A revision of the endemic South African long-jawed ground spider genus Drassodella Hewitt, 1916 (Araneae: Gallieniellidae) Author Mbo, Zingisile Author Haddad, Charles R. text Zootaxa 2019 2019-04-10 4582 1 1 62 journal article 27120 10.11646/zootaxa.4582.1.1 c79b6327-48fb-47bf-b17b-d20d2de295e3 1175-5326 2643724 9DC61794-4BD7-4F6D-BB8C-84D9855C8151 Genus Drassodella Hewitt, 1916 Drassodella Hewitt, 1916 : 209 ; Tucker, 1923 : 309 ; Roewer, 1954 : 384 , 1467; Bonnet, 1956 : 1556 ; Platnick, 1989 : 457 ; Dippenaar-Schoeman & Jocqué, 1997 : 166 ; Jocqué & Dippenaar-Schoeman, 2006 : 126 . Type species: Drassodella salisburyi Hewitt, 1916 , by original designation. Diagnosis: Drassodella can be recognised from Austrachelas by the following: 1) carapace almost always with an median upsilon marking of white feathery setae and the lateral margins with a narrow fringe of white feathery setae, which are absent in Austrachelas ; 2) usually with striped or spotted dorsal markings on the abdomen, which are rarely absent (e.g. Figs 1–4 ), while Austrachelas have chevron markings (see Haddad et al. 2009 : figs 1–15); 3) shield-shaped sternum ( Fig. 26 ), while oval in Austrachelas (see Haddad et al. 2009 : fig. 17); 4) anterior legs with weakly scopulate metatarsi and tarsi ( Figs 34 , 39–42 ), while densely scopulate in Austrachelas (see Haddad et al. 2009 : figs 24–26, 33); 5) posterior metatarsi with the pro- and retrolateral dorsal spines occurring singly (i.e. pl 3 rl 3), while usually paired in Austrachelas (i.e. pl 6 rl 4-6). FIGURES 1–4. General habitus photographs of live Drassodella spiders: 1 D. amatola sp. nov. female from Hogsback, Eastern Cape; 2 D. quinquelabecula Tucker, 1923 female from Aardvark Nature Reserve, Western Cape; 3 D. septemmaculata (Strand, 1909) male from Fisherhaven, Western Cape; 4 D. vasivulva Tucker, 1923 male from De Hoop Nature Reserve, Western Cape. Photographs 1, 3, 4 by C. Haddad, 2 by P. Webb. Description: Small to medium spiders, body length 2.56–9.85 mm . Carapace broad, oval, cephalic region slightly narrowed, broadest between coxae II and III ( Figs 5–8 ); in lateral profile raised evenly along midline, sloping gently posteriorly ( Fig. 9 ); posterior margin rebordered, moderately concave above pedicel ( Figs 10 , 18 ); eight eyes in two rows (4:4); lateral eyes set on low tubercles; all eyes surrounded by black rings; AER slightly recurved when viewed dorsally ( Figs 5 , 19 ), straight or slightly recurved when viewed from front; AME smaller or equal to ALE; AME separated by more than half their diameter, closer to ALE than to each other; PER slightly recurved, broader than AER ( Fig. 19 ); PME flattened, transversely oval, more or less the same distance between each other and PLE; PME smaller than PLE. Clypeus slightly larger than AME diameter, with few long erect setae; surface smooth, sparsely covered in short straight and feathery setae; usually black, rarely yellow-orange or brown with reticulate black mottling, usually with white feathery setae forming an inverted Y-shaped (upsilon) dorsal marking along midline, with split posterior to fovea ( Fig. 18 ); lateral margins with narrow fringe composed of white feathery setae ( Figs 20, 21 ), rarely without dorsal markings; several long erect setae scattered between fovea and posterior margin ( Fig. 18 ); fovea short to elongate, narrow. FIGURES 5–17. Digital microscope photographs of Drassodella somatic morphology: 5–8 dorsal view of carapace and chelicerae of D. flava sp. nov. female (5), D. guttata sp. nov. male (6), D. transversa sp. nov. male (7) and D. purcelli Tucker, 1923 female (8); 9 D. tenebrosa Lawrence, 1938 female, carapace, lateral view; 10 D. purcelli female, posterodorsal view of carapace, showing pedicel and sclerites; 11 same, ventral pedicel sclerite, ventral view; 12–14 dorsal view of abdomens of D. septemmaculata (Strand, 1909) male (12), D. aurostriata sp. nov. male (13) and D. amatola sp. nov. female (14); 15 D. tenebrosa female, abdomen, anterodorsal view; 16 D. septemmaculata male, abdomen, showing sclerites on venter; 17 D. tenebrosa female, spinnerets, posteroventral view. Abbreviations: AAS—anterior abdominal sclerite; CGS—cylindrical gland spigots; CON—condyle; DPS—dorsal pedicel sclerite; IS—inframamillary sclerite; PB—pleural bars; PES—post-epigastric sclerite; PPS—posterior pleural sclerite; PT—petiole; SC—dorsal scutum; SI—sigillum; VPS—ventral