysis short, thumb-like, directed at 2-o’clock in ventral
view (Fig. 3B). Distal part of cymbium slender, cylindriform; cymbium with a short baso-retrolateral, lamella-like
outgrowth (Fig. 3B–C). Tegulum nearly oval. Subtegulum small. MA nearly globular, medio-retrolateral (Fig. 3B).
Embolus long, with well sclerotised median part over membranous part, with wide, transversely oriented apical
part, with hook-like tip, with a short, flat apophysis (Fig. 3A–B). Conductor broad, hyaline, apico-retrolateral,
anterior part slightly overlying the embolar tip (Fig. 3B–C).
Female: Unknown.
Etymology. The specific epithet, from the Latin unum = one, is an adjective and refers to the single inferior
denticle of the RTA of the new species.
Va ri at io n . Males (n=3): body length 10.87–14.99.
Distribution. India (Kerala) (Fig. 4).
FIGURE 3. Africactenus unumus sp. nov. A–C Male left pedipalp: A. Prolateral; B. Ventral; C. Retrolateral. Abbreviations: C
conductor, Cl lamella-like outgrowth of cymbium, E embolus, EA embolic apophysis, EB embolic base, MA median
apophysis, RTA retrolateral tibial apophysis, ST subtegulum, T tegulum, VTA ventral tibial apophysis. Scale bars 2 mm. Arrow
indicate inferior denticle of RTA.
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FIGURE 4. Geographic distribution records of Africactenus unumus sp. nov. Red circle, type locality.
Genus Ctenus Walckenaer, 1805
Ctenus cochinensis Gravely, 1931
Figs 1B, D–E, 5, 6
Ctenus cochinensis Gravely, 1931: 235, figs 1E, 3C (description and illustrations of male and female). Tikader & Malhotra
1981: 112, fig. 5A–D (description and illustrations of male and female).
Type material. Male and 2 female syntypes of Ctenus cochinensis Gravely, 1931 from INDIA: Kerala: Palakkad:
Parambikulam (10
o
23'34.29''N, 76
o
46'32.24''E), 597 m alt. (male) and Thrissur: Kavalai (10
o
16'23.89''N,
76
o
32'13.03''E), 390 m alt. (females); F.H. Gravely leg.; 16–24 September 1914; repository ZSI; voucher numbers
1506/18 (male) and 1550/18 (females), not examined.
Note. The species was identified using the illustrations given by Gravely (1931) and Tikader & Malhotra (1981).
Material examined. INDIA, Kerala: Thrissur, Chimmony Wildlife Sanctuary, 10
o
25'6.80''N, 76
o
32'7.25''E,
709 m alt., 8 May 2013, M.S. Pradeep leg., from ground, by hand: 1 subadult male, 2 subadult females,
(ADSH20191) and 23 July 2013: 1 female with egg sac (ADSH20192); Ernakulam, Kothamangalam, Salim Ali
Bird Sanctuary (Thattekkadu Bird Sanctuary), 10
o
7'48.3''N, 76
o
41'43.24''E, 96 m alt., 22 May 2013, M.S. Pradeep
leg., from ground, by hand: 1 subadult male, 1 female, (ADSH20193); Malappuram, Akambadam, 11
o
18'37.42''N,
76
o
12'31.68''E, 41 m alt., 23 August 2013, M.S. Pradeep leg., from ground, by hand: 2 females with egg sacs
(ADSH20194); Kottayam, Areeppara in Edappady, 9
o
42'35.62''N, 76
o
42'48.42''E, 27 m alt., 16 August 2016, M.S.
Pradeep leg., from ground, by hand: 1 male (ADSH20195) and 25 March 2017: 1 female (ADSH20196).
Diagnosis. Ctenus cochinensis can be separated from other congeners by the following combination of
characters: hook-shaped RTA without apical bifurcation, MA short, roughly squarish, embolus with well-defined,
transversely oriented apical part, flat, as wide as long epigynal plate, posteriorly placed epigynal teeth, nearly
straight lateral folds of copulatory ducts and peanut-like spermathecae (Fig. 5A–F).
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INDIAN CTENID SPIDERS
FIGURE 5. Ctenus cochinensis Gravely, 1931. A–C. Male left pedipalp: A. prolateral; B. ventral; C. retrolateral. D–G. Female
genitalia: D. Epigynum, ventral; E. Internal genitalia, dorsal; F. Epigynum, posterior; G. Epigynum of female from Salim Ali
Bird Sanctuary, illustrating variation. Abbreviations: C conductor, CD copulatory duct, E embolus, EB embolic base, EF
epigynal field, EP epigynal plate, ET epigynal tooth, FD fertilization duct, LF lateral fold of copulatory duct, MA median
apophysis, RCT retrolateral cymbial tooth, RTA retrolateral tibial apophysis, S spermatheca, ST subtegulum, T tegulum, Tmp
membranous pit of tegulum, VTA ventral tibial apophysis. Scale bars: A–C, 1 mm; D–G, 0.5 mm. Arrows pointing to
copulatory opening. Digits 1 and 2, spermathecal chambers.
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Redescription. Male (based on ADSH20195, Fig. 1D). Carapace pale brownish with yellowish-brown median
band; thoracic part provided with narrow, longitudinal yellowish-brown medio-lateral bands and patches. Fovea
long (1.41), longitudinal, reddish. Eye field black except between PLEs, which is yellowish-brown; ALEs, PLEs
and PMEs basally covered with fine, short white hairs (Fig. 1D). Clypeus, chelicerae, fangs, endites, labium,
sternum, and spinnerets pale brownish. Dorsum of chelicerae covered with long brownish hairs; promargin with a
thick layer of inwardly curved brownish hairs, with three teeth; retromargin with four. Sternum covered with black
hairs. Opisthosoma roughly oval, hirsute; dorsum, laterals, and venter greyish-brown; venter with a broad, inverted
triangular, creamy patch. Leg segments pale brownish to brownish with black shades and annulations; all tarsi
scopulated, tarsal scopula entire; metatarsus I with entire scopula, distal 2/3
rd
of metatarsus II scopulated, metatarsi
III and IV lack scopula. Body length 10.71. Carapace length 6.39, width 5.44. Opisthosoma length 4.32, width
2.91. Eye diameters: ALE 0.19. AME 0.32. PLE 0.39. PME 0.45. Eye interdistances: ALE–ALE 0.25. ALE–PME
0.08. AME–ALE 0.16. AME–AME 0.18. PLE–PLE 1.52. PME–PME 0.22. Clypeus height at AMEs 0.16, at ALEs
0.67. Chelicerae length 2.06. Measurements of pedipalp and legs: pedipalp 8.42 [3.19, 1.20, 1.28, 2.75], I 23.26
[6.21, 2.81, 6.28, 6.23, 1.73], II 20.12 [5.65, 2.61, 5.02, 5.34, 1.50], III 16.70 [4.93, 2.14, 3.62, 4.65, 1.36], IV
23.89 [6.40, 2.40, 5.37, 7.97, 1.75]. Leg formula: 4123. Spination of pedipalp: femur pld 1 do 4 rld 1, patella pl 1,
tibia pl 1 plv 1 rld 1 rlv 1, tarsus/cymbium spineless; legs: femur I pl 1 pld 2 do 3 rld 4, II pl 1 pld 3 do 3 rld 4, III
pl 5 do 3 rl 4, IV pld 4 do 3, rld 3; patellae I–IV pl 1 rl 1; tibia I pl 1 plv 5 do 3 rld 2 rlv 5, II pl 1 plv 5 do 3 rl 2 rlv
5, III pl 2 plv 3 do 2 rl 2 rlv 3, IV pl 2 plv 4 do 3 rl 2 rlv 3; metatarsus I pld 2 plv 3 rld 3 rlv 3, II pld 3 plv 3 rld 3 rlv
3 vt 1, III pl 1 pld 3 plv 2 rl 1 rld 3 rlv 3, IV pl 3 pld 1 plv 3 rl 2 rld 3 rlv 4; tarsi I–IV spineless. Pedipalp (Fig. 5A–
C): palpal segments brownish with black shades. RTA stout, with broad proximal part, short, distal part with acute
tip, appears hook-like in retrolateral view (Fig. 5B–C). Ventro-lateral tibial apophysis short, thumb-like, directed at
2-o’clock in ventral view (Fig. 5B). Cymbium tip conical, without basal extension, with short, acute, retrolateral
cymbial tooth (Fig. 5B–C). Tegulum highly sclerotised, with a median membranous pit (Fig. 5B). Subtegulum
highly sclerotised. MA short, nearly rectangular, baso-retrolateral, with a slight apico-prolateral extension (Fig.
5B). Embolus apically with a prolateral curvature, with blunt tip, narrow in ventral view (Fig. 5A–B). Conductor
transparent, apico-retrolateral, rounded in ventral view (Fig. 5A–C).
Female (based on ADSH20196) (Fig. 1B, E). Like male except the following. Carapace brownish with pale
brown median band; thoracic part provided with narrow, longitudinal, pale brown medio-lateral bands. Fovea
(1.08) dark. Eye field brownish except between PLEs, which is pale brown. Clypeus, chelicerae, fangs, endites,
labium, and sternum brownish. Dorsum, laterals and venter of opisthosoma blackish-brown with inconspicuous
black spots and W-shaped markings; venter with paired median longitudinal bands formed of pale yellowish spots.