pedicel sclerite. Chelicerae usually moderately protruding in males, slightly angled and/or vertically orientated in females, slightly tapering distally ( Figs 5–8 ); surface finely granulate, with many moderately long erect setae on anterior surface of paturon; cheliceral escort seta ( sensu Ramírez 2014 : fig. 15B) present, promarginally near fang base ( Fig. 22 ); condyle (boss) small to large ( Fig. 9 ); fangs moderately elongate ( Fig. 22 ), with fine ridges near opening of venom ducts; promargin with three teeth, retromargin with two teeth ( Fig. 23 ). Chilum single, distinct, triangular in shape; endites obliquely depressed, very broad, slightly sinuous, widest medially, without median groove, with distinct dense maxillar hair tuft and serrula ( Figs 24, 25 ); labium slightly longer than broad or as long, posterolateral margins of labium with distinct sclerotized groove; anterior portion slightly inclined, posterior portion short; anterior margin with erect setae, slightly concave medially. FIGURES 18–26. Scanning electron microscope photographs of Drassodella guttata sp. nov. female (18–21, 24–26) and male (23), and D. tolkieni sp. nov. female (22): 18 posterior half of carapace, dorsal view, indicating fovea, erect and feathery setae, and rebordered concave posterior margin; 19 eye region, dorsal view; 20 feathery setae on posterior carapace margin and coxa IV; 21 same, feathery setae enlarged; 22 chelicerae, ventral view; 23 chelicerae teeth; 24 endites and labium, ventral view; 25 serrula on endites, ventral view; 26 sternum. Abbreviations: CES—promarginal escort seta; ES—erect setae; FS—feathery setae; MHT—maxillar hair tuft. Sternum shield-shaped, longer than broad, with blunt apex posteriorly ( Figs 11 , 26 ); anterior margin concave or straight, widest at coxae II. Intercoxal sclerites present between coxae I & II, II & III, III & IV; precoxal triangles present; pleural bars weakly to moderately sclerotised, isolated or fused ( Fig. 9 ). Pedicel composed of broad dorsal sclerite and inverted Y- or V-shaped ventral sclerite ( Figs 9–11 ). FIGURES 27–38. Scanning electron microscope photographs of male (27, 28, 35–38) and female (29–31, 34) Drassodella guttata sp. nov. and female D. tolkieni sp. nov. (32, 33). 27 femur I, curved long ventral setae, lateral view; 28 femur IV, curved long ventral setae, lateral view; 29 leg II, patellar indentation, retrolateral view; 30 same, detail of lyriform organ; 31 lyriform organ at distal end of tibia II, retrolateral view; 32 metatarsus III with spine, feathery and chemosensory setae, and trichobothria, dorsal view; 33 distal end of metatarsus III, dorsal view; 34 metatarsus III, scopula, prolateral view; 35 distal end of tarsus I, dorsal view; 36 tarsus III, trichobothrium; 37 tarsus I, chemosensory seta; 38 same, tarsal organ. Abbreviations: CS—chemosensory setae; FS—feathery setae; MS—metatarsal stopper; SP—spine; SS—slit sensilla; TC—tarsal claws; TOtarsal organ; TR—trichobothria. Legs long and moderately thin ( Figs 1–4 ), leg formula 4123; anterior pairs of legs weakly spined, posterior pairs of legs moderately to strongly spined; trochanters notched; femora with curved elongate ventral setae ( Figs 27, 28 ); patellar indentation narrow, with large lyriform organ at proximal end ( Figs 29, 30 ); tibiae with distal lyriform organ retrolaterally ( Fig. 31 ); tibiae, metatarsi and tarsi with several trichobothria, feathery and chemosensory setae ( Figs 31–33, 35–37 ); posterior legs with single or paired ventral metatarsal spines; feathery setae present on all leg segments (e.g. Figs 29, 33, 34 ), usually dorsally and laterally situated on metatarsi and tarsi; metatarsi and tarsi weakly scopulate ( Figs 34 , 39–42 ), anterior metatarsi usually scopulate in distal half only, tarsi scopulate for entire length; tarsal organ oval and slightly elevated ( Fig. 38 ); tarsi with paired dentate claws, usually with 3-5 teeth, situated dorsally to claw tufts comprising tenent setae ( Figs 35 , 43 ); claw tufts denser on anterior ( Fig. 40 ) than posterior tarsi ( Fig. 43 ); females with ventral row of erect setae on palpal femur. FIGURES 39–50. Scanning electron microscope photographs of female (39, 41, 44, 46, 48–50) and male (43) Drassodella guttata sp. nov. , and female D. tolkieni sp. nov. (40, 42, 45, 47): 39 tarsus III, scopula, ventrolateral view; 40 tarsus II, claws and scopulate setae, lateral view; 41 