Spinnerets pale yellowish with black shades. Leg segments brownish. Body length 12.72. Carapace length 6.49,
width 5.11. Opisthosoma length 6.23, width 4.11. Eye diameters: ALE 0.21. AME 0.25. PLE 0.36. PME 0.47. Eye
interdistances: ALE–ALE 1.36. ALE–PME 0.09. AME–ALE 0.51. AME–AME 0.15. PLE–PLE 1.58. PME–PME
0.25. Clypeus height at AMEs 0.14, at ALEs 0.58. Chelicerae length 2.85. Measurements of palp and legs: palp
6.79 [2.46, 1.26, 1.43, 1.64], I 16.45 [4.74, 2.69, 4.15, 3.76, 1.11], II 15.20 [4.53, 2.55, 3.54, 3.48, 1.10], III 13.26
[4.05, 1.97, 2.87, 3.44, 0.93], IV 19.18 [5.37, 2.36, 4.30, 5.76, 1.39]. Leg formula: 4123. Spination of palp: femur
pld 2 do 2 rld 1, patella pl 1, tibia pl 2 rl 1 rld 2, tarsus pl 2 pld 1 rl 2; legs: femur I pld 2 plv 1 do 3 rld 3, II pl 2 pld
2 do 3 rld 3, III plv 4 do 4 rlv 4, IV pld 4 do 3, rld 1; patellae I–II spineless, III–IV pl 1 rl 1; tibiae I–II plv 5 rlv 5,
III pl 2 plv 3 do 2 rld 2 rlv 3, IV pl 2 plv 3 do 3 rl 2 rlv 3; metatarsus I plv 3 rlv 2, II plv 3 rlv 3, III pl 3 pld 1 plv 3
rl 2 rld 2 rlv 3, IV pl 3 pld 1 plv 3 rl 3 rld 2 rlv 4; tarsi I–IV spineless. Epigynum (Fig. 5D–F): epigynal plate flat,
nearly as wide as long, with distinct short, wide epigynal field (Fig. 5D, F). Copulatory openings postero-lateral to
the epigynal plate, masked by epigynal plate (Fig. 5E, arrows). Epigynal teeth simple, placed postero-laterally to
epigynal plate (Fig. 5D–E). Copulatory ducts short, wide (Fig. 5E); lateral folds well marked, slightly diagonally
oriented, continuous with epigynal teeth (Fig. 5E). Spermathecae peanut-shaped, obliquely oriented, close to each
other, with two differently sized chambers (Fig. 5E, 1 & 2). Fertilization ducts short, narrow, with slightly upturned
apical part (Fig. 5E).
Va ri at io n . Adult females (n=5): body length 10.43–12.72. The epigynum of the female specimen from the
Salim Ali Bird Sanctuary (ADSH20193) has wavy antero-lateral margins and its epigynal teeth are placed close to
the posterior epigynal border line (Fig. 5G).
Distribution. India (Kerala) (Fig. 6).
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FIGURE 6. Geographic distribution records of Ctenus cochinensis Gravely, 1931. Empty circles, new record; full circles,
record from literature.
Ctenus indicus Gravely, 1931
Figs 1C, F, 2C–E, 7, 8
Ctenus indicus Gravely, 1931: 234, fig. 2J–L (description and illustrations of female). Tikader & Malhotra 1981: 114, fig. 6A–
C (description and illustrations of female).
Type material. Two female syntypes of Ctenus indicus Gravely, 1931 from INDIA: Kerala: Palakkad:
Parambikulam (10
o
23'34.29''N, 76
o
46'32.24''E), 597 m alt.; F.H. Gravely leg.; 16–24 September 1914; repository
ZSI; voucher numbers 1505/18; unspecified number of female syntypes from Thrissur: Kavalai (10
o
16'23.89''N,
76
o
32'13.03''E), 390 m alt.; F.H. Gravely leg.; date and repository unknown; 1 female syntype from Tamil Nadu:
Ooty (formerly Ootacamund) (11
o
24'23.09''N, 76
o
41'35.68''E), 2246 m alt.; F.H. Gravely leg.; date and repository
unknown, all not examined.
Note. The species was identified using the illustrations given by Gravely (1931) and Tikader & Malhotra
(1981).
Material examined. INDIA, Kerala: Ernakulam, Kothamangalam, Salim Ali Bird Sanctuary (Thattekkadu
Bird Sanctuary),10
o
7'48.3''N, 76
o
41'43.24''E, 96 m alt., 22 May 2013, M.S. Pradeep leg., from ground, by hand: 1
subadult female (ADSH201031); Kollam, Kulathupuzha Forest Reserve, 8
o
54'6.37''N, 77
o
3'51.70''E, 134 m alt., 31
July 2013, M.S. Pradeep leg., from ground, by hand: 1 subadult female (ADSH201032) and 11 January 2015: 1
subadult female (ADSH201033); Pathanamthitta, Gavi, 9
o
26'9.07''N, 77
o
9'56.78''E, 1201 m alt., 21 December
2013, M.S. Pradeep leg., from ground, by hand: 1 subadult female, (ADSH201034); Trivandrum, Kallar in
Ponmudi, 8
o
45'35.79''N, 77
o
7'00.75''E, 920 m alt., 4 October 2014, M.S. Pradeep leg., from ground, by hand: 1
female (ADSH201035); Kollam, Thenmala, 8
o
57'30.7''N, 77
o
10'38.9''E, 567 m alt., 10 January 2015, M.S. Pradeep
leg., from ground, by hand: 1 female (ADSH201036).
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FIGURE 7. Ctenus indicus Gravely, 1931. A–C. Female genitalia: A. Epigynum, ventral; B. Internal genitalia, dorsal; C.
Epigynum, posterior. Abbreviations: EF epigynal field, EP epigynal plate, ET epigynal tooth, FD fertilization duct, LF lateral
fold of copulatory duct, S spermatheca. Scale bars: A–C, 1 mm. Arrows: 1 indicates concave plate, 2 straight plate, 3
copulatory opening. Digits 1 and 2, spermathecal chambers.
Diagnosis. Females of C. indicus can be separated from the females of other species by the following
combination of characters: epigynal teeth lying away from the lateral epigynal margin, epigynal plate with broad
anterior part and epigynum with strongly sclerotized lateral plates (Fig. 7A–C).
Redescription. Female (based on ADSH201037, Figs 1C, F, 2C–E). Carapace reddish-brown with broad pale
brown median band; thoracic part provided with narrow, longitudinal, pale brown medio-lateral bands and patches.
Fovea long (1.87), dark. Eye field blackish except between PLEs, which is pale brown. Clypeus, chelicerae, fangs,
endites, labium, and sternum brownish. Dorsum of chelicerae covered with long brownish hairs; promargin with a
thick layer of short, inwardly curved hairs, with three teeth; retromargin with four teeth. Sternum covered with
black hairs. Opisthosoma oval, hirsute, greyish-brown; dorsum antero-laterally with a pair of black patches.
Spinnerets brownish. Leg and palpal segments brownish to blackish-brown. All femora with white spots and
patches, femur IV retro-basally with a few small humps, each consists of numerous tiny, smooth tubercles (Fig.
2C–E); 2/3
rd
of tibia I and 1/3
rd
of tibia II scopulated, tibiae III and IV lack scopulae; all metatarsi with scopulae, I
and II entire, III and IV only on distal half; all tarsi scopulated, tarsal scopula complete. Body length 21.10.
Carapace length 10.92, width 8.76. Opisthosoma length 10.18, width 6.91. Eye diameters: ALE 0.32. AME 0.42.
PLE 0.51. PME 0.60. Eye interdistances: ALE–ALE 1.88. ALE–PME 0.25. AME–ALE 0.18. AME–AME 0.28.
PLE–PLE 2.28. PME–PME 0.32. Clypeus height at AMEs 0.28, at ALEs 1.04. Chelicerae length 4.77.
Measurements of palp and legs: palp 11.57 [4.13, 2.11, 2.69, 2.64], I 28.16 [8.01, 4.24, 7.32, 6.64, 1.95], II 26.59
[7.81, 4.16, 6.31, 6.42, 1.89], III 22.60 [6.71, 3.48, 4.78, 5.94, 1.69], IV 31.69 [8.70, 3.73, 7.11, 9.92, 2.23]. Leg
formula: 4123. Spination of palp: femur pld 2 do 2 rld 1, patella pl 1, tibia pl 2 rl 1 rld 1, tarsus pl 2 pld 1 rl 1 rld 1;
legs: femur I pld 2 plv 1 do 3 rld 3, II pl 1 pld 3 do 3 rld 3, III pld 4 do 3 rld 4, IV pld 4 do 3, rld 2; patellae I–II
spineless, III–IV pl 1 rl 1; tibiae I–II plv 5 rlv 5, III pl 2 pld 2 plv 3 do 1 rl 2 rlv 3, IV pl 2 plv 3 do 3 rl 2 rlv 3;
metatarsi I–II plv 3 rlv 3, III pl 3 pld 1plv 3 rl 3 rld 2 rlv 3, IV pl 2 pld 1 plv 3 rl 2 rld 3 rlv 4; tarsi I–IV spineless.
Epigynum (Fig. 7A–C): Median field of epigynal plate nearly triangular with circular epigynal field, with a median
bulging (Fig. 7A, C), with strongly sclerotized lateral plates, first concave, second straight (Fig. 7A, 1
st
& 2
nd
arrows); epigynal lateral folds moderately distinct (Fig. 7B), with a broad tooth (Fig. 7A). Copulatory openings
Zootaxa 4388 (3) © 2018 Magnolia Press
·
405
INDIAN CTENID SPIDERS
hidden behind proximal part of epigynal teeth (Fig. 7A, 3
rd
arrow). Copulatory ducts short, lying beneath the lateral
folds (Fig. 7B). Spermathecae nearly peanut-shaped, obliquely oriented, widely separated, with two differently
sized chambers (Fig. 7B). Fertilization ducts short, narrow with slightly upturned apical part (Fig. 7B).