detail of scopulate setae, ventral view; 42 same, lateral view; 43 tarsus III, detail of tarsal claws and associated setae, arrow indicating tenant setae forming claw tuft; 44 dorsal abdominal feathery setae, arrow indicating white patch composed of dense modified feathery setae; 45 ventral abdominal feathery setae; 46, 47 spinnerets, general view; 48 left ALS, ventral view; 49 left PMS, ventral view; 50 left PLS, ventral view. Abbreviations as follows: Acaciniform gland spigot(s); Cy—cylindrical gland spigot(s); maAmp—major ampullate gland spigot(s); miAmp—minor ampullate gland spigot(s); nu—nubbin(s); Pi—piriform gland spigot(s). Abdomen oval-elongate, with yellow, white or cream dorsal stripes or spots, or without markings, on black or grey background ( Figs 1–4 , 12–14 ); dorsum covered in short erect and feathery setae, markings comprising dense feathery setae ( Fig. 44 ); dorsum with two pairs of small sigilla, posterior pair more widely separated ( Figs 12, 14 ), often with additional pair of sclerites on anterior margin above pedicel ( Fig. 15 ) and laterally above booklungs; males with short dorsal abdominal scutum anteriorly, absent in females; several fine straight erect setae on anterior margin above pedicel; pale ventrally ( Fig. 16 ), with one or two lines of tiny sclerites usually present, sometimes absent, with few scattered feathery setae between short straight setae ( Fig. 45 ); epigastric region weakly sclerotised, and post-epigastric sclerites weakly to strongly sclerotized ( Fig. 16 ), between short straight setae. Spinnerets (analysed with SEM observations only in females of D. guttata sp. nov. and D. tolkieni sp. nov. ): ALS conical, nearly cylindrical, not widely separated, with sclerotized apical segment ( Figs 17 , 47 ); single major ampullate gland spigot and numerous piriform gland spigots present ( Fig. 48 ); PMS with two ( Fig. 17 ) or three rows of cylindrical gland spigots, single minor ampullate gland spigot and many aciniform gland spigots ( Fig. 49 ); PLS with aciniform gland spigots, cylindrical gland spigots, single minor ampullate gland spigot and nubbins ( Fig. 50 ); PLS longer and more widely separated than ALS ( Fig. 17 ). FIGURES 51–59. Scanning electron microscope photographs of female (51, 52) and male (55–59) Drassodella guttata sp. nov. , and female D. tolkieni sp. nov. (53, 54): 51 epigyne, slightly ventrolateral view; 52 detail of epigynal muscle sigilla; 53 epigyne, dorsal view; 54 detail of glandular pores on spermathecal head; 55 right palp, prolateral view; 56 same, distal end of palp and cymbial scopula; 57 left palp, ventral view; 58 detail of distal end of tegulum, right palp, prolateral view; 59 same, left palp, ventral view. Abbreviations: AH—anterior hood; CD—copulatory duct; CO—copulatory opening; CS—cymbial scopula; DP—dorsal embolic prong; EM—embolus; EMS—epigynal muscle sigilla; LH—lateral hood; MA—median apophysis; MSmedian septum; ST—spermathecae; VP—ventral embolic prong. Female epigynes with distinct anterior hoods, lateral hoods only present in D. melana species group ( Fig. 51 ); anterior hood transverse, median septum broad and subrectangular in D. melana species group ( Figs 60–69 ), variable but generally more longitudinal in D. salisburyi species group ( Figs 70–77 ); epigyne muscle sigilla ( Figs 51, 52 ) broad or elongate, sometimes absent; longitudinal ridge of median septum sometimes broadened posteriorly; copulatory openings forming narrow slit laterally on median septum ( Fig. 51 ); copulatory ducts very short, entering ST II along mesal margin ( Fig. 53 ); ST II anterior to ST I, with broad, medially-curving duct linking them ( Figs 60–69 ); ST II with widely spaced glandular pores on spermathecal head ( Fig. 54 ). Male palp with broad oval cymbium, tapering distally, with sparse to dense cymbial scopula present dorsally ( Figs 55–57 , 78–92 ); embolus in D. melana species group originating prolaterally, forming broad arch, with fine retrolateral tip ( Figs 57–59 , 78–85 ), subtriangular and variably orientated in D. salisburyi species group ( Figs 86– 92 ); embolus with one or two basal prongs prolaterally in D. melana species group ( Figs 57–59 ), absent in D. salisburyi species group; median apophysis hook-shaped in D. melana species group ( Fig. 59 ), subtriangular or forming stout hook in D. salisburyi species group ( Figs 86–92 ); retrolateral tibial apophysis simple, triangular.