Male: Unknown.
Distribution. India (Kerala, Tamil Nadu) (Fig. 8).
FIGURE 8. Geographic distribution records of Ctenus indicus Gravely, 1931. Empty circles, new records; full circles, records
from literature.
Acknowledgements
We are grateful to Rev. Fr. Prasanth Palackappillil CMI, Principal, Sacred Heart College, Thevara, Cochin for
providing all facilities for completing this work. We thank the Chief Conservator of Forests, Kerala State Forest
Division for the collecting permit. Many thanks to Dr M.J. Mathew, Sacred Heart College, Ernakulam, Kerala for
the distribution maps. We extend our heartfelt thanks to two anonymous reviewers for their constructive comments
on an earlier draft of the MS and to Dr Adalberto J. Santos (Universidade Federal de Minas Gerais, Brazil) for his
editorial efforts. We especially acknowledge the Science and Engineering Research Board (SERB)-DST, New
Delhi for providing funding support under the Major Research Project No. SR/SO/AS-99/2012.
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wsc.nmbe.ch (accessed 26 August 2017)
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... Female. For redescription and illustrations of the female, see Sankaran and Sebastian (2018). ...
... India: Kerala, Tamil Nadu (Gravely 1931;Tikader and Malhotra 1981;Sankaran and Sebastian 2018;Karthika and Sunil Jose 2020;Jäger 2022). ...
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A new genus in the spider family Ctenidae Keyserling, 1877 is described from Asia: Bowie gen. nov. belongs to the subfamily Cteninae and all members formerly described were placed so far in the genus Ctenus Walckenaer, 1805. After morphological as well as preliminary molecular characters were checked, it was clear that a new genus had to be erected to accommodate this predominantly Asian lineage of ground-dwelling spiders. As is the case in most Ctenidae generally, it was not easy to find apomorphic characters diagnosing this new taxon. Therefore, a combination of morphological characters is used to define all congeners. An important and newly introduced character in this respect is –among others– the fused patellar crack of the male palp. Forty-nine valid species are transferred to the new genus (first country/province records of particular species underlined): Bowie martensi (Jäger, 2012) comb. nov. (Nepal), B. bomdilaensis (Tikader & Malhotra, 1981) comb. nov. (India), B. indicus (Gravely, 1931) comb. nov. (India), B. cladarus (Jäger, 2012) comb. nov., (Myanmar), B. pingu (Jäger & Minn, 2015) comb. nov. (Myanmar), B. natmataung (Jäger & Minn, 2015) comb. nov., (Myanmar), B. sikkimensis (Gravely, 1931) comb. nov. (India), B. ramosus (Thorell, 1887) comb. nov. (Myanmar), B. goaensis (Bastawade & Borkar, 2008) comb. nov. (India), B. himalayensis (Gravely, 1931) comb. nov. (India), B. meghalayaensis (Tikader, 1976) comb. nov. (India), B. narashinhai (Patel & Reddy, 1988) comb. nov. (India), B. ceylonensis (F.O. Pickard-Cambridge, 1897) comb. nov. (Sri Lanka), B. andamanensis (Gravely, 1931) comb. nov. (Andaman Islands), B. kapuri (Tikader, 1973) comb. nov. (Andaman Islands), B. cochinensis (Gravely, 1931) comb. nov., (India), B. thorelli (F.O. Pickard-Cambridge, 1897) comb. nov. (Sri Lanka), B. lishuqiang (Jäger, 2012) comb. nov., (China: Sichuan), B. banna (Yao & Li in Chu et al. 2022) comb. nov. (China: Yunnan), B. theodorianum (Jäger, 2012), comb. nov. (Thailand, Laos, Vietnam), B. robustus (Thorell, 1897) comb. nov. (Myanmar, Thailand, Laos), B. yaeyamensis (Yoshida, 1998) comb. nov. (Taiwan), B. yulin (Yao & Li in Chu et al. 2022) comb. nov. (China: Yunnan), B. simplex (Thorell, 1897) comb. nov. (Myanmar, Thailand, Laos), B. bayeri (Jäger 2012) comb. nov. (Laos), B. holthoffi (Jäger, 2012) comb. nov. (Laos), B. saci (Ono, 2010) comb. nov. (Vietnam), B. floweri (F.O. Pickard-Cambridge, 1897) comb. nov. (Malaysia Peninsula), B. argentipes (Hasselt, 1893) comb. nov. (Malaysia Peninsula, Singapore, Indonesia: Sumatra), B. palembangensis (Strand, 1906) comb. nov. (Indonesia: Sumatra), B. angigitanus (Roewer, 1938) comb. nov. (Papua New Guinea), B. pulvinatus (Thorell, 1890) comb. nov. (Malaysia: Sarawak), B. hosei (F.O. Pickard-Cambridge, 1897) comb. nov. (Malaysia: Sarawak, Brunei), B. monaghani (Jäger, 2013) comb. nov. (Laos), B. javanus (Pocock, 1897) comb. nov. (Indonesia: Sumatra, Java, Bali), B. fungifer (Thorell, 1890) comb. nov. (Malaysia Peninsula), B. valvularis (Hasselt, 1882) comb. nov. (Indonesia: Sumatra), B. bicostatus (Thorell, 1890) comb. nov. (Malaysia: Sarawak), B. bantaengi (Merian, 1911) comb. nov. (Indonesia: Sulawesi), B. bowonglangi (Merian, 1911) comb. nov. (Indonesia: Sulawesi), B. celebensis (Pocock, 1897) comb. nov. (Indonesia: Sulawesi), B. sagittatus (Giltay, 1935) comb. nov. (Indonesia: Sulawesi), B. kochi (Simon, 1897b) comb. nov. (Indonesia: West Papua), B. sarawakensis (F.O. Pickard-Cambridge, 1897) comb. nov. (Malaysia: Sarawak), B. philippinensis (F.O. Pickard-Cambridge, 1897) comb. nov. (Philippines), B. aruanus (Strand, 1911) comb. nov. (Indonesia: Maluku), B. angularis (Roewer, 1938) comb. nov. (Indonesia: Maluku), B. rufisternis (Pocock, 1898) comb. nov. (Papua New Guinea: New Britain), and B. corniger (F.O. Pickard-Cambridge, 1898) comb. nov. (South Africa). For thirty-two species, illustrations of their respective copulatory organs, as well as habitus photos, are provided. Fifty-five new species are described, these are listed, together with the already described species, according to their geographic occurrence and to their affiliation to species groups as far as the latter could be recognised (type species indicated by an asterisk): Bowie hunkydory spec. nov. (Nepal), B. ziggystardust spec. nov. (Nepal), B. ladystardust spec. nov. (Nepal), B. aladdinsane spec. nov. (India), B. majortom spec. nov. (Nepal), B. jeangenie spec. nov. (India), B. heroes spec. nov. (India), B. fascination spec. nov. (Vietnam), B. low spec. nov. (Thailand), B. dodo spec. nov. (Vietnam), B. stationtostation spec. nov. (Myanmar), B. candidate spec. nov. (Vietnam), B. diamonddogs spec. nov. (Vietnam), B. yassassin spec. nov. (Taiwan), B. bemywife spec. nov. (Thailand), B. subterraneans spec. nov. (Thailand), B. afterall spec. nov. (Thailand), B. warszawa spec. nov. (Thailand), B. artdecade spec. nov. (Cambodia), B. bigbrother spec. nov. (Vietnam), *B. rebelrebel spec. nov. (Vietnam), B. youngamericans spec. nov. (Vietnam), B. right spec. nov. (Vietnam), B. stay spec. nov. (Vietnam), B. fame spec. nov. (Vietnam), B. win spec. nov. (Vietnam), B. joethelion spec. nov. (Malaysia Peninsula), B. mossgarden spec. nov. (Malaysia Peninsula), B. neukoeln spec. nov. (Malaysia Peninsula), B. scarymonsters spec. nov. (Indonesia: Sumatra), B. teenagewildlife spec. nov. (Indonesia: Sumatra), B. letsdance spec. nov. (Indonesia: Java), B. crystaljapan spec. nov. (Indonesia: Sumatra), B. tonight spec. nov. (Malaysia: Sarawak), B. catpeople spec. nov. (Malaysia: Sabah), B. ricochet spec. nov. (Indonesia: Kalimantan), B. fashion spec. nov. (Malaysia Peninsula), B. withinyou spec. nov. (Malaysia: Sarawak), B. abdulmajid spec. nov. (Singapore), B. blackout spec. nov. (Malaysia Peninsula), B. modernlove spec. nov. (Malaysia: Sabah), B. chinagirl spec. nov. (Malaysia: Sabah), B. withoutyou spec. nov. (Malaysia: Sabah), B. magicdance spec. nov. (Indonesia: Sulawesi), B. bluejean spec. nov. (Malaysia: Sabah), B. criminalworld spec. nov. (Malaysia: Sabah), B. shakeit spec. nov. (Malaysia: Sabah), B. ashestoashes spec. nov. (Indonesia: Kalimantan), B. underground spec. nov. (Indonesia: Kalimantan), B. lodger spec. nov. (Philippines), B. redsails spec. nov. (Philippines), B. thenextday spec. nov. (Indonesia: Papua), B. lazarus spec. nov. (Papua New Guinea), B. thiesi spec. nov. (Papua New Guinea) and B. blackstar spec. nov. (Papua New Guinea). Formerly unknown sexes are described for the first time for the following species: Bowie martensi comb. nov. (female), B. indicus comb. nov. (only male RTA tip), B. narashinhai comb. nov. (male), B. argentipes comb. nov. (female) and B. celebensis comb. nov. (male). Fourteen species groups are proposed on the basis of morphological characters: cladarus-species group (17 species), robustus-species group (14 species), bemywife-species group (2 species), rebelrebel-species group (10 species), youngamericans-species group (3 species), floweri-species group (3 species), scarymonsters-species group (2 species), teenagewildlife-species group (2 species), argentipes-species group (10 species), javanus-species group (5 species), chinagirl-species group (11 species), shakeit-species group (5 species), lodger-species group (3 species) and blackstar-species group (7 species). Thus, 93 species are grouped, leaving 11 species without a current assignment to any of these groups. Ctenus kandyensis Kim & Ye, 2014 syn. nov. is recognised as a junior synonym of Bowie thorelli comb. nov., and C. calcarifer F.O. Pickard-Cambridge, 1902 syn. nov. as junior synonym of B. sarawakensis comb. nov. Five species previously placed in Ctenus are considered nomina dubia, four of them are transferred to Bowie gen. nov., one to Nilus: Bowie barbatus (Thorell, 1895) comb. nov., B. tumidulus (Simon, 1887b) comb. nov., B. flavidus (Hogg, 1922) comb. nov., B. pollii (Hasselt, 1893) comb. nov. and Nilus marginatus (Walckenaer, 1847) comb. nov. Beside the latter species, three species are transferred from Ctenus, all of them to Anahita: A. periculosa (Bristowe, 1931) comb. nov., A. dangsa (Reddy & Patel, 1994) comb. nov. and A. tuniensis (Patel & Reddy, 1988) comb. nov. All in all, Bowie gen. nov. is the second largest genus within the family Ctenidae, with 108 species in total including nomina dubia. It occurs from Nepal in the Himalayas and South India across large parts of South and South East Asia to Papua New Guinea. One undescribed species is known from northern Australia (Queensland). Representatives are known to live in the leaf litter of forests, with most species having small distribution ranges, usually occurring within a 100 km radius. With this revision, the family Ctenidae contains now 586 species and 48 genera, and the number of species assigned to the genus Ctenus, so far used as nomenclatural “waste bin”, is reduced to 164.
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A cladistic analysis based on parsimony is undertaken to test hypotheses concerning the monophyly of the ctenid spiders and the relationships among their various clades. The data matrix comprised a total of 98 species representing 16 families scored for 146 characters, with all but six taken from various morphological systems; the remaining are behavioral attributes. Ctenidae is shown to be polyphyletic as currently delimited, but the most severely questioned taxa (i.e., Acanthoctenus, Cupiennius, and Ancylometes) are indeed members of this family. The so-called ctenid eye pattern is shown to be a synapomorphy uniting a more restricted set of taxa. This clade is recognized as Ctenidae sensu stricto and it comprises at least five major lineages: Acantheinae, Acanthocteninae, Calocteninae, Cteninae, and Viridasiinae. The latter taxa are hypothesized to be the sister group of all other ctenids. In turn, a clade formed by Miturgidae s.s. and Zoridae s.s. is proposed as the sister group of Ctenidae. The cladistic analysis also refutes the monophyly of Ctenus, Anahita, Enoploctenus, Celaetycheus, and Leptoctenus. The following taxonomic changes are proposed: Anahita isaloensis Ono, 1993 is transferred to Vulsor Simon, 1888; Diallomus Simon, 1897, currently placed in Zoridae, is transferred to Ctenidae; Tunabo Chamberlin, 1916 is synonymized with Neoctenus Simon, 1897; Neoctenus is removed from Zoridae and transferred to Trechaleidae; Xenoctenus Mello-Leitão, 1938 is removed from the synonymy of Tunabo and considered incertae sedis, within Ctenoidea.
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The new ctenid genus Afroneutria is proposed to include five species: Afroneutria velox (Blackwall, 1865) comb. nov. (type species), A. immortalis (Arts, 1912) comb. nov., A. erythrochelis (Simon, 1876) comb. nov., A. quadrimaculata sp. nov. and A. hybrida sp. nov., all collected in central to southeast Africa. Afroneutria species can be distinguished from the remaining Ctenidae by the presence of a synapomorphic laminar projection on the embolus tip. The genus can also be distinguished by the presence of the following set of characters: elongated cup-shaped median apophysis and elongated and laminar embolus on the male palp; large lateral projections on the epigynum and the spermathecae divided into two parts, dorsal and ventral, in the vulva.
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A cladistic analysis based on parsimony was performed to test the monophyly of Cteninae. The data matrix comprises 72 terminal taxa scored for 88 characters from external morphology, female internal genitalia, and two behavioural characters. Cteninae, as currently delimited, is polyphyletic. The monoplyly of Cteninae requires the transfer of two Ctenus species to Enoploctenus (Acantheinae) and two Ctenus species to Asthenoctenus (Viridasiinae). Ctenus, the genus type, comprises approximately half of the described species of the family, and is polyphyletic as currently delimited. From the 32 Ctenus species added to this analysis, only 13 are recognized as Ctenus s.s.: (C. rectipes (C. pauloterrai (C. manauara, C. nigritus ((C. dubius, C. crulsi) ((C. amphora, C. minor) (C. medius, C. paubrasil (C. fernandae (C. ornatus, C. vehemens)))))))). The results suggest that at least eight new genera should be described to accommodate species misplaced in Ctenus. The result also refutes the monophyly of Amauropelma Raven, Stumkat & Gray, 2001, Celaetycheus Simon, 1897, Leptoctenus L. Koch, 1878, and Thoriosa Simon, 1910. The following taxonomic changes are proposed: Ctenus inazensis Strand, 1909 and Ctenus miserabilis Strand, 1916 are transferred to Enoploctenus; Ctenus tarsalis F.O. Pickard-Cambridge, 1902 and Ctenus bulimus Strand, 1909 are transferred to Asthenoctenus Simon, 1897; Celaetycheus modestus Bryant, 1942 is transferred to Ohvida Polotow & Brescovit, 2009.
Article
The African Ctenidae in the Collections of the British Museum (Natural History) are revised. Thirty species are contained in the subfamilies Cteninae and Acantheinae. Ctenus batesii Pocock, 1903, is a synonym of Ctenus capulinus (Karsch), 1879, Ctenus dirus Thorell, 1899, is a synonym of Ctenus kingsleyi F. O. P.-Cambridge, 1898, Ctenus marshalli F. O. P.-Cambridge, 1898, is a synonym of Ctenus melanogastra (Bösenberg & Lenz), 1895, and Ctenus spenceri F. O. P.-Cambridge, 1898, is a synonym of Ctenus pulchriventris Simon, 1896.The subfamily Acantheinae is recorded from Africa for the first time. A new genus, Africactenus, is erected for a group of allied species. The genotype is Ctenus agilior Pocock, 1899, and the following new species are described:—A. depressus, A. desartsi, A kribiensis, A. pococki, A. sandersoni, A. simoni, A. sladeni, A. tridentatus and A. trilateralis. Ctenus longurio Simon, 1909, is transferred to the genus Africactenus on examination of the type-material.Certain species from the Simon Collection are listed in the Appendix and figured for the first time.
Abbreviations: EF epigynal field, EP epigynal plate, ET epigynal tooth, FD fertilization duct, LF lateral fold of copulatory duct, S spermatheca
  • Posterior Epigynum
Epigynum, posterior. Abbreviations: EF epigynal field, EP epigynal plate, ET epigynal tooth, FD fertilization duct, LF lateral fold of copulatory duct, S spermatheca. Scale bars: A-C, 1 mm. Arrows: 1 indicates concave plate, 2 straight plate, 3 copulatory opening. Digits 1 and 2, spermathecal chambers.
Leg formula: 4123. Spination of palp: femur pld 2 do 2 rld 1, patella pl 1, tibia pl 2 rl 1 rld 1, tarsus pl 2 pld 1 rl 1 rld 1; legs: femur I pld 2 plv 1 do 3 rld 3
  • Metatarsi With Scopulae
2/3 rd of tibia I and 1/3 rd of tibia II scopulated, tibiae III and IV lack scopulae; all metatarsi with scopulae, I and II entire, III and IV only on distal half; all tarsi scopulated, tarsal scopula complete. Body length 21.10. Carapace length 10.92, width 8.76. Opisthosoma length 10.18, width 6.91. Eye diameters: ALE 0.32. AME 0.42. PLE 0.51. PME 0.60. Eye interdistances: ALE-ALE 1.88. ALE-PME 0.25. AME-ALE 0.18. AME-AME 0.28. PLE-PLE 2.28. PME-PME 0.32. Clypeus height at AMEs 0.28, at ALEs 1.04. Chelicerae length 4.77. Measurements of palp and legs: palp 11.57 [4.13, 2.11, 2.69, 2.64], I 28.16 [8.01, 4.24, 7.32, 6.64, 1.95], II 26.59 [7.81, 4.16, 6.31, 6.42, 1.89], III 22.60 [6.71, 3.48, 4.78, 5.94, 1.69], IV 31.69 [8.70, 3.73, 7.11, 9.92, 2.23]. Leg formula: 4123. Spination of palp: femur pld 2 do 2 rld 1, patella pl 1, tibia pl 2 rl 1 rld 1, tarsus pl 2 pld 1 rl 1 rld 1; legs: femur I pld 2 plv 1 do 3 rld 3, II pl 1 pld 3 do 3 rld 3, III pld 4 do 3 rld 4, IV pld 4 do 3, rld 2; patellae I-II spineless, III-IV pl 1 rl 1; tibiae I-II plv 5 rlv 5, III pl 2 pld 2 plv 3 do 1 rl 2 rlv 3, IV pl 2 plv 3 do 3 rl 2 rlv 